Role of macroalgal forests within Mediterranean shallow bays in blue carbon storage.

Although seaweeds rank among the most productive vegetated habitats globally, their inclusion within Blue Carbon frameworks is at its onset, partially because they usually grow in rocky substrates and their organic carbon (Corg) is mostly exported and stored beyond their habitat and thus, demonstrating its long-term storage is challenging. Here, we studied the sedimentary Corg storage in macroalgal forests dominated by Gongolaria barbata and in adjacent seagrass Cymodocea nodosa mixed with Caulerpa prolifera algae meadows, and bare sand habitats in Mediterranean shallow coastal embayments. We characterized the biogeochemistry of top 30 cm sedimentary deposits, including sediment grain-size, organic matter and Corg contents, Corg burial rates and the provenance of sedimentary Corg throughout stable carbon isotopes (δ13Corg) and pyrolysis analyses. Sediment Corg stocks and burial rates (since 1950) in G. barbata forests (mean ± SE, 3.5 ± 0.2 kg Corg m-2 accumulated at 15.5 ± 1.6 g Corg m-2 y-1) fall within the range of those reported for traditional Blue Carbon Ecosystems. Although the main species contributing to sedimentary Corg stocks in all vegetated habitats examined was C. nodosa (36 ± 2 %), macroalgae contributed 49 % (19 ± 2 % by G. barbata and 30 ± 3 % by C. prolifera) based on isotope mixing model results. Analytical pyrolysis confirmed the presence of macroalgae-derived compounds in the sediments, including N-compounds and α-tocopherol linked to G. barbata and C. prolifera, respectively. The sedimentary Corg burial rate linked to macroalgae within the macroalgal forests examined ranged from 5.4 to 9.5 g Corg m-2 y-1 (7.4 ± 2 g Corg m-2 y-1). This study provides empirical evidence for the long-term (∼70 years) sequestration of macroalgae-derived Corg within and beyond seaweed forests in Mediterranean shallow coastal embayments and thereby, supports the inclusion of macroalgae in Blue Carbon frameworks.

Multiple scale assessment of habitat, landscape, and geographic-specific attributes driving decapod assemblages in Posidonia oceanica seagrass meadows.

Seagrass meadows are biodiversity hotspots for invertebrate species including decapods. Understanding the drivers of species abundance, richness and diversity of decapod assemblages is crucial for the conservation of such hotspots, but how drivers act across multiple spatial scales remains unexplored. Here we describe the decapod assemblages of Posidonia oceanica seagrass meadows and assess the influence of attributes from three increasing spatial scales (habitat, landscape, and geographical levels) on the assemblages' structure and composition, as well as the variability partitioning among each one of these levels. Overall, geographical level attributes (i.e., inlet aperture, confinement) affected the most the decapod assemblages, while we only found a modest contribution from habitat (e.g., detritus biomass, sediment organic matter) and landscape attributes (e.g., fragmentation). We suggest that decapod assemblages are driven by the interaction of multiple processes occurring at different scales and other highly stochastic phenomena such as larval dispersion and recruitment.

Optimizing marine macrophyte capacity to locally ameliorate ocean acidification under variable light and flow regimes: Insights from an experimental approach.

The urgent need to remediate ocean acidification has brought attention to the ability of marine macrophytes (seagrasses and seaweeds) to take up carbon dioxide (CO2) and locally raise seawater pH via primary production. This physiological process may represent a powerful ocean acidification mitigation tool in coastal areas. However, highly variable nearshore environmental conditions pose uncertainty in the extent of the amelioration effect. We developed experiments in aquaria to address two interconnected goals. First, we explored the individual capacities of four species of marine macrophytes (Ulva lactuca, Zostera marina, Fucus vesiculosus and Saccharina latissima) to ameliorate seawater acidity in experimentally elevated pCO2. Second, we used the most responsive species (i.e., S. latissima) to assess the effects of high and low water residence time on the amelioration of seawater acidity in ambient and simulated future scenarios of climate change across a gradient of irradiance. We measured changes in dissolved oxygen, pH, and total alkalinity, and derived resultant changes to dissolved inorganic carbon (DIC) and calcium carbonate saturation state (Ω). While all species increased productivity under elevated CO2, S. latissima was able to remove DIC and alter pH and Ω more substantially as CO2 increased. Additionally, the amelioration of seawater acidity by S. latissima was optimized under high irradiance and high residence time. However, the influence of water residence time was insignificant under future scenarios. Finally, we applied predictive models as a function of macrophyte biomass, irradiance, and residence time conditions in ambient and future climatic scenarios to allow projections at the ecosystem level. This research contributes to understanding the biological and physical drivers of the coastal CO2 system.

Exploring coexistence mechanisms in a three-species assemblage.

Interactions among species are essential in shaping ecological communities, although it is not always clear under what conditions they can persist when the number of species involved is higher than two. Here we describe a three-species assemblage involving the seagrass Cymodocea nodosa, the pen shell Pinna nobilis and the herbivore sea urchin Paracentrotus lividus, and we explore the mechanisms allowing its persistence through field observations and manipulative experiments. The abundance of pen shells was higher in seagrass beds than in bare sand, suggesting a recruitment facilitation. The presence of sea urchins, almost exclusively attached or around pen shells, indicated habitat facilitation for sea urchins, which overgrazed the meadow around the pen shells forming seagrass-free halos. Our results suggest that this system persists thanks to: (i) the behavioral reluctance of sea urchins to move far from pen shells, making their impact on seagrass strictly local, (ii) the sparse distribution of pen shells and (iii) the plant's resistance mechanisms to herbivory. Unpacking these mechanisms allows a better understanding of how ecological communities are assembled.

Seagrass-driven changes in carbonate chemistry enhance oyster shell growth.

Quantifying the strength of non-trophic interactions exerted by foundation species is critical to understanding how natural communities respond to environmental stress. In the case of ocean acidification (OA), submerged marine macrophytes, such as seagrasses, may create local areas of elevated pH due to their capacity to sequester dissolved inorganic carbon through photosynthesis. However, although seagrasses may increase seawater pH during the day, they can also decrease pH at night due to respiration. Therefore, it remains unclear how consequences of such diel fluctuations may unfold for organisms vulnerable to OA. We established mesocosms containing different levels of seagrass biomass (Zostera marina) to create a gradient of carbonate chemistry conditions and explored consequences for growth of juvenile and adult oysters (Crassostrea gigas), a non-native species widely used in aquaculture that can co-occur, and is often grown, in proximity to seagrass beds. In particular, we investigated whether increased diel fluctuations in pH due to seagrass metabolism affected oyster growth. Seagrasses increased daytime pH up to 0.4 units but had little effect on nighttime pH (reductions less than 0.02 units). Thus, both the average pH and the amplitude of diel pH fluctuations increased with greater seagrass biomass. The highest seagrass biomass increased oyster shell growth rate (mm day-1) up to 40%. Oyster somatic tissue weight and oyster condition index exhibited a different pattern, peaking at intermediate levels of seagrass biomass. This work demonstrates the ability of seagrasses to facilitate oyster calcification and illustrates how non-trophic metabolic interactions can modulate effects of environmental change.

Coast-wide evidence of low pH amelioration by seagrass ecosystems.

Global-scale ocean acidification has spurred interest in the capacity of seagrass ecosystems to increase seawater pH within crucial shoreline habitats through photosynthetic activity. However, the dynamic variability of the coastal carbonate system has impeded generalization into whether seagrass aerobic metabolism ameliorates low pH on physiologically and ecologically relevant timescales. Here we present results of the most extensive study to date of pH modulation by seagrasses, spanning seven meadows (Zostera marina) and 1000 km of U.S. west coast over 6 years. Amelioration by seagrass ecosystems compared to non-vegetated areas occurred 65% of the time (mean increase 0.07 ± 0.008 SE). Events of continuous elevation in pH within seagrass ecosystems, indicating amelioration of low pH, were longer and of greater magnitude than opposing cases of reduced pH or exacerbation. Sustained elevations in pH of >0.1, comparable to a 30% decrease in [H+ ], were not restricted only to daylight hours but instead persisted for up to 21 days. Maximal pH elevations occurred in spring and summer during the seagrass growth season, with a tendency for stronger effects in higher latitude meadows. These results indicate that seagrass meadows can locally alleviate low pH conditions for extended periods of time with important implications for the conservation and management of coastal ecosystems.

High variability of Blue Carbon storage in seagrass meadows at the estuary scale.

Seagrass meadows are considered important natural carbon sinks due to their capacity to store organic carbon (Corg) in sediments. However, the spatial heterogeneity of carbon storage in seagrass sediments needs to be better understood to improve accuracy of Blue Carbon assessments, particularly when strong gradients are present. We performed an intensive coring study within a sub-tropical estuary to assess the spatial variability in sedimentary Corg associated with seagrasses, and to identify the key factors promoting this variability. We found a strong spatial pattern within the estuary, from 52.16 mg Corg cm-3 in seagrass meadows in the upper parts, declining to 1.06 mg Corg cm-3 in seagrass meadows at the estuary mouth, despite a general gradient of increasing seagrass cover and seagrass habitat extent in the opposite direction. The sedimentary Corg underneath seagrass meadows came principally from allochthonous (non-seagrass) sources (~70-90 %), while the contribution of seagrasses was low (~10-30 %) throughout the entire estuary. Our results showed that Corg stored in sediments of seagrass meadows can be highly variable within an estuary, attributed largely to accumulation of fine sediments and inputs of allochthonous sources. Local features and the existence of spatial gradients must be considered in Blue Carbon estimates in coastal ecosystems.

Multilevel assessments reveal spatially scaled landscape patterns driving coastal fish assemblages.

Ecological research, particularly in marine environments, tends to focus on single habitats and often single spatial scales, and thus not account for ecological processes operating at multiple spatial scales. Here we aim to explore how coastal fish assemblages are influenced by landscape patterns integrating multiple spatial scales, to assess the strength of these associations and to identify the most relevant spatial scales at which these associations occur. We use a multiscale approach through multilevel modelling to evaluate the association of landscape metrics with fish assemblages, at three nested spatial scales, in temperate coastal seascapes composed of seagrass meadows, sandy bottoms and rocky reefs. Landscape composition metrics, expressed as cover of vegetated habitats, significantly influenced fish assemblages at small (metres) and intermediate (hundred of metres) scales, while landscape configuration metrics did it at all three scales assessed (from metres to kilometres). Species richness was only influenced by small scale landscape patterns (cover of rocky reefs, positive association), whereas total abundance was associated with landscape patterns measured at small and intermediate scales, encompassing metrics associated with landscape composition (rocky reef cover, positive association), and those indicating landscape heterogeneity (negative association). Similarly, the abundances of different functional groups were influenced by metrics consistent with their mobility and their ecological and behavioural traits at all the spatial scales assessed. These results show how landscape patterns influence coastal fish assemblages, and particularly show that spatially scaled landscape patterns, measured in complex ecological systems as a whole, act simultaneously but not always equally on species assemblages.

Long-term shifts in the north western Mediterranean coastal seascape: The habitat-forming seaweed Codium vermilara.

Long-term ecological studies are crucial to understand how and why natural ecosystems change over time and space. Through a revision of historical data and a comparison with current in situ field data, we contribute to the understanding of how the Mediterranean coastal seascape has changed in the last decades. Here we describe the large decrease of the main habitat-forming species Codium vermilara along the Catalan coast (NW Mediterranean). We have analyzed data on presence/absence, abundance and biomass. Since the 70s-80s, when the species reached its highest abundances, the species has totally disappeared from 45% of the revisited sites, and showed a decrease in 95% of its abundance and 97% of its biomass. Codium vermilara has also shown a reduction in its depth range, from 30 to the first 20m depth. This study highlights the importance of having historical data to detect and describe changes in ecological systems.

Variability of sedimentary organic carbon in patchy seagrass landscapes.

Seagrass ecosystems, considered among the most efficient carbon sinks worldwide, encompass a wide variety of spatial configurations in the coastal landscape. Here we evaluated the influence of the spatial configuration of seagrass meadows at small scales (metres) on carbon storage in seagrass sediments. We intensively sampled carbon stocks and other geochemical properties (δ(13)C, particle size, depositional fluxes) across seagrass-sand edges in a Zostera muelleri patchy seagrass landscape. Carbon stocks were significantly higher (ca. 20%) inside seagrass patches than at seagrass-sand edges and bare sediments. Deposition was similar among all positions and most of the carbon was from allochthonous sources. Patch level attributes (e.g. edge distance) represent important determinants of the spatial heterogeneity of carbon stocks within seagrass ecosystems. Our findings indicate that carbon stocks of seagrass areas have likely been overestimated by not considering the influence of meadow landscapes, and have important relevance for the design of seagrass carbon stock assessments.