Assessment of temperature optimum signatures of corals at both latitudinal extremes of the Red Sea.

Rising ocean temperatures are pushing reef-building corals beyond their temperature optima (Topt ), resulting in reduced physiological performances and increased risk of bleaching. Identifying refugia with thermally resistant corals and understanding their thermal adaptation strategy is therefore urgent to guide conservation actions. The Gulf of Aqaba (GoA, northern Red Sea) is considered a climate refuge, hosting corals that may originate from populations selected for thermal resistance in the warmer waters of the Gulf of Tadjoura (GoT, entrance to the Red Sea and 2000 km south of the GoA). To better understand the thermal adaptation strategy of GoA corals, we compared the temperature optima (Topt ) of six common reef-building coral species from the GoA and the GoT by measuring oxygen production and consumption rates as well as photophysiological performance (i.e. chlorophyll fluorescence) in response to a short heat stress. Most species displayed similar Topt between the two locations, highlighting an exceptional continuity in their respective physiological performances across such a large latitudinal range, supporting the GoA refuge theory. Stylophora pistillata showed a significantly lower Topt in the GoA, which may suggest an ongoing population-level selection (i.e. adaptation) to the cooler waters of the GoA and subsequent loss of thermal resistance. Interestingly, all Topt were significantly above the local maximum monthly mean seawater temperatures in the GoA (27.1°C) and close or below in the GoT (30.9°C), indicating that GoA corals, unlike those in the GoT, may survive ocean warming in the next few decades. Finally, Acropora muricata and Porites lobata displayed higher photophysiological performance than most species, which may translate to dominance in local reef communities under future thermal scenarios. Overall, this study is the first to compare the Topt of common reef-building coral species over such a latitudinal range and provides insights into their thermal adaptation in the Red Sea.

Elevated temperatures reduce the resilience of the Red Sea branching coral stylophora pistillata to copper pollution.

Copper (Cu) is a common marine pollutant of coastal environments and can cause severe impacts on coral organisms. To date, only a few studies assessed the effects of Cu contamination in combination with elevated seawater temperatures on corals. Furthermore, experiments focusing on coral recovery during a depuration phase, and under thermal stress, are lacking. The present study investigated the physiological response of the common and thermally tolerant scleractinian coral Stylophora pistillata from the northern Red Sea to Cu contamination (2.5, 5 or 10 µg L - 1) in combination with thermal stress (5 °C above local ambient temperatures (26 °C)) for 23 days, and assessed the impact of elevated temperatures on its ability to recover from such pollution during a one-week depuration period. Variation in coral photo-physiological biomarkers including antioxidant defense capacity, were dose, time and temperature-dependent, and revealed additive effects of elevated temperatures. Successful recovery was achieved in ambient temperature only and was mediated by antioxidant defenses. Elevation of temperature altered the recovery dynamics during depuration, causing reduced Cu bioaccumulation and photosynthetic yield. The present study provides novel information on the effects of elevated temperature on the resilience (resistance and recovery processes) of a scleractinian coral exposed to a common marine pollutant. Our findings suggest that ocean warming may alter the resilience strategies of corals when exposed to local pollution, an impact that might have long-term consequences on the chances of survival of reefs in increasingly populated and warming coastal environments.

Desert dust deposition supplies essential bioelements to Red Sea corals.

Climate change-related increase in seawater temperature has become a leading cause of coral bleaching and mortality. However, corals from the northern Red Sea show high thermal tolerance and no recorded massive bleaching event. This specific region is frequently subjected to intense dust storms, coming from the surrounding arid deserts, which are expected to increase in frequency and intensity in the future. The aerial dust deposition supplies essential bioelements to the water column. Here, we investigated the effect of dust deposition on the physiology of a Red Sea coral, Stylophora pistillata. We measured the modifications in coral and Symbiodiniaceae metallome (cellular metal content), as well as the changes in photosynthesis and oxidative stress status of colonies exposed during few weeks to dust deposition. Our results show that 1 mg L-1 of dust supplied nanomolar amounts of nitrate and other essential bioelements, such as iron, manganese, zinc and copper, rapidly assimilated by the symbionts. At 25°C, metal bioaccumulation enhanced the chlorophyll concentration and photosynthesis of dust-exposed corals compared to control corals. These results suggest that primary production was limited by metal availability in seawater. A 5°C increase in seawater temperature enhanced iron assimilation in both control and dust-enriched corals. Temperature rise increased the photosynthesis of control corals only, dust-exposed ones having already reached maximal photosynthesis rates at 25°C. Finally, we observed a combined effect of temperature and bioelement concentration on the assimilation of molybdenum, cadmium, manganese and copper, which were in higher concentrations in symbionts of dust-exposed corals maintained at 30°C. All together these observations highlight the importance of dust deposition in the supply of essential bioelements, such as iron, to corals and its role in sustaining coral productivity in Red Sea reefs.

Instability and Stasis Among the Microbiome of Seagrass Leaves, Roots and Rhizomes, and Nearby Sediments Within a Natural pH Gradient.

Seagrass meadows are hotspots of biodiversity with considerable economic and ecological value. The health of seagrass ecosystems is influenced in part by the makeup and stability of their microbiome, but microbiome composition can be sensitive to environmental change such as nutrient availability, elevated temperatures, and reduced pH. The objective of the present study was to characterize the bacterial community of the leaves, bulk samples of roots and rhizomes, and proximal sediment of the seagrass species Cymodocea nodosa along the natural pH gradient of Levante Bay, Vulcano Island, Italy. The bacterial community was determined by characterizing the 16S rRNA amplicon sequencing and analyzing the operational taxonomic unit classification of bacterial DNA within samples. Statistical analyses were used to explore how life-long exposure to different pH/pCO2 conditions may be associated with significant differences in microbial communities, dominant bacterial classes, and microbial diversity within each plant section and sediment. The microbiome of C. nodosa significantly differed among all sample types and site-specific differences were detected within sediment and root/rhizome microbial communities, but not the leaves. These results show that C. nodosa leaves have a consistent microbial community even across a pH range of 8.15 to 6.05. The ability for C. nodosa to regulate and maintain microbial structure may indicate a semblance of resilience within these vital ecosystems under projected changes in environmental conditions such as ocean acidification.

Contrasting heat stress response patterns of coral holobionts across the Red Sea suggest distinct mechanisms of thermal tolerance.

Corals from the northern Red Sea, in particular the Gulf of Aqaba (GoA), have exceptionally high bleaching thresholds approaching >5℃ above their maximum monthly mean (MMM) temperatures. These elevated thresholds are thought to be due to historical selection, as corals passed through the warmer Southern Red Sea during recolonization from the Arabian Sea. To test this hypothesis, we determined thermal tolerance thresholds of GoA versus central Red Sea (CRS) Stylophora pistillata corals using multi-temperature acute thermal stress assays to determine thermal thresholds. Relative thermal thresholds of GoA and CRS corals were indeed similar and exceptionally high (~7℃ above MMM). However, absolute thermal thresholds of CRS corals were on average 3℃ above those of GoA corals. To explore the molecular underpinnings, we determined gene expression and microbiome response of the coral holobiont. Transcriptomic responses differed markedly, with a strong response to the thermal stress in GoA corals and their symbiotic algae versus a remarkably muted response in CRS colonies. Concomitant to this, coral and algal genes showed temperature-induced expression in GoA corals, while exhibiting fixed high expression (front-loading) in CRS corals. Bacterial community composition of GoA corals changed dramatically under heat stress, whereas CRS corals displayed stable assemblages. We interpret the response of GoA corals as that of a resilient population approaching a tipping point in contrast to a pattern of consistently elevated thermal resistance in CRS corals that cannot further attune. Such response differences suggest distinct thermal tolerance mechanisms that may affect the response of coral populations to ocean warming.

Unravelling the Importance of Diazotrophy in Corals - Combined Assessment of Nitrogen Assimilation, Diazotrophic Community and Natural Stable Isotope Signatures.

There is an increasing interest in understanding the structure and function of the microbiota associated with marine and terrestrial organisms, because it can play a major role in host nutrition and resistance to environmental stress. Reef-building corals live in association with diazotrophs, which are microbes able to fix dinitrogen. Corals are known to assimilate diazotrophically-derived nitrogen (DDN), but it is still not clear whether this nitrogen source is derived from coral-associated diazotrophs and whether it substantially contributes to the coral's nitrogen budget. In this study, we aimed to provide a better understanding of the importance of DDN for corals using a holistic approach by simultaneously assessing DDN assimilation rates (using 15N2 tracer technique), the diazotrophic bacterial community (using nifH gene amplicon sequencing) and the natural δ15N signature in Stylophora pistillata corals from the Northern Red Sea along a depth gradient in winter and summer. Overall, our results show a discrepancy between the three parameters. DDN was assimilated by the coral holobiont during winter only, with an increased assimilation with depth. Assimilation rates were, however, not linked to the presence of coral-associated diazotrophs, suggesting that the presence of nifH genes does not necessarily imply functionality. It also suggests that DDN assimilation was independent from coral-associated diazotrophs and may instead result from nitrogen derived from planktonic diazotrophs. In addition, the δ15N signature presented negative values in almost all coral samples in both seasons, suggesting that nitrogen sources other than DDN contribute to the nitrogen budget of corals from this region. This study yields novel insight into the origin and importance of diazotrophy for scleractinian corals from the Northern Red Sea using multiple proxies.

Dissolved Nitrogen Acquisition in the Symbioses of Soft and Hard Corals With Symbiodiniaceae: A Key to Understanding Their Different Nutritional Strategies?

Nitrogen is one of the limiting nutrients for coral growth and primary productivity. Therefore, the capacity of different associations between corals and their algal symbionts (Symbiodiniaceae) to efficiently exploit the available nitrogen sources will influence their distribution and abundance. Recent studies have advanced our understanding of nitrogen assimilation in reef-building scleractinian (hard) coral-Symbiodiniaceae symbioses. However, the nutrient metabolism of other coral taxa, such as Alcyoniina (soft corals), remains underexplored. Using stable isotope labeling, we investigated the assimilation of dissolved nitrogen (i.e., ammonium, nitrate, and free amino acids) by multiple species of soft and hard corals sampled in the Gulf of Aqaba in shallow (8-10 m) and mesophotic (40-50 m) reefs. Our results show that dissolved nitrogen assimilation rates per tissue biomass were up to 10-fold higher in hard than in soft coral symbioses for all sources of nitrogen. Although such differences in assimilation rates could be linked to the Symbiodiniaceae density, Symbiodiniaceae species, or the C:N ratio of the host and algal symbiont fractions, none of these parameters were different between the two coral taxa. Instead, the lower assimilation rates in soft coral symbioses might be explained by their different nutritional strategy: whereas soft corals may obtain most of their nitrogen via the capture of planktonic prey by the coral host (heterotrophic feeding), hard corals may rely more on dissolved nitrogen assimilation by their algal symbionts to fulfill their needs. This study highlights different nutritional strategies in soft and hard coral symbioses. A higher reliance on heterotrophy may help soft corals to grow in reefs with higher turbidity, which have a high concentration of particles in suspension in seawater. Further, soft corals may benefit from lower dissolved nitrogen assimilation rates in areas with low water quality.

Fast and pervasive transcriptomic resilience and acclimation of extremely heat-tolerant coral holobionts from the northern Red Sea.

Corals from the northern Red Sea and Gulf of Aqaba exhibit extreme thermal tolerance. To examine the underlying gene expression dynamics, we exposed Stylophora pistillata from the Gulf of Aqaba to short-term (hours) and long-term (weeks) heat stress with peak seawater temperatures ranging from their maximum monthly mean of 27 °C (baseline) to 29.5 °C, 32 °C, and 34.5 °C. Corals were sampled at the end of the heat stress as well as after a recovery period at baseline temperature. Changes in coral host and symbiotic algal gene expression were determined via RNA-sequencing (RNA-Seq). Shifts in coral microbiome composition were detected by complementary DNA (cDNA)-based 16S ribosomal RNA (rRNA) gene sequencing. In all experiments up to 32 °C, RNA-Seq revealed fast and pervasive changes in gene expression, primarily in the coral host, followed by a return to baseline gene expression for the majority of coral (>94%) and algal (>71%) genes during recovery. At 34.5 °C, large differences in gene expression were observed with minimal recovery, high coral mortality, and a microbiome dominated by opportunistic bacteria (including Vibrio species), indicating that a lethal temperature threshold had been crossed. Our results show that the S. pistillata holobiont can mount a rapid and pervasive gene expression response contingent on the amplitude and duration of the thermal stress. We propose that the transcriptomic resilience and transcriptomic acclimation observed are key to the extraordinary thermal tolerance of this holobiont and, by inference, of other northern Red Sea coral holobionts, up to seawater temperatures of at least 32 °C, that is, 5 °C above their current maximum monthly mean.

Warming resistant corals from the Gulf of Aqaba live close to their cold-water bleaching threshold.

Global climate change is causing increasing variability and extremes in weather worldwide, a trend set to continue. In recent decades both anomalously warm and cold seawater temperatures have resulted in mass coral bleaching events. Whilst corals' response to elevated temperature has justifiably attracted substantial research interest, coral physiology under cold water stress is relatively unfamiliar. The response to below typical winter water temperature was tested for two common reef building species from the Gulf of Aqaba in an ex situ experiment. Stylophora pistillata and Acropora eurystoma were exposed to 1 or 3 °C below average winter temperature and a suite of physiological parameters were assessed. At 3 °C below winter minima (ca. 18.6 °C), both species had significant declines in photosynthetic indices (maximum quantum yield, electron transport rate, saturation irradiance, and photochemical efficiency) and chlorophyll concentration compared to corals at ambient winter temperatures. It was previously unknown that corals at this site live close to their cold-water bleaching threshold and may be vulnerable as climate variability increases in magnitude. In order to determine if a cold winter reduces the known heat resistance of this population, the corals were subsequently exposed to an acute warm period at 30 °C the following summer. Exposed to above typical summer temperatures, both species showed fewer physiological deviations compared to the cold-water stress. Therefore, the cold winter experience did not increase corals' susceptibility to above ambient summer temperatures. This study provides further support for the selection of heat tolerant genotypes colonising the Red Sea basin and thereby support the mechanism behind the Reef Refuge Hypothesis.

Simulated climate change scenarios impact the reproduction and early life stages of a soft coral.

Coral reefs are threatened worldwide by global climate change, manifested in anthropogenic ocean warming and acidification. Despite the importance of coral sexual reproduction for the continuity of coral reefs, our understanding of the extent of the impact of climate change on coral sexual reproduction, particularly on coral reproductive phenology and early life stages, is limited. Here, we experimentally examined the effects of predicted end-of-the-century seawater conditions on the sexual reproduction and photosynthetic capacity of a Red-Sea zooxanthellate octocoral, Rhytisma fulvum. Sexually mature colonies were exposed to ambient temperature and pH conditions and to Representative Concentration Pathway (RCP) conditions (4.5 and 8.5), five weeks prior to their expected surface-brooding event. The reproductive phenology of the colonies under the simulated seawater conditions was compared to that on the natural reef. In addition, subsequent planulae development and their metamorphosis into primary polyps under the same RCP conditions as their parent colonies were monitored in a running seawater system. The results reveal that both RCP conditions led to a change in the timing of onset of the surface-brooding event and its synchronicity. In contrast, the surface-brooding event under ambient conditions co-occurred with that of the in-situ reef colonies and maintained its synchrony. Similarly, planula survival and polyp metamorphosis rate were significantly reduced under both RCP conditions compared to propagules reared under ambient conditions. In addition, the photosynthetic capacity of the parent colonies under both RCPs showed a reduction relative to that under the ambient conditions in the experiment, suggesting a reduction in carbon fixation during the late stages of gametogenesis. While our findings indicate that octocoral reproductive phenology is affected by environmental changes, further work is required in order to elucidate the long-term implications for the R. fulvum population in the northern Red Sea.

Recovery assessment of the branching coral Stylophora pistillata following copper contamination and depuration.

Most contemporary coral reefs live under both global (e.g. warming and acidification) and local (e.g. overfishing, pollution) stressors, which may synergistically undermine their resilience to thermal bleaching and diseases. While heavy metal toxicity in reefs has been well characterized, information on corals recovery from acute contamination is lacking. We studied for 42 days the ability of the coral Stylophora pistillata from the Gulf of Aqaba (northern Red Sea) to recover from a short (3 days) and prolonged (14 days) copper (Cu) contamination (1 µg L-1), after 11 ('Exp3/D11') and 28 ('Exp14/D28') days of depuration, respectively. Cu caused a decrease in chlorophyll content after 3 days, and in net photosynthesis (Pn) after 14 and 42 days. 'Exp14/D28' showed successful recovery based on Pn and relative electron transport rate, as opposed to 'Exp3/D11'. Results suggest the depuration time may be of greater importance than the exposure period to recover from such contamination.

Seasonal Variations in the Culturable Mycobiome of Acropora loripes along a Depth Gradient.

Coral associated fungi are widespread, highly diverse and are part and parcel of the coral holobiont. To study how environmental conditions prevailing near the coral-host may affect fungal diversity, the culturable (isolated on potato dextrose agar) mycobiome associated with Acropora loripes colonies was seasonally sampled along a depth gradient in the Gulf of Aqaba. Fragments were sampled from both apparently healthy coral colonies as well as those exhibiting observable lesions. Based on phylogenetic analysis of 197 fungal sequences, Ascomycota were the most prevalent (91.9%). The abundance of fungi increased with increasing water depth, where corals sampled at 25 m yielded up to 70% more fungal colony forming units (CFUs) than those isolated at 6 m. Fungal diversity at 25 m was also markedly higher, with over 2-fold more fungal families represented. Diversity was also higher in lesioned coral samples, when compared to apparently healthy colonies. In winter, concurrent with water column mixing and increased levels of available nutrients, at the shallow depths, Saccharomytacea and Sporidiobolacea were more prevalent, while in spring and fall Trichocomacea (overall, the most prevalent family isolated throughout this study) were the most abundant taxa isolated at these depths as well as at deeper sampling sites. Our results highlight the dynamic nature of the culturable coral mycobiome and its sensitivity to environmental conditions and coral health.

Intracellular competition for nitrogen controls dinoflagellate population density in corals.

The density of dinoflagellate microalgae in the tissue of symbiotic corals is an important determinant for health and productivity of the coral animal. Yet, the specific mechanism for their regulation and the consequence for coral nutrition are insufficiently understood due to past methodological limitations to resolve the fine-scale metabolic consequences of fluctuating densities. Here, we characterized the physiological and nutritional consequences of symbiont density variations on the colony and tissue level in Stylophora pistillata from the Red Sea. Alterations in symbiont photophysiology maintained coral productivity and host nutrition across a broad range of symbiont densities. However, we demonstrate that density-dependent nutrient competition between individual symbiont cells, manifested as reduced nitrogen assimilation and cell biomass, probably creates the negative feedback mechanism for symbiont population growth that ultimately defines the steady-state density. Despite fundamental changes in symbiont nitrogen assimilation, we found no density-related metabolic optimum beyond which host nutrient assimilation or tissue biomass declined, indicating that host nutrient demand is sufficiently met across the typically observed range of symbiont densities under ambient conditions.

A method to disentangle and quantify host anabolic turnover in photosymbiotic holobionts with subcellular resolution.

A wide range of organisms host photosynthesizing symbionts. In these animals the metabolic exchange between host and symbionts has prevented in situ host anabolic turnover to be studied without the confounding effect of translocated photosynthates. Using the symbiotic coral Stylophora pistillata as a model organism and [1-13C]-pyruvate and [2,3-13C]-pyruvate in different incubation conditions (light, light + DCMU, and darkness), we employed NanoSIMS isotopic imaging to quantify host anabolism, with and without translocated metabolites from their photosynthesizing dinoflagellate symbionts. Under our experimental conditions, host de novo lipid synthesis accounted for ~40% of the total holobiont lipid reserve, and dinoflagellate recycling of metabolic 13CO2 enhanced host tissue 13C-enrichment by 13-22% in the epidermis, 40-58% in the gastrodermis, and 135-169% in host lipid bodies. Furthermore, we show that host anabolic turnover in different tissue structures differs, in a manner consistent with the localisation, function and cellular composition of these structures.

Feeding increases the number of offspring but decreases parental investment of Red Sea coral Stylophora pistillata.

Successful reproductive output and recruitment is crucial to coral persistence and recovery following anthropogenic stress. Feeding is known to alter coral physiology and increase resilience to bleaching.The goal of the study was to address the knowledge gap of the influence of feeding on reproductive output and offspring phenotype.Colonies of Stylophora pistillata from the Northern Gulf of Aqaba (Red Sea) were fed an Artemia diet or unfed for 5 months during gametogenesis, fertilization, and brooding. In addition, time to settlement and mortality of planulae were assessed at water temperatures ranging from winter temperature (22°C) to three degrees above average peak summer temperature (31°C). A range of physiological parameters was measured in parents and offspring.In brooding parents, feeding significantly increased protein concentration and more than tripled the number of released planulae. Planulae from unfed colonies had higher chlorophyll per symbiont concentration and concomitantly higher photosynthetic efficiency compared to planulae from fed parents. In settlement assays, planulae showed a similar thermal resistance as known for this Red Sea adult population. Mortality was greater in planulae from unfed parents at ambient and 3°C above ambient temperature despite higher per offspring investment in terms of total fatty acid content. Fatty acid profiles and relative abundances were generally conserved between different fed and unfed colonies but planulae were enriched in monounsaturated fatty acids relative to adults, that is, 16:1, 18:1, 20:1, 22:1, and 24:1 isomers.Ultimately the availability of zooplankton could influence population physiology and recruitment in corals.

Productivity and carbon fluxes depend on species and symbiont density in soft coral symbioses.

Soft corals often constitute one of the major benthic groups of coral reefs. Although they have been documented to outcompete reef-building corals following environmental disturbances, their physiological performance and thus their functional importance in reefs are still poorly understood. In particular, the acclimatization to depth of soft corals harboring dinoflagellate symbionts and the metabolic interactions between these two partners have received little attention. We performed stable isotope tracer experiments on two soft coral species (Litophyton sp. and Rhytisma fulvum fulvum) from shallow and upper mesophotic Red Sea coral reefs to quantify the acquisition and allocation of autotrophic carbon within the symbiotic association. Carbon acquisition and respiration measurements distinguish Litophyton sp. as mainly autotrophic and Rhytisma fulvum fulvum as rather heterotrophic species. In both species, carbon acquisition was constant at the two investigated depths. This is a major difference from scleractinian corals, whose carbon acquisition decreases with depth. In addition, carbon acquisition and photosynthate translocation to the host decreased with an increase in symbiont density, suggesting that nutrient provision to octocoral symbionts can quickly become a limiting factor of their productivity. These findings improve our understanding of the biology of soft corals at the organism-scale and further highlight the need to investigate how their nutrition will be affected under changing environmental conditions.

Ecological changes over 90 years at Low Isles on the Great Barrier Reef.

Coral reefs are under increasing stress from local and global factors. Long-term perspectives are becoming increasingly important for understanding ecosystem responses. Here, we provide insights from a 91-year study of the Low Isles on the northern Great Barrier Reef (GBR) that begins with the pioneering Great Barrier Reef Expedition (1928-29). We show that intertidal communities have experienced major phase-shifts since 1928, with few signs of a return to the initial state. Coral communities demolished by cyclones 50 years ago and exposed to multiple stressors have yet to recover. Richness and diversity of these communities systematically declined for corals and other invertebrates. Specifically, massive corals have replaced branching corals, and soft corals have become much more numerous. The long-term perspective of this study illustrates the importance of considering multiple factors in reef decline, and potential recovery, of coral reefs, and the importance of tracking changes in community structure as well as coral abundance over long periods.

Divergent Capacity of Scleractinian and Soft Corals to Assimilate and Transfer Diazotrophically Derived Nitrogen to the Reef Environment.

Corals are associated with dinitrogen (N2)-fixing bacteria that potentially represent an additional nitrogen (N) source for the coral holobiont in oligotrophic reef environments. Nevertheless, the few studies investigating the assimilation of diazotrophically derived nitrogen (DDN) by tropical corals are limited to a single scleractinian species (i.e., Stylophora pistillata). The present study quantified DDN assimilation rates in four scleractinian and three soft coral species from the shallow waters of the oligotrophic Northern Red Sea using the 15N2 tracer technique. All scleractinian species significantly stimulated N2 fixation in the coral-surrounding seawater (and mucus) and assimilated DDN into their tissue. Interestingly, N2 fixation was not detected in the tissue and surrounding seawater of soft corals, despite the fact that soft corals were able to take up DDN from a culture of free-living diazotrophs. Soft coral mucus likely represents an unfavorable habitat for the colonization and activity of diazotrophs as it contains a low amount of particulate organic matter, with a relatively high N content, compared to the mucus of scleractinian corals. In addition, it is known to present antimicrobial properties. Overall, this study suggests that DDN assimilation into coral tissues depends on the presence of active diazotrophs in the coral's mucus layer and/or surrounding seawater. Since N is often a limiting nutrient for primary productivity in oligotrophic reef waters, the divergent capacity of scleractinian and soft corals to promote N2 fixation may have implications for N availability and reef biogeochemistry in scleractinian versus soft coral-dominated reefs.

Impact of ocean acidification on crystallographic vital effect of the coral skeleton.

Distinguishing between environmental and species-specific physiological signals, recorded in coral skeletons, is one of the fundamental challenges in their reliable use as (paleo)climate proxies. To date, characteristic biological bias in skeleton-recorded environmental signatures (vital effect) was shown in shifts in geochemical signatures. Herein, for the first time, we have assessed crystallographic parameters of bio-aragonite to study the response of the reef-building coral Stylophora pistillata to experimental seawater acidification (pH 8.2, 7.6 and 7.3). Skeletons formed under high pCO2 conditions show systematic crystallographic changes such as better constrained crystal orientation and anisotropic distortions of bio-aragonite lattice parameters due to increased amount of intracrystalline organic matrix and water content. These variations in crystallographic features that seem to reflect physiological adjustments of biomineralizing organisms to environmental change, are herein called crystallographic vital effect (CVE). CVE may register those changes in the biomineralization process that may not yet be perceived at the macromorphological skeletal level.