Limiting factors for seedling emergence and establishment of the invasive non-native Rosa rugosa in a coastal dune system

The relative importance of seed, habitat and microhabitat limitation has rarely been investigated for invasive non-native species, although this is critical for their effective management and for predicting future range expansion. Rosa rugosa is an abundant non-native shrub in coastal habitats of NW Europe; it is common along the Baltic coast but more scarce in exposed dunes of the North Sea. To investigate whether invasion of exposed dunes is limited by seed, habitat and microhabitat limitation, seedling emergence and establishment were examined in a factorial sowing, transplant and disturbance experiment. Twenty plots were randomly placed in each of five dune habitats (white dune, Empetrum dune, grey dune, outer dune heath, inner dune heath), and studied over 2 years. Seedling emergence in control subplots was zero in all habitats, whereas 96% and 98% of the undisturbed and disturbed seeded subplots produced seedlings. Disturbance had a positive effect on emergence and subsequent survival in white dune, outer and inner dune heath. Seedling survival and growth, and growth of transplanted seedlings, were markedly lower in grey dune. These findings indicate that establishment of R. rugosa is seed-limited in coastal dune habitats, and that the species is able to establish in both active and fixed dunes once seeds have arrived. Although differential seedling emergence and growth indicate that habitats differ in their degree of invasion susceptibility by R. rugosa, the positive influence of small-scale disturbance suggests microhabitat limitation in some dune habitats as well. Dune management should aim to reduce seed production and dispersal of R. rugosa in near-natural sites, and anthropogenic changes of habitat dynamics should be prevented.     

Responses of Dune Plant Communities to Continental Uplift from a Major Earthquake: Sudden Releases from Coastal Squeeze

Vegetated dunes are recognized as important natural barriers that shelter inland ecosystems and coastlines suffering daily erosive impacts of the sea and extreme events, such as tsunamis. However, societal responses to erosion and shoreline retreat often result in man-made coastal defence structures that cover part of the intertidal and upper shore zones causing coastal squeeze and habitat loss, especially for upper shore biota, such as dune plants. Coseismic uplift of up to 2.0 m on the Peninsula de Arauco (South central Chile, ca. 37.5º S) caused by the 2010 Maule earthquake drastically modified the coastal landscape, including major increases in the width of uplifted beaches and the immediate conversion of mid to low sandy intertidal habitat to supralittoral sandy habitat above the reach of average tides and waves. To investigate the early stage responses in species richness, cover and across-shore distribution of the hitherto absent dune plants, we surveyed two formerly intertidal armoured sites and a nearby intertidal unarmoured site on a sandy beach located on the uplifted coast of Llico (Peninsula de Arauco) over two years. Almost 2 years after the 2010 earthquake, dune plants began to recruit, then rapidly grew and produced dune hummocks in the new upper beach habitats created by uplift at the three sites. Initial vegetation responses were very similar among sites. However, over the course of the study, the emerging vegetated dunes of the armoured sites suffered a slowdown in the development of the spatial distribution process, and remained impoverished in species richness and cover compared to the unarmoured site. Our results suggest that when released from the effects of coastal squeeze, vegetated dunes can recover without restoration actions. However, subsequent human activities and management of newly created beach and dune habitats can significantly alter the trajectory of vegetated dune development. Management that integrates the effects of natural and human induced disturbances, and promotes the development of dune vegetation as natural barriers can provide societal and conservation benefits in coastal ecosystems.     

Regional and local variation of spider assemblages (Araneae) from coastal grey dunes along the North Sea

Aim This study aims to determine the underlying causes of local and regional patterns of variation in community structure of spiders in coastal grey dunes, and especially whether ecological time constrains the species composition in young and isolated grey dune habitats. Location The study was conducted in coastal dunes from northern France (Boulonnais, Nord‐Pas‐de‐Calais), Belgium (Flemish coastal dunes) and the Netherlands [Amsterdam Water Supply (AWD) dunes and Dunes from the Provincial Water Company North‐Holland]. Methods Spiders were collected with pitfall traps in twenty‐eight grey dune patches in the four areas under investigation. Species composition and environmental parameters (vegetation structure, distance to the sea and the patch‐edge, eolic dynamics and lime richness) were determined. Assemblage composition was related to the regional and local environmental factors with the Primer software package to determine the assemblage‐determining parameters. Differences in species presence were analysed as a function of their habitat preference and distribution range. Results Differences in grey dune spider assemblage structure can mainly be attributed to differences in local sand dynamics and the region. Species from dynamic dunes are mainly present in grey dunes from Belgium and France, while species from non‐dunal xerothermic habitats (chalk grasslands and heathland) occur in both the Boulonnais and the north Holland dune region. These species are absent from geologically young or other xerotherm habitat isolated Flemish coastal dunes. Main conclusion The data show that regional variation in spider assemblage composition results from local landscape characteristics (dynamics in the dune area), the latitude and the connectivity to non‐dunal xerothermic habitats. The strong and moderate geological isolation of dune areas from the Flemish coast and the AWD dunes, respectively, results in the absence of (at least some) species that are primarily bound to heathland and/or chalk grassland. This indicates the importance of ecological time for the assemblage structure. The limited dispersal capacity of the heathland and/or chalk grassland species is probably the main reason for their absence.     

Sexual polymorphism,growth, and reproductive effort in Egyptian Thymelaea hirsuta (Thymelaeaceae)

Gender phenotypes of Thymelaea hirsuta (L.) Endl. were surveyed in six major habitats in the western Mediterranean region of Egypt. Five gender phenotypes were observed at all sites: subandroecious (male), subgynoecious (female), protogynous, protandrous, and gender-labile individuals. Patterns of vegetative growth, reproductive effort, and sex-size relationships were also determined. Females and males were comparable in overall abundance (35.5% and 30.2%, respectively). However, the frequency of sex forms varied significantly among habitats. Plant size (canopy volume) was used as a measure of environmental quality for the species: the coastal dunes were by far the most favorable habitat, followed by the nonsaline depressions, inland plateau, and inland siliceous deposits. The smallest plants were associated with the inland ridges and saline depressions. Protandrous individuals, and female shrubs at the more favorable habitats (e.g., the coastal dunes and nonsaline depressions), had greater canopy volume than males. In contrast, males were larger than females at the less favorable habitats (e.g., the inland plateau and saline depressions). Advantageous growth conditions at the coastal dunes are demonstrated by the greater rate of increase in crown diameter and crown volume noted there, for each size class and sex form. The greatest rates of increase were present in smaller-sized individuals. Size differences between the sex forms were considered in terms of trade-offs between individual growth rate (GR) and reproductive effort (RE); evidence of a trade-off was mixed. Indeed, in a comparison of GR and RE at two extremes of habitat (coastal dune vs. inland plateau), females had greater GR and greater RE at the relatively benign coastal dunes, while at the more harsh, inland plateau site males showed greater GR and RE than females. Peak reproductive effort varied significantly with gender phenotype and habitat type.     

How does dune morphology shape coastal EC habitats occurrence? A remote sensing approach using airborne LiDAR on the Mediterranean coast

We examined the relationship between coastal habitats (sensu European Union Habitats Directive) and local dune morphology along a Mediterranean coastal dune system by integrating field collected vegetation data and remotely sensed imagery. Specifically, we described the morphological profile of each EC habitat based on the morphological variables that are most likely to affect their occurrence, including elevation, slope, curvature, northness, eastness and sea distance. In addition, we assessed the role and strength of each morphological variable in determining the occurrence of EC habitats.We used 394 random vegetation plots representative of six EC habitats (Habitat 1210: “Annual vegetation of drift lines”; Habitat 2110: “Embryonic shifting dunes”; Habitat 2120: “Shifting dunes along the shoreline with Ammophila arenaria”; Habitat 2210 and 2230: “Crucianellion maritimae fixed beach dunes” and “Malcolmietalia dune grasslands”; Habitat 2250: “Coastal dunes with Juniperus spp.”; Habitat 2260: “Cisto-Lavanduletalia dune sclerophyllous scrubs”) found along the Tyrrhenian coast of central Italy. We derived each morphological variable from a DTM (Digital Terrain Model) obtained from 2-m resolution LiDAR (Light Detection And Range) images. The mean value of each variable was calculated at different spatial scales using buffer areas of increasing radius (2 m, 4 m, 8 m) around each vegetation plot. Mean morphological values for each EC habitat were compared using Kruskal-Wallis rank test. The role and strength of the relationship between habitat type and the morphological variables were assessed using Generalized Linear Models.EC habitats occur differentially across dune morphology, and the role and strength of each morphological variable define habitat specificity. Dune elevation and sea distance were determined to be the key factors in shaping EC habitat occurrence along this section of the Mediterranean coast. Identification of the close relationship between habitat type and morphological variables deriving from airborne LiDAR imagery points to the high potential of such remote sensing tool for analyzing and monitoring the integrity of coastal dune ecosystems. As airborne LiDAR enables the rapid collection of extremely accurate topographic data over large areas, it also offers useful information for the management of these threatened and fragile ecosystems.     

Habitat preferences of an endangered insect species,Cepero’s ground-hopper (<Emphasis Type="Italic">Tetrix ceperoi</Emphasis>)

Preventing the global decline in biodiversity is a major task for conservation biologists. Although habitat loss has been identified as a key factor driving extinction processes, our knowledge on the habitat requirements of many endangered species, particularly invertebrates, is still sparse. We present a feasible method to study the microhabitat preferences of insect species. In Central Europe, the endangered Cepero’s ground-hopper, Tetrix ceperoi, is believed to have its only remaining natural habitats in dune slacks of the Wadden Sea Islands. Our results suggest that this species performs an active habitat choice of damp, bare patches with high temperatures. While ponds and fens in dune slacks provide large areas of damp bare ground and algal mats, grasslands, degraded dune slacks and the transitional zone between salt marsh and dunes are less suited as habitats. The major threat for T. ceperoi is found in the succession of its pioneer habitats due to the reduced natural dynamics. In industrialized countries, pioneer habitats and species are threatened substantially by coastal protection and floodplain regulation. This is only compensated in part by anthropogenic creation of secondary habitats, such as different kinds of pits or coal heaps. Nevertheless, there is a strong need for restoration of dynamic habitats by floodplain revitalization and dune slack restoration.     

Are there habitats that contribute best to plant species diversity in coastal dunes?

The following paper describes patterns of diversity across major habitat types in a relatively well preserved coastal dune system in central Italy. The research addresses the following questions: (a) whether different habitats defined on the base of a land cover map support similar levels of biodiversity in terms of vascular flora richness and number of rare and endangered species, and (b) how each habitat contributes to the total species diversity of the coastal environment. A random stratified sampling approach based on a detailed land cover map was applied to construct rarefaction curves for each habitat type and to estimate total species richness. In addition, the number of exclusive, rare and endangered species was calculated for each habitat type. Results highlight the importance of the coastal dune zonation (embryo-dune, main dune, transition and stabilized dune) in species conservation because they harbour progressively higher species richness. However, differences among these habitats were not significant, so no particular species rich “hotspots” could be evidenced. On the contrary, rarefaction curves show that the upper beach (strand) habitat sustains significantly smaller number of species, but surprisingly, it shows the highest rarity values and highest proportion of endangered species. Therefore, for the establishment of successful biodiversity conservation programs in these coastal environments, it is imperative not only to conserve biologically rich hotspots but also to include species poor habitats containing endangered or unique elements. Thus, the complete coastal vegetation mosaic including all coastal habitats is important to adequately characterize the plant species diversity of coastal dune ecosystems.     

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.     

Environmental factors and ground disturbance affecting the composition of species and functional traits of ground layer lichens on grey dunes and dune heaths of Estonia

A unique, species‐rich and endangered lichen biota can be found on European coastal and inland sand dunes. However, it is increasingly affected by natural succession as well as by anthropogenic disturbances. We studied lichen diversity on the grey dunes and dune heaths of coastal and inland regions of Estonia. A total of 28 study plots were investigated; in each 0.1 ha study plot general environmental variables and anthropogenic disturbances were described and all epigeic lichen species were identified. We found 66 lichenized fungus (lichen) species, including several rare and ten red‐listed lichens. Multivariate analysis (DCA, CCA) was performed to examine gradients in species composition and to relate variation in species data to environmental factors. In addition, we used redundancy analysis (RDA) to relate variation in species’ trait composition to environmental factors. Species composition on grey dunes differed significantly from that on dune heaths. The characteristic species for grey dunes are, besides several Cladonia species, foliose lichens, e.g. Hypogymnia physodes, Parmelia sulcata and Peltigera spp. Also species’ traits composition was different for either habitat, indicating that sorediate lichens, foliose lichens, lichens with cyanobacterium as the main photobiont, and sparsely branched Cladonia species dominate on grey dunes, while esorediate, green‐algal, crustose and richly branched fruticose lichens are common on dune heaths. Soil pH was the most essential environmental variable for determining both species composition and species’ traits composition. The composition of lichen species was also significantly influenced by forest closeness, soil Mg content and cover of bare sand; the effect of ground disturbances was low compared to the effect of these environmental factors. To protect and conserve the species‐rich lichen biota, it is necessary to protect the dune habitats from building activity, to avoid overtrampling in recreation areas and to regularly remove shrubs and trees.     

Habitat Use and Mobility of Two Threatened Coastal Dune Insects: Implications for Conservation

We studied the habitat use and mobility of the Grayling butterfly (Hipparchia semele) and the Blue-Winged Grasshopper (Oedipoda caerulescens), two threatened insects within spatially structured blond and grey dunes in a nature reserve along the Belgian coast. Although both species occur in the same biotope, H. semele were more abundant in open, dynamic sites with a relatively high amount of bare sand, while O. caerulescens preferred sheltered, more stable environments with a lower amount of bare sand. Unlike H. semele, substrate use varied in accordance to body colouration in O. caerulescens, especially on cold days, with light-coloured animals being more abundant on sand and dark-coloured animals more abundant on moss. During a mark-recapture-study, we marked 493 Grayling butterflies and 1289 Blue-Winged Grasshoppers. On average, both sexes of H. semele were equally mobile (about 150 m/day; maximum recorded distance of about 1700 m) while male O. caerulescens were significantly more mobile than females (daily average 47 vs. 5 m; maximum distances observed for O. caerulescens were about 800 m). The importance of habitat heterogeneity (within and among patches) and the consequences of habitat use and mobility of both species for the conservation of typical coastal dune habitats are discussed. The complementary use of species-specific information to site-based management measures is advocated.     

Habitat suitability models for the sand lizard Liolaemus wiegmannii based on landscape characteristics in temperate coastal dunes in Argentina

The knowledge of environmental variables associated with the species occurrence allows the recognition of sites which fulfil ecological requirements eventually used for conservation of species. The coastal dunes of Argentina are inhabited by sand lizards. Anthropic activities have severely degraded this ecosystem, affecting the habitat structure at a large scale. In this context, the effects of landscape characteristics on the sand lizard's (Liolaemus wiegmannii, Liolaemidae) presence were analysed to build habitat suitability maps along the coastal dunes of Argentina. A thematic map of study area was obtained from supervised classification of satellite images to identify landscape characteristics. Surveys were conducted during the lizard activity season, and landscape variables were measured in two spatial units. All information collected was compiled into a Geographic Information System. The relationship between the presence of lizards and landscape variables was evaluated by Generalized Linear Models. The predictions of these models were transferred by using Geographic Information System to habitat suitability maps. Almost all individuals (80%) were observed in semi‐fixed dunes. The analysis of landscape metrics in the two spatial extents showed complementary results. The habitat suitability models suggest that: (i) heterogeneous landscapes composed by disaggregated patches of semi‐fixed dunes and low or null percentages of active dunes distant from the coastline are the preferred environments, and (ii) human modifications such as urbanizations and forestation of dunes, have a negative impact on species occurrence. Suitable habitats were almost absent in those sectors of coastal dunes with highest level of urbanization, whereas they were distributed almost continuously in those areas without human disturbances.     

Climate and coastal dune vegetation: disturbance, recovery, and succession

The sand dune habitats found on barrier islands and other coastal areas support a dynamic plant community while protecting areas further inland from waves and wind. Foredune, interdune, and backdune habitats common to most coastal dunes have very different vegetation, likely because of the interplay among plant succession, exposure, disturbance, and resource availability. However, surprisingly few long-term data are available describing dune vegetation patterns. A nine-year census of 294 plots on St. George Island, Florida suggests that the major climatic drivers of vegetation patterns vary with habitat. Community structure is correlated with the elevation, soil moisture, and percent soil ash of each 1 m² plot. Major storms reduce species richness in all three habitats. Principle coordinate analysis suggests that changes in the plant communities through time are caused by climatic events: changes in foredune vegetation are correlated with temperature and summer precipitation, interdune vegetation with storm surge, and backdune vegetation with precipitation and storm surge. We suggest that the plant communities in foredune, interdune, and backdune habitats tend to undergo succession toward particular compositions of species, with climatic disturbances pushing the communities away from these more deterministic trajectories.     

Hard times for Italian coastal dunes: insights from a diachronic analysis based on random plots

Multi-year temporal studies are invaluable tools for monitoring changes in biodiversity through time. However, their applications in coastal ecosystems are still scarce. We investigated temporal trends in coastal dunes analyzing a set of 858 randomly-sampled georeferenced relevés performed between 2002 and 2015 along Central Italy’s sandy coastlines. Specifically, we explored changes in species richness and cover of targeted sandy habitats, we investigated trends in the cover of selected psammophilous native species and we assessed patterns of invasion by means of regression techniques. We observed a significant decrease in species richness and cover of the dune grasslands habitat. The species-level analysis confirmed a negative trend for two characteristic species of dune grasslands, Cutandia maritima and Medicago littoralis, while revealing a similar decline for Crucianella maritima and for Ammophila arenaria subsp. australis, key species of mobile dunes. The most striking trends emerged analyzing patterns in the cover of an invasive alien species, Carpobrotus sp., which showed a concerning increase in shifting dunes. In conclusion, our analyses reveal concerning changes involving dune grasslands, and at the same time hint at “early warnings” of degradation processes traceable in shifting dunes.     

Evidence for ecological release over a fine spatial scale in a lizard from the White Sands formation

When a population experiences relaxation of selective pressures due to reduced predation or competition, it may undergo ecological release. Ecological release often manifests as increased niche breadth and trait variation, as individuals have the opportunity to exploit a wider variety of resources. At the recently‐colonized White Sands dune formation in New Mexico, lizards from white sand habitat have fewer predators and competitors than their dark soil counterparts, and show evidence of ecological release compared to dark soil lizards. To determine whether the dynamics of ecological release also play out over an even finer habitat gradient, we studied southwestern fence lizards in the center of the White Sands dune formation and the narrow ecotone between White Sands and the surrounding dark soil habitats. We predicted that lizards from the central dunes would exhibit ecological release in terms of broadened resource use compared to lizards from the ecotone. We first conducted avian surveys in both the central dunes and ecotone habitats to measure abundance, richness and diversity of avian species that could act as lizard predators. Next, we measured microhabitat‐scale resource use by comparing perch selection of lizards in both habitats. Finally, we measured landscape‐scale resource use by quantifying home range sizes and daily distances traveled by lizards in both habitats. We found that central dunes lizards used a greater diversity of perch types, and had greater variation in both home range size and daily distances traveled, than ecotone lizards. There were fewer predatory bird species in the central dunes than on the ecotone, and there was a tendency for fewer predation events in the central dunes than on the ecotone. Our results demonstrate that ecological release is detectable over a fine habitat gradient, such as between the center of a recently‐colonized habitat and its ecotone with the surrounding ecosystem.     

Effects of long‐term grazing management on sand dune vegetation of high conservation interest

Question: Can long‐term grazing management maintain and restore species‐rich sand dune plant communities within a sand dune site of high conservation interest? Location: Newborough Warren, North Wales, UK. Methods: Vegetation changes that occurred between 1987 and 2003, subsequent to grazing by domestic livestock being introduced to the site after decades with little or no stock grazing, were analysed using data collected from permanent monitoring quadrats over a 16‐year period. Results: At the plant community level, grazing brought about a shift from a tall‐grass dominated, species‐poor community to a more species‐rich community in the dry dunes, but did not change community type in dune slacks. However, at the species level, grazing enhanced the abundance of some desired perennial, annual and biennial species, graminoids and bryophytes in both habitat types. The increased frequency of positive indicator species for habitat condition suggests that grazing was beneficial for species of conservation interest. Ellenberg nitrogen (N) values decreased after grazing in dry habitats but showed no long‐term change independent of grazing, suggesting no increase in site fertility over the study period. Surprisingly, light (L) values also decreased in the dry dunes after grazing. Conclusions: Long‐term grazing management can play an important role for the conservation of dune communities and associated species. Because of its positive effects on species diversity, plant communities and habitat condition in sand dunes, livestock grazing is recommended for conservation management.     

Patch quality and connectivity influence spatial dynamics in a dune wolfspider

The spatial population dynamics of the wolfspider Pardosa monticola, inhabiting patchily distributed grasslands in the Flemish coastal dunes of Belgium and Northern France were investigated with incidence function models using field survey data from 1998 and 2000. Vegetation height and patch size were related to habitat quality. Mark-recapture experiments revealed maximum cursorial dispersal distances of 280 m for moss dunes and 185 m for higher dune grassland. Higher shrub vegetation appeared to be dispersal barriers. These habitat-dependant cursorial distances and the theoretically estimated ballooning distance were included with patch distances into a connectivity index for both dispersal modes. Forward multiple regression indicated that patch occurrence was influenced by habitat quality and ballooning connectivity. Habitat quality and cursorial connectivity explained patterns in short-term colonisation. Extinction appeared to be stochastic and not related to habitat quality and connectivity. Genetic differentiation and variability was low. The discrepancy between the estimated low dispersal capacity and the indirect estimate of gene flow ( F(ST)) indicates that historical population dynamics and/or historical ballooning dispersal influence the genetic structure in this species.     

Bryophytes in Mediterranean coastal dunes: ecological strategies and distribution along the vegetation zonation

The bryophytic flora has been widely studied in Atlantic coastal dune environments while bryological literature addressing the Mediterranean bryophytes is still scarce and fragmented, although they are an important component of this area, especially considering their ability to colonize and stabilize sandy soils. The aims of this study were: (i) to analyze the taxonomy, chorology and ecology of coastal dunes bryoflora; (ii) to analyze distribution patterns of bryophytic species along a coastal dune vegetation zonation. We used as coastal dune model system a particularly well-preserved study area of the Sardinian coast (Italy), which includes several habitats listed in European Habitats Directive 92/43/EEC. We conducted a general survey of the area and a random sampling. A total of 20 bryophytic species were detected most of which were acrocarpous mosses and, to a lesser extent, pleurocarpous and liverworts. Most bryophytes are pioneer colonizers, reflecting their adaptation to environments with high levels of stress/disturbance conditions such as those of coastal dunes. Finally, in this study we described for the first time how bryophytic species are distributed along a Mediterranean coastal dune vegetation zonation, focusing also on their chorological and ecological characteristics and on how they seem to follow the sea-land gradient, probably in response to different environmental conditions developing along this gradient.     

The impact of human pressure on landscape patterns and plant species richness in Mediterranean coastal dunes

We analyze the spatial patterns of natural dune cover patches and their plant richness, comparing coastal sites with different levels of human pressure in central Italy. We created a detailed land cover map of dune sites. The spatial pattern of natural dune cover types was characterized by computing a set of patch-based metrics. To quantify patch plant richness, we used 16 m² vegetation plots, randomly distributed on coastal dune cover types. For each patch, the richness of the entire pool of species and of three guilds (i.e., typical dune, ruderal, and alien species) was considered. We compared different levels of human pressure on coastal dunes focusing on pattern metrics and floristic information by using the nonparametric Kruskal–Wallis test. In sites with high human pressure, we have observed a general simplification in the natural dune spatial pattern and a decline of plant richness but with a specific response for each cover type. Alien and ruderal species presented low richness in all patches. In coastal dunes, the harsh ecological conditions and the strong sea–inland gradient shape the distribution of human activities and control the number of ruderal species. The approach effectively describes fragmentation and biodiversity in dune ecosystems.     

Green beach vegetation dynamics explained by embryo dune development

Sandy coastlines are dynamic environments with potential for biodiverse habitats, such as green beaches. Green beach vegetation can develop on nutrient-poor beaches landward from embryo dunes. It is characterised by low-dynamic coastal wetland habitat such as salt marshes and dune slacks. It has been hypothesised that the establishment of green beach vegetation is facilitated by the shelter provided by embryo dunes, however evidence is lacking.We explored the importance of geomorphology and soil conditions on the species richness and turnover of green beach vegetation over a time period of 10 years. We recorded 107 plots along 11 transects over a gradient from beach to dune on the island of Schiermonnikoog, the Netherlands. We characterised transect geomorphology at transect level and soil conditions and vegetation at plot level in 2006 and 2016.We found that the green beach vegetation was highly dynamic, total plant cover increased by 62% within 10 years. In 2006 beach width was an important factor in explaining species richness, with the highest number of species occurring on narrow beaches with a large volume of embryo dunes. In 2016, species richness was positively associated with the build-up of organic matter. Overall species richness declined relative to 2006 and was accompanied by an increase in elevation due to sand burial and the expansion of embryo dune volume.Our data suggests that geomorphology influenced the vegetation indirectly by affecting sand burial rate. Plant species richness declined less at sheltered conditions where sand burial was limited, allowing the build-up of organic matter. This indicates a time-dependent relationship between the development of embryo dunes and plant species richness: embryo dunes can be a source of shelter, thus increasing species richness, but can compete for space over time, lowering species richness again. Our results are relevant for engineering and management of biodiverse sandy shores.