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Coral reefs in the northern Red Sea experience strong seasonality. This affects reef carbon (C) cycling, but ecosystem-wide quantification of C fluxes in such reefs is limited. This study quantified seasonal reef community C fluxes with incubations. Resulting data were then incorporated into seasonal linear inverse models (LIM). For spring, additional sponge incubation results allowed for unique assessment of the contribution of sponges to C cycling. The coral reef ecosystem was heterotrophic throughout all seasons as gross community primary production (GPP; 136–200, range of seasonal means in mmol C m−2 d−1) was less than community respiration (R; 192–279), and balanced by import of organic carbon (52–100), 88‒92% of which being dissolved organic carbon (DOC). Hard coral GPP (74–110) and R (100–137), as well as pelagic bacteria DOC uptake (58–101) and R (42–86), were the largest C fluxes across seasons. The ecosystem was least heterotrophic in spring (highest irradiance) (GPP:R 0.81), but most heterotrophic in summer and fall with higher water temperatures (0.68 and 0.60, respectively). Adding the sponge community to the model increased community R (247 ± 8 without to 353 ± 13 with sponges (mean ± SD)). Sponges balanced this demand primarily with DOC uptake (105 ± 6, 97% by cryptic sponges). This rate is comparable to the uptake of DOC by pelagic bacteria (104 ± 5) placing the cryptic sponges among the dominant C cycling groups in the reef.

Mesophotic coral ecosystems (30–150 m depth) present a high oceanic biodiversity, but remain one of the most understudied reef habitats, especially below 60 m depth. Here, we have assessed the rates of photosynthesis and dissolved inorganic carbon (DIC) and nitrogen (DIN) assimilation by Symbiodiniaceae associated with four soft coral species of the genus Sinularia and two stony coral species of the genus Leptoseris collected respectively at 65 and 80–90 m depth in the Gulf of Eilat. Our study demonstrates that both Leptoseris and Sinularia species have limited autotrophic capacities at mid-lower mesophotic depths. DIC and DIN assimilation rates were overall ~ 10 times lower compared to shallow corals from 10 m depth in the same reef. While Leptoseris symbionts transferred at least 50% of the acquired nitrogen to their host after 8-h incubation, most of the nitrogen was retained in the symbionts of Sinularia. In addition, the host tissue of Sinularia species presented a very high structural carbon to nitrogen ratio (C : N) compared to Leptoseris or to the shallow coral species, suggesting nitrogen limitation in these mesophotic soft corals. The limited capacity of soft coral symbionts to acquire DIN and transfer it to the coral animal, as well as the high C : N ratios, might explain the scarcity of symbiotic soft corals at mid-lower mesophotic depths compared to their prevalence in the shallower reef. Overall, this study highlights the significance of DIN for the distribution of the Cnidarian- Symbiodiniaceae association at mesophotic depth.

Coral reefs are one of the most endangered habitats due to climate change, but not enough attention has been paid to how plastic pollution affects coral reef health. Plastics are massively produced worldwide for many purposes and they degrade very slowly, breaking down into tiny, invisible particles of 5 mm or less, called microplastics. When these tiny particles reach coral reefs, they harm corals by constantly rubbing on them through the action of waves and currents. Corals may also ingest microplastics and get a false sense of “fullness,” which results in the coral not feeding on nutritious food. Within the coral, microplastics may block the gut and cause internal damage. Also, microplastics, which are already made of chemicals, can pick up pollutants and harmful microorganisms from the seawater and transfer them to the coral. A reduction of microplastics pollution is therefore urgent.

Dinitrogen (N2 ) fixing bacteria (diazotrophs) are an important source of new nitrogen in oligotrophic environments and represent stable members of the microbiome in tropical corals, while information on corals from temperate oligotrophic regions is lacking. Therefore, this study provides new insights into the diversity and activity of diazotrophs associated with the temperate coral Oculina patagonica from the Mediterranean Sea by combining metabarcoding sequencing of amplicons of both the 16S rRNA and nifH genes and 15 N2 stable isotope tracer analysis to assess diazotroph-derived nitrogen (DDN) assimilation by the coral. Results show that the diazotrophic community of O. patagonica is dominated by autotrophic bacteria (i.e. Cyanobacteria and Chlorobia). The majority of DDN was assimilated into the tissue and skeletal matrix, and DDN assimilation significantly increased in bleached corals. Thus, diazotrophs may constitute an additional nitrogen source for the coral host, when nutrient exchange with Symbiodinium is disrupted (e.g. bleaching) and external food supply is limited (e.g. oligotrophic summer season). Furthermore, we hypothesize that DDN can facilitate the fast proliferation of endolithic algae, which provide an alternative carbon source for bleached O. patagonica. Overall, O. patagonica could serve as a good model for investigating the importance of diazotrophs in coral recovery from bleaching.

The tropical seagrass Halophila stipulacea exhibits a limited capacity to use nitrate, but has a high uptake capacity and efficiency for ammonium. Consequently, N2 fixation by associated diazotrophic epiphytes may be important in providing ammonium needed for seagrass photosynthesis and growth. When exposed to seasonality, this association could provide a competitive advantage for H. stipulacea in relation to other seagrass species, but related knowledge is scarce. Here, we thus report measurements of net photosynthesis (O2 fluxes) and N2 fixation (acetylene reduction) associated with H. stipulacea from the Northern Red Sea over all four seasons of 2013. In parallel, we characterized the seagrass meadow areal extent, shoot and leaves density, and the leaf area index. Using this data, we quantified meadow net primary production and N2 fixation and estimated the photo-metabolic N demand met by N2 fixation at the community level. Results revealed a marked seasonality of meadow N2 fixation, with rates ranging from 0.06 in winter to 4.60 mmol N m−2 d−1 in summer, i.e., an increase by two orders of magnitude. In summer, when nutrient concentrations in the water column were lowest and light intensity and temperature highest, there was a significant positive linear relationship between meadow net primary production and N2 fixation, with the latter accounting for ca. 20% of the photo-metabolic N demand. These findings suggest that N2 fixation by associated diazotrophic epiphytes can mitigate N limitation of H. stipulacea meadows under N scarcity. This trait may also favor the ongoing geographical expansion of H. stipulacea.

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.

Coral holobionts (i.e., coral-algal-prokaryote symbioses) exhibit dissimilar thermal sensitivities that may determine which coral species will adapt to global warming. Nonetheless, studies simultaneously investigating the effects of warming on all holobiont members are lacking. Here we show that exposure to increased temperature affects key physiological traits of all members (herein: animal host, zooxanthellae and diazotrophs) of both Stylophora pistillata and Acropora hemprichii during and after thermal stress. S. pistillata experienced severe loss of zooxanthellae (i.e., bleaching) with no net photosynthesis at the end of the experiment. Conversely, A. hemprichii was more resilient to thermal stress. Exposure to increased temperature (+ 6°C) resulted in a drastic increase in daylight dinitrogen (N2 ) fixation, particularly in A. hemprichii (threefold compared with controls). After the temperature was reduced again to in situ levels, diazotrophs exhibited a reversed diel pattern of activity, with increased N2 fixation rates recorded only in the dark, particularly in bleached S. pistillata (twofold compared to controls). Concurrently, both animal hosts, but particularly bleached S. pistillata, reduced both organic matter release and heterotrophic feeding on picoplankton. Our findings indicate that physiological plasticity by coral-associated diazotrophs may play an important role in determining the response of coral holobionts to ocean warming.

We investigated physiological responses including calcification, photosynthesis and alterations to polar metabolites, in the scleractinian coral Stylophora pistillata exposed to different concentrations of polyethylene microplastics. Results showed that at high plastic concentrations (50 particles/mL nominal concentration) the photosynthetic efficiency of photosystem II in the coral symbiont was affected after 4 weeks of exposure. Both moderate and high (5 and 50 particles/mL nominal) concentrations of microplastics caused subtle but significant alterations to metabolite profiles of coral, as determined by Nuclear Magnetic Resonance (NMR) spectroscopy. Specifically, exposed corals were found to have increased levels of phosphorylated sugars and pyrimidine nucleobases that make up nucleotides, scyllo-inositol and a region containing overlapping proline and glutamate signals, compared to control animals. Together with the photo-physiological stress response observed and previously published literature, these findings support the hypothesis that microplastics disrupt host-symbiont signaling and that corals respond to this interference by increasing signaling and chemical support to the symbiotic zooxanthellae algae. These findings are also consistent with increased mucus production in corals exposed to microplastics described in previous studies. Considering the importance of coral reefs to marine ecosystems and their sensitivity to anthropogenic stressors, more research is needed to elucidate coral response mechanisms to microplastics under realistic exposure conditions.

Marine plastic debris can act as a reservoir of chemical additives that can pose a potential threat to sensitive ecosystems such as coral reefs. A survey of foam macrodebris collected on beaches indeed revealed high concentrations of hexabromocyclododecanes (ΣHBCDD) in polystyrene (PS) samples (up to 1940 μg g−1). Results also showed that PS fragments can still leach over 150 ng g−1 d−1 of ΣHBCDD (primarily as the α-isomer) for relatively long durations, and that these additives are readily bioaccumulated and well-retained by corals. Despite significant HBCDD bioaccumulation in coral tissue, short-term exposure to HBCDD or PS leachate had no considerable effect on coral photosynthetic activity, symbiont concentration and chlorophyll content. Exposure to the PS leachate did however cause consistent polyp retraction in nubbins over the 5-day exposure. This response was not observed in animals exposed to HBCDD alone, suggesting that another constituent of the leachate stressed corals.

Labile dissolved organic carbon (DOC) is a major pollutant in coastal marine environments affected by anthropogenic impacts, and may significantly contribute to coral bleaching and subsequent mortality on coastal reefs. DOC can cause bleaching indirectly through the rapid proliferation of copiotrophic and pathogenic bacteria. Here we demonstrate that labile DOC compounds can also impair the coral-dinoflagellate symbiosis by directly affecting coral physiology on both the host and algal symbiont level. In a controlled aquarium experiment, we monitored over several weeks key physiological parameters of the tropical coral Stylophora pistillata exposed to ambient and elevated labile DOC levels (0.1 and 1.0 mM) in combination with low and high nitrogen (i.e. ammonium) conditions (0.2 and 4.0 μM). At the symbiont level, DOC exposure under low ammonium availability decreased the photosynthetic efficiency accompanied by ∼75 % Chl a and ∼50 % symbiont cell reduction. The photosynthetic functioning of the symbionts recovered once the DOC enrichment ceased indicating a reversible shift between autotrophic and heterotrophic metabolism. At the host level, the assimilation of exogenous DOC sustained the tissue carbon reserves, but induced a depletion of the nitrogen reserves, indicated by ∼35 % decreased protein levels. This suggests an imbalanced exogenous carbon to nitrogen supply with nitrogen potentially limiting host metabolism on the long-term. We also demonstrate that increased ammonium availability delayed DOC-induced bleaching likely by keeping symbionts in a photosynthetically competent state, which is crucial for symbiosis maintenance and coral survival. Overall, the present study provides further insights into how coastal pollution can de-stabilize the coral-algal symbiosis and cause coral bleaching. Therefore, reducing coastal pollution and sustaining ecological integrity are critical to strengthen the resilience of coral reefs facing climate change.

N-2 fixation by coral reef benthic substrates may support primary productivity on oligo trophic coral reefs. However, little is known regarding the influence of environmental para meters on coral reef benthic N-2 fixation. This study quantified N-2 fixation and photosynthesis in 3 common reef framework substrates: turf algae, coral rock, and the abundant encrusting sponge Mycale fistulifera over 4 seasons in the northern Gulf of Aqaba. N-2 fixation activity was detected during day and night for all substrates, but on an annual average was significantly higher for turf algae (4.4 +/- 3.9 nmol C2H4 cm(-2) h(-1)) and coral rock (3.5 +/- 2.8 nmol C2H4 cm(-2) h(-1)) compared to M. fistulifera (0.2 +/- 0.2 nmol C2H4 cm(-2) h(-1)). There was strong seasonality in N-2 fixation, with rates one order of magnitude higher in summer when temperature and irradiance were highest but inorganic nutrient concentrations lowest. During summer and fall, when nutrients were low, we found a significant positive linear relationship between gross photosynthesis (P-gross) and N-2 fixation in turf algae and coral rock. Further, we estimate N-2 fixation can supply up to 20 and 27% of the N demand for net photosynthesis (P-net) in coral rock and turf algae, respectively. By contrast there was no significant relationship between N-2 fixation and Pgross in M. fistulifera, which displayed negative Pnet and heterotrophic metabolism (P-gross: respiration

International audience; Coral reefs are highly productive ecosystems thriving in nutrient-poor waters. Their productivity depends largely on the availability of nitrogen, the proximate limiting nutrient for primary production. In reefs, the major nitrogen pathways include regeneration, nitrification, ammonification and dinitrogen (N 2) fixation. N 2 fixation is performed by prokaryotes called " diazotrophs " that abound in coral rubbles, sandy bottoms, microbial mats, or seagrass meadows. Corals, which are the main reef builders, have developed a partnership with dinoflagellates which transform dissolved inorganic nitrogen into amino acids and protein, and with diazotrophs to gain diazotrophically-derived nitrogen (DDN). Pioneering studies found active diazotrophic cyanobacteria in the corals' mucus and/or tissue, and later high throughput sequencing efforts have described diverse communities of non-cyanobacterial diazotrophs associated with scleractinian corals. Yet, the metabolic processes behind these associations and how they benefit corals are currently not well understood. While genomic studies describe the diversity of diazotrophs and isotopic tracer experiments quantify N 2 fixation rates, combined advanced methods are needed to elucidate the mechanisms behind the transfer of DDN to the coral holobiont and whether these mechanisms change according to the identity of the diazotrophs or coral species. Here we review our current knowledge on N 2 fixation in corals: the diversity and localization of diazotrophs in the coral holobiont, the environmental factors controlling N 2 fixation, the fate of DDN within the coral symbiosis as well as its potential role in coral resilience. We finally summarize the unknowns: are the diversity, abundance and localization of diazotrophs within the holobiont species-and/or site-specific? Do they have an impact on DDN production? What are the metabolic mechanisms implicated? Do they change spatially, temporally or according to environmental factors? We encourage scientists to undertake research efforts to tackle these questions in order to shed light on nitrogen cycling in reef ecosystems and to understand if coral-associated N 2 fixation can improve coral's resilience in the face of climate change.

Northern Red Sea coral reefs experience pronounced seasonal variations in environmental factors such as water temperature, light intensity, and nutrient availability. This allows studying related effects on primary production by different functional groups. The present study therefore quantified primary production of all dominant benthic primary producers from a Jordanian fringing reef (29° 27′ 31″ N, 34° 58′ 26″ E) by measuring net photosynthesis (P n) and dark respiration (R) using stirred respirometry chamber incubations during all four seasons of 2013. Annual mean P n was highest for the macroalga Caulerpa (901 nmol O2 cm−2 h−1) and lowest for both the soft coral Sarcophyton and sedimentary microphytobenthos (212 and 223 nmol O2 cm−2 h−1, respectively). Sedimentary microphytobenthos exhibited the strongest response to seasonality with 5.7 times higher P n in spring than in winter. R was highest in hard corals among all groups in every season, likely due to nighttime calcification and heterotrophic activity. Gross photosynthesis-to-respiration ratios (P g:R) were highest for turf algae and macroalgae as well as cyanobacterial mats. While R was primarily positively related to PAR and temperature and P g:R to inorganic nutrients, few groups revealed significant relations between P n and environmental parameters. Seasonal budgets found daily gross primary production and respiration to be dominated by hard and soft corals. Total reef gross primary production was comparable between less oligotrophic spring and more oligotrophic summer. This indicates that processes such as heterotrophic feeding and microbial dinitrogen fixation may help the functional groups overcome potential nutrient limitation of primary production in summer.

Biological dinitrogen (N2) fixation (diazotrophy, BNF) relieves marine primary producers of nitrogen (N) limitation in a large part of the world oceans. N concentrations are particularly low in tropical regions where coral reefs are located, and N is therefore a key limiting nutrient for these productive ecosystems. In this context, the importance of diazotrophy for reef productivity is still not resolved, with studies up to now lacking organismal and seasonal resolution. Here, we present a budget of gross primary production (GPP) and BNF for a highly seasonal Red Sea fringing reef, based on ecophysiological and benthic cover measurements combined with geospatial analyses. Benthic GPP varied from 215 to 262 mmol C m−2 reef d−1, with hard corals making the largest contribution (41–76%). Diazotrophy was omnipresent in space and time, and benthic BNF varied from 0.16 to 0.92 mmol N m−2 reef d−1. Planktonic GPP and BNF rates were respectively approximately 60- and 20-fold lower than those of the benthos, emphasizing the importance of the benthic compartment in reef biogeochemical cycling. BNF showed higher sensitivity to seasonality than GPP, implying greater climatic control on reef BNF. Up to about 20% of net reef primary production could be supported by BNF during summer, suggesting a strong biogeochemical coupling between diazotrophy and the reef carbon cycle.

Coral reef ecosystems fringing the coastline of Dahab (South Sinai, Egypt) have experienced increasing anthropogenic disturbance as an emergent international tourism destination. Previous reports covering tourism-related impacts on coastal environments, particularly mechanical damage and destructive fishing, have highlighted the vital necessity for regular ecosystem monitoring of coral reefs near Dahab. However, a continuous scientific monitoring programme of permanent survey sites has not been established to date. Thus, this study conducted in situ monitoring surveys to investigate spatio-temporal variability of benthic reef communities and selected reef-associated herbivores along with reef health indicator organisms by revisiting three of the locally most frequented dive sites during expeditions in March 2010, September 2011 and February 2013. In addition, inorganic nutrient concentrations in reef-surrounding waters were determined to evaluate bottom-up effects of key environmental parameters on benthic reef community shifts in relation to grazer-induced top-down control. Findings revealed that from 2010 to 2013, live hard coral cover declined significantly by 12 % at the current-sheltered site Three Pools (TP), while showing negative trends for the Blue Hole (BH) and Lighthouse (LH) sites. Hard coral cover decline was significantly and highly correlated to a substantial increase in turf algae cover (up to 57 % at TP) at all sites, replacing hard corals as dominant benthic space occupiers in 2013. These changes were correlated to ambient phosphate and ammonium concentrations that exhibited highest values (0.64 ± 0.07 μmol PO4 3− l−1, 1.05 ± 0.07 μmol NH4 + l−1) at the degraded site TP. While macroalgae appeared to respond to both bottom-up and top-down factors, change in turf algae was consistent with expected indications for bottom-up control. Temporal variability measured in herbivorous reef fish stocks reflected seasonal impacts by local fisheries, with concomitant changes in macroalgal cover. These findings represent the first record of rapid, localised change in benthic reef communities near Dahab, consistent with indications for bottom-up controlled early-stage phase shifts, underlining the necessity for efficient regional wastewater management for coastal facilities.

Tropical coral reefs are among the most productive and diverse ecosystems, despite being surrounded by ocean waters where nutrients are in short supply. Benthic dinitrogen (N2) fixation is a significant internal source of “new” nitrogen (N) in reef ecosystems, but related information appears to be sparse. Here, we review the current state (and gaps) of knowledge on N2 fixation associated with coral reef organisms and their ecosystems. By summarizing the existing literature, we show that benthic N2 fixation is an omnipresent process in tropical reef environments. Highest N2 fixation rates are detected in reef-associated cyanobacterial mats and sea grass meadows, clearly showing the significance of these functional groups, if present, to the input of new N in reef ecosystems. Nonetheless, key benthic organisms such as hard corals also importantly contribute to benthic N2 fixation in the reef. Given the usually high coral coverage of healthy reef systems, these results indicate that benthic symbiotic associations may be more important than previously thought. In fact, mutualisms between carbon (C) and N2 fixers have likely evolved that may enable reef communities to mitigate N limitation. We then explore the potential effects of the increasing human interferences on the process of benthic reef N2 fixation via changes in diazotrophic populations, enzymatic activities, or availability of benthic substrates favorable to these microorganisms. Current knowledge indicates positive effects of ocean acidification, warming, and deoxygenation and negative effects of increased ultraviolet radiation on the amount of N fixed in coral reefs. Eutrophication may either boost or suppress N2 fixation, depending on the nutrient becoming limiting. As N2 fixation appears to play a fundamental role in nutrient-limited reef ecosystems, these assumptions need to be expanded and confirmed by future research efforts addressing the knowledge gaps identified in this review.

Since productivity and growth of coral-associated dinoflagellate algae is nitrogen (N)-limited, dinitrogen (N2) fixation by coral-associated microbes is likely crucial for maintaining the coraldinoflagellate symbiosis. It is thus essential to understand the effects future climate change will have on N2 fixation by the coral holobiont. This laboratory study is the first to investi- gate short-term effects of ocean acidification on N2 fixation activity associated with the tropical, hermatypic coral Seriatopora hystrix using the acetylene reduction assay in combination with cal- cification measurements. Findings reveal that simulated ocean acidification (p CO2 1080 µatm) caused a rapid and significant decrease (53%) in N2 fixation rates associated with S. hystrix com- pared to the present day scenario (p CO2 486 µatm). In addition, N2 fixation associated with the coral holobiont showed a positive exponential relationship with its calcification rates. This sug- gests that even small declines in calcification rates of hermatypic corals under high CO2 conditions may result in decreased N2 fixation activity, since these 2 processes may compete for energy in the coral holobiont. Ultimately, an intensified N limitation in combination with a decline in skeletal growth may trigger a negative feedback loop on coral productivity exacerbating the negative long-term effects of ocean acidification.

Summary The release of organic matter (OM) by scleractinian corals represents a key physiological process that importantly contributes to coral reef ecosystem functioning, and that is affected by inorganic nutrient availability. While OM fluxes have been studied for several dominant reef taxa, no information is available for soft corals, one of the major benthic groups in tropical reef environments. Thus, this study investigates OM fluxes along with other key physiological parameters (i.e. photosynthesis, respiration and chlorophyll a tissue content) in the common soft coral genus Xenia after a 4-week exposure period to elevated ammonium (N, 20.0 µM), phosphate (P, 2.0 µM) and combined inorganic nutrient enrichment treatment (NP). Corals maintained without nutrient enrichment served as non-treated controls and revealed constant uptake rates for particulate organic carbon (POC) (-0.32 ± 0.16 mg POC m-2 coral surface area h-1), particulate nitrogen (PN) (-0.05 ± 0.02 mg PN m-2 h-1) and dissolved organic carbon (DOC) (-4.8 ± 2.1 mg DOC m-2 h-1). While DOC uptake significantly increased in the N treatment, POC flux was not affected. The P treatment significantly enhanced PN release as well as photosynthesis and respiration rates suggesting an influence by autotrophic carbon acquisition of zooxanthellae endosymbionts on OM fluxes by the coral host. Our physiological findings confirm the significant effect of inorganic nutrient availability on OM fluxes and key metabolic processes for the soft coral Xenia, and provide first clues on OM cycles initiated by soft corals in reef environments exposed to ambient and elevated inorganic nutrient concentrations.

Functional traits define species by their ecological role in the ecosystem. Animals themselves are host–microbe ecosystems (holobionts), and the application of ecophysiological approaches can help to understand their functioning. In hard coral holobionts, communities of dinitrogen (N 2 )-fixing prokaryotes (diazotrophs) may contribute a functional trait by providing bioavailable nitrogen (N) that could sustain coral productivity under oligotrophic conditions. This study quantified N 2 fixation by diazotrophs associated with four genera of hermatypic corals on a northern Red Sea fringing reef exposed to high seasonality. We found N 2 fixation activity to be 5- to 10-fold higher in summer, when inorganic nutrient concentrations were lowest and water temperature and light availability highest. Concurrently, coral gross primary productivity remained stable despite lower Symbiodinium densities and tissue chlorophyll a contents. In contrast, chlorophyll a content per Symbiodinium cell increased from spring to summer, suggesting that algal cells overcame limitation of N, an essential element for chlorophyll synthesis. In fact, N 2 fixation was positively correlated with coral productivity in summer, when its contribution was estimated to meet 11% of the Symbiodinium N requirements. These results provide evidence of an important functional role of diazotrophs in sustaining coral productivity when alternative external N sources are scarce.