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The limiting effects of stressors like desiccation, light and salinity on seagrass growth and distribution are well-studied. However, little is known about their interactive effects, and whether such effects might differ among populations that are adapted to different local conditions. In two laboratory experiments we tested (a) if growth and development of intertidal, temperate Zostera noltii is affected by emergence time (experiment 1 and 2), and (b) how this is affected by an additional, second stressor, namely shading (experiment 1) or high salinity (25, 30 and 35, experiment 2). In addition, we tested (c) whether the effects of emergence time and salinity varied between three different European seagrass populations (Saint-Jacut/France, Oosterschelde/The Netherlands, and Sylt/Germany), which are likely adapted to different salinity levels (experiment 2). In both experiments, emergence of 8 h per tidal cycle (of 12 h) had a negative effect on seagrass relative growth rate (RGR), and aboveground biomass. Emergence furthermore reduced either rhizome length (experiment 1) or belowground biomass (experiment 2). Shading (experiment 1) resulted in lower RGR and a two-fold higher aboveground/belowground ratio. We found no interactive effects of emergence and shading stress. Salinity (experiment 2) did not affect seagrass growth or morphology of any of the three populations. The three tested populations differed greatly in morphology but showed no differential response to emergence or salinity level (experiment 2). Our results indicate that emergence time and shading show an additive negative effect (no synergistic or antagonistic effect), making the plants still vulnerable to such combination, a combination that may occur as a consequence of self-shading during emergence or resulting from algal cover. Emergence time likely determines the upper limit of Z. noltii and such shading will likely lower the upper limit. Shading resulted in higher aboveground/belowground ratios as is a general response in seagrass. Z. noltii of different populations originating from salinity 30 and 35 seem tolerant to variations in salinity within the tested range. Our results indicate that the three tested populations show morphotypic rather than ecotypic variation, at least regarding the salinity and emergence, as there were no interactive effects with origin. For restoration, this implies that the salinity regime of the donor and receptor site of Z. noltii is of no concern within the salinity range 25–35.

Changes in the seascape often result in altered hydrodynamics that lead to coinciding changes in sediment dynamics. Little is known on how altered sediment dynamics affect long-term seagrass persistence. We studied the thresholds of sediment dynamics in relation to seagrass presence by comparing sediment characteristics and seagrass presence data of seven separate seagrass meadows. All meadows had a long-term (>20 years) presence. Within these meadows, we distinguish so-called “hotspots” (areas within a meadow where seagrass was found during all mapping campaigns) and “coldspots” (with infrequent seagrass presence). We monitored static sediment characteristics (median grain size, bulk density, silt content) and sediment dynamics (that is, bed level change and maximum sediment disturbance depth), bioturbation (that is, lugworm densities and induced fecal pit and mound relief), and seagrass cover. We statistically analyzed which sediment characteristic best explains seagrass cover. Densely vegetated hotspots were shown to have lower sediment dynamics than sparsely vegetated hotspots and coldspots, whereas static sediment characteristics were similar (grain size, bulk density). The vegetation cover was either low (2–15%) or high (>30%) and sediment dynamics showed a threshold for vegetation cover. From this correlative finding, we postulate a self-sustaining feedback of relatively dense seagrass via sediment stabilization and accordingly a runaway feedback once the seagrass cover becomes too sparse. The sensitivity for sediment dynamics shown in our study implies that future existence of seagrass meadows may be at risk as ongoing climate change might directly (increased environmental extremes) or indirectly (changing seascapes) negatively affect seagrass beds.

1. Restoration of key species in dynamic coastal ecosystems benefits from reduction in environmentalstress. This can be realized by promoting positive feedback (intrinsic processes) orby reducing extrinsic negative forcing.2. In a seagrass (Zostera noltii) restoration project in the south-western Netherlands, weinvestigated transplantation success in relation to intrinsic processes (i.e. comparing sods vs.single shoots, transplant size, transplant configuration and transplant density) and extrinsicforcing (i.e. bioturbation by Arenicola marina, desiccation and exposure to water dynamics).In total, 2600 m2 of seagrass sods were mechanically transplanted to six intertidal flats overthe course of 5 years.3. In total, 43% of sod transplants (2?25 m2) survived at the long term, whereas single shoottransplants failed within the first 3 months. The use of larger, or more compact (sod), transplantconfigurations had no long-term effect on survival, and initial densities did not affecttransplantation success either. Reducing desiccation stress increased the transplantation successduring the first growing season. Shielding transplants from bioturbating lugworms had apositive effect on long-term survival.4. Seagrass abundance in summer was related to spring abundance, whereas winter survivalwas not related to prior seagrass abundance. At four of the six intertidal flats, transplantsgradually decreased in size over time. At the other two, extensive colonization occurredaround the transplant areas in some years and was still partly present in 2015. A correlationwith the studied environmental parameters was not found.5. Synthesis and applications. Intrinsic processes favour transplantation development duringthe growing season, allowing positive feedback. Extrinsic processes favour the development ata longer time-scale (i.e. reduction in bioturbation, thus breaking the positive feedback of thebare state). Most surprisingly, the starting colonization of two out of six tidal flats could not berelated to environmental factors (hydrodynamics, light, emergence time, sediment characteristics,macro-algae and grazing). Environmental managers can improve transplantation successby restoring the positive feedback, reducing stress, but also via risk spreading by performingtransplants over wider areas. They thereby accept the complexity of processes and unpredictabletemporal and spatial variation in which transplantation sites turn out to be successful.

Key-ecosystems such as seagrass beds are disappearing on a global scale. In order to counter-act local loss of seagrass beds, seagrass restoration projects have been carried out with varying degrees of success. As seagrass biomass peaks in summer, most restoration projects are monitored during this period, while wintering processes are largely ignored. We here attempted to elucidate some important bottlenecks for wintering survival in temperate areas of the intertidal model species Zostera noltii, using a three-year transplant-monitoring dataset and a field experiment. We found that next year's transplant success could not be predicted based on the preceding year's growth success, emphasizing the winter as a crucial period for survival of seagrass transplants. In addition, transplant success was neither determined by abiotic site characteristics. Low autumn rhizome starch concentrations in unsuccessful transplants, compared to successful transplants and natural beds, hinted at the importance of starch for winter survival. Hence, we tested the importance of starch, accumulated in autumn, versus the importance of the presence of sparse aboveground photosynthetic winter biomass for winter survival of seagrass transplants in a field experiment. We clipped the overwintering-leaves of three natural beds that naturally varied in their autumn rhizome starch concentrations. Decreased leaf densities in winter did not affect seagrass biomass in June, nor did this treatment affect rhizome starch concentration in June. Autumn rhizome starch reserves did however provide a good indication of next year's growth success, confirming the importance of starch reserves for winter survival. We thus conclude that autumn rhizome starch can be a good predictor of next year's growth success, whereas the preceding growing season shoot density and the presence of leaves during the winter were bad indicators of next year's growth success.