Description: Benthic fluxes of dissolved nutrients and oxygen measured in the southern North Sea using ex situ incubation chambers indicate a prominent annual cycle characterized by low level from mid-autumn (Oct) to early spring (Mar) and enhanced values from mid-spring (Apr) to early autumn (Sep) with peak in late summer (late Aug/early Sep). The same cycle is also shown in the budget of total organic carbon (TOC) and macrobenthic biomass in surface sediments. The significant positive correlations between the benthic nutrient fluxes, oxygen, sedimentary TOC and macrobenthos suggest that their variation might respond to a common source, i.e. the primary production. However, the linkages between these quantities and pelagic primary production, which exhibits a dominant bloom in early spring (Mar/Apr) and a secondary bloom in early summer (Jun/Jul) in the study area, is not straightforward. We present a numerical study to unravel the complex linkages. A 3-D coupled hydrodynamic-biogeochemical model (ECOSMO) was used to provide benthic boundary conditions for a 1-D biogeochemical model in the sediment (TOCMAIM) that mechanistically resolves the interaction between macrobenthos and organic matter through bioturbation. Simulation results based on a satisfactory hindcast from 1948 to 2015 reveal that although the spring algal bloom normally starts in late winter (Feb) and peaks in early spring (Mar/Apr), deposition of labile OC to seafloor is limited in this period due to energetic hydrodynamic conditions. Sedimentation and accumulation of labile OC (originated from fresh planktonic detritus) in seafloor surface sediments are facilitated in summer when wind-waves become weak enough. This drives the blooming of macrobenthos, with peak of biomass in late summer (Aug). Bioturbation intensity, which is dependent upon macrobenthic biomass, community structure as well as local food resource, peaks also in later summer. Enhanced bioturbation and benthic metabolism result in an increased oxygen flux into sediments, promoting remineralization of OC and release of nutrients. The following period (late Sep/Oct) is characterized by low level of pelagic primary production in combination with enhanced wind-waves, which not only reduce the input of labile OC into sediments substantially but also remobilize surface material (sediments and OC) on a major part of the shallow coastal seafloor. Depletion of labile OC in the uppermost centimeters of sediments by a combined effect of erosion, macrobenthic uptake and downward mixing (through bioturbation) accounts for the rapid decline of benthic nutrient fluxes in Oct, which remain low through the stormy winter until the next spring.

Description: The importance of macrobenthos in benthic‐pelagic coupling and early diagenesis of organic carbon (OC) has long been recognized but has not been quantified at a regional scale. By using the southern North Sea as an exemplary area we present a modelling attempt to quantify the budget of total organic carbon (TOC) reworked by macrobenthos in seafloor surface sediments. Vertical profiles in sediments collected in the field indicate a significant but nonlinear correlation between TOC and macrobenthic biomass. A mechanistic model is used to resolve the bi‐directional interaction between TOC and macrobenthos. A novelty of this model is that bioturbation is resolved dynamically depending on variations in local food resource and macrobenthic biomass. The model is coupled to 3D hydrodynamic‐biogeochemical simulations to hindcast the mutual dependence between sedimentary TOC and macrobenthos from 1948 to 2015. Agreement with field data reveals a satisfactory model performance. Our simulations show that the preservation of TOC in the North Sea sediments is not only determined by pelagic conditions (hydrodynamic regime and primary production) but also by the vertical distribution of TOC, bioturbation intensity, and the vertical positioning of macrobenthos. Macrobenthos annually ingest 20%–35% and in addition vertically diffuse 11%–22% of the total budget of TOC in the upper‐most 30 cm sediments in the southern North Sea. This result indicates a central role of benthic animals in modulating the OC cycling at the sediment‐water interface of continental margins.

Description: Field data collected for the North Sea indicate a prominent seasonal variation in the vertical distribution of total organic carbon (TOC) and macrobenthic biomass in sediments. The vertical TOC profiles classify into three modes, with maximum at surface, middle and deep part of sediments, respectively. We here present a mechanistic model to quantify, for the first time, the dynamic interaction between sedimentary TOC and benthic fauna. The major model principles include that (i) the vertical distribution of macrobenthic biomass is a trade-off between nutritional benefit (quantity and quality of TOC) and the costs of burial (respiration) and mortality, and (ii) the vertical transport of TOC is in turn modulated by macrobenthos through bioturbation. A novelty of our model is that bioturbation is resolved dynamically depending on variation of local food resources and macrobenthic biomass. This allows capturing of the benthic response to both depositional and erosional conditions and improving estimates of the material exchange flux at the sediment-water interface. The coupling of the TOC-benthos model with 3D hydrodynamic-ecological simulations reveals that the three profile modes of sedimentary TOC (in both quantify and quality) can be explained as a combined response to pelagic conditions (shear stress and primary production) and the synergy between bioturbation, vertical redistribution of higher quality TOC and vertical positioning of benthic organisms. A model reconstruction of the benthic status in the North Sea from 1950s to 2010s indicates that despite a relatively stable pattern at decadal and regional scales, significant variations exist at smaller scales characterized by seasons and local areas. In addition, inter-annual and multi-year cycle-like variations are also prominent especially in coastal areas.

Description: Trait based indices constitute a versatile tool for the prediction of ecosystem functioning over large spatial scales and represent a promising approach to meet societal, political and regulatory demands. Here we investigate for the first time the ability of different trait based indices to predict nutrient fluxes of ammonium, nitrate, nitrite, silicate and phosphate under different environmental conditions. We hypothesize that irrigation traits, as applied in the newly proposed index “Community Irrigation Potential” (IPc), will increase the predictability of macrofaunal impact on nutrient fluxes compared to commonly used sediment reworking traits, as in the index “Community Bioturbation Potential” BPc. We correlate IPc and BPc with experimental nutrient flux data measured under different environmental conditions. Both trait based indices and environmental conditions significantly affected all analysed nutrient fluxes. We therefore conclude that neither the trait based indices nor the environmental conditions suffice for quantitative modelling of sediment biogeochemical turnover. Accordingly, information on of macrofaunal activity is needed to reliably predict biogeochemical turnover. Our results further demonstrate that generally nutrient fluxes of ammonium, nitrate, nitrite, silicate and phosphate are more closely linked to irrigation traits than to sediment reworking traits. In conclusion, linking macrofaunal bioirrigation to important environmental factors such as permeability, changing nutrient gradients in the water column and organic matter concentrations may strongly enhance performance of ecosystem models.

Description: This study shows that macrofaunal irrigation traits constitute a valuable complement to sediment reworking traits in estimating macrofaunal impact on nutrient fluxes across the sediment-water interface. We correlated density, biomass, community bioturbation potential (BPc, an index based on reworking traits, body mass and density) and community irrigation potential (IPc, an index based on irrigation traits, body mass and density) with nitrite, nitrate, ammonium, silicate and phosphate flux data under different environmental conditions. Generalized linear models performed best with a combination of environmental conditions and irrigation trait-based indices. This was not only a direct effect of the irrigation traits, but also of the scaling factor 0.75 employed in IPc to infer metabolic activity from body mass. Accordingly, predictive models of nutrient flux across the sediment-water interface will profit greatly from incorporating macrofaunal irrigation behaviour by means of trait-based indices.

Description: Bioturbation is one of the key mediators of biogeochemical processes in benthic habitats that can have a high contribution to seafloor functioning and benthic pelagic coupling in coastal waters. Previous studies on bioturbation were limited to point locations and extrapolations in single regions, but have not accounted for regional differences under changing environmental conditions, though there are indications that species contributions will differ across regions or with biotic and abiotic context. To capture those differences and assess global patterns and commonalities, multi-regional analyses are imperative. Here for the first time, bioturbation potential (BPc), a functional indicator of benthic community bioturbation, was estimated based on macrofauna data from four regions (i.e. German Baltic Sea, German North Sea, Belgian part of the North Sea and the Eastern English Channel). For each region and sediment type we identified key species contributing to BPc. Comparison within and across regions demonstrated regional differences, and both overlap and mismatch between species that are functionally important and those that are dominant in biomass. Knowledge on the functionally important species is crucial when management objectives include the protection of certain ecosystem functions. Available environmental layers were used as predictors to model the spatial distribution of BPc for each area and to explore the underlying drivers of differences. Random forest models were trained using as response variables either i) BPc initially calculated per station; or ii) BPp – the species-specific contribution to BPc – for key species (with subsequent summation of their predicted full-coverage distributions to BPc). Maps of BPc distribution predicted by random forest were compared with those generated using natural neighbour interpolation. Overall, derived BPc values increased towards the German parts of the North and Baltic Seas. The relevance of BPc for ecosystem processes and functions, however, vary with biotic and abiotic settings. Results revealed a strong association of BPc with species diversity and region, but less with sediment grain size. A large range of BPc occurred when species richness was low. This suggests that the provisioning of high bioturbation activity is possible also under low diversity, where it is vulnerable due to reduced resilience. The executed multi-regional analysis allowed identifying regional differences in performance of macrofauna, suggesting the need for regionspecific conservation and management strategies. https://doi.org/10.1016/j.ecolind.2019.105945 Received 26 July 2019; Received in revised form 12 November 2019; Accepted 14 November 2019 ⁎ Corresponding author. E-mail address: mayya.gogina@io-warnemuende.de (M. Gogina). Ecological Indicators 110 (2020) 105945 1470-160X/ Crown Copyright © 2019 Published by Elsevier Ltd. All rights reserved. T

Description: Coastal morphodynamic and hydrodynamic processes impose a first-order control on biogeochemical cycling and distribution of benthic habitats. However, little is known on how biota in turn affect sediment transport, biogeochemical cycling and morphodynamics at a regional scale and long-term. Here we firstly present a mini-review of recent progress on numerical modeling of interactions between benthic biota, biogeochemical cycling and coastal morphodynamics. Special focus is laid on 1) the synergetic effects of dominant benthic functional groups on sedimentation process, and 2) the macrobenthic response to changing morphological, hydrodynamic and biogeochemical conditions. In a second part we present preliminary results from a case study area (Jade Bay, Wadden Sea) based on simulations using a coupled hydrodynamic-biogeochemical-morphodynamic model and comparison with biogeochemical and taxonomic field monitoring data. The aim is to understand benthic ecosystem functioning and to identify the main drivers of coastal ecosystem changes for better prediction of their future development.

Description: Loss of macrofaunal bioturbation and bioirrigation activity may strongly reduce benthic biogeochemical cycling and thus ecosystem functioning. The identification of bioturbating key species in the marine benthic realm is therefore of high importance for ecosystem management purposes. In this study top bioturbators in the German Bight were identified by mapping the trait based bioturbation potential (BPc) for 423 North Sea stations. BPc mapping highlighted the importance of Amphiura filiformis, Echinocardium cordatum and Nucula nitidosa as major bioturbating species in the German Bight. Effects of their bioturbation and bioirrigation activity on silicate, ammonium and nitrate flux were investigated in laboratory experiments. While E. cordatum significantly influenced biogeochemical cycling, effects of A. filiformis remained inconclusive probably due to arm regeneration. N. nitidosa showed little impact on biogeochemical cycling, although the bivalve was found to be an important bioturbator. E. cordatum may thus be considered an essential mediator of biogeochemical cycling in the sediment water interface as well as one of the most important bioturbators in the German Bight.

Description: Increasing anthropogenic activities on land and at sea underline the demand for easily applicable indices to effectively predict human mediated changes in ecosystem functioning. Here, we propose a novel bioirrigation index (IPc) that is based on body mass, abundance, burrow type, feeding type and injection pocket depth of bottom dwelling animals. The index was validated with combined field (in situ communities) and manipulative (single species) experiments. Results from both community and single-species experimental incubations indicate that IPc is able to predict the bioirrigation rate in different sediment types (mud, fine sand, sand). The trait-based index thus demonstrates robustness in the prediction of animal-mediated functional processes that support biogeochemical functions under variable environmental conditions. Accordingly, we argue that trait-based indices provide a useful tool for the quantitative prediction of ecosystem processes as effect traits provide a direct link to the behavioral mechanisms that drive ecosystem functioning.