Unusual karyotype diversity in the European spiders of the genus Atypus (Araneae: Atypidae)

Compared with araneomorph spiders, karyotypes of the spider infraorder Mygalomorphae are nearly unknown. In this study we investigated karyotypes of European species of the genus Atypus (Atypidae). The male karyotype of A. muralis and A. piceus comprises 41 chromosomes, whereas female complements contain 42 chromosomes. On the other hand, both sexes of A. affinis possess 14 chromosomes only. It is the lowest diploid number found in mygalomorph spiders so far. Furthermore, obtained data suggest X0 sex chromosome system in A. piceus, A. muralis and neo-XY system in A. affinis. Karyotypes of all three Atypus species are composed of biarmed chromosomes only. Thus they differ significantly from the karyotype of A. karschi, the only other species of this genus studied so far. Its karyotype was reported to be composed of acrocentric chromosomes and possesses X(1)X(2)0 sex chromosome system. All this shows that unlike in most genera of araneomorph spiders, mygalomorphs of the genus Atypus exhibit unusual diversity in the number, morphology of chromosomes, and the sex chromosome system. Considering high number of chromosomes being plesiomorphic character in spiders, then karyotypes of A. muralis and A. piceus represent ancestral situation and that of A. affinis being derived by multiple fusions. Karyotype differences in Atypus correspond with morphological differences, namely the number of segments of the posterior lateral spinnerets. Thus in contrast to published hypothesis, the 3-segmented posterior lateral spinnerets of A. affinis should present a derived state.     

Conservation status of the only representative of infraorder Mygalomorphae (Araneae) in cultivated regions of Central Europe

Purse-web spiders Atypus spp. (Araneae: Atypidae) are the only representatives of the archaic spider infraorder Mygalomorphae in the Central European fauna. These burrow-dwelling spiders are red-listed in all Central European countries. Atypus muralis occurs on relics of continental steppes, Atypus piceus on dry submediterranean grassland and dry herb-rich forest fringes, while Atypus affinis inhabits sparse dwarf dry forests, usually those dominated by oaks. Long-term sustainability of their habitats in the relatively humid Central European climate was maintained by grazing. Since the 1950s, pasturing was terminated across most of Central Europe, which led to immediate overgrowth of pastures by aggressive grasses and woody plants. Grazing was later reinstated in some reserves, but in cases of poorly controlled grazing intensity, footsteps had immediate detrimental effects on the residual populations of purse-web spiders and other invertebrates. Therefore, grazing intensity needs to be regulated, considering its double-edge effects on the biota of xerothermophilic habitats. Nests of European Atypus species have never been found in habitats subject to recent or even regular distribution. Therefore, they may serve as reliable indicators of well preserved xerothermic habitats. Their nests are easy to detect, and the European Atypus spiders can be utilized as model species for monitoring impacts of conservation management. Monitoring population dynamics of Atypus requires establishing permanent study plots of uniform size, documentation of vegetation by phytocenological relevé, counting their density (number of purse-webs found in the plot) and age structure (diameter of purse webs).     

Missing the target? A critical view on butterfly conservation efforts on calcareous grasslands in south-western Germany

Butterflies are strongly declining on grassland habitats of Central Europe. Therefore, the success of conservation measures on high quality grassland habitats is controversially discussed. We compared the changes in butterfly diversity and community structure on six managed calcareous grasslands with eight unmanaged vineyard fallows. We obtained strong losses of species diversity and remarkable shifts of community compositions on both habitat types. However, the changes on vineyard fallows were only slightly more severe but more stochastic than on the calcareous grasslands. The shifts in community composition with respect to functional species traits were rather similar between the two different grassland types so that complex butterfly communities evolved into generalist-dominated ones. Connectivity was higher among vineyard fallows than among calcareous grasslands. Consequently, conservation measures on calcareous grasslands only partly achieved their goal to maintain the high species diversity and functional complexity still observed in the 1970s. The negative impacts of eutrophication and monotonisation of the landscape as well as climate change are affecting all habitats, independently from management concepts. Therefore, management on conservation sites can buffer against these effects, but is not sufficient for a full compensation.     

Habitat associations of epigeal spiders in upland calcareous grassland landscapes: the importance for conservation

Upland calcareous grassland landscapes are typically comprised of a matrix of calcareous grassland, acid grassland and limestone heath plant communities. This matrix of habitats is produced by a combination of underlying geology, climate and management. These landscapes are typically managed through grazing, with management targeted to maintain particular plant communities in the calcareous grassland habitat, whilst patches of acid grassland and limestone heath are not targeted by conservation management. The biodiversity value of acid grassland and limestone heath patches within the calcareous grassland matrix are unknown. This study provides the first assessment of their biodiversity value by examining aspects of epigeal spider diversity supported by these non-target habitat patches in comparison to calcareous grassland. Spiders were sampled in each habitat from April to August 2014 using pitfall traps across three upland regions in Great Britain. Spider species assemblages were distinct between limestone heath and both grassland types. Distinction in species assemblages are likely due to differences in vegetation structure and microclimate, e.g., humidity, degree of shade. Each habitat type supported several rare species (e.g., Jacksonella falconeri, Agyneta subtilis) revealing the contribution to spider fauna. The distinct spider species assemblage and presence of rare species in limestone heath patches demonstrate their importance in the upland calcareous grassland matrix. This study highlights the value of monitoring biodiversity in non-target habitats within a habitat matrix alongside those that are actively targeted by management.     

Conservation genetics of peripheral populations of the mygalomorph spider Atypus affinis (Atypidae) in northern Europe

Genetic diversity and differentiation were studied in peripheral populations of the recently rediscovered purse-web spider Atypus affinis in Denmark and Sweden using allozyme electrophoresis. Because of the very narrow environmental niche exploited by this species, only a limited number of potential habitats are available. Furthermore, fragmentation has reduced the number and size of potential habitats within the last century. The level of genetic diversity in A. affinis was intermediate compared with other nonsocial spiders, but low compared with invertebrates in general. Significant genetic differentiation was found within distances of only 1-10 km with FST estimates ranging from 0.020 to 0.075. Within distances of 30-60 km FST ranged from 0.081 to 0.312. Hierarchical FST revealed that genetic variability was partitioned at 89.8% within populations, 9.5% among populations within regions and only 0.7% among the four main regions in Denmark and Sweden. Comparing the result of the genetic analysis with the life history of A. affinis, it is concluded that the level of successful dispersal is low, and that the species has not recently reinvaded northern Europe but prevailed undiscovered for several decades. Finally, it is suggested that the genetic scenario found for A. affinis might represent that for a wide range of other arthropods with similar life history characteristics.     

Exceptionally poor land snail fauna of central Yakutia (NE Russia): climatic and habitat determinants of species richness

There is an ongoing debate on the causes of the latitudinal diversity gradient, but diversity decline towards high latitudes is poorly documented for many invertebrate taxa. Therefore, we sampled land snail assemblages at 79 sites and in various habitat types in central Yakutia, a region with extremely continental, cool and dry climate. We tested whether habitats lacking suitable shelters for winter survival harbour less species than those with vegetation cover that softens climatic extremes. Both local species diversity and regional species diversity were extremely low: 13 species were recorded in total with an average of 1.4 species per site. While the majority of grassland sites were without snails (26 of 34 sites), forest sites supported at least one snail species in most cases (38 of 45 sites). Within grasslands, snail occurrences were associated with a higher herb-layer biomass. Numbers of snail species correlated with the amount of available calcium only in forests, in which species accumulation towards more favourable habitats was possible due to softening of climate harshness. As minute snails are known to be effective passive dispersers and the study area was not glaciated during the last glacial stage, there was certainly enough time for colonization of all favourable habitats. Our results suggest climatically driven limitations of both local and regional land snail diversity in central Yakutia. We conclude that the hypothesis of climate harshness remains the most probable explanation of a sharp drop in land snail diversity in high-latitude areas with cold climate.     

Paleoenvironmental reconstruction of middle Miocene paleosols bearing Kenyapithecus and Victoriapithecus, Nyakach Formation, southwestern Kenya

Paleosols in the middle Miocene (15 Ma) Nyakach Formation at Kaimogool, near Sondu, southwestern Kenya have yielded specimens of the early cercopithecoid Victoriapithecus macinnesi and the early kenyapithecine Kenyapithecus africanus, and can be used as evidence for the environmental mosaic occupied by these primates. Five distinct types of paleosols (pedotypes) are recognized in the Nyakach Formation section at Kaimogool South. The most common paleosols are reddish brown, silty calcareous profiles with blocky structure and large root traces (Ratong pedotype) which are interpreted as soils of well-drained, dry bushland or thicket (nyika). Weakly developed paleosols associated with paleochannels (Dhero pedotype) represent wooded grassland early in the ecological succession from streamside flooding. One of these paleosols has yielded a fossil flora of grasses and small-leaved dicots like those of modern semi-arid wooded grassland. Crumb structured, calcareous paleosols with iron-manganese nodules (Yom pedotype) are interpreted to represent seasonally waterlogged, wooded grassland (dambo or vlei). Thick, red clayey, calcareous paleosols with blocky ped structure and large root traces (Tut pedotype) are interpreted as soils of well-drained dry woodland. Other blocky-structured, gray to brown calcareous paleosols with iron-manganese nodules (Chido pedotype) are interpreted as soils of seasonally waterlogged, riparian dry woodland. Fossil soils, plants and gastropods are evidence of an unusually dry (300-500 mm mean annual precipitation) habitat for apes, consisting of a vegetational mosaic dominated by dry woodland, bushland and thickets with few areas of seasonally waterlogged grassland. Fossils of V. macinnesi are rare from Nyakach, but were found in paleosols representative of bushland and thicket habitats (Ratong). Fossils of the ape K. africanus were found within paleosols indicative of dry woodland (Tut). Other paleosol types representative of seasonally dry dambo grassland (Yom), colonizing grassland (Dhero) or riparian woodland (Chido) are also represented, but have not yet produced primate fossils.     

From sediment to soil: floodplain phosphorus transformations at the Danube River

Substantial transformations of biogeochemical phosphorus (P) fractions can occur within a few hundreds of years under humid and tropical conditions; however, slower changes are expected under dry and temperate climate. The objective of this study was to infer P transformations over time by comparing suspended sediments in the Danube River to floodplain soils developed from such sediments over different time periods in the continental climate of Central Europe. We analyzed suspended sediments from 20 flood events between 1990 and 2006, and floodplain soils from seven sites along a chronosequence covering about 500 yrs. The studied flood sediments had similar characteristics over the 16-yr observation period. Total phosphorus (TP) averaged 732 mg kg⁻¹, and biogeochemical fractionation yielded important primary mineral contributions (apatite phosphorus, AP, ~80% of TP). The TP concentrations of the floodplain soils were in the range of the Danube sediments and showed little variation along the chronosequence. However, the distribution of P among biogeochemical fractions changed considerably in less than 500 yrs of soil development. The youngest soils (<20 yrs) were dominated by AP, as was observed for the Danube sediments. In less than 250 yrs of pedogenesis, AP markedly decreased and organic phosphorus (OP) increased, and in less than 500 yrs, OP reached AP levels. This shows that while P biogeochemistry in very young floodplain soils is strongly related to the river sediments, significant transformations can occur in less than 250 yrs of soil development in the dry and temperate climate of Central Europe.     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.     

Explaining Rarity of the Dry Grassland Perennial Astragalus exscapus

In Central Europe several plant species of dry grasslands are particularly rare. Here I investigate whether habitat requirements, reproduction, and dispersal potential can contribute to the rarity of Astragalus exscapus (Fabaceae) growing in dry grassland habitats in dry regions of Europe. In addition, I question whether historic events might have contributed to the present-day rarity of A. exscapus. To assess habitat requirements of A. exscapus, vegetation composition and soil characteristics were studied in 37 populations in central Germany. Production and dispersal potential of seeds were investigated in 10 populations, and germination and recruitment were assessed in experimental plots in three populations. Vegetation of the habitats included most dry grassland community types occurring in the central German dry region indicating a broad ecological niche of the species within dry subcontinental grasslands. Soil characteristics of the habitats also spanned a wide range. Seed production was moderate. 98% of the seeds sown in the laboratory germinated whereas under natural conditions 20% of the seeds developed seedlings. Half of these seedlings survived for one year but only 4.5% for two years. 90% of the seeds were dispersed less than 50 cm distance indicating a low dispersal potential. I conclude that A. exscapus is mainly limited in dispersal but recruitment limitation might also be important in explaining its rarity. Furthermore, former climate change and postglacial reforestation of the area very likely contribute to the rarity of A. exscapus.     

Spontaneous regeneration of dry grasslands on set-aside fields

Dry grasslands are among the most threatened plant communities of Central Europe. We explore the time scale of spontaneous regeneration of dry grasslands on abandoned fields in an area of Central Europe, where also ancient grassland communities occur (Saxony-Anhalt, Germany). On three permanent plots with shallow soils we monitored during 10 years change of species composition and analysed whether spontaneous succession leads to assemblages similar to the ancient dry grassland communities in the direct surroundings. We found that dry grassland species are able to invade the permanent plots and during the 10 years of succession the number of dry grassland species increased. But even after 10 years there was a clear difference between ancient dry grassland communities and the assemblages on the permanent plots. Our findings suggest two important conclusions: First, spontaneous succession on abandoned fields is a cheap possibility for the conservation of some dry grassland species, at least on shallow soils. Second, the time scale of the regeneration process, however, is rather long. Hence, conservation of remnants of ancient grassland communities needs special attention.     

Ecological indicators of Tuber aestivum habitats in temperate European beech forests

Although Tuber aestivum is widely distributed across Europe, little is known about its biology and ecology. We assess the vegetation composition of wild T. aestivum sites and use this information to characterise suitable habitats in Fagus sylvatica, Carpinus betulus and Ostrya carpinifolia (FCO) forests. Plant species composition at 16 naturally colonised T. aestivum sites in southwestern Germany and Switzerland was compared with that of 232 reference sites in Swiss FCO forest communities. The vegetation composition of truffle sites exhibits a high proportion of herb and shrub species that are indicators of a warm and dry climate, alkaline soil, and open tree canopy and thus typically occur in thermophilous FCO communities on calcareous bedrock. We conclude that T. aestivum is associated with a set of key species in addition to known host species. Based on their floristic composition, we identified FCO forest communities with high and low suitability for T. aestivum.     

The effects of habitat fragmentation on niche requirements of the marsh fritillary, Euphydryas aurinia, (Rottemburg, 1775) on calcareous grasslands in southern UK

The marsh fritillary, Euphydryas aurinia, has declined greatly in distribution across its range within Europe, resulting in its designation as a protected species under Annex II of the 1979 Bern Convention and the EC Habitats and Species Directive. The decline has been linked to a marked reduction in the extent of suitable calcareous and wet grassland habitats, habitats which have been lost through conversion of land to agriculture or urban areas, or reduced in quality due to inappropriate management. The UK is now one of the major strongholds for this butterfly in Europe, although much of the remaining habitat is small, isolated and highly fragmented. E. aurinia populations fluctuate greatly due to the combined effects of biotic (e.g. parasitoids) and abiotic (e.g. climate change) factors. We quantified the habitat associations of larval webs of E. aurinia on fragmented versus extensive (unfragmented) calcareous grassland habitat in southern England to test the hypothesis that habitat requirements of E. aurinia are more constrained within fragmented landscapes. Within both fragmented and unfragmented landscapes the quality and quantity of its main host plant in the UK, Succisa pratensis, was positively related to numbers of E. aurinia larval webs found. The sward height was also important at predicting the distribution of larval webs in both landscapes, although the heights were greater within sites in the unfragmented (≈20 cm) compared to fragmented (≈15 cm) landscape. We also found significant effects of elevation and the cover of bare ground on numbers of larval webs. Elevation was strongly correlated with the availability of host plant, whilst bare ground was only significant on sites within the fragmented landscape, showing a negative relationship with number of larval webs. Our results further emphasise the importance of not only maintaining the habitat quality of extant calcareous grassland sites for E. aurinia in the UK, but also increasing the size and connectivity of these sites to increase the chances and rate of (re)colonisation of unoccupied but suitable habitat. In addition, we show that the habitat requirements of E. aurinia on sites in a large unfragmented landscape may be less specific and thus require less extensive management than that required to create optimal conditions necessary at smaller, more isolated sites in fragmented landscapes.     

Predicting microscale shifts in the distribution of the butterfly Plebejus argus at the northern edge of its range

Species are often observed to occur in restricted patches of particularly warm microclimate at their high latitude/altitude geographic range margin. In these areas, global warming is expected to cause small‐scale expansion of the occupied area, but most previous studies of range expansion have used very coarse scale data. Using high resolution microclimate models together with detailed field surveys, we tested whether the butterfly Plebejus argus, occurring on limestone grassland in north Wales, was responding as might be expected due to climate change in the last 30–40 yr. The abundance of adult Plebejus argus at 100 m resolution in 2011 was strongly affected by elevation and near‐ground temperatures in May. A statistical model including microclimate, fitted to 2011 data, was successful (67% correct) at hindcasting the occurrence of Plebejus argus in 1983 when the average May air temperature was 1.4°C cooler. However, the model was less accurate at hindcasting occurrences in 1972 (50% correct). Given the distribution of micro‐sites in this landscape, we predict that further warming of approximately 1°C would make the majority of sites highly microclimatically suitable for this species. There are a growing number of long‐term studies of range change, and investigations into the mechanisms driving them, but still surprisingly few that explicitly make and test predictions with independent data. Our tests are a valuable example of how accurate predictions of distribution change can be, but also of the inevitable uncertainties. Improved understanding of how well models predict will be very important to plan robust climate change adaptation measures.     

Soil nitrogen and carbon heterogeneity in woodlands and grasslands: contrasts between temperate and tropical regions

Aim  Soil resource heterogeneity is linked to several ecological processes including invasion of woody species into grasslands. Studies from the temperate zone have demonstrated greater soil heterogeneity beneath woody vegetation than beneath grasslands. Woody species have a more widespread and coarser root system than herbaceous species, and may have a competitive advantage in relatively heterogeneous soils. We tested the global generality of greater soil heterogeneity beneath woody vegetation.
Location  Global.
Methods  We used data from published literature for soil nitrogen and carbon heterogeneity from paired woodland and grassland sites around the world.
Results  Woodland and grassland soil heterogeneities from paired observations were strongly correlated. There was, however, significant geographical variability in the relationship. Soils were more heterogeneous in woodlands than grasslands in temperate areas, but the opposite was true for tropical habitats. Grassland soils were more heterogeneous at lower than higher latitudes. Woodland soil heterogeneity did not vary with latitude.
Main conclusions  The previously described high soil heterogeneity in woody vegetation compared to grasslands holds only for temperate regions. Consequently, the relationship between soil resource heterogeneity and vegetation type is dependent on the study region. Macroecological studies should test the generality of relationships between soil and vegetation at the global scale.     

Habitats of relict terrestrial snails in southern Siberia: lessons for the reconstruction of palaeoenvironments of full‐glacial Europe

Aim Shells of fossil molluscs are important for palaeoecological reconstructions. However, the habitat requirements of snail species typical of central European full‐glacial loess sediments are poorly known because most of them became very rare or extinct in Europe. The recent discovery of an almost complete extant assemblage of such species in mountainous regions of central Asia enables more precise characterization of their habitats, which may significantly improve reconstructions of Pleistocene environments. Location Altai Mountains, Russia. Methods Terrestrial snail assemblages, vegetation composition and selected environmental variables were recorded at 118 sites along a gradient of climatic continentality in the Russian Altai. Habitat characteristics of sites where species typical of the full‐glacial period occurred were described using a classification tree. Results Seven of the eight species that are typical of central European full‐glacial loess sediments were found in the study area. They were confined to cool areas with January mean temperatures below ?17 °C, but occurred mainly in sheltered habitats with a warmer microclimate, such as scrub or open woodland. Pupilla loessica and Vallonia tenuilabris had the broadest habitat range, occurring from woodland to dry steppe. Unexpectedly, Columella columella, Pupilla alpicola, Vertigo genesii, V. parcedentata and V. pseudosubstriata were found mainly in wooded fens and shrubby tundra rather than in open steppe. Most of these seven species were recorded in base‐rich wooded fens. Very dry open steppe habitats usually supported no snails. Main conclusions Habitat ranges of the studied snails in the Altai indicate that the full‐glacial landscapes of central European lowlands that harboured these species were not completely dominated by open and dry loess steppe. Most probably they contained a significant component of shrubby vegetation, patches of wet habitats, and probably also areas of woodland at sites with a favourable mesoclimate.     

Alien and native plants show contrasting responses to climate and land use in Europe

Aim We tested whether the distribution and cover of alien plant species in Europe was related to human disturbance and microclimate. Location Surveys were conducted at 13 sites across Europe, each containing a pair of landscapes with different land‐use intensities. Methods Sampling locations were chosen based on land use and microclimate at two scales: land use was characterized at the patch and landscape scale; climate was expressed as regional and local temperature. The slope of each sample location was derived from a digital elevation model. Cover of plant species was measured using point counts and analysed using mixed effect models. Species were classified as native, archaeophytes and neophytes (pre‐ versus post‐ad 1500 immigrants). Due to the zero inflation observed in the alien groups, their cover was analysed conditional on their presence. Results Anthropogenic disturbance was a significant explanatory variable, increasing the presence and cover of alien species and decreasing the cover of native species. Alien presence was increased in sites under agricultural management, while their cover responded to land use at both local and landscape scales (and to their interaction), such that only natural habitats in semi‐natural landscapes had low alien cover. Microclimate was important for neophytes, with presence concentrated around mesic conditions. Slope was relevant for archaeophytes and native species, suppressing the former group and promoting the latter one. Main conclusions We found that, at the European scale, the distribution of alien plants is related to anthropogenic disturbance more than to microclimatic differences. The presence of neophytes, however, was influenced by climate at local and regional scales, with the highest incidence under mesic conditions. The different patterns observed for the presence and cover of alien species suggest different mechanisms acting during their establishment and spread. They also suggest that to counteract the expansion of alien species natural habitats may need to be maintained at landscape scales.     

Is the lack of leguminous savanna trees in grasslands of South Africa related to nutritional constraints?

As with many grasslands globally, the Highveld grasslands of South Africa are tree-less, despite having a climate that can support tree growth. Models predict that fire maintains these grasslands. The question arises as to why fire-tolerant savanna trees do not survive in these ecosystems? Savanna tree survival in mesic areas is restricted by demographic bottlenecks, specifically limitations to sapling-escape from fire. It was hypothesised that ancient highly leached soils from grassland areas would prevent saplings from growing fast enough to escape the fire-trap. Growth rates of savanna tree seedlings (Acacia karroo Hayne and Acacia sieberiana Burtt Davy) were measured in a common garden experiment using soils from ten sites collected along a savanna-grassland continuum. Soils from grassland sites were relatively nutrient-poor compared to those from savannas with lower pH, and associated cations. A. sieberiana growth rates responded to pH and these nutrients, whereas A. karroo growth was less strongly linked to specific nutrients. Even so, both species accumulated more biomass when grown in soils from savanna sites compared to grassland sites. An exception was a low elevation low nutrient savanna site that resulted in poor growth, yet sustains high tree biomass in situ. Differences between growth in grassland and savanna soils were small. They may contribute to, but are unlikely to explain, the treeless nature of these grasslands.     

Plant communities and conservation of Botrychium-rich grasslands in the Bitcherland (Northern Vosges Biosphere Reserve, France)

Two species of Botrychium are present in the Bitcherland grasslands, B. lunaria and B. matricariifolium, and both are protected in this territory. They occur in five different continental grassland communities, (i) a dune grassland (Diantho-Armerietum), (ii) a mesotrophic sandy grassland (Viscario-Avenetum), composed of three sub-units, therophitic ( V.A. airetosum), meso-eutrophic (V.A. saxifragetosum) and acidic (V.A. violetosum caninae), and (iii) an oligotrophic sandy grassland (Aveno-Genistetum). Both species of Botrychium appear in the three sub-units of Viscario-Avenetum, but only one in Diantho-Armerietum (B. matricariifolium) and in Aveno-Genistetum (B. lunaria). These Botrychium species are present in quite similar habitats in other lowland areas in Central Europe. The conservation of these threatened species, sensitive to plant competition, requires the maintenance or restoration of extensive agricultural management with late cutting and no fertilization.     

Disturbance,microclimate and historical habitat connectivity determine the population performance of the threatened grassland specialist Carex caryophyllea in remnant grasslands

Many grassland specialist plant populations in Europe have become restricted to remnant habitats. The performance of these populations depends on both species‐specific traits and local and landscape level aspects of habitat quality. Understanding which specific local or landscape level conditions determine the performance of grassland species populations in remnant habitats would help design the restoration of the habitats and to detect the conditions that favour the long‐term persistence of grassland species in them. Such information is especially needed in urbanised landscapes, where remnant habitats engulfed by urban land use types may experience increased erosion, higher temperatures and invasion by alien species. This study investigates the population performance determinants of Carex caryophyllea (VU), a grassland specialist, in 43 remnant grasslands in an urban‐rural gradient in Finland. The population performance was assessed with metrics of persistence, establishment and reproduction, and related to environmental conditions with generalized additive models and redundancy analysis. The most important positive determinants for the performance of C. caryophyllea populations were disturbance through management or ground erosion, a warm microclimate, large habitat area and high historical connectivity to suitable grassland habitats. Present connectivity to other C. caryophyllea populations had a weak and near‐significant positive relationship with population performance. Urbanisation of the surrounding landscape correlated with population performance as well, possibly due to the high historical cover of grasslands in presently urbanised landscapes. The results imply that the most effective restoration method of remnant C. caryophyllea populations would be reinstating disturbance regimes in overgrown habitats with warm microclimates close to suitable habitats and other existing populations, whether urban or rural. This would counteract the species future decline due to possible extinction debts and help the species persist in the study area in the long term.