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With its geographically isolated location and geological history, the Mediterranean Sea harbors well-known biodiversity hotspots, such as Posidonia oceanica seagrass meadows. Recently, long-living mats formed by the fleshy red alga Phyllophora crispa have been described to be associated with a high diversity of sessile invertebrates in the Tyrrhenian Sea. One of the key taxa among these sessile invertebrates are bryozoans: their abundance, diversity, and spatial distribution in P. crispa mats represent a gap in scientific knowledge. Thus, we conducted a pilot study on bryozoan assemblages associated with P. crispa mats around Giglio Island (Tuscan Archipelago, Italy) in 2018, followed by a comparative study on four sites distributed around the island in the subsequent year, 2019. We compared these findings to bryozoan abundance and diversity on P. oceanica shoots and leaves during the second expedition. The findings revealed more than 46 families, with a significantly higher number of taxa identified in P. crispa mats (33) than in P. oceanica meadows (29). The Shannon diversity index was similar between P. crispa and P. oceanica shoots, while Pielou’s evenness index was lower in P. crispa mats. The most abundant families reported across all habitats were Crisiidae, Aetidae, and Lichenoporidae; but the most abundant family on P. crispa was Chlidoniidae (Chlidonia pyriformis). The assemblages associated with P. crispa differed among sites, with higher abundances but lower diversity on the exposed southernmost site. The total bryozoan abundance was significantly higher on P. crispa (average 2.83 × 106 ± 1.99 × 106 colonies per m2 seafloor) compared to P. oceanica meadows (average 0.54 × 106 ± 0.34 × 106 colonies per m2 seafloor). Our results show a high diversity of bryozoans on P. crispa thalli compared to P. oceanica meadows, which was consistent throughout the study. These findings confirm the value of the red alga-generated habitat for associated bryozoans and may have implications for future biodiversity assessments and conservation measures.

Biological imaging is an essential tool to visualise and obtain reference data. In this context, the programme ImageJ has been widely used in many disciplines to determine the surface areas of planar biological samples in marine and aquatic experimental biology. Despite its range of advantages, ImageJ is relatively time-consuming, because of the need to manually select the target areas for quantification. Hence, we here evaluated the freeware programme Photopea as a potential alternative by comparing the accuracy and time required for the surface area quantification of exemplary algae compared with established ImageJ analysis. Our results show that Photopea is equally accurate as ImageJ, but 45% more time efficient. This time efficiency originates from using colour contrast that reduces the time needed to analyse each picture. Photopea thus offers an accurate, rapid, and cost-free tool to easily obtain reference data from field and laboratory experiments. This tool is particularly useful for experiments with an extensive sample size of specimens and thus can increase the power of study results.

Coral reefs are threatened by global and local stressors. Yet, reefs appear to respond differently to different environmental stressors. Using a global dataset of coral reef occurrence as a proxy for the long-term adaptation of corals to environmental conditions in combination with global environmental data, we show here how global (warming: sea surface temperature; acidification: aragonite saturation state, Ωarag ) and local (eutrophication: nitrate concentration, and phosphate concentration) stressors influence coral reef habitat suitability. We analyse the relative distance of coral communities to their regional environmental optima. In addition, we calculate the expected change of coral reef habitat suitability across the tropics in relation to an increase of 0.1°C in temperature, an increase of 0.02 μmol/L in nitrate, an increase of 0.01 μmol/L in phosphate and a decrease of 0.04 in Ωarag . Our findings reveal that only 6% of the reefs worldwide will be unaffected by local and global stressors and can thus act as temporary refugia. Local stressors, driven by nutrient increase, will affect 22% of the reefs worldwide, whereas global stressors will affect 11% of these reefs. The remaining 61% of the reefs will be simultaneously affected by local and global stressors. Appropriate wastewater treatments can mitigate local eutrophication and could increase areas of temporary refugia to 28%, allowing us to 'buy time', while international agreements are found to abate global stressors.

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.

Mutualistic nutrient cycling in the coral-algae symbiosis depends on limited nitrogen (N) availability for algal symbionts. Denitrifying prokaryotes capable of reducing nitrate or nitrite to dinitrogen could thus support coral holobiont functioning by limiting N availability. Octocorals show some of the highest denitrification rates among reef organisms, however little is known about the community structures of associated denitrifiers and their response to environmental fluctuations. Combining 16S rRNA gene amplicon sequencing with nirS in-silico PCR and quantitative PCR, we found differences in bacterial community dynamics between two octocorals exposed to excess dissolved organic carbon (DOC) and concomitant warming. While bacterial communities of the gorgonian Pinnigorgia flava remained largely unaffected by DOC and warming, the soft coral Xenia umbellata exhibited a pronounced shift towards Alphaproteobacteria dominance under excess DOC. Likewise, the relative abundance of denitrifiers was not altered in P. flava, but decreased by one order of magnitude in X. umbellata under excess DOC likely due to decreased proportions of Ruegeria spp. Given that holobiont C:N ratios remained stable in P. flava but showed a pronounced increase with excess DOC in X. umbellata host, our results suggest that microbial community dynamics may reflect the nutritional status of the holobiont. Hence, denitrifier abundance may be directly linked to N availability. This suggests a passive regulation of N cycling microbes, which could help stabilize nutrient limitation in the coral-algal symbiosis and thereby support holobiont functioning in a changing environment. Importance Octocorals are important members of reef-associated benthic communities that can rapidly replace scleractinian corals as the dominant ecosystem engineers on degraded reefs. Considering the substantial change in the (a)biotic environment that is commonly driving reef degradation, maintaining a dynamic and metabolically diverse microbial community might contribute to octocoral acclimatization and ecological adaptation. Nitrogen (N) cycling microbes, in particular denitrifying prokaryotes, may support holobiont functioning by limiting internal N availability, but little is known about the identity and (a)biotic drivers of octocoral-associated denitrifiers. Here, we show contrasting dynamics of bacterial communities associated with two common octocoral species, the soft coral Xenia umbellata and the gorgonian Pinnigorgia flava after a six-week exposure to excess dissolved organic carbon (DOC) under concomitant warming conditions. The specific responses of denitrifier communities associated with the two octocoral species aligned with the nutritional status of holobiont members. This suggests a passive regulation of this microbial trait based on N availability in the coral holobiont.

Efficient nutrient cycling in the coral-algal symbiosis requires constant but limited nitrogen availability. Coral-associated diazotrophs, i.e., prokaryotes capable of fixing dinitrogen, may thus support productivity in a stable coral-algal symbiosis but could contribute to its breakdown when overstimulated. However, the effects of environmental conditions on diazotroph communities and their interaction with other members of the coral holobiont remain poorly understood. Here we assessed the effects of heat stress on diazotroph diversity and their contribution to holobiont nutrient cycling in the reef-building coral Stylophora pistillata from the central Red Sea. In a stable symbiotic state, we found that nitrogen fixation by coral-associated diazotrophs constitutes a source of nitrogen to the algal symbionts. Heat stress caused an increase in nitrogen fixation concomitant with a change in diazotroph communities. Yet, this additional fixed nitrogen was not assimilated by the coral tissue or the algal symbionts. We conclude that although diazotrophs may support coral holobiont functioning under low nitrogen availability, altered nutrient cycling during heat stress abates the dependence of the coral host and its algal symbionts on diazotroph-derived nitrogen. Consequently, the role of nitrogen fixation in the coral holobiont is strongly dependent on its nutritional status and varies dynamically with environmental conditions.

Background Many coral reefs worldwide are experiencing declines in hard corals, resulting in other benthic organisms, e.g., soft corals, becoming more dominant. As such, more studies on the ecophysiology of soft corals are needed. Despite many methods for asexual reproduction of hard corals, effective methods for soft corals, i.e., without a hard skeleton, are scarce. This study, thus, assessed four fragmentation methods, the glue, rubber band, tunnel mesh, and plug mesh method for the pulsating soft coral Xenia umbellata that is widely distributed in the tropical Indo-Pacific. Methods Methods were comparatively assessed by determining the required time and labor for the fragmentation plus the health status of the fragmented corals by measuring their oxygen fluxes and pulsation rates, i.e., a special feature of this soft coral that can be used as a proxy for its health. Results There were no significant health status differences between methods. This was indicated by similar gross photosynthesis (between 7.4 to 9.7 μg O2 polyp−1 h−1) and pulsating rates (between 35 and 44 pulses min−1) among methods. In terms of time/labor intensity and success rates, i.e., the percentage of fragments attached to the desired surface, the plug mesh method was the most efficient method with a significantly higher success rate (95 ± 5%), while the others had a success rate between 5 ± 5 and 45 ± 15%. The time needed for fragmentation, though not significant, was also the shortest (78 ± 11 s fragment−1), while other methods required between 84 ± 14 and 126 ± 8 s frag−1. The plug mesh method may thus be a valuable tool related to the reproduction of soft corals for use in subsequent experimental work.

Coral reefs experience phase shifts from coral- to algae-dominated benthic communities, which could affect the interplay between processes introducing and removing bioavailable nitrogen. However, the magnitude of such processes, i.e., dinitrogen (N2) fixation and denitrification levels, and their responses to phase shifts remain unknown in coral reefs. We assessed both processes for the dominant species of six benthic categories (hard corals, soft corals, turf algae, coral rubble, biogenic rock, and reef sands) accounting for > 98% of the benthic cover of a central Red Sea coral reef. Rates were extrapolated to the relative benthic cover of the studied organisms in co-occurring coral- and algae-dominated areas of the same reef. In general, benthic categories with high N2 fixation exhibited low denitrification activity. Extrapolated to the respective reef area, turf algae and coral rubble accounted for > 90% of overall N2 fixation, whereas corals contributed to more than half of reef denitrification. Total N2 fixation was twice as high in algae- compared to coral-dominated areas, whereas denitrification levels were similar. We conclude that algae-dominated reefs promote new nitrogen input through enhanced N2 fixation and comparatively low denitrification. The subsequent increased nitrogen availability could support net productivity, resulting in a positive feedback loop that increases the competitive advantage of algae over corals in reefs that experienced a phase shift.

The Eastern Tropical Pacific (ETP) is one of the most isolated and least studied regions in the world. This particularly applies to the coast of El Salvador, where the only reef between Guatemala and Nicaragua, called Los Cóbanos reef, is located. There is very little published information about the reef’s biodiversity, and to our knowledge, no research on its ecology and responses to anthropogenic impacts, such as overfishing, has been conducted. The present study, therefore, described the benthic community of Los Cóbanos reef, El Salvador, using the Line-Point-Intercept-Transect method and investigated changes in the benthic community following the exclusion of piscine macroherbivores over a period of seven weeks. Results showed high benthic algae cover (up to 98%), dominated by turf and green algae, and low coral cover (0–4%). Porites lobata was the only hermatypic coral species found during the surveys. Surprisingly, crustose coralline algae (CCA) showed a remarkable total cover increase by 58%, while turf algae cover decreased by 82%, in experimental plots after seven weeks of piscine macroherbivore exclusion. These findings apparently contradict the results of most previous similar studies. While it was not possible to ascertain the exact mechanisms leading to these drastic community changes, the most likely explanation is grazing on turf by small grazing macroherbivores that had access to the cages during the experiment and clearing of CCA initially covered by epiphytes and sediments. A higher CCA cover would promote the succesful settlement by corals and prevent further erosion of the reef framework. Therefore it is crucial to better understand algal dynamics, herbivory, and implications of overfishing at Los Cóbanos to avoid further reef deterioration. This could be achieved through video surveys of the fish community, night-time observations of the macroinvertebrate community, exclusion experiments that also keep out herbivorous macroinvertebrates, and/or experimental assessments of turf algae/CCA interactions.

Prolonged aerial exposure can be detrimental for coral health. Here, we present an observation of a scleractinian corals' outstanding regeneration after exposure to air for several months. This observation may lead to new ways to investigate the regenerative capacity of reef-building corals.

This is the data to "High summer temperatures amplify functional differences between coral- and algae-dominated reef communities". All information on how the dataset was collected can be found in the manuscript. Abstract of the manuscript: Shifts from coral to algal dominance are expected to increase in tropical coral reefs as a result of anthropogenic disturbances. The consequences for key ecosystem functions such as primary productivity, calcification, and nutrient recycling are poorly understood, particularly under changing environmental conditions. We used a novel in situ incubation approach to compare functions of coral- and algae-dominated communities in the central Red Sea bi-monthly over an entire year. In situ gross and net community primary productivity, calcification, dissolved organic carbon fluxes, dissolved inorganic nitrogen fluxes, and their respective activation energies were quantified to describe the effects of seasonal changes. Overall, coral-dominated communities exhibited 30% lower net productivity and 10 times higher calcification than algae-dominated communities. Estimated activation energies indicated a higher thermal sensitivity of coral-dominated communities. In these communities, net productivity and calcification were negatively correlated with temperature (>40% and >65% reduction, respectively, with +5°C increase from winter to summer), while carbon losses via respiration and dissolved organic carbon release were more than doubled at higher temperatures. In contrast, algae-dominated communities doubled net productivity in summer, while calcification and dissolved organic carbon fluxes were unaffected. These results suggest pronounced changes in community functioning associated with phase shifts. Algae-dominated communities may outcompete coral-dominated communities due to their higher productivity and carbon retention to support fast biomass accumulation while compromising the formation of important reef framework structures. Higher temperatures likely amplify these functional differences, indicating a high vulnerability of ecosystem functions of coral-dominated communities to temperatures even below coral bleaching thresholds. Our results suggest that ocean warming may not only cause but also amplify coral-algal phase shifts in coral reefs. Usage information: The data Excel file contains two data sheets: Sheet 1 (Community composition): This sheet gives the community composition of the assessed benthic communities in % cover of functional groups. Sheet 2 (Metabolism): This sheet contains the metabolic data of the benthic communities. All abbreviations and units are in the related publication.

Human activities are changing ecosystems at an unprecedented rate, yet large-scale studies into how local human impacts alter natural systems and interact with other aspects of global change are still lacking. Here we provide empirical evidence that local human impacts fundamentally alter relationships between ecological communities and environmental drivers. Using tropical coral reefs as a study system, we investigated the influence of contrasting levels of local human impact using a spatially extensive dataset spanning 62 outer reefs around inhabited Pacific islands. We tested how local human impacts (low versus high determined using a threshold of 25 people km−2 reef) affected benthic community (i) structure, and (ii) relationships with environmental predictors using pre-defined models and model selection tools. Data on reef depth, benthic assemblages, and herbivorous fish communities were collected from field surveys. Additional data on thermal stress, storm exposure, and market gravity (a function of human population size and reef accessibility) were extracted from public repositories. Findings revealed that reefs subject to high local human impact were characterised by relatively more turf algae (>10% higher mean absolute coverage) and lower live coral cover (9% less mean absolute coverage) than reefs subject to low local human impact, but had similar macroalgal cover and coral morphological composition. Models based on spatio-physical predictors were significantly more accurate in explaining the variation of benthic assemblages at sites with low (mean adjusted-R2 = 0.35) rather than high local human impact, where relationships became much weaker (mean adjusted-R2 = 0.10). Model selection procedures also identified a distinct shift in the relative importance of different herbivorous fish functional groups in explaining benthic communities depending on the local human impact level. These results demonstrate that local human impacts alter natural systems and indicate that projecting climate change impacts may be particularly challenging at reefs close to higher human populations, where dependency and pressure on ecosystem services are highest.

Recent research indicates that synchronicity of sexual reproduction in coral spawning events is breaking down, leading to aging populations and decreased recruitment success. In this perspective, we develop a hypothesis that this phenomenon could be caused by ongoing ocean acidification (OA). We hypothesize, that the underlying physiological machinery could be the carbon concentrating mechanism (CCM). The endosymbiotic zooxanthellae of corals could use this mechanism to sense calm water motion states in a comparable way to that known from macroalgae. In macroalgae, it is well-established that dissolved inorganic carbon (DIC) acts as the trigger for signaling low water motion. Hence, evolutionarily developed signals of low water motion, suited for gamete-release, may be misleading in the future, potentially favoring opportunistic species in a broad range of marine organisms.

The loss of coral cover is often accompanied by an increase of benthic algae, a decline in biodiversity and habitat complexity. However, it remains unclear how surrounding communities influence the trajectories of re-colonization between pulse disturbance events. Over a 12-month field experiment in the central Red Sea, we examined how healthy (hard-coral dominated) and degraded (algae-dominated) reef areas influence recruitment and succession patterns of benthic reef foundation communities on bare substrates. Crustose coralline algae and other calcifiers were important colonizers in the healthy reef area, promoting the accumulation of inorganic carbon. Contrary, substrates in the degraded area were predominantly colonized by turf algae, lowering the accumulation of inorganic carbon by 178%. While coral larvae settlement similarly occurred in both habitats, degraded areas showed 50% fewer recruits. Our findings suggest that in degraded reefs the replenishment of adult coral populations is reduced due to recruitment inhibition through limited habitat complexity and grazing pressure, thereby restraining reef recovery.

The Red Sea is one of the warmest and most saline seas on the planet. Yet, scleractinian corals have managed to flourish under these distinct conditions supporting one of the largest networks of coral reef ecosystems worldwide. Here, we summarize current knowledge on the ecophysiology of reef-building corals gained from 60 years of research in the Red Sea starting from insights in the 1960s to the most recent studies of the past few years. We provide a brief overview over seasonal dynamics and environmental gradients in the Red Sea that are used to study ecophysiological processes of corals under changing environmental and extreme conditions (i.e., temperature, salinity, nutrient, and light availability). We then focus on how this environmental variability shapes the central processes of coral physiology in the Red Sea covering the topics of photosynthesis, calcification, nutrient cycling, and reproduction. We continue by reporting the first physiological measurements of Red Sea deep-sea corals. Last, we discuss how, through the integration of traditional methods with recent developments in the omics field and model systems, we are now beginning to understand the complexity of processes that contribute to the ecological success of corals under these variable conditions. This synthesis may serve as a basis for future studies that aim to contribute to a better understanding of the impacts of environmental change on coral reefs in the Red Sea and the rest of the world.

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.

Epilithic algal communities play critical ecological roles on coral reefs, but their response to individual and interactive effects of ocean warming (OW) and ocean acidification (OA) is still largely unknown. We investigated growth, photosynthesis and calcification of early epilithic algal community assemblages exposed for 6 months to four temperature profiles (−1.1, ±0.0, +0.9, +1.6 °C) that were crossed with four carbon dioxide partial pressure (pCO2) levels (360, 440, 650, 940 µatm), under flow-through conditions and natural light regimes. Additionally, we compared the cover of heavily calcified crustose coralline algae (CCA) and lightly calcified red algae of the genus Peyssonnelia among treatments. Increase in cover of epilithic communities showed optima under moderately elevated temperatures and present pCO2, while cover strongly decreased under high temperatures and high-pCO2 conditions, particularly due to decreasing cover of CCA. Similarly, community calcification rates were strongly decreased at high pCO2 under both measured temperatures. While final cover of CCA decreased under high temperature and pCO2 (additive negative effects), cover of Peyssonnelia spp. increased at high compared to annual average and moderately elevated temperatures. Thus, cover of Peyssonnelia spp. increased in treatment combinations with less CCA, which was supported by a significant negative correlation between organism groups. The different susceptibility to stressors most likely derived from a different calcification intensity and/or mineral. Notably, growth of the epilithic communities and final cover of CCA were strongly decreased under reduced-pCO2 conditions compared to the present. Thus, CCA may have acclimatized from past to present-day pCO2 conditions, and changes in carbonate chemistry, regardless in which direction, negatively affect them. However, if epilithic organisms cannot further acclimatize to OW and OA, the interacting effects of both factors may change epilithic communities in the future, thereby likely leading to reduced reef stability and recovery.

Observations of coral–algal competition can provide valuable information about the state of coral reef ecosystems. Here, we report contact rates and apparent competition states for six shallow lagoonal reefs in Fiji. A total of 81.4% of examined coral perimeters were found to be in contact with algae, with turf algae (54.7%) and macroalgae of the genus Lobophora (16.8%) representing the most frequently observed contacts. Turf algae competitiveness was low, with 21.8% of coral–turf contacts being won by the algae (i.e. overgrowth or bleaching of coral tissue). In contrast, Lobophora competitiveness against corals was high, with 62.5% of contacts being won by the alga. The presence of epiphytic algae on Lobophora was associated with significantly greater algal competitiveness against corals, with 75.8% and 21.1% of interactions recorded as algal wins in the presence and absence of epiphytes, respectively. Sedimentation rate, herbivorous fish biomass, and coral colony size did not have a significant effect on Lobophora–coral interactions. This research indicates a novel and important role of epiphytes in driving the outcome of coral–algal contacts.

The Spermonde Archipelago is a complex of ~70 mostly populated islands off Southwest Sulawesi, Indonesia, in the center of the Coral Triangle. The reefs in this area are exposed to a high level of anthropogenic disturbances. Previous studies have shown that variation in the benthos is strongly linked to water quality and distance from the mainland. However, little is known about the fish assemblages of the region and if their community structure also follows a relationship with benthic structure and distance from shore. In this study, we used eight islands of the archipelago, varying in distance from 1-55 km relative to the mainland, and three years of surveys, to describe benthic and fish assemblages and to examine the spatial and temporal influence of benthic composition on the structure of the fish assemblages. Cluster analysis indicated that distinct groups of fish were associated with distance, while few species were present across the entire range of sites. Relating fish communities to benthic composition using a multivariate generalized linear model confirmed that fish groups relate to structural complexity (rugosity) or differing benthic groups; either algae, reef builders (coral and crustose coralline algae) or invertebrates and rubble. From these relationships we can identify sets of fish species that may be lost given continued degradation of the Spermonde reefs. Lastly, the incorporation of water quality, benthic and fish indices indicates that local coral reefs responded positively after an acute disturbance in 2013 with increases in reef builders and fish diversity over relatively short (one year) time frames. This study contributes an important, missing component (fish community structure) to the growing literature on the Spermonde Archipelago, a system that features environmental pressures common in the greater Southeast Asian region.

With increasing stressors to coral reefs, defining tools that evaluate their dynamics and resilience is important to interpret system trajectories and direct conservation efforts. In this context, surveys must go beyond conventional monitoring approaches that focus on abundance and biomass of key groups and quantify metrics that better assess ecological processes and ecosystem trajectories. By measuring a variety of conventional (e.g. proportional cover of broad benthic groups, biomass of herbivorous fish) and complementary resilience-based metrics (e.g. algal turf height, coral recruitment rates, juvenile coral densities, herbivorous fish grazing rates), this study evaluated the ecosystem responses to community-based management in Fiji. The study was conducted across three paired tabu areas (periodically closed to fishing) and adjacent fished sites. Conventional metrics reflected no management effect on benthic or herbivorous fish assemblages. In contrast, the complementary metrics generally indicated positive effects of management, particularly within the benthos. Significant differences were observed for turf height (33% lower), coral recruitment rate (159% higher) and juvenile coral density (42% higher) within areas closed to fishing compared to adjacent open reefs. In addition, turf height was inversely related to coral recruitment and juvenile coral density, and longer turfs (≥5 mm) were more competitive in interaction with corals. These results emphasise that conventional metrics may overlook benefits of local management to inshore reefs, and that incorporating complementary resilience-based metrics such as turf height into reef survey protocols will strengthen their capacity to predict the plausible future condition of reefs and their responses to disturbances., This work was supported by the (German) Federal Ministry of Education and Research (BMBF) through the ‘Nachwuchsgruppen Globaler Wandel 4 + 1’ (REPICORE, grant number 01LN1303A).

The importance of Symbiodinium algal endosymbionts and a diverse suite of bacteria for coral holobiont health and functioning are widely acknowledged. Yet, we know surprisingly little about microbial community dynamics and the stability of host-microbe associations under adverse environmental conditions. To gain insight into the stability of coral host-microbe associations and holobiont structure, we assessed changes in the community structure of Symbiodinium and bacteria associated with the coral Pocillopora verrucosa under excess organic nutrient conditions. Pocillopora-associated microbial communities were monitored over 14 days in two independent experiments. We assessed the effect of excess dissolved organic nitrogen (DON) and excess dissolved organic carbon (DOC). Exposure to excess nutrients rapidly affected coral health, resulting in two distinct stress phenotypes: coral bleaching under excess DOC and severe tissue sloughing (>90% tissue loss resulting in host mortality) under excess DON. These phenotypes were accompanied by structural changes in the Symbiodinium community. In contrast, the associated bacterial community remained remarkably stable and was dominated by two Endozoicomonas phylotypes, comprising on average 90% of 16S rRNA gene sequences. This dominance of Endozoicomonas even under conditions of coral bleaching and mortality suggests the bacterial community of P. verrucosa may be rather inflexible and thereby unable to respond or acclimatize to rapid changes in the environment, contrary to what was previously observed in other corals. In this light, our results suggest that coral holobionts might occupy structural landscapes ranging from a highly flexible to a rather inflexible composition with consequences for their ability to respond to environmental change. published

Browsing fishes have been identified as an important component of coral reef resilience, because in contrast to other herbivorous fishes they are able to feed on established macroalgae. Climate change and local anthropogenic impacts have contributed to phase shifts in many coral reefs from coral to macroalgae dominance, and recent research suggests the potential ability of browsers to reverse such phase shifts. However, there is high variation among studies and some contradicting findings exist. Here, we review the relevant literature to assemble a list of species currently known to contribute to browsing in the Indo-Pacific. Furthermore, we identify spatial and temporal patterns, outline factors influencing browsing, and discuss the probability of phase shift reversal. We formulate research recommendations addressing the identified gaps in knowledge about the interactions of browsing fishes and their environment. To date, 37 species of fishes have been observed consuming macroalgae in the Indo-Pacific. The most important groups are the family Siganidae, the subfamily Scarinae (Labridae), and the subfamily Nasinae (Acanthuridae). Browsing species vary between studies depending on location, season and macroalgae species examined. Several influencing factors, such as structural complexity, palatability of macroalgae and ecosystem connectivity have been suggested to cause these discrepancies. The most promising avenues for future research are the effect of structural complexity, the importance of mobile link species and influences of food availability on the selectivity of browsing species. Increasing our knowledge in these fields will provide a better basis for successful management strategies directed at increasing the resilience of coral reefs.

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

Corals are animals that form close mutualistic associations with endosymbiotic photosynthetic algae of the genus Symbiodinium. Together they provide the calcium carbonate framework of coral reef ecosystems. The importance of the microbiome (i.e., bacteria, archaea, fungi, and viruses) to holobiont functioning has only recently been recognized. Given that growth and density of Symbiodinium within the coral host is highly dependent on nitrogen availability, nitrogen-cycling microbes may be of fundamental importance to the stability of the coral-algae symbiosis and holobiont functioning, in particular under nutrient-enriched and -depleted scenarios. We summarize what is known about nitrogen cycling in corals and conclude that disturbance of microbial nitrogen cycling may be tightly linked to coral bleaching and disease. published

A recent review of crown-of-thorns starfish (Acanthaster planci; COTS) in Indonesia has suggested that their impacts have gone under-reported. In 2012–2013, we surveyed COTS at permanent transects within 12 sites of the Spermonde Archipelago, Indonesia, a coastal region close to two rivers and the heavily urbanised city of Makassar. Evidence of COTS was apparent at 8 of 12 sites surveyed with highest densities (37 starfish per 250 m−2 at Barrang Lompo) comparable to those reported in the Indonesian historical literature. At Barrang Lompo and Bonetambung, the COTS outbreak resulted in the loss of half the live coral. Terrestrial effluents have reduced water quality in the Spermonde Archipelago, which further supports recent work linking water quality and COTS outbreaks, thus providing a warning of future outbreaks to Indonesian coastal managers given the country’s increased urbanisation.

Localized impacts, such as eutrophication and overfishing, have been identified as major threats to reefs worldwide, resulting in changes in coral reef community composition, from coral-dominance to algal-dominance. Herbivory by fishes is frequently cited as a key process structuring benthic community response to stressors. Here, we experimentally tested the capacity of herbivorous fishes to remove macroalgae along an onshore–offshore spatial gradient in the Spermonde Archipelago, Indonesia; a coral reef system exposed to long-term anthropogenic disturbances. Bioassays of Sargassum and Padina spp. were collected from the reef flat and transplanted onto the reef slope at five islands varying in distance (1–55 km) from the coastline and city of Makassar, South Sulawesi. Supplementary underwater video deployment and fish surveys were used to identify species-specific differences in herbivore communities and rates of algal removal. Over a 24 h period, Padina was almost completely removed from the two outermost islands, but removal declined toward shore. With the exception of the site closest to Makassar, more than 50% of all transplanted macroalgae were removed. Species diversity of algal-consuming fishes and total bites increased toward off-shore, with only one species, Siganus virgatus (Siganidae), feeding at all sites. These changes in herbivore communities underline the capacity for functional resilience under varying levels of reef degradation. Our results exemplify the complexities of ecological functioning along environmental gradients.

Coral reefs in the Gulf of Thailand are highly under-investigated regarding responses to anthropogenic stressors. Thus, this study simulated overfishing and eutrophication using herbivore exclosure cages and slow-release fertilizer to study the in-situ effects on benthic algae and invertebrate settlement in a coral reef of Koh Phangan, Thailand. Settlement of organisms and the development of organic matter on light-exposed and shaded tiles were quantified weekly/biweekly over a study period of 12 weeks. Simulated eutrophication did not significantly influence response parameters, while simulated overfishing positively affected dry mass, turf algae height and fleshy macroalgae occurrence on light-exposed tiles. On shaded tiles, settlement of crustose coralline algae decreased, while abundances of ascidians increased compared to controls. An interactive effect of both stressors was not observed. These results hint to herbivory as actual key controlling factor on the benthic community, and fleshy macroalgae together with ascidians as potential bioindicators for local overfishing.

Sea cucumbers play an important role in the recycling and remineralization of organic matter (OM) in reef sands through feeding, excretion, and bioturbation processes. Growing demand from Asian markets has driven the overexploitation of these animals globally. The implications of sea cucumber fisheries for shallow coastal ecosystems and their management remain poorly understood. To address this knowledge gap, the current study manipulated densities of Holothuria scabra within enclosures on a reef flat in Fiji, between August 2015 and February 2016, to study the effects of sea cucumber removal on sedimentary function as a biocatalytic filter system. Three treatments were investigated: (i) high density (350 g m−2 wet weight; ca. 15 individuals); (ii) natural density (60 g m−2; ca. 3 individuals); and (iii) exclusion (0 g m−2). Quantity of sediment reworked through ingestion by H. scabra, grain size distribution, O2 penetration depth, and sedimentary oxygen consumption (SOC) were quantified within each treatment. Findings revealed that the natural population of H. scabra at the study site can rework ca. 10,590 kg dry sediment 1,000 m−2 year−1; more than twice the turnover rate recorded for H. atra and Stichopus chloronotus. There was a shift towards finer fraction grains in the high treatment. In the exclusion treatment, the O2 penetration depth decreased by 63% following a 6 °C increase in water temperature over the course of two months, while in the high treatment no such change was observed. SOC rates increased ca. two-fold in the exclusion treatment within the first month, and were consistently higher than in the high treatment. These results suggest that the removal of sea cucumbers can reduce the capacity of sediments to buffer OM pulses, impeding the function and productivity of shallow coastal ecosystems.

The invasion of Indo-Pacific lionfish is one of the most pressing concerns in the context of coral reef conservation throughout the Caribbean. Invasive lionfish threaten Caribbean fish communities by feeding on a wide range of native prey species, some of which have high ecological and economic value. In Roatan (Honduras) a local non-governmental organisation (i.e. Roatan Marine Park) trains residents and tourists in the use of spears to remove invasive lionfish. Here, we assess the effectiveness of local removal efforts in reducing lionfish populations. We ask whether reefs subject to relatively frequent removals support more diverse and abundant native fish assemblages compared to sites were no removals take place. Lionfish biomass, as well as density and diversity of native prey species were quantified on reefs subject to regular and no removal efforts. Reefs subject to regular lionfish removals (two to three removals month−1) with a mean catch per unit effort of 2.76 ± 1.72 lionfish fisher−1 h−1 had 95% lower lionfish biomass compared to non-removal sites. Sites subject to lionfish removals supported 30% higher densities of native prey-sized fishes compared to sites subject to no removal efforts. We found no evidence that species richness and diversity of native fish communities differ between removal and non-removal sites. We conclude that opportunistic voluntary removals are an effective management intervention to reduce lionfish populations locally and might alleviate negative impacts of lionfish predation. We recommend that local management and the diving industry cooperate to cost-effectively extend the spatial scale at which removal regimes are currently sustained.

The endosymbiosis between cnidarians and dinoflagellates of the genus Symbiodinium is key to the high productivity of tropical coral reefs. In this endosymbiosis, Symbiodinium translocate most of their photosynthates to their animal host in exchange for inorganic nutrients. Among these, carbon dioxide (CO2) derived from host respiration helps to meet the carbon requirements to sustain photosynthesis of the dinoflagellates. Nonetheless, recent studies suggest that productivity in symbiotic cnidarians such as corals is CO2-limited. Here we show that glucose enrichment stimulates respiration and gross photosynthesis rates by 80 and 140%, respectively, in the symbiotic upside-down jellyfish Cassiopeia sp. from the Central Red Sea. Our findings show that glucose was rapidly consumed and respired within the Cassiopeia sp. holobiont. The resulting increase of CO2 availability in hospite in turn likely stimulated photosynthesis in Symbiodinium. Hence, the increase of photosynthesis under these conditions suggests that CO2 limitation of Symbiodinium is a common feature of stable cnidarian holobionts and that the stimulation of holobiont metabolism may attenuate this CO2 limitation. published

Coastal eutrophication is a key driver of shifts in bacterial communities on coral reefs. With fringing and patch reefs at varying distances from the coast the Spermonde Archipelago in southern Sulawesi, Indonesia offers ideal conditions to study the effects of coastal eutrophication along a spatially defined gradient. The present study investigated bacterial community composition of three coral reef habitats: the water column, sediments, and mucus of the hard coral genus Fungia, along that cross-shelf environmental and water quality gradient. The main research questions were: (1) How do water quality and bacterial community composition change along a coastal shelf gradient? (2) Which water quality parameters influence bacterial community composition? (3) Is there a difference in bacterial community composition among the investigated habitats? For this purpose, a range of key water parameters were measured at eight stations in distances from 2 to 55 km from urban Makassar. This was supplemented by sampling of bacterial communities of important microbial habitats using 454 pyrosequencing. Findings revealed that the population center Makassar had a strong effect on the concentrations of Chlorophyll a, suspended particulate matter (SPM), and transparent exopolymer particles (TEP), which were all significantly elevated at the inshore compared the other seven sites. Shifts in the bacterial communities were specific to each sampled habitat. Two OTUs, belonging to the genera Escherichia/Shigella (Gammaproteobacteria) and Ralstonia (Betaproteobacteria), respectively, both dominated the bacterial community composition of the both size fractions of the water column and coral mucus. The sampled reef sediments were more diverse, and no single OTUs was dominant. There was no gradual shift in bacterial classes or OTUs within the sampled habitats. In addition, we observed very distinct communities between the investigated habitats. Our data show strong changes in the bacterial community composition at the inshore site for water column and sediment samples. Alarmingly, there was generally a high prevalence of potentially pathogenic bacteria across the entire gradient.

Microbial nitrogen fixation (diazotrophy) is a functional trait widely associated with tropical reef-building (scleractinian) corals. While the integral role of nitrogen fixation in coral nutrient dynamics is recognized, its ecological significance across different coral functional groups remains yet to be evaluated. Here we set out to compare molecular and physiological patterns of diazotrophy (i.e., nifH gene abundance and expression as well as nitrogen fixation rates) in two coral families with contrasting trophic strategies: highly heterotrophic, free-living members of the family Fungiidae (Pleuractis granulosa, Ctenactis echinata), and mostly autotrophic coral holobionts with low heterotrophic capacity (Pocilloporidae: Pocillopora verrucosa, Stylophora pistillata). The Fungiidae exhibited low diazotroph abundance (based on nifH gene copy numbers) and activity (based on nifH gene expression and the absence of detectable nitrogen fixation rates). In contrast, the mostly autotrophic Pocilloporidae exhibited nifH gene copy numbers and gene expression two orders of magnitude higher than in the Fungiidae, which coincided with detectable nitrogen fixation activity. Based on these data, we suggest that nitrogen fixation compensates for the low heterotrophic nitrogen uptake in autotrophic corals. Consequently, the ecological importance of diazotrophy in coral holobionts may be determined by the trophic functional group of the host. published

The disruption of the coral-algae symbiosis (coral bleaching) due to rising sea surface temperatures has become an unprecedented global threat to coral reefs. Despite decades of research, our ability to manage mass bleaching events remains hampered by an incomplete mechanistic understanding of the processes involved. In this study, we induced a coral bleaching phenotype in the absence of heat and light stress by adding sugars. The sugar addition resulted in coral symbiotic breakdown accompanied by a fourfold increase of coral-associated microbial nitrogen fixation. Concomitantly, increased N:P ratios by the coral host and algal symbionts suggest excess availability of nitrogen and a disruption of the nitrogen limitation within the coral holobiont. As nitrogen fixation is similarly stimulated in ocean warming scenarios, here we propose a refined coral bleaching model integrating the cascading effects of stimulated microbial nitrogen fixation. This model highlights the putative role of nitrogen-fixing microbes in coral holobiont functioning and breakdown. published

SUMMARYSmall-island coral reef ecosystems are usually closely coupled to the activities of human inhabitants. Ahus Island (Papua New Guinea) is an isolated Pacific island with a rapidly growing population, heavy reliance on marine resources and limited infrastructure. We hypothesized that untreated sewage was driving distinct benthic assemblages around Ahus and neighbouring uninhabited Onetah. At sites with varying proximities to beach toilets, fore-reef herbivorous fish biomass and benthic composition were measured alongside reef-flat sedimentary oxygen consumption (SOC); a high SOC rate reflects high organic input into coastal waters, thus serving as a potential indicator of sewage input. Fish biomass was low (17.1–20.1 g m–2), but consistent between sites. However, cyanobacteria dominated the fore-reef closest to toilets (62 ± 3%) with highest reef-flat SOC, whereas hard corals dominated furthest away (63 ± 1%), where SOC was lowest. To our knowledge, this is the first study that used SOC to detect local differences in sewage pollution. The results indicate that whilst corals can maintain their dominance on overfished reefs, additional sewage stress may drive pronounced benthic shifts, highlighting the urgency to improve small-island waste management.

Coastal pollution and algal cover are increasing on many coral reefs, resulting in higher dissolved organic carbon (DOC) concentrations. High DOC concentrations strongly affect microbial activity in reef waters and select for copiotrophic, often potentially virulent microbial populations. High DOC concentrations on coral reefs are also hypothesized to be a determinant for switching microbial lifestyles from commensal to pathogenic, thereby contributing to coral reef degradation, but evidence is missing. In this study, we conducted ex situ incubations to assess gene expression of planktonic microbial populations under elevated concentrations of naturally abundant monosaccharides (glucose, galactose, mannose, and xylose) in algal exudates and sewage inflows. We assembled 27 near-complete (>70%) microbial genomes through metagenomic sequencing and determined associated expression patterns through metatranscriptomic sequencing. Differential gene expression analysis revealed a shift in the central carbohydrate metabolism and the induction of metalloproteases, siderophores, and toxins in Alteromonas, Erythrobacter, Oceanicola, and Alcanivorax populations. Sugar-specific induction of virulence factors suggests a mechanistic link for the switch from a commensal to a pathogenic lifestyle, particularly relevant during increased algal cover and human-derived pollution on coral reefs. Although an explicit test remains to be performed, our data support the hypothesis that increased availability of specific sugars changes net microbial community activity in ways that increase the emergence and abundance of opportunistic pathogens, potentially contributing to coral reef degradation. published

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 Curacao, 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.

One of the major threats facing coral reefs is intense benthic algal growth that can result in overgrowth and mass mortality of corals if not controlled by herbivore grazing. Unlike the well-studied coastlines of the Caribbean, there is currently a lack of knowledge regarding the effects of herbivory on benthic communities in the Red Sea. This is particularly relevant today as the local impacts in the Red Sea are increasing due to growing population and tourism. Over 4 mo, this study investigated the impact of herbivory as a potential key factor controlling algal growth on a reef flat in the Egyptian northern Red Sea. The main experiment consisted of in situ deploy- ment of exclosure cages in combination with quantification of sea urchins and herbivorous fish. When all herbivores were excluded, our findings showed a significant 17-fold increase of algal dry mass within 4 mo. Although herbivorous fish occurred in much lower abundance (0.6 ± 0.1 ind. m �2 ; mean ± SE) compared to sea urchins (3.4 ± 0.2 ind. m �2 ), they were 5-fold more efficient in reducing algal dry mass and 22-fold more efficient in reducing autotrophic production of nitrogen. A significant shift from benthic turf to macroalgae (mostly Padina sp. and Hydroclathrus clathrathus) was observed when grazers were excluded. These algae may serve as early warning indicators for overfishing. Findings suggest that herbivorous fish act as an important top-down factor controlling both benthic algal biomass and composition at the study location. Results also indicate the potential of rapid benthic community change at the study site if herbivory is impeded.

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.

Shallow warm-water and deep-sea cold-water corals engineer the coral reef framework and fertilize reef communities by releasing coral mucus, a source of reef dissolved organic matter (DOM). By transforming DOM into particulate detritus, sponges play a key role in transferring the energy and nutrients in DOM to higher trophic levels on Caribbean reefs via the so-called sponge loop. Coral mucus may be a major DOM source for the sponge loop, but mucus uptake by sponges has not been demonstrated. Here we used laboratory stable isotope tracer experiments to show the transfer of coral mucus into the bulk tissue and phospholipid fatty acids of the warm-water sponge Mycale fistulifera and cold-water sponge Hymedesmia coriacea, demonstrating a direct trophic link between corals and reef sponges. Furthermore, 21–40% of the mucus carbon and 32–39% of the nitrogen assimilated by the sponges was subsequently released as detritus, confirming a sponge loop on Red Sea warm-water and north Atlantic cold-water coral reefs. The presence of a sponge loop in two vastly different reef environments suggests it is a ubiquitous feature of reef ecosystems contributing to the high biogeochemical cycling that may enable coral reefs to thrive in nutrient-limited (warm-water) and energy-limited (cold-water) environments.

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.

16S rRNA gene molecular analysis elucidated the spatiotemporal distribution of bacterial biofilm communities along a water quality gradient. Multivariate statistics indicated that terrestrial runoff, in particular dissolved organic carbon and chlorophyll a concentrations, induced shifts of specific bacterial communities between locations and seasons, suggesting microbial biofilms could be suitable bioindicators for water quality.

There has been an exponential increase in coral mucus research over the last 5 years, attracting attention from coral biologists and reef ecologists. The most active area has been the study of microbial structure and function associated with mucus, and very recent findings have increased our understanding of the roles of microbes in coral health and disease, and also on the ecosystem level. Here we overview some of the latest findings, but also identify scientific gaps. A priority area for future research is understanding the structure and dynamics of the surface mucus layer in relation to microbial community development. Environmental factors including climate change impacts affect the release of mucus/organic matter by the coral engineer and may therefore also have profound effects on entire reef ecosystem function, so it is vital that we also gain a better understanding of these responses.

Natural and anthropogenic impacts such as terrestrial runoff, influence the water quality along the coast of the Great Barrier Reef (GBR) and may in turn affect coral reef communities. Associated bacterial biofilms respond rapidly to environmental conditions and are potential bioindicators for changes in water quality. As a prerequisite to study the effects of water quality on biofilm communities, appropriate biofilm substrates for deployment in the field must be developed and evaluated. This study investigates the effect of different settlement substrates (i.e. glass slides, ceramic tiles, coral skeletons and reef sediments) on bacterial biofilm communities grown in situ for 48 days at two locations in the Whitsunday Island Group (Central GBR) during two sampling times. Bacterial communities associated with the biofilms were analysed using terminal restriction fragment length polymorphism (T-RFLP) and clone library analyses of 16S rRNA genes. Findings revealed that substrate type had little influence on bacterial community composition. Of particular relevance, glass slides and coral skeletons exhibited very similar communities during both sampling times, suggesting the suitability of standardized glass slides for long-term biofilm indicator studies in tropical coral reef ecosystems.

Permeable sediments and associated microbial communities play a fundamental role in nutrient recycling within coral reef ecosystems by ensuring high levels of primary production in oligotrophic environments. A previous study on organic matter degradation within biogenic carbonate and terrigenous silicate reef sands in the Red Sea suggested that observed sand-specific differences in microbial activity could be caused by variations in microbial biomass and diversity. Here, we tested this hypothesis by comparing bacterial abundance and community structure in both sand types, and by further exploring the structuring effects of time (season) and space (sediment depth, in/out-reef). Changes in bacterial community structure, as determined via automated ribosomal intergenic spacer analysis (ARISA), were primarily driven by sand mineralogy at specific seasons, sediment depths and reef locations. By coupling ARISA with 16S-ITS rRNA sequencing, we detected significant community shifts already at the bacterial class level, with Proteobacteria (Gamma-, Delta-, Alpha-) and Actinobacteria being prominent members of the highly diverse communities. Overall, our findings suggest that reef sand-associated bacterial communities vary substantially with sand type. Especially in synergy with environmental variation over time and space, mineralogical differences seem to play a central role in maintaining high levels of bacterial community heterogeneity. The local co-occurrence of carbonate and silicate sands may thus significantly increase the availability of microbial niches within a single coral reef ecosystem.