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Coastal ecosystems act as filters of nutrients from land to the open sea. We investigated the role of eelgrass (Zostera marina) metabolism in the coastal filter transforming nitrogen, phosphorus, and organic carbon. Field campaigns following identical methodologies were carried out at two contrasting coastal locations: the mesohaline and nutrient-rich Roskilde Fjord, Denmark, and the mesotrophic brackish Tvärminne archipelago, Finland. Over the 24-h in situ benthic incubations, we measured oxygen concentrations continuously and assessed changes in DOM characteristics and net fluxes of carbon, nitrogen, and phosphorus. Ecosystem metabolism modeled on the basis of the O2 data showed that the systems were either net heterotrophic (Roskilde Fjord; − 1.6 and − 2.4 g O2 m−2 day−1 in eelgrass meadow and bare sand, respectively) or had balanced primary production and respiration (Tvärminne; 0.0 and 0.2 g O2 m−2 day−1). Overall, initial nutrient stoichiometry was a key factor determining benthic–pelagic fluxes of nutrients, which exacerbated the deviations from Redfield ratios of N and P, indicating an efficient use of the limiting nutrient. A net diel uptake of dissolved inorganic N was observed at both locations (− 2.3 μmol l−1 day−1 in Roskilde Fjord and − 0.1 μmol l−1 day−1 in Tvärminne). Despite minor changes in dissolved organic carbon concentrations during the incubations, a marked increase of fluorescent DOM was observed at both locations, suggesting rapid heterotrophic processing of the DOM pool. Our results underline that the biogeochemical role of eelgrass in the coastal filter is not inherent, but strongly dependent on the environmental conditions.

Understanding the ecological interactions that enhance the resilience of threatened ecosystems is essential in assuring their conservation and restoration. Top-down trophic interactions can increase resilience to bottom-up nutrient enrichment, however, as many seagrass ecosystems are threatened by both eutrophication and trophic modifications, understanding how these processes interact is important. Using a combination of approaches, we explored how bottom-up and top-down processes, acting individually or in conjunction, can affect eelgrass meadows and associated communities in the northern Baltic Sea. Field surveys along with fish diet and stable isotope analyses revealed that the eelgrass trophic network included two main top predatory fish species, each of which feeds on a separate group of invertebrate mesograzers (crustaceans or gastropods). Mesograzer abundance in the study area was high, and capable of mitigating the effects of increased algal biomass that resulted from experimental nutrient enrichment in the field. When crustacean mesograzers were experimentally excluded, gastropod mesograzers were able to compensate and limit the effects of nutrient enrichment on eelgrass biomass and growth. Our results suggest that top-down processes (i.e., suppression of algae by different mesograzer groups) may ensure eelgrass resilience to nutrient enrichment in the northern Baltic Sea, and the existence of multiple trophic pathways can provide additional resilience in the face of trophic modifications. However, the future resilience of these meadows is likely threatened by additional local stressors and global environmental change. Understanding the trophic links and interactions that ensure resilience is essential for managing and conserving these important ecosystems and the services they provide.

Functional diversity (FD) experiments are highly effective for investigating how a community interacts with its environment. However, such experiments using morphological and chemical traits have not been conducted for submerged aquatic plants and their insights would be highly valuable for understanding the ecology of these communities. We conducted a 15-week field experiment in the Baltic Sea where we manipulated the species composition of aquatic plant communities to investigate functional diversity. We constructed artificial triculture communities with different species compositions to change the Community Weighted Means (CWMs) and variability of traits. We measured nine plant traits and tested how community productivity (CP) was related to FD, trait CWMs and community trait ranges. CP varied by more than four times across treatments and functional richness was significantly related to CP. Functional evenness and functional divergence were not significantly related to CR Height, leaf area and delta C-13 were significantly related to CP. Leaf delta C-13 trends with CP suggested that the carbon supply is not replete, yet species composition was partly responsible for the relationship. Plant height likely had multifaceted benefits to CP because there was evidence of a competitive height interaction between the tallest and 2 nd tallest species, therefore the effects of plant height to CP would have been disproportionally large. The height of the tallest species significantly drove the variability of the community height range, which was significantly related to CP and it had a relatively large influence on the calculation of FD indices. Leaf area, which was strongly correlated to plant height, was also significantly related to CR The significant relationship between functional richness and CP was most likely driven by the presence of taller plants. FD likely enhanced CP, by selecting for extreme trait values which enhanced production (selection effect), while niche complementarity effects were not observed. This study provides experimental evidence and mechanistic insights into the role of FD and specific traits for CP in submerged aquatic plant communities. To conclude, FD was significantly related to CP of temperate aquatic plant communities likely by selecting for traits which enhanced light capture, with consequences for carbon supply.

Aquatic plant meadows are valuable components to the ‘coastal filter’ and it is important to understand the processes that drive their ability to cycle nutrients. However, at present, the field-based evidence for understanding the drivers of nutrient uptake by plants is lacking. This study aimed to investigate how well individual shoots of aquatic plants could meet their nitrogen demands using the sediment nutrient pool (porewater ammonium) and to explore which traits helped to facilitate such uptake. Several species were investigated in shallow, submerged (2–4 m) mixed-species communities in the northern Baltic Sea using incubation experiments with enriched ammonium. After a 3.5 h incubation time, individuals were collected and analysed for nitrogen (% DW) and 15N (at-%) concentrations. Uptake by plants was calculated per unit nitrogen in response to the 15 N-labelled source and to overall nitrogen availability. Background porewater ammonium availability was highly variable between individual plants. Species identity did not significantly affect uptake metrics and the effect of ambient porewater availability was weak. As biomass increased there were significant logarithmic declines in the 95th quantiles of nutrient uptake rates, ambient porewater nutrient availability and aboveground nitrogen tissue concentrations (% DW). Such findings suggested that uptake rates of plants were significantly demand-driven and the nutrient conditions of the porewater were significantly driven by the demands of the plant. Findings parameterised the unfulfilled potential for some aquatic plants to cycle nutrients more efficiently and highlighted the potential importance of access to new nutrient sources as a way of enhancing nutrient cycling by aquatic plants. Plant traits and community properties such as the activity of infauna could facilitate such an access and are likely important for nutrient uptake.

1. Aquatic plant meadows are important coastal habitats that sustain many ecosystem functions such as primary production and carbon sequestration. Currently, there is a knowledge gap in understanding which plant functional traits, for example, leaf size or plant height underlie primary production in aquatic plant communities. 2. To study how plant traits are related to primary production, we conducted a field survey in the Baltic Sea, Finland, which is characterized by high plant species and functional diversity. Thirty sites along an exposure gradient were sampled (150 plots), and nine plant morphological and chemical traits measured. The aim was to discern how community-weighted mean traits affect community production and whether this relationship changes along an environmental gradient using structural equation modelling (SEM). 3. Plant height had a direct positive effect on production along an exposure gradient (r = 0.33) and indirect effects through two leaf chemical traits, leaf δ15N and leaf δ13C (r = 0.24 and 0.18, respectively) resulting in a total effect of 0.28. In plant communities experiencing varying exposure, traits such as root N concentration and leaf δ15N had positive and negative effects on production, respectively. 4. Synthesis. Our results demonstrate that the relationship between aquatic plant functional traits and community production is variable and changes over environmental gradients. Plant height generally has a positive effect on community production along an exposure gradient, while the link between other traits and production changes in plant communities experiencing varying degrees of exposure. Thus, the underlying biological mechanisms influencing production differ in plant communities, emphasizing the need to resolve variability and its drivers in real-world communities. Importantly, functionally diverse plant communities sustain ecosystem functioning differently and

Harmful algal blooms are considered to be one of the biggest threats to benthic vegetation. The amount of annual algal mats has increased due to eutrophication but their effects on perennial vegetation have rarely been tested in field experiments. In regions where eelgrass ( Zostera marina ) grows in mixed meadows, plant species richness may ameliorate algal-induced stress through facilitative mechanisms and improve eelgrass growth and survival. To test this, we conducted a 3 mo. field experiment at 3.5 m depth in the Baltic Sea, where we applied filamentous brown algae (300 g ww) to eelgrass monocultures and polycultures ( Z. marina , Ruppia cirrhosa , Stuckenia pectinata and Potamogeton perfoliatus ). The algal stress period lasted for 13 d and plots were sampled after a 5 wk recovery period. In both algal-stressed mono- and polycultures, the presence of algal mats significantly reduced eelgrass shoot and root biomass production and affected the leaf growth negatively. Plant richness did not have strong effects on eelgrass growth and eelgrass in both algal-stressed mono- and polycultures showed significantly reduced productivity. Our results clearly demonstrate that loose-lying algal mats pose a serious threat to seagrass meadows and that plant diversity cannot alone ameliorate negative effects by macroalgal mats.

Studies conducted in terrestrial ecosystems have shown that increasing plant diversity enhances ecosystem processes such as primary production. In marine systems, knowledge of how plant diversity influences ecosystem processes and higher trophic levels is still limited. To examine how plant richness and composition influence recruitment and colonization processes, defaunated eelgrass ( Zostera marina ), sago pondweed ( Potamogeton pectinatus ) and perfoliate pondweed ( Potamogeton perfoliatus ) were planted on an unvegetated, sandy bottom in all possible combinations in July and August. Both field experiments lasted 1 week. Our results showed that in these plant assemblages plant richness had a negative effect on faunal diversity ( H′ ) and evenness ( J′ ), while total abundance showed strong temporal variation with weak, positive relationships with plant species richness in both August and July. Plant species composition had strong effects on amphipods ( Gammarus spp.), thus both the abundance and biomass were higher in treatments containing P. perfoliatus . The colonization process was significantly influenced by the numerical dominance of a few faunal species, e.g. the settlement of lagoon cockles ( Cerastoderma glaucum ), and by the timing of the experiments. The results indicate that faunal colonization in these communities is rapid and significantly influenced by the traits of particular plant species.

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom-up and top-down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top-down and bottom-up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top-down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross-site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large-scale patterns corresponded strikingly with prior small-scale experiments. Our results link global and local evidence that biodiversity and top-down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors. National Science Foundation [OCE-1031061] info:eu-repo/semantics/publishedVersion

This paper focuses on the marine foundation eelgrass species, Zostera marina, along a gradient from the northern Baltic Sea to the north-east Atlantic. This vast region supports a minimum of 1480 km2 eelgrass (maximum >2100 km2), which corresponds to more than four times the previously quantified area of eelgrass in Western Europe.Eelgrass meadows in the low salinity Baltic Sea support the highest diversity (4–6 spp.) of angiosperms overall, but eelgrass productivity is low (

Stressful environments may enhance the occurrence of facilitative interspecific interactions between plants. In several regions, Zostera marina occurs in mixed assemblages. However, the potential effects of plant diversity on stress responses and stability properties of Z. marina are poorly understood. We investigated the resistance and recovery of Z. marina subjected to shading (1 mo) in a field experiment lasting 2.5 mo. We shaded Z. marina planted in mono- and polycultures (Potamogeton perfoliatus, P. pectinatus, P. filiformis) in a factorial design (Shading×Richness) at 2 m depth. We estimated the resistance and recovery of Z. marina by measuring four response variables. Polyculture Z. marina lost proportionally less biomass than monocultures, thus having a greater resistance to shading. In contrast, after a 1 mo recovery period, monocultures exhibited higher biomass gain, and a faster recovery than polycultures. Our results suggest that plant species richness enhances the resistance of Z. marina through facilitative mechanisms, while the faster recovery in monocultures is possibly due to interspecific competition. Our results highlight the need of a much better understanding of the effects of interspecific interactions on ecosystem processes in mixed seagrass meadows, and the preservation of diverse plant assemblages to maintain ecosystem functioning.