Benthic microbial biogeographic trends in the North Sea are shaped by an interplay of environmental drivers and bottom trawling effort.

Microbial composition and diversity in marine sediments are shaped by environmental, biological, and anthropogenic processes operating at different scales. However, our understanding of benthic microbial biogeography remains limited. Here, we used 16S rDNA amplicon sequencing to characterize benthic microbiota in the North Sea from the top centimeter of 339 sediment samples. We utilized spatially explicit statistical models, to disentangle the effects of the different predictors, including bottom trawling intensity, a prevalent industrial fishing practice which heavily impacts benthic ecosystems. Fitted models demonstrate how the geographic interplay of different environmental and anthropogenic drivers shapes the diversity, structure and potential metabolism of benthic microbial communities. Sediment properties were the primary determinants, with diversity increasing with sediment permeability but also with mud content, highlighting different underlying processes. Additionally, diversity and structure varied with total organic matter content, temperature, bottom shear stress and bottom trawling. Changes in diversity associated with bottom trawling intensity were accompanied by shifts in predicted energy metabolism. Specifically, with increasing trawling intensity, we observed a transition toward more aerobic heterotrophic and less denitrifying predicted metabolism. Our findings provide first insights into benthic microbial biogeographic patterns on a large spatial scale and illustrate how anthropogenic activity such as bottom trawling may influence the distribution and abundances of microbes and potential metabolism at macroecological scales.

Coral reef benthic community changes in the Anthropocene: Biogeographic heterogeneity, overlooked configurations, and methodology.

Non-random community changes are becoming more frequent in many ecosystems. In coral reefs, changes towards communities dominated by other than hard corals are increasing in frequency, with severe impacts on ecosystem functioning and provision of ecosystem services. Although new research suggests that a variety of alternative communities (i.e. not dominated by hard corals) exist, knowledge on the global diversity and functioning of alternative coral reef benthic communities, especially those not dominated by algae, remains scattered. In this systematic review and meta-analysis of 523 articles, we analyse the different coral reef benthic community changes reported to date and discuss the advantages and limitations of the methods used to study these changes. Furthermore, we used field cover data (1116 reefs from the ReefCheck database) to explore the biogeographic and latitudinal patterns in dominant benthic organisms. We found a mismatch between literature focus on coral-algal changes (over half of the studies analysed) and observed global natural patterns. We identified strong biogeographic patterns, with the largest and most biodiverse biogeographic regions (Western and Central Indo-Pacific) presenting previously overlooked soft-coral-dominated communities as the most abundant alternative community. Finally, we discuss the potential biases associated with methods that overlook ecologically important cryptobenthic communities and the potential of new technological advances in improving monitoring efforts. As coral reef communities inevitably and swiftly change under changing ocean conditions, there is an urgent need to better understand the distribution, dynamics as well as the ecological and societal impacts of these new communities.

Nutrient History Affects the Response and Resilience of the Tropical Seagrass Halophila stipulacea to Further Enrichment in Its Native Habitat.

Eutrophication is one of the main threats to seagrass meadows, but there is limited knowledge on the interactive effects of nutrients under a changing climate, particularly for tropical seagrass species. This study aimed to detect the onset of stress in the tropical seagrass, Halophila stipulacea, by investigating the effect of in situ nutrient addition during an unusually warm summer over a 6-month period. We measured a suite of different morphological and biochemical community metrics and individual plant traits from two different sites with contrasting levels of eutrophication history before and after in situ fertilization in the Gulf of Aqaba. Nutrient stress combined with summer temperatures that surpassed the threshold for optimal growth negatively affected seagrass plants from South Beach (SB), an oligotrophic marine protected area, while H. stipulacea populations from North Beach (NB), a eutrophic and anthropogenically impacted area, benefited from the additional nutrient input. Lower aboveground (AG) and belowground (BG) biomass, reduced Leaf Area Index (LAI), smaller internodal distances, high sexual reproductive effort and the increasing occurrence of apical shoots in seagrasses from SB sites indicated that the plants were under stress and not growing under optimal conditions. Moreover, AG and BG biomass and internodal distances decreased further with the addition of fertilizer in SB sites. Results presented here highlight the fact that H. stipulacea is one of the most tolerant and plastic seagrass species. Our study further demonstrates that the effects of fertilization differ significantly between meadows that are growing exposed to different levels of anthropogenic pressures. Thus, the meadow's "history" affects it resilience and response to further stress. Our results suggest that monitoring efforts on H. stipulacea populations in its native range should focus especially on carbohydrate reserves in leaves and rhizomes, LAI, internodal length and percentage of apical shoots as suitable warning indicators for nutrient stress in this seagrass species to minimize future impacts on these valuable ecosystems.

Unusually Warm Summer Temperatures Exacerbate Population and Plant Level Response of Posidonia oceanica to Anthropogenic Nutrient Stress.

Posidonia oceanica is a key foundation species in the Mediterranean providing valuable ecosystem services. However, this species is particularly vulnerable towards high coastal nutrient inputs and the rising frequency of intense summer heat waves, but their combined effect in situ has received little attention so far. Here, we investigated the effects of in situ nutrient addition during an unusually warm summer over a 4-month period, comparing different morphological, physiological and biochemical population metrics of seagrass meadows growing in protected areas (Ischia) with meadows already exposed to significant anthropogenic pressure (Baia - Gulf of Pozzuoli). Our study highlights that the effects of warmer than usual summer temperatures on the population level of seagrass meadows can be exacerbated if the plants are already exposed to higher anthropogenic pressures. Morphological and population level indicators mainly changed over time, possibly impacted by season and the warmer temperatures, and displayed more pronounced reductions in seagrasses from impacted sites. The additional nutrient supply had even more deleterious effects, as shown by a decrease in approximately 67% in cover in fertilized plots at high impacted sites and 33% at low impacted sites. Moreover, while rhizome starch concentration showed a seasonal increase in plants from low impacted sites it displayed a trend of a 27% decrease in fertilized plots of the high impacted sites. Epiphyte biomass was approximately four-fold higher on leaves of plants growing in impacted sites and even doubled with the additional nutrient input. Predicting and anticipating stress in P. oceanica is of crucial importance for conservation and management efforts, given the limited colonizing and reproductive ability and extremely slow growth of this ecosystem engineer. Our results suggest that monitoring efforts should focus especially on leaf area index (LAI), carbohydrate concentrations in the rhizomes, and epiphyte cover on leaves as indicators of the onset of stress in Posidonia oceanica, which can be used by decision makers to take appropriate measures before damage to the ecosystem becomes irreversible, minimize future human interference and strengthen the resilience of these important ecosystems.

Sponges from Zanzibar host diverse prokaryotic communities with potential for natural product synthesis.

Sponges are one of the most dominant organisms in marine ecosystems. One reason for their success is their association with microorganisms that are besides the host itself responsible for the chemical defence. Sponge abundances have been increasing on coral reefs in the Western Indian Ocean (WIO) and are predicted to increase further with rising anthropogenic impacts on coral reefs. However, there is a paucity of information on chemical ecology of sponges from the WIO and their prokaryotic community composition. We used a combination of Illumina sequencing and a predictive metagenomic analysis to (i) assess the prokaryotic community composition of sponges from Zanzibar, (ii) predict the presence of KEGG metabolic pathways responsible for bioactive compound production and (iii) relate their presence to the degree of observed chemical defence in their respective sponge host. We found that sponges from Zanzibar host diverse prokaryotic communities that are host species-specific. Sponge-species and respective specimens that showed strong chemical defences in previous studies were also predicted to be highly enriched in various pathways responsible for secondary metabolite production. Hence, the combined sequencing and predictive metagenomic approach proved to be a useful indicator for the metabolic potential of sponge holobionts.

Sponge chemical defenses are a possible mechanism for increasing sponge abundance on reefs in Zanzibar.

Coral reefs are experiencing increasing anthropogenic impacts that result in substantial declines of reef-building corals and a change of community structure towards other benthic invertebrates or macroalgae. Reefs around Zanzibar are exposed to untreated sewage and runoff from the main city Stonetown. At many of these sites, sponge cover has increased over the last years. Sponges are one of the top spatial competitors on reefs worldwide. Their success is, in part, dependent on their strong chemical defenses against predators, microbial attacks and other sessile benthic competitors. This is the first study that investigates the bioactive properties of sponge species in the Western Indian Ocean region. Crude extracts of the ten most dominant sponge species were assessed for their chemical defenses against 35 bacterial strains (nine known as marine pathogens) using disc diffusion assays and general cytotoxic activities were assessed with brine shrimp lethality assays. The three chemically most active sponge species were additionally tested for their allelopathic properties against the scleractinian coral competitor Porites sp.. The antimicrobial assays revealed that all tested sponge extracts had strong antimicrobial properties and that the majority (80%) of the tested sponges were equally defended against pathogenic and environmental bacterial strains. Additionally, seven out of ten sponge species exhibited cytotoxic activities in the brine shrimp assay. Moreover, we could also show that the three most bioactive sponge species were able to decrease the photosynthetic performance of the coral symbionts and thus were likely to impair the coral physiology.