Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Semi‐natural grasslands, among them thin‐soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red‐listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its size and with the proportion of dead branches, and was twice higher in plate alvars than in ryhk alvars. Also, the species composition in plate alvars differed from ryhk alvars by having 42 characteristic lichen species in plate alvars vs three indicators of ryhk alvars. The composition of lichens was significantly influenced by encroachment of alvars, e.g. by high juniper cover and shrub layer height, as well as by the proportion of dead branches and stem circumference of juniper. We conclude that the epiphytic lichen assemblies on junipers are threatened by grassland encroachment similarly to ground layer lichen assemblies. We suggest that some old and scencent junipers should be preserved during the restoration of alvar grasslands.     

Vegetation,environmental characteristics and ideas on the maintenance of alvars on the Bruce Peninsula,Canada

Abstract. Despite international recognition that alvar habitats are important reservoirs of biodiversity, they remain little studied in North America. In this paper, the results are reported on an investigation of alvars in the central portion of their known distribution on this continent. 210 plots were distributed among seven sites and were quantitatively sampled for vascular plants, lichens, bryophytes and a suite of environmental variables. Detrended and Canonical Correspondence Analyses and other methods were used to investigate differences among alvars, within alvars and between alvars and adjacent forested habitats. The plant communities and environmental conditions were highly similar among sites in the study region, yet very different from surrounding habitats. There were abrupt changes in vegetation and environmental conditions from alvar to forest, without the presence of transition zones in the vegetation or environmental gradients as the forest was approached. The environmental factors associated with the change from alvar to forest and with variation within alvar habitat were examined. Some alvars in the study were found to contain stunted, slow-growing trees reaching ages of 524 yr. These same sites appear to have remained unburned for several centuries, while other sites likely burned 90 yr ago. The plant communities were very similar between the alvars that lacked a major, biomass-removing disturbance in centuries and alvars that had experienced catastrophic fire relatively recently. Maintenance of the plant communities and open nature of alvars appears site-specific rather than habitat specific.     

The conservation of ground layer lichen communities in alvar grasslands and the relevance of substitution habitats

Semi-natural calcareous grasslands (alvars) are biodiversity hotspots in Northern Europe, particularly for herb layer plants. In the last century, traditional management has ceased, and the area of grasslands has declined due to extensive encroachment. We were interested in the drivers of ground layer (alias terricolous or epigeic) lichen communities. Our survey consisted of 86 habitat fragments in western Estonia, covering four types of historic alvar grasslands and three types of alvar-like habitats. We found that the ground lichen communities were primarily soil-type-specific, but were also affected by historic disturbances and land use change. In contrast to knowledge about herb layer communities, for which shrub encroachment has been shown to be main driver, the increased density of the herb layer and the reduced diversity of microhabitats were major drivers for the ground layer lichen community. These drivers caused a decrease in species richness, but only within the species of conservation value, and also led to a shift in the composition of lichen growth form from the dominance of squamulose and crustose towards fruticose lichens. We conclude that the traditional practice of restoring alvars by cutting shrubs is insufficient to maintain ground layer lichen biodiversity. Alvar maintenance practices should include grazing, which creates various small-scale ground disturbances and increases microhabitat heterogeneity. Alvar-like habitats originating from large-scale historic disturbances appeared to be suitable for calcicolous epigeic lichens, and can therefore be considered to be temporary substitution habitats, i.e. refugia for the regional species pool.     

The ecological equivalence of quarry floors to alvars

Question: Are the biophysical conditions of abandoned limestone quarry floors and natural alvars sufficiently similar to each other for alvars to be used as a model for quarry floor restoration? Location: Ontario, Canada. Methods: We measured plant species frequency and environmental and soil variables in 13 abandoned limestone quarries and used ANOVA to compare them with data previously collected from seven natural alvars. We used multivariate ordinations on the quarry floor data alone and on the combined quarry floor and alvar data to determine how plant community structure was controlled by the abiotic environment in both habitats. Results: Except for higher levels of many nutrients, the physical characteristics were similar between quarry floors and alvars. 246 plant species were found on quarry floors as compared to 283 on alvars, with 79 species in common between the two habitat types. While quarry floors supported fewer bryophytes and more exotic vascular plants compared with alvars, five alvar endemics and 24 characteristic alvar species were found to grow there. The age of the site, nutrient levels, and presence of standing water and bare rock were important factors influencing species composition in both habitats. Conclusions: Through natural revegetation alone, the abandoned quarry floors surveyed in this study have already taken on many physical and vegetation characteristics of natural alvars. This makes alvars very suitable as the restoration goal for abandoned limestone quarries.     

Use of vegetation classification and plant indicators to assess grazing abandonment in Estonian coastal wetlands

Question: What are the effects of grazing abandonment on the vegetation composition of Estonian coastal wetlands? Location: Vormsi Island and Silma Nature Reserve in western Estonia, Europe. Methods: Local knowledge and field reconnaissance were used to identify current and historical management levels of wetland sites within the west Estonian study area. Nine study sites, with varying management histories, were selected comprising an area of 287 ha. A total of 198 quadrats were taken from 43 distinct vegetation patches in five of the sites. TWINSPAN analysis was used to identify community type, and a phytosociological key was constructed for character taxa. This vegetation classification was then applied within a GIS‐based context to classify all the study sites, using a ground survey technique and 1:2000 scale air photos. Results: We identified 11 different brackish coastal wetland community types. Indicator species were defined with community characteristics for the seven main vegetation types readily recognisable in the field. Coastal wet grasslands were most extensive in grazed sites, or sites that had been more intensively grazed, while abandoned sites were largely composed of Phragmites australis stands, tall grassland, and scrub. Site variations based on vegetation composition were significantly correlated with past grazing intensity. Plant community types showed significant edaphic differences, with particularly low soil moisture and high conductivity and pH for open pioneer patches compared to other vegetation types. Conclusion: Abandonment of traditionally grazed coastal grasslands threatens their characteristic biodiversity. This study found that grazing abandonment reduced the extent of coastal wetland grasslands of particular conservation value. Nevertheless, plant species of conservation interest were found across the sequence of community types described. The study shows that grazing is an important factor influencing coastal wetland plant communities but suggests that vegetation distribution is affected by environmental variables, such as topography.     

Relationships between seed banks and spatial heterogeneity of North American alvar vegetation

Abstract. This is the first quantitative study of seed bank characteristics in North American alvar habitats. We assessed seed bank density, species richness, and species composition in 75 plots distributed among five alvar sites in Bruce Peninsula National Park, Ontario, Canada, each of which displayed areas of high and low vegetation cover within the alvar and a fully forested perimeter area. Forested habitats immediately adjacent to alvar patches contained minimal seed banks for species restricted to the alvar patches. Open alvars contained less than 1% seeds from woody forest species. This suggests that forest is not invading adjacent alvar habitat via seeds and that adjacent forest does not contain a reservoir of alvar seeds. When compared to areas on the alvar with high vascular plant cover, areas with low cover contained a slightly smaller viable seed bank, but seed banks from high and low vegetation cover plots had similar species composition and species richness. High vegetation cover plots had slightly higher mean and maximum soil depths compared with low cover plots, but no differences in other physical and chemical parameters. Thus, spatial heterogeneity in plant cover is associated only weakly with heterogeneity in below‐ground factors. Despite the availability of seed and soil resources, vegetation dynamics are constrained in areas with low plant cover, and thus alvar community development seems to respond non‐linearly to resource availability.     

Slow response of plant species richness to habitat loss and fragmentation

We examined the response of vascular plant species richness to long-term habitat loss and fragmentation of Estonian calcareous grasslands (alvars). The current number of habitat specialist species in 35 alvars was not explained by their current areas and connectivities but it was explained by their areas and connectivities 70 years ago ( R ² = 0.27). We estimated the magnitude of extinction debt in local communities by assuming an equilibrium species richness in 14 alvars that had lost only a small amount of area and by applying this model to the remaining alvars, in which the average area has declined from 3.64 km² in the 1930s to 0.21 km² at present. The extinction debt estimated for individual alvars was around 40% of their current species number. Our conclusions are applicable to temperate grasslands in general, which have lost much area because of agricultural intensification and cessation of traditional management.     

Radar remote sensing as a novel tool to assess the performance of an agri-environment scheme in coastal grasslands

Coastal grasslands are semi-natural habitats used for agriculture, but they are also vital habitats for many taxa, such as waders. The importance of this habitat for wildlife has found recognition in the policies of the European Union, resulting in the implementation of biodiversity-friendly agri-environment schemes (AES) to ensure the sustainable management of these areas. However, the performance of AESs has often been questioned and data to analyse their effectiveness is mostly lacking.To fill this knowledge gap, we used freely available European Space Agency's Sentinel-1 Synthetic Aperture Radar (SAR) images and ground truth data to describe the vegetation height based habitat suitability of coastal grasslands as breeding sites for waders at a country-wide scale in Estonia, where the main threat to this habitat is overgrowing with high sward plant species.We show that satellite remote sensing can be used as a straight-forward and reliable tool to estimate the grazed coastal grassland habitat suitability for breeding waders in terms of vegetation height in two broad classes.We demonstrate that the AES applied on Estonian coastal grasslands is important for improving the suitability of this habitat for the breeding wader community: compared to 24% of suitable habitat in areas where no agricultural subsidies are paid, 60% of the land under the targeted AES is suitably managed for breeding waders. The highest proportion of habitat suitability (76%) is achieved in areas receiving targeted top-up subsidy for wader conservation.Continued monitoring of the performance of the AES is essential for taking timely and targeted conservation actions and we strongly recommend the wider use of remotely sensed data for this purpose. We promote the continuation of the AES scheme applied on Estonian coastal grasslands while improving its efficiency through carefully raised stocking rates and increased uptake of the top-up scheme.     

Formation and succession of alvar communities in the Baltic land uplift area

Calciphilous plant communities, soil and geomorphology along the gradient of elevation in land uplift area in the coast of western Estonia were described. In grazed areas, species rich alvar grassland is a final stage in successional sere. The differences in site history in various points of the transect were reflected by the variability of underlying deposits. When soils with different ages but having similar parent material were compared, some successional trends (decrease of the content of Ca, increase of the content of P and humus supply) became evident. Species composition of plant communities was determined mainly by local ecological conditions (moisture level, soil pH, also grazing) at any particular site. Differences between communities of the same association but with different successional age were negligible.     

Hybrid ecosystems can contribute to local biodiversity conservation

Calcareous grasslands have become severely threatened habitats in Europe. The aim of this study was to investigate the changes in plant species richness, and functional and phylogenetic diversity in northern Estonian calcareous (alvar) grasslands resampled after 90 years of land-use change. Functional traits characterizing species that have benefited most from decreased habitat area and altered environmental conditions, and additional species that can potentially inhabit the remaining grassland patches were identified. Also changes in the relative amount of habitat-specific species were studied to detect a possible decrease in habitat integrity. Although grasslands in the studied region had lost most of their original area (~90 %), species richness had substantially increased due to invasion by more competitive, nutrient-demanding native species. Functional diversity generally increased, whereas phylogenetic diversity showed no response to altered conditions. Overall, these grasslands have lost their integrity as calcareous grassland habitat type in the region, because the relative amount of habitat-specific characteristic species has declined significantly. However, although the grasslands have transformed to a ‘hybrid’ habitat type and restoration to their previous state is likely not reasonable, such degraded species-rich grassland fragments can still be recognized as important habitats to preserve high local biodiversity and several characteristic species of calcareous grasslands. As current landscapes consist of an increasing number of hybrid and novel communities, new tools to supplement traditional conservation or restoration practices are necessary to recognize and maintain regions and habitats of high local biodiversity.     

The seed bank in an estonian calcareous grassland: Comparison of different successional stages

Seed bank species-composition and seed-density were determined in a successional calcareous (alvar) grassland in western Estonia. Three similar study areas were chosen to compare two different successional stages: open alvar grassland and overgrown areas with young pine forest 30 to 40 years old. In both successional stages, the centre and the edge of a relatively uniform stand were examined. Fifteen soil samples (7 cm in diameter, 5 cm deep) were taken from each of twelve sampling sites. The seedling emergence method was used to estimate seeds in the soil samples. A total of 69 species were detected in the seed bank, of which 18 did not occur in the vegetation. Eighty-nine taxa were recorded in the vegetation and of these 38 were not detected in the seed bank. Fifty-one species occurred both in the seed bank and in the vegetation. The three most abundant taxa in the seed bank wereCarex tomentosa, Linum catharticum andPlantago media, which together made up 49% of the seedlings recorded. Differences in the species compositions of seed bank samples from grassland and forest sites were negligible, although the species richness per area of the above-ground vegetation was significantly higher in the open grassland. The only species tending to be lost from forest site vegetation but still occurring in the forest soil seed bank wereArenaria serpylifolia, Cerastium fontanum andLinum catharticum. About half of all the emerged species from all samples belonged to the transient or short-term persistent seed bank. In the grassland sites there were more species which belonged to the transient seed bank than in the forest sites, where the seed bank contained more short-term persistent type seeds. The seed density was significantly higher in forest sites and lower in grassland sites, which may be explained by the better germination conditions in well-illuminated communities. On the basis of the current study it might be assumed that the soil seed banks of overgrown alvar grasslands which include young pine forests can play a certain role in grassland restoration management.     

Functional ecology of rare and common epigeic lichens in alvar grasslands

The autecology of rare species can be derived using similarities among functional traits and environmental conditions observed for common species, i.e. we employed the ‘matching analogy approach’ with the analytical scheme ‘common species → driver → trait → driver → rare species’. We addressed the driver–trait relationship for common epigeic lichens of thin-soil calcareous grasslands, which are endangered by cessation of traditional land use. Common lichens were suppressed by encroaching herbs and shrubs, and were supported by ground disturbances. The lichens of open low-productivity alvars are predominantly calciphilous, epibryic, crustose-squamulose, sexually reproducing and contain UV-protective pigments. Lichens of encroached alvars are soil-type generalists, fruticose, reproduce vegetatively and contain herbivore-deterring compounds. Rare lichens resemble the species of open low-productivity alvars, except their more limited niche space, i.e. they are restricted to arctic-alpine habitats. The conservation practices on alvars should support the formation of crust-forming communities by suppressing the growth of shrubs and herbs, and by promoting recurrent small-scale soil disturbances.     

Restoration of alvar vegetation on Öland,Sweden

Abstract. Alvars in the Baltic region, particularly on the Swedish islands of Öland and Gotland and in western Estonia, are well‐known for their plant species richness and extensive populations of rare species. Grasslands make up most of alvar vegetation. The extent of these grasslands decreases because of bush enchroachment which takes place in most alvars when traditional land use practice changes, notably when grazing is ceased. The main threat for the alvar grassland is formed by the rapid expansion of Juniperus communis and Potentilla fruticosa in dry and wet sites, respectively. Applied research has been carried out during several years in order to develop plans for the restoration of alvar grassland. Thus it was shown that species richness in areas with a mixture of grassland and shrubland decreases in relation to the increased cover by shrubs. In the case of Juniperus there is a distinct drop in species number as soon as the cover exceeds 75‐80%, while the decrease is more gradual with increased cover of Potentilla. The seed banks under dense stands of these shrubs only contain ca. 20% of the species found in dry and wet alvar meadows. This was confirmed by clearing experiments. Long‐term recordings have shown that establishment of juniper seedlings takes place mainly in half‐open areas between already existing junipers. Intermediate‐sized junipers expanded faster than old and big shrubs. Potentilla shrubs recover fast from cutting or burning. After 2‐4 yr they have almost attained their former size. This recovery can be prevented when cattle grazing is introduced. In the framework of a local alvar restoration project on Öland, covering 7000 ha, grazing regimes are re‐established, fences erected and large‐scale clearings carried out. Recently established juniper shrubs are being cleared, intermediate sized junipers (30‐50 yr old) are selectively removed while creating a mosaic landscape with high biological diversity. Older dense juniper stands are left alone or are only partly cleared. Potentilla stands in moist areas are cut to create moist meadows which are breeding grounds for waders, and to establish corridors between remaining open areas. Three items are discussed (1) the importance of the Stora Alvar area; (2) re‐introduction of grazing and (3) re‐introduction of species. The outstanding importance of the area regarding species richness and endemism should be recognized through a ‘golden list’ to be used along with red and blue lists.     

Small-scale plant species richness in calcareous grasslands determined by the species pool, community age and shoot density

Variation in small-scale plant species richness was examined between twenty discrete sites of calcareous alvar grasslands in the western Estonian coastal area. This community type is very species rich at the small-scale. The pattern of diversity has often been explained by variation in ecological conditions which makes it possible to avoid competitive exclusion. Here we test an alternative species pool hypothesis. The species pool was defined as the set of species present in a community, but excluding all those species whose presence may be connected with exceptional environmental conditions for this community. Monte Carlo modelling was used to determine the strengths of the relationships between the non-independent variables: richness vs species pool and average shoot density vs richness. Site age was characterized by its elevation above sea level, since there is a continuous land uplift in the study area. Species richness was positively correlated to the size of the species pool, community age and vegetation density. We conclude that historical processes on both the regional and local levels, which determine the arrival of any particular species to a target community, and not environmental heterogeneity, are responsible for the variations in species richness between communities of this type.     

Specialist taxa restricted to threatened habitats contribute significantly to the regional diversity of Peltigera (Lecanoromycetes,Ascomycota) in Estonia

The widespread cyanolichen genus Peltigera comprises many insufficiently known poorly delimited and/or undescribed species. Phylogenetic analysis of 252 Peltigera specimens from a wide range of habitat types in Estonia revealed 31 putative taxa (OTUs). Multivariate analysis revealed habitat-specific segregation between the Peltigera species along a gradient from humid eutrophic forests to dry oligotrophic forests and grasslands and along a soil pH gradient from alkaline soils of alvar grasslands to acidic soils of conifer forests. The diversity of Peltigera was the highest on roadsides and dunes and the lowest in alvar habitats which, however, supported the unique assemblage of undescribed Peltigera taxa. Deciduous broad-leaved forests, too, included several undescribed or rare and red-listed species. The results demonstrate that in Estonia many Peltigera species have narrow habitat requirements and are at present threatened by habitat loss and degradation.     

Rare and threatened plant communitiesof Estonia

This paper introduces the criteria for identifying rare and threatened plant communities in Estonia. An overview of such communities is given and the general problems associated with their protection are briefly discussed. Of forest communities, alvar forests and boreo-nemoral forests must be preserved very carefully. Once widely distributed minerotrophic mobile-water swamp forests and flood plain forests nowadays cover only a small area. Numerous rare communities are associated with coastal regions and islands. Wooded meadows, the most species rich vascular plant communities in northern Europe, as well as flooded meadows and many other communities require continuation of traditional management for their preservation.     

Determinants of bryophyte species composition and diversity on the Great Alvar of Öland,Sweden

Factors driving the species richness and distribution of bryophytes are poorly studied and not well understood, particularly in grasslands. We analysed the occurrence of bryophyte species and variation in species richness across 674 plots (0.5?m?×?0.5?m) in alvar vegetation (grassland on limestone pavement with thin or no soil) on Öland (Sweden) in relation to substrate characteristics and chemistry, inundation frequency, grazing pressure and geographical variables. We found 148 taxa, including 11 nationally red-listed ones. Species richness per plot was significantly associated with substrate type, positively associated with pH and grazing intensity, but negatively associated with soil depth. However, richness of species typical of, or restricted to, alvar habitats responded differently to richness of species more common in other habitats. Typical alvar species were favoured by high pH, shallow soil and low phosphate availability, while generalists preferred relatively low pH, higher phosphate availability and organic or mull soil types. Distance from the alvar margin had only weak effects. Concerning the effects on individual species and community composition, inundation frequency and pH were found to have the largest effects, although other factors (substrate type, soil depth, bare soil, bare stone, phosphate availability and grazing pressure) were more important for some individual species, stressing the importance of microsite variability and variability in management for regional species richness. From a conservation perspective, it is concluded that grazing is generally positive whilst factors increasing phosphate availability may disadvantage the typical alvar species, and proximity to the alvar margin is not a major problem.     

The plant cover of Russia in the area of the Braun-Blanquet syntaxonomy: the development of an approach and the results

Theoretical basis of ecologo-floristic classification according to Braun-Blanquet approach is discussed with the special attention to the contribution of Russian scientists. Principal results concern presentation of vegetation continuum in discrete classification units: polymodal conception of plant communities, principle of plural syntaxonomic decisions, pragmatic approach to classification, method of homological rows of plant communities, etc. Current ecologo-floristic classification of vegetation in Russia consists of 77 classes. This system is a good basement for estimation of rarity of plant communities and biodiversity protection. The first variant of "Red Book" of plant communities of Russia and other eastern European countries is discussed.     

The role of a mound-building ecosystem engineer for a grassland butterfly

Both land use intensification and abandonment within grasslands lead to a homogenisation of vegetation structure. Therefore, specially structured microsites such as vegetation gaps with bare ground play an important role for species conservation within grasslands. Vegetation gaps are crucial for the establishment of low-competitive plant species and offer special microclimatic conditions essential for the development of the immature stages of many invertebrate species. The influence of small-scale soil disturbance in the form of mounds created by ecosystem engineers such as ants or moles on biodiversity is therefore of special scientific concern. The effects of mound-building species on plant species diversity have been extensively studied. However, knowledge on the significance of these species for the conservation of other animals is rare. In this study we analyse the importance of mounds created by the European mole (Talpa europaea) as an oviposition habitat for the small copper (Lycaena phlaeas) within Central European mesotrophic grasslands. Our study showed that host plants occurring at molehills were preferred for oviposition. Oviposition sites were characterised by an open vegetation structure with a high proportion of bare ground (with a mean coverage of about 50 %), a low cover of herbs and low-growing vegetation (mean height: 4.5 cm). Our study clearly illustrates the importance of small-scale soil disturbance for immature stages of L. phlaeas and the conservation of this species within mesotrophic grasslands. Mound-building ecosystem engineers, such as T. europaea, act as important substitutes for missing dynamics within mesotrophic grasslands by diversifying vegetation structure and creating small patches of bare soil.