The sands of Forvie (Scotland): A case study in geomorphology and conservational management

The Sands of Forvie, National Nature Reserve, Scotland is a complex coastal sand dune system which is associated with the dynamic estuary of the River Ythan. The dune system has developed over more than 5000 years. The south part is a peninsula of dunes, sand hills, sand arcs, erosion and deflation surfaces. The north part is superimposed on a rock plateau with a cliff coastline. This plateau is covered in glacial deposits and is essentially an upland heath landscape. Some of the best examples of large active parabolic dunes in Britain are found in North Forvie. The sequence of geomorphic development is described. As a nature reserve with a rich ecology it has been managed for conservational purposes since 1960s. As such it is an excellent case study of how conservational management has changed to become more flexible and more aware of the importance of dynamic processes.     

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.     

Dynamic preservation of the coastline in the Netherlands

In 1990, the Government and Parliament of the Netherlands decided on a new national policy for coastal defence. To ensure the enduring safety of the low-lying polders as well as the sustainable preservation of the coastal dunes, it was decided to maintain the coastline in its 1990 position. The main method for coastline preservation in the Netherlands is beach nourishment. Since 1991, approximately 7 million m³ of sand is added to the Dutch beaches annually. From an evaluation study of sand nourishment projects it is concluded that nourishment is an effective and efficient method to preserve the coastline. The evaluation study also proves that nourishment projects should be ‘tailor-made’, taking the local morphological and hydraulic conditions into account. The first results of an experimental shoreface nourishment are discussed. The choice for nourishment as the principal method to combat erosion is well in line with the intention to restore the natural dynamics of the dune-coast in the Netherlands. The key probably is a less strict policy with regard to the maintenance of the foredunes. Plans for the development of typical coastal features such as slufters^*, wash-overs, blow-outs and mobile dunes are currently under discussion. A slufter is a breakthrough in the first dune-ridge whereby the sea invades former dune slacks or beach plains situated behind the duneridge. In this way a tidal marsh is formed within the dune zone.     

Landscape ecology of the Dutch coast

This paper is a summary and elaboration of an earlier publication in Dutch on the compilation of a landscape-ecological map, scale 1 : 50 000, of the Dutch coast. It is argued that such an integrated map is the best basis for the conservation and management of the coastal dunes and salt marshes. It may be combined with local more detailed vegetation maps, some examples of which are mentioned in the context of management. The Dutch North Sea coast is a ca. 350 km long chain of sandy beaches and sand dunes, from only 100 m to more than 10 km wide. On sheltered stretches of dune coasts along estuaries in the Southwest and on the Wadden Sea islands, salt marshes have developed. The small-scale gradient structure of the beach-dune-salt marsh complex is emphasized.     

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.     

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.     

Variation in populations of the coastal dune building grassLeymus arenarius in Iceland revealed by endospermal prolamins

Variation in the endospermal protein patterns of seeds amongst and between inland and coastal populations of the dune building grassLeymus arenarius was examined in Iceland. Seeds were collected from six coastal populations and five inland populations in Iceland. Endospermal proteins (prolamins) of seeds were extracted with Tris-buffered 2-propanol (50%) and β-mercaptoethanol (0.5% v/v). We used 8% and 12% sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS-PAGE) to separate the prolamin protein bands, which were stained with Coomassie Brilliant Blue R. Coastal populations were differentiated from inland populations on the basis of high frequency of high molar weight (HMW) (110–150 kg mol⁻¹) prolamins. Coastal populations had significantly higher proportions of their seed with more than two HMW prolamins, than those of inland populations. Subtle differences were found among the HMW prolamins of coastal and inland populations indicating limited intrapopulation variation. The dynamic environment of sand dunes probably influences the genetic composition of these populations. The results suggest that seeds of the inland and coastal populations should be treated separately in sand dune reclamation and restoration works     

Floristic composition and vegetation ecology of the Malindi bay coastal dune field, Kenya

A short outline is given of the floristic composition, structure and distribution of coastal dune vegetation found at Malindi Bay, Kenya. The area was studied by air photo interpretation and field sampling to determine the relationship of plants to aeolian features. TWINSPAN classification was used to distinguish geomorphological units on the basis of their species composition. In this paper, an inventory and first quantitative analysis of vegetation distribution is presented. We identified 174 plant species from 62 families in the sand dunes and several plant communities are distinguished based on the species content and the connection with morphological units.Papilionaceae with 18 species andPoaceae with 17 species were the most represented families. A distinct zonal distribution of the plant communities was found. The most important plant species are the pioneer vegetation consisting ofHalopyrum mucronatum, Ipomoea pescaprae andScaevola plumieri. The woody shrub species which have colonized the established primary dunes and hummock dunes areCordia somaliensis, Pluchea discoridis, Tephrosia purpurea (dunensis). Succulent herbs were identified in the dune slacks and salt marsh that are moist and damp environments.     

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.     

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).     

Trampling impact on vegetation of embryonic and stabilised sand dunes in Montenegro

Trampling is one of the human activities that are harmful for plant species and communities of sand dune ecosystems. The aim of this study was to compare the vegetation of embryonic and shifting Ammophila sand dunes with and without fencing to limit trampling. Fenced sand dunes appeared to be richer in species but differences were more prominent in embryonic sand dunes. Some species (Cakile maritima, Pancratium maritimum) were missing on trampled embryonic dunes. The positive impact of trampling exclusion on embryonic sand dunes was indicated by a lowered slope in a Whittaker graph as well as by rarefaction curves that showed higher species richness on the lower slope. Changes in the vegetation of more stabilised shifting Ammophila sand dunes due to trampling are not evident, although species composition is also impoverished. Fencing of parts of sand dunes proved to be an effective measure for vegetation conservation. In addition to physical exclusion of visitors, fences can also have symbolic value for raising public awareness.     

Changes in landscape and vegetation of coastal dunes in northwest Europe: a review

In coastal dunes, landscape changes are a rule, rather than an exception. This paper gives an overview of changes in landscape and vegetation with a focus on the past century. The history of dunes is characterised by phases of sand drift, alternated with geomorphological stability. The historical development of dune woodland during these stable phases has been documented for sites all over Europe. Vegetation reconstructions of historical open dune habitats however is very difficult due to limited preservation of fossil remains. People have drastically altered coastal dune landscapes through centuries of exploitation and more recently development of the coast. Historical land use has generally pushed vegetation back into a semi-natural state. During roughly the past century a tendency of increasing fixation and succession is observed on coastal dunes throughout northwest Europe. Six causes of change are discussed. 1) Changes in land use, mainly abandonment of agricultural practices, have led to the development of late successional stages such as scrub and woodland. 2) Crashing rabbit populations due to myxomatosis in the 1950s caused vigorous grass growth and probably stimulated scrub development. 3) A general tendency of landscape fixation is observed due to both natural and anthropogenic factors. 4) Eutrophication, mainly due to atmospheric nitrogen deposition is clearly linked to grass encroachment on acidic but also on some calcareous dunes. 5) The impact of climate change on vegetation is still unclear but probably lengthening of growing season and maybe enhanced CO₂ concentrations have led to an acceleration of succession. 6) A general anthropogenisation of the landscape occurs with rapid spread of non-native species as an important consequence. The reconstruction of a natural reference landscape is considered largely unattainable because of irreversible changes and the long tradition of human impact, in many cases since the development of the dunes. Two contradictory elements need reconciliation. First, the general acceleration of succession and scrub and woodland development in particular is partly caused by a decreased anthropogenic interference in the landscape and deserves more appreciation. Second, most biodiversity values are largely linked to open, early succession dune habitats and are threatened by the same tendency. Apart from internal nature management, in which grazing plays an important part, re-mobilisation of stable, senescent dunes is an important challenge for dune management.     

Integrated shoreline management and tourism development on the cross-border World Heritage Site: A case study from the Curonian spit (Lithuania/Russia)

The varied and very dynamic landscape with a high biological diversity is a distinctive feature of the Curonian spit at the regional scale. Throughout the 20th century the main morphodynamic trend in the littoral of the Curonian spit was shoreline grading on the lagoon side, whereas on the marine side the increasing erosion at the foot of the spit, and increasing accretion at the head of the spit prevailed. The results of a discriminant analysis (Wilkes’ λ = 0.001626 and F = 29.267 when p < 0.001) show that sites with prevailing erosion, accretion and sediment input from the drifting dunes form regular inter-related spatial structures in the littoral with distinctive resulting discontinuities of the sediment drift along the lagoon coast. Dune littoral cells are characterized by Aeolian sediment input and distribution ‘down-drift’, (usually northwards) from the source. The most likely changes in the current development trends of the lagoon shore zone of the Curonian spit are related to expected climate changes and further slowing down of the dune drift. The probability of storms and ice-drift events, and their impact on coasts is expected to increase as a result of climate change. The dune advance will gradually slow down, and with it, the sand input to the coastal zone will decline. In this paper, we define integrated shoreline management as a system of long-term shoreline management measures, which is based on a littoral cell approach and aimed at harmonizing human activity in the coastal zone with the natural development of the shoreline. We propose an integrated management program for the lagoon shoreline of the Curonian spit, which is site-specific for each littoral cell as a coastal management unit. Drifting dunes and seaside beaches are the natural amenities, which are best known and best appreciated on the Curonian spit by 49% of the respondents representing the total Lithuanian population. A responsible tourism development should be considered as the key means for proper appreciation of the drifting dunes and natural coasts by society, which means to acknowledge and cherish the aesthetic and conservation values of dune and coastal landscapes of the Curonian spit as a World Heritage Site. There are at least two pre-conditions for this: (1) to provide visitors with sufficient information about diverse values and functions of the Curonian dunes and coasts within a broader regional and global heritage conservation context; (2) to enable tourists to enjoy the most impressive dune and coastal landscapes at close range. In this paper we propose to encourage active dune tourism, to reintroduce grazing into the Curonian dune areas, to restore and to preserve the most impressive landscapes of the highest white drifting dunes by bringing the blown out sand from the leeward foot of the dunes back to the crest artificially.     

Evolution of a sediment-starved,over-stabilised dunefield: Kenfig Burrows,South Wales,UK

The past few decades have seen a progressive reduction in the extent of mobile dune systems and early successional stage habitats in many parts of NW Europe. The evidence suggests that a number of factors have contributed to this process, but their relative importance remains uncertain. This paper examines the nature and possible causes of geomorphological and vegetation changes at Kenfig Burrows, South Wales, a site of European nature conservation importance. This dune system is interpreted to have evolved from a sand barrier system which formerly existed to seaward of the present shoreline and which moved landwards and broke down during the later Holocene, driven by rising sea level and periods of more frequent storms. Most of the inland sand invasion occurred during the Little Ice Age, and large-scale sand-blowing continued until the early twentieth century, by which time the shoreline in the area had reached a state of quasi-equilibrium and a trend towards stabilization began, encouraged by exhaustion of marine sand supply and a change towards warmer, wetter and less windy conditions. Stabilization after the 1940–50s was also favoured by other factors, including reduced grazing by livestock and rabbits, reduced physical disturbance, increased nitrogen deposition, and sand dune management measures. By the 1980s the dunes were almost entirely stabilized. Natural reactivation of the surface sand is unlikely given the existing balance between the factors which favour dune mobility (frequent occurrence of sand-moving winds and high sand supply), and those which encourage vegetation growth and dune stability (high precipitation, high temperatures, low wind speeds, high rates of nitrogen deposition and low grazing / disturbance pressure). Since 2012 trials have been underway to determine if it is possible to increase the mobility of the dune system through intervention measures, including turf stripping and creation of artificial ‘notches’ to increase local wind speeds and sand-transport, but it will be several years before the effectiveness of these measures is known.     

The floristics and conservation status of sand-dune communities in Wales

An inventory of the vascular plants, bryophytes, fungi and terricolous lichens recorded within the different sand-dune communities in Wales has been compiled and a summary of the importance of these different taxonomic groups to the ecology and conservation of sand dunes is provided. The total floristic assemblage amounted to 945 species, with vascular plants representing 439 of these. Fungi unexpectedly formed the next most important group with ca. 289 species, followed by bryophytes (171 species) and terricolous lichens (66 species). However, very few of these species are confined to sand dunes. Of the vascular plants only ca. 7 % (32 species) could be classed as either wholly dependent or strongly associated with sand dunes, whereas up to 13% (22 species) of the bryophytes fell within this category. Of the fungi only 4% (10 species) appear to be restricted to sand dunes, but none of the lichen species could be classed as being either limited to, or strongly associated with, this habitat. Over 9% (91 species) of the total flora are considered to be rare, scarce or endangered within the UK or Europe, and ca. 8% of the vascular plant species are considered to be endemic or near-endemic to Europe. The inventory therefore not only provides an insight in the overall plant diversity of dune systems in Wales, it also gives an indication of the proportion of species that are under threat, and the numbers of species that are more-or-less totally dependent on sand dunes. In the discussion various relevant sand-dune management issues are addressed.     

The development and application of remote sensing to monitor sand dune habitats

Sand dunes are complex systems that contain several habitats, often as mosaics or transitions between types. Several of these habitats are afforded protection under European Legislation and in the UK nationally within Special Areas of Conservation (SAC) and Sites of Special Scientific Interest (SSSI). Natural England has a statutory duty to report to Europe on the conservation status and condition of sand dunes; and is required to report to the UK Government on designated sites. To achieve this we have sought ways of capturing, analysing and interpreting data on the extent and location of sand dune habitats. This requires an ability to be able to obtain data over large areas of coastline in an efficient way. Natural England and Environment Agency Geomatics have worked collaboratively for over 16 years, sharing data and ecological knowledge. In 2012 work started to evaluate the use of remote sensing to map UK BAP and Annex I sand dune habitats. A methodology has now been developed and tested to map sand dune habitats. The key objective was to provide an operational tool that will help to map these habitats and understand change on sites around England. This has been achieved through analysis of LIDAR and Compact Airborne Spectrographic Imager (CASI) data using Object Orientated Image Analysis. Quality Control (QC) and accuracy assessments have shown this approach to be successful and 11 sites have been mapped to date. These techniques are providing a new approach to monitoring change in coastal vegetation communities and informing management of protected sites.     

The formation of protective dune ridge along the Southeast Baltic Sea coast: historical and social aspects

This paper is based on literature-based studies, as well as material collected by the author over more than 30 years on the shores of the Southeast Baltic Sea. In order to protect roads running along the shoreline from East Prussia to Königsberg, local shore reinforcement works commenced in the fifteenth century, although some individual attempts had been made before. In 1684, M. Ch. Hartknoch theoretically substantiated the necessity for, and methods of, shore reinforcement. The state of the shores deteriorated considerably during the Seven Years War (1756–1763), when woodlands growing along the shores were completely destroyed; hence, nearly 100 km length of the coast of the Kur?i? Nerija (Curonian) sand spit was left without any vegetation. Aeolian sand buried as many as fourteen settlements. This situation forced the authorities to organise maintenance of the coast and the dunes at the governmental level, and to start the formation of an uninterrupted dune ridge. The second half of the nineteenth century and the first half of the twentieth century are notable for the extensive construction of new ports. Along the shores running from the Sambian Peninsula to the Gulf of Riga, eight port engineering structures had been constructed with the intervals of approximately 26 km between the ports of Klaip?da and Liepaja, and every 13 km between Palanga and Pape. Shore management measures were being implemented simultaneously, thus ensuring the functioning of the above-mentioned coastal protection structures and the reduction of impacts upon the shores. In the second half of the twentieth century, ports were dredged and developed significantly. Their entrance channels reduced the long-shore sediment transport. Moreover, total sand reserves in the shore zone also diminished. The processes of erosion of the beaches and the dune ridge made it necessary to actively reinforce the shore. A number of countries passed legal acts governing the protection and use of the shore zone. At the turn of the twenty-first century, the incompatibility of such priorities in human activities as sea transport and recreation with nature protection has become evident. The conflict of interests in the Kur?i? Nerija National Park, the port of Klaip?da, and the resort of Palanga serve as good examples of such incompatibility. The conclusion that can be drawn that in twenty-first century is that the priorities in general coastal management should be as follows: harmonisation of interests among the states and individual fields of activities, as well as measures aimed at neutralisation of negative natural trends (higher frequency of storms, global rise of water level, etc.). The protective beach dune ridge will further play the role of efficient means for reducing shore erosion.     

Conservation and management of the mediterranean coastal sand dunes in Israel

The aims of this study are to review the current situation of the Israeli Mediterranean coastal sand dunes, to examine the causes for this situation, and to propose options for future conservation and management of the protected dune areas based on ecological, environmental, landscape and recreational demands and interests. The coastal dunes of Israel are characterized by diverse plant communities, with 173 plant species occurring on sand (8.2% of the total flora of Israel) including many endemic species (26% of all endemic species in Israel). Most of the species are annuals. The importance of the coastal strip as a centre of floral and faunal speciation is also manifested in the existing sand-bound animals. However, many species are rare. This is mainly due to the extensive industrial and urban development along the coastal plain and the direct and indirect destruction of the remaining open dune areas by tourism, recreation and sand mining. Only ca. 17% of the Israeli coastal dunes are still of good or reasonable ecological value, while < 5% of this area has been designated as protected area. Management policies differ from place to place and depend on local objectives. These objectives derive mainly from the knowledge and data that exist for each location, and its statutory status. Since 1995 several projects, which aim to develop integrated management tools for nature conservation and recreation uses for all coastal sand dunes in Israel have been conducted. These projects are summarized in the present paper.     

A CFD study of wind patterns over a desert dune and the effect on seed dispersion

In the Namib Desert seed distribution is greatly influenced by wind patterns. Existing literature regarding wind patterns over dunes focuses on two-dimensional simulations of flow over simplified dune structures. The three-dimensional geometries of the sand dunes suggests far more complex flow features exist, which are not captured by two-dimensional simulations. Computational fluid dynamics (CFD) was used to reproduce the three-dimensional near surface wind patterns around a dune with the aim to learn more about seed distribution. Field work included terrain mapping, wind speed, direction and temperature metering. The CFD results show the expected two-dimensional flow features of high pressure at the dune toe, low pressure at the crest and flow acceleration up windward slope. Also observed are some three-dimensional flow features such as a spiral vortex near the crest and transverse flow due to crest-line curvature of the dune. It was also observed how the wall shear stress differs due to the three-dimensional shape of the dune. The wall shear stress suggests that seed accumulation is more likely to occur behind trailing (down-wind) crest edges. Particle tracking showed how seeds tend to move over the dune crest and recirculate towards the crest on the lee-side. The study showed that adding the third dimension makes the simulations more complex, adds to computational requirements and increases simulation time but also provides vital flow information which is not possible with two-dimensional simulations.     

Coastal sand dune vegetation: An extreme case of species invasion

The coastal sand-dune flora of the Gulf and Caribbean region of Mexico was analyzed to understand differences in floristic composition and richness found along the coast. Each of the 655 species reported was classified according to its ecology and distribution range by checking herbaria specimens, literature and specialists. Three groups were formed: (a) species with predominantly coastal distribution; (b) ruderal or secondary species frequently found inland, common of disturbed areas such as roadsides, abandoned fields or forming part of secondary growths; (c) inland species frequently found in other vegetation types such as tropical dry or seasonal forest and grassland. A total of 71 coastal species, 237 ruderal/secondary and 336 species from other community types were found. The distribution of these groups was analyzed along 44 sites of the Gulf and Caribbean, in the different dune habitats and for the dominant growth forms. Coastal species are more widely distributed; they predominate in habitats with sand movement and the herbaceous component prevails. Ruderal/secondary species and especially those belonging to other vegetation types frequently appear in only one or two sites occupying more protected or stabilized habitats. The two latter groups considerably increase species richness of sand dune flora, but also pose interesting problems for dune conservation.