Coastal vegetated habitats maintain highly diverse communities, where the contribution of macrophyte production is significant for macroinvertebrate primary consumers. In the brackish-waters of the Baltic Sea, the taxonomical diversity of different macrophytes includes both marine and limnic species. To study the basal food-web differences of two key vegetated habitat types, either dominated by a perennial brown macroalgae (Fucus vesiculosus) or by angiosperm plants, ¹³C and ¹⁵N compositions of different primary producers and macroinvertebrate consumers were examined, and their diets were estimated by Bayesian mixing models. Carbon isotope diversity of primary producers was high especially in the hard-bottom Fucus-dominated habitats, which was also reflected in a larger consumer isotope niche. However, consumer isotope niche among sites was similar within the same habitat type. Our models indicated that the perennial macrophyte dietary median contribution was about 25% for deposit feeders and omnivores in both habitat types, while epigrazers preferred filamentous algae (30–60%). The niche positions of the abundant clams L. balthica, M. arenaria and C. glaucum differed between the two habitats, but they showed only small (<10% units) differences in their macrophyte dietary contributions. The isotopic compositions of the dominating primary producer assemblage reflected significantly in the isotope niche structure of the associated primary consumers.

The contributions of habitat-forming species to the biodiversity and ecosystem processes of marine and terrestrial ecosystems are widely recognized. Aquatic plants are considered foundation species in shallow ecosystems, as they maintain biodiversity and sustain many ecosystem functions such as primary production and respiration. Despite the increasing amount of biodiversity-ecosystem functioning experiments in seagrass habitats, the effects of benthic variability on ecosystem functioning are rarely investigated across spatially variable aquatic plant habitats. Here, we quantitatively link seasonal variability in seafloor metabolism (i.e. gross primary production and community respiration) with major benthic community components (i.e. microphytobenthos, aquatic plants and macrofauna) across a structural complexity gradient of habitat-forming species (in terms of shoot density and biomass), ranging from bare sand, to a sparse mixture of plants to a dense monospecific seagrass meadow. The increasing complexity gradient enhanced the magnitude of the relationships between benthic community and seafloor metabolism. The daily average seafloor metabolism per season at the bare site was similar to the sparse site, highlighting the role of microphytobenthos for seafloor metabolism in shallow unvegetated sediments. The contribution of the associated macrofauna to the seafloor respiration was similar to the aquatic plant community contribution. Infauna was the main macrofaunal component significantly explaining the seasonal variability of seafloor respiration. However, benthic community-metabolism relationships were stronger within the plant community than within the macrofauna community (i.e. steepest slopes and lowest p-values). Understanding these relationships are a priority since climate change and biodiversity loss are reducing habitat complexity around the world, jeopardizing valuable ecosystem functions and services.

Shallow coastal zones may provide cross-habitat nutrient subsidies for benthic communities offshore, as macrophyte matter can drift to deeper sediments. To study the relative importance of carbon and nutrient flows derived from different primary food sources in a coastal ecosystem, the diets of clam Macoma balthica, polychaete Marenzelleria spp. and mussel Mytilus trossulus were examined across environmental gradients in the northern Baltic Sea using a triple-isotope approach (i.e. 13C, 15N and 34S) and Bayesian mixing models (MixSIAR). Our results suggest that in shallow habitats, production from Fucus vesiculosus is the primary energy source for M. balthica. The proportion of macroalgae-derived matter in the diet of M. balthica and Marenzelleria spp. decreased following a depth gradient. Our models for M. trossulus indicate that the pelagic POM dominates its diet. Our results indicate a trophic connectivity between shallow macrophyte-dominated and deeper habitats, which receive significant amounts of nutrient subsidies from shallower areas.

Mytilid mussels form abundant, species-rich reefs on rocky substrates, but the role of this key habitat in carbon (C) cycling remains poorly understood. We performed a seasonal study on a 5 m deep photic Mytilus trossulus reef in the Central Baltic Sea to investigate pathways and rates of organic C flow. Reef gross primary production (GPP) and respiration (R) were estimated seasonally using underwater O2 eddy covariance on hourly and daily timescales. Photogrammetry and biotic sampling were used to quantify reef rugosity and mussel coverage, and to derive mussel filtration and biodeposition. Mussels were highly abundant, reaching ~50000 ind. m-2, and the reef structure increased the seabed surface area by 44%. GPPhourly was up to 20 mmol O2 m-2 h-1 and GPPdaily was up to 107 mmol O2 m-2 d-1, comparable to a nearby seagrass canopy. Hourly eddy fluxes responded linearly to light intensity and flow velocity, with higher velocities enhancing reef O2 uptake at night. Reef Rdaily exceeded GPPdaily on 12 of 13 measurement days, and Rannual (29 mol O2 m-2 yr-1) was 3-fold larger than GPPannual. The reef sustained a productive community of microbes and fauna whose activities accounted for ~50% of Rannual. Horizontal water advection promoted food supply to the reef and likely facilitated substantial lateral C export of mussel biodeposits. Our analyses suggest that a reduction in mussel reef extent due to ongoing environmental change will have major implications for the transport and transformation of C and nutrients within the coastal Baltic Sea.

The important role of macroalgal canopies in the oceanic carbon (C) cycle is increasingly being recognized, but direct assessments of community productivity remain scarce. We conducted a seasonal study on a sublittoral Baltic Sea canopy of the brown alga Fucus vesiculosus, a prominent species in temperate and Arctic waters. We investigated community production on hourly, daily, and seasonal timescales. Aquatic eddy covariance (AEC) oxygen flux measurements integrated similar to 40 m(2) of the seabed surface area and documented considerable oxygen production by the canopy year-round. High net oxygen production rates of up to 35 +/- 9 mmol m(-2) h(-1) were measured under peak irradiance of similar to 1200 mu mol photosynthetically active radiation (PAR) m(-2) s(-1) in summer. However, high rates > 15 mmol m(-2) h(-1) were also measured in late winter (March) under low light intensities < 250 mu mol PAR m(-2) s(-1) and water temperatures of similar to 1 degrees C. In some cases, hourly AEC fluxes documented an apparent release of oxygen by the canopy under dark conditions, which may be due to gas storage dynamics within internal air spaces of F. vesiculosus. Daily net ecosystem metabolism (NEM) was positive (net autotrophic) in all but one of the five measurement campaigns (December). A simple regression model predicted a net autotrophic canopy for two-thirds of the year, and annual canopy NEM amounted to 25 mol O-2 m(-2) yr(-1), approximately six-fold higher than net phytoplankton production. Canopy C export was similar to 0.3 kg C m(-2) yr(-1), comparable to canopy standing biomass in summer. Macroalgal canopies thus represent regions of intensified C assimilation and export in coastal waters.

The Aquatic Eddy Covariance (AEC) technique has emerged as an important method to quantify in situ seafloor metabolism over large areas of heterogeneous benthic communities, enabling cross-habitat comparisons of seafloor productivity. However, the lack of a corresponding sampling protocol to perform biodiversity comparisons across habitats is impeding a full assessment of marine ecosystem metabolism. Here, we study a range of coastal benthic habitats, from rocky-bed communities defined by either perennial macroalgae or blue mussel beds to soft-sediment communities comprised of either seagrass, patches of different macrophyte species or bare sand. We estimated that the maximum contribution to the AEC metabolic flux can be found for a seafloor area of approximately 80 m(2) with a 5 meter upstream distance of the instrument across all the habitats. We conducted a sampling approach to characterize and quantify the dominant features of biodiversity (i.e., community biomass) within the main seafloor area of maximum metabolic contribution (i.e., gross primary production and community respiration) measured by the AEC. We documented a high biomass contribution of the macroalgal Fucus vesiculosus, the seagrass Zostera marina and the macroinvertebrate Mytilus edulis to the net ecosystem metabolism of the habitats. We also documented a significant role of the bare sediments for primary productivity compared to vegetated canopies of the soft sediments. The AEC also provided insight into dynamic short-term drivers of productivity such as PAR availability and water flow velocity for the productivity estimate. We regard this study as an important step forward, setting a framework for upcoming research focusing on linking biodiversity metrics and AEC flux measurements across habitats.

We studied the contribution of environmental and spatial factors in determining the metacommunity dynamics of benthic macroinvertebrates in ocean-exposed sandy beaches. A combination of different metacommunity models contributed to the structure of the benthic species, suggesting that the interplay of environmental and spatial factors played a key role in determining the beach community structure. Our study highlights the sensitivity of beach invertebrates to environmental factors such as morphodynamic descriptors, and to oceanographic-related variables (e.g., sea-water temperature). The results also suggest significant spatial signals at a large geographical scale. We applied two different species categorizations, high dispersive vs low dispersive and generalist vs specialist, to disentangle the roles of dispersal mode and habitat specialization in the beach metacommunity structure. The strength of the environmental and spatial factors varied depending on the category of species traits considered, emphasizing the value of using different groups of species in explaining variation in metacommunity dynamics. Low dispersive species showed a better ability to track environmental variability than high dispersive species, which were more spatially constrained. Habitat specialists were better able to track environmental variability than generalists, which were mainly predicted by pure spatial factors. A better understanding of the metacommunity dynamics using different species categorizations can help to improve our predictions about exposed beach community structure, and to prioritize management actions to cope with biodiversity loss in such superlative marine environment.