Mediterranean Lithophyllum stictiforme (Corallinales, Rhodophyta) is a genetically diverse species complex: implications for species circumscription, biogeography and conservation of coralligenous habitats.

Lithophyllum species in the Mediterranean Sea function as algal bioconstructors, contributing to the formation of biogenic habitats such as coralligenous concretions. In such habitats, thalli of Lithophyllum, consisting of crusts or lamellae with entire or lobed margins, have been variously referred to as either one species, L. stictiforme, or two species, L. stictiforme and L. cabiochiae, in the recent literature. We investigated species diversity and phylogenetic relationships in these algae by sequencing three markers (psbA and rbcL genes, cox2,3 spacer), in conjunction with methods for algorithmic delimitation of species (ABGD and GMYC). Mediterranean subtidal Lithophyllum belong to a well-supported lineage, hereby called the L. stictiforme complex, which also includes two species described from the Atlantic, L. lobatum and L. searlesii. Our results indicate that the L. stictiforme complex consists of at least 13 species. Among the Mediterranean species, some are widely distributed and span most of the western and central Mediterranean, whereas others appear to be restricted to specific localities. These patterns are interpreted as possibly resulting from allopatric speciation events that took place during the Messinian Salinity Crisis and subsequent glacial periods. A partial rbcL sequence from the lectotype of L. stictiforme unambiguously indicates that this name applies to the most common subtidal Lithophyllum in the central Mediterranean. We agree with recent treatments that considered L. cabiochiae and L. stictiforme conspecific. The diversity of Lithophyllum in Mediterranean coralligenous habitats has been substantially underestimated, and future work on these and other Mediterranean corallines should use identifications based on DNA sequences.

Physical effects of habitat-forming species override latitudinal trends in temperature.

Latitudinal and elevational temperature gradients (LTG and ETG) play central roles in biogeographical theory, underpinning predictions of large-scale patterns in organismal thermal stress, species' ranges and distributional responses to climate change. Yet an enormous fraction of Earth's taxa live exclusively in habitats where foundation species modify temperatures. We examine little-explored implications of this widespread trend using a classic model system for understanding heat stresses - rocky intertidal shores. Through integrated field measurements and laboratory trials, we demonstrate that thermal buffering by centimetre-thick mussel and seaweed beds eliminates differences in stress-inducing high temperatures and associated mortality risk that would otherwise arise over 14° of latitude and ~ 1 m of shore elevation. These results reveal the extent to which physical effects of habitat-formers can overwhelm broad-scale thermal trends, suggesting a need to re-evaluate climate change predictions for many species. Notably, inhabitant populations may exhibit deceptive resilience to warming until refuge-forming taxa become imperiled.

Disentangling the effect of space, time, and environmental and anthropogenic drivers on coastal macrobenthic ß diversity in contrasting habitats over 15 years.

Coastal zones are biodiversity hotspots and deliver essential ecosystem functions and services, yet they are exposed to multiple and interacting anthropogenic and environmental constraints. The individual and cumulative effects of these constraints on benthic communities, a key component of coastal ecosystems, and their variability across space and time, remains to be thoroughly quantified to guide conservation actions. Here, we explored how the presence of biogenic habitats influences the response of benthic communities to natural and anthropogenic constraints. We investigated this effect in both intertidal and subtidal habitats exposed to different pressures. We used data collected in the North-East Atlantic over 15 years (2005-2019) as part of the REBENT monitoring program, covering 38 sites of bare sediments, intertidal seagrass beds and maerl beds. We collected a range of environmental variables and proxies of anthropogenic pressures and used variation and hierarchical partitioning with redundancy analyses to estimate their relative effect on macrobenthic communities. We used descriptors modeling spatial and temporal structures (dbMEMs) to explore the scale of their effects and potential missing predictors. The selected variables explained between 53 % and 64 % of macrobenthic ß diversity depending on habitat and depth. Fishing pressures, sedimentary and hydrodynamics variables stood out as the most important predictors across all habitats while proxies of anthropogenic pressures were overall more important in intertidal habitats. In the intertidal, presence of biogenic habitat strongly modulated the amount of explained variance and the identity of the selected variable. Across both tidal levels, analysis of models' residuals further indicated that biogenic habitats might mitigate the effect of extreme environmental events. Our study provides a hierarchy of the most important drivers of benthic communities across different habitats and tidal levels, emphasizing the prominence of anthropogenic pressures on intertidal communities and the role of biogenic habitats in mitigating environmental changes.

Identifying potential high-risk zones for land-derived plastic litter to marine megafauna and key habitats within the North Atlantic.

The pervasive use of plastic in modern society has led to plastic litter becoming ubiquitous within the ocean. Land-based sources of plastic litter are thought to account for the majority of plastic pollution in the marine environment, with plastic bags, bottles, wrappers, food containers and cutlery among the most common items found. In the marine environment, plastic is a transboundary pollutant, with the potential to cause damage far beyond the political borders from where it originated, making the management of this global pollutant particularly complex. In this study, the risks of land-derived plastic litter (LDPL) to major groups of marine megafauna - seabirds, cetaceans, pinnipeds, elasmobranchs, turtles, sirenians, tuna and billfish - and a selection of productive and biodiverse biogenic habitats - coral reefs, mangroves, seagrass, saltmarsh and kelp beds - were analysed using a Spatial Risk Assessment approach. The approach combines metrics for vulnerability (mechanism of harm for megafauna group or habitat), hazard (plastic abundance) and exposure (distribution of group or habitat). Several potential high-risk zones (HRZs) across the North Atlantic were highlighted, including the Azores, the UK, the French and US Atlantic coasts, and the US Gulf of Mexico. Whilst much of the modelled LDPL driving risk in the UK originated from domestic sources, in other HRZs, such as the Azores archipelago and the US Gulf of Mexico, plastic originated almost exclusively from external (non-domestic) sources. LDPL from Caribbean islands - some of the largest generators of marine plastic pollution in the dataset of river plastic emissions used in the study - was noted as a significant input to HRZs across both sides of the Atlantic. These findings highlight the potential of Spatial Risk Assessment analyses to determine the location of HRZs and understand where plastic debris monitoring and management should be prioritised, enabling more efficient deployment of interventions and mitigation measures.

Distribution and Characterization of Deep Rhodolith Beds off the Campania coast (SW Italy, Mediterranean Sea).

Rhodolith beds (RBs) are bioconstructions characterized by coralline algae, which provide habitat for several associated species. Mediterranean RBs are usually located in the mesophotic zone (below 40 m), and thus are frequently remote and unexplored. Recently, the importance and vulnerability of these habitats have been recognized by the European Community and more attention has been drawn to their investigation and conservation. This study reports the results of an extensive monitoring program, carried out within the Marine Strategy Framework Directive (2008/56/EC), in six sites off the Campania coast (Italy, Mediterranean Sea). New insights were given into the distribution, cover, vitality (i.e., live/dead rhodolith ratio), structural complexity, and coralline algae composition of RBs. Remotely operated vehicles (ROV) investigations allowed the description of several RBs, and the discovery of a RB with rhodolith cover >65% offshore the Capri Island. Only two sites (Secchitiello and Punta Campanella) showed a very low mean cover of live rhodoliths (<10%); hence, not being classifiable as RBs. The collected rhodoliths were mostly small pralines (~2 cm), spheroidal to ellipsoidal, with growth-forms ranging from encrusting/warty to fruticose/lumpy. Coralline algae identification revealed a high diversity within each bed, with a total of 13 identified taxa. The genus Lithothamnion dominated all sites, and Phymatolithon calcareum and Lithothamnion corallioides, protected by the Habitats Directive (92/43/EEC), were detected in all RBs.

Sublethal effects of contaminants on marine habitat-forming species: a review and meta-analysis.

Contaminants may affect ecosystem functioning by reducing the fitness of organisms and these impacts may cascade through ecosystems, particularly if the sensitive organisms are also habitat-forming species. Understanding how sub-lethal effects of toxicants can affect the quality and functions of biogenic habitats is critical if we are to establish effective guidelines for protecting ecosystems. We carried out a global systematic review and meta-analysis critically evaluating contaminant effects on properties of habitat-formers linked to ecosystem functioning. We reviewed a total of 95 publications. However, 40% of publications initially captured by the literature search were identified as having flaws in experimental design and ~11% did not present results in an appropriate way and thus were excluded from the quantitative meta-analysis. We quantitatively reviewed 410 studies from 46 publications, of which 313 (~76%) were on plants and seaweeds, that is macro-algae, saltmarsh plants and seagrasses, 58 (~14%) studied corals and 39 (~10%) looked at toxicant impacts on bivalves, with 70% of those on mussels and the remaining studies on oysters. Response variables analysed were photosynthetic efficiency, amount of chlorophyll a (as a proxy for primary production) and growth of plants, seaweeds and corals as well as leaf area of plants. We also analysed filtration, growth and respiration rates of bivalves. Our meta-analysis found that chemical contaminants have a significant negative impact on most of the analysed functional variables, with the exception of the amount of chlorophyll a. Metals were the most widely harmful type of contaminant, significantly decreasing photosynthetic efficiency of kelps, leaf area of saltmarsh plants, growth of fucoids, corals and saltmarsh plants and the filtration rates of bivalves. Organic contaminants decreased the photosynthetic efficiency of seagrass, but had no significant effects on bivalve filtration. We did not find significant effects of polycyclic aromatic hydrocarbons on any of the analysed functional variables or habitat-forming taxa, but this could be due to the low number of studies available. A meta-regression revealed that relationships between concentrations of metal contaminants and the magnitude of functional responses varied with the type of metal and habitat-former. Increasing concentrations of contaminants significantly increased the negative effects on the photosynthetic efficiency of habitat-formers. There was, however, no apparent relationship between ecologically relevant concentrations of metals and effect sizes of photosynthetic efficiency of corals and seaweeds. A qualitative analysis of all relevant studies found slightly different patterns when compared to our quantitative analysis, emphasising the need for studies to meet critical inclusion criteria for meta-analyses. Our study highlights links between effects of contaminants at lower levels of organisation (i.e. at the biochemical and/or physiological level of individuals) and ecological, large-scale impacts, through effects on habitat-forming species. Contaminants can clearly reduce the functioning of many habitat-forming marine species. We therefore recommend the adoption of routine measures of functional endpoints in monitoring and conservation programs to complement structural measures.

The multiple roles of ß-diversity help untangle community assembly processes affecting recovery of temperate rocky shores.

Metacommunity theory highlights the potential of ß-diversity as a useful link to empirical research, especially in diverse systems where species exhibit a range of stage-dependent dispersal characteristics. To investigate the importance of different components and scales of ß-diversity in community assembly, we conducted a large-scale disturbance experiment and compared relative recovery across multiple sites and among plots within sites on the rocky shore. Six sites were spread along 80 km of coastline and, at each site, five plots were established, matching disturbed and undisturbed quadrats. Recovery was not complete at any of the sites after 1 year for either epibenthos (mostly composed of macroalgae and, locally, mussels) or infauna. Significant differences in recovery among sites were observed for epibenthos but not for infauna, suggesting that different community assembly processes were operating. This was supported by epibenthos in the recovering plots having higher species turnover than in undisturbed sediment, and recovery well predicted by local diversity, while infaunal recovery was strongly influenced by the epibenthic community's habitat complexity. However, infaunal community recovery did not simply track formation of habitat by recovering epibenthos, but appeared to be overlain by within-site and among-site aspects of infaunal ß-diversity. These results suggest that documenting changes in the large plants and animals alone will be a poor surrogate for rocky shore community assembly processes. No role for ecological connectivity (negative effect of among-site ß-diversity) in driving recovery was observed, suggesting a low risk of effects from multiple disturbances propagating along the coast, but a limited resilience at the site scale to large-scale disturbances such as landslides or oil spills.