Denitrification may potentially alleviate excess nitrogen (N) availability in coral holobionts to maintain a favourable N to phosphorous ratio in the coral tissue. However, little is known about the abundance and activity of denitrifiers in the coral holobiont. The present study used the nirS marker gene as a proxy for denitrification potential along with measurements of denitrification rates in a comparative coral taxonomic framework from the Red Sea: Acropora hemprichii, Millepora dichotoma, and Pleuractis granulosa. Relative nirS gene copy numbers associated with the tissues of these common corals were assessed and compared with denitrification rates on the holobiont level. In addition, dinitrogen (N2) fixation rates, Symbiodiniaceae cell density, and oxygen evolution were assessed to provide an environmental context for denitrification. We found that relative abundances of the nirS gene were 16- and 17-fold higher in A. hemprichii compared to M. dichotoma and P. granulosa, respectively. In concordance, highest denitrification rates were measured in A. hemprichii, followed by M. dichotoma and P. granulosa. Denitrification rates were positively correlated with N2 fixation rates and Symbiodiniaceae cell densities. Our results suggest that denitrification may counterbalance the N input from N2 fixation in the coral holobiont, and we hypothesize that these processes may be limited by photosynthates released by the Symbiodiniaceae.

Coral reefs in the wider Caribbean declined in hard coral cover by ~80% since the 1970s, but spatiotemporal analyses for sub-regions are lacking. Here, we explored benthic change patterns in the Mexican Caribbean reefs through meta-analysis between 1978 and 2016 including 125 coral reef sites. Findings revealed that hard coral cover decreased from ~26% in the 1970s to 16% in 2016, whereas macroalgae cover increased to ~30% in 2016. Both groups showed high spatiotemporal variability. Hard coral cover declined in total by 12% from 1978 to 2004 but increased again by 5% between 2005 and 2016 indicating some coral recovery after the 2005 mass bleaching event and hurricane impacts. In 2016, more than 80% of studied reefs were dominated by macroalgae, while only 15% were dominated by hard corals. This stands in contrast to 1978 when all reef sites surveyed were dominated by hard corals. This study is among the first within the Caribbean region that reports local recovery in coral cover in the caribbean, while other caribbean reefs have failed to recover. Most Mexican caribbean coral reefs are now no longer dominated by hard corals. in order to prevent further reef degradation, viable and reliable conservation alternatives are required. Monitoring change in coral reef ecosystems is essential in an era when humanity is having a widespread and long-term impact on nature. Current anthropogenic climate change and local stressors (such as overfishing and a mix of pollution and sedimentation from coastal development 1) place coral reefs as the most endangered ecosystems on earth 2. Rapid reversals in their health have been reported globally 3 , including reefs from the Caribbean region, where declines of the live hard coral cover of ~80% between 1975 and 2000 have been documented 4-6. In the late 1970s, entire populations of reef-building coral species (i.e. Acropora palmata and Acropora cervicornis) collapsed as a result of the white-band disease 7. Furthermore, the mass mortality of black sea urchins (Diadema antillarum), overfishing and eutrophication 8 have resulted in a proliferation of more opportunistic, fast-growing organisms such as (macro)algae that outcompete reef-building corals 8-11. As a result, many Caribbean benthic coral reef communities changed drastically from low coral cover to persistent states of high cover (macro)algae in the process of so-called phase shifts 11-15. Efforts to mitigate or reverse phase shifts and reef degradation in the Caribbean include the development of new coral reef monitoring and managing strategies 16-18. Monitoring efforts of Caribbean reefs began in the late 1970s at various reef locations for short durations 19. It was until 1980 when coral reef monitoring programs first began for some countries due to the evident reef degradation and increasing threats 19. In the Mesoamerican Reef System (MAR), the monitoring officially began in 2005 with the Healthy Reefs for Healthy People Initiative 20. The MAR is recognized by the World Wildlife Fund (WWF) as one of 200 global priority ec...

Eutrophication (i.e. the increase of [in-]organic nutrients) may affect the functioning of coral reefs, but knowledge about the effects on nitrogen (N) cycling and its relationship to productivity within benthic reef communities is scarce. Thus, we investigated how in situ manipulated eutrophication impacted productivity along with 2 counteracting N-cycling pathways (dinitrogen [N2]-fixation, denitrification), using a combined acetylene assay. We hypothesised that N2-fixation would decrease and denitrification increase in response to eutrophication. N fluxes and productivity (measured as dark and light oxygen fluxes assessed in incubation experiments) were determined for 3 dominant coral reef functional groups (reef sediments, turf algae, and the scleractinian coral Pocillopora verrucosa) after 8 wk of in situ nutrient enrichment in the central Red Sea. Using slow-release fertiliser, we increased the dissolved inorganic N concentration by up to 7-fold compared to ambient concentrations. Experimental nutrient enrichment stimulated both N2-fixation and denitrification across all functional groups 2- to 7-fold and 2- to 4-fold, respectively. Productivity doubled in reef sediments and remained stable for turf algae and P. verrucosa. Our data therefore suggest that (1) turf algae are major N2-fixers in coral reefs, while denitrification is widespread among all investigated groups; (2) surprisingly, and contrary to our hypothesis, both N2-fixation and denitrification are involved in the response to moderate N eutrophication, and (3) stimulated N2-fixation and denitrification are not directly influenced by productivity. Our findings underline the importance and ubiquity of microbial N cycling in (Red Sea) coral reefs along with its sensitivity to eutrophication.

While various sources increasingly release nutrients to the Red Sea, knowledge about their effects on benthic coral reef communities is scarce. Here, we provide the first comparative assessment of the response of all major benthic groups (hard and soft corals, turf algae and reef sands—together accounting for 80% of the benthic reef community) to in-situ eutrophication in a central Red Sea coral reef. For 8 weeks, dissolved inorganic nitrogen (DIN) concentrations were experimentally increased 3-fold above environmental background concentrations around natural benthic reef communities using a slow release fertilizer with 15% total nitrogen (N) content. We investigated which major functional groups took up the available N, and how this changed organic carbon (Corg) and N contents using elemental and stable isotope measurements. Findings revealed that hard corals (in their tissue), soft corals and turf algae incorporated fertilizer N as indicated by significant increases in δ15N by 8%, 27% and 28%, respectively. Among the investigated groups, Corg content significantly increased in sediments (+24%) and in turf algae (+33%). Altogether, this suggests that among the benthic organisms only turf algae were limited by N availability and thus benefited most from N addition. Thereby, based on higher Corg content, turf algae potentially gained competitive advantage over, for example, hard corals. Local management should, thus, particularly address DIN eutrophication by coastal development and consider the role of turf algae as potential bioindicator for eutrophication.

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The resistance of hard corals to warming can be negatively affected by nitrate eutrophication, but related knowledge for soft corals is scarce. We thus investigated the ecophysiological response of the pulsating soft coral Xenia umbellata to different levels of nitrate eutrophication (control = 0.6, medium = 6, high = 37 μM nitrate) in a laboratory experiment, with additional warming (27.7 to 32.8 °C) from days 17 to 37. High nitrate eutrophication enhanced cellular chlorophyll a content of Symbiodiniaceae by 168%, while it reduced gross photosynthesis by 56%. After additional warming, polyp pulsation rate was reduced by 100% in both nitrate eutrophication treatments, and additional polyp loss of 7% d−1 and total fragment mortality of 26% was observed in the high nitrate eutrophication treatment. Warming alone did not affect any of the investigated response parameters. These results suggest that X. umbellata exhibits resistance to warming, which may facilitate ecological dominance over some hard corals as ocean temperatures warm, though a clear negative physiological response occurs when combined with nitrate eutrophication. This study thus confirms the importance of investigating combinations of global and local factors to understand and manage changing coral reefs.

The northern Red Sea experiences strong annual differences in environmental conditions due to its relative high-latitude location for coral reefs. This allows the study of regulatory effects by key environmental parameters (i.e., temperature, inorganic nutrient, and organic matter concentrations) on reef primary production and dinitrogen (N2) fixation, but related knowledge is scarce. Therefore, this study measured environmental parameters, primary production and N2 fixation of phytoplankton groups in the water overlying a coral reef in the Gulf of Aqaba. To this end, we used a comparative approach between mixed and stratified water column scenarios in a full year of seasonal observations. Findings revealed that inorganic nutrient concentrations were significantly higher in the mixed compared to the stratified period. While gross photosynthesis and N2 fixation rates remained similar, net photosynthesis decreased from mixed to stratified period. Net heterotrophic activity of the planktonic community increased significantly during the stratified compared to the mixed period. While inorganic nitrogen (N) availability was correlated with net photosynthesis over the year, N2 fixation only correlated with N availability during the mixed period. This emphasizes the complexity of planktonic trophodynamics in northern Red Sea coral reefs. Comparing mixed and stratified planktonic N2 fixation rates with those of benthic organisms and substrates revealed a close seasonal activity similarity between free-living pelagic and benthic diazotrophs. During the mixed period, N2 fixation potentially contributed up to 3% of planktonic primary production N demand. This contribution increased by ca. one order of magnitude to 21% during the stratified period. Planktonic N2 fixation is likely a significant N source for phytoplankton to maintain high photosynthesis under oligotrophic conditions in coral reefs, especially during stratified conditions.

Recent research suggests that prior exposure of several months to elevated irradiance induces enhanced thermal tolerance in scleractinian corals. While this tolerance has been reported at the species level, individual coral colonies may react differently due to individual variability in thermal tolerance. As thermal anomalies are predicted to become common in the upcoming future, intraspecific variation may be key to the survival of coral populations. In order to study light-history based thermal stress responses on individual colonies, we developed a preliminary microcosm experiment where three randomly chosen, aquacultured colonies of the model coral Stylophora pistillata were exposed to two irradiance treatments (200 and 400 μmol photons m−2 s−1) for 31 days, followed by artificially induced heat stress (∼33.4 °C). We found different responses to occur at both the intraspecific and the intracolonial levels, as indicated by either equal, less severe, delayed, and/or even non-necrotic responses of corals previously exposed to the irradiance of 400 compared to 200 μmol photons m−2 s−1. In addition, all individual colonies revealed light-enhanced calcification. Finally, elevated irradiance resulted in a lower chlorophyll a concentration in one colony compared to the control treatment, and the same colony displayed more rapid bleaching compared to the other ones. Taken together, this study highlights the potential importance of intra-individual variability in physiological responses of scleractinian corals and provides recommendations for improving methodological designs for future studies.