The role of European marram grass in dune stabilization and succession near Cape Agulhas, South Africa

The coastline near the southern tip of Africa is characterized by large mobile dunes that are driven along wide beaches by strong winds throughout the year. This results in the blockage of the river mouths causing severe flooding of the low-lying farmland of the Agulhas Plain during the rainy winter season. Large parts of the driftsands were stabilized with the European dune pioneer species Marram grass (Ammophila arenaria), which has proved highly invasive along the North American west coast. In order to establish the potential invasiveness ofA. arenaria in South African coastal dune systems and its role in the succession of a large stabilization area, studies were carried out on De Mond Nature Reverve. Using aerial photos, maps and planting records, the vegetation of sites of various ages were sampled. By means of this chronosequence of stands, there is clear evidence that succession takes place at De Mond. Four communities are distinguished, varying from recent plantings ofA. arenaria to mature dune thicket or dune fynbos (heath) vegetation. These relate to four different stages of succession,A. arenaria occurring in reduced abundance in the older communities. After 50 years, formerA. arenaria areas are usually covered by dense dune scrub and in some places even in asteraceous dune fynbos. Succession is most rapid in sheltered, moist dune slacks, butA. arenaria remains vigorous in conditions favourable for its growht, i.e. on exposed, steep dune slopes with strong sand movement.A. arenaria does not appear to spread unaidedly at De Mond and has been successfully used for temporary dune stabilization.     

The importance of coastal sand dunes for terrestrial invertebrates in Wales and the UK,with particular reference to aculeate Hymenoptera (bees,wasps &; ants)

Although most UK sand dune systems are now fossilized, with little mobility and reducing amounts of bare sand, they support important populations and assemblages of terrestrial invertebrates. Offering open conditions, warm substrates and a range of habitats and habitat structures, they have become increasingly significant as other coastal habitats have been lost. In Wales, 680 Red Data Book and Nationally Scarce species have been recorded from dunes. 109 species in the UK are restricted to dunes, and in Wales there are an additional 145 species confined to dunes and 208 species strongly associated with dunes. Of these, 172 species are dependent upon bare and sparsely-vegetated sand, in grey dunes and early-successional dune grassland, at some stage of their life cycle, rising to 292 species if those associated with the strandline, foredunes, yellow dunes and pioneer dune slacks are included, equating to 63% of the 462 dune species. Bees and wasps are particularly well represented, with 278 species (68% of the Welsh fauna) recorded on Welsh dunes, including 17 obligates and 44 species with a strong dependence, 52 of which are associated with bare and sparsely-vegetated sand. Key to maintaining invertebrate populations on UK dunes is the provision of bare sand but in Wales, bare sand accounts for only 1.7% of the total sand dune resource. As a more appropriate bare sand threshold is likely to range between 10 and 30%, radical action is required to re-mobilize at least the key sand dune systems.     

Management of basiphilous dune slack communities in relation to carbonate accumulation and hydrological conditions

In dune slacks in The Netherlands, a decline of rare mesotrophic basiphilous plant species and their plant communities has been observed in combination with an increase of more productive systems with common, taller acidophilous plant species. This has been attributed to both natural and anthropogenic changes. In a humid climate with a precipitation surplus, as in The Netherlands, the calcium carbonate content of a calcareous soil increases with depth. However, soils in coastal dune slacks, may have a higher carbonate content in the topsoil horizon than in the underlying layers. Carbonates which buffer the pH can prolong the presence of mesotrophic basiphilous plant communities which are of high conservation value. To explain the occurrence of calcareous surface horizons in dune slacks, hydrological and micromorphological analyses were carried out in three dune slacks. Two slacks are situated on the Wadden Sea islands in the northern part of The Netherlands; one on Schiermonnikoog and one on Texel. The third slack is situated in the dunes on the island of Goeree in the southwestern part of The Netherlands. In all three slacks, carbonate occurs as mollusc and gastropod fragments (silt- or sand-sized) and as micritic nodules in the topsoil layer, due to aeolian deposition and sedimentation by water.In situ carbonate accumulation (calcitans and calcareous crusts) due to CO₂ release in inundated and/or capillary rise of calcareous groundwater near, or at the soil surface. Accumulation of carbonate also occurs as a result of biological activity by algae in the topsoil of the Goeree site. In general, hydrological processes maintaining high levels of calcareous groundwater are a prerequisite for the maintenance of high carbonate levels in topsoils. Such levels are necessary for the conservation and management of basiphilous pioneer vegetation.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species.     

Sustainability of sand dune restoration along the coast of the Tyrrhenian sea

This study reports the results of restoration management on sand dune environments along the coastal belt of the Castelporziano nature reserve (Rome, Italy) and the subsequent monitoring phases to test the sustainability of the ‘soft techniques’ applied. In the area concerned, over a length of ca. 3 km, 40 dunes were built up along with three belts located at <40 m, 40–70 m, and >70 m, respectively, from the shoreline. On each of 38 dunes 20 individuals ofAmmophila littoralis were planted; this species is one of the local autochthonous species considered particularly suitable for stabilizing sand dunes. After one year, two years and five years, respectively the changes in height and surface of each dune, the survival rates ofA. littoralis, and its changes in cover, the appearance of new shoots and the establishment of new species were observed. A progressive increase in species number, which five years after the restoration amounted to about 60% of those characterizing the natural dunes, was reported indicating a progressive trend towards populations similar to natural ones. In the colonization of new species there is a prevalence of theSporobolus-Elymetum farcti and theSalsolo Kali-Cakiletum maritimae association, while the species established successively refer to theEchinophoro spinosae-Ammophiletum arundinaceae association and theCrucianellion maritimae alliance as occurring in natural successions. This succession runs parallel to the natural dune colonization processes. In particular, the data regarding survival, cover and number of vegetative shoots indicate that the dune belt between 40 and 70 m from the sea is the one most suitable for restoration. Some changes in dune morphology was observed: the height of the artificial dunes tended to decrease considerably in the five years of observation, whereas a progressive increase in their surface area was observed. During the study period.A littoralis favoured the establishment of new species, but as yet exercises no action on increasing dune height.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species. Nomenclature: Roberts (1987, 1995) forAraneae; van der Meijden et al. (1990) for vascular plants; Corly et al. (1981) for bryophytes; Schaminée et al. (1996) for vegetation associations.     

Nutrient deficiency in dune slack pioneer vegetation: a review

A review of results of fertilization experiments in wet dune slacks is presented. In most cases the above-ground biomass appeared to be limited by nitrogen availability. Primary phosphorus limitation was assessed only once in a dune slack where sod cutting had been applied very recently. In most other case studies phosphorus limits biomass production after nitrogen deficiency was lifted. Potassium availability is of minor importance for biomass production in this type of ecosystem. Singular nitrogen additions led to increased dominance ofCarex andJuncus species as well as perennial grasses, such asAgrostis stolonifera andCalamagrostis epigejos. A combined addition of nitrogen and phosphorus led to total dominance of grasses, while the characteristic basiphilous pioneer species (including mosses) decreased or even disappeared. Certain mechanisms are considered which may maintain nutrient availability in slacks with basiphilous pioneer vegetation at a low level, despite of the accumulation of nutrients in the developing organic soil layer. Some implications for management and further research are discussed.     

Distribution and conservation status of littoral vascular plant species along the European coasts

A comprehensive list of 1068 typical littoral plant species and subspecies has been composed. They are considered endemic in a wide sense and are subdivided into widespread, transregional, regional and local endemics, the latter three categories being considered as endemics s.s. For each taxon the distribution, habitat preference, endemic status and conservation status are given. The list, which is available upon request, is summarized in a number of figures and tables, from which it appears that 61% of all species are endemics s.s., that ca. 30% of all species are dune and beach species and another nearly 30% are maritime rock species. Species of wet habitats are concentrated in northern and northwestern Europe, dune species in western and southwestern Europe, western Mediterranean and Black Sea. The conservation status of most species is indicated; 37% is considered threatened. It is concluded that the Bern Convention and the European Habitat Directive offer an entirely insufficient framework for effective conservation action. It is suggested to take the present list as a starting point for a geographical/taxonomical/ecological data base of European coastal endemics.     

Contrasting ecology of prograding coastal dunes on the northwest coast of Ireland

This paper examines the influence of dune morphological evolution on plant species diversity and composition on the Magheramore dune system, Donegal, north-west Ireland. It aims to demonstrate the need for understanding of local geomorphological factors and their affect on ecological processes for the enhanced conservation management of coastal dunes. Vegetation surveys and cluster analysis of species associations, in combination with GIS analysis focused on two prograding dune sites within the Magheramore system. Contrasting geomorphic evolution over similar timescales has imposed site-specific environmental conditions that have had a direct impact on the range of plant species communities present. Cluster analysis revealed 5 main species community types that defined specific dune habitats, the organisation of which is strongly influenced by geomorphic evolution. This study demonstrates the importance of geomorphic evolution and morphological heterogeneity in habitat and species diversity.     

Hydrogeology of a restored coastal dune system in northeastern Scotland

This paper reports the results of 12 years of hydrological monitoring at the St. Fergus dune system in northeastern Scotland. The site is adjacent to the UK’s largest gas terminal and the dunes are crossed in five places by North Sea gas pipelines which were constructed between 1976 and 1990. These are buried beneath the dune system which was restored after pipeline installation. The dunes include a substantial freshwater wetland which is seasonally flooded and provides an important habitat for waterfowl. The hydrogeology of the site is characterized and the hydrogeological processes that sustain this wetland feature are considered including recent climatic fluctuations. Nomenclature: Names of vascular plants follow Clapham et al. (1962).     

From overexploitation to sustainable use,an overview of 160 years of water extraction in the Amsterdam dunes,the Netherlands

Dune slacks are low-lying, nutrient-poor, species-rich, inter-dunal, seasonally flooded wetlands, are amongst the most threatened habitats in the Dutch coastal dunes. Since 1853 Waternet has been extracting groundwater from the coastal dunes southwest of Haarlem to produce its drinking water. Dune slacks largely disappeared due to the desiccation caused by this water abstraction, over more than a century biodiversity declined as a consequence. Increased societal concern pushed habitat restoration high on the political agenda by the end of the 1980s. It was agreed to do what is possible to restore original dune slacks without endangering the water supply. Far reaching interventions in the dune water system were foreseen to achieve this mutual goal. To allow reliable decision making, the entire hydrological history of the drinking water production in the Amsterdam Dunes since 1853 and its ecological consequences were evaluated over a 10-year study period. The main tool was a 3D groundwater model constructed using all information gathered to date and calibrated using the long-term monitoring data available and widely extended for the purpose, to which ecological modeling was added and calibrated with the available long-term and extended vegetational inventories. These scientific tools were used to assess proposed interventions to be decided upon, which aimed at finding a new balance between groundwater extraction and nature restoration. In 1996 and 2007 large-scale measures were taken, which include filling in of recovery canals, mowing, grazing and sod-cutting to support the native plant communities of wet dune slack habitats. Results of these measures in terms of the restoration of natural hydrological conditions are shown together with the first results for the recovery of wet slacks vegetation that resulted from the combined hydrological and ecological restoration measures that were taken since 1995.     

Using earth observation data from multiple sources to map rare habitats in a coastal conservation area

Kenfig NNR (National Nature Reserve) is a coastal sand dune system in south Wales, UK. The site is an important location for the conservation of the fen orchidLiparis loeselii, a significant proportion of the UK population is found solely on the site. Approaches to the mapping and monitoring of the habitats at Kenfig NNR using EO (Earth Observation) methods are investigated. Typical airborne EO missions over such sites produce more than a single source of EO data; these may include various optical imaging sensors with different spectral ranges, film cameras and ranging devices to measure topography. Conservation managers are thus presented with the problem of which sources of data to use when producing a land cover map of the site of interest. Using a data set gathered over the Kenfig NNR site, we investigate land cover mapping methods for conservation. The land cover types of interest typically cover small areas within a much larger site so they present a hard problem for the EO data and associated classification methods to solve. Land cover classifications produced from the data sets provide a set of competing hypotheses of land cover type for the site. Methods we use to resolve this competition between the data sets include voting methods, data fusion methods and a method utilising fuzzy logic to aggregate information. This paper is intended to act as an introduction to some of the issues involved in using EO data for habitat mapping in highly heterogeneous coastal dune environments and to present some preliminary results of the performance of each method.     

Vegetation dynamics on the Łeba Bar/Poland: a comparison of the vegetation in 1932 and 2006 with special regard to endangered habitats

Coastal dune ecosystems are among the most dynamic habitats with high conservation value in Europe but are also under strong anthropogenic pressure regarding coastal protection and recreation. Hence, it is of high importance to know about long-term natural and anthropogenic changes and development of protected dune habitats for nature conservation, as well as for coastal management. This paper aims to identify the vegetation dynamics on the Łeba Bar/Poland over a period of 74 years by cartometric comparison using modern Geographical Information System (GIS). To quantify the rate of vegetation dynamics two aerial photographs dating from 1932 and 2006 were digitalized and analyzed with GIS to produce digital vegetation maps. Information about decrease, increase and stability of vegetation types of this area are discussed. The results show that there has been a clear reduction in the total area of bare sand and a considerable increase of woodlands and dense grass communities. Nevertheless, the remaining extensive drift sand areas and deflation hollows on the Łeba Bar offer one of the most important habitats for pioneer vegetation on bare sand of migrating dunes and dunes at the Southern Baltic coast. The present work proved the need to observe the future development of the vegetation communities and to implement management measures to maintain the dynamic of this unique dune landscape.     

Factors affecting vegetation establishment and development in a sand dune chronosequence at Newborough Warren,North Wales

Newborough Warren is a large calcareous west coast UK dune system, which has experienced rapid vegetation spread in the last 70 years. Information from two high resolution chronosequences for dry and wet dune habitats, 0–145 years, was used to answer the following questions: Does climate influence colonisation of vegetation on bare sand? What are the timescales and sequences of successional change in the vegetation? Analysis of aerial photographs showed that stabilisation of the dune system since 1945 has occurred in three main phases. The onset of stabilisation predated myxomatosis by 10 years; while stabilisation virtually halted during the period 1964–1978. Periods of rapid stabilisation were coincident with higher values of Talbot’s Mobility index (M)?>?0.3. Successional development was apparent in both dry and wet habitats. Fixed dune grassland started to replace earlier successional communities at around 40 years, and could persist to 145 years. Linear succession in dune slacks was less apparent, but a separation between communities typically regarded as ‘younger’ and ‘older’ occurred at around 40 years. Species richness in dry dune habitats increased with age to a maximum on soils around 60 years old, then declined again. Species richness was unrelated to age or soil development in wet dune slacks. The influence of climate suggests that conservation managers can only operate within the constraints imposed by natural climatic conditions. Vegetation growth and soil development are closely linked and maintaining some open areas is key to preventing soil development and over-stabilisation.     

Ecohydrological studies of dune slack vegetation at Kenfig dunes,South Wales,UK

Dune slacks are important coastal sand dune habitats and seasonal changes in water levels within dune aquifers control both their formation and the specific hydrological conditions which then govern the floristic composition of their characteristic plant communities. Kenfig Dunes National Nature Reserve is one of the largest dune sites in South Wales and Southern Britain. It supports an exceptional range of dune slack communities, including most of those recognised in the British National Vegetation Classification scheme. Detailed studies of the vegetation ecology and hydrology of dune slacks reveal the important influence of hydrological variables in controlling the composition of dune slack vegetation and also valuable information on water table profile and the key factors governing the annual hydrological budget of the dune system aquifer.     

Dynamic coastal dune spit: the impact of morphological change on dune slacks at Whiteford Burrows, South Wales, UK

Whiteford Burrows is a coastal dune spit wetland in South Wales that is susceptible to morphological change. The height of the ridge of groundwater within the sand aquifer is essentially proportional to the width of the spit. The water table elevation impacts both the frequency and duration of slack flooding events and, therefore, slack ecology. A severe late winter storm event on 17 March 1995 caused extensive erosion of the foreshore, reducing the effective width of the dune system by 4 % and the water table elevation by up to 1 m. This observed relationship allows water level elevations in the dune system to be hindcast using historical maps and air photos which record past change in dune morphology. These historical data indicate that the dunes were relatively broad in the nineteenth century and the slacks were humid and liable to regular winter flooding. The system slowly dried out towards the 1940s as the spit thinned, when subsequent widening allowed the water table to rise and once again flood slack floors in winter. Despite these changes, the alkalinity of the Whiteford Burrows dune system has inhibited organic matter accumulation and maintained conditions needed for the persistence of a diverse basiphilous vegetation assemblage in many of the slacks.     

Early development of vegetation in restored dune plant microhabitats on a nourished beach at Ocean City, New Jersey

Topography and vegetation of restored dunes on a developed barrier island were examined after a large-scale beach nourishment project. Restoration began in 1993 using sand-trapping fences andAmmophila breviligulata Fern. plantings. Subsequent growth of dunes was favored by installing new fences and suspending beach raking to accommodate nesting birds. Plant species richness, percent cover of vegetation, and height ofA. breviligulata were sampled in 1999 on seven shore perpendicular transects in six dune microhabitats (backdune, primary crest, mid-foredune, swale, seaward-most fenced ridge, incipient dune on the backbeach). A total of 26 plant taxa were found at all seven sites. Richness and percent cover were greatest in the backdune and crest, especially in locations that predated the 1992 nourishment. Richness was greater where fences enhanced stabilization. Fences initially compensate for time and space and allow vegetation to develop rapidly, but maintenance nourishment is required to protect against wave erosion and ensure long-term viability of habitat. An expanded environmental gradient is an option, where beach nourishment provides space for a species-rich crest and backdune to develop, while the incipient dune remains dynamic. Options where space is restricted include a dynamic, full-sized seaward section of a naturally functioning dune (truncated gradient) or a spatially restricted sampler of a wider natural dune (compressed gradient) maintained using fences. Expanded and truncated gradients may become self-sustaining and provide examples of natural cycles of change. Compressed gradients provide greater species richness and flood protection for the available space, but habitats are vulnerable to erosion, and resident views may be impaired.     

Sponge-feeding by the Caribbean starfish Oreaster reticulatus

The common Caribbean starfish Oreaster reticulatus (Linnaeus) feeds on sponges by everting its stomach onto a sponge and digesting the tissue, leaving behind the sponge skeleton. In the San Blas Islands, Republic of Panama, 54.2% of the 1549 starfish examined from February 1987 to June 1990 at eight sites were feeding, and 61.4% of these were feeding on sponges, representing 51 species. Sponges were fed on disproportionately heavily in comparison to their abundance, which was only 9.7% of available prey. In feeding choice experiments, 736 pieces of 34 species of common sponges from a variety of shallow-water habitats, and also 9 ind of a coral, were offered to starfish in individual underwater cages. Acceptance or rejection of sponge species was unambiguous for 31 of the 34 species, and there was a clear relationship between sponge acceptability and sponge habitat. Starfish ate 16 of 20 species that normally grow only on the reefs, but only 1 of 14 species that live in the seagrass meadows and rubble flats surrounding the reefs. The starfish live in the seagrass meadows and rubble flats, and avoid the reefs, and so the acceptable reef sponges are generally inaccessible until a storm fragments and transports them into starfish habitat. After Huricane Joan washed fragments of reef sponges into a seagrass meadow in October 1988, starfish consumed the edible species. When the seagrass meadow was experimentally seeded with tagged reef sponge fragments in June 1994, O. reticulatus consumed edible species and accumulated in the area seeded. Reef sponges that were living in a seagrass meadow, from which O. reticulatus had been absent for at least 4 yr (from 1978 to 1982), were eliminated when the starfish migrated into the area, and the sponges have been unable to recolonize up to June 1994. O. reticulatus feeding and habitat preferences appear to restrict distributions of many Caribbean reef sponge species to habitats without O. reticulatus and may have exerted significant selective pressure on defences of those sponges that live in O. reticulatus habitats.     

A framework for the management of sand dune systems in Wales

The Countryside Council for Wales (CCW) is developing a management framework with the primary aim of restoring favourable conservation status to the sand dune resource of Wales. It will take onboard the requirements of both national and international conservation legislation and will also help CCW integrate its responsibilities for biodiversity, geodiversity, landscape, access and recreation for this habitat. In order to achieve certain conservation goals it will be necessary to have in place a variety of different types of management ranging from non- or minimal intervention through to intensive single species management and habitat re-creation. However, it will not provide a comprehensive framework for all aspects of site management, but only those that are deemed to be of strategic importance, and have significance within an all-Wales perspective for their nature conservation importance.