Coastal vegetation on the Western, Southern, and Eastern coasts of South Korea

We used the Braun-Blanquet method to study the vegetation of coastal wetlands in South Korea. Three habitat types were found, i.e., salt marshes, salt swamps, and sand dunes. These plant communities were classified as: 1) two groups (five associations each) in the salt marshes that comprised either annual herbaceous halophytes (ClassThero-Salicornietea), or biennial/perennial herbaceous species (ClassAsteretea tripolii); 2) one group in the salt swamps consisting of five hydrophilous halo-tolerant associations (ClassPhragmitetea); and 3) three groups in the sand dunes, including one association of annual herbaceous halophytes (ClassSalsoletea komarovii), seven associations of herbaceous perennial halophytes (ClassGlehnietea littoralis), and one association of shrub perennial halophytes (ClassVrticetea rotundifoliae). These three habitat types accounted for the majority of the six main classifications of coastal vegetation distributed in South Korea.     

Factors responsible for Honckenya peploides (Caryophyllaceae) and Leymus mollis (Poaceae) spatial segregation on subarctic coastal dunes

Low water and nutrient availability and significant sand movement, salt spray, and soil salinity are typical of coastal dunes. These conditions are generally unfavorable for the various life stages of plants and especially for seedlings. However, the intensity of these stresses decreases landward, even over short distances, with significant effects on community composition. On coastal dunes in subarctic Québec, Canada, Honckenya peploides (Caryophyllaceae) colonizes the upper beach where it forms small mounds called embryo dunes. Leymus mollis (Poaceae) is mostly restricted to the foredune; however, a few individuals successfully establish on the upper beach, particularly on embryo dunes. We hypothesized that this differential distribution is associated with differences in the tolerance of the two species' seedlings to physical stresses. Honckenya peploides and L. mollis seedling tolerance to sand burial, salt spray, soil salinity, and nutrient and water availability was assessed in greenhouse experiments. Unexpectedly, our results showed that tolerance to sand burial, salt spray, and soil salinity was lower for H. peploides than for L. mollis. If seeds are available and seedlings tolerate the conditions prevailing on the upper beach well, why are mature L. mollis individuals rare in this habitat? We suggest that massive abrasion events (e.g., violent storm waves and ice thrust) restrict the presence of the species on the upper beach.     

Vegetation analysis of the Mediterranean region of Nile Delta

Thirty-four vegetation clusters identified in the present study, after the application of TWINSPAN and DCA multivariate techniques, were assigned into 8 vegetation types, each of definite vegetation and habitat characters. The suggested vegetation types are well segregated along the DCA axis one which reflects soil moisture, salinity (as indicated by EC values), fertility (as indicated by the organic matter and nitrogen contents) and species diversity gradients. In general, soil moisture and soil fertility increase and species diversity decreases with the following sequence of vegetation types:Echinops spinosissimus-Ononis serrata on inland sand dunes,Pancratium maritimum on coastal sand dunes,Halocnemum strobilaceum-Salsola kali in saline sand deposits,Atriplex halimus-Chenopodium murale along the terraces and slopes of drains,Arthrocnemum glaucum-Tamarix nilotica in salt marshes,Chenopodium murale along the slopes of drains,Phragmites australis along the littoral zones of drains, andLemna gibba-Potamogeton crispus in the water zone. This sequence reflects also a gradient of human interference, starting with the vegetation of the less disturbed habitats (sand dunes and saline sand deposits) and ending with the fully man-made habitats (drain zones).     

A mixed strategy in the annual endemic Aster laurentianus (Asteraceae)--a stress-tolerant, yet opportunistic species

Several environmental factors influence the distribution of plants in coastal salt marshes. Substrate salinity is among the major factors preventing several species from establishing near the water line. However, interspecific competition for light and nutrients is often significant in determining the upper limit of plants along the salt marsh gradient. In this study, we tested the effects of substrate salinity and light and nutrient availability on the performance of the annual Aster laurentianus (Asteraceae), an endangered species of eastern Canadian salt marshes. This species is typically found in a narrow band along the shores of shallow lagoons, cornered between the high water line and the dense, herbaceous community of the upper marsh. Low light availability was the most significant factor limiting plant performance. Salinity had little effect on A. laurentianus as, unexpectedly, did nutrient availability. Yet plants were able to absorb nutrients when these were made more available. Luxury consumption, the uptake of excess nutrients, may make sense for this annual plant because the habitat in which it grows is subject to frequent disturbances (e.g., sand accretion and salinity pulses) that may kill canopy species and release suppressed A. laurentianus individuals. These results suggest that interspecific competition for light may play a significant role in restraining A. laurentianus from the upper part of salt marshes. Luxury consumption may help the species to opportunistically take advantage of release from taller species, particularly towards the upper edge of the salt marsh gradient.     

Are biotic factors significant in influencing the distribution of halophytes in saline habitats?

The influence of biotic factors on the distribution and establishment of halophytes is being considered in this review. Physicochemical factors, such as salinity and flooding, often are considered to be the determining factors controlling the establishment and zonational patterns of species in salt marsh and salt desert environments. Sharp boundaries commonly are found between halophyte communities even though there is a gradual change in the physicochemical environment, which indicates that biotic interactions may play a significant role in deterining the distribution pattern of species and the composition of zonal communities. Competition is hypothesized to play a key role in determining both the upper and lower limits of species distribution along a salinity gradient. Field and laboratory experiments indicate that the upper limits of distribution of halophytes into less saline or nonsaline habitats is often determined by competition. There appears to be a reciprocal relationship between the level of salt tolerance of species and their ability to compete with glycophytes in less saline habitats. Halophytes are not competitive in nonsaline habitats, but their competitive ability increases sharply in saline habitats. Allelopathic effects have been reported in salt desert habitats, but have not been reported along salinity gradients in salt marshes. Some species of halophytes that are salt accumulators have the ability to change soil chemistry. Chemical inhibition of intolerant species occurs when high concentrations of sodium are concentrated in the surface soils of salt desert plant communities that are dominated by salt-accumulating species. Establishment of less salt-tolerant species is inhibited in the vicinity of these salt-accumulating species. Herbivory is reported to cause both an increase and a decrease in plant diversity in salt marsh habitats. Heavy grazing is reported to eliminate sensitive species and produce a dense cover of graminoids in high marsh coastal habitats. However, in other marshes, grazing produced bare patches that allowed annuals and other low marsh species to invade upper marsh zonal communities. A retrogression in plant succession may occur in salt marshes and salt deserts because of heavy grazing. Intermediate levels of grazing by sheep, cattle, and horses could produce communities with the highest species richness and heterogeneity. Grazing by geese produced bare areas that had soils with higher salinity and lower soil moisture than vegetated areas, allowing only the more salt-tolerant species to persist. Removal of geese from areas by use of inclosures caused an increase in species richness in subarctic salt marshes. Invertebrate herbivores could also inhibit the survival of seeds and the ability of plants to establish in marshes. Parasites could play a significant role in determining the species composition of zonal communities, because uninfected rarer species are able to establish in the gaps produced by the death of parasitized species.     

Recognition of co-existence pattern of salt marshes and mangroves for littoral forest restoration

Climate-change driven sea level rise causes a increase in salinity in coastal wetlands accelerating the alteration of the species composition. It triggers the gradual extinction of species, particularly the mangrove population which is intolerant of excessive salinity. Thus despite being crucial to a wide range of ecosystem services, mangroves have been identified as a vulnerable coastal biome. Hence restoration strategy of mangroves is undergoing rigorous research and experiments in literature at an interdisciplinary level. From a data-driven perspective, analysis of mangrove occurrence data could be the key to comprehend and predict mangrove behavior along different environmental parameters, and it could be important in formulating management strategy for mangrove rehabilitation and restoration. As salt marshes are the natural salt-accumulating halophytes, mitigating excessive salinity could be achieved by incorporating salt-marshes in mangrove restoration activities. This study intends to find a novel restoration strategy by assessing the frequent co-existence status of salt marshes, with the mangroves, and mangrove associates in different zones of degraded mangrove patches for species-rich plantation. To achieve this, we primarily design a novel methodological framework for the practice of knowledge discovery concerning the coexistence pattern of salt marshes, mangroves, and mangrove associates along with environmental parameters using a data mining paradigm of association rule mining. The proposed approach has the capability to uncover underlying facts and forecast likely facts that could automate the study in the field of ecological research to comprehend the occurrence of inter-species relationships. Our findings are based on published data gathered on the Sundarban Mangrove Forest, one of the world’s most important littoral forests. The existing literature reinforces the findings that include all the sets of frequently co-occurring mangroves, their associates, and salt marshes along the salinity gradient of coastal Sundarbans. A detailed understanding of the occurrence patterns of all these, along with the environmental variables, would be able to promote decision-making strategy. This framework is effective for both academia and stakeholders, especially the foresters/ conservation planners, to regulate the spread of salt marshes and the restoration of mangroves as well.     

Effects of sandblasting and salt spray on inland plants transplanted to coastal sand dunes

A transplant experiment was conducted on a sandy beach to elucidate whether salt spray and sandblasting are the major factors inhibiting inland plants from becoming established on coastal sand dunes. Potted inland plants of Miscanthus sinensis and Imperata cylindrica var. koenigii were transplanted in two zones on the beach and in one area far inland from the beach. One zone on the beach (sea side) was located on a front dune that was occupied by native sand-dune plants; the other zone (land side) was located behind the sand dunes, where grassland comprised both sand-dune and inland species. To assess the condition of transplants, we measured changes in the canopy leaf area periodically at all sites. The final dry weight at each site was determined at the end of the experiment. Seasonal changes in sandblasting and salt spray intensities were evaluated periodically at the sites by measurement of the opaqueness of exposed transparent plastic sheets and the amount of sodium trapped in exposed filter papers, respectively. All transplants died in the sea-side zone, where both salt spray and sandblasting were most frequent and intense. The final dry weight was greatest at the inland site, which lacked salt spray and sandblasting. Although salt spray was intense in the land-side zone, the canopy leaf area decreased considerably only in seasons during which salt spray was accompanied by intense sandblasting. We concluded that sandblasting accompanied by salt spray is one of the main factors inhibiting the survival and growth of inland plants on coastal sand dunes.     

Germination ecology of coastal plants in relation to salt environment

Responses of seed germination to salinity were examined using 37 species collected from salt marshes, cliffs, and fore (unstable) and hind (stable) sand dunes along Japanese coasts. For comparison, seed germination of nine inland species was also examined. The soil salinities in salt marshes ranged from 150 to 300 mmol/L NaCl, whereas those in fore and hind dunes ranged from 0 to 150 mmol/L NaCl, with a few exceptions. Cliff soils showed relatively high salinities up to 300 mmol/L NaCl. Ciff and foredune soils that encountered a typhoon and storm showed high salinities >300 mmol/L NaCl. Salt tolerance in seed germination of coastal plants was ordered by comparing the responses of percentage and rate of germination to salinity conditions up to 200 mmol/L NaCl, being in the order of salt marsh>cliff>foredune≅hind dune≅inland. Thse results indicate that salt tolerance in seed germination of coastal plants is closely related to the salinity conditions of their habitats. Germination experiments under favorable conditions showed that a high percentage of the seeds of salt marsh species germinate rapidly, those of diff species germinate slowly and those of foredune species exhibit a low percentage and low rate of germination. It seems that these germination characteristics contribute to the success of germination at the ‘safe site’ and the subsequent survivorship of emerged plants in their natural habitats.     

Salt-spray stimulated growth in strand-line species

The response to salt spray and soil salinity of two sand dune strandline species ( Cakile maritima Scop. and Salsola kali L.) and two salt marsh strand-line species ( Atriplex hastata L. and A. littoralis L.) was compared in sand-compost cultures. The growth of the salt-marsh species remained unaffected, while the growth of the sand dune species Cakile maritima was strongly reduced by NaCl (150 and 300 m M ) absorbed via the root system. All four species were resistant to airborne salinity, and under conditions of low soil fertility, salt spray increased the dry matter production, especially of the sand dune species. Mineral analysis revealed foliar uptake of Na, K, Cl, Ca and Mg. Na and Cl ions absorbed from seawater droplets induced succulence. Both salt spray and soil salt increased the methylated quaternary ammonium compound content in the shoot tissue. Under non-saline conditions a considerable amount of these osmotic solutes was still present, while turgor pressure potential in these plants was rather low. The relation between salt, compatible osmotic solutes, turgor pressure potential and growth is discussed. Next to the major constituents of seawater, Na and Cl, especially magnesium and to a lesser extent, calcium, accumulated in the shoot tissue. Based on the positive growth response of the sand dune species to airborne salt, they should be termed 'aerohalophytes', whereas 'soil halophytes' should be used when referring to the Atriplex species, which are more specifically adapted to the increased salinity of salt marsh soils.     

Plant functional traits and diversity in sand dune ecosystems across different biogeographic regions

Plant species of a functional group respond similarly to environmental pressures and may be expected to act similarly on ecosystem processes and habitat properties. However, feasibility and applicability of functional groups in ecosystems across very different climatic regions have not yet been studied. In our approach we specified the functional groups in sand dune ecosystems of the Mediterranean, Hyrcanian and Irano-Turanian phytogeographic regions. We examined whether functional groups are more influenced by region or rather by habitat characteristics, and identified trait syndromes associated with common habitat types in sand dunes (mobile dunes, stabilized dunes, salt marshes, semi-wet sands, disturbed habitats). A database of 14 traits, 309 species and 314 relevés was examined and trait-species, trait-plot and species-plot matrices were built. Cluster analysis revealed similar plant functional groups in sand dune ecosystems across regions of very different species composition and climate. Specifically, our study showed that plant traits in sand dune ecosystems are grouped reflecting habitat affiliation rather than region and species pool. Environmental factors and constraints such as sand mobility, soil salinity, water availability, nutrient status and disturbance are more important for the occurrence and distribution of plant functional groups than regional belonging. Each habitat is shown to be equipped with specific functional groups and can be described by specific sets of traits. In restoration ecology the completeness of functional groups and traits in a site may serve as a guideline for maintaining or restoring the habitat.     

Variable response of three Trifolium repens ecotypes to soil flooding by seawater

Background and Aims

Despite concerns about the impact of rising sea levels and storm surge events on coastal ecosystems, there is remarkably little information on the response of terrestrial coastal plant species to seawater inundation. The aim of this study was to elucidate responses of a glycophyte (white clover, Trifolium repens) to short-duration soil flooding by seawater and recovery following leaching of salts.

Methods

Using plants cultivated from parent ecotypes collected from a natural soil salinity gradient, the impact of short-duration seawater soil flooding (8 or 24 h) on short-term changes in leaf salt ion and organic solute concentrations was examined, together with longer term impacts on plant growth (stolon elongation) and flowering.

Key Results

There was substantial Cl^– and Na⁺ accumulation in leaves, especially for plants subjected to 24 h soil flooding with seawater, but no consistent variation linked to parent plant provenance. Proline and sucrose concentrations also increased in plants following seawater flooding of the soil. Plant growth and flowering were reduced by longer soil immersion times (seawater flooding followed by drainage and freshwater inputs), but plants originating from more saline soil responded less negatively than those from lower salinity soil.

Conclusions

The accumulation of proline and sucrose indicates a potential for solute accumulation as a response to the osmotic imbalance caused by salt ions, while variation in growth and flowering responses between ecotypes points to a natural adaptive capacity for tolerance of short-duration seawater soil flooding in T. repens. Consequently, it is suggested that selection for tolerant ecotypes is possible should the predicted increase in frequency of storm surge flooding events occur.     

Does salt stress constrain spatial distribution of dune building grasses Ammophila arenaria and Elytrichia juncea on the beach?

Rising sea levels threaten coastal safety by increasing the risk of flooding. Coastal dunes provide a natural form of coastal protection. Understanding drivers that constrain early development of dunes is necessary to assess whether dune development may keep pace with sea‐level rise. In this study, we explored to what extent salt stress experienced by dune building plant species constrains their spatial distribution at the Dutch sandy coast. We conducted a field transplantation experiment and a glasshouse experiment with two dune building grasses Ammophila arenaria and Elytrigia juncea. In the field, we measured salinity and monitored growth of transplanted grasses in four vegetation zones: (I) nonvegetated beach, (II) E. juncea occurring, (III) both species co‐occurring, and (IV) A. arenaria dominant. In the glasshouse, we subjected the two species to six soil salinity treatments, with and without salt spray. We monitored biomass, photosynthesis, leaf sodium, and nutrient concentrations over a growing season. The vegetation zones were weakly associated with summer soil salinity; zone I and II were significantly more saline than zones III and IV. Ammophila arenaria performed equally (zone II) or better (zones III, IV) than E. juncea, suggesting soil salinity did not limit species performance. Both species showed severe winter mortality. In the glasshouse, A. arenaria biomass decreased linearly with soil salinity, presumably as a result of osmotic stress. Elytrigia juncea showed a nonlinear response to soil salinity with an optimum at 0.75% soil salinity. Our findings suggest that soil salinity stress either takes place in winter, or that development of vegetated dunes is less sensitive to soil salinity than hitherto expected.     

盐度和淹水对长江口潮滩盐沼植物碳储量的影响

盐生植物是盐沼有机碳储存的"临时库",也是土壤有机碳累积的主要来源,其碳储量大小对盐沼生态系统"碳汇"功能的发挥十分重要。以长江口潮滩本地种芦苇(Phragmites australis)和海三棱藨草(Scirpus mariqueter),及入侵种互花米草(Spartina alterniflora)为研究对象,采用单因素盆栽实验,模拟分析淹水盐度(0、5、10、15、25和35)、淹水深度(0、10、20、40、60cm和80cm)和淹水频率(每天、每3天、每7天、每10天和每15天)变化对各盐生植物地上、地下和总体碳储量大小的影响。研究结果表明,随着淹水盐度增加,芦苇、互花米草和海三棱藨草地上部分与总体碳储量均显著降低。土壤盐度可分别解释其地上部分碳储量变异的47.2%、66.5%和72.7%,与总体碳储量变异的34.7%、45.0%和62.0%。随着淹水深度增加,芦苇地上部分、总体碳储量和海三棱藨草地上部分碳储量均显著降低,其变异的68.6%、28.5%和71.1%可由淹水深度变化(10—80cm)解释。互花米草在80cm淹水深度下仍有较高的地上部分碳储量和总体碳储量。3种盐生植物碳储量对淹水频率变化的响应差异均不显著,所有处理地下部分碳储量差异也未达到显著水平。总体而言,互花米草对水盐胁迫的耐受性要强于本地种芦苇和海三棱藨草。尽管互花米草和芦苇具有相对较高的碳储量,但水盐胁迫对其碳储量的显著抑制作用不容忽视。海三棱藨草碳储量本就不高,输入土壤的有机碳量较为有限,海平面上升及盐水入侵等逆境胁迫会使其对盐沼"碳汇"贡献更加微弱。     

Nature management in coastal areas of Western Europe

The general environmental characteristics of West European coastal dunes and those of salt marshes are outlined. It is pointed out that the dynamical character of the dune system with its interaction between plants and sand in building and degeneration phases has been known for a much longer period of time than generally believed.As to the salt marshes the special significance is discussed of ungrazed marshes; the majority of the marshes having been under a grazing regime for a long time is elucidated.The distinction between internal management is mentioned, i.e. the measures taken to maintain or improve the qualities of an area, versus external management, i.e. the measures to prevent or counteract negative influences from outside the area.Nomenclature follows Heukels & Van Ooststroom (1977).     

Données synchorologiques sur la végétation littorale européenne

Four aspects of the synchorology of plant communites along the European coastline are treated: (1) with examples from sand dunes (Agropyreta and Ammophileta) and salt marshes (Puccinellieta maritimae) synvicariance is elucidated; (2) The increase in synvicariance towards the inner sand dunes is discussed as a result of an increasing effect of autochthonous climatic features; (3) with examples from sand dunes and cliffs the concept of synendemism is elucidated; (4) synvicariance at the landscape (geosigmetum) is discussed with examples from the French coastline.

     

Growth, sodium uptake and antioxidant responses of coastal plants differing in their ecological status under increasing salinity

In the present study, we compared the response to salinity of three plants from Brittany coast with contrasted ecological status: Limonium latifolium (salt marshes), Matricaria maritima (beach tops and sand dunes) and Crambe maritima (fixed dunes). Under controlled glasshouse conditions, the growth of the three plants decreased with increasing external salinity. L. latifolium and C. maritima exhibited the highest and lowest resistance to severe salt stress (400 mM), respectively. M. maritima could be considered as an intermediate species, since it tolerated salinity up to 200 mM. The same observation could be made with sodium absorption and acuumulation in plant tissues, the most tolerant species (L. latifolium being the least Na accumulator. Hydrogen peroxide (H₂O₂) and malondialdehyde (MDA), commonly produced in conditions of stress, accumulated significantly in salt treated C. maritima and M. maritima while not in the tolerant L. latifolium. The latter used glutathione reductase to maintain constant H₂O₂ levels under salt stress while peroxidases were very low and ascorbate peroxidase did not respond to salinity stimulation. The medium tolerant halophyte M. maritima used peroxidases to protect from NaCl-induced H₂O₂, while the sensitive C. maritima failed to detoxify H₂O₂ despite a sharp increase in catalase activity. Results showed that the three coastal species differ in resistance to salinity. They also suggested that the level of plant resistance to salinity could be attributed to differing mechanisms to manage the accumulation of sodium and cope with the oxidative damages.     

Effects of salinity on growth,membrane permeability and root hydraulic conductivity in three saltbush species

The species of the genus Atriplex have been introduced in West Asia and North Africa to determine their adaptability for use as fodder species. These halophytes are well adapted to extreme environmental conditions and may possess interesting properties for soil rehabilitation. The effect of NaCl stress on growth, water relation and mineral nutrition were investigated in three xero-halophyte species of Atriplex used for rehabilitation of arid steppe in Algeria. Atriplex halimus, Atriplex canescens and Atriplex nummularia, were cultivated in hydroponic conditions and treated with increasing doses of NaCl (0–300 mM). All species showed positive plant growth for low and moderate levels of salinity. A. halimus had higher dry weight production than A. nummularia and A. canescens in high salinity concentration. Increasing concentration of salinity induced decrease in chlorophyll content (Chl a and b) and root hydraulic conductivity (L₀) in all species, especially in A. canescens. All three species showed marked increase in electrolyte leakage across the salinity gradient. In addition all species were able to accumulate a large quantity of sodium (Na), chloride (Cl) and proline and to maintain higher relative water content, which was probably associated with a greater capacity for osmotic adjustment, whereas potassium (K) and calcium (Ca) decreased with increase salinity. The data suggest that salt tolerance strategies in all Atriplex species could involve a delicate balance among ion accumulation, osmotic adjustment, production of osmotica and maintenance of relative water content and growth.     

Effects of short- and long-term salinity on leaf water relations, gas exchange, and growth in Ipomoea pes-caprae

Ipomoea pes-caprae is widespread in pantropical coastal areas along the beach. The aim of this study was to investigate the salinity tolerance level and physiological mechanisms that allow I. pes-caprae to endure abrupt increases in salinity under brief or prolonged exposure to salinity variations. Xylem sap osmolality (X_osm), leaf water relations, gas exchange, and number of produced and dead leaves were measured at short- (1-7 d) and long- (22-46 d) term after a sudden increase in soil salinity of 0, 85, 170, and 255 mM NaCl. In the short-term, X_osm was not affected by salinity, but in the long-term there was a significant increase in plants grown in presence of salt compared with control plants. After salt addition, the plants showed osmotic stress with temporal cell turgor loss. However, the water potential gradient for water uptake was re-established at 4, 7 and 22 d after salt addition, at 85, 170 and 255 mM NaCl, respectively. In the short-term I. pes-caprae was able to tolerate salinities of up to 255 mM NaCl without significant reduction in carbon assimilation or growth. With the duration of stress, leaf ion concentration continued to increase and reached toxic levels at high salinity with a progressive decrease in photosynthetic rate, reduced leaf formation and accelerated senescence. Then, if high levels of soil salts from tidal inundation occur for short periods, the survival of I. pes-caprae is possible, but prolonged exposure to salinity may induce metabolic damage and reduce drastically the plant growth.     

Implication of nutrient and salinity interaction on the productivity of <Emphasis Type="Italic">Spartina patens</Emphasis>

Reintroduction of fresh water to coastal systems with altered hydrologic regimes is a management option for restoring degraded wetland habitats. Plant production in these systems is believed to be enhanced by increased nutrient availability and reduced salinity. Although studies have documented nutrient limitation and salinity stress in coastal marshes, interpreting the effects of freshwater reintroduction on plant production is difficult because high nutrient availability often is confounded with low salinity. We tested the hypothesis that plant growth response to nutrients does not vary with salinity in a greenhouse study. Treatments consisted of four nutrient concentrations and four non-lethal salinity levels; plant response was measured as biomass accumulation after 144 days of exposure. The significant interaction between salinity and nutrient concentrations indicates that response of Spartina patens marshes to freshwater inflows would vary by site-specific soil conditions. Biomass decreased with increased salinity at all four nutrient concentrations with variation among the nutrient concentrations decreasing as salinity increased. We demonstrate the importance of considering ambient salinity and nutrient soil conditions in restoration planning involving freshwater inflow. We propose salinity should remain a primary concern in restoration plans targeted at improving degraded S. patens-dominated marsh habitat.     

Green beach vegetation dynamics explained by embryo dune development

Sandy coastlines are dynamic environments with potential for biodiverse habitats, such as green beaches. Green beach vegetation can develop on nutrient-poor beaches landward from embryo dunes. It is characterised by low-dynamic coastal wetland habitat such as salt marshes and dune slacks. It has been hypothesised that the establishment of green beach vegetation is facilitated by the shelter provided by embryo dunes, however evidence is lacking.We explored the importance of geomorphology and soil conditions on the species richness and turnover of green beach vegetation over a time period of 10 years. We recorded 107 plots along 11 transects over a gradient from beach to dune on the island of Schiermonnikoog, the Netherlands. We characterised transect geomorphology at transect level and soil conditions and vegetation at plot level in 2006 and 2016.We found that the green beach vegetation was highly dynamic, total plant cover increased by 62% within 10 years. In 2006 beach width was an important factor in explaining species richness, with the highest number of species occurring on narrow beaches with a large volume of embryo dunes. In 2016, species richness was positively associated with the build-up of organic matter. Overall species richness declined relative to 2006 and was accompanied by an increase in elevation due to sand burial and the expansion of embryo dune volume.Our data suggests that geomorphology influenced the vegetation indirectly by affecting sand burial rate. Plant species richness declined less at sheltered conditions where sand burial was limited, allowing the build-up of organic matter. This indicates a time-dependent relationship between the development of embryo dunes and plant species richness: embryo dunes can be a source of shelter, thus increasing species richness, but can compete for space over time, lowering species richness again. Our results are relevant for engineering and management of biodiverse sandy shores.