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Subtropical seagrass meadows play a major role in the coastal carbon cycle, but the nature of air–water CO2exchanges over these ecosystems is still poorly understood. The complex physical forcing of air–water exchange in coastal waters challenges our ability to quantify bulk exchanges of CO2and water (evaporation), emphasizing the need for direct measurements. We describe the first direct measurements of evaporation and CO2flux over a calcifying seagrass meadow near Bob Allen Keys, Florida. Over the 78‐d study, CO2emissions were 36% greater during the day than at night, and the site was a net CO2source to the atmosphere of 0.27 ± 0.17 μmol m−2s−1(x̅ ± standard deviation). A quarter (23%) of the diurnal variability in CO2flux was caused by the effect of changing water temperature on gas solubility. Furthermore, evaporation rates were ~ 10 times greater than precipitation, causing a 14% increase in salinity, a potential precursor of seagrass die‐offs. Evaporation rates were not correlated with solar radiation, but instead with air–water temperature gradient and wind shear. We also confirm the role of convective forcing on night‐time enhancement and day‐time suppression of gas transfer. At this site, temperature trends are regulated by solar heating, combined with shallow water depth and relatively consistent air temperature. Our findings indicate that evaporation and air–water CO2exchange over shallow, tropical, and subtropical seagrass ecosystems may be fundamentally different than in submerged vegetated environments elsewhere, in part due to the complex physical forcing of coastal air–sea gas transfer.

Microphytobenthos (MPB) resuspension is a key mechanism in the transfer of organic matter from productive intertidal mudflats to terrestrial and marine systems. In this study, we infer on the contribution of physical and biological factors involved in the MPB resuspension. We use a physical‐biological coupled model forced by realistic meteorological and hydrodynamical forcings to simulate chronic (without any concomitant sediment resuspension) and massive (driven by bed failure) resuspension over a year. The model simulates mud surface temperature, MPB growth, and grazing by the gastropod Peringia ulvae. The model suggests that MPB resuspension is the highest in spring tides and at the flood beginning due to high current velocity and low water heights that promote waves‐sea bottom interactions. The seasonal export of MPB biomass is the highest in spring, up to threefold higher than in summer when the export is the lowest. The simulated seasonal dynamics of MPB resuspension results from the MPB biomass concentration in the sediment, physical disturbances, and the bioturbation activity by P. ulvae. Annually, 43% of the simulated MPB primary production is resuspended. The MPB resuspension (60.8 g C·m−2·yr−1) exceeds the loss by P. ulvaegrazing (41.1 g C·m−2·yr−1). The model suggests that chronic and massive resuspension events are important in the synoptic to seasonal MPB dynamics in temperate intertidal mudflats. Accounting for such processes in the carbon budget assessment in the land‐ocean interface could bring new insights to our understanding of the role played by MPB in the coastal carbon cycle. Intertidal mudflats support a high biological productivity sustained mainly by microalgae living in the sediment. Microalgae form a dense biofilm at the surface of the mud during daytime low tides and fix a high quantity of inorganic carbon into organic carbon through photosynthesis. Microalgae can be resuspended along with the sediment into the sea water during the high tides. Such a transfer is facilitated by episodic physical disturbances like waves and currents modulated by biological drivers. Our study aims to simulate with a numerical model the biologically mediated chronic and physically driven episodic massive resuspension of microalgae on an intertidal mudflat for year 2012. The model is a useful tool to disentangle and estimate the respective contribution of physical and biological factors involved in the microalgae resuspension. At the tidal scale, the resuspension of microalgae is the highest at the flood beginning and during spring tides due to higher current velocity and low water heights that promote waves‐sea bottom interactions and sediment erosion. In 2012, the simulated microalgae resuspension is the highest in spring and the lowest in summer. In the model, the seasonal dynamics of microalgae resuspension results from the microalgae biomass in the sediment, physical disturbances, and the grazers activity at the mud surface that remobilized and facilitated the microalgae resuspension, a process called bioturbation. Annually, 43% of the simulated organic carbon produced by microalgae is resuspended over the year. As microalgae sustain the food web in the mud and in the sea water, their production and resuspension are of key importance for the functioning of coastal ecosystems and the sustainability of economic activity such as shellfish farming. Moreover, estimating the export of organic carbon from the mud to the water is essential to understand the contribution of microalgae biofilms to carbon cycle of the global coastal ocean. Seventy‐six percent of microphytobenthos biomass resuspended at flood beginningSpring massive and chronic events represent 20% and 17.5% of the annual microphytobenthos export, respectivelyForty‐three percent of annual microphytobenthos primary production is resuspended

Seagrass meadows play an important role in “blue carbon” sequestration and storage, but their dynamic metabolism is not fully understood. In a dense Zostera marinameadow, we measured benthic O2fluxes by aquatic eddy covariance, water column concentrations of O2, and partial pressures of CO2(pCO2) over 21 full days during peak growing season in April and June. Seagrass metabolism, derived from the O2flux, varied markedly between the 2 months as biomass accumulated and water temperature increased from 16°C to 28°C, triggering a twofold increase in respiration and a trophic shift of the seagrass meadow from being a carbon sink to a carbon source. Seagrass metabolism was the major driver of diurnal fluctuations in water column O2concentration and pCO2, ranging from 173 to 377 μmol L−1and 193 to 859 ppmv, respectively. This 4.5‐fold variation in pCO2was observed despite buffering by the carbonate system. Hysteresis in diurnal water column pCO2vs. O2concentration was attributed to storage of O2and CO2in seagrass tissue, air–water exchange of O2and CO2, and CO2storage in surface sediment. There was a ~ 1:1 mol‐to‐mol stoichiometric relationship between diurnal fluctuations in concentrations of O2and dissolved inorganic carbon. Our measurements showed no stimulation of photosynthesis at high CO2and low O2concentrations, even though CO2reached levels used in IPCC ocean acidification scenarios. This field study does not support the notion that seagrass meadows may be “winners” in future oceans with elevated CO2concentrations and more frequent temperature extremes.

Microphytobenthos (MPB) are a key primary producer of intertidal mudflats. MPB face strong variability in incident irradiance during low tides. Despite photoprotection and photoacclimation, such variations can translate into the photoinhibition of MPB cells. This study explores the effect of photoinhibition on MPB primary production (PP) over a large and productive temperate mudflat (Brouage mudflat, NW France). We used a regional and high‐resolution tri‐dimensional hydrodynamic model coupled to an MPB model with or without photoinhibition. Photoinhibition leads to a 20% (−0.79 × 103t C) decrease of the simulated MPB PP over the entire mudflat. As the upper shore is exposed to light more frequently and longer than the lower shore, the decrease of MPB PP is higher on the upper shore (−29%) than on the lower shore (−5%). With the highest photosynthetically active radiation cumulated over the mudflat, the decrease of MPB PP due to photoinhibition is the highest during spring and spring tides (−22% and −23%, respectively). The model suggests MPB photoinhibition is sensitive to the photoacclimation status of MPB cells through the light saturation parameter. This first modeling attempt to account for MPB photoinhibition is highly constrained by our current theoretical knowledge and limitations on the MPB growth physiology, but it suggests that this process can have a substantial impact on the MPB PP. As such, assessing the MPB photosynthetic response to the highly variable environmental conditions that prevail in large and productive intertidal mudflats is a real challenge for quantifying MPB PP from a synoptic to inter‐annual time scale. Benthic micro‐algae or microphytobenthos (MPB) inhabiting the surficial sediment sustain the high biological production of intertidal mudflats. MPB achieve photosynthesis by aggregating into a dense biofilm at the mud surface during daytime low tides. As MPB can be exposed to short‐term variations and high light levels, they change their short‐term physiology and position within the sediment to protect themselves. However, such strategies can be outbalanced by a too long stressful light exposure. In this study, we explore with a numerical model the impact of photoinhibition on MPB primary production (PP) over a large and very productive mudflat (NW France). The model suggests that photoinhibition can strongly impact MPB PP. With photoinhibition, the yearly PP decreases by 20% over the whole mudflat. The model suggests MPB PP is sensitive to the photoacclimation status of MPB cells, that is, their light use efficiency at a given light level. This first modeling attempt to account for MPB photoinhibition is highly constrained by our current theoretical knowledge and limitations on the MPB growth physiology, but it suggests that this process can have a substantial impact on the MPB PP. With or without photoinhibition, simulated yearly MPB PP is higher on the upper and middle shores than on the lower shorePhotoinhibition leads to a 20% (−0.79×103t C yr−1) decrease of MPB PP over the entire mudflatThe photoinhibition process in the model is sensitive to the photoacclimation status of MPB cells With or without photoinhibition, simulated yearly MPB PP is higher on the upper and middle shores than on the lower shore Photoinhibition leads to a 20% (−0.79×103t C yr−1) decrease of MPB PP over the entire mudflat The photoinhibition process in the model is sensitive to the photoacclimation status of MPB cells

Coastal vegetated habitats like seagrass meadows can mitigate anthropogenic carbon emissions by sequestering CO2as “blue carbon” (BC). Already, some coastal ecosystems are actively managed to enhance BC storage, with associated BC stocks included in national greenhouse gas inventories. However, the extent to which BC burial fluxes are enhanced or counteracted by other carbon fluxes, especially air‐water CO2flux (FCO2) remains poorly understood. In this study, we synthesized all available direct FCO2measurements over seagrass meadows made using atmospheric Eddy Covariance, across a globally representative range of ecotypes. Of the four sites with seasonal data coverage, two were net CO2sources, with average FCO2equivalent to 44%–115% of the global average BC burial rate. At the remaining sites, net CO2uptake was 101%–888% of average BC burial. A wavelet coherence analysis demonstrated that FCO2was most strongly related to physical factors like temperature, wind, and tides. In particular, tidal forcing was a key driver of global‐scale patterns in FCO2, likely due to a combination of lateral carbon exchange, bottom‐driven turbulence, and pore‐water pumping. Lastly, sea‐surface drag coefficients were always greater than the prediction for the open ocean, supporting a universal enhancement of gas‐transfer in shallow coastal waters. Our study points to the need for a more comprehensive approach to BC assessments, considering not only organic carbon storage, but also air‐water CO2exchange, and its complex biogeochemical and physical drivers. Carbon storage is a valuable ecosystem service of seagrass meadows, serving as a possible pathway to draw down atmospheric carbon dioxide (CO2) levels. However, this approach may be unsuccessful if carbon storage in sediments is exceeded by the release of CO2from the water. To better understand the scope of this problem, we compiled all available measurements of air‐water CO2exchange over seagrass meadows. We found that rates of CO2release or uptake were indeed large, even when compared with potential rates of carbon storage in seagrass soils. However, these large air‐water exchanges of CO2did not occur for the same reason everywhere. While light availability was sometimes a strong predictor of air‐water CO2exchange, tidal mixing and temperature were also very important, revealing a much more complex network of drivers than previously thought. Despite these diverse conditions, we found one key similarity across all sites, in that rates of air‐water gas transfer appear to always be greater than would be expected for the open ocean. Taken together, the results of our study show that assessments of carbon storage in coastal seagrass ecosystems will be incomplete if they do not consider exchanges of CO2between the water and air. Direct measurements show that air‐water CO2exchange over seagrass meadows is of similar magnitude to carbon burial ratesKey drivers are tides, temperature, light, and wind, which off in importance over hourly seasonal time scalesSurface drag coefficients were greater than open water prediction, suggesting a near‐universal gas transfer enhancement across all sites Direct measurements show that air‐water CO2exchange over seagrass meadows is of similar magnitude to carbon burial rates Key drivers are tides, temperature, light, and wind, which off in importance over hourly seasonal time scales Surface drag coefficients were greater than open water prediction, suggesting a near‐universal gas transfer enhancement across all sites