Microbiome variation in corals with distinct depth distribution ranges across a shallow–mesophotic gradient (15–85 m)

Mesophotic coral ecosystems (MCEs) are generally poorly studied, and our knowledge of lower MCEs (below 60 m depth) is largely limited to visual surveys. Here, we provide a first detailed assessment of the prokaryotic community associated with scleractinian corals over a depth gradient to the lower mesophotic realm (15–85 m). Specimens of three Caribbean coral species exhibiting differences in their depth distribution ranges (Agaricia grahamae, Madracis pharensis and Stephanocoenia intersepta) were collected with a manned submersible on the island of Curaçao, and their prokaryotic communities assessed using 16S rRNA gene sequencing analysis. Corals with narrower depth distribution ranges (depth-specialists) were associated with a stable prokaryotic community, whereas corals with a broader niche range (depth-generalists) revealed a higher variability in their prokaryotic community. The observed depth effects match previously described patterns in Symbiodinium depth zonation. This highlights the contribution of structured microbial communities over depth to the coral’s ability to colonize a broader depth range.     

Pleistocene reef environments, constituent grains, and coral community structure: Curaçao, Netherlands Antilles

We investigated the degree to which component grains vary with depositional environment in sediments from three reef habitats from the Pleistocene (125?ka) Hato Unit of the Lower Terrace, Curaçao, Netherlands Antilles: windward reef crest, windward back reef, and leeward reef crest. The windward reef crest sediment is the most distinctive, dominated by fragments of encrusting and branching coralline red algae, coral fragments and the encrusting foraminiferan Carpenteria sp. Windward back reef and leeward reef crest sediments are more similar compositionally, only showing significant differences in relative abundance of coral fragments and Homotrema rubrum. Although lacking high taxonomic resolution and subject to modification by transport, relative abundance of constituent grain types offers a way of assessing ancient skeletal reef community composition, and one which is not limited to a single taxonomic group. The strong correlation between grain type and environment we found in the Pleistocene of Curaçao suggests that constituent grain analysis may be an effective tool in delineating Pleistocene Caribbean reef environments. However, it will not be a sufficient indicator where communities vary significantly within reef environments or where evolutionary and/or biogeographical processes lead to different relationships between community composition and reef environment. Detailed interpretation of geological, biological, and physical characteristics of the Pleistocene reefs of Curaçao reveals that the abundance of the single coral species, Acropora palmata, is not a good predictor of the ecological structure of the ancient reef coral communities. This coral was the predominant species in two of the three reef habitats (windward and leeward reef crest), but the taxonomic composition (based on species relative abundance data) of the reef coral communities was substantially different in these two environments. We conclude that qualitative estimates of coral distribution patterns (presence of a key coral species or the use of a distinctive coral skeletal architecture), when used as a component in a multi-component analysis of ancient reef environments, probably introduces minimal circular reasoning into quantitative paleoecological studies of reef coral community structure.     

An assessment of shallow and mesophotic reef brachyuran crab assemblages on the south shore of O‘ahu,Hawai‘i

Shallow coral reefs are extensively studied but, although scleractinian corals have been recorded to 165 m, little is known about other mesophotic coral reef ecosystem (MCE) inhabitants. Brachyuran crabs fill many ecological and trophic niches on reefs, making them ideal candidates for evaluating species composition among depths to ask whether MCEs host the same communities as shallower reef communities that have been well studied. Here we deployed autonomous reef monitoring structures for 2 yr on the south shore of O‘ahu along a depth gradient (12, 30, 60, and 90 m) to sample and assess brachyuran crab communities. A total of 663 brachyuran crabs representing 69 morphospecies (16 families) were found. Community composition was not significantly different within depths, but was highly stratified by depth. Each depth was distinct, but the 30 and 60 m depths were least dissimilar from one another. We show that deeper reefs host significantly different brachyuran communities, and at much lower total abundance, than shallow reefs in Hawai‘i, with 4–27 unique morphospecies per depth and only 3 of 69 morphospecies (~4 %) occurring across the entire depth range sampled.     

Zoantharia (Anthozoa: Hexacorallia) abundance and associations with Porifera and Hydrozoa across a depth gradient on the west coast of Curaçao

Despite much research on zoantharians, little work has been done on the abundance of these animals in depths greater than 5 m. In this study, we examined the abundance of zoantharian species and their associated fauna at four depths (5, 10, 20, and 30 m) at multiple sites on the leeward west side of Curaçao in the southern Caribbean in order to provide baseline data. We aimed to quantify zoantharian-host associations in Curaçao to examine if the associations present vary according to different sites and across depths. Results showed that zooxanthellate Palythoa and Zoanthus were most common in shallow waters (5 m), although perhaps less common than observed 35–45 years ago, while sponge-associated species of Macrocnemina were commonly observed at greater depths. There was increasing zoantharian abundance and diversity with greater depth, and two major clusters of zoantharian assemblages in multi-dimensional scaling analyses, one cluster of shallow 5-m datasets, and the other of the large majority of 10-, 20-, and 30-m datasets, demonstrating that assemblages differed significantly by depth (5 m vs. 10, 20, 30 m). In contrast, no significant differences in zoantharian abundance or assemblage were seen across different sites. The data from this study establish a baseline of zoantharian assemblage data that can be compared with other regions and to datasets from Curaçao in the future, which will be of importance in tracking ecological shifts in the face of on-going climate change.     

Spatial variability in reef-fish assemblages in shallow and upper mesophotic coral ecosystems in the Philippines

The variability in reef-fish species assemblages was examined at three geographic locations in the Philippines (Apo, Abra and Patn), each showing varying levels of disturbances (low to high) at two depths, shallow-water reef (SWR; 8–20 m) and the upper mesophotic coral ecosystem (MCE; 30–35 m). Fish species assemblages varied among locations and between depths. Differences in fish assemblages among locations corresponded to the variability in benthic assemblages and levels of disturbances, wherein locations with higher coral cover and less disturbances had the highest fish species richness, abundance and biomass. Variation in fish assemblages between depths was also associated with changes in benthic assemblages and possibly inaccessibility to local fishing techniques. Fish species richness decreased with depth in all locations, but biomass increased only in the MCEs of Apo and Abra, which is a similar pattern exhibited in many MCEs. Our results suggest that despite location differences, depth had a relatively consistent influence on fish species assemblages, particularly in locations exposed to low and intermediate disturbance. Under high disturbance, MCEs exhibit similar vulnerability to SWRs.     

Nitrogen fixation rates in algal turf communities of a degraded versus less degraded coral reef

Algal turf communities are ubiquitous on coral reefs in the Caribbean and are often dominated by N₂-fixing cyanobacteria. However, it is largely unknown (1) how much N₂ is actually fixed by turf communities and (2) which factors affect their N₂ fixation rates. Therefore, we compared N₂ fixation activity by turf communities at different depths and during day and night-time on a degraded versus a less degraded coral reef site on the island of Curaçao. N₂ fixation rates measured with the acetylene reduction assay were slightly higher in shallow (5–10-m depth) than in deep turf communities (30-m depth), and N₂ fixation rates during the daytime significantly exceeded those during the night. N₂ fixation rates by the turf communities did not differ between the degraded and less degraded reef. Both our study and a literature survey of earlier studies indicated that turf communities tend to have lower N₂ fixation rates than cyanobacterial mats. However, at least in our study area, turf communities were more abundant than cyanobacterial mats. Our results therefore suggest that turf communities play an important role in the nitrogen cycle of coral reefs. N₂ fixation by turfs may contribute to an undesirable positive feedback that promotes the proliferation of algal turf communities while accelerating coral reef degradation.     

Mesophotic depths as refuge areas for fishery-targeted species on coral reefs

Coral reefs are subjected to unprecedented levels of disturbance with population growth and climate change combining to reduce standing coral cover and stocks of reef fishes. Most of the damage is concentrated in shallow waters (<30 m deep) where humans can comfortably operate and where physical disturbances are most disruptive to marine organisms. Yet coral reefs can extend to depths exceeding 100 m, potentially offering refuge from the threats facing shallower reefs. We deployed baited remote underwater stereo-video systems (stereo-BRUVs) at depths of 10–90 m around the southern Mariana Islands to investigate whether fish species targeted by fishing in the shallows may be accruing benefits from being at depth. We show that biomass, abundance and species richness of fishery-targeted species increased from shallow reef areas to a depth of 60 m, whereas at greater depths, a lack of live coral habitat corresponded to lower numbers of fish. The majority of targeted species were found to have distributions that ranged from shallow depths (10 m) to depths of at least 70 m, emphasising that habitat, not depth, is the limiting factor in their vertical distribution. While the gradient of abundance and biomass versus depth was steepest for predatory species, the first species usually targeted by fishing, we also found that fishery-targeted herbivores prevailed in similar biomass and species richness to 60 m. Compared to shallow marine protected areas, there was clearly greater biomass of fishery-targeted species accrued in mesophotic depths. Particularly some species typically harvested by depth-limited fishing methods (e.g., spearfishing), such as the endangered humphead wrasse Cheilinus undulatus, were found in greater abundance on deeper reefs. We conclude that mesophotic depths provide essential fish habitat and refuge for fishery-targeted species, representing crucial zones for fishery management and research into the resilience of disturbed coral reef ecosystems.     

Geomorphology and benthic cover of mesophotic coral ecosystems of the upper insular slope of southwest Puerto Rico

The upper insular slope of southwest Puerto Rico is defined as extending from the shelf break at ~20 m water depth down to a depth of ~160 m where there is a pronounced change in geomorphic character and the basal slope begins. The upper slope is divided into two geomorphic zones separated by a pronounced break in slope gradient at ~90 m water depth. Descending from the shelf break, these are Zone I (20–90 m) and Zone II (90–160 m). As orientation of the shelf margin changes, geomorphology of Zone I shows systematic variations consistent with changes in exposure to prevailing waves. Within Zone I, exposed southeast-facing slopes have a gentler gradient and lower relief than more sheltered southwest-facing slopes, which are steep and irregular. Mesophotic coral ecosystems (MCEs) are largely restricted to Zone I and concentrated on topographic highs removed from the influence of active downslope sediment transport. Accordingly, MCEs are more abundant, extensive and diverse on southwest-facing slopes where irregular topography funnels downslope sediment transport into steep narrow grooves. MCEs are more sporadic and widely spaced on southeast-facing slopes where topographic highs are more widely spaced and downslope sediment transport is spread over open, low-relief slopes inhibiting coral recruitment and growth. Relict features formed during preexisting sea levels lower than present include deep buttresses at ~45–65 m water depth and a prominent terrace at ~80 m. Based on correlations with existing reef accretion and sea-level records, it is proposed that the 80-m terrace formed during the last deglaciation ~14–15 ka and subsequently drowned during a period of rapid sea-level rise associated with meltwater pulse 1A at ~14 ka and deep buttresses at ~45–65 m formed between ~11.5 and 13.5 ka and then drowned during a period of rapid sea-level rise associated with meltwater pulse 1B at ~11.3 ka.     

Community ecology of mesophotic coral reef ecosystems

Given the global degradation of shallow-water coral reef ecosystems resulting from anthropogenic activities, mesophotic coral reef ecosystems (MCEs) are gaining attention because they are generally considered a de facto refuge for shallow-water species. Despite their inferred importance, MCEs remain one of the most understudied reef habitats, and basic information on the taxonomic composition, depth range, habitat preferences, and abundance and distribution of MCE taxa is scarce. The processes that structure these communities are virtually unknown. Here, we provide a review of what is known about MCEs community ecology and outline essential gaps in our knowledge of these deeper water coral reef ecosystems. The primary findings of this review are as follows: (1) many dominant shallow-water species are absent from MCEs; (2) compared to shallow reefs, herbivores are relatively scarce, perhaps due to limited habitat complexity at depth; (3) changes in the dominant photosynthetic taxa with depth suggest adaptation and specialization to depth; (4) evidence regarding the importance of heterotrophy for zooxanthellate corals at depth is conflicting and inconclusive; and (5) decreased light with depth, but not temperature, appears to be the primary factor limiting the depth of MCEs. The majority of research done to date has been performed in the Caribbean, where some generalization can be made about the community structure and distribution of MCEs. The larger and more diverse Indo-Pacific remains largely unexplored with no apparent generalizations from the few sites that have been comparatively well studied. For MCEs, large gaps in knowledge remain on fundamental aspects of ecology. Advanced technologies must be harnessed and logistical challenges overcome to close this knowledge gap and empower resource managers to make informed decisions on conserving shallow-water and mesophotic coral reef ecosystems.     

Population Structure of Montastraea cavernosa on Shallow versus Mesophotic Reefs in Bermuda

Mesophotic coral reef ecosystems remain largely unexplored with only limited information available on taxonomic composition, abundance and distribution. Yet, mesophotic reefs may serve as potential refugia for shallow-water species and thus understanding biodiversity, ecology and connectivity of deep reef communities is integral for resource management and conservation. The Caribbean coral, Montastraea cavernosa, is considered a depth generalist and is commonly found at mesophotic depths. We surveyed abundance and size-frequency of M. cavernosa populations at six shallow (10m) and six upper mesophotic (45m) sites in Bermuda and found population structure was depth dependent. The mean surface area of colonies at mesophotic sites was significantly smaller than at shallow sites, suggesting that growth rates and maximum colony surface area are limited on mesophotic reefs. Colony density was significantly higher at mesophotic sites, however, resulting in equal contributions to overall percent cover. Size-frequency distributions between shallow and mesophotic sites were also significantly different with populations at mesophotic reefs skewed towards smaller individuals. Overall, the results of this study provide valuable baseline data on population structure, which indicate that the mesophotic reefs of Bermuda support an established population of M. cavernosa.     

40 Years of benthic community change on the Caribbean reefs of Curaçao and Bonaire: the rise of slimy cyanobacterial mats

Over the past decades numerous studies have reported declines in stony corals and, in many cases, phase shifts to fleshy macroalgae. However, long-term studies documenting changes in other benthic reef organisms are scarce. Here, we studied changes in cover of corals, algal turfs, benthic cyanobacterial mats, macroalgae, sponges and crustose coralline algae at four reef sites of the Caribbean islands of Curaçao and Bonaire over a time span of 40 yr. Permanent 9 m² quadrats at 10, 20, 30 and 40 m depth were photographed at 3- to 6-yr intervals from 1973 to 2013. The temporal and spatial dynamics in the six dominant benthic groups were assessed based on image point-analysis. Our results show consistent patterns of benthic community change with a decrease in the cover of calcifying organisms across all sites and depths from 32.6 (1973) to 9.2% (2013) for corals and from 6.4 to 1% for crustose coralline algae. Initially, coral cover was replaced by algal turfs increasing from 24.5 (1973) to 38% around the early 1990s. Fleshy macroalgae, still absent in 1973, also proliferated covering 12% of the substratum approximately 20 yr later. However, these new dominants largely declined in abundance from 2002 to 2013 (11 and 2%, respectively), marking the rise of benthic cyanobacterial mats. Cyanobacterial mats became the most dominant benthic component increasing from a mere 7.1 (2002) to 22.2% (2013). The observed increase was paralleled by a small but significant increase in sponge cover (0.5 to 2.3%). Strikingly, this pattern of degradation and phase change occurred over the reef slope down to mesophotic depths of 40 m. These findings suggest that reefs dominated by algae may be less stable than previously thought and that the next phase may be the dominance of slimy cyanobacterial mats with some sponges.     

Mesophotic coral ecosystems in the Hawaiian Archipelago

Efforts to map coral reef ecosystems in the Hawaiian Archipelago using optical imagery have revealed the presence of numerous scleractinian, zoothanthellate coral reefs at depths of 30–130+ m, most of which were previously undiscovered. Such coral reefs and their associated communities have been recently defined as mesophotic coral ecosystems (MCEs). Several types of MCEs are found in Hawai‘i, each of which dominates a different depth range and is characterized by a unique pattern of coral community structure and colony morphology. Although MCEs are documented near both ends of the archipelago and on many of the islands in between, the maximum depth and prevalence of MCEs in Hawai‘i were found to decline with increasing latitude. The Main Hawaiian Islands (MHI) had significantly deeper and greater percentages of scleractinian coral, and peaks in cover of both scleractinian corals and macroalgae occurred within depth bins 20 m deeper than in the Northwestern Hawaiian Islands (NWHI). Across the archipelago, as depth increased the combined percentage of living cover of mega benthic taxa declined sharply with increasing depth below 70 m, despite the widespread availability of hard substrate.     

Long-term dynamics of the brown macroalga <Emphasis Type="Italic">Lobophora variegata</Emphasis> on deep reefs in Curaçao

Lobophora variegata occurs in the eulittoral zone and in deep water on coral reefs in Curaçao. An analysis of the long-term (1979–2006) changes in the vertical distribution of the macroalga in permanent quadrats indicated a significant increase in cover of the deepwater community. In 1998, Lobophora covered 1 and 5% of the quadrats at 20 and 30 m, respectively. By 2006, these values had risen to 25 and 18%, precipitating a shift in abundance of corals and macroalgae at both depths. This increase coincided with losses in coral cover, possibly linked to bleaching, disease and storm-related mortality in deep water plating Agaricia corals. In contrast, macroalgae remained relatively rare (<6% cover) on shallower (10 m) and deeper (40 m) reefs despite declines in coral cover also occurring at these depths, illustrating the depth-dependent dynamics of coral reefs. Several hypotheses are suggested to explain these changes.     

Character release following extinction in a Caribbean reef coral species complex

Abstract The Pleistocene extinction of the widespread organ‐pipe Montastraea coral had measurable morphological and ecological effects on surviving lineages of the Montastraea“annularis” species complex. Extinction of the organ‐pipe Montastraea occurred after more than 500,000 years of dominance in the shallow‐water reef habitat of Barbados. Extinction resulted in a morphological shift of the columnar Montastraea lineage from thick to thin columns in modern reef environments. Pleistocene colonies of the columnar morphotype sympatric with organ‐pipe Montastraea showed greater column widths than those in allopatry. We subjected our data to a number of criteria for interpreting the morphological shift as character release following lifting of competitive pressure after extinction. The morphological differences do not appear to be due either to chance or to physical properties of the marine environment. Differential local extinction and recolonization of four members of the species complex did not occur on Barbados, so that the species coexisted and appear to have coevolved between more than 600,000 and 82,000 years ago. The morphological shift is related to coral growth form and growth rate, and thus reflects the acquisition of a primary resource in corals‐light. Character release occurred at the same oceanic Caribbean island (Barbados) where environments have fluctuated with similar variance throughout the period of coexistence. Not only has competition among living members of the Montastraea“annularis” species complex been convincingly demonstrated, but trends in relative abundance among fossil members of the species complex strongly suggest that a competitive hierarchy was operating during their Pleistocene coexistence on Barbados. We also observed an ecological analogue to character release on another Caribbean island, Curaçao. The distribution and abundance of living columnarM. annularis s.s. and massive M. faveolata from the leeward reef crest in Curaçao is greater now than in the Pleistocene, when organ‐pipe Montastraea dominated this shallow‐water reef habitat. Extinction of the faster growing, shallow‐water organ‐pipe Montastraea resulted in higher abundance of the columnar Montastraea lineage in shallow‐water habitats, where it shifted its morphology to one adapted to high light levels. The species extinction released surviving lineages from a competitive network that had resulted in lower rank abundance in the Pleistocene community and enhanced abundance of both columnar M. annularis s.s. and M. faveolata in modern communities. Full validation of our interpretation of character release must await experiments that demonstrate whether phenotypic differences between populations have a genetic basis. However, we believe the results of this study point to the important, yet heretofore neglected, role that biological interactions have played in the evolution of closely related reef coral species.     

Structure of Caribbean coral reef communities across a large gradient of fish biomass

The collapse of Caribbean coral reefs has been attributed in part to historic overfishing, but whether fish assemblages can recover and how such recovery might affect the benthic reef community has not been tested across appropriate scales. We surveyed the biomass of reef communities across a range in fish abundance from 14 to 593 g m⁻², a gradient exceeding that of any previously reported for coral reefs. Increased fish biomass was correlated with an increased proportion of apex predators, which were abundant only inside large marine reserves. Increased herbivorous fish biomass was correlated with a decrease in fleshy algal biomass but corals have not yet recovered.     

The influence of multiple factors upon reef fish abundance and species richness in a tropical coral complex

The present study was conducted on Tamandaré reefs, northeast Brazil and aimed to analyse the importance of different factors (e.g. tourism activity, fishing activity, coral abundance and algal abundance) on reef fish abundance and species richness. Two distinct reef areas (A ver o mar and Caieiras) with different levels of influence were studied. A total of 8239 reef fish individuals were registered, including 59 species. Site 1 (A ver o mar) presented higher reef fish abundance and richness, with dominance of roving herbivores (29.9 %) and mobile invertebrate feeders (28.7 %). In contrast, at Site 2 (Caieiras) territorial herbivores (40.9 %) predominated, followed by mobile invertebrate feeders (24.6 %). Concerning the benthic community, at Site 1 macroalgae were recorded as the main category (49.3 %); however, Site 2 was dominated by calcareous algae (36.0 %). The most important variable explaining more than 90 % of variance on reef fish abundance and species richness was macroalgae abundance, followed by fishing activity. Phase shifts on coral reefs are evident, resulting in the replacement of coral by macroalgae and greatly influencing reef fish communities. In this context, it is important to understand the burden of the factors that affect reef fish communities and, therefore, influence the extinction vulnerability of coral reef fishes.     

Distribution,food preference,and trophic position of the corallivorous fireworm Hermodice carunculata in a Caribbean coral reef

The fireworm Hermodice carunculata is a facultative corallivore on coral reefs. It can interact with algal overgrowth to cause coral mortality. However, because of its cryptic nature, little is known about its ecology. We used micropredator attracting devices (MADs) and stable isotope analyses to provide insights into the distribution and diet of H. carunculata in a coral reef on Curaçao, southern Caribbean. MADs consisted of algal clumps inside accessible mesh nets which H. carunculata could use as refuge. To obtain indications on its distribution pattern, MADs filled with Halimeda opuntia were deployed in different reef habitats ranging from 0 to 16 m water depth. Fireworms were found inside MADs in all reef habitats, indicating that they have a widespread horizontal and vertical distribution, ranging from the shoreline to the deeper reef slope. On the reef crest, MADs were filled using different algal species and deployed on dead or live scleractinian corals. MADs hosted more fireworms when placed on live corals, regardless of algal species used, suggesting that algal-induced corallivory may be widespread. To test for food preferences, different food sources were added inside the MADs. Fireworms detected potential prey within 6 h and were significantly more attracted by decaying corals and raw fish than by live corals, hydrozoans, or gorgonians. Stable isotope analyses indicated detritus, macroalgae, and scleractinian corals as potential food sources and revealed an ontogenetic dietary shift toward enriched δ ¹³C and δ ¹⁵N values with increasing fireworm size, suggesting that large-sized individuals feed on food sources of higher trophic levels. Our findings highlight H. carunculata as a widespread, and omnivorous scavenger that has the potential to switch feeding toward weakened or stressed corals, thereby likely acting as a harmful corallivore on degraded reefs.     

Abundant betaines in reef-building corals and ecological indicators of a photoprotective role

Betaines are well known as compatible solutes that exert protein- and membrane-stabilizing effects, including protective effects on photosynthesis in plants and free-living algae stressed by high irradiance or unusual temperatures. Betaines, however, have received minimal attention in reef-building corals. One goal of this research was to identify and quantify the betaines of reef-building corals with chemically definitive methods. Metabolite profiling was conducted on 10 species (6 genera) of Curaçao corals by liquid chromatography/time-of-flight mass spectrometry calibrated using six stable-isotope-labeled internal standards. Glycine betaine (GlyB), proline betaine (ProB), alanine betaine (AlaB), β-alanine betaine, hydroxyproline betaine (HProB), taurine betaine (TauB), trigonelline (Trig) and the chemically related sulfonium compound dimethylsulfoniopropionate were found in all species. Relative levels of betaines varied across species, with GlyB and ProB being most prominent. Betaines were collectively abundant; estimated total concentrations were 12 to 204 (mean = 75) mmol per liter of tissue. A second goal was to examine ecological patterns in betaine concentrations in field populations of Curaçao corals. Betaine concentrations exhibited intraspecific patterns that matched a priori predictions for molecules that defend photosynthesis against negative effects of high irradiance. In Madracis mirabilis—which occupies unshaded locations—GlyB, ProB, AlaB, HProB, and Trig were 37–94% more abundant in colonies at 5 m depth (high irradiance) than 20 m. In M. pharensis—which occupies exposed and shaded locations—GlyB, ProB, and AlaB were 30–44% more abundant in unshaded than shaded colonies at one depth. M. senaria exhibited 45–93% increases in concentrations of betaines—GlyB, AlaB, HProB, TauB, and Trig—between early and late in the day, although M. mirabilis and pharensis did not. The results indicate that multiple betaines occur commonly in reef-building coral species, and betaine concentrations are modulated in response to growth light conditions in ways consistent with betaines acting as agents of photoprotection of coral photosynthesis.     

<Emphasis Type="Italic">Euphyllia paradivisa</Emphasis>, a successful mesophotic coral in the northern Gulf of Eilat/Aqaba,Red Sea

Mesophotic coral ecosystems (MCEs) host a thriving community of biota that has remained virtually unexplored. Here we report for the first time on a large population of the endangered coral species Euphyllia paradivisa from the MCEs of the Gulf of Eilat/Aqaba (GOE/A), Red Sea. The mesophotic zone in some parts of the study site harbors a specialized coral community predominantly comprising E. paradivisa (73 % of the total coral cover), distributed from 36 to 72 m depth. Here we sought to elucidate the strict distribution but high abundance of E. paradivisa in the MCEs at the GOE/A. We present 4 yr of observations and experiments that provide insight into the physiological plasticity of E. paradivisa: its low mortality rates at high light intensities, high competitive abilities, successful symbiont adaptation to the shallow-water environment, and tolerance to bleaching conditions or survival during prolonged bleaching. Despite its ability to survive under high irradiance in shallow water, E. paradivisa is not found in the shallow reef of the GOE/A. We suggest several factors that may explain the high abundance and exclusivity of E. paradivisa in the MCE: its heterotrophic capabilities; its high competition abilities; the possibility of it finding a deep-reef refuge there from fish predation; and its concomitant adaptation to this environment.