Moderate predation on meiofauna by the macrobenthos of the Wadden Sea

The role of meiofauna in the trophic web of marine ecosystems is a controversial topic. During an experimental study on tidal flats in the eastern part of the North Sea, potential predators on meiofauna were kept in enclosures. A gobiid fish, shrimp, crabs, an amphipod, four polychaete worms and a nemertine were tested for their ability to decimate nematodes, turbellarians, ostracods and benthic copepods. Only few macrobenthic species preyed heavily on permanent meiofauna: juveniles of the shore crabCarcinus maenas, the hermit crabPagurus bernhardus with the colonial hydrozoanHydractinia echinata on its shells, and the rag wormNereis diversicolor. When benthic infauna was protected from these predators with exclosures, juvenile macrofauna responded with a marked increase in number, while the permanent meiofauna remained unaltered except for a limited increase in nematodes. It is concluded that in the Wadden Sea abundance of permanent meiofauna (Nematoda, Turbellaria, Ostracoda, Copepoda) is only locally or temporarily regulated by macrobenthic predators.I acknowledge a grant from the Deutsche Forschungsgemeinschaft (DFG).     

Growth and photosynthetic responses to zinc stress of an invasive cordgrass, Spartina densiflora

Spartina densiflora Brongn. is found in coastal marshes of southwest Spain, growing over sediments containing 100-4800 ppm Zn. A glasshouse experiment was designed to investigate the effect of Zn from 0 to 100 mmol.l(-1) on the growth and photosynthetic apparatus of S. densiflora, by measuring relative growth rate, leaf elongation rate, number of tillers, height of tillers, chlorophyll fluorescence parameters, gas exchange and photosynthetic pigment concentrations. We also determined total ash, Zn, calcium, magnesium and phosphorus concentrations, and the C/N ratio. At 100 mmol.l(-1) Zn, S. densiflora showed a 48% biomass reduction after 1 month of treatment. Long-term effects of Zn on growth of S. densiflora consisted mainly of variations in net photosynthesis. Modification of the Zn/Mg ratio was linked to a strong decrease in RuBP carboxylase (Zn was favoured in local competition with Mg, so that the affinity of RuBisCO for CO(2) decreased), oxygenase activity of RuBisCO acting as a substitute for the photosynthetic function. Also, Zn had a marked overall effect on the photochemical (PSII) apparatus and the synthesis of photosynthetic pigments. However, the results indicate that S. densiflora is capable of tolerating very high and continued exposure to Zn, as this species lowers the translocation of Zn from the nutrient solution to roots and controls Zn ion transport into leaves. Therefore, S. densiflora could be useful in the phytostabilization of soils.     

Monitoring harbour seal (Phoca vitulina) in the Danish Wadden Sea

Monitoring of harbour seal (Phoca vitulina) has been carried out in the Danis part of the Wadden Sea during the past 8 years by the Fishery and Maritime Museum. In addition to this, field observations over three breeding seasons have been undertaken. The census shows a steady increase of approx. 14% per yeart. The distribution of the seals in the area can be split up into two different types of haulout localities, and a varied pattern of haulouts is observed during the year. The mass die-off in 1988 is discussed within the framework of the knowledge on the population, and it is recommended that more effort be put into monitoring and population studies of the harbour seal in the entire International Wadden Sea. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, 1–4 November 1988)     

Geographical variation in germination traits of the salt-marsh cordgrass Spartina alterniflora in its invasive and native ranges

入侵地和原产地盐沼植物互花米草种子萌发性状的地理变异 种子萌发是植物早期生活史中最重要的阶段,决定了植物的生态位和地理分布范围,对外来植物的入侵潜力有重要影响。盐沼植物互花米草(Spartina alterniflora)在中国沿海滩涂的入侵范围最大,并已入侵到比原产地更低的纬度范围,这为我们研究互花米草在不同地理区域之间以及沿纬度梯度的萌发性状差异和适应提供了契机。在控温培养箱中淡水培养条件下,我们比较研究了来自入侵地(19°–40° N)10 个地点和原产地(27°–43° N)16个地点不同纬度互花米草种群的种子萌发性状差异,以及这种差异与各种 群来源地潮差和气候因素的相关性。原产地互花米草种群种子的萌发率和萌发指数比入侵地种群分别高10%和20%,但入侵地互花米草种群的萌发速度比原产地快3 d。入侵地互花米草种群的萌发率和萌发 指数随着纬度升高呈现线性递增的变化趋势,而原产地呈现线性递减的变化趋势。入侵地和原产地互花米草种群的平均萌发时间都与纬度呈现线性负相关。入侵地互花米草种群的萌发率和萌发指数与年日均温、年日最低均温、和年日最高均温呈现负相关,而在原产地呈现相反的相关关系。入侵地和原产地互花米草种群的平均萌发时间分别与年日均温、年日最低均温和年日最高均温呈现正相关关系。我们的研究结果表明,入侵地和原产地互花米草种群的萌发率和萌发指数已沿纬度进化出不同的渐变群格局,但平均萌发时间进化出与原产地一致的纬度渐变群格局,即在生物入侵过程中沿纬度梯度种子萌发策略会随着入侵时间和过程而发生变化。     

The relationship between silicon availability,and growth and silicon concentration of the salt marsh halophyte Spartina anglica

Incorporation of cover crops into cropping systems may contribute to a more efficient utilization of soil and fertilizer P by less P-efficient crops through exudation of P-mobilizing compounds by the roots of P-efficient plant species. The main objective of the present work was to test this hypothesis. First a method has been developed which allows the quantification of organic anion exudation from individual cluster roots formed by P-deficient white lupin (Lupinus albus L.). Lupin plants were grown in nutrient solution at 1 μM P and in a low P loess in small rhizotrons. Organic anions exuded from intact plants grown in nutrient solution were collected from individual cluster roots and root tips sealed in small compartments by an anion-exchange resin placed in nylon bags (resin-bags). Succinate was the dominant organic anion exuded followed by citrate and malate. The mean of citrate exudation-rate was 0.06 pmol mm⁻¹ s⁻¹ with exudation highly dependent on the citrate concentration and on the age of the cluster roots. Exudates from cluster roots and root tips grown at the soil surface (rhizotron-grown plants) were collected using overlayered resin–agar (resin mixed with agar). Citrate exudation from cluster roots was 10 times higher than that from root tips. Fractionation of P in the cluster root rhizosphere-soil indicates that white lupin can mobilize P not only from the available and acid-soluble P, but also from the stable residual soil P fractions. In pot experiments with an acid luvisol derived from loess low in available P, growth of wheat was significantly improved when mixed-cropped with white lupin due to improved P uptake. Both in mixed culture and in rotation wheat could benefit from the P mobilization capacity of white lupin, supporting the hypothesis above. Nine tropical leguminous cover crops and maize were grown in a pot experiment using a luvisol from Northern Nigeria low in available P. All plant species derived most of their P from the resin and bicarbonate-extractable inorganic P. Organic P (P_o) accumulated particularly in the rhizosphere of all plant species. There was a significant negative correlation between the species-specific rhizosphere acid phosphatase activity and P_o accumulation. Growth and P uptake of maize grown in rotation after legumes were enhanced indicating that improved P nutrition was a contributing factor. This revised version was published online in June 2006 with corrections to the Cover Date.     

Human impacts on fisheries resources and abundance in the Danish Wadden Sea, c1520 to the present

For the past 30 years, the Danish Wadden Sea area has been protected by fishery and hunting regulations, pollution controls and settlement restrictions. Nevertheless, todays Wadden Sea is a depleted ecosystem when we compare it with past abundance and diversity of marine animal populations. This review indicates that the abundance of finfishes has undergone a long-term decline since the seventeenth century. The review also indicates a trajectory of ecological decline since the early twentieth century that seems to be related to: (1) fishing extractions which focused on undersized fish throughout the first half of the twentieth century; (2) habitat destruction, which was most marked in the first half of the twentieth century; and (3) pollution, which was worst in the third quarter of the twentieth century. Historical investigation reveals that we need to fundamentally revise present-day baselines about the potential species richness and abundance of the Wadden Sea.     

Molecular analysis of diazotroph diversity in the rhizosphere of the smooth cordgrass, Spartina alterniflora

N(2) fixation by diazotrophic bacteria associated with the roots of the smooth cordgrass, Spartina alterniflora, is an important source of new nitrogen in many salt marsh ecosystems. However, the diversity and phylogenetic affiliations of these rhizosphere diazotrophs are unknown. Denaturing gradient gel electrophoresis (DGGE) of PCR-amplified nifH sequence segments was used in previous studies to examine the stability and dynamics of the Spartina rhizosphere diazotroph assemblages in the North Inlet salt marsh, near Georgetown, S.C. In this study, plugs were taken from gel bands from representative DGGE gels, the nifH amplimers were recovered and cloned, and their sequences were determined. A total of 59 sequences were recovered, and the amino acid sequences predicted from them were aligned with sequences from known and unknown diazotrophs in order to determine the types of organisms present in the Spartina rhizosphere. We recovered numerous sequences from diazotrophs in the gamma subdivision of the division Proteobacteria (gamma-Proteobacteria) and from various anaerobic diazotrophs. Diazotrophs in the alpha-Proteobacteria were poorly represented. None of the Spartina rhizosphere DGGE band sequences were identical to any known or previously recovered environmental nifH sequences. The Spartina rhizosphere diazotroph assemblage is very diverse and apparently consists mainly of unknown organisms.     

Hybridization between introduced smooth cordgrass (Spartina alterniflora; Poaceae) and native California cordgrass (S. foliosa) in San Francisco Bay, California, USA

Introduced Spartina alterniflora (smooth cordgrass) is rapidly invading intertidal mudflats in San Francisco Bay, California. At several sites, S. alterniflora co-occurs with native S. foliosa (California cordgrass), a species endemic to California salt marshes. In this study, random amplified polymorphic DNA markers (RAPDs) specific to each Spartina species were identified and used to test for hybridization between the native and introduced Spartina species in the greenhouse and in the field. Greenhouse crosses were made using S. alterniflora as the pollen donor and S. foliosa as the maternal plant, and these crosses produced viable seeds. The hybrid status of the crossed offspring was confirmed with the RAPD markers. Hybrids had low self-fertility but high fertility when back-crossed with S. foliosa pollen. Hybrids were also found established at two field sites in San Francisco Bay; these hybrids appeared vigorous and morphologically intermediate between the parental species. Field observations suggested that hybrids were recruiting more rapidly than the native S. foliosa. Previous work identified competition from introduced S. alterniflora as a threat to native S. foliosa. In this study, we identify introgression and the spread of hybrids as an additional, perhaps even more serious threat to conservation of S. foliosa in San Francisco Bay.     

Climatic change and the Wadden Sea,The Netherlands

The predicted increase in atmospheric carbon dioxide and the effects of global warming will influence the Wadden Sea, The Netherlands, an area of exceptional ecological value. The direct effect of elevated atmospheric CO₂ on terrestrial coastal ecosystems is either marginal or unknown. The slight acidification of the sea which is predicted might have an impact on primary producers and juvenile animals. The effect of CO₂ fertilization on marine primary production remains to be elucidated. Profound changes will occur if sea level rises at the predicted rate of 60 cm per century, as sedimentation rates will be insufficient to maintain the salt marshes on the barrier islands. The marshes of the mainland coast will be impoverished, as high and low marshes are not expected to continue to coexist at the same locations. As sediment supply to the Wadden Sea is sufficient to compensate for sea level rise, the estuarine character of the Wadden Sea, with sand- and mudflats, is expected to remain largely unchanged.     

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.     

The macrobenthos in Spartina alterniflora salt marshes of the Wanggang tidal-flat,Jiangsu coast,China

On the Jiangsu coast, eastern China, Spartina alterniflora, which was introduced artificially into the region, is becoming a dominant plant species in the inter-tidal salt marshes. In order to evaluate the environmental and ecological impact of the colonization of S. alterniflora, we carried out investigations into the benthic macrofauna of the Spartina marshes of the Wanggang area, central Jiangsu coast, in 2006 and 2007. Based on analysis of the data sets obtained, 12 species of macrobenthos have been identified for the Wanggang salt marsh, including S. alterniflora, Cerithidea cingulata, Littorna scabra, Bithynia fuchsiana, Macrophthalmus japonicus, Uca arcuata, Nereis sp., Boleophthalmus petinirostris, Cyclina sinensis, Bullacta exarata, Angustassiminea castanea and Glaucomya chinensis. The results indicate that some of the native species have adapted to the new ecological environment associated with the cordgrass S. alterniflora. The biomass of macrobenthos varies significantly over different parts of the salt marsh. Further, there is a seasonal change in bio-density, with the density in summer (July and August) > autumn (November) > early summer (May). At the landward edge and over the central part of the S. alterniflora marsh, the bio-diversity is higher than the other areas of the marsh.     

On the dynamics of the stocks of blue mussels (Mytilus edulis L.) in the Danish Wadden Sea

As biological basis for the monitoring programme for the commercially exploited stock(s) of mussels (Mytilus edulis L.) in the Danish Wadden Sea, samples of mussels have been collected regularly since 1986, both from sub-tidal and inter-tidal mussel beds. These samples are the basis for the estimation of total biomass. They also provide data on size frequency distributions, which have been analysed for cohort identification resulting in length at age data, which again have been used for estimating parameters (L and K) for the von Bertalanffy growth equation (VBGE) as well as mortality parameters. By applying these in the Beverton & Holt model, estimates of average biomass and annual production (P) of the mussels have been obtained together with possible fisheries yields from the beds. The growth and mortality parameters and the figures for annual production and P/B are compared with figures from other investigations. These analyses have been the basis for annual assessments of the mussel stocks, which again are used in the current management of mussel fishery in the Danish Wadden Sea.     

Predictive ecosystem research in the Wadden Sea

Predictive ecosystem research needs a pluralistic approach. Retrospective studies reveal the initial causes of ongoing ecological change. In the Wadden Sea, inherent ecosystem stability may be falsely assumed, because the effects of modern coastal architecture and of anthropogenic eutrophication to some extent complement each other. Expected environmental changes often have corresponding phases in the past which may serve as a model to predict ecological implications. Historically, quantitative ecology entered the Wadden Sea, via fisheries research, from the oceanic side. Quantified material fluxes may reveal imbalances which are indicative of the rough direction of ecosystem change. For ecosystem research to contribute to the maintenance of the Wadden Sea as a centre of coastal organisms, quantitative knowledge of resources and ecosystem metabolism must be supplemented by qualitative knowledge of habitat requirements and species interdependences. Qualitative ecology entered the Wadden Sea from the landward side. Extending this approach to anticipatory field experiments may help to predict ecological changes at the species level.     

Inbreeding depression in smooth cordgrass (Spartina alterniflora, Poaceae) invading San Francisco Bay

The magnitude of inbreeding depression in invading plant populations is often presumed to be small and of little consequence. The purpose of this study was to assess the magnitude of inbreeding depression in a pollen-limited, partially self-incompatible, invading plant population. The magnitude and timing of inbreeding depression were compared among ten maternal plants sampled from a population of smooth cordgrass (Spartina alterniflora) invading San Francisco Bay. Selfed and outcrossed progeny were compared for embryo abortion, survival of seedlings, and growth/survival at the end of the first growing season in three greenhouse environments. Estimates of inbreeding depression varied among environments, with competitive environment > high-nutrient environment > low-nutrient environment. Population-level estimates of inbreeding depression ranged from 0.61 to 0.81; however, maternal plants varied significantly in their magnitude of inbreeding depression, ranging from 0.1 to 0.97. The 95% confidence interval for inbreeding depression for some maternal plants included zero. There was a significant negative correlation between the overall magnitude of inbreeding depression and self-fertility rate among maternal plants. The few maternal plants with high self-fertility carried relatively little genetic load, and their selfed progeny are likely to survive on open mudflats. The noncompetitive, pollen-limited growing conditions associated with invasion may allow self-fertility to spread in this population.     

Effects of salinity and temperature stress on Ecophysiological characteristics of exotic cordgrass, Spartina alterniflora

Salintiy and temperature are two important ecological factors which affect the distribution and abundance of Spartina alerniflora Loisel. To find out how cordgrass adapts to the environmental conditions in the introduced range, we studied the dynamics of a series of important physiological components including superoxide dismutase (SOD), guaiacol peroxidase (POD), catalase (CAT), malondialdehyde (MDA), soluble sugar and free proline under different salinity and temperature stresses. The results showed that low NaCl concentration (lower than 100 mmol·L^?1) enhanced the growth of S. alterniflora. On the contrary, high NaCl concentration (higher than 100 mmol·L^?1) inhibited the growth of S. alterniflora. To a certain extent, S. alterniflora was able to be acclimated to the osmotic pressure created by external solution concentration by adjusting the activities of POD, SOD and CAT, and the contents of free proline and soluble sugar. S. alterniflora varied in its responses to environment in different parts of the plant under 5°C and 38°C temperature stress. Compared with roots, leaves accumulated more soluble sugar, and CAT activities in leaves were higher, whereas SOD and POD activities in leaves were much lower than those in roots.     

NaCl胁迫对大米草光合·蒸腾等生理特性的影响

研究了NaCl胁迫下大米草净光合速率（Pn）、蒸腾速率（Tr）,叶片气孔导度（Gs）、细胞间CO2浓度（Ci）、株高、叶长、叶宽、茎粗、叶绿素含量、气孔限制值和水分利用效率。结果表明：当NaCl浓度高于300mmol／L时,大米草Pn、Tr、Gs、株高、叶长以及叶绿素含量受到显著抑制（P〈0．05）,叶宽及茎粗则无显著性差异。NaCl胁迫下,大米草光合速率的降低是气孔因素和非气孔因素综合导致的结果,Pn、Tr、Gs、株高以及叶绿素含量的降低可作为大米草受NaCl胁迫的症状．而WUE则保持在较高的水平．因此在防治大米草荨延时．排水处理不是最佳选择。     

Interactive effects between physical forces and ecosystem engineers on seed burial: a case study using Spartina anglica

Seed burial (i.e. vertical seed dispersal) has become increasingly valued for its relevance for seed fate and plant recruitment. While ecosystem engineers have been generally considered as the most important drivers of seed burial, the role of physical forces, such as wind or water flow, has been largely overlooked. Using tidal habitats as a model system, and a combination of flume and mesocosm experiments, we investigated the effects of 1) currents, 2) benthic animals with different engineering activities and 3) their interplay on seed burial of a common salt marsh pioneer plant, Spartina anglica. Our results reveal that in such systems, water flow can be of equal or higher importance than ecosystem engineers for seed burial. For passive seed‐burying engineers (PSE), coupling their actions with currents produced synergistic seed burial effects, whereas the interactive effects were only additive for active seed‐burying engineers (ASE). This paper extends current understanding of seed burial and seed bank formation by revealing the need to incorporate physical forces into seed burial mechanisms. We provide the first empirical evidence that physical forces influence seed burial by synergistically interacting with ecosystem engineers, thus highlighting the role of biophysical interactions as important drivers for vertical seed movement.