Near-shore bathymetry and reef biotopes of Henderson Island, Pitcairn Group

The coral reef which surrounds two thirds of Henderson Island, Pitcairn Group, consists of a reef flat, reef margin and fore-reef which slopes gradually into deeper water. The range of sublittoral habitats provided by this topography is limited, reflecting a low level of coral diversity. Spur and groove formations of the fore-reef are present around the northern end of the island which contrasts with the south-western corner, where, in the absence of a fringing reef platform, subaerial cliff faces are fronted by a cliff foot submarine 'trench'. In general, live coral cover on the fore-reef was estimated at between 10–30%, though at sites off the east coast it reached 80%. The associated communities, particularly of sessile filter-feeding groups, were found to be impoverished. The two factors of biogeographical isolation and of local bioerosion processes are proposed as being the main reasons for the depauperate nature of the island's reef. Brief notes are included on the lagoon and fore-reef bathymetry and habitats of Ducie and Oeno Atolls for comparative purposes.     

Macroborers within coral framework at Discovery Bay, north Jamaica: species distribution and abundance, and effects on coral preservation

 Macroboring organisms are recognised as key agents of reef framework modification and destruction, and while recent studies in the Pacific have improved understanding of spatial variations in macroboring community structure, and rates of macroboring within individual reefs, comparable studies from the Caribbean are largely lacking. This study assesses the distribution of macroboring species and the degree of framework infestation across the reefs at Discovery Bay, north Jamaica. Although individual species of borers exhibit variable distributions across the reef, relative abundances of the main groups of macroborers (sponges, bivalves, worms) illustrate clear distributional trends. Sponges are dominant at fore-reef sites, while sipunculan and polychaete worms are only of importance at back-reef/lagoon and shallow fore-reef sites. Bivalves are locally important within back-reef and lagoon patch reef framework. Average percentages of internal bioerosion (macroboring) vary between sites, but are highest at back-reef and deep fore-reef sites. No systematic pattern of variation occurs within back-reef/lagoon samples, but a significant trend of increased macroboring is recognised with increased water depth on the fore-reef. In addition, significant differences in terms of the susceptibility of individual coral species are recognised. These factors are likely to result in biasing of the fossil record, with variable styles of preservation evident both between sites (i.e. with depth/environment) and within sites (i.e. between coral species). Accepted: 1 June 1998     

Sub-rubble communities of Curaçao and Bonaire coral reefs

The distribution and abundance of sessile organisms under coral rubble has been studied at Bonaire and Curaçao, Netherlands Antilles. Species richness under rubble is extremely high with at least 367 species of which sponges, tunicates and bryozoans are the most important. Shallow sub-rubble communities can be considered refuges as the majority of these species are crypt-obligate. Sub-rubble communities may also have a preserve function for sponges, but do not harbour enough corals to ensure a quick coral recolonization of the reef surface after a major disaster. Cryptic community composition is affected by depth and pollution, and differs substantially between the two neighbouring islands, possibly as a result of different bottom characteristics. Biomass of the sub-rubble communities may contribute considerably to total reef biomass. Diversity varies inversely with increased depth and increased rubble size, possibly indicating abiotic control (e.g. physical disturbance by wave action and reef slope substrate collapse).     

Holocene coral reef rubble and its binding agents

A literature review regarding reef rubble (defined as mechanically or chemically abraded parts of framebuilders or reef rock larger than sand fraction) and its binding agents is presented. Rubble is produced by natural and man-made events such as storms, wave agitation, earthquakes, bioerosion, ship groundings, and dynamite fisheries. The regeneration of reefs after rubble-forming processes requires rigid rubble binding, which is always preceded by preliminary stabilization. Preliminary stabilization can be achieved by a decline in hydrodynamic energy, interlocking of components, seagrass, and overgrowth by sponges or algae. Rigid binding is primarily achieved by diagenetic cementation. The literature indicates that binding by coralline algae or other organisms (corals, worms, bryozoans) is only of subordinate importance. Highest rates of rigid rubble binding are known from fore-reef areas with low sloping angles above fair-weather wave base; rigid rubble binding is particularly rare in deeper fore-reef environments and not described from the reef crest. Rigid binding by diagenetic cementation is generally known from inter- and supratidal near-shore ramparts as well as back-reef, reef-flat, and shallow fore-reef rubble accumulations, while coralline algae rigidly bind rubble only in very shallow fore-reef environments. Rubble binding does not appear to be easily achieved and fewer reports of bound rubble were found than of loose rubble.     

Halimeda growth and diversity on the deep fore-reef of Enewetak Atoll

The deep fore-reef at Enewetak has been examined from the submersible Makali'i. Green algae grow to about-150 m at photon flux densities of approximately 1 Em^-2s^-1. Halimeda cover is 50% at many sites down to-90 m. Halimeda populations are important within the zone of scleractinian corals down to about-65 m, while a Halimeda zone with low coral cover or lacking corals between-65 m and-150 m probably is an important source of reef carbonate. Halimedas of the deep fore-reef, like those of the lagoon, constitute an important structural component in reef building. Other calcareous green algae such as Tydemania are less important on the deep fore-reef, but growth of coralline red algae continues to over-200m. Halimeda diversity is high down to near the base of the euphotic zone.     

Will increased storm disturbance affect the biodiversity of intertidal,nonscleractinian sessile fauna on coral reefs?

Relatively little is known about how the future effects of climatic change, including increases in sea level, temperature and storm severity and frequency, will impact on patterns of biodiversity on coral reefs, with the notable exception of recent work on corals and fish in tropical reef ecosystems. Sessile invertebrates such as ascidians, sponges and bryozoans occupying intertidal rubble habitats on coral reefs contribute significantly to the overall biodiversity and ecosystem function, but there is little or no information available on the likely impacts on these species from climate change. The existing strong physical gradients in these intertidal habitats will be exacerbated under predicted climatic change. By examining the distribution and abundance of nonscleractinian, sessile invertebrate assemblages exposed to different levels of wave action and at different heights on the shore around a coral reef, we show that coral reef intertidal biodiversity is particularly sensitive to physical disturbance. As physical disturbance regimes increase due to more intense storms and wave action associated with global warming, we can expect to see a corresponding decrease in the diversity of these cryptic sessile assemblages. This could impact negatively on the future health and productivity of coral reef ecosystems, given the ecosystem services these organisms provide.     

Coral community structure and risk assessment of high-latitude reefs at Sodwana Bay,South Africa

The importance of studying coral communities at different spatial scales is acknowledged in a growing volume of scientific literature, and principles of landscape ecology were thus used to elucidate the patterns in coral community structure on the high-latitude reefs in South Africa. These reefs are at the southernmost distribution of this fauna in Africa, are surprisingly species rich, and represent a biodiversity peak in this fauna south of the equator, regardless of the marginal nature of the environment. Coral community patterns were identified on and between the reefs at Sodwana Bay, justifying the grouping of reef areas in distinct zones. A number of landscape components were identified, ranging from the entire reef complex (10 km scale), individual reefs (1 km scales) and reef zones, to components that were separated using multivariate statistical analysis of transect data. These components transcended spatial similarities, e.g. the fore-reef on Five-mile Reef was not similar to the fore-reef on Seven-mile Reef, but was rather grouped with the reef flat on Two-mile Reef. This information was “translated” into an index of management intervention, based on risk assessment, and was generated using parameters that measure susceptibility to crown-of-thorns feeding, bleaching, diver-related damage and swell-induced breakage. We also assessed was the time elapsed since the last major disturbance and the proximity to the only boat launch site, a proxy measure of continuous disturbance. The risk assessment suggested that conservation management is most needed in the stable and “climax” coral communities that are usually characterised by a near-equal mix of hard and soft corals at maximal coral species diversity.     

Spatial heterogeneity of benthic community assemblages with an emphasis on reef algae at French Frigate Shoals, Northwestern Hawai‘ian Islands

Reefs in tropical atoll systems have historically been described on a geomorphic basis, and segregated into loosely defined fore-reef, back-reef, and lagoonal reef zones. However, recent oceanographic monitoring data have shown that physical factors within a single geomorphic zone can vary significantly, calling into question whether benthic communities within a single zone are biologically similar. To determine the amount of benthic variability that may occur in a geomorphic zone, percent cover of benthic organisms was determined at the species level for 28 sites in three geomorphic zones at French Frigate Shoals, Northwestern Hawai‘ian Islands. Multivariate statistical analyses found most windward fore-reef and back-reef sites to be statistically similar, but considerable variation to exist among sites within calmer lagoonal areas. Surveys revealed macroalgae to dominate over scleractinian coral species at the majority of sites in this healthy, subtropical reef system, although select lagoonal areas were dominated by dense coral communities.     

Production and supply of larvae as determinants of zonation in a brooding tropical coral

Production and settlement of planktonic larvae of the coral Favia fragum (Esper) were studied. The species is restricted to shallow back- and fore-reef habitats throughout the Caribbean Sea. Adults are in their greatest abundance on the reef-crest and shallow reef slope (<3 m) at Tague Bay, St. Croix, in the US Virgin Islands. Because F. fragum broods larvae that are capable of immediate settlement, this distribution pattern may be due to variation in fecundity among depths. Corals were collected from shallow (1.0 m) and deep depths (10-13 m) and cultured in individual containers exposed to shaded ambient light. Corals from shallow depths had greater fecundity (polyp⁻¹ lunar cycle⁻¹) and were larger than deep corals. To test the hypothesis that fecundity was related to successful fertilization, corals were kept in different densities in an area with sea-grass, where there were no natural adults. Production of larvae 6 months later was not affected by density of adults, possibly due to self-fertilization. Larval choice of habitat was also examined. In the laboratory, twice as many larvae settled on coral rubble fragments collected from depths where adults were common (1.5 and 3 m) than on those from depths where adults were rare (10 m). Larval supply may establish the vertical distribution of adults on St. Croix.     

A coral damage index and its application to diving sites in the Egyptian Red Sea

 A coral damage index (CDI) is provided, to screen sites to obtain a perspective on the extent and severity of physical damage to coral. Sites are listed as “hot spots” if in any transect the percent of broken coral colonies (BCC) is greater than or equal to 4% or if the percent cover of coral rubble (CR) is greater than or equal to 3%. To demonstrate its utility, the CDI is applied to a real-life management situation off Hurghada and Safaga, Egypt in the Red Sea. The extent of coral damage covered all four diving sites. Forty percent of all the transects were “hot spots” that required management action. Thirty-one percent of the 16 “hot spot” transects were identified by both broken coral and rubble criteria, 25% by only broken coral criterion and 44% by only coral rubble criterion of the CDI, suggesting that past breakage was responsible for most of the observed damage. Sixty-three percent of the “hot spot” transects were at 4 m depth versus 37% at 8 m depth, suggesting that most of the damage was caused by anchors dragging across the reef in shallow water. The severity of coral damage, reflected by CR, was the greatest at Small Giftun in transect 5 at 4 m depth (333% above the CDI). EI Fanous experienced the most severe degree of broken coral damage (325% above the CDI) at 8 m depth along transect 2. Estimates of the number of dives per year show diving carrying capacities for El Fanous, Gotta Abu Ramada, Ras Abu Soma and Small Giftun being exceeded by large amounts. The CDI can be used globally to; gauge the severity and extent of damage, focus managers on areas that need mooring buoys and associated dive site management programs, and provide a starting point from which to focus more detailed coral reef assessments and restoration programs. Accepted: 30 June 1999     

Morphologie, Ökologie und Zonierung von Korallenriffen bei Aqaba, (Golf von Aqaba,Rotes Meer)

The coral reefs of the Gulf of Aqaba are among the most northern ones of the world. This study, the first concerning the east coast of this topographically and hydrographically peculiar sea, considers relationships of biophysiographical and structural reef zones to fundamental abiotic environmental factors. An introduction to paleogeography, geology, petrography, topography, climate and hydrography is followed by terminological definitions used to describe the different reef areas. The investigations were carried out on two transects crossing fringing reefs of different shape. Each transect was 20 m wide and run from the shore over nearly 200 m to the fore reef in about 30 m depth. One reef, a “coastal-fringing reef”, represents an unaltered straight reef flat from shore to the reef edge 60 m away; two large pinnacles reach the surface some 125 m off the shore. The other reef, a “lagoon-fringing reef”, is divided into a 100 m wide lagoon of 0.5–2.3 m depth and a reef crest separated from the former by a rear reef. The reef platform of the lagoon-fringing reef is cut by a system of channels and tunnels; the reef edge is about 135 m off shore. Such water depth, substrate, temperature, illumination and water movement were recorded, about 200 common or dominant species (plants and animals) were collected, their distribution plotted and, together with other data and structural items, charted. Indicator species characterize the biophysiographical zones. Their variation as well as that of the structural and substrate zones depend on different zones of water movement. This basic factor also controls other ecological parameters such as food and oxygen supply as well as temperature and salinity gradients between fore reef and shore. From this point of view the ecological requirements of some indicator and other species and conversely the ecological settings of different reef areas are discussed. The different shapes of both reefs are explained on the basis of a “reef development cycle” — a hypothesis applicable to fringing reefs at unchanging sea level and based on the fact that only a small surf-influenced area of “living reef” is able to compensate for reef destruction: While a young coastal fringing reef is growing outwards, its back reef is gradually altered to a reef lagoon by erosion. After stillstand of seaward expansion the reef crest, too, is cut by a channel system eroded by rip currents. This stage is represented by the lagoon-fringing reef. Isolated pinnacles remain as remnants of the former reef crest; young coastal-fringing reefs develop from the shore. This stage is examplified by the first reef studied. Extension, growth intensity, dominant frame building corals, and the number of species of the Aqaba reefs are compared with those of Eilat and with reefs of the middle Red Sea, South India, Southwest-Pacific and Jamaica.     

The area of potential shallow-water tropical and subtropical coral ecosystems in the United States

Geographic information system-based analysis was used to derive comprehensive, consistent estimates of the potential area of broadly defined, shallow-water, tropical and subtropical coral ecosystems within the territorial sea and exclusive economic zone of the United States. A coral ecosystem is composed of habitats including unconsolidated sediment, mangrove, hermatypic coral, colonized hardbottom, and submerged vegetation, and major structural zones like reef crest, lagoon, and fore reef. This broad definition reflects the importance of both reef and non-reef habitats and structural zones in the function of these ecosystems. Nautical charts, published by the National Oceanic and Atmospheric Administration’s Office of the Coast Survey, provide a consistent source of 10-fathom (∼18 m) and 100-fathom (∼183 m) depth curve information. The 10-fathom or 100-fathom depth curves are used as surrogates for the potential distribution and extent of shallow-water coral ecosystems in tropical and subtropical U.S. waters. An estimated 36,813 sq·km area has been identified where coral ecosystems can potentially be found in waters less than 10 fathoms (18 m) deep. In addition, an estimated 143,059 sq·km area has been identified where coral ecosystems potentially can be found in U.S. waters at depths down to 100 fathoms (183 m). Results also indicate that previous studies underestimated the extent of potential coral ecosystems for some locations in U.S. tropical and subtropical waters by as much as 100% and that the regional distribution of coral ecosystems has been incorrectly reported.     

Foraminiferal distribution and diversity,Madang Reef and Lagoon,Papua New Guinea

In the Madang Lagoon, on the northern coast of Papua New Guinea (PNG), distinct groups of foraminifera, defined by numerical Q-mode cluster analysis of foraminiferal species occurrences, occupy four major environments and sedimentary regimes, generally aligned parallel to the coast: (1) the harbor and bay inlets, which have large fresh-water runoff and organic detrital inputs; (2) the fringing reefs along the west side of the lagoon which are influenced by coastal factors such as overhanging mangroves or fresh-water runoff; (3) the central lagoon floor which is over 50 m deep and covered with fine sand and patch reefs rising from it; and (4) the reef barrier with adjacent live coral-covered fore-reef slope and generally sandy back-reef slope. The four clusters are also mirrored in both species richness and Fisher alpha diversity analysis. Cluster 4 includes 79 species of large, thick-shelled miliolids, robust agglutinated species, calcarinids, and amphisteginids (Fisher α ≥20) that occur on the coral-rich barrier reef and back-reef. Cluster 3 has 50 species (Fisher α=8–20) and occupies the central lagoon floor. Cluster 2 has 25 or fewer species (Fisher α=2–6) and occurs on the shallow fringing reefs. Cluster 1 is the least diverse (≤7 species, Fisher α ≤2) and occurs in the harbors and bays in the mouths of larger rivers and streams. The larger, endosymbiont-bearing foraminifera (alveolinellids, soritids, amphisteginids, nummulitids, and calcarinids) generally live on the back- and fore-reef slopes and in the lagoon, avoid the organic-rich coastal and harbor habitats, and preferentially dwell in well-lit environments to the bottom of the lagoon. The river mouths and bays are unusual for reef systems because of their high organic content, which creates low-oxygen and nutrient-rich conditions. Here the foraminiferal fauna is dominated by only a few and, for the most part, particularly thin-shelled and highly fragile species. Each faunal group contains a number of numerically abundant indicator species that do not occur in other faunal clusters. This implies low horizontal transport rates within the reef and lagoon complex and signifies that faunal mixing among the cluster groups is limited. Foraminiferal death assemblages may thus be autochthonous and retain information regarding the original community structure. They may also preserve environmental information useful in paleoecological studies and they are good ecological indicators of reef and lagoon habitats.     

Biodiversity of <Emphasis Type="Italic">Spongosorites coralliophaga</Emphasis> (Stephens, 1915) on coral rubble at two contrasting cold-water coral reef settings

Cold-water coral reefs (CWRs) in the northeast Atlantic harbor diverse sponge communities. Knowledge of deep-sea sponge ecology is limited and this leaves us with a fragmented understanding of the ecological roles that sponges play in CWR ecosystems. We present the first study of faunal biodiversity associated with the massive demosponge Spongosorites coralliophaga (Stephens, 1915) that typically colonizes coral debris fields of CWRs. Our study focused on the sessile fauna inhabiting sponges mixed with coral rubble at two contrasting settings in the northeast Atlantic: the shallow inshore (120–190 m water depth) Mingulay Reef Complex (MRC) and the deep offshore (500–1200 m) Logachev Mound (LM) coral province. MRC is dominated by the scleractinian Lophelia pertusa, while LM is dominated by L. pertusa and Madrepora oculata. Nine sponge–coral rubble associations were collected from MRC and four from LM. Measurements of abundance, species richness, diversity, evenness, dry biomass, and composition of sessile fauna on sponge and coral rubble microhabitats were undertaken. Differences in community composition between the two regions were mainly a response to changes in fauna with depth. Fauna composition was also different between sponge and coral rubble within each region. Infauna constituted a minor component of the sponge-associated fauna in MRC but had a higher contribution in LM. Sponge and coral rubble sessile fauna in both regions was mainly composed of cnidarians and molluscs, similarly to some previous studies. Sponges’ outer surfaces at MRC were colonized by a species-rich community with high abundance and biomass suggesting that S. coralliophaga at MRC acts as a settlement surface for various organisms but such a role is not the case at LM. This difference in the role of S. coralliophaga as a biological structure is probably related to differences in fauna composition with depth, bottom current speed, and the quantity/quality of food supplied to the benthos.     

Sponge distribution across Davies Reef,Great Barrier Reef,relative to location,depth, and water movement

Sponge populations were surveyed at different depths in three zones of Davies Reef, a large platform reef of the central Great Barrier Reef. Depth is the major discriminatory factor as few sponges are found within the first 10 m depth and maximal populations occur between 15 m and 30 m on fore-reef, lagoon and back-reef slopes. Reef location is another major factor, with the lagoon containing a significantly different sponge population to either the fore-reef or the back-reef slopes. Physical factors are considered to be the major influences behind these patterns. Physical turbulence is strongest within the first 10 m and apparently limits sponge growth within these shallow zones. Insufficient photosynthetic radiation limits the growth of the sponge population below 30 m depth as many of the species are phototrophic with a dependence on cyanobacterial symbionts for nutrition. Sponge populations on the outer (fore- and back-) reef slopes are comparable with each other but different from those on lagoon slopes where currents are reduced and fine sediment loads are higher. The largest populations occur on the back-reef slope where currents are stronger and there are possibly higher concentrations of organic nutrients originating from the more productive shallow parts of the reef. While there are correlations between sponge populations and environmental parameters, data are insufficient to enable more definitive conclusions to be drawn. Most sponge species are distributed widely over the reef, however, some are restricted to a few habitats and, hence, may be used to characterize those habitats.     

Distribution of a nematocyst-bearing sponge in relation to potential coral donors

Haliclona sp. 628 (Demospongiae, Haplosclerida, Chalinidae), a sponge found on the reef slope below 5 m depth on the Great Barrier Reef, has two unusual characteristics. It contains a symbiotic dinoflagellate, Symbiodinium sp., similar in structure to the dinoflagellate found within Acropora nobilis (S. microadriaticum), and it contains coral nematocysts randomly distributed between the ectosome and endosome and usually undischarged in intact sponge tissue. Given the unusual occurrence of nematocysts in Haliclona sp. 628, the focus of this study was to determine the distribution of this species of sponge on the reef slope at Heron Island Reef in relation to the distribution of potential coral donors. A combination of line and belt transects was used to estimate the abundance of Haliclona sp. 628 and a co-occurring congener, Haliclona sp. 1031, which does not contain nematocysts, at three widely separated sites on the reef slope at Heron Island Reef. The abundance of different types of substratum (sand, sand-covered coral rubble, dead A. nobilis, live A. nobilis, other live coral, and other dead coral) along the transects and the substratum to which each sponge colony was attached were also recorded. Despite the predominance of live A. nobilis and sand-covered rubble at all sites, between 30 and 55% of Haliclona sp. 628 colonies were attached to dead A. nobilis which comprised less than 8% of the available substratum along any transect. In contrast, Haliclona sp. 1031 was found significantly more frequently on other dead corals and less frequently on live A. nobilis than would be expected based on the availability of the different substrata in the sites. Potential explanations to account for the distribution of Haliclona sp. 628 in relation to potential coral donors are discussed.     

Abundance and reproductive patterns of the excavating sponge Cliona vermifera: a threat to Pacific coral reefs?

Cliona vermifera is a common excavating sponge in coral reefs from the East Pacific. Abundance and reproductive patterns of the sponge in a Mexican Pacific coral reef over a 4-year period are herein described. Sponge abundance was estimated along three transects 50 m long which were randomly placed on the reef, and along each one, a piece of coral rubble and a branch of a live coral from the Pocillopora spp. coral colony closest to the transect were collected at random, approximately every 2 m, yielding 25 pieces of each category per transect (and 75 pieces total of each category). A 2-way ANOVA revealed that invasion was significantly higher in living coral colonies (34.8 %) than in rubble (13.7 %). It also indicated that the abundance in both coralline substrates showed a temporal variation without a clear pattern of increase over the years. It was estimated that 60–85 % of sponges in the population reproduced sexually every year. The sponge proved gonochoristic, with a sex ratio strongly departing from parity (1 male: 3 females). Over the 4-year study period, at least two cohorts of oocytes with densities of up to 3.5 oocytes per mm² tissue were observed. Spermatogenesis lasted about a month, but often producing more than a pulse from July to November, coupled with peaks of oocyte maturation. Fertilization occurred internally to produce encapsulated zygotes that were released in one or more spawning events from July to November. In the following months (December to February), which were the periods of lowest temperature (~18.5–20 °C), no gametic activity occurred in the sponges. Because anomalous temperature rises that are detrimental to corals do not appear to negatively affect the reproduction and abundance of C. vermifera, it is likely that the excavating activity of this sponge may be compromising the health of those coral reefs that are recurrently affected by episodes of thermal stress.     

Catastrophic impact of hurricanes on atoll outer reef slopes in the Tuamotu (French Polynesia)

Underwater effects on coral reefs of the six hurricanes which ravaged French Polynesia between December 82 and April 83 were observed by SCUBA diving around high islands and atolls during September and October 1983. Special attention was paid to Tikehau atoll reef formations (Tuamotu archipelago) where quantitative studies on scleractinians, cryptofauna and fishes were conducted in 1982 immediatly prior to the hurricanes. On outer reef slopes coral destruction, varying from 50 to 100%, was a function of depth. Upper slope coral communities composed of small colonies well adapted to high energy level environments, suffered less than deeper formations. However, there is a narrow erosional trough in this zone at a depth of 6 m that was probably the result of storm-wave action (plunge point). Coral destruction was spectacular at depths greater than 12 m: 60 to 80% between 12 m and 30 m and 100% beyond 35 m, whereas earlier living coral coverage ranged from 60 to 75% in these zones. The outer slope was transformed into a scree zone covered with coarse sand and dead coral rubble. Dives on different sites around steep outer slopes (>45°) of the atolls and more gentle slopes (<25°) of some parts of the high islands permitted the formulation of an explanatory hypothesis: direct coral destruction by hurricane-induced waves occurred between the surface and 18–20 m; on low-angle slopes broken colonies were thrown up on reef flats and beaches; on steep slopes avalanches destroyed much of the living corals and left scree slopes of rubble and sand.     

Holocene growth history of an eastern Pacific fringing reef,Punta Islotes,Costa Rica

Rock and sediment cores reveal that a well-developed fringing reef in Golfo Dulce, Pacific Costa Rica, up to 9 m thick was established on Cretaceous basalt about 5500 y BP. It is presently being smothered with fine sediments and is almost completely dead. This reef is made up of three main facies that are represented by comparable extant reef zones: reef-flat branching coral, fore-reef slope massive coral, and fore-reef talus sediment facies. Reef growth began with the establishment of small patch reefs dominantly formed by the branching coral Pocillopora damicornis. P. damicornis spread across the basalt bench and massive colonies of Porites lobata grew on the outer slopes, eventually blocking the seaward transport of Pocillopora fragments to the fore-reef talus sediments. The reef flourished until 500 years ago. Lower accumulation rates during the past 500 years may be due to deteriorating environmental conditions rather than slower growth after the reef reached sea level. Present-day reef communities are severely degraded with less than 2% living coral cover. The increased turbidity associated with the final stage of degradation of this reef is probably related to human activity on the adjacent shores, including deforestation, mining, and road construction.