The challenge of setting restoration targets for macroalgal forests under climate changes.

The process of site selection and spatial planning has received scarce attention in the scientific literature dealing with marine restoration, suggesting the need to better address how spatial planning tools could guide restoration interventions. In this study, for the first time, the consequences of adopting different restoration targets and criteria on spatial restoration prioritization have been assessed at a regional scale, including the consideration of climate changes. We applied the decision-support tool Marxan, widely used in systematic conservation planning on Mediterranean macroalgal forests. The loss of this habitat has been largely documented, with limited evidences of natural recovery. Spatial priorities were identified under six planning scenarios, considering three main restoration targets to reflect the objectives of the EU Biodiversity Strategy for 2030. Results show that the number of suitable sites for restoration is very limited at basin scale, and targets are only achieved when the recovery of 10% of regressing and extinct macroalgal forests is planned. Increasing targets translates into including unsuitable areas for restoration in Marxan solutions, amplifying the risk of ineffective interventions. Our analysis supports macroalgal forests restoration and provides guiding principles and criteria to strengthen the effectiveness of restoration actions across habitats. The constraints in finding suitable areas for restoration are discussed, and recommendations to guide planning to support future restoration interventions are also included.

Chronic and intensive bottom trawling impairs deep-sea biodiversity and ecosystem functioning.

Bottom trawling has many impacts on marine ecosystems, including seafood stock impoverishment, benthos mortality, and sediment resuspension. Historical records of this fishing practice date back to the mid-1300s. Trawling became a widespread practice in the late 19th century, and it is now progressively expanding to greater depths, with the concerns about its sustainability that emerged during the first half of the 20th century now increasing. We show here that compared with untrawled areas, chronically trawled sediments along the continental slope of the north-western Mediterranean Sea are characterized by significant decreases in organic matter content (up to 52%), slower organic carbon turnover (ca. 37%), and reduced meiofauna abundance (80%), biodiversity (50%), and nematode species richness (25%). We estimate that the organic carbon removed daily by trawling in the region under scrutiny represents as much as 60-100% of the input flux. We anticipate that such an impact is causing the degradation of deep-sea sedimentary habitats and an infaunal depauperation. With deep-sea trawling currently conducted along most continental margins, we conclude that trawling represents a major threat to the deep seafloor ecosystem at the global scale.

Effects of Local Acidification on Benthic Communities at Shallow Hydrothermal Vents of the Aeolian Islands (Southern Tyrrhenian, Mediterranean Sea).

The Aeolian Islands (Mediterranean Sea) host a unique hydrothermal system called the "Smoking Land" due to the presence of over 200 volcanic CO2-vents, resulting in water acidification phenomena and the creation of an acidified benthic environment. Here, we report the results of a study conducted at three sites located at ca. 16, 40, and 80 m of depth, and characterized by CO2 emissions to assess the effects of acidification on meio- and macrobenthic assemblages. Acidification caused significant changes in both meio- and macrofaunal assemblages, with a clear decrease in terms of abundance and a shift in community composition. A noticeable reduction in biomass was observed only for macrofauna. The most sensitive meiofaunal taxa were kinorhynchs and turbellarians that disappeared at the CO2 sites, while the abundance of halacarids and ostracods increased, possibly as a result of the larger food availability and the lower predatory pressures by the sensitive meiofaunal and macrofaunal taxa. Sediment acidification also causes the disappearance of more sensitive macrofaunal taxa, such as gastropods, and the increase in tolerant taxa such as oligochaetes. We conclude that the effects of shallow CO2-vents result in the progressive simplification of community structure and biodiversity loss due to the disappearance of the most sensitive meio- and macrofaunal taxa.

Impairment of microbial and meiofaunal ecosystem functions linked to algal forest loss.

Habitat loss is jeopardizing marine biodiversity. In the Mediterranean Sea, the algal forests of Cystoseira spp. form one of the most complex, productive and vulnerable shallow-water habitats. These forests are rapidly regressing with negative impact on the associated biodiversity, and potential consequences in terms of ecosystem functioning. Here, by comparing healthy Cystoseira forests and barren grounds (i.e., habitats where the macroalgal forests disappeared), we assessed the effects of habitat loss on meiofaunal and nematode biodiversity, and on some ecosystem functions (here measured in terms of prokaryotic and meiofaunal biomass). Overall, our results suggest that the loss of Cystoseira forests and the consequent barren formation is associated with the loss of meiofaunal higher taxa and a decrease of nematode biodiversity, leading to the collapse of the microbial and meiofaunal variables of ecosystem functions. We conclude that, given the very limited resilience of these ecosystems, active restoration of these vulnerable habitats is needed, in order to recover their biodiversity, ecosystem functions and associated services.

Potentially combined effect of the invasive seaweed Caulerpa cylindracea (Sonder) and sediment deposition rates on organic matter and meiofaunal assemblages.

The seaweed Caulerpa cylindracea (Sonder) is one of the most successful marine bioinvaders worldwide. Caulerpa cylindracea can influence the quantity and biochemical composition of sedimentary organic matter (OM). However, it is still unknown if the effects of C. cylindracea on both OM and small metazoans (i.e. meiofauna) can change according to different sediment deposition rates. To provide insights on this, we investigated the biochemical composition of sediments along with the abundance and composition of meiofaunal assemblages in sediments colonized and not-colonized by the seaweed C. cylindracea under different regimes of sediment deposition. Our results show that the presence of the invasive alga C. cylindracea could alter quantity, biochemical composition, and nutritional quality of organic detritus and influence the overall functioning of the benthic system, but also that the observed effects could be context-dependent. In particular, we show that the presence of C. cylindracea could have a positive effect on meiofaunal abundance wherever the sediment deposition rates are low, whereas the contextual presence of high to medium sedimentation rates can provoke an accumulation of sedimentary organic matter, less favourable bioavailability of food for the benthos, and consequent negative effects on meiofauna.

Denitrification, Nitrogen Uptake, and Organic Matter Quality Undergo Different Seasonality in Sandy and Muddy Sediments of a Turbid Estuary.

The interaction between microbial communities and benthic algae as nitrogen (N) regulators in poorly illuminated sediments is scarcely investigated in the literature. The role of sediments as sources or sinks of N was analyzed in spring and summer in sandy and muddy sediments in a turbid freshwater estuary, the Curonian Lagoon, Lithuania. Seasonality in this ecosystem is strongly marked by phytoplankton community succession with diatoms dominating in spring and cyanobacteria dominating in summer. Fluxes of dissolved gas and inorganic N and rates of denitrification of water column nitrate (Dw) and of nitrate produced by nitrification (Dn) and sedimentary features, including the macromolecular quality of organic matter (OM), were measured. Shallow/sandy sites had benthic diatoms, while at deep/muddy sites, settled pelagic microalgae were found. The OM in surface sediments was always higher at muddy than at sandy sites, and biochemical analyses revealed that at muddy sites the OM nutritional value changed seasonally. In spring, sandy sediments were net autotrophic and retained N, while muddy sediments were net heterotrophic and displayed higher rates of denitrification, mostly sustained by Dw. In summer, benthic oxygen demand increased dramatically, whereas denitrification, mostly sustained by Dn, decreased in muddy and remained unchanged in sandy sediments. The ratio between denitrification and oxygen demand was significantly lower in sandy compared with muddy sediments and in summer compared with spring. Muddy sediments displayed seasonally distinct biochemical composition with a larger fraction of lipids coinciding with cyanobacteria blooms and a seasonal switch from inorganic N sink to source. Sandy sediments had similar composition in both seasons and retained inorganic N also in summer. Nitrogen uptake by microphytobenthos at sandy sites always exceeded the amount loss via denitrification, and benthic diatoms appeared to inhibit denitrification, even in the dark and under conditions of elevated N availability. In spring, denitrification attenuated N delivery from the estuary to the coastal area by nearly 35%. In summer, denitrification was comparable (~100%) with the much lower N export from the watershed, but N loss was probably offset by large rates of N-fixation.

Trophic status and meiofauna biodiversity in the Northern Adriatic Sea: Insights for the assessment of good environmental status.

The Descriptor 5 (Eutrophication) of the EU Marine Strategy Framework Directive aims at preventing the negative effects of eutrophication. However, in coastal systems all indicators based on water column parameters fail in identifying the trophic status and its effects on biodiversity and ecosystem functioning. We investigated benthic trophic status, in terms of sedimentary organic matter quantity, composition and quality, along with meiofaunal abundance, richness of taxa and community composition in three coastal sites (N Adriatic Sea) affected by different levels of anthropogenic stressors. We show that, on the basis of organic matter quantity and composition, the investigated areas can be classified from oligo-to mesotrophic, whereas using meiofauna as a descriptor, their environmental quality ranged from sufficient to moderately impacted. Our results show that the benthic trophic status based on organic matter variables, is not sufficient to provide a sound assessment of the environmental quality in marine coastal ecosystems. However, data reported here indicate that the integration of the meiofaunal variable allows providing robust assessments of the marine environmental status.

Biodiversity loss and turnover in alternative states in the Mediterranean Sea: a case study on meiofauna.

In the Mediterranean Sea hard-bottom macroalgal meadows may switch to alternative and less-productive barrens grounds, as a result of sea urchins overgrazing. Meiofauna (and especially nematodes) represent key components of benthic ecosystems, are highly-diversified, sensitive to environmental change and anthropogenic impacts, but, so-far, have been neglected in studies on regime shifts. We report here that sedimentary organic matter contents, meiofaunal taxa richness and community composition, nematode α- and ß-biodiversity vary significantly between alternative macroalgal and barren states. The observed differences are consistent in six areas spread across the Mediterranean Sea, irrespective of barren extent. Our results suggest also that the low biodiversity levels in barren states are the result of habitat loss/fragmentation, which is associated also with a lower availability of trophic resources. Furthermore, differences in meiofaunal and nematode abundance, biomass and diversity between macroalgal meadow and barren states persist when the latter is not fully formed, or consists of patches interspersed in macroalgal meadows. Since barren grounds are expanding rapidly along the Mediterranean Sea and meiofauna are a key trophic component in marine ecosystems, we suggest that the extension and persistence of barrens at the expenses of macroalgal meadows could also affect resilience of higher trophic level.

High Meiofaunal and Nematodes Diversity around Mesophotic Coral Oases in the Mediterranean Sea.

Although the mesophotic zone of the Mediterranean Sea has been poorly investigated, there is an increasing awareness about its ecological importance for its biodiversity, as fish nursery and for the recruitment of shallow water species. Along with coastal rocky cliffs, isolated coralligenous concretions emerging from muddy bottoms are typical structures of the Mediterranean Sea mesophotic zone. Coralligenous concretions at mesophotic depths in the South Tyrrhenian Sea were investigated to assess the role of these coralligenous oases in relation to the biodiversity of surrounding soft sediments. We show here that the complex structures of the coralligenous concretions at ca. 110 m depth influence the trophic conditions, the biodiversity and assemblage composition in the surrounding sediments even at considerable distances. Coral concretions not only represent deep oases of coral biodiversity but they also promote a higher biodiversity of the fauna inhabiting the surrounding soft sediments. Using the biodiversity of nematodes as a proxy of the total benthic biodiversity, a high turnover biodiversity within a 200 m distance from the coralligenous concretions was observed. Such turnover is even more evident when only rare taxa are considered and seems related to specific trophic conditions, which are influenced by the presence of the coralligenous structures. The presence of a high topographic complexity and the trophic enrichment make these habitats highly biodiverse, nowadays endangered by human activities (such as exploitation of commercial species such as Corallium rubrum, or trawling fisheries, which directly causes habitat destruction or indirectly causes modification in the sedimentation and re-suspension rates). We stress that the protection of the coralligenous sea concretions is a priority for future conservation policies at the scale of large marine ecosystems and that a complete census of these mesophotic oases of biodiversity should be a priority for future investigations in the Mediterranean Sea.

Red coral extinction risk enhanced by ocean acidification.

The red coral Corallium rubrum is a habitat-forming species with a prominent and structural role in mesophotic habitats, which sustains biodiversity hotspots. This precious coral is threatened by both over-exploitation and temperature driven mass mortality events. We report here that biocalcification, growth rates and polyps' (feeding) activity of Corallium rubrum are significantly reduced at pCO2 scenarios predicted for the end of this century (0.2 pH decrease). Since C. rubrum is a long-living species (>200 years), our results suggest that ocean acidification predicted for 2100 will significantly increases the risk of extinction of present populations. Given the functional role of these corals in the mesophotic zone, we predict that ocean acidification might have cascading effects on the functioning of these habitats worldwide.

Fish-farm impact on metazoan meiofauna in the Mediterranean Sea: Analysis of regional vs. habitat effects.

The worldwide exponential growth of off-shore mariculture is raising severe concerns about the impacts of this industry on marine habitats and their biodiversity. We investigated the metazoan meiofaunal response to fish-farm impact in four regions of the Mediterranean Sea. Meiofaunal assemblages were investigated in two habitats (seagrass meadows of Posidonia oceanica and non-vegetated soft bottoms) comparing sites receiving faeces and uneaten food pellets from fish farms to control sites. We report here that, consistently across different regions, the meiofaunal abundance typically responded positively to fish-farm effluents. Biodeposition caused also significant changes in assemblage structure and the reduction in the richness of higher meiofaunal taxa, but the multivariate analysis of variance revealed that the effects were region- and habitat-specific. In non-vegetated systems, three of the four regions investigated displayed significant effects of the fish farms on richness of meiofaunal taxa. In vegetated habitats, meiofauna did not respond to biodeposition (except in one region), suggesting that seagrass meadows can mask the effects of fish-farm effluents on benthic biodiversity. We conclude that different indicators of fish-farm impact are needed in vegetated and non-vegetated benthic systems.