Invasive alien plants in marine protected areas: the <Emphasis Type="Italic">Spartina anglica</Emphasis> affair in the European Wadden Sea

The common cord-grass Spartina anglica, a fertile hybrid of S. maritima and S. alterniflora, was planted in the European Wadden Sea extensively during the late 1920s and 1930s to promote sediment accretion. After establishment, it colonised as a pioneer plant in the upper tidal zone, where it occurs frequently in coherent swards at the seaward front of saltmarshes and in patches on the tidal flats. Often, a conspicuous, almost monotypic, belt of S. anglica is formed. Over the last two decades, an increase in abundance and accelerated spread of S. anglica was observed, possibly promoted by warmer spring temperatures. This alien species may benefit from global warming, and there is considerable concern about its harmful impacts on the native biocoenoses and native biodiversity of the unique Wadden Sea ecosystem, encompassing effects on hydromorphodynamics and coastal protection. For a definitive assessment, however, an adequate quantification and comparison of documented and potential effects of S. anglica is important, but currently unavailable. Consequently, no management strategy exists for the prevention or restoration of the Wadden Sea ecosystem. Thus, the development of an alien species plan on the level of the Trilateral Cooperation on the Protection of the Wadden Sea is essential.     

Density-dependent effects on the dieback of exotic species Spartina anglica in coastal China

The exotic species Spartina anglica C.E. Hubbard has been experiencing dieback in coastal China during the past decade, while it is invading other places in the world. Thus, an explanation for the dieback of the populations in coastal China would be helpful for controlling invasions of S. anglica in other places of the world. Both field and greenhouse experiments were conducted to determine the possible role of density in the dieback of the populations in China. The density effects were tested by maintaining six density levels in field experiments and three density levels in greenhouse experiments over 7 months. The results showed that growth, asexual reproduction and biomass accumulation in medium density were significantly higher than those in other density treatments for field experiments. The growth and asexual reproduction in high-density patches were lowest among the treatments in the field. Data showed the same pattern for the greenhouse experiments. The results implied that density regulation had a great effect on most of the studied parameters. Because the species is surviving at high density along the Chinese coast, and has a much lower growth rate due to strong intraspecific competition, and also because high-density populations may decline into low density populations due to resource exhaustion, high plant density might be one of the important factors in the dieback of S. anglica.     

Remedial habitat creation: does Nereis diversicolor play a confounding role in the colonisation and establishment of the pioneering saltmarsh plant, Spartina anglica?

Increasing concerns over global warming and expected sea level rises have led to the adoption of new coastal management strategies around the south-east coast of England. This paper explores the role played by the estuarine invertebrate Nereis diversicolor in limiting the colonisation and establishment of the invasive pioneering salt marsh plant, Spartina anglica. The biology of N. diversicolor is briefly reviewed and data from field experiments are presented demonstrating significant negative effects of worm abundance on transplanted S. anglica biomass. Laboratory-based experiments demonstrated significant negative effects of N. diversicolor abundance on the survival of S. anglica seeds transplanted to sediment cores. The importance of estuarine invertebrates in engineering the mudflat habitat may confound the foreseen ecosystem services and function provided by saltmarsh management schemes. Received: 15 February 1999 / Received in revised form: 4 June 1999 / Accepted: 11 June 1999     

Limited seed retention during winter inhibits vegetation establishment in spring,affecting lateral marsh expansion capacity

Coastal systems worldwide deliver vital ecosystem services, such as biodiversity, carbon sequestration, and coastal protection. Effectivity of these ecosystem services increases when vegetation is present. Understanding the mechanisms behind vegetation establishment in bio‐geomorphic systems is necessary to understand their ability to recover after erosive events and potential adaptations to climate change. In this study, we examined how seed availability affects vegetation establishment in the salt marsh–intertidal flat transition zone: the area with capacity for lateral marsh expansion. This requires vegetation establishment; therefore, seed availability is essential. In a 6‐month field experiment, we simulated a before and after winter seed dispersal at two locations, the salt‐marsh vegetation edge and the intertidal flat, and studied seed retention, the seed bank, and the seed viability of three pioneer marsh species: Salicornia procumbens, Aster tripolium, and Spartina anglica. During winter storm conditions, all supplied seeds eroded away with the sediment surface layer. After winter, supplied seeds from all three species were retained, mostly at the surface while 9% was bioturbated downwards. In the natural seed bank, A. tripolium and S. anglica were practically absent while S. procumbens occurred more frequently. The viability of S. procumbens seeds was highest at the surface, between 80% and 90%. The viability quickly decreased with depth, although viable S. procumbens seeds occurred up to 15 cm depth. Only when seeds were supplied after winter, many S. procumbens and some S. anglica individuals did establish successfully in the transition zone. Viable seed availability formed a vegetation establishment threshold, even with a local seed source. Our results suggest that, although boundary conditions such as elevation, inundation, and weather conditions were appropriate for vegetation establishment in spring, the soil surface in winter can be so dynamic that it limits lateral marsh expansion. These insights can be used for designing effective nature‐based coastal protection.     

Eco-physiological responses of the declining population Spartina anglica to N and P fertilizer addition

Nitrogen and phosphorus are both important life elements. N, P and combined N-P fertilizers were added to the declining population Spartina anglica Hubbard in coastal China. Some growth parameters and eco-physiological responses of S. anglica to different fertilizer treatments (N, P and combined N-P fertilizer addition with high, medium and low levels, respectively) were measured. The fertilizer addition had a highly significant effect on the dynamics of its height-growth, number of leaves, number of roots and total biomass. Only N addition had a significant effect on leaf area and leaf thickness in all fertilizer treatments. On the dynamics of its height-growth, the effect of N addition was the most apparent, and the effect of N-P addition was not greater than those of N and P addition separately. The photosynthesis rate was enhanced and the yield was the highest with the highest N, the highest N-P and the medium P addition. The rates were higher than those of CK by 19.08 μmol·m^?2·s^?1, 15.47 μmol·m^?2·s^?1 and 11.23 μmol·m^?2·s^?1, respectively. The activity of SOD and POD increased with the treatments after freshwater stress for 14 days. Effects of medium N and P addition were significant for SOD activity. However, POD activity was significantly higher with the treatment of higher N and higher N-P addition. In a word, fertilizer addition improved the growth of the declining population S. anglica. The results indicated that the decline of S. anglica was correlated with the nutriment deficiency in soil, especially with the lack of N.     

Effects of sea level rise on mangrove Avicennia population growth,colonization and establishment: Evidence from a field survey and greenhouse manipulation experiment

Mangroves will either face longer tidal inundation or retreat landwards in response to on-going accelerated sea level rise. However, little is known about the growth, regeneration or colonization of mangrove seedlings under the different tidal inundation regimes associated with accelerated sea level rise. In the present study, a field survey and a greenhouse mesocosm experiment were conducted to evaluate possible effects of accelerated sea level rise on colonization, establishment and seedling growth of a mangrove pioneer species, Avicennia marina. Avicennia populations at different elevations of the intertidal zone on Xiamen Island in Fujian Province, China showed distinctive patterns in both seedling density and plant age. The seedlings at lower elevations had less annual biomass accumulation and population productivity, but higher shoot to root ratios, suggesting that elevation has positive effects on seedling growth. The greenhouse mesocosm experiment with 1-year-old A. marina seedlings utilized five inundation periods (0, 2, 4, 6 and 12 h in a semidiurnal tidal cycle) and two inundation depths (root immersion and canopy immersion). Both inundation period and depth exerted significant and negative effects on biomass accumulation, photosynthetic rate, leaf electron transportation and water use efficiency. However, the negative effects of canopy immersion were more profound than root immersion. Canopy immersion exacerbated the effects of prolonged inundation, with no seedlings surviving under canopy immersion at the 12-hour treatment. These results suggest that at lower elevations with higher sea level, canopy immersion will have greater negative effects on seedling colonization, establishment and early growth of A. marina. This finding is instrumental in predicting the future dynamics of mangrove forests under increasing sea levels.     

Morphological and biomechanical responses of floodplain willows to tidal flooding and salinity

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Photosynthetic and physiological responses of Kandelia candel L. Druce seedlings to duration of tidal immersion in artificial seawater

The present study demonstrates the influence of the duration of periodical waterlogging with artificial seawater on the photosynthetic and physiological responses of Kandelia candel L. Druce seedlings, the pre-dominant species of subtropical mangroves in China. Artificial tidal fluctuations applied here closely mimicked the twice daily tidal inundation which mangroves experience in the field. All the seedlings were immersed in artificial seawater during 70-day cultivation. Similar trends with increasing duration of immersion occurred in photosynthetic rate, transpiration rate, stomatal conductance and intercellular CO₂ concentration, where significant decreases occurred only in long time treatments of 10 or 12 h. Water used efficiency and chlorophyll contents showed lower in medium periods and higher in long periods of immersion. This indicates that the increase in pigment contents of leaves was ineffective in promoting P_n under long time immersion. Light saturation points under short time waterlogging (0–4 h) occurred at light intensities of 800–1000 μmol/m²/s, and at around 400 μmol/m²/s in long time treatments (8–12 h). Long periods of tidal immersion therefore significantly inhibited photosynthesis of mature leaves. Alcohol dehydrogenase and oxidase activity in roots both increased under longer immersion periods, suggesting that roots are sensitive to anaerobiosis under long term waterlogging. The activities of peroxidase and superoxide dismutase in mature leaves increased in 8 h and 10 h treatments, respectively. The content of malondialdehyde in mature leaves increased under long time treatments. Abscisic acid accumulation in mature leaves also had a sharp increase from 8 h to 12 h inundation. Even though the anti-oxidative enzymes were induced by waterlogging, this was not sufficient to protect the seedlings from senescence. The results suggested that K. candel seedlings completely tolerated tidal immersion by seawater up to about 8 h in each cycle, which matches the natural distribution of K. candel in inter-tidal zones of China.     

Cultivation of the brown alga Sargassum horneri: sexual reproduction and seedling production in tank culture under reduced solar irradiance in ambient temperature

As a large conspicuous intertidal brown alga, individuals of Sargassum horneri can reach a length of more than 7 m with a fresh weight of 3 kg along the coasts of the Eastern China Sea. The biomass of this alga as a vital component in coastal water ecology has been well documented. In recent years, a steady disappearance of the algal biomass along the once densely populated coastal areas of the Eastern China Sea has drawn attention in China. Efforts have been made to reconstruct the subtidal algal flora or even to grow the alga by use of long-lines. As part of the efforts to establish an efficient technique for producing seedlings of S. horneri, in this investigation a series of culture experiments were carried out in indoor raceway and rectangular tanks under reduced solar irradiance at ambient temperature in 2007–2008. The investigation demonstrated that: (1) sexual reproduction of S. horneri could be accelerated in elevated temperature and light climates, at least 3 months earlier than in the wild; (2) eggs of S. horneri had the potential to be fertilized up to 48 h, much longer than that of known related species; (3) suspension and fixed culture methods were both effective in growing the seedlings to the long-line cultivation stage; and (4) the life cycle of S. horneri in culture could be shortened to 4.5 months, thus establishing this alga as an appropriate model for investigating sexual reproduction in dieocious species of this genus.     

Nitrogen level changes the interactions between a native (Scirpus triqueter) and an exotic species (Spartina anglica) in Coastal China

The exotic species Spartina anglica, introduced from Europe in 1963, has been experiencing a decline in the past decade in coastal China, but the reasons for the decline are still not clear. It is hypothesized that competition with the native species Scirpus triqueter may have played an important role in the decline due to niche overlap in the field. We measured biomass, leaf number and area, asexual reproduction and relative neighborhood effect (RNE) of the two species in both monoculture and mixture under three nitrogen levels (control, low and high). S. anglica showed significantly lower biomass accumulation, leaf number and asexual reproduction in mixture than in monoculture. The inter- and intra-specific RNE of S. anglica were all positive, and the inter-specific RNE was significantly higher than the intra-specific RNE in the control. For S. triqueter, inter- and intra-specific RNE were negative at the high nitrogen level but positive in the control and at the low nitrogen level. This indicates that S. triqueter exerted an asymmetric competitive advantage over S. anglica in the control and low nitrogen conditions; however, S. anglica facilitated growth of S. triqueter in high nitrogen conditions. Nitrogen level changed the interactions between the two species because S. triqueter better tolerated low nitrogen. Since S. anglica is increasingly confined to upper, more nitrogen-limited marsh areas in coastal China, increased competition from S. triqueter may help explain its decline.