Grey Dunes Research Articles

Conservation management of coastal dunes for Orthoptera has to consider oviposition and nymphal preferences

Dune and heathland ecosystems can harbour a multitude of specialized insect species. To assess the habitat quality of these ecosystems, the presence of specialized insect species may act as a useful indicator. The Orthoptera species Myrmeleotettix maculatus, Decticus verrucivorus, and Platycleis albopunctata present such umbrella species. Because knowledge of oviposition and nymphal habitats of these species is insufficient, we conducted a combined study consisting of an outdoor oviposition experiment and a field survey analyzing nymphal habitat preferences during summer 2009 on the Baltic island of Hiddensee, Germany. The oviposition experiment showed, that all three species mostly avoid oviposition under lichens (= mature grey dunes). M. maculatus preferred bare ground for oviposition, D. verrucivorus favoured both bare ground and mosses, and P. albopunctata laid most eggs into mosses. Young nymphs of both M. maculatus and P. albopunctata preferred initial grey dunes with a high proportion of bare ground and moss-rich grey dunes. Old nymphs were related to moss-rich and lichen-rich grey dunes with more dense vegetation. Based on our results, early seral stages of dune succession with bare ground and mosses as keystone structures are crucial for the conservation of the three studied umbrella species. Because old nymphs and adults additionally require more dense grey dune vegetation or adjacent heath stands, practical dune and heathland management measures should aim to maintain a mosaic-like pattern of different grey dune and dwarf-shrub vegetation stands.

Patterns in Species Diversity during Succession of Coastal Dunes

The humped-back relationship in species diversity during succession was tested using vegetation in a coastal sand dune system of the German Wadden Sea island of Spiekeroog. Permanent plots were studied over 15 years along a spatial chronosequence from young grey dunes to old brown dunes. Species diversity, succession rate, and environmental indicator value were used to evaluate multitemporal dynamics of the ecosystem. Long-term development of the dune vegetation was reflected along the chronosequence, whereas fluctuations of plant communities were analyzed by short-term changes of each permanent plot. The study confirmed the intermediate stress theory, whereas highest species diversity was reached at the transition zone of the environmental gradients. Total species richness showed humped-back relations along the xerosere. Hotspots in species diversity varied with the life form group. Highest richness of herbaceous plants was reached in semidynamic young grey dunes, whereas highest richness of bryophytes and lichens shifted due to competition effects to the driest parts of the dune gradient in older successional stages. Moreover, the study confirmed the biodiversity-stability theory, with highest ecosystem stability at highest diversity. More stable environments showed lower succession rates than dynamic, disturbed environments. Thus, during secondary succession with probably repeated disturbance, the succession rate was higher and no clear direction of vegetation development was found in comparison to primary succession. The succession rate in a primary series represented a unimodal relation with total species richness. Thus, species-poor vegetation often dominated by one species, such as heathlands, as well as species-rich vegetation, showed lowest succession rates. These communities probably are more stable due to a balanced species combination represented by higher evenness.

Response of carabid beetles (Coleoptera: Carabidae) and spiders (Araneae) to coastal heathland succession

We analyzed the impacts of succession and grass encroachment on carabid beetle and spider assemblages in a coastal heathland. Further, indicator species for different successional stages (grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, birch forest) were identified, and their relations to habitat parameters were analyzed. The study was conducted on the Baltic island of Hiddensee, Germany. Ground-dwelling arthropods were sampled using pitfall traps along a successional gradient containing five stages. Ordination by nonmetric multidimensional scaling (NMDS) and analysis of similarities (ANOSIM) revealed a clear separation of species composition among the successional stages. Both in carabid beetles and spiders, most indicator species were obtained in the youngest stage (grey dunes) and fewest in the intermediate stages (grassy heath, heath with shrubs). Also the proportion of endangered species was highest in grey dunes. Based on our results, conservation management of coastal heathlands should preserve a mosaic of different successional stages with a clear preference on younger stages (grey dunes and dwarf-shrub heath).

Impacts of succession and grass encroachment on heathland Orthoptera

For the conservation of biodiversity, heathlands present important ecosystems throughout Europe. The formerly widespread habitats are nowadays restricted to small and isolated remnants. Without land use heathland vegetation undergoes succession and, in addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Orthoptera in a coastal heathland on the Baltic island of Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong differences among the successional stages. Orthoptera species richness was highest in transitional stages. The high proportion of grasses offer favourable habitat conditions for graminivorous, chorto- and thamnobiont species. Orthoptera density was highest in grey dunes. Threatened and specialised species were restricted to the young stages grey dunes and dwarf-shrub heath. Hence, in order to maintain a high diversity of Orthoptera in heathlands, maintaining different successional stages is of critical importance and this should be integrated into heathland management practices.

Impact of the invasive moss Campylopus introflexus on carabid beetles (Coleoptera: Carabidae) and spiders (Araneae) in acidic coastal dunes at the southern Baltic Sea

Campylopus introflexus is an invasive moss in Europe and North America that is adapted to acidic and nutrient-poor sandy soils with sparse vegetation. In habitats like acidic coastal dunes (grey dunes) it can reach high densities, build dense carpets and modify habitat conditions. While the impact of the moss invasion on the vegetation is well analyzed, there is a lack of knowledge regarding possible effects on arthropods. In the present study we analyzed the impact of Campylopus introflexus on the ground-dwelling arthropods carabid beetles and spiders, as both taxa are known to be useful indicator taxa even on a small-scale level. In 2009 we compared species composition in a) invaded, moss-rich (C. introflexus) and b) native, lichen-rich (Cladonia spp.) acidic coastal dunes by using pitfall traps. A total of 1,846 carabid beetles (39 species) and 2,682 spiders (66 species) were caught. Species richness of both taxa and activity densities of spiders were lower in invaded sites. Species assemblages of carabids and spiders differed clearly between the two habitat types and single species were displaced by the moss encroachment. Phytophagous carabid beetles, web-building spiders and wolf spiders were more abundant in native, lichen-rich sites. Shifts in species composition can be explained by differences in the vegetation structure, microclimate conditions and most likely a reduced food supply in invaded sites. By forming dense carpets and covering large areas, the moss invasion strongly alters typical arthropod assemblages of endangered and protected (EU-directive) acidic coastal dunes.

Patch connectivity and sand dynamics affect dispersal‐related morphology of the blue‐winged grasshopperOedipoda caerulescensin coastal grey dunes

Abstract. 1. Anthropogenic land use leads to increasing habitat fragmentation and habitat loss for many species. Exchange of individuals between and recolonisation of habitat patches is essential for the stability and long‐term survival of populations in fragmented landscapes. Inter‐patch dispersal depends on the dispersal ability of species and on landscape structure.2. In this study, we analysed the influence of patch size, patch connectivity and sand dynamics on thorax‐ and wing length as proxies for dispersal ability of the blue‐winged grasshopperOedipoda caerulescensin coastal grey dunes.3. Our study revealed that sand dynamics and patch configuration had clear though sex‐specific effects on dispersal‐related morphology. Males were smaller and had shorter wings if patches became larger and less connected. In addition, both sexes were larger in habitat patches with high sand dynamics compared to those in patches with lower dynamics. Investments in wing length were only larger in connected populations when sand dynamics were low, indicating that both landscape and patch‐related environmental factors are of importance.4. The observed effect of landscape characteristics on morphology agrees with predictions from theoretical models on the evolution of dispersal in metapopulations. However, as effects appear to be less pronounced in females, it is possible that selection pressures act differently on both sexes. An alternative explanation, however, would be that the observed patterns could be due to sex‐specific phenotypic plasticity. The presented results add to the evidence that dispersal‐related morphology varies according to expectations on dispersal evolution in recently structured populations at small spatial scales.

The importance of habitat mosaics for Orthoptera (Caelifera and Ensifera) in dry heathlands

In summer 2008, the Orthopteran species Chorthippus biguttulus, Myrmeleotettix maculatus (Caelifera), Decticus verru- civorus and Platycleis albopunctata (Ensifera) were sampled in coastal heathland on the German Baltic Sea island of Hiddensee. The aim of this study was to assess differences in abundance of Orthoptera in three different habitats and determine the importance of habitat mosaics. Distribution patterns varied among species and total abundance of Orthoptera differed significantly among the three habitats. Due to species-specific habitat preferences the Caelifera were most abundant in grey dunes and the Ensifera in dwarf-shrub heath adjoining grey dunes. In conclusion, grey dunes are a suitable habitat for the Caelifera studied, while the Ensifera require a heathland mosaic consisting of both grey dunes and dwarf shrub vegetation.

Is pitfall trapping a valuable sampling method for grassland Orthoptera?

Common methods to assess diversity and abundance of Orthoptera are sweep netting, transect counts and box-quadrat sampling. Pitfall trapping, by contrast, is rarely used, and the value of this method is still being questioned. In 2008, we studied Orthoptera species richness and abundance in five vegetation types along a gradient of dune succession on the Baltic Sea island of Hiddensee (NE Germany) by comparing transect-count and pitfall-trapping data. Using transect counts, 12 species were detected in the study area. With pitfall traps, three chorto- and thamnobiont Ensifera species (C. dorsalis, M. roeselii and T. viridissima) were not caught at all, and it was only in low-growing and sparsely-vegetated grey dunes that all present species were detected. With pitfall traps, the proportion of present species recorded strongly declined with increasing height and density of the vegetation type. Assuming that transect counts are a good proxy for relative Orthoptera densities, densities ascertained by pitfall traps are strongly biased by vegetation structure and locomotive behaviour of the species. More than 80% of all individuals were caught in sparsely-vegetated grey dunes. Frequency patterns of the species also differed. Using pitfall traps, especially chortobiont species were significantly underrepresented. Qualitative and quantitative sampling of Orthoptera using pitfall traps seems only reasonable in habitats with low and sparse vegetation and a high proportion of geobiont species.

Interactive effects of rabbit grazing and environmental factors on plant species-richness on dunes of Norderney

On the German island Norderney rabbit grazing effects on vegetation, soil and microclimate were investigated with exclosures in different dune vegetation. Differences in species composition between grazed and ungrazed plots increased with grazing activity. A decline in growth height resulted in higher light availability and temperature on the grazed sites. One of the most visible effects was the decline in flowering plants. Most grazing effects varied between habitat types. Grey dunes were distinctly affected by rabbit grazing. In relation to the habitat type, life forms were differently affected: in closed vegetation, rabbit activities further open patches, resulting in more therophytes. In contrast, in open vegetation rabbit activities were too strong and following sand dynamics too high for many therophytes. In general, grazing diversified habitats, slightly supported the total species-richness, and enhanced the heterogeneity of species composition. Highest species-richness in total and of herbs was reached at intermediate levels of grazing pressure. The study is in line that highest species-diversity will be found at intermediate levels of disturbance and environmental stress and that grazing effects vary with the habitat type.

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.

Invasion of coastal dunes by the alien shrub Rosa rugosa is associated with roads, tracks and houses

Mapping the distribution of invasive alien plant species is significant for testing ecological hypotheses and for guiding effective management. Little is known about the distribution of invasive plants at landscape scale, and the factors controlling their dispersal and establishment are still poorly understood. This is the case for Rosa rugosa, an invasive shrub with negative effects on biodiversity in dune ecosystems of NW Europe. The aim of the study was to identify the factors which determine the distribution of R. rugosa in coastal dunes. In a large semi-natural dune area of NW Denmark (2364 ha) all patches of the species (1321, 1.3–59.1 m 2) were GPS mapped. Patch distribution was GIS analysed, based on aerial photographs and vegetation maps. The distance of R. rugosa from the most conspicuous landscape elements was compared with randomly placed reference points. The species had invaded 0.35% of the dune landscape, and it was present in all vegetation types and all parts of the study area. Rosa rugosa patches were slightly smaller in grey dunes than in white dunes and brown dunes, and some of the largest patches were found close to villages. The occurrence of the shrub was positively correlated with distance to the coastline, to roads, tracks and houses. We conclude that the distribution of R. rugosa is determined by both natural and anthropogenic factors. It is further discussed how these factors may control dispersal and establishment of the species, and how the results can be used for improved management of coastal dunes.

Distribution of earthworms and influence of soil properties across a successional sand dune ecosystem in NW England

An investigation of earthworms across a sand dune system in NW England examined species distribution and abundance with respect to soil physico-chemical conditions and management. Replicated 0.1 m 2 quadrats were examined every 50 m along 700 m transects across areas of vegetation succession and samples were then taken every 10 m where earthworms were first encountered. A translocation of Aporrectodea longa assessed the ability of this species to live in soils with a high sand content and laboratory work examined growth, maturation and survival of this species in soils with increasing proportions of sand (0–100%). Nine earthworm species were found on the dunes, but none in yellow dunes where organic matter (OM) content was <1%. Dendrobaena octaedra and Lumbricus rubellus were located 300 m from the strand line in grey dunes with an OM content of 3.9%. Allolobophora chlorotica and Lumbricus castaneus occurred within a wet dune slack at 340 m (OM content 11%). In areas of human disturbance (dune car park), A. longa, Aporrectodea caliginosa and Lumbricus terrestris were present, and in soil below pine trees L. castaneus and L. rubellus were present in litter. Laboratory growth of A. longa demonstrated significant ( p < 0.05) increases in growth (2–2.75 g) with 25 and 50% sand compared with 0, 75 and 100% over 24 weeks and a greater rate of maturation. Initial results show earthworm colonisation to be influenced by dune successional stage. Laboratory findings suggested A. longa could exist in higher sand content areas but experimental design needs development. The translocation was unsuccessful. Future investigations could examine soil properties more closely and undertake monitoring on site throughout the year.

Expansion of Rosa rugosa and Hippophaë rhamnoides in coastal grey dunes: Effects at different spatial scales

The study analysed the effects of shrub expansion on vegetation composition and plant species diversity in coastal grey dunes on the North Sea island Spiekeroog, comparing Rosa rugosa and Hippophaë rhamnoides. Species composition was recorded in plots of two spatial scales, 1 and 16 m 2, considering the full range of shrub cover from less than 10 to almost 100%. Although R. rugosa and H. rhamnoides established and spread in the same grey dune environment, the vegetation of the two shrubland types was much different. While the H. rhamnoides plots were relatively species-rich, characterised by remnant grey dune vegetation with many small, often annual, light-demanding species except in the densest shrubs, the R. rugosa plots were clearly species-poorer due to the loss of many typical grey dune species, including only few shade-tolerant taxa. The total number of species, the number of herbaceous species and of species typical for grasslands decreased with increasing cover of H. rhamnoides and R. rugosa at both spatial scales. For the number of shrubs and shrubland species, hardly any significant effects of shrub cover were observed in R. rugosa, while there were positive effects in H. rhamnoides. Both the Shannon index and evenness decreased with increasing cover of the two shrub species at both spatial scales. Here, the decline in species diversity was more improved in R. rugosa than in H. rhamnoides.

Trampling affects the distribution of specialised coastal dune arthropods

From a conservation point of view, species’ tolerances towards disturbance are often generalised and lack reference to spatial scales and underlying processes. In order to investigate how average typical species react to habitat fragmentation and disturbance, we adopted a multi-species approach to address occupancy patterns of five specialised dune arthropods (butterflies Hipparchia semele, Issoria lathonia; grasshopper Oedipoda caerulescens; spiders Alopecosa fabrilis, Xysticus sabulosus) in recently fragmented coastal dune habitats which are subjected to varying levels and modes of local disturbance, i.e. trampling by cattle or people. Occupancy patterns were assessed during two successive years in 133 grey dune fragments of the Flemish coastal dunes (Belgium, France). By treating species as a random factor in our models, emphasis was placed on generalisations rather than documenting species-specific patterns. Our study demonstrates that deteriorating effects of local disturbance on arthropod incidence cannot be interpreted independent of its landscape context, and appear to be more severe when patch area and connectivity decrease. When controlled for patch area and trampling intensity, the probability of species occupancy in poorly connected patches is higher under cattle trampling than under recreation. Incidences additionally decrease with increasing intensity of cattle trampling, but increases with trampling by tourists. This study provides evidence of mode- and landscape-dependent effects of local disturbance on species occupancy patterns. Most importantly, it demonstrates that trampling of sensitive dune fragments will lead to local and metapopulation extinction in landscapes where trampling occurs in a spatially autocorrelated way, but that the outcome (spatial patterns) varies in relation to disturbance mode, indicating that effects of disturbance cannot be generalised. Aus Sicht des Naturschutzes werden die Toleranzen von Arten gegenüber Störungen häufig verallgemeinert und lassen einen Bezug zu räumlichen Skalen und den zugrundeliegenden Prozessen vermissen. Um zu untersuchen, wie typische Durchschnittsarten auf Habitatfragmentierung und Störung reagieren, verwendeten wir einen Vielartenansatz, um die Nutzungsmuster von fünf spezialisierten Dünenarthropoden (Schmetterlinge: Hipparchia semele, Issoria lathonia; Heuschrecken: Oedipoda caerulescens; Spinnen: Alopecosa fabrilis, Xysticus sabulosus) in kürzlich fragmentierten Küstendünenhabitaten zu behandeln, die unterschiedlichen Arten und verschieden starken lokalen Störungen ausgesetzt waren, wie z. B. Vertritt durch Weidevieh oder Menschen. Die Nutzungsmuster wurden in zwei aufeinander folgenden Jahren in 133 Graudünenfragmenten der flämischen Küstendünen (Belgien, Frankreich) untersucht. Durch die Berücksichtigung der Arten als zufällige Faktoren legten wir den Schwerpunkt eher auf Generalisierung als auf die Dokumentation artspezifischer Muster. Unsere Untersuchung zeigt, dass die negativen Effekte lokaler Störungen auf die Arthropodenvorkommen nicht unabhängig vom Landschaftskontext interpretiert werden können, und dass sie ernster sind, wenn die Flächengröße und -vernetzung abnehmen. Wenn wir die Flächengröße und die Vertrittintensität kontrollieren, ist die Wahrscheinlichkeit des Artvorkommens in wenig vernetzten Flächen bei Beweidung größer als bei Freizeitnutzung. Die Vorkommen nehmen außerdem mit zunehmender Intensität des Weideviehvertritts ab, nehmen jedoch mit zunehmendem Vertritt durch Touristen zu. Diese Untersuchung liefert Belege für art- und landschaftsabhängige Effekte lokaler Störungen auf die Nutzungsmuster von Arten. Besonders wichtig ist, dass gezeigt wird, dass der Vertritt von sensiblen Dünenfragmenten zum Aussterben von lokalen und Metapopulationen in Landschaften führen wird, wenn der Vertritt in einer räumlich autokorrelierten Form stattfindet. Das Ergebnis (die räumlichen Muster) variiert jedoch in Relation zur Störungsart und weist darauf hin, dass die Effekte von Störungen nicht generalisiert werden können.

Analyse de la végétation des dunes littorales de Zemmouri (Boumerdès, Algérie)

Research has been carried out on the Eastern Algerian vegetation dunes coast area. These plant communities undergo climatic and anthropic degradations. Using multivariate analysis (correspondence analysis, ascending hierarchical clustering and canonical correspondence analysis), the spatial distribution of the species is studied by considering the various ecological gradients sites. The edaphic (salinity and granulometry) and topographic (distance with the sea) factors are the parameters which explain the best the floristic composition and the space organization of the phytocoenosis. Three sets of species are pointed out which correspond to the well defined units. Because of their low floristic richness, of 9 species on the embryonic dunes to 28 on the stabilized grey dunes, these natural floristic inheritances deserve protection measures and the restoration at the limits of grey dunes may be possible.

Using distribution patterns of five threatened invertebrates in a highly fragmented dune landscape to develop a multispecies conservation approach

We analysed the patterns of occupancy of five threatened invertebrates in a highly fragmented dynamic grey dune landscape. During two years (2003–2004), 133 dune patches between Nieuwpoort (Belgium) and Bray-Dunes (France) varying in area, connectivity, eolian sand dynamics and trampling disturbance were sampled for five focal species: two spiders ( Alopecosa fabrilis and Xysticus sabulosus), two butterflies ( Issoria lathonia and Hipparchia semele) and one grasshopper ( Oedipoda caerulescens). Overall diversity was highest in large and well connected patches that were characterised by high eolian sand dynamics and an intermediate trampling intensity. Patch occupancy differed greatly among species: all species significantly occurred more often in large and connected patches. High trampling intensity (by cattle and/or tourists) negatively affected the two ground dwelling spiders, but not the grasshopper or the butterfly species. High eolian sand dynamics positively affected the presence of the spider X. sabulosus, the grasshopper O. caerulescens and the butterfly H. semele, but had no significant effect on both other species. Colonisation was mainly explained by connectivity and never by patch area, while extinction events in H. semele were explained by small patch area. We discuss the implications of using a suite of focal species for management and restoration purposes in the highly fragmented dune area in Belgium and we promote the use of a multispecies approach for evaluating and monitoring conservation efforts in general.

A multivariate approach to plant community distribution in the coastal dune zonation of NW Denmark

A near-natural coastal dune system in NW Jutland, Denmark was investigated. The area contains a gradient in natural soil mobility with the highest values in dynamic white dunes dominated by Ammophila arenaria, and low mobility in fixed decalcified dune heath with Empetrum nigrum and Calluna vulgaris. A total of 267 plots was investigated and presence of mosses and rooted vascular plants recorded. Cluster analysis identified five communities: white dune, grey dune, outer dune heath, inner dune heath and humid dune heath. These communities were ordered along two main axes in a Detrended Correspondence Analysis (DCA). The relationship between DCA axes and several environmental variables, including soil characteristics and Ellenberg indicator values, were investigated by correlation. Two major complex gradients underlying compositional variation were identified; one short coenocline with an underlying steep gradient in habitat maturity, soil stability and nutrient-limited productivity, and one longer gradient in water-limited productivity and peat accumulation only applying to the older successional stages. Distance from the coast was an important correlate of several of the measured variables. Microclimatic indices, i.e. heat index and wind index, had limited value as descriptors of community patterns as they were not correlated with the DCA axes or with other environmental variables. However, within communities the microclimatic indices correlated with several explanatory variables. (Less)

Assessment of Damages from Recreational Activities on Coastal Dunes of the Southern Baltic Sea

A study of the changes in natural phytodiversity that can be linked to recreational activities on tertiary dunes (grey dunes) is presented. Ten neighboring areas on the Islands of Wolin (Poland) and Usedom (Germany), representing all major disturbance factors (holiday resorts, campgrounds, parking), were chosen with two appropriate reference areas from Usedom and Wolin Island (Poland) that are situated in remote parts of the coast and showed no permanent signs of human disturbance. The results of recreational activities, especially trampling, eutrophication, and the neighboring effects of nearby gardens, parks, or fallow land, were assumed to be the main factors influencing natural dunal phytodiversity. Mechanical stress from trampling could result in a decrease in phytodiversity, whereas rising nutrient levels (e.g., fecal deposits), as well as neighboring effects from urban development and a growing species pool, could lead to an increase in diversity, including the occurrence of generalist or ruderal species not typical of unmodified dunes. Hence, changes within natural dunal phytodiversity depend on character and intensity of recreational activities. Results show that the location of the beach largely influences visitor numbers and hence the level of disturbance. At holiday resorts, damage to dunes can be low, even though tourist numbers on the beach are very high, if the dunes can be overlooked from the main promenade. Visitors are reluctant to trespass onto the dunes if they are being watched by many people. Nevertheless, areas without major signs of human disturbance were mostly found on remote dunes that were only accessible by foot or bicycle. Application of established phytodiversity indices (e.g., H′) showed drawbacks in detecting the different types of disturbance; however, sensitivity is improved by a modified index (H′dune).

Sand dynamics in coastal dune landscapes constrain diversity and life‐history characteristics of spiders

SummaryIn fragmented landscapes where natural disturbance acts at the local level, community composition can be altered either by ‘species‐by‐species matching’, with community‐wide character displacement, or by ‘constrained species sorting’, whereby disturbance favours species with distinct ecological traits. It is necessary to understand such species shifts when aiming to detect bottlenecks associated with changing disturbance dynamics at different spatial levels. Such an understanding also provides insight into the underlying ecological mechanisms in operation and allows identification of species of conservation concern.This approach to understanding species’ shifts was applied to a study of spider diversity and life‐history variation in relation to sand dynamics in stabilized and dynamic grey dune landscapes located along the coasts of France, Belgium and the Netherlands.Local (α) diversity appeared to decrease with increasing local sand dynamics in both stable and dynamic landscapes. In dynamic dune landscapes, β‐diversity decreased significantly and approached regional (γ) diversity under local grey dune stabilization; in stabilized landscapes, β‐diversity was not affected by local sand dynamics.Comparative analyses of ecological traits revealed shifts in life‐history patterns, suggesting that patterns in local diversity resulted from species sorting. Species from fragments characterized by high sand dynamics showed narrower niche breadths, larger body sizes and longer generation times, while summer‐active species tended to become residential after sand stabilization. This provides evidence for constrained species sorting in which natural disturbance (through local sand dynamics) allows only species with distinct ecological traits to persist.Synthesis and applications. Species with larger body sizes, longer generation times and a higher degree of habitat specialization (i.e. related to dispersal ability) would be expected to be more vulnerable at dynamic sites and more prone to extinction than their counterparts from stabilized fragments. In particular, species with a burrowing lifestyle would be expected to go extinct locally as a result of increasing soil development and soil hardness. Shifts in species composition were found to be more pronounced in dynamic landscapes. Hence ensuring conservation of sand dynamics at a landscape level rather than at a local level is of prime importance when aiming to conserve typical psammophilous spider species within local assemblages in grey dune habitats.

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.