Restoration engineering measures, such as managed realignments or building groins, modify the environmental characteristics of coastal intertidal ecosystems. Creating physical modifications that are beneficial to an intertidal system's ecology necessitates an in-depth understanding of the relationships between the abiotic and biotic components of a given intertidal habitat. In this study, we evaluate how hydrodynamics and sediment characteristics drive the development of the benthic macrofauna community during the first 5 years following engineering measures to enhance benthic macrofauna diversity at three locations. The creation of low-energy habitats through groins (Knuitershoek and Baalhoek) and a managed realignment dike breach (Perkpolder) led to the accumulation of fine sediments in all three impact sites. Biomass of benthic macrofauna quickly increased between 2016 and 2020, with successional processes being more important in Perkpolder, where the habitat was started completely from scratch due to a managed realignment, than at Knuitershoek or Baalhoek, where habitat conditions were improved by adding groins. In addition, the density of benthos-eating birds, especially oystercatchers, increased at some of the modified sites. While a low-energy habitat may harbor more diverse assemblages of benthic macrofauna than a highly dynamic one, the extremely high silt content, which is typical for low-energy habitats, may slow benthic community development. The observed increase of biomass at our impact sites highlights the value of the interventions, while the delays in the response of the benthic macrofauna community emphasizes the need for extensive monitoring both in time and space and the identification of underlying abiotic–biotic mechanisms.

Tidal flats are biogeomorphic landscapes, shaped by physical forces and interaction with benthic biota. We used a metabolic approach to assess the overarching effect of bioturbators on tidal landscapes. The benthic bivalve common cockle (Cerastoderma edule) was used as model organism. The effect of C. edule on sediment resuspension was approximated as a function of the overall population metabolic rate per unit of area. We combined i) laboratory observations on how C. edule affect sediment resuspension along gradients of bioturbation activity, sediment cohesiveness and hydrodynamic force with ii) spatial data on the natural distribution of intertidal C. edule populations. This allowed us to build an integrated model of the C. edule effect on sediment resuspension along the tidal gradient. Owing to the temperature dependence of metabolic rate, the model also accounted for seasonal variation in bioturbators activity. Laboratory experiments indicated that sediment resuspension is positively related to the metabolic rate of the C. edule population especially in cohesive sediments. Based on this observation, we predicted a clear spatial and seasonal pattern in the relative importance of C. edule contribution to sediment resuspension along a tidal transect. At lower elevations, our model indicates that hydrodynamics overrules biotic effects; at higher elevations, inter-tidal hydrodynamics should be too low to suspend bioturbated sediments. The influence of C. edule on sediment resuspension is expected to be maximal at the intermediate elevation of a mudflat, owing to the combination of moderate hydrodynamic stress and high bioturbator activity. Also, bio-mediated sediment resuspension is predicted to be particularly high in the warm season. Research into metabolic dependency of bio-mediated sediment resuspension may help to place phenomenological observations in the broader framework of metabolic theories in ecology and to formulate general expectations on the coastal ecosystem functioning.

Predictive models accounting for the effect of bioturbation on sediment resuspension must be based on ecological theory as well as on empirical parametrizations. The scaling trend of individual metabolic and activity rates with body mass may be a key to the mechanistic understanding of the observed patterns. With this study we tested if general size scaling rules in bio-mediated sediment resuspension may apply to a broad range of physical contexts for the endobenthic bivalve Cerastoderma edule. The effect on sediment resuspension of populations of C. edule differing by individual size was measured across physical gradients of current velocity and sediment composition in terms of fraction of fine particles. C. edule were able to enhance the resuspension of sediment containing silt, while they had scarce effect on the resuspension of coarse sediment. The effect of bioturbation was maximal at intermediate current velocity, when the hydrodynamic forcing is not strong enough to overcome the abiotic sediment resistance but it is able to suspend the bioturbated sediment. Although differences in sediment silt content and intensities of hydrodynamic stress have a relevant influence in determining the bioturbators individual contribution to sediment resuspension, the observed mass scaling trend is consistent across all treatments and close to theoretical expectation for size scaling of individual metabolic rates. This observation supports the hypothesis that the contribution of individual bioturbators to sediment resuspension is directly related to their energy use. Therefore, the proposed approach allows the formulation of expectations of biotic contribution to sediment resuspension based on the general size scaling laws of individual energy use.