Anthropogenic nitrogen pollution impacts saltmarsh resilience with inhibition of seedling establishment and population dispersal.

Saltmarsh, a prominent buffer ecosystem, has been identified as an important sink for nitrogen (N) pollutants from marine- and land-based anthropogenic activities. However, how the enriched anthropogenic N impacts saltmarsh sustainability has been neglected due to limited understanding of marsh resilience based on seedling establishment and population dispersal under anthropogenic N inputs. This study combined mesocosm experiments and model simulations to quantify the effects of increased anthropogenic N on the seedling-based vegetation expansion of Spartina alterniflora. The results indicated that seedling survivals, growth rates, and morphological indicators were inhibited by 20.08 %, 37.14 %, and > 35.56 %, respectively, under 1.5 gN/kg anthropogenic N. The sensitivity rate of vegetation expansion was increased by 70 % with 1 gN/kg increased N concentration under the scenario of low seedling density (< 15 m/yr). These findings revealed an important unidentified weakness of the marsh development process to anthropogenic N inputs. Finally, we highlighted the importance of appropriate protection measures to control nutrient pollution in salt marshes. Our study provides new insights for enhancing the resilience and sustainability of saltmarsh ecosystems.

Tidal channel meanders serve as stepping-stones to facilitate cordgrass landward spread by creating invasion windows.

Understanding the mechanisms by which the geomorphic structures affect habitat invasibility by mediating various abiotic and biotic factors is essential for predicting whether these geomorphic structures may provide spatial windows of opportunity to facilitate range-expansion of invasive species in salt marshes. Many studies have linked geomorphic landscape features such as tidal channels to invasion by exotic plants, but the role of tidal channel meanders (i.e., convex and concave sides) in regulating the Spartina alterniflora invasion remains unclear. Here, we examined the combined effects of tidal channel meander-mediated hydrodynamic variables, soil abiotic stresses, and propagule pressure on the colonization of Spartina in the Yellow River Delta, China, by conducting field observations and experiments. The results showed that lower hydrodynamic disturbance, bed shear stress, and higher propagule pressure triggered by eddies due to the convex structure of channel meanders facilitated Spartina seedling establishment and growth, whereas the concave side considerably inhibited the Spartina invasion. Lower soil abiotic stresses also significantly promoted the invasibility of the channel meanders by Spartina. Based on these findings, we propose a conceptual framework to illustrate the effects of the meandering geomorphology of tidal channels on the mechanisms that might allow the landward spread of Spartina and related processes. Our results demonstrate that the meandering geomorphic structures of tidal channels could act as stepping-stones to significantly facilitate the landward invasion of Spartina along tidal channels. This implies that geomorphic characteristics of tidal channels should be integrated into invasive species control and salt marsh management strategies.

Invasive Spartina alterniflora habitat forms high energy fluxes but low food web stability compared to adjacent native vegetated habitats.

Invasive Spartina spp. mostly colonizes a bare tidal flat and then establishes a new vegetated habitat, where it promotes the productivity of local ecosystems. However, it was unclear whether the invasive habitat could well exhibit ecosystem functioning, e.g. how its high productivity propagates throughout the food web and whether it thereby develops a high food web stability relative to native vegetated habitats. By developing quantitative food webs for a long-established invasive Spartina alterniflora habitat and adjacent native salt marsh (Suaeda salsa) and seagrass (Zostera japonica) habitats in China's Yellow River Delta, we investigated the distributions of energy fluxes, assessed the stability of food webs, and investigated the net trophic effects between trophic groups by combining all direct and indirect trophic interactions. Results showed that the total energy flux in the invasive S. alterniflora habitat was comparable to that in the Z. japonica habitat, whereas 4.5 times higher than that in the S. salsa habitat. While, the invasive habitat had the lowest trophic transfer efficiencies. Food web stability in the invasive habitat was about 3 and 40 times lower than that in the S. salsa and Z. japonica habitats, respectively. Additionally, there were strong net effects caused by intermediate invertebrate species in the invasive habitat rather than by fish species in both native habitats. This study revealed the contradiction between the promotion of energy fluxes and the decrease of food web stability resulting from the invasion of S. alterniflora, which provides new insights into the community-based management of plant invasions.

A new perspective on the impacts of Spartina alterniflora invasion on Chinese wetlands in the context of climate change: A case study of the Jiuduansha Shoals, Yangtze Estuary.

Spartina alterniflora, an invasive plant, was introduced to the Chinese coastal zone in the early 90s. As an eco-engineering species, S. alterniflora not only alters saltmarsh species distributions, previously described as habitat degradation, but it also plays a vital role in coastal protection, especially for the development of recently emerged intertidal shoals. To provide a reference for coastal management under global change, we quantified the impact of the invasion process on provided ecological and coastal protection functions, exemplified at the emerging Jiuduansha Shoals (JDS) in the Yangtze Estuary. Results obtained by high-precision satellite monitoring and numerical modelling showed that the establishment and growth of S. alterniflora can exert considerable changes on local environment. The invasion of S. alterniflora to JDS wetland can be divided into three distinct phases, (1) establishment 1998-2003, (2) expansion 2003-2009, and (3) dominant 2009-2018 stages according to the changes in saltmarsh composition. Spatially, S. alterniflora continuously replaced Scirpus mariqueter, forcing S. mariqueter and Phragmites australis slowly to the lower and higher intertidal habitats, respectively. Notably, S. alterniflora expansion was the main driver that contributed to over 70 % of recent JDS wetland expansion even under sediment deficit conditions. Established S. alterniflora marsh (directly) dampens more waves because of aboveground stems, but it also causes more accretion and indirectly leads to higher "morphological" wave dampening. Thus, it increases coastal defense provided by the saltmarsh in the context of sea-level rise and strengthening storms. In conclusion, the role of S. alterniflora invasion to the local environment under global changes is controversial. For sustainable coastal management, we need context-dependent S. alterniflora management to maximize the benefit of coastal protection and minimize the impact on local ecology, especially in sediment-starving estuaries with expected coastline retreat.

Effects of waves, burial depth and material density on microplastic retention in coastal sediments.

Coastal sediments, recognized as a major sink for microplastics (MPs), are subject to frequent physical disturbances, such as wave disturbance and associated sediment dynamics. Yet it remains poorly understood how wave disturbance regulates MPs accumulation in such a dynamic environment. Here, we examined the effects of waves and their interactions with material density and burial depth on the retention of MPs in coastal sediments, through manipulative experiments in a mangrove habitat along the coast of South China. The results clearly revealed that stronger waves removed more buried MPs from the sediments. Moreover, storms can have disproportional effects on MPs retention by inducing large waves and strong sediment erosion. We also demonstrated that MPs retention generally increased linearly with growing material density and non-linearly with raised burial depth in the sediment. Overall, our findings highlight the importance of both external and internal factors in shaping MPs retention in coastal ecosystems like mangroves, which is essential to assess and predict MPs accumulation patterns as well as its impacts on ecosystem functioning of such blue carbon habitats.

Bioaccumulation of emerging persistent organic pollutants in the deep-sea cold seep ecosystems: Evidence from chlorinated paraffin.

Persistent organic pollutants (POPs) are highly toxic and can accumulate in marine organisms, causing nonnegligible harm to the global marine ecosystem. The Cold seep is an essential marine ecosystem with the critical ecological function of maintaining the deep-sea carbon cycle and buffering global climate change. However, the environmental impact of emerging POPs in the deep-sea cold seep ecosystem is unknown. Here, we investigated the potential pollution of chlorinated paraffins (CPs) and their bioaccumulation in the cold seep ecosystem. High concentrations of CPs were detected in the cold seep ecosystems, where CPs bioaccumulated by the keystone species of deep-sea mussels can be released into the surface sediment and vertically migrate into the deeper sediment. Furthermore, more toxic CPs were accumulated from transforming other CPs in the cold seep ecosystem. Our study provides the first evidence that high concentrations of POPs are bioaccumulated by deep-sea mussels in the cold seep ecosystem, causing adverse ecological effects. The discovery of CPs bioaccumulation in the deep-sea cold seep ecosystem is a crucial mechanism affecting deep-sea carbon transport and cycling. This study has important guiding significance for revealing the deep-sea carbon cycle process, addressing global climate change, and making deep-sea ecological and environmental protection policies.

The role of bio-geomorphic feedbacks in shaping microplastic burial in blue carbon habitats.

Coastal sediments are considered as hotspots of microplastics (MPs), with substantial MPs stocks found in blue carbon habitats such as mangroves and tidal marshes, where wave-damping vegetation reduces sediment erosion and enhances accretion. Here, we examined the effects of such bio-geomorphic feedbacks in shaping MPs burial, through a year-round field study in a mangrove habitat along the coast of South China. The results revealed that MPs abundance decreased significantly with the increase of cumulative sediment erosion as the strength of bio-geomorphic feedbacks declined. More shapes and colors of MPs were found at locations with weaker waves and less sediment erosion, where the average particle size was also higher. Our findings highlight the importance of bio-geomorphic feedbacks in affecting both the abundance and characteristics of the buried MPs. Such knowledge extends our understanding of MPs transport and burial from the perspective of bio-geomorphology, which is essential to assess and predict MPs accumulation patterns as well as its impacts on ecosystem functioning of the blue carbon habitats.

Morphological wave attenuation of the nature-based flood defense: A case study from Chongming Dongtan Shoal, China.

The risk of coastal storm flooding is deteriorating under global warming, especially for the heavily urbanized deltaic cities, like Shanghai. The Nature-Based Flood Defense (NBFD), as an eco-friendly design alternative for hard infrastructure against coastal flooding, is gaining attention. Nevertheless, the vulnerability of saltmarsh due to the biological instability, resulting in the uncertainties on coastal protection, is considered the bottleneck challenge that hinders the broad application of the NBFD concept. We argue that except for direct wave attenuations by the above-ground vegetation during storms, the gradual sediment trapping and consolidating during the non-storm period is a more crucial function of coastal saltmarsh, which mitigates storm waves by forming a broader and higher intertidal morphology. This benefit is an important value of saltmarsh-based coastal protection but is largely neglected in many NBFD studies. Taking Chongming Dongtan Shoal (CDS) as a case study, we demonstrated that over 2/3th wave attenuation during storms is contributed by the saltmarsh morphology, and less than 1/3th is from the saltmarsh vegetation. The relative contribution of the saltmarsh morphology on wave mitigation is even enhanced under the increasing storm grades from 100 yrs. to 5000 yrs. return levels. To promote this idea for broader application, the cost-benefit analysis of three artificial NBFD solutions (e.g., submerged breakwater, timber piles, and sand nourishment) are compared. We identified an optimal measure of the submerged breakwater for CDS, which minimizes the ecological impact and maximizes the cost-benefit. Moreover, the wave-free zone behind the breakwater increases the chance of vegetation establishment, helps suspended sediment trapping, hence fostering a beneficent cycle for saltmarsh restoration. In summary, ignoring the contribution of saltmarsh morphology on wave attenuation largely underestimated the benefits of vegetation-based coastal protection, which should be greatly emphasized to provide a solid basis for developing NBFD.

Impact of off-bottom seaweed cultivation on turbulent variation in the hydrodynamic environment: A flume experiment study with mimic and natural Saccharina latissima thalli.

The seaweed industry is growing worldwide to meet future resource needs in terms of food and fuel. In the meantime, the impact of expanding off-bottom seaweed cultivation on its environment is unclear. For example, it remains poorly understood how off-bottom seaweeds affect the local hydrodynamic environment, especially concerning turbulence that is more important for nutrient transport and availability than the mean flow velocity. Here, we carried out well-controlled flume experiments with mimic seaweed thalli, which are available, controllable, and stable, to investigate the impact of off-bottom seaweed canopies on whole-depth flow velocities in terms of both mean flow and turbulence velocity profiles. A careful comparison of behavior in the flow between natural and mimic seaweed thalli was made before these experiments. The results show that the floating seaweed thalli generate a surface boundary layer and have a profound impact on the velocity structure in the bottom boundary layer. More importantly, the generation, growth and dissipation of turbulence in the seaweed thalli area deeply affect the downstream distribution of near-bed turbulent strength and associated bed shear stress. Ignoring this turbulent variation would cause inaccurate predictions of morphological changes of the seabed. Our findings suggest that expanding the seaweed cultivation area may cause high risks of bed degradation and low diffusion in the downstream cultivation area. These findings provide novel insights into the environmental influence of off-bottom seaweed cultivation, with important implications for optimizing management strategies to promote seaweed productivity while minimizing seabed destabilization.

Can the native faunal communities be restored from removal of invasive plants in coastal ecosystems? A global meta-analysis.

Coastal ecosystems worldwide are being threatened by invasive plants in the context of global changes. However, how invasive plants influence native faunal communities and whether native faunal communities can recover following the invader removals/controls across global coastal ecosystems are still poorly understood. Here, we present the first global meta-analysis to quantify the impacts of Spartina species invasions on coastal faunal communities and further to evaluate the outcomes of Spartina species removals on faunal community recovery based on 74 independent studies. We found that invasive Spartina species generally decreased the biodiversity (e.g., species richness), but increased coastal faunal abundance (e.g., individual number) and fitness (e.g., biomass), though the effect on abundance was insignificant. The pattern of influence was strongly dependent on habitat types, faunal taxa, trophic levels, and feeding types. Specifically, Spartina species invasion of mudflats caused greater impacts than invasion of vegetated habitats. Insects and birds at higher trophic levels were strongly affected by invasive Spartina, indicating that invasive plant effects can cascade upward along the food chain. Additionally, impacts of Spartina invasions were more obvious on food specialists such as herbivores and carnivores. Furthermore, our analyses revealed that invader removals were overall beneficial for native faunal communities to recover from the displacement caused by Spartina invasions, but this recovery process depended on specific removal measure and time. For example, the long-term waterlogging had strong negative impacts on faunal recovery, so it should not be encouraged. Our findings suggest that invasive plants could have contrasting effects on functional responses of native faunal communities. Although invasive plant removals could restore native faunal communities, future functional restorations of invaded ecosystems should take the legacy effects of invasive species on native communities into account. These findings provide insightful implications for future scientific controls of invasive species and ecosystem restoration under intensifying global changes.

Saltmarsh resilience controlled by patch size and plant density of habitat-forming species that trap shells.

Habitat fragmentaion into small patches is regarded as a vital cause of biodiversity loss. Fragmentationof habitat-forming species is especially harmful, as patchiness of such species often controls ecosystem stability and resilience by density and patch size-dependent self-reinforcing feedbacks. Although fragmentation are expected to weaken or even break such feedbacks, it remains unclear how the resulting patchiness of habitat-forming species affect ecosystem resilience to environmental stresses. Here, using Spartian alterniflora, the habitat-forming species in saltmarshes as a model, we investigate how patch size, plant density, and shell aggregation interactively control the persistence of a degrading salt marsh that suffered from erosion induced by hydrodynamics. Our results demonstrate that large patches can trap more shells along the patch edge than the smaller ones, therefore significantly facilitating plant re-growth within the patch. Shell removal experiments further reveal that large patches trapping more shells along patch edges reinforce their own persistence by decreasing erosion and thus facilitating plant recovery. By contrast, small patches with lesser plants cannot persist as they trap less shells along patch edges but are able to accumulate more shells at interior locations where they hinder plant re-growth, indicating a critical threshold of patch size ~20 m2 below which ecosystem collapses. The current study highlights the importance to identify critical threshold of stress-resistant patch sizes in transition-prone ecosystems as early-warning to alert undesired ecosystem collapse and restoration practice.

Biomechanical properties of marsh vegetation in space and time: effects of salinity, inundation and seasonality.

BACKGROUND AND AIMS: Over the last decade, the importance of plant biomechanical properties in shaping wave dissipation efficiency of marsh vegetation has gained growing attention. Here we provide the first analyses of how biomechanical stem properties vary with seasons and along environmental gradients in coastal and estuarine marshes, which is essential to enable accurate assessments of flood defence value of marsh vegetation. METHODS: We quantified both spatial and seasonal variation in stem flexibility and breakability for a variety of common marsh vegetation (Spartina anglica, Scirpus maritimus, Phragmites australis, Elymus athericus, Suaeda maritima, Aster tripolium, Saliconia procumbens) distributed along both salinity and inundation gradients. KEY RESULTS: Increasing salinity tends to induce a shift from species with tall shoots, high flexural stiffness (stem resistance to bending; N mm2) towards species with shorter and more flexible stems. The same trend was found with increasing inundation stress (i.e. decreasing elevation) from the higher part of the low marsh towards the pioneer zone. Stem breakability (the force required to break or fold a stem, N) followed the same pattern of stem stiffness due to the positive relationship between flexural strength (material resistance to flexure, N mm-2) and Young's bending modulus (material resistance to bending; N mm-2). Shifts in stem stiffness and breakability at the community level were found to relate positively to the variation in canopy height between species, highlighting the concurrence of changes in morphological and biomechanical traits under environmental changes. Compared to the differences between species, within-species variability between sampling locations and between seasons is generally minor. CONCLUSIONS: Our findings imply that environmental changes may significantly modify wave attenuation capacity of coastal vegetation by inducing species shifts. This emphasizes the need to understand the response of community composition to climate change and human disturbances, when using nature-based flood protection by coastal vegetation as an adaptive response to global change.

Wave effects on seedling establishment of three pioneer marsh species: survival, morphology and biomechanics.

BACKGROUND AND AIMS: It is important to have an in-depth mechanistic understanding of tidal marsh establishment and dynamics to ensure the long-term persistence of these valuable ecosystems. As wave forcing may be expected to impact seedling establishment, we studied the effect of water-imposed drag forces on seedling survival, morphology and biomechanical properties of three marsh pioneer species that are dominant along the salinity gradient in many areas around the world: Spartina anglica (salt to brackish), Scirpus maritimus (brackish) and Phragmites australis (brackish to fresh). METHODS: Using a newly developed plant-shaking mesocosm (PSM) that mimicked water-imposed wave drag forces, the effect of wave stress on seedling survival was examined, together with impacts on morphology and biomechanical properties. KEY RESULTS: After 7 weeks of exposure to wave stress, lowered seedling survival and growth for all species was revealed. Wave treatments increased the root/shoot biomass ratio to enhance anchorage and made seedlings more flexible (i.e. reduced flexural rigidity), which might be regarded as a mixed outcome between a stress avoidance and stress tolerance strategy. CONCLUSIONS: The different biomechanical responses between the three dominant marsh pioneer species, overall, make them less resistant to external stress. Therefore, our results indicate that the likelihood of marshes becoming established is reduced if wave energy increases. Despite the different biomechanical response of these three pioneer species to waves, the seedlings of all species were found to have low resistance to external stresses.

Sprouting as a gardening strategy to obtain superior supplementary food: evidence from a seed-caching marine worm.

Only a handful of non-human animals are known to grow their own food by cultivating high-yield fungal or algal crops as staple food. Here we report an alternative strategy utilized by an omnivorous marine worm, Hediste diversicolor, to supplement its diet: gardening by sprouting seeds. In addition to having many other known feeding modes, we showed using video recordings and manipulative mesocosm experiments that this species can also behave like gardeners by deliberately burying cordgrass seeds in their burrows, which has been previously shown to reduce the loss of seeds to water. These seeds, however, are protected by the seed husk, and we used feeding experiments to show that they were not edible for H. diversicolor until they had sprouted or the seed husk had been artificially removed. Additionally, sprouts were shown to be highly nutritious, permitting higher growth rates in H. diversicolor than the low-quality basal food, detritus. We propose both a proximate cause (seed husk as a physical barrier) and ultimate cause (nutritional demand) for this peculiar feeding behavior. Our findings suggest that sprouting may be a common strategy used by seed-collecting animals to exploit nutrients from well-protected seeds.

[Reinvasion of exotic plant species Spartina alterniflora in Chongming Dongtan Nature Reserve of Shanghai].

A demonstration plot in Chongming Dongtan for controlling exotic plant species Spartina alterniflora by using an integrated treatment technique of tussock cutting plus water level controlling was selected to make a 2-year monitoring on the reinvasion process of this plant species. A large number of S. alterniflora seedlings could proliferate and settle down in the hydrological restoring plots by tidal water in spring, and, after two years, the reinvasion community was established, with no obvious differences in its density as compared to the communities in neighboring areas. However, the distance of extension via vegetative propagation was limited. At the plots maintaining waterlogging, the distance of 2-year lateral extension was less than 1 m; at physically isolated plots, there was no any reinvasion of S. alterniflora. The rapid spreading of the seedlings in spring was the key of the fast reinvasion of S. alterniflora, and thus, to completely eradicate the spreading source from neighboring areas, to build isolation barrier to stop the spreading of this plant, and to adopt biological substitute by planting a native species such as Phragmites australis would be the main countermeasures to effectively prevent the reinvasion of S. alterniflora.