Recent research indicates that many ecosystems, including intertidal marshes, follow the alternative stable states theory. This theory implies that thresholds of environmental factors can mark a limit between two opposing stable ecosystem states, e.g. vegetated marshes and bare mudflats. While elevation relative to mean sea level is considered as the overall threshold condition for colonization of mudflats by vegetation, little is known about the individual driving mechanisms, in particular the impact of waves, and more specifically of wave period. We studied the impact of different wave regimes on plants in a full scale flume experiment. Seedlings and adult shoots of the pioneer Scirpus maritimus were subjected to two wave periods at two water levels. Drag forces acting on, and sediment scouring occurring around the plants were quantified, as these are the two main mechanisms determining plant establishment and survival. Depending on life stage, two distinct survival strategies emerge: seedlings present a stress avoidance strategy by being extremely flexible, thus limiting the drag forces and thereby the risk of breaking. Adult shoots present a stress tolerance strategy by having stiffer stems, which gives them a higher resistance to breaking. These strategies work well under natural, short period wind wave conditions. For long period waves, however, caused e.g. by ships, these survival strategies have a high chance to fail as the flexibility of seedlings and stiffness of adults lead to plant tissue failure and extreme drag forces respectively. This results in both cases in strongly bent plant stems, potentially limiting their survival.

Nature-based mitigation is increasingly proposed as a strategy to cope with global change and related risks for coastal flooding and erosion. Tidal marshes are known to provide shoreline protection as their aboveground biomass attenuates waves and their belowground biomass contributes to reducing erosion rates. The aim of this study was to quantify how effectively wave attenuation rates and erosion reduction rates are sustained throughout seasons in pioneer tidal marshes in the Elbe estuary (Germany). Changes in hydrodynamics and sediment dynamics were measured during 17 months along three sea-to-land transects of 50 m length. Simultaneously, changes in biomass of the monospecific pioneer vegetation (Bolboschoenus maritimus) were measured monthly. This study shows that wave and flow attenuation rates positively correlate with seasonal variations in aboveground biomass, that is: in summer, aboveground biomass and associated wave and flow attenuation rates are highest; while aboveground biomass is washed away during the first storms in autumn or winter, resulting in low wave and flow attenuation rates. Contrastingly, maximum incoming wave heights and flow velocities occur during winter, indicating that wave and flow attenuation is most needed then. However, hibernating root biomass assures low erosion rates in winter, especially at sandy sites. Although wave attenuation by pioneer marshes is highly variable throughout seasons and pioneer marshes alone are not so effective, they might facilitate the survival of higher marshes. Therefore, it is important to conserve or restore a gradual sea-to-land gradient from tidal flats, over pioneer marsh to high marsh to provide nature-based shoreline protection.

In hydrodynamically stressful environments, some species—known as ecosystem engineers—are able to modify the environment for their own benefit. Little is known however, about the interaction between functional plant traits and ecosystem engineering. We studied the responses of Scirpus tabernaemontani and Scirpus maritimus to wave impact in full-scale flume experiments. Stem density and biomass were used to predict the ecosystem engineering effect of wave attenuation. Also the drag force on plants, their bending angle after wave impact and the stem biomechanical properties were quantified as both responses of stress experienced and effects on ecosystem engineering. We analyzed lignin, cellulose, and silica contents as traits likely effecting stress resistance (avoidance, tolerance). Stem density and biomass were strong predictors for wave attenuation, S. maritimus showing a higher effect than S. tabernaemontani. The drag force and drag force per wet frontal area both differed significantly between the species at shallow water depths (20 cm). At greater depths (35 cm), drag forces and bending angles were significantly higher for S. maritimus than for S. tabernaemontani. However, they do not differ in drag force per wet frontal area due to the larger plant surface of S. maritimus. Stem resistance to breaking and stem flexibility were significantly higher in S. tabernaemontani, having a higher cellulose concentration and a larger cross-section in its basal stem parts. S. maritimus had clearly more lignin and silica contents in the basal stem parts than S. tabernaemontani. We concluded that the effect of biomass seems more relevant for the engineering effect of emergent macrophytes with leaves than species morphology: S. tabernaemontani has avoiding traits with minor effects on wave attenuation; S. maritimus has tolerating traits with larger effects. This implies that ecosystem engineering effects are directly linked with traits affecting species stress resistance and responding to stress experienced.

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