Bioturbation, the biogenic modification of sediments through particle reworking and burrow ventilation, is a key mediator of many important geochemical processes in marine systems. In situ quantification of bioturbation can be achieved in a myriad of ways, requiring expert knowledge, technology, and resources not always available, and not feasible in some settings. Where dedicated research programmes do not exist, a practical alternative is the adoption of a trait-based approach to estimate community bioturbation potential (BPc). This index can be calculated from inventories of species, abundance and biomass data (routinely available for many systems), and a functional classification of organism traits associated with sediment mixing (less available). Presently, however, there is no agreed standard categorization for the reworking mode and mobility of benthic species. Based on information from the literature and expert opinion, we provide a functional classification for 1033 benthic invertebrate species from the northwest European continental shelf, as a tool to enable the standardized calculation of BPc in the region. Future uses of this classification table will increase the comparability and utility of large-scale assessments of ecosystem processes and functioning influenced by bioturbation (e.g., to support legislation). The key strengths, assumptions, and limitations of BPc as a metric are critically reviewed, offering guidelines for its calculation and application.

Shelf seas and their associated benthic habitats represent key systems in the global carbon cycle. However, the quantification of the related stocks and flows of carbon are often poorly constrained. To address benthic carbon storage in the North-West European continental shelf, we have spatially predicted the mass of particulate organic carbon (POC) stored in the top 10 cm of shelf sediments in parts of the North Sea, English Channel and Celtic Sea using a Random Forest model, POC measurements on surface sediments from those seas and relevant predictor variables. The presented model explains 78% of the variance in the data and we estimate that approximately 250 Mt of POC are stored in surficial sediments of the study area (633,000 km 2 ). Upscaling to the North-West European continental shelf area (1,111,812 km 2 ) yielded a range of 230-882 Mt of POC with the most likely estimate being on the order of 476 Mt. We demonstrate that the largest POC stocks are associated with coarse-grained sediments due to their wide-spread occurrence and high dry bulk densities. Our results also highlight the importance of coastal sediments for carbon storage and sequestration. Important predictors for POC include mud content in surficial sediments, annual average bottom temperature and distance to shoreline, with the latter possibly a proxy for terrestrial inputs. Now that key variables in determining the spatial distribution of POC have been identified, it is possible to predict future changes to the POC stock, with the presented maps providing an accurate baseline against which to assess predicted changes.

A carbon budget for the northwest European continental shelf seas (NWES) was synthesized using available estimates for coastal, pelagic and benthic carbon stocks and flows. Key uncertainties were identified and the effect of future impacts on the carbon budget were assessed. The water of the shelf seas contains between 210 and 230 Tmol of carbon and absorbs between 1.3 and 3.3 Tmol from the atmosphere annually. Offshelf transport and burial in the sediments account for 60-100 and 0-40% of carbon outputs from the NWES, respectively. Both of these fluxes remain poorly constrained by observations and resolving their magnitudes and relative importance is a key research priority. Pelagic and benthic carbon stocks are dominated by inorganic carbon. Shelf sediments contain the largest stock of carbon, with between 520 and 1600 Tmol stored in the top 0.1 m of the sea bed. Coastal habitats such as salt marshes and mud flats contain large amounts of carbon per unit area but their total carbon stocks are small compared to pelagic and benthic stocks due to their smaller spatial extent. The large pelagic stock of carbon will continue to increase due to the rising concentration of atmospheric CO 2 , with associated pH decrease. Pelagic carbon stocks and flows are also likely to be significantly affected by increasing acidity and temperature, and circulation changes but the net impact is uncertain. Benthic carbon stocks will be affected by increasing temperature and acidity, and decreasing oxygen concentrations, although the net impact of these interrelated changes on carbon stocks is uncertain and a major knowledge gap. The impact of bottom trawling on benthic carbon stocks Frontiers in Marine Science | www.frontiersin.org 1 March 2020 | Volume 7 | Article 143Legge et al.Carbon on the Northwest European Shelf is unique amongst the impacts we consider in that it is widespread and also directly manageable, although its net effect on the carbon budget is uncertain. Coastal habitats are vulnerable to sea level rise and are strongly impacted by management decisions. Local, national and regional actions have the potential to protect or enhance carbon storage, but ultimately global governance, via controls on emissions, has the greatest potential to influence the long-term fate of carbon stocks in the northwestern European continental shelf.

We measured denitrification, anaerobic ammonium oxidation (anammox), oxygen uptake, nutrient exchange, and pore-water profiles of oxygen in intact sediments at three sites in the southern North Sea, which we experimentally exposed to different oxygen saturations (ambient and , 33% of air-saturation for oxygen [i.e., our hypoxic treatment]) over 14 months. Denitrification ranged from 1 mmol N m 22 h 21 to 21 mmol N m 22 h 21 , anammox 0.2 mmol N m 22 h 21 to 5.7 mmol N m 22 h 21 , and oxygen uptake 47 mmol O 2 m 22 h 21 to 631 mmol O 2 m 22 h 21 . The seasonal patterns under ambient oxygen were correlated with those in the hypoxic treatment; though, on the whole, the magnitude of flux was different. On average, under hypoxia, both the penetration and consumption of oxygen decreased by , 50%, denitrification increased by 32%, and anammox remained constant. Anammox accounted for between 10% and 20% of the total N 2 production, which agrees with expectations for waters of these depths (30-80 m). Under ambient oxygen the sediments were strong sources of nitrate to the overlying water, 12 mmol NO { 3 m 22 h 21 on average, but under hypoxia total N mineralization decreased by 46% and nitrate exchanged ceased. Short-term hypoxia alters the balance between available N returned to the overlying water, primarily as NO { 3 , and that removed from the ecosystem as N 2 gas.

Tickler-chain SumWing and electrode-fitted PulseWing trawls were compared to assess seabed impacts. Multi-beam echo sounder (MBES) bathymetry confirmed that the SumWing trawl tracks were consistently and uniformly deepened to 1.5 cm depth in contrast to 0.7 cm following PulseWing trawling. MBES backscatter strength analysis showed that SumWing trawls (3.11 dB) flattened seabed roughness significantly more than PulseWing trawls (2.37 dB). Sediment Profile Imagery (SPI) showed that SumWing trawls (mean, SD) homogenised the sediment deeper (3.4 cm, 0.9 cm) and removed more of the oxidised layer than PulseWing trawls (1 cm, 0.8 cm). The reduced PulseWing trawling impacts allowed a faster re-establishment of the oxidised layer and micro-topography. Particle size analysis suggested that SumWing trawls injected finer particles into the deeper sediment layers (∼4 cm depth), while PulseWing trawling only caused coarsening of the top layers (winnowing effect). Total penetration depth (mean, SD) of the SumWing trawls (4.1 cm, 0.9 cm) and PulseWing trawls (1.8 cm, 0.8 cm) was estimated by the depth of the disturbance layer and the layer of mobilized sediment (SumWing = 0.7 cm; PulseWing trawl = 0.8 cm). PulseWing trawls reduced most of the mechanical seabed impacts compared to SumWing trawls for this substrate and area characteristics.