Corals of the world

Losses of corals worldwide emphasize the need to understand what drives reef decline. Stressors such as overfishing and nutrient pollution may reduce resilience of coral reefs by increasing coral–algal competition and reducing coral recruitment, growth and survivorship. Such effects may themselves develop via several mechanisms, including disruption of coral microbiomes. Here we report the results of a 3-year field experiment simulating overfishing and nutrient pollution. These stressors increase turf and macroalgal cover, destabilizing microbiomes, elevating putative pathogen loads, increasing disease more than twofold and increasing mortality up to eightfold. Above-average temperatures exacerbate these effects, further disrupting microbiomes of unhealthy corals and concentrating 80% of mortality in the warmest seasons. Surprisingly, nutrients also increase bacterial opportunism and mortality in corals bitten by parrotfish, turning normal trophic interactions deadly for corals. Thus, overfishing and nutrient pollution impact reefs down to microbial scales, killing corals by sensitizing them to predation, above-average temperatures and bacterial opportunism.

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Overfishing and nutrient pollution interact with temperature to disrupt coral reefs down to microbial scales

Endozoicomonas bacteria were found highly associated with the coral Stylophora pistillata, and these bacteria are also ubiquitously associated with diverse corals worldwide. Novel Endozoicomonas-specific probes revealed that Endozoicomonas bacteria were abundant in the endodermal tissues of S. pistillata and appear to have an intimate relationship with the coral.

https://journals.asm.org/doi/pdf/10.1128/AEM.03365-13

The Microbiome of the Red Sea Coral Stylophora pistillata Is Dominated by Tissue-Associated Endozoicomonas Bacteria

Endozoicomonas bacteria are emerging as extremely diverse and flexible symbionts of numerous marine hosts inhabiting oceans worldwide. Their hosts range from simple invertebrate species, such as sponges and corals, to complex vertebrates, such as fish. Although widely distributed, the functional role of Endozoicomonas within their host microenvironment is not well understood. In this review, we provide a summary of the currently recognized hosts of Endozoicomonas and their global distribution. Next, the potential functional roles of Endozoicomonas, particularly in light of recent microscopic, genomic, and genetic analyses, are discussed. These analyses suggest that Endozoicomonas typically reside in aggregates within host tissues, have a free-living stage due to their large genome sizes, show signs of host and local adaptation, participate in host-associated protein and carbohydrate transport and cycling, and harbour a high degree of genomic plasticity due to the large proportion of transposable elements residing in their genomes. This review will finish with a discussion on the methodological tools currently employed to study Endozoicomonas and host interactions and review future avenues for studying complex host-microbial symbioses.

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Diversity and function of prevalent symbiotic marine bacteria in the genus Endozoicomonas

Reef-building corals are well regarded not only for their obligate association with endosymbiotic algae, but also with prokaryotic symbionts, the specificity of which remains elusive. To identify the central microbial symbionts of corals, their specificity across species and conservation over geographic regions, we sequenced partial SSU ribosomal RNA genes of Bacteria and Archaea from the common corals Stylophora pistillata and Pocillopora verrucosa across 28 reefs within seven major geographical regions. We demonstrate that both corals harbor Endozoicomonas bacteria as their prevalent symbiont. Importantly, catalyzed reporter deposition–fluorescence in situ hybridization (CARD–FISH) with Endozoicomonas-specific probes confirmed their residence as large aggregations deep within coral tissues. Using fine-scale genotyping techniques and single-cell genomics, we demonstrate that P. verrucosa harbors the same Endozoicomonas, whereas S. pistillata associates with geographically distinct genotypes. This specificity may be shaped by the different reproductive strategies of the hosts, potentially uncovering a pattern of symbiont selection that is linked to life history. Spawning corals such as P. verrucosa acquire prokaryotes from the environment. In contrast, brooding corals such as S. pistillata release symbiont-packed planula larvae, which may explain a strong regional signature in their microbiome. Our work contributes to the factors underlying microbiome specificity and adds detail to coral holobiont functioning.

/pdf/differential-specificity-between-closely-related-corals-and-5000ykcfwp.pdf

Differential specificity between closely related corals and abundant Endozoicomonas endosymbionts across global scales.

Coral reefs of the Central Red Sea display a high degree of endemism, and are increasingly threatened by anthropogenic effects due to intense local coastal development measures. Overfishing and eutrophication are among the most significant local pressures on these reefs, but there is no information available about their potential effects on the associated microbial community. Therefore, we compared holobiont physiology and 16S-based bacterial communities of tissue and mucus of the hard coral Acropora hemprichii after 1 and 16 weeks of in-situ inorganic nutrient enrichment (via fertilizer diffusion) and/or herbivore exclusion (via caging) in an offshore reef of the Central Red Sea. Simulated eutrophication and/or overfishing treatments did not affect coral physiology with respect to coral respiration rates, chlorophyll a content, zooxanthellae abundance, or δ 15N isotopic signatures. The bacterial community of A. hemprichii was rich and uneven, and diversity increased over time in all treatments. While distinct bacterial species were identified as a consequence of eutrophication, overfishing, or both, two bacterial species that could be classified to the genus Endozoicomonas were consistently abundant and constituted two thirds of bacteria in the coral. Several nitrogen-fixing and denitrifying bacteria were found in the coral specimens that were exposed to experimentally increased nutrients. However, no particular bacterial species was consistently associated with the coral under a given treatment and the single effects of manipulated eutrophication and overfishing could not predict the combined effect. Our data underlines the importance of conducting field studies in a holobiont framework, taking both, physiological and molecular measures into account.

In-situ Effects of Eutrophication and Overfishing on Physiology and Bacterial Diversity of the Red Sea Coral Acropora hemprichii

Overfishing and land-derived eutrophication are major local threats to coral reefs and may affect benthic communities, moving them from coral dominated reefs to algal dominated ones. The Central Red Sea is a highly under-investigated area, where healthy coral reefs are contending against intense coastal development. This in-situ study investigated both the independent and combined effects of manipulated inorganic nutrient enrichment (simulation of eutrophication) and herbivore exclosure (simulation of overfishing) on benthic algae development. Light-exposed and shaded terracotta tiles were positioned at an offshore patch reef close to Thuwal, Saudi Arabia and sampled over a period of 4 months. Findings revealed that nutrient enrichment alone affected neither algal dry mass nor algae-derived C or N production. In contrast, herbivore exclusion significantly increased algal dry mass up to 300-fold, and in conjunction with nutrient enrichment, this total increased to 500-fold. Though the increase in dry mass led to a 7 and 8-fold increase in organic C and N content, respectively, the algal C/N ratio (18±1) was significantly lowered in the combined treatment relative to controls (26±2). Furthermore, exclusion of herbivores significantly increased the relative abundance of filamentous algae on the light-exposed tiles and reduced crustose coralline algae and non-coralline red crusts on the shaded tiles. The combination of the herbivore exclusion and nutrient enrichment treatments pronounced these effects. The results of our study suggest that herbivore reduction, particularly when coupled with nutrient enrichment, favors non-calcifying, filamentous algae growth with high biomass production, which thoroughly outcompetes the encrusting (calcifying) algae that dominates in undisturbed conditions. These results suggest that the healthy reefs of the Central Red Sea may experience rapid shifts in benthic community composition with ensuing effects for biogeochemical cycles if anthropogenic impacts, particularly overfishing, are not controlled.

/pdf/in-situ-effects-of-simulated-overfishing-and-eutrophication-481vzqa923.pdf

In-Situ Effects of Simulated Overfishing and Eutrophication on Benthic Coral Reef Algae Growth, Succession, and Composition in the Central Red Sea

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.

/pdf/herbivory-effects-on-benthic-algal-composition-and-growth-on-31xcctrmid.pdf

Herbivory effects on benthic algal composition and growth on a coral reef flat in the Egyptian Red Sea

In the Central Red Sea, healthy coral reefs meet intense coastal development, but data on the effects of related stressors for reef functioning are lacking. This in situ study therefore investigated the independent and combined effects of simulated overfishing through predator/grazer exclusion and simulated eutrophication through fertilizer addition on settlement of reef associated invertebrates on light-exposed and -shaded tiles over 4 months. At the end of the study period invertebrates had almost exclusively colonized shaded tiles. Algae were superior settling competitors on light-exposed tiles. On the shaded tiles, simulated overfishing prevented settlement of hard corals, but significantly increased settlement of polychaetes, while simulated eutrophication only significantly decreased hard coral settlement relative to controls. The combined treatment significantly increased settlement of bryozoans and bivalves compared to controls and individual manipulations, but significantly decreased polychaetes compared to simulated overfishing. These results suggest settlement of polychaetes and hard corals as potential bioindicators for overfishing and eutrophication, respectively, and settlement of bivalves and bryozoans for a combination of both. Therefore, if the investigated stressors are not controlled, phase shifts from dominance by hard corals to that by other invertebrates may occur at shaded reef locations in the Central Red Sea.

Christian Jessen

Papers

In-situ Effects of Eutrophication and Overfishing on Physiology and Bacterial Diversity of the Red Sea Coral Acropora hemprichii

In-Situ Effects of Simulated Overfishing and Eutrophication on Benthic Coral Reef Algae Growth, Succession, and Composition in the Central Red Sea

Herbivory effects on benthic algal composition and growth on a coral reef flat in the Egyptian Red Sea

In situ effects of simulated overfishing and eutrophication on settlement of benthic coral reef invertebrates in the Central Red Sea.