Effects of storm frequency on dune vegetation

In the Gulf of Mexico, barrier islands absorb the majority of wind and wave action from storms, which modifies their dune morphology and vegetation dynamics. Storm frequency is predicted to increase as a result of climate change, yet the effects of this change on coastal ecosystems remain poorly understood. Using estimates of plant growth in storm and nonstorm years from long‐term census data describing the dynamics of dune vegetation on St. George Island, FL, we built a first‐order model that predicts how dune communities will respond to a change in storm frequency. It predicts that an increasing frequency of storms will result in a change in the vegetation across the dunes. The fore‐ and interdune communities are predicted to become more similar to one another through the dominance of a small number of common storm‐resilient species. Alternatively, the backdune community is predicted to become more distinct through an increase in rare species that represent primary succession. Finally, the model predicts that many species will not respond to an increase in the number of storms per year in the same manner in which they respond to current storm frequency. This model is beneficial both for the development of more complex approaches to predicting effects of climate change and for informing preventative management techniques.     

古尔班通古特沙漠土壤水分与化学性质的空间分布

区域尺度上,气象和水文状况是影响植被分布格局和土壤性状的主要因子,当然,局部的地貌特征及植被分布对土壤性质的影响也是不能忽视的。以古尔班通古特沙漠为研究对象,对其南缘至腹地约25 km的土壤水分与化学性质的空间分布及其相互关系进行研究,结果显示:水平方向上,从沙漠南缘至腹地,土壤水分在丘底呈减小趋势,尤其是距边缘10 km以外,且在距边缘6—7 km出现1个峰值。同时土壤pH值和电导率的大小以及有机碳,全氮,有效氮,全磷和有效磷含量与土壤水分的变化状况相似,在丘底呈减小趋势,而在丘坡和丘顶上均呈波动分布。这说明丘底土壤性质的空间格局受到气象和水文状况的显著影响,而丘坡和丘顶土壤性质并没有受到气象和水文状况的显著影响,这种波动分布主要由采样点的设置及植物的分布状况所引起。丘底土壤水分及化学性质均显著高于丘坡和丘顶,这说明地貌特征对土壤性质的空间分布有显著影响。垂直方向上,不同深度土层土壤性质也存在显著差异,土壤水分,pH值和电导率随着土层深度增加而增加,而土壤养分随土层深度增加而减小。相关分析结果表明:仅4、5月份的土壤含水量及年平均土壤含水量与丘底土壤化学性质显著正相关,这可能由于冬季积雪的融化导致了土壤水分在4、5月份的聚集比较明显,使土壤水分对土壤化学性质发挥作用。总之,土壤空间异质性是由气象和水文以及地貌特征共同影响的,而土壤空间异质性的变异进而影响到植被的分布,植被的分布反过来又影响土壤的空间分布,因此土壤的空间分布和植被的空间分布是相互影响,共同作用的。     

A conceptual model of coastal dune ecology synthesizing spatial gradients of vegetation, soil, and geomorphology

Patterns of coastal dune vegetation are closely related with soil conditions controlled by geomorphic forms and processes. This study developed a conceptual model integrating these relationships in a spatially explicit manner. A rectangle of 180 × 280 m containing 126 grids of 20 × 20 m was established in the Sindu coastal dunefield in west Korean Peninsula. Sampling from each grid determined 11 soil properties and identified percent cover of 21 woody and herbaceous plant species. Digital elevation models were generated by topographic survey and used to derive eight topographic parameters. Redundancy analysis and canonical correspondence analysis examined the effect of geomorphic factors on edaphic characteristics and the edaphic influence on spatial distribution of vegetation, respectively. The spatial pattern of soil properties and plant species were inferred from spatial interpolation techniques. In the foredune area, distance from the coastline was a significant indicator of soil nutrients derived from the marine sources by aeolian processes. This favored the dominance by Elymus mollis. Moisture-tolerant species (e.g., Calamagrostis epigeios) had high cover in the acidic soils of dune slacks, which covaried with wetness index, an indirect measure of the depth to the freshwater table. Vegetation–soil interactions (e.g., nitrogen fixation by legumes) were important in secondary dune areas, with topographic effects less significant. Vegetation, soil, and geomorphic factors are closely connected in a causal chain across a whole dune area. Our model thus addresses the importance of integrating foredune, dune slack, and secondary dune into one continuous system.     

Structure, Pattern and Mechanisms of Formation of Seed Banks in Sand Dune Systems in Northeastern Inner Mongolia, China

A comparison of structure and pattern of the soil seed bank was made between active and stabilized sand dunes in northeastern Inner Mongolia, China. The objective of this paper was to determine the significance of seed bank in vegetation restoration of sand dunes. The results showed that (1) average seed density decreased from stabilized sand dune to interdune lowland of stabilized sand dune, to interdune lowland of active sand dune, and to active sand dune; (2) horizontally, along the transect from interdune lowland to ecotone and to sand dune top, a ‘V’ shaped pattern was presented in the active dune system, and a reverse ‘V’ shaped pattern in the stabilized sand dune system; (3) vertically, the proportion (accounting for the total seeds) of seeds found in 0–20 mm soil profile decreased from stabilized sand dune to interdune lowland of stabilized sand dune, to interdune lowland of active sand dune, and to active sand dune. The same order was also found in 20–50 mm and 50–100 mm soil profiles; (4) the Sokal and Sneath similarity indices in the species-composition between soil seed bank and above-ground vegetation were ranked as: the stabilized sand dune (24%) > the interdune lowland of active sand dune (21%) > the interdune lowland of stabilized sand dune (18%) > the active sand dune (5%); and (5) vegetation restoration of active sand dunes depends on the dispersal of seeds from nearby plant communities on the interdune lowlands. Much effort must be made to preserve the lowlands, as lowlands are the most important seed reservoir in the active sand dune field.     

Spatial connectivity and boundary patterns in coastal dune vegetation in the Circeo National Park,Central Italy

Abstract. Distribution patterns of coastal sand dune plant communities in the Circeo National Park (Central Italy) are quantified and compared by measuring spatial connectivity and richness of community boundaries along the dune profile. The purpose of this study is: (1) to evaluate patchiness and frequency of spatial links between communities; (2) to identify the communities most sensitive to disturbance; and (3) to predict probable changes due to modification of spatial zonation. Data were obtained using belt transects across the Holocene coastal dune zone. Vegetation zonation from the seashore to the foredune slacks is analysed in relation to chorological, phytosociological and life‐form types. We found that under relatively undisturbed environmental conditions communities formed a sequence (communities 1 to 7), with the exception of a replacement community, which occupied gaps in the perennial vegetation. The spatial distribution of some communities was reduced as a consequence of disturbance; others became fragmented in small patches or even disappeared. In coastal environments with strong, complex gradients, the existence of certain communities depends on specific links (neighbourhood effects) and high connectivity values do not indicate better conservation conditions.     

Biogeographic variability of coastal perennial grasslands at the European scale

Question

Coastal environments have often been described as azonal. While this characteristic is clear for the foredune system, it seems less evident for more inland fixed dunes, which host habitats of major conservation concern, whose features seem to be more related to local climatic conditions. We hypothesized that, unlike other coastal habitats, dune perennial grasslands differ floristically and structurally across their European range and that patterns of variation are linked to the corresponding climate.

Location

European coasts (Atlantic Ocean, Baltic, Mediterranean, Black Sea).

Methods

We used a large data set of phytosociological relevés, representative of coastal grasslands throughout their European range. The role of climatic variables (temperature, precipitation and continentality) in determining the variability in species composition and vegetation structure (by means of life forms) was investigated through CCA, DCA and GLM. The degree of concentration of species occurrences within groups was calculated through the Phi coefficient.

Results

Through multivariate analyses we identified seven major types of coastal grassland, corresponding to different geographic areas. The groups significantly differed in their climatic envelope, as well as in their species composition and community structure.

Conclusion

Our results confirm the hypothesis that coastal dune perennial grasslands are subjected to local climate, which exerts significant effects on both floristic composition and community structure. As a consequence, coastal grasslands are particularly prone to the effect of possible climate change, which may alter species composition and distribution, and lead to shifts in the distribution of native plant communities.

     

Plant community response following the removal of the invasive Lupinus arboreus in a coastal dune system

The removal of invasive species is common in restoration projects, yet the long‐term effects of pest management programs are seldom assessed. We present results of a long‐term program to remove the invasive species Lupinus arboreus (lupin) from sand dunes in New Zealand. We evaluate the response of plant communities to lupin removal, by comparing total plant cover, the cover of non‐native and native plant species, and species richness between sand dune sites where lupin removal has occurred, not occurred, and where lupin has never been present. Neither lupin presence nor removal had a significant impact on the foredune environment. Following removal, total and other non‐native plant cover remained higher, and the cover of several native sand dune species remained lower compared with uninvaded sites in the deflation and backdune environments. These changes can be attributed to persistent effects associated with the invasion of lupin, but have also developed in response to lupin removal. The results of this study have implications for restoration projects in sand dunes. Pest management alone is unlikely to be sufficient to restore plant communities. Given the difficulties in restoring plant communities once an invasive species has established, managers should prioritize actions to prevent the spread of invasive species into uninvaded areas of sand dunes. Finally, the response to lupin invasion and removal differed between dune habitats. This highlights the importance of tailoring a pest management program to restoration goals by, for example, prioritizing areas in which the impacts of the invading species are greatest.     

A comparison of seed banks across a sand dune successional gradient at Lake Michigan dunes (Indiana,USA)

In habitats where disturbance is frequent, seed banks are important for the regeneration of vegetation. Sand dune systems are dynamic habitats in which sand movement provides intermittent disturbance. As succession proceeds from bare sand to forest, the disturbance decreases. At Indiana Dunes National Lakeshore, we examined the seed banks of three habitat types across a successional gradient: foredunes, secondary dunes, and oak savanna. There were differences among the types of species that germinated from each of the habitats. The mean seed bank density increased across the successional gradient by habitat, from 376 to 433 to 968 seeds m⁻², but with foredune and secondary dune seed bank densities being significantly lower than the savanna seed bank density. The number of seeds germinated was significantly correlated with soil organic carbon, demonstrating for this primary successional sequence that seed density increases with stage and age. The seed bank had much lower species richness than that of the aboveground vegetation across all habitats. Among sites within a habitat type, the similarity of species germinated from the seed banks was very low, illustrating the variability of the seed bank even in similar habitat types. These results suggest that restoration of these habitats cannot rely on seed banks alone.     

Above and belowground community development in a marine sand dune ecosystem

Transects across undisturbed marine sand dunes sequentially traversed the following plant communities: littoral, foredunes, intradune complex of ridges and hollows, deflation plain, myrtle forest, and mature conifer forest. Organic carbon levels were low in the littoral zone and increased across the dune ecosystem landward to the forest communities. The highest percentage of nutrients was isolated from the heavy fraction of soil residues. Soil microfloral populations responded to vegetation, physical dune characteristics, and seasonal moisture patterns. Populations of bacteria and actinomycetes were higher in winter than during summer sampling periods in all communities. Populations of microscopic fungi were higher in winter in all the communities except the hollows. The distribution of vesicular-arbuscular mycorrhizal fungi responded to vegetation and sand dune succession but did not display seasonality. Species of Gigaspora and Acaulospora were the most commonly isolated VAM fungi.Sand aggregation increased along the dune transect and was correlated to plant community succession: the most highly aggregated soil was found in the two forest communities. With scanning electron microscopy, sand grains and organic residues were observed entangled by strands of filamentous mciroorganisms. Many of the filaments were of the dimensions of VAM fungi, which may be important for the release of nutrients associated with the cementing agents of sand aggregates and for the survival of early pioneer plants of sand dunes.     

Inter-island but not intra-island divergence among populations of sea oats, Uniola paniculata L. (Poaceae)

Understanding the underlying causes of phenotypic trait variation among populations is important for informing conservation decisions. This knowledge can be used to determine whether locality matters when sourcing populations for habitat restoration. Uniola paniculata is a federally protected coastal dune grass native to the southeastern Atlantic and the Gulf coasts of the USA that is often used to stabilize restored dune habitats. This study uses neutral genetic markers (allozymes) and a greenhouse common garden study to determine the relative contributions of neutral evolutionary processes and natural selection to patterns of phenotypic variation among natural populations of U. paniculata. Seeds were sourced from foredune and backdune populations spanning shoreline-to-landward environmental gradients on each of four Georgia barrier islands. Based on previous work, we expected to find evidence of divergent selection among populations located on the shoreline-to-landward environmental gradient. However, differences among islands, rather than intra-island habitat differences, drive divergent selection on aboveground and total biomass. The lack of evidence for divergent selection across the shoreline-to-landward gradient suggests that previously documented intra-island trait variation is likely due to phenotypic plasticity. Our findings have implications for conservation and restoration efforts involving U. paniculata, as there is evidence for divergent selection among populations located on neighboring islands.     

Environmental Factors Influencing Coastal Vegetation Pattern: New Insights from the Mediterranean Basin

Coastal dune ecosystems show strongly dynamic interactions between abiotic and biotic factors. The relationship between plant communities and environmental factors has been previously studied in oceanic dune systems, but few studies have been conducted along Mediterranean coasts. In this study we analyze the relative contribution of environmental factors associated with two of the main drivers of vegetation zonation: soil and wind. We chose two representative coastal dune systems in the western Mediterranean Basin subject to low levels of human disturbance. Within 54 plots we recorded floristic and environmental data. Vegetation zonation and relationships with environmental variables were investigated through unconstrained and constrained ordinations, correlation, and variance partitioning. Environmental factors shift along the gradient from coastal to inland dunes, concomitantly with the pattern of community types from annual beach communities to shrub-covered fixed dunes. This general gradient is similar both in the Mediterranean and in the oceanic coastal ecosystems, with the same factors that show similar trends along the dune profile. However, our results highlight some peculiarities of the Mediterranean dune systems in relation to the amount of variation explained by environmental factors. While most studies conducted in oceanic ecosystems find that wind-related parameters may control the vegetation zonation, in our study areas we observed a minor importance of the wind-related variables when compared to soil properties. In particular, organic matter and grain-size variability were found to be closely correlated with the distribution of plant communities along the gradient.     

Richness, diversity, and rate of primary succession over 20?year in tropical coastal dunes

The tropical coastal dunes in central Gulf of Mexico have been stabilizing over the last decades resulting in reduced substrate mobility, and promoting primary succession. We describe changes in species richness and diversity in dune vegetation during 20?years. Our questions: (a) Do species richness and diversity increase over time as predicted by models of ecological succession or do they show a hump-backed manner similar to the observations in temperate coastal dunes?, (b) What is the interaction between vegetation cover and diversity and species richness?, (c) Is there a relationship between species diversity and succession rate and does succession rate change over time?, and (d) How do plant functional types change during succession? In order to answer these questions, we set 140 4?×?4?m permanent plots in a mobile dune area and monitored vegetation cover and species richness from 1991 to 2011. In time, diversity increased in a logistic manner toward an asymptotic value once vegetation cover surpassed 60?%. Species richness increased in a humped-back shape, also reaching a maximum peak at 60?% vegetation cover. The succession rate of diversity was measured by the Euclidean distance, and showed a significant humped-back relation, meaning that it was slower in early and late successional stages. The study supports the intermediate disturbance theory. The conservation of coastal dunes vegetation should focus on all, species-poor and species-rich habitats that help to maintain the ecological integrity of these ecosystems. The understanding of community dynamics and diversity patterns becomes an essential component of coastal dune management and conservation.     

The impact of grazing by macropods on coastal foredune vegetation in southeast Queensland

Abstract The extent of grazing by two macropodids, the agile wallaby (Macropus agilis) and the swamp wallaby (Wallabia bicolor) on coastal foredunes on South Stradbroke Island in southeast Queensland was investigated to determine potential impacts on the principal sand colonizing species, sand spinifex grass (Spinifex sericeus). Grazing on spinifex grass on the foredunes of South Stradbroke island can be attributed principally to agile wallabies. Foraging activity by wallabies was higher in areas of high spinifex abundance, however, grazing intensity and impact on spinifex was only important on foredunes with low spinifex abundance. Spinifex consumption by wallabies was also related to a number of factors, especially composition and structure of vegetation in adjacent habitats. Spinifex consumption increased when the abundance of ground cover components (grasses, sedges, forbs) in adjacent habitats was low and structural complexity was high. Grazing on foredunes by wallabies significantly affects the species composition of the foredune community by excluding the establishment of a number of perennial foredune plant species. This may have implications for community succession in coastal ecosystems.     

Changes in soil and vegetation during dune slack succession

Many rare plant species occur in Dutch wet dune slacks, particularly in the Junco baltici‐Schoenetum nigricantis. For nature management it is important to understand the processes controlling the presence of these basiphilous early successional communities, which is why we investigated vegetation and soil development during succession in coastal dune slacks. We compared 12 chronosequential stages of 0, 2, 4, 9, 10, 13, 25, 30, 43, 60, 70 and 85 yr in five different dune slack systems. In four of these locations turf had earlier been removed in order to restore the basiphilous pioneer stage. The main variation in the vegetation is related to the acidification/soil enrichment gradient and the salinity/maritime gradient. During succession, organic matter accumulates and acidification takes place. Maritime influence can buffer the soil and postpone the succession of basiphilous pioneer communities for many years. A significant correlation with age was found for 18 variables. Multiple regressions predicted changes in the vegetation (dependent variables: biomass, cover of Salix repens, Calamagrostis epigejos and Schoenus nigricans) as a function of acidification, organic matter accumulation, increase in available P and presence of Na in the soil. We conclude that natural ageing of the vegetation and the associated process of accumulation of biomass drive succession in this hydrosere. The underlying soil processes are acidification and organic matter accumulation. During succession dominance shifts from S. nigricans to S. repens or C. epigejos. Maintenance of the pioneer character of the habitat is only possible by local intervention or by natural or man‐induced dune forming. The effect of sod‐stripping depends on the environmental conditions and, in case of acidification, success is limited. Succession can be postponed by mowing.     

Multi-scale phylogenetic structure in coastal dune plant communities across the globe

Aims Studies integrating phylogenetic history and large-scale community assembly are few, and many questions remain unanswered. Here, we use a global coastal dune plant data set to uncover the important factors in community assembly across scales from the local filtering processes to the global long-term diversification and dispersal dynamics. Coastal dune plant communities occur worldwide under a wide range of climatic and geologic conditions as well as in all biogeographic regions. However, global patterns in the phylogenetic composition of coastal dune plant communities have not previously been studied.Methods The data set comprised vegetation data from 18463 plots in New Zealand, South Africa, South America, North America and Europe. The phylogenetic tree comprised 2241 plant species from 149 families. We calculated phylogenetic clustering (Net Relatedness Index, NRI, and Nearest Taxon Index, NTI) of regional dune floras to estimate the amount of in situ diversification relative to the global dune species pool and evaluated the relative importance of land and climate barriers for these diversification patterns by geographic analyses of phylogenetic similarity. We then tested whether dune plant communities exhibit similar patterns of phylogenetic structure within regions. Finally, we calculated NRI for local communities relative to the regional species pool and tested for an association with functional traits (plant height and seed mass) thought to vary along sea–inland gradients.Important findings Regional species pools were phylogenetically clustered relative to the global pool, indicating regional diversification. NTI showed stronger clustering than NRI pointing to the importance of especially recent diversifications within regions. The species pools grouped phylogenetically into two clusters on either side of the tropics suggesting greater dispersal rates within hemispheres than between hemispheres. Local NRI plot values confirmed that most communities were also phylogenetically clustered within regions. NRI values decreased with increasing plant height and seed mass, indicating greater phylogenetic clustering in communities with short maximum height and good dispersers prone to wind and tidal disturbance as well as salt spray, consistent with environmental filtering along sea–inland gradients. Height and seed mass both showed significant phylogenetic signal, and NRI tended to correlate negatively with both at the plot level. Low NRI plots tended to represent coastal scrub and forest, whereas high NRI plots tended to represent herb-dominated vegetation. We conclude that regional diversification processes play a role in dune plant community assembly, with convergence in local phylogenetic community structure and local variation in community structure probably reflecting consistent coastal-inland gradients. Our study contributes to a better understanding of the globally distributed dynamic coastal ecosystems and the structuring factors working on dune plant communities across spatial scales and regions.     

Woody plant encroachment impacts on soil carbon and microbial processes: results from a hierarchical Bayesian analysis of soil incubation data

Quantitative methods were used to examine soil properties and their spatial heterogeneity in a 0-year fenced mobile dune (MD0), an 11-year fenced mobile dune (MD11) and a 20-year fenced mobile dune (MD20) in Horqin Sandy Land, Northern China. The objective of the study was to assess the effect of vegetation restoration on heterogeneity of soil properties in sand dunes and to provide a concept model to describe the relationship between vegetation succession and spatial heterogeneity variation of soil properties in the dunes. The results showed that the average values of vegetation cover, species number and diversity, soil organic carbon (C), total nitrogen (N), and electrical conductivity (EC) increased with the increase in fenced age of mobile dunes, while soil water content (0–20 cm) showed the reverse trend. Geostatistical analysis revealed that the spatial heterogeneity of soil organic C, total N, EC, very fine sand content, and soil water content (0–20 cm) increased from MD0 to MD11 with succession from sand pioneer plant to shrub species then decreased from MD11 to MD20 due to continuous development of herbaceous plants. Canonical correspondence analysis (CCA) showed that there was a relatively high correspondence between vegetation and soil factors, suggesting that the major gradients relating soil organic C, total N, EC, pH, slope, very fine sand content, and soil water content are the main factors for the distribution of dune plants and account for 68.1% of the species-environment relationship among the three sites. In addition, the distribution of the sand pioneer plant was positively related to the relative height of the sampling site and soil water content, and that of most herbaceous plants were determined by soil organic C, total N, EC, pH, and very fine sand content in mobile dunes. The conceptual model of relationship between vegetation succession and spatial heterogeneity of soil properties in mobile dunes suggests spatial patterns of soil properties are most strongly related to plant-induced heterogeneity in dune ecosystems prone to wind erosion, and conversely, the magnitude and degree of spatial heterogeneity in soil properties can influence the plant distribution pattern and vegetation succession of mobile dunes.     

An annotated flora of the Monach Isles national nature reserve,Outer Hebrides

Summary

The term machair is commonly applied to the landform/vegetation systems of many dune pasture areas of parts of the Highlands and Islands of Scotland. Definition of such areas appears to rest on locational, morphological, vegetational and soil criteria. Closer examination suggests that the main diagnostic feature is the soil parent material, shell-rich blown sand, which gives rise to high calcium carbonate and pH conditions. Secondary criteria include the morphological maturity of the machair surface, the absence of depositional processes, specific plant associations (notably the elimination of long dune grasses) and a history of grazing and, possibly, cultivation. Machair areas must also be set in a cool, moist maritime climatic regime.     

Concordance between plant species,oribatid mites and soil in a Mediterranean stone pine forest

Biological interactions between above-ground and below-ground organisms are not clearly defined among communities with regard to compositional patterns. The study investigates the concordance of species assemblages between vascular plants and oribatid mites and soil chemical properties with special attention to the role of vegetation structure, i.e. tree, shrub and herbaceous cover, for biological components. Data were collected in a Mediterranean coastal Nature Reserve using sampling design based on random selection of plots with cover of stone pine (Pinus pinea L.) exceeding 15%. Agreement of distribution patterns was verified by Spearman’s rank correlation coefficient applied to pairs of matrices of plot scores by principal component analysis (plants, mites and soil) and the Mantel test. The feasible role of vegetation cover on plant and mite assemblages was tested by redundancy analysis (RDA). Significant correlations were found for biological assemblages, indicating congruent plant–mite compositional patterns. On the other hand, the hypothesis of concordance between biological communities and soil was rejected. Moreover, RDA showed that vegetation cover was a driver of both plant and oribatid mite assemblages. In particular, herbaceous cover proved to be a good proxy for the two biological communities investigated, with different taxa linked to forest clearings and to areas with denser tree cover. Our results indicate that soil features were not of primary importance for below-ground and above-ground community assemblages in the study area. In the light of our findings and ongoing threats in coastal areas, we recommend that management measures be directed at maintenance of diversified vegetation structure, which may ensure above-ground and below-ground biodiversity with diverse biological community assemblages.     

Soil particle sizes and plant communities on coastal dunes

To identify and assess the distribution patterns of coastal dune vegetation along the eastern, southern, and western coasts of South Korea, we investigated the plant communities and soil factors at 30 sites. In all, 12 communities on CCA (canonical correspondence analysis) Axis 1 and 2 could be arranged into 3 groups: 1 ) 2 communities ofElymus mollis andIschaemum anthephoroides, with medium sand contents; 2) 6 communities ofCarex pumila, Carex kobomugi, Ixeris repens, Zoysia mac-rostachya, Calystegia soldanella, andVitex rotundifolia, with coarse sand contents; and 3) 4 communities ofLathyrus japonicus,Glehnia littoralis, Messerschmidia sibirica, andRosa rugosa, with very coarse sand contents. As identified via CCA ordination, the distribution of these groups was positively correlated with soil particle sizes.     

Nutrient limitation and vegetation changes in a coastal dune slack

Abstract. Basiphilous pioneer species are among the most endangered plant species in The Netherlands. They find most of their refuges in young coastal dune slacks, especially on the Wadden Sea islands. For the purpose of nature management it is important to know which processes control the presence of basiphilous pioneer communities, and to learn about the nature of slacks harbouring the concerning successional sequences. In a large dune slack on the Island of Terschelling, we assessed soil nutrient status and tested for nutrient limitation in four chronosequential stages: 2, 6, 37 and ca. 80 yr of age. Stage 2 harboured a basiphilous pioneer vegetation; in the stages 3 and 4 a dense vegetation of dwarf shrubs and grasses occurred. Soil organic matter and nutrient concentrations in each stage were measured in 1991. In 1992 and 1993 fertilizers were applied to all stages to detect nutrient limitation. Rates of accumulation of organic matter, nutrients and above-ground biomass were estimated. When interpreted as successional stages, the different stages represent a sequence as expected on the basis of general successional theory. There was a peak in yearly nutrient accumulation between the 6- and 37-yr old stage and a steady state after ca. 80 yr. Between the first two and the latter two stages a shift occurred from allogenic to autogenic succession which correlated with a shift in emphasis from available nutrients to light availability as limiting resources. Basiphilous pioneer species suffered only deficiency of nitrogen, probably because of their low phosphorus requirements. It is concluded that in dune slack habitats, in addition to a low nutrient availability in general, a very low phosphorus availability favours basiphilous pioneer species to species showing co-limitation of nitrogen and phosphorus as found in some grasses and dwarf shrubs. A comparison between the effects of lime addition and the effects of nitrogen and phosphorus additions suggests that, in the early stages, soil buffering increases the availability of nitrogen and inhibits the availability of phosphorus. Sod cutting is an effective technique for restoring basiphilous pioneer vegetation, when slacks are acidified only superficially and buffering-mechanisms can be reactivated. Yearly mowing and removing of standing crop may prolong the life-span of basiphilous pioneer vegetation, when soil acidification has not yet dropped below pH 6.