No abstract

34In the face of ongoing habitat fragmentation, species-area relationships (SARs) have 35 gained renewed interest and are increasingly used to set conservation priorities. An 36 important question is how large habitat areas need to be to optimize biodiversity 37 conservation. The relationship between area and species richness is explained by 38 colonization-extinction dynamics, whereby smaller sites harbour smaller populations, 39 which are more prone to extinction than the larger populations sustained by larger sites. 40These colonization-extinction dynamics are predicted to vary with trophic rank, habitat 41 affinity and dispersal ability of the species. However, empirical evidence for the effect of 42 these species characteristics on SARs remains inconclusive. 43In this study we used carabid beetle data from 58 calcareous grassland sites to 44 investigate how calcareous grassland area affects species richness and activity density for 45 3 species differing in trophic rank, habitat affinity and dispersal ability. In addition, we 46 investigated how SARs are affected by the availability of additional calcareous grassland 47 in the surrounding landscape. 48Our results demonstrate that beetle species richness and activity density increase 49 with calcareous grassland area for zoophagous species that are specialists for dry 50 grasslands and to a lesser extent for zoophagous habitat generalists. Phytophagous 51 species and zoophagous forest and wet grassland specialists were not affected by 52 calcareous grassland area. The dependence of species on large single sites increased with 53 decreasing dispersal ability for species already vulnerable to calcareous grassland area. 54Additional calcareous grassland in the landscape had a positive effect on local species 55 richness of both dry grassland specialists and generalists, but this effect was restricted to 56 a few hundred meters. 57Our results demonstrate that SARs are affected by trophic rank, habitat affinity 58 and dispersal ability. These species characteristics do not operate independently but 59 should be viewed in concert. In addition, species' responses depend on the landscape 60 context. Our study suggests that the impact of habitat area on trophic interactions may be 61 larger than previously anticipated. In small habitat fragments surrounded by a hostile 62 matrix, food chains may be strongly disrupted. This highlights the need to conserve 63 continuous calcareous grassland patches of at least several hectares in size. 64 65

Semi-natural grasslands are increasingly grazed by large herbivores for nature conservation purposes. For many insects such grazing is essential for the conservation of their habitat, but at the same time, populations decrease at high grazing intensity. We hypothesised that grazing management may cause increased butterfly mortality, especially for life-stages with low mobility, such as hibernating caterpillars. To test this, we measured the effect of sheep grazing on overwinter larval survival. We used the Glanville fritillary (Melitaea cinxia), which has gregarious caterpillars hibernating in silk nests, as a model species. Caterpillar nests were monitored throughout the hibernating period in calcareous grassland reserves with low and high intensity sheep grazing and in an ungrazed control treatment. After grazing, 64 % of the nests at the high intensity grazing treatment were damaged or missing, compared to 8 and 12 % at the ungrazed and low intensity grazing treatment, respectively. Nest volume and caterpillar survival were 50 % lower at the high intensity grazing treatment compared to both ungrazed and low intensity grazing treatments. Nest damage and increased mortality were mainly caused by incidental ingestion of the caterpillars by the sheep. It is likely that grazing similarly affects other invertebrates, depending on their location within the vegetation and their ability to actively avoid herbivores. This implies that the impact of grazing strongly depends on the timing of this management in relation to the phenology of the species. A greater focus on immature and inactive life-stages in conservation policy in general and particularly in action plans for endangered species is required to effectively preserve invertebrate diversity.

We present an overview of high nitrogen deposition effects on coastal dune grasslands in NW-Europe (H2130), especially concerning grass encroachment in calcareous and acidic Grey Dunes. The problem is larger than previously assumed, because critical loads are still too high, and extra N-input from the sea may amount to 10 kg ha −1 yr −1 . Grass encroachment clearly leads to loss of characteristic plant species, from approximately 16 in open dune grassland to 2 in tall-grass vegetation. Dune zones differ in grass encroachment, due to the chemical status of the soil. In calcareous and iron-rich dunes (Renodunal district), grass encroachment showed a clear gradient over the dune area. Grass encroachment is low in calcareous foredunes, due to low P-availability, and large grazers were not needed to counteract grass encroachment after 2001. In partly decalcified middle dunes, P-availability and grass encroachment are high due to dissolution of calcium phosphates, and grazing only partially helped to control this. In acidic, iron-rich hinterdunes, grass encroachment gradually increased between 1990 and 2014, possibly because P-availability increased with time due to increased soil organic matter content. In acidic, iron-poor dunes (Wadden district), grass encroachment is a large problem, because chemical P-fixation with Ca or Fe does not occur. Large grazers may however reduce tall-grass cover. High cumulative Ndeposition could theoretically lead to increased N-storage and N-mineralization in the soil. Mineralization indeed increased with N-deposition, but in 15 N experiments, most ammonium was converted to nitrate, and storage in soil organic matter was low. Soil N-storage is probably reduced by high nitrate leaching, which will favour dune restoration when N-deposition levels decrease.

The impact of atmospheric N-deposition on succession from open sand to dry, lichen-rich, short grassland, and tall grass vegetation dominated by Carex arenaria was surveyed in 19 coastal dune sites along the Baltic Sea. Coastal dunes with acid or slightly calcareous sand reacted differently to atmospheric wet deposition of 5-8 kg N ha -1 y -1 . Accelerated acidification, as well as increased growth of Carex and accumulation of organic matter, was observed only at acid sites with pH NaCl of the parent material below 6.0. At sites with slightly calcareous parent material, increased N-deposition had no effect. A trigger for grass encroachment seems to be high acidification in early successional stages to below pH NaCl 4.0. Metals like Al or Fe become freely available and may hamper intolerant species. At acid sites, N-mineralization increases with elevated N-deposition, which may further stimulate Carex arenaria. Due to high growth plasticity, efficient resource allocation and tolerance of high metal concentrations, C. arenaria is a superior competitor under these conditions and can start to dominate the dune system. Carex-dominated vegetation is species-poor. Even at the moderate Nloads in this study, foliose lichens, forbs and grasses were reduced in short grass vegetation at acid sites. Species indicating these first effects of atmospheric deposition on dry, lichen-rich, short grasslands are identified and recommendations for restoration of grass-encroached sites given.

Conservation management is expected to increase local biodiversity, but uniform management may lead to biotic homogenization and diversity losses at the regional scale. We evaluated the effects of renewed grazing and cutting management carried out across a whole region, on the diversity of plants and seven arthropod groups. Changes in occurrence over 17 years of intensive calcareous grassland management were analysed at the species level, which gave insight into the exact species contributing to regional homogenization or differentiation. Reponses were compared between species differing in habitat affinity, dispersal ability, food specialisation and trophic level. Local species richness increased over the sampling period for true bugs and millipedes, while carabid beetles and weevils declined in local species richness. Species richness remained unchanged for plants, woodlice, ants and spiders. Regional diversity and compositional variation generally followed local patterns. Diversity shifts were only to a limited extent explained by species' habitat affinity, dispersal ability, trophic level and food specialisation. We conclude that implementation of relatively uniform conservation management across a region did not lead to uniform changes in local species composition. This is an encouraging result for conservation managers, as it shows that there is not necessarily a conflict of interest between local and regional conservation goals. Our study also demonstrates that shifts in diversity patterns differ markedly between taxonomic groups. Single traits provide only limited understanding of these differences. This highlights the need for a wide taxonomic scope when evaluating conservation management and demonstrates the need to understand the mechanisms underlying occurrence shifts.

Encroachment of tall grasses and shrubs in coastal dunes has resulted in loss of vegetation heterogeneity. This is expected to have negative effects on animal diversity. To counteract encroachment and develop structural heterogeneity grazing is a widely used management practice. Here, we aim to functionally interpret changes in vegetation composition and configuration following grazing management on habitat suitability for sand lizards. Aerial photographs taken over a period of 16 years were used to quantify changes in vegetation composition. A GISbased method was developed to calculate habitat suitability for sand lizards in a spatially explicit manner, encompassing differences in vegetation structure and patch size. Coast Conserv (2012) 16:89-99 DOI 10.1007 different scale levels to fully capture the natural landscape dynamics.

At three coastal dune sites at the island of Hiddensee, north-east Germany, vegetation cover was mapped during 2002 and compared to vegetation surveys from the late 1980s and 1930s. Abiotic and biotic factors, which have been identified as being critical for coastal dunes in former studies such as disturbance, salt spray or nutrient availability, were measured. Grazing and land-use history were reviewed by literature and interviews. Tall graminoid communities, mainly Carex arenaria, are a common vegetation unit today. Development, distribution of these dominances and possible causes for its occurrence have not been analysed. Generally, older successional vegetation units increased and pioneer stages decreased from the 1930s until 2002. At the geologically youngest site, the southern dunes, grass encroachment by Carex arenaria was highest (ca. 50% cover in 2002), and age and density of trees lower than at the older, central dunes. Landuse changes such as decrease in grazing pressure, additional feeding of livestock, increase in coastal protection measures and subsequent decrease in shifting sands as well as varying availability of groundwater and amount of salt spray are relevant factors for vegetation changes in coastal dunes over the past 70 years. Site-specific land-use differences such as livestock density and land-use history have a stronger influence than atmospheric N-pollution on the vegetation composition of these acidic, coastal dunes under low to moderate N-deposition loads of 6-8 kg N ha −1 yr −1 .