Molluscan assemblages on coral reefs and associated hard substrata in the northern Red Sea

Information on spatial variability and distribution patterns of organisms in coral reef environments is necessary to evaluate the increasing anthropogenic disturbance of marine environments (Richmond 1993; Wilkinson 1993; Dayton 1994). Therefore different types of subtidal, reef-associated hard substrata (reef flats, reef slopes, coral carpets, coral patches, rock grounds), each with different coral associations, were investigated to determine the distribution pattern of molluscs and their life habits (feeding strategies and substrate relations). The molluscs were strongly dominated by taxa with distinct relations to corals, and five assemblages were differentiated. The Dendropoma maxima assemblage on reef flats is a discrete entity, strongly dominated by this encrusting and suspension-feeding gastropod. All other assemblages are arranged along a substrate gradient of changing coral associations and potential molluscan habitats. The Coralliophila neritoidea-Barbatia foliata assemblage depends on the presence of Porites and shows a dominance of gastropods feeding on corals and of bivalves associated with living corals. The Chamoidea-Cerithium spp. assemblage on rock grounds is strongly dominated by encrusting bivalves. The Drupella cornus-Pteriidae assemblage occurs on Millepora-Acropora reef slopes and is strongly dominated by bivalves associated with living corals. The Barbatia setigera-Ctenoides annulata assemblage includes a broad variety of taxa, molluscan life habits and bottom types, but occurs mainly on faviid carpets and is transitional among the other three assemblages. A predicted degradation of coral coverage to rock bottoms due to increasing eutrophication and physical damage in the study area (Riegl and Piller 2000) will result in a loss of coral-associated molluscs in favor of bivalve crevice dwellers in dead coral heads and of encrusters on dead hard substrata.     

Book review of Littler DM. Littler MM (2000) Caribbean Reef Plants An Identification Guide to the Reef Plants of the Caribbean,Bahamas, Florida and Gulf of Mexico

Information on spatial variability and distribution patterns of organisms in coral reef environments is necessary to evaluate the increasing anthropogenic disturbance of marine environments (Richmond 1993; Wilkinson 1993; Dayton 1994). Therefore different types of subtidal, reef-associated hard substrata (reef flats, reef slopes, coral carpets, coral patches, rock grounds), each with different coral associations, were investigated to determine the distribution pattern of molluscs and their life habits (feeding strategies and substrate relations). The molluscs were strongly dominated by taxa with distinct relations to corals, and five assemblages were differentiated. The Dendropoma maxima assemblage on reef flats is a discrete entity, strongly dominated by this encrusting and suspension-feeding gastropod. All other assemblages are arranged along a substrate gradient of changing coral associations and potential molluscan habitats. The Coralliophila neritoidea–Barbatia foliata assemblage depends on the presence of Porites and shows a dominance of gastropods feeding on corals and of bivalves associated with living corals. The Chamoidea–Cerithium spp. assemblage on rock grounds is strongly dominated by encrusting bivalves. The Drupella cornus–Pteriidae assemblage occurs on Millepora–Acropora reef slopes and is strongly dominated by bivalves associated with living corals. The Barbatia setigera–Ctenoides annulata assemblage includes a broad variety of taxa, molluscan life habits and bottom types, but occurs mainly on faviid carpets and is transitional among the other three assemblages. A predicted degradation of coral coverage to rock bottoms due to increasing eutrophication and physical damage in the study area (Riegl and Piller 2000) will result in a loss of coral-associated molluscs in favor of bivalve crevice dwellers in dead coral heads and of encrusters on dead hard substrata.     

Influence of setting,sieve size,and sediment depth on multivariate and univariate assemblage attributes of coral reef-associated mollusc death assemblages from the Gulf of Aqaba

Measures of diversity and ecology of marine invertebrate assemblages depend on a variety of factors including environmental conditions and methodological decisions. In this study, the influence of such factors on multi- and univariate assemblage parameters of molluscan death assemblages from the Gulf of Aqaba (Red Sea, Jordan) was evaluated. Sediment samples were collected at two coral reef types, a patch reef at 13 m of water depth characterized by fine-grained sediments and a Millepora-fringing reef with coarse-grained sediments at 5 m of water depth. The upper and lower 10 cm of the sediment column were separately removed and sieved with mesh sizes of 1 and 2 mm. A large dataset of 6400 bivalve and gastropod shells was compiled to evaluate how setting, sediment depth, and sieve size influenced taxonomic composition and species richness, species-abundance patterns and the Shannon–Wiener index, the number of drilled shells per species and drilling frequency (DF) of the assemblage. Setting had the strongest impact on all aspects, followed by sieve size, but sediment depth was insignificant, probably due to complete homogenization of the sediments by reworking and bioturbation. Multivariate assemblage parameters distinguished much better between categories (setting, sieve size) than univariate measures. Sieve size-related disagreements recognized between the two higher taxa are mostly due to the underlying difference in body-size distribution of bivalve and gastropod assemblages. We conclude that species richness and other ecological characteristics of molluscan death assemblages in coral reef-associated sediments will most strongly reflect habitat complexity of the sites chosen, are significantly influenced by methodological decisions (i.e., sieve size), will only poorly preserve temporal patterns, and the results will differ between bivalves and gastropods.     

Initial phases in reef development,studied at artificial reef types off Eilat, (Red Sea)

The initial steps in the formation of a coral reef (as distincr from the enlarging of an already existing reef) have been observed at block moles off Eilat (Red Sea). Within a period of 11 years, thus far four phases can be distinguished: (1) start-phase: rapid and homogeneous colonization by fouling organisms, still indeterminant for the final reef development; (2) preparation-phase: settling of shells, calcareous red algae and foraminiferans not affected by grazing animals which largely consume the initial settlers and subsequently attaching larvae; (3) phase of pioneer frame-building: growth of scleractinians and hydrocorals, settled on the remains of rock-attached shells or on other places inaccessible to grazers; secondary frame-builders (e.g.Tridacna) grow in their shelter; (4) phase of frame-binding: dead coral colonies are overgrown by calcareous foraminiferans, algae and bryozoans consolidating the coralline structures by their deposits. Living corals are successively colonized by a specific community (mainly decapods and molluscs); among these, borers are the earliest settlers. Upon death of the coral, there is a complete change in associated organisms forming a less specific but more diverse community.Supported by the Deutsche Forschungsgemeinschaft.     

Heavy metals distribution in the coral reef ecosystems of the Northern Red Sea

Concentrations of seven heavy metals (Cu, Zn, Pb, Cd, Ni, Co and Fe) were measured in the seawater, sediments, common scleractinian reef-building corals and soft corals (Octocorallia : Alcyonacea) at seven reef sites in the Northern Red Sea: I (Hurghada), II (Ras Za’farana), III (El-Ain Al-Sukhna), IV (El-Tur), V (Sha’b Rashdan), VI (Sharm El-Sheikh) and VII (Dahab). Levels of heavy metals were considerably elevated in seawater, sediments and corals collected from reef sites exposed to increased environmental contamination, as a result of diversified natural and anthropogenic inputs. Soft corals of genera Lithophyton, Sarcophyton and Sinularia showed higher concentrations of Zn, Pb, Cd and Ni than hard coral genera Acropora and Stylophora. Soft coral Sarcophyton trocheliophorum collected from El Ain Al-Suhkna (Gulf of Suez) had greater concentration of Cu, followed by hard corals Acropora pharaonis and Acropora hemprichi. The elevated levels of Zn, Cd and Ni were reported in the dry tissue of soft coral Sinularia spp. On the other hand, the soft coral Lithophyton arboreum displayed the highest concentration of Pb at Sha’b Rashdan (Gulf of Suez) and elevated concentration of Zn at Sharm El-Sheikh. Sediments showed significantly higher concentration of Fe than corals. The higher levels of Fe in hard corals than soft corals reflected the incorporation of Fe into the aragonite and the chelation with the organic matrix of the skeleton. The greater abundance of soft corals in metal-contaminated reef sites and the elevated levels of metals in their tissue suggesting that the soft corals could develop a tolerance mechanism to relatively high concentrations of metals. Although the effects of heavy metals on reef corals were not isolated from the possible effects of other stresses, the percentage cover of dead corals were significantly higher as the concentrations of heavy metals increased.     

Ecological relations between hermit crabs and their shell-supplying gastropods: Constrained consumers

Hermit crabs are critically dependent upon gastropod shells for their survival and reproductive fitness. While anecdotal reports have suggested that hermit crabs may be capable of removing live gastropods from their shells to access the essential shell resource, no systematic experiments have been conducted to investigate this possibility. This paper reports experiments on both marine (Pagurus bernhardus) and terrestrial (Coenobita compressus) hermit crabs in which crabs were paired in the laboratory with the gastropods whose shells they inhabit in the field. Pairings included both shelled and naked crabs and spanned the full range of the gastropod life cycle. Neither marine nor terrestrial hermit crabs were successful at removing live gastropods from their shells. Furthermore, only a small fraction of the crabs (5.7%) were capable of accessing shells in which the gastropod had been killed in advance, with its body left intact inside the shell. Finally, although hermit crabs readily entered empty shells positioned on the surface, few crabs (14.3%) were able to access empty shells that were buried just centimeters beneath them. These results suggest that hermit crabs are constrained consumers, with the shells they seek only being accessible during a narrow time window, which begins following natural gastropod death and bodily decomposition and which typically ends when the gastropod's remnant shell has been buried by tidal forces. Further experiments are needed on more species of hermit crabs as well as fine-grained measurements of (i) the mechanical force required to pull a gastropod body from its shell and (ii) the maximum corresponding force that can be generated by different hermit crab species' chelipeds.     

Thermal refugia against coral bleaching throughout the northern Red Sea

Tropical reefs have been impacted by thermal anomalies caused by global warming that induced coral bleaching and mortality events globally. However, there have only been very few recordings of bleaching within the Red Sea despite covering a latitudinal range of 15° and consequently it has been considered a region that is less sensitive to thermal anomalies. We therefore examined historical patterns of sea surface temperature (SST) and associated anomalies (1982–2012) and compared warming trends with a unique compilation of corresponding coral bleaching records from throughout the region. These data indicated that the northern Red Sea has not experienced mass bleaching despite intensive Degree Heating Weeks (DHW) of >15°C‐weeks. Severe bleaching was restricted to the central and southern Red Sea where DHWs have been more frequent, but far less intense (DHWs <4°C‐weeks). A similar pattern was observed during the 2015–2016 El Niño event during which time corals in the northern Red Sea did not bleach despite high thermal stress (i.e. DHWs >8°C‐weeks), and bleaching was restricted to the central and southern Red Sea despite the lower thermal stress (DHWs < 8°C‐weeks). Heat stress assays carried out in the northern (Hurghada) and central (Thuwal) Red Sea on four key reef‐building species confirmed different regional thermal susceptibility, and that central Red Sea corals are more sensitive to thermal anomalies as compared to those from the north. Together, our data demonstrate that corals in the northern Red Sea have a much higher heat tolerance than their prevailing temperature regime would suggest. In contrast, corals from the central Red Sea are close to their thermal limits, which closely match the maximum annual water temperatures. The northern Red Sea harbours reef‐building corals that live well below their bleaching thresholds and thus we propose that the region represents a thermal refuge of global importance.     

Marine heatwaves reveal coral reef zones susceptible to bleaching in the Red Sea

As the Earth's temperature continues to rise, coral bleaching events become more frequent. Some of the most affected reef ecosystems are located in poorly monitored waters, and thus, the extent of the damage is unknown. We propose the use of marine heatwaves (MHWs) as a new approach for detecting coral reef zones susceptible to bleaching, using the Red Sea as a model system. Red Sea corals are exceptionally heat‐resistant, yet bleaching events have increased in frequency. By applying a strict definition of MHWs on >30 year satellite‐derived sea surface temperature observations (1985–2015), we provide an atlas of MHW hotspots over the Red Sea coral reef zones, which includes all MHWs that caused major coral bleaching. We found that: (a) if tuned to a specific set of conditions, MHWs identify all areas where coral bleaching has previously been reported; (b) those conditions extended farther and occurred more often than bleaching was reported; and (c) an emergent pattern of extreme warming events is evident in the northern Red Sea (since 1998), a region until now thought to be a thermal refuge for corals. We argue that bleaching in the Red Sea may be vastly underrepresented. Additionally, although northern Red Sea corals exhibit remarkably high thermal resistance, the rapidly rising incidence of MHWs of high intensity indicates this region may not remain a thermal refuge much longer. As our regionally tuned MHW algorithm was capable of isolating all extreme warming events that have led to documented coral bleaching in the Red Sea, we propose that this approach could be used to reveal bleaching‐prone regions in other data‐limited tropical regions. It may thus prove a highly valuable tool for policymakers to optimize the sustainable management of coastal economic zones.     

Cyclone effects on coral reef habitats in New Caledonia (South Pacific)

The impacts of the unusually strong Cyclone Erica (March 2003) on coral reef habitats at a site located on the northwest coast of New Caledonia (South Pacific) were assessed using a 6-year data set (2002–2007). We examined the interannual variations of key variables describing reef habitats (live hard and soft corals, dead corals in place, coral debris, algae and relative proportion of mechanically vulnerable and resistant live hard corals). The cyclone-induced disturbances of habitats differed according to three reef types: patch reefs, barrier reefs far from passes (more than 3 km from the nearest pass) and barrier reefs near passes (less than 3 km from the nearest pass). Short-term mechanical damage was detected on the three-dimensional structure of reef habitats with a notable shift from a community dominated by mechanically vulnerable corals to one dominated by resistant corals on barrier reefs far from passes. The history of habitats and their pre-disturbance characteristics, in link with local hydrodynamics, was found to influence their short-term susceptibility to extreme events such as cyclones. However, the most significant effects appeared in the midterm (within 2 years after the cyclone) as the cover of live hard corals significantly decreased by approximately 45% between 2002 and 2004 on all reef types. The short- and midterm disturbances of coral reef habitats are discussed with regard to published temporal variations in reef fish assemblages, underlining the delayed effects of this cyclonic event on fish as well as benthic habitats. Coral reef habitats and live corals had shown significant patterns of recovery 4 years after the cyclone, followed by similar recovery in fish community, suggesting good resilience in a face of this major natural disturbance in an area under moderate anthropogenic pressure.     

Patterns in the use of space by benthic communities on two coral reefs of the Great Barrier Reef

A rapid benthic line-transect survey method for use by non-specialist observers is described. At both Davies Reef (mid-continental shelf) and Myrmidon Reef (outer-continental shelf) in the central Great Barrier Reef a set of 6 sites of varying depths on the reef flat, crest and slope were sampled using this method. At least 10 contiguous 10 m transects were made at each site. Benthic organisms were recorded as life forms with categories based on both high level taxa and morphologies, and including scleractinian corals, alcyonarians, sponges, algae and others. Percentage cover data for 19 benthic categories are presented for all sites. Coral cover on both reefs is high on the crest and slope but low on the reef flat. At all sites the cover of soft corals and sponges is much less than cover of hard corals and algae. Abundances of soft corals and sponges increase with depth. Analysis of gaps between hard corals show that many colonies grow close to each other (<1 cm)even when total coral cover is low.     

Symbiosis of sea anemones and hermit crabs: different resource utilization patterns in the Aegean Sea

The small-scale distribution and resource utilization patterns of hermit crabs living in symbiosis with sea anemones were investigated in the Aegean Sea. Four hermit crab species, occupying shells of nine gastropod species, were found in symbiosis with the sea anemone Calliactis parasitica. Shell resource utilization patterns varied among hermit crabs, with Dardanus species utilizing a wide variety of shells. The size structure of hermit crab populations also affected shell resource utilization, with small-sized individuals inhabiting a larger variety of shells. Sea anemone utilization patterns varied both among hermit crab species and among residence shells, with larger crabs and shells hosting an increased abundance and biomass of C. parasitica. The examined biometric relationships suggested that small-sized crabs carry, proportionally to their weight, heavier shells and increased anemone biomass than larger ones. Exceptions to the above patterns are related either to local resource availability or to other environmental factors.     

Marine molluscs from the Iztuzu beach near Dalyan (Mediterranean coast of Turkey)

Abstract

A total of l84 species of marine molluscs has been identified from shells collected on the beach of Iztuzu near Dalyan (Mu?la province, Turkey). Of these, 16 species of gastropod and 4 species of bivalvia are new For the extreme Eastern Mediterranean. Only three species of Lessepsian migrants (from the Red Sea via the Suez canal) have been found.     

Quality of shells occupied by the hermit crabPagurus geminus: How many hermit crabs are satisfied with their shells?

The rate of changing shells in the hermit crabPagurus geminus was investigated to determine how many hermit crabs are satisfied with their shells. Animal collected from the coast of Oya, Tanabe, Wakayama Prefecture, were presented with fresh gastropod shells newly made by removing the soft parts. Approximately 80% of the hermit crabs changed from their original shells acquired in the natural habitat into fresh shells experimentally given to them and, thus, were regarded as dissatisfied with the shells possessed in the natural habitat. This condition was thought to lead to the fact that hermit crabs occasionally attempted to exchange shells between 2 individuals and even to attack living snails in the natural habitat.     

Patterns in the distribution of soft corals across the central Great Barrier Reef

Distribution patterns of soft coral genera were examined at 11 reefs situated in a broad transect from inshore to the Coral Sea in the central region of the Great Barrier Reef. Twenty-five genera representing the Orders Alcyonacea and Stolonifera were recorded, and the survey also included one genus of the Order Gorgonacea. Total living soft coral cover is greatest on outershelf reef slopes, and is often less than and inversely related to the cover by stony corals. Soft coral diversity is generally low on reef flats, where soft coral cover is low or nil except in protected, inshore areas. The most diverse assemblages occur on reef slopes in midshelf and outershelf areas, where Efflatounaria and nephtheid genera predominate, and widely distributed alcyoniid genera are common. These richer assemblages are less well represented in the Coral Sea, while innershelf reefs support a less diverse fauna of somewhat different generic composition. Distribution patterns of soft corals across the transect broadly match similar variations in the distributions of stony corals and fishes, inshore reefs being generally depauperate. Such variations across the continental shelf are closely associated with changes in prevailing environmental conditions, but further research will be required to elucidate the effects of environmental parameters on benthic community structure.     

Substrate specificity in the Devonian tabulate coral Pleurodictyum

The tabulate coral Pleurodictyum americanum Roemer has been cited as an example of a host-specific organism occurring exclusively on the shells of gastropods, particularly Palaeozygopieura hamiltoniae (Hall). Examination of over 1600 specimens of P. americanum, from the Middle Devonian Hamilton Group of western New York, reveals additional complexities which require reinterpretation. While substrate selectivity for Palaeozygopieura shells is evident in all 42 subsamples, a variety of other substrates were also utilized by Pleurodictyum including corals, brachiopods, other molluscs and pebbles. Recent scleractinian corals inhabiting soft bottoms show similar substrate preference, selecting for the tubes of live serpulids, or gastropod shells (invariably with a secondary sipunculid host), but also occasionally settling on unoccupied shells or pebbles. Shell surfaces of P. hamiltoniae, preserved as external molds on the Pleurodictyum epitheca, exhibit encrustation by worm tubes and bryozoans as well as borings and mechanical shell damage, suggesting that these were not the shells of live gastropods. However, the invariant aperture-downward orientation and the high degree of selectivity of P. americanum strongly suggest that the shells were occupied by secondary hosts. □ Substrate specificity, commen-salism, tabulate coral, gastropod, sipunculid, Devonian, Hamilton Group, New York.     

Interference and exploitation components in interespecific competition between sympatric intertidal hermit crabs

This study was designed to evaluate the effect of interference and exploitation competition in shell partitioning between two hermit crab species (Pagurus criniticornis and Clibanarius antillensis). Field samples revealed that shells of the gastropod Cerithium atratum were the main resource used by both hermit crab species and that Pagurus used eroded or damaged shells in higher frequency than Clibanarius. The exploitative ability of each species was compared between species in the laboratory using dead gastropod (Cerithium) baits to simulate predation events and signalize newly available shells to hermit crabs. Pagurus reached the baits more rapidly than Clibanarius, but this higher exploitative ability did not explain shell utilization patterns in nature. Another experiment evaluated the dominance hierarchy between these two hermit crab species and revealed that Clibanarius was able to outcompete Pagurus for higher quality shells in agonistic encounters. This higher interference competitive ability of Clibanarius in relation to Pagurus may explain field observations. Nevertheless, Pagurus may be responsible to enhance shell availability to other hermit crab species that have lower ability to find and use newly available shells. Differently, the poorer condition of shells used by Pagurus, the higher ability of this species to attend gastropod predation events and its higher consumption rate by shell-breaking crabs (Menippe nodifrons) may increase its predation risks, thus revealing the disadvantages of such an exploitative competitive strategy for hermit crabs.     

Changes in coral assemblages during an outbreak of Acanthaster planci at Lizard Island, northern Great Barrier Reef (1995–1999)

Population outbreaks of crown-of-thorns starfish (Acanthaster planci L.) represent one of the most significant biological disturbances on tropical coral reefs and have the potential to devastate coral communities, thereby altering the biological and physical structure of reef habitats. This study reports on changes in area cover, species diversity and taxonomic composition of corals during an outbreak of A. planci at Lizard Island, in the northern Great Barrier Reef, Australia. Mean coral cover declined by 28.8% across ten locations studied. However, densities of A. planci, and their effects on local coral assemblages, were very patchy. Declines in coral cover were mostly due to the selective removal of certain coral taxa (mainly Acropora and Pocilloporidae corals); such that the greatest coral loss occurred at locations with highest initial cover of preferred coral prey. Most notably, coral assemblages in back-reef locations were transformed from topographically complex staghorn Acropora-dominated habitats, to relatively depauperate assemblages dominated by alcyonacean soft corals. Although coral loss was greatest among formerly dominant taxa (especially Acropora), effects were sufficiently widespread across different coral taxa, such that overall coral diversity tended to decline. Clearly, moderate outbreaks of A. planci have the potential to greatly alter community structure of coral communities even if they do not devastate live corals. Recovery in this instance is expected to be very rapid given that all coral taxa persisted, and effects were greatest among fast growing corals.     

Population status of giant clams (Mollusca: Tridacnidae) in the northern Red Sea,Egypt

Throughout their range, giant clams (family Tridacnidae) are increasingly threatened by anthropogenic impacts and natural disasters, but little is known about their population status. In this first assessment of the tridacnid population at Abu Sauatir in the northern Red Sea, a total of 491 m² were surveyed and >200 clams recorded. Tridacna maxima was the only species found. The population's live:dead ratio was 3:1. Overall clam density was 0.08?±?0.008 live and 0.02?±?0.007 dead individuals per 0.25 m². Greatest densities occurred on the reef flat in 1 m depth (live), and on the northern reef slope in deeper waters (dead). On the slope, live clam density decreased significantly, whereas dead clam density increased significantly with depth. Sizes of live and dead individuals differed significantly. Live clams ranged from 1 to 30 cm (median 5 cm). Juveniles ≤2 cm (8.2% of the population) and individuals >11 cm occurred on the reef slope but not on the reef flat. Live clam sizes did not differ significantly between reef sites. Dead clam size ranged from 2 to 15 cm (median 6.5 cm). 2.1% of the empty shells were ≤2 cm long. Dead clam sizes differed significantly between 5 and 10 m depth on the northern reef slope. The low clam abundance (live and dead) in the shallowest and most easily accessible areas of the reef flat, combined with small sizes, strongly suggest artisanal reef-top gathering for meat and shells.     

Space partitioning by stony corals soft corals and benthic algae on the coral reefs of the northern Gulf of Eilat (Red Sea)

The major faunistic and floristic components occupying space on the coral reefs of the northern Gulf of Eilat (Red Sea) are stony corals, soft corals and benthic algae. The percent living coverage of the three components and the relative abundance of the different species of each component were studied by line transects, on the reef flats and the upper forereef zones of nine localities. A wider and higher range of living coverage values of stony corals were recorded at the upper fore-reef zones (18.30–49.09%) compared with the reef flats (5.50–31.66%) at the different stations. The most abundant stony corals on the reef flats areCyphastrea microphthalma, Stylophora pistillata, Favia favus, Porites lutea, Platygyra lamellina and the hydrozoanMillepora dichotoma. The fire coralM. dichotoma dominates the upper fore-reef zone in most of the stations. The average percent living coverage of soft corals on the reef flats ranged between 0.20 and 17.06%, and on the upper fore-reef zones between 1.68 and 15.13%. Seventy percent of the total living coverage of the soft coral community is contributed by 2 to 3 species. They tend to form large monospecific carpets, such as those composed ofSinularia sp.,Sarcophyton glaucum andLobophytum pauciflorum. The common benthic algae on the coral reef studied occur as turfs or macroscopic noncalcareous algae. They play a significant role in occupying space, especially on the reef flats. The most abundant algae recorded in all localities are the turfsSphacelaria tribuloides, Jania sp. and the macroscopic non-calcareous algaeTurbinaria elatensis andColpomenia sinuosa. Comparison between reef flats and upper fore-reef zones, in terms of average living cover of stony corals, shows that the variation among the reef flats is grater than the variation among the upper fore-reef zones. However, there is no significant variation in the average living coverage of soft corals between these two zones. Annual living-coverage values of algae on the reef flats are significantly higher than those of the upper fore-reef zones. Extremely low tides occurring periodically but unpredictably at Eilat cause mass mortality of the benthic communities on the reef flats reopening new spaces for settlement. The coexistence of stony corals, soft corals and algae on the reef ecosystem is due to different biological properties of each component. Opportunistic life histories of certain stony corals and most algae enable quick colonization of newly opened spaces. Lack of predators, high tolerance against abiotic factors and ability to form large aggregates of colonies are suggested as possible factors supporting the existence of soft corals in shallow water. Biological factors such as competition, predation and grazing pressure play an increasingly important role in controlling space utilization by the components studied with the advancement of succession.