Yossi Loya

Community structure and species diversity of hermatypic corals at Eilat, Red Sea

This study investigates the community structure of reefbulding corals in terms of species composition, zonation and diversity patterns, as well as possible factors affecting the observed distributions. The study was carried out by a series of line transects run underwater with SCUBA apparatus from the reef flat to a depth of 30 m. The reefs of Eilat are of the fringing type, with seleractinian corals as the most important hermatypic organisms. The different zones of the reef are analyzed on the basis of topographical characteristics of the reef, as well as from the numerical data on abundance and living coverage, using cluster analysis of all transects surveyed. The present knowledge concerning species diversity is reviewed and analyzed in the context of hermatypic coral data. Three different diversity indices (the species count, Simpsons index and Shannon and Weavers index) were calculated for estimating the diversity obtained on different zones of the reef. It was found that there is a successive increase in diversity of hermatypic corals from shallow water to a depth of 30 m. Species diversity and living voverage of corals were significantly greater in steeper zones as compared to flatter zones of the reef. A possible explanation for this phenomenon is the accumulation of sediments in the flat zones. It is proposed that the severe and umpredictable nature of the reef flat may account for low abundance and living coverage of corals. It is also proposed that deep-water species which do not invade shallow water are species which have developed high specialization to their local environment. The idea that light intensity is a significant factor in calcium-carbonate deposition by scleractinian corals is supported by field measurements of individual colonies at different depths.

Global human footprint on the linkage between biodiversity and ecosystem functioning in reef fishes.

A global survey of reef fishes shows that the consequences of biodiversity loss are greater than previously anticipated as ecosystem functioning remained unsaturated with the addition of new species. Additionally, reefs worldwide, particularly those most diverse, are highly vulnerable to human impacts that are widespread and likely to worsen due to ongoing coastal overpopulation.

Vibrio shiloi sp. nov., the causative agent of bleaching of the coral Oculina patagonica

The aetiological agent of bleaching of the coral Oculina patagonica was characterized as a new Vibrio species on the basis of 16S rDNA sequence, DNA-DNA hybridization data and phenotypic properties, including the cellular fatty acid profile. Based on its 16S rDNA and DNA-DNA hybridization, the new Vibrio species is closely related to Vibrio mediterranei. The name Vibrio shiloi sp. nov. is proposed for the new coral-bleaching species, the type strain being AK1T (= ATCC BAA-91T = DSM 13774T).

A coral oxygen isotope record from the northern Red Sea documenting NAO, ENSO, and North Pacific teleconnections on Middle East climate variability since the year 1750

A 245-year coral oxygen isotope record from the northern Red Sea (Ras Umm Sidd/Egypt, ∼28°N) in bimonthly resolution is presented. The mean annual coral δ18O signal apparently reflects varying proportions of both sea surface temperature and δ18Oseawater variability. In conjunction with instrumental observations of climate the coral record suggests for interannual and longer timescales that colder periods are accompanied by more arid conditions in the northern Red Sea but increased rainfall in the southeastern Mediterranean, whereas warmer periods are accompanied by decreased rainfall in the latter and less arid conditions in the northern Red Sea. A ∼70-year oscillation of probably North Atlantic origin dominates the coral time series. Interannual to interdecadal variability is correlated with instrumental indices of the North Atlantic Oscillation (NAO), the El Nino-Southern Oscillation (ENSO), and North Pacific climate variability. The results suggest that these modes contributed consistently to Middle East climate variability since at least 1750, preferentially at a period of ∼5.7 years.

Extreme Diel Fluctuations of Oxygen in Diffusive Boundary Layers Surrounding Stony Corals

The diffusive boundary layers surrounding sessile marine organisms have been implicated in controlling an organisms metabolism and growth. We studied boundary layers surrounding hermatypic corals by monitoring oxygen concentrations on a submillimetric scale. Oxygen concentration within the boundary layers varied from supersaturation during the day to anoxia at night, although the ambient water composition remained constant. Detailed mapping and oxygen measurements revealed diel oxygen fluctuations from supersaturation (373% air saturation) in the light to complete oxygen depletion at darkness in the massive coral Favia favus. Exposure to a 5-cm/s current reduced the boundary layer thickness from 2.44 mm to 1.90 mm, allowing more rapid oxygen exchange across the diffusive boundary layer. Similar patterns were found in the branching coral Stylophora pistillata. In massive corals, the thickness of the diffusive boundary layer was negatively correlated with the size of the polyp. We suggest that the distribution of corals in areas of differential turbulence is related to the thickness of the diffusive boundary layers surrounding them.

Penetration of the Coral-Bleaching Bacterium Vibrio shiloi into Oculina patagonica

ABSTRACT Inoculation of the coral-bleaching bacterium Vibrio shiloi into seawater containing its host Oculina patagonica led to adhesion of the bacteria to the coral surface via a β-d-galactose receptor, followed by penetration of the bacteria into the coral tissue. The internalized V. shiloi cells were observed inside the exodermal layer of the coral by electron microscopy and fluorescence microscopy using specific anti-V. shiloi antibodies to stain the intracellular bacteria. At 29°C, 80% of the bacteria bound to the coral within 8 h. Penetration, measured by the viable count (gentamicin invasion assay) inside the coral tissue, was 5.6, 20.9, and 21.7% of the initial inoculum at 8, 12, and 24 h, respectively. The viable count in the coral tissue decreased to 5.3% at 48 h, and none could be detected at 72 h. Determination of V. shiloi total counts (using the anti-V. shiloiantibodies) in the coral tissue showed results similar to viable counts for the first 12 h of infection. After 12 h, however, the total count more than doubled from 12 to 24 h and continued to rise, reaching a value 6 times that of the initial inoculum at 72 h. Thus, the intracellular V. shiloi organisms were transformed into a form that could multiply inside the coral tissue but did not form colonies on agar medium. Internalization of the bacteria was accompanied by the production of high concentrations of V. shiloi toxin P activity in the coral tissue. Internalization and multiplication of V. shiloi are discussed in terms of the mechanism of bacterial bleaching of corals.

Climate change impedes scleractinian corals as primary reef ecosystem engineers.

Coral reefs are among the most diverse and productive ecosystems on our planet. Scleractinian corals function as the primary reef ecosystem engineers, constructing the framework that serves as a habitat for all other coral reef-associated organisms. However, the corals engineering role is particularly susceptible to global climate change. Ocean warming can cause extensive mass coral bleaching, which triggers dysfunction of major engineering processes. Sub-lethal bleaching results in the reduction of both primary productivity and coral calcification. This may lead to changes in the release of organic and inorganic products, thereby altering critical biogeochemical and recycling processes in reef ecosystems. Thermal stress-induced bleaching and subsequent coral mortality, along with ocean acidification, further lead to long-term shifts in benthic community structure, changes in topographic reef complexity, and the modification of reef functioning. Such shifts may cause negative feedback loops and further modification of coral-derived inorganic and organic products. This review emphasises the critical role of scleractinian corals as reef ecosystem engineers and highlights the control of corals over key reef ecosystem goods and services, including high biodiversity, coastal protection, fishing, and tourism. Thus, climate change by impeding coral ecosystem engineers will impair the ecosystem functioning of entire reefs.

An updated 18S rRNA phylogeny of tunicates based on mixture and secondary structure models.

BackgroundTunicates have been recently revealed to be the closest living relatives of vertebrates. Yet, with more than 2500 described species, details of their evolutionary history are still obscure. From a molecular point of view, tunicate phylogenetic relationships have been mostly studied based on analyses of 18S rRNA sequences, which indicate several major clades at odds with the traditional class-level arrangements. Nonetheless, substantial uncertainty remains about the phylogenetic relationships and taxonomic status of key groups such as the Aplousobranchia, Appendicularia, and Thaliacea.ResultsThirty new complete 18S rRNA sequences were acquired from previously unsampled tunicate species, with special focus on groups presenting high evolutionary rate. The updated 18S rRNA dataset has been aligned with respect to the constraint on homology imposed by the rRNA secondary structure. A probabilistic framework of phylogenetic reconstruction was adopted to accommodate the particular evolutionary dynamics of this ribosomal marker. Detailed Bayesian analyses were conducted under the non-parametric CAT mixture model accounting for site-specific heterogeneity of the evolutionary process, and under RNA-specific doublet models accommodating the occurrence of compensatory substitutions in stem regions. Our results support the division of tunicates into three major clades: 1) Phlebobranchia + Thaliacea + Aplousobranchia, 2) Appendicularia, and 3) Stolidobranchia, but the position of Appendicularia could not be firmly resolved. Our study additionally reveals that most Aplousobranchia evolve at extremely high rates involving changes in secondary structure of their 18S rRNA, with the exception of the family Clavelinidae, which appears to be slowly evolving. This extreme rate heterogeneity precluded resolving with certainty the exact phylogenetic placement of Aplousobranchia. Finally, the best fitting secondary-structure and CAT-mixture models suggest a sister-group relationship between Salpida and Pyrosomatida within Thaliacea.ConclusionAn updated phylogenetic framework for tunicates is provided based on phylogenetic analyses using the most realistic evolutionary models currently available for ribosomal molecules and an unprecedented taxonomic sampling. Detailed analyses of the 18S rRNA gene allowed a clear definition of the major tunicate groups and revealed contrasting evolutionary dynamics among major lineages. The resolving power of this gene nevertheless appears limited within the clades composed of Phlebobranchia + Thaliacea + Aplousobranchia and Pyuridae + Styelidae, which were delineated as spots of low resolution. These limitations underline the need to develop new nuclear markers in order to further resolve the phylogeny of this keystone group in chordate evolution.

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.

Endolithic algae: an alternative source of photoassimilates during coral bleaching

Recent reports of worldwide coral bleaching events leading to devastating coral mortality have caused alarm among scientists and resource managers. Differential survival of coral species through bleaching events has been widely documented. We suggest that among the possible factors contributing to survival of coral species during such events are endolithic algae harboured in their skeleton, providing an alternative source of energy. We studied the dynamics of photosynthetic pigment concentrations and biomass of endoliths in the skeleton of the encrusting coral Oculina patagonica throughout a bleaching event. During repeated summer bleaching events these endolithic algae receive increased photosynthetically active radiation, increase markedly in biomass, and produce increasing amounts of photoassimilates, which are translocated to the coral. Chlorophyll concentrations and biomass of endoliths were 4.6 ± 1.57 and 1570 ± 427 μg cm−2 respectively, in skeletons of relatively healthy colonies (0–40%bleaching) but up to 14.8± 2.5 and 4036 ± 764 g cm−2 endolith chlorophyll and biomass respectively, in skeletons of bleached colonies (greater than 40% bleaching). The translocation dynamics of 14C–labelled photoassimilates from the endoliths to bleached coral tissue showed significantly higher 14C activity of the endoliths harboured within the skeletons of bleached corals than that of the endoliths in non–bleached corals. This alternative source of energy may be vital for the survivorship of O. patagonica, allowing gradual recruitment of zooxanthellae and subsequent recovery during the following winter.