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Seagrass and macroalgae beds are key blue carbon ecosystems in the ocean. However, coastal development and climate change are sparking a growing concern about the vulnerability of sediment organic carbon (OC) to remineralization after macrophyte perturbation. Thus, the aim of this study was to assess the potential of long-term remineralization of sediment OC stocks (1 year) in coastal vegetated habitats (i.e., seagrasses Zostera noltei and Cymodocea nodosa, macroalgae Caulerpa prolifera and unvegetated sediment) after complete disturbance of macrophyte meadows under conducive conditions to microorganisms growth (i.e., oxygen saturated, non-nutrient limitation, turbulence and dark). Leached dissolved organic carbon (DOC) from particulate organic carbon (POC) remineralization, carbonate dissolution and photo-reactivity of long-term persistent DOC were also evaluated. Our results evidenced that, sediment OC from Z. noltei and unvegetated habitats were entirely remineralized to CO2. However, sediment OC from C. nodosa and C. prolifera communities exhibited a significant fraction of recalcitrant OC, and therefore, a 42 and 46% of the sediment OC still remained after 1 year of culture, respectively. POC remineralization released relevant amounts of both labile and recalcitrant DOC, which showed low photo-reactivity. Finally, we discuss that the main argument to promote management, monitoring, and restoration programs for macrophytes is usually based on their sediment OC deposit, which favor larger species. The study presented here adds arguments to also include small macrophyte species, since their sediment OC may be highly labile and entirely remineralized to CO2 once these habitats are disturbed.

Seagrasses form one of the most productive and threatened ecosystems worldwide because of global change and anthropogenic pressures. The frequency of extreme climatic events, such as heat waves, are expected to increase and may drive even more adverse effects than gradual warming. This study explores for the first time the effects of a sudden and temporary increase of temperature in situ on carbon metabolism and dissolved organic carbon (DOC) fluxes in a community dominated by a seagrass (Cymodocea nodosa) during two contrasting seasons (winter and summer). Results showed a positive correlation between temperature and seagrass production between seasons, while the experimental sudden and temporary increase in water temperature did not produce significant differences in carbon community metabolism and DOC fluxes in winter. In contrast, high temperature conditions in summer enhanced significantly the net community production and affected positively to DOC fluxes. Hence, this study indicates that a sudden and temporary increase in water temperature, which characterize marine heat waves, in temperate areas may enhance the autotrophic metabolism of seagrass communities and can yield an increase in the DOC released, in contrast to previous researches suggesting solely negative effects on seagrasses.

Eutrophication affects seagrasses negatively by increasing light attenuation through stimulation of biomass of fast-growing, bloom-forming algae and because high concentrations of ammonium in the water can be toxic to higher plants. We hypothesized nevertheless, that moderate amounts of nitrophilic macroalgae that coexists with seagrasses under eutrophic conditions, can alleviate the harmful effects of eutrophication on seagrasses by reducing ammonium concentrations in the seawater to non-toxic levels because such algae have a very large capacity to take up inorganic nutrients. We studied therefore how combinations of different ammonium concentrations (0, 25 and 50 μM) and different standing stocks of macroalgae (i.e. 0, 1 and 6 layers of Ulva sp.) affected survival, growth and net production of the seagrass Zostera noltei. In the absence of Ulva sp., increasing ammonium concentrations had a negative influence on the performance of Z. noltei. The presence of Ulva sp. without ammonium supply had a similar, but slightly smaller, negative effect on seagrass fitness due to light attenuation. When ammonium enrichment was combined with presence of Ulva sp., Ulva sp. ameliorated some of negative effects caused by high ammonium availability although Ulva sp. lowered the availability of light. Benthic microalgae, which increased in biomass during the experiment, seemed to play a similar role as Ulva sp.--they contributed to remove ammonium from the water, and thus, aided to keep the ammonium concentrations experienced by Z. noltei at relatively non-toxic levels. Our findings show that moderate amounts of drift macroalgae, eventually combined with increasing stocks of benthic microalgae, may aid seagrasses to alleviate toxic effects of ammonium under eutrophic conditions, which highlights the importance of high functional diversity for ecosystem resistance to anthropogenic disturbance.

Global change, such as warming and ocean acidification, and local anthropogenic disturbances, such as eutrophication, can have profound impacts on marine organisms. However, we are far from being able to predict the outcome of multiple interacting disturbances on seagrass communities. Herbivores are key in determining plant community structure and the transfer of energy up the food web. Global and local disturbances may alter the ecological role of herbivory by modifying leaf palatability (i.e. leaf traits) and consequently, the feeding patterns of herbivores. This study evaluates the main and interactive effects of factors related to global change (i.e. elevated temperature, lower pH levels and associated ocean acidification) and local disturbance (i.e. eutrophication through ammonium enrichment) on a broad spectrum of leaf traits using the temperate seagrass Cymodocea nodosa, including structural, nutritional, biomechanical and chemical traits. The effect of these traits on the consumption rates of the generalist herbivore Paracentrotus lividus (purple sea urchin) is evaluated. The three disturbances of warming, low pH level and eutrophication, alone and in combination, increased the consumption rate of seagrass by modifying all leaf traits. Leaf nutritional quality, measured as nitrogen content, was positively correlated to consumption rate. In contrast, a negative correlation was found between feeding decisions by sea urchins and structural, biomechanical and chemical leaf traits. In addition, a notable accomplishment of this work is the identification of phenolic compounds not previously reported for C. nodosa. Our results suggest that global and local disturbances may trigger a major shift in the herbivory of seagrass communities, with important implications for the resilience of seagrass ecosystems.

Posidonia oceanica is an endemic seagrass species from the Mediterranean Sea that provides critical ecological services to coastal environments. This species is distributed from the Turkish to the Spanish coast, where its westernmost record was documented in Punta Chullera, Malaga (36°18′36.45′′ N, 5°14′54.31′′ W). Nevertheless, previous studies suggested that its distribution was even further west, although these populations were never described. In this study, we documented and characterised the only known P. oceanica population on the coast of Cadiz, in Cala Sardina (36°18′38.80′′ N, 5°15′15.13′′ W). The newly documented population of P. oceanica presented a fragmented structure, consisting of nine patches found in a rocky shallow area surrounded by the invasive algae Rugulopteryx okamurae, with a total size of 61.14 m2. Shoots had a relatively small size (21.0 ± 2.9 cm) in comparison with centrally-distributed populations. The relatively small size of the plants, alongside the observed low shoot density (437 ± 42 shoots m−2) and leaf area index (4.8 ± 0.7 m2 m−2), may indicate that this meadow could be exposed to sub-optimal environmental conditions for plant development. By contrast, the meadow showed relatively high production rates (0.03 ± 0.01 leaf day−1 shoot−1) in comparison with other Mediterranean populations. The percentage of carbon in plant leaves was 38.73 ± 1.38%, while the nitrogen and C/N were 1.38 ± 0.37% and 29.93 ± 6.57, respectively. The documentation of this meadow extends the distribution of this species to the Mediterranean coast of Cadiz, making this region the place with the highest seagrass biodiversity (four species) in the Iberian Peninsula, and potentially in Europe. This exploratory study provides a baseline to examine the potential effects of climate change, anthropogenic disturbances or the presence of invasive species.

Global change has been acknowledged as one of the main threats to the biosphere and its provision of ecosystem services, especially in marine ecosystems. Seagrasses play a critical ecological role in coastal ecosystems, but their responses to ocean acidification (OA) and climate change are not well understood. There have been previous studies focused on the effects of OA, but the outcome of interactions with co-factors predicted to alter during climate change still needs to be addressed. For example, the impact of higher CO2 and different hydrodynamic regimes on seagrass performance remains unknown. We studied the effects of OA under different current velocities on productivity of the seagrass Zostera noltei, using changes in dissolved oxygen as a proxy for the seagrass carbon metabolism, and release of dissolved organic carbon (DOC) in a four-week experiment using an open-water outdoor mesocosm. Under current pH conditions, increasing current velocity had a positive effect on productivity, but this depended on shoot density. However, this positive effect of current velocity disappeared under OA conditions. OA conditions led to a significant increase in gross production rate and respiration, suggesting that Z. noltei is carbon-limited under the current inorganic carbon concentration of seawater. In addition, an increase in non-structural carbohydrates was found, which may lead to better growing conditions and higher resilience in seagrasses subjected to environmental stress. Regarding DOC flux, a direct and positive relationship was found between current velocity and DOC release, both under current pH and OA conditions. We conclude that OA and high current velocity may lead to favourable growth scenarios for Z. noltei populations, increasing their productivity, non-structural carbohydrate concentrations and DOC release. Our results add new dimensions to predictions on how seagrass ecosystems will respond to climate change, with important implications for the resilience and conservation of these threatened ecosystems.

Non-structural carbohydrates (NSCs) are energetic compounds that can be accumulated in tissues and mobilized during periods of unfavorable conditions to maintain the biological functions of plants. The balance of these biochemical compounds is controlled by environmental factors such as temperature and irradiance. Zostera noltei and Zostera marina find one of their southern distribution limits in southern Spain, where relatively high seawater temperatures are reached during summer (23–24 °C). To better understand the effects of elevated temperatures on the concentration of NSCs, we conducted a seasonal study at Cadiz Bay, representing warm-adapted populations of these species. Our results showed a bimodal pattern in both species, with the highest NSC content observed in December and June, followed by a depletion in March and August. In addition, the NSC content observed in the leaves of Z. noltei (71.26 ± 30.77 mg g−1 dry weight) was higher than in the rhizomes and roots (52.14 ± 38.86 mg g−1 DW). The observed patterns suggest that these species accumulated NSCs to cope with periods of unfavorable environmental conditions. We also suggest that the limited concentration of NSCs in Z. noltei rhizomes and roots indicates that this population may be suffering physiological stress.

Seagrass shoots interact with hydrodynamic forces and thereby a positively or negatively influence the survival of associated species. The modification of these forces indirectly alters the physical transport and flux of edible particles within seagrass meadows, which will influence the growth and survivorship of associated filter-feeding organisms. The present work contributes to gaining insight into the mechanisms controlling the availability of resources for filter feeders inhabiting seagrass canopies, both from physical (influenced by seagrass density and patchiness) and biological (regulated by filter feeder density) perspectives. A factorial experiment was conducted in a large racetrack flume, which combined changes in hydrodynamic conditions, chlorophyll a concentration in the water and food intake rate (FIR) in a model active filter-feeding organism (the cockle). Results showed that seagrass density and patchiness modified both hydrodynamic forces and availability of resources for filter feeders. Chlorophyll a water content decreased to 50% of the initial value when densities of both seagrass shoots and cockles were high. Also, filter feeder density controlled resource availability within seagrass patches, depending on its spatial position within the racetrack flume. Under high density of filter-feeding organisms, chlorophyll a levels were lower between patches. This suggests that the pumping activity of cockles (i.e. biomixing) is an emergent key factor affecting both resource availability and FIR for filter feeders in dense canopies. Applying our results to natural conditions, we suggest the existence of a direct correlation between habitat complexity (i.e. shoot density and degree of patchiness) and filter feeders density. Fragmented and low-density patches seem to offer both greater protection from hydrodynamic forces and higher resource availability. In denser patches, however, resources are allocated mostly within the canopy, which would benefit filter feeders if they occurred at low densities, but would be limiting when filter feeder were at high densities.

In an 8-week aquarium experiment, we investigated the interactive effects of waves (present vs. absent) and water-column nutrient level (high vs. low) on the survival, growth, morphology, and biomechanics of the seagrass, Zostera noltii. Survival was reduced when plants were exposed to both waves and high nutrient levels. Wave and nutrient interaction significantly reduced aboveground biomass and leaf lengths, whereas waves independently reduced growth rate, internode abundance, elongation, and appearance rates. Nutrient supply significantly reduced the strength of the leaves. Wave and nutrient interaction was the main driving force affecting survival and morphological properties of seagrass, whereas dynamical characteristics were independently affected by waves, and nutrient supply affected mainly biomechanical properties. In conclusion, this experiment revealed that the combination of exposure to waves and high nutrient levels was detrimental for Z. noltii, which indicates that this could be an important unexplored force involved in seagrass declines.

Leaf decay in seagrasses is enhanced in some seasons since large green senescent beach-cast seagrass leaves are frequently recorded during autumn and winter seasons. Here, we explore if senescence is operating in seagrass leaf decay or if hydrodynamic stress is responsible for the seasonal leaf abscission. A seasonal study on the temperate seagrass Cymodocea nodosa was carried out in four locations with contrasting hydrodynamic regimes. The morphological, biomechanical and material properties of C. nodosa were measured. The force required to break the ligule was always lower than that required to break the blade. This could be considered an adaptive strategy to reduce acute drag forces and thus lessen the chance of plant uprooting. The absolute force needed to dislodge the blade at the ligule level varied with season and location, with the lowest forces recorded in autumn. This may indicate that senescence is operating in this species. On the other hand, the minimum estimated failure velocities for leaf abscission were also recorded in autumn. Consequently, this may cause the premature shedding of leaves in this season before the senescence process has finished and can probably explain the occurrence of green beach-cast seagrass leaves usually found during autumn and winter.