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Calcareous alvar grasslands are one of the most species-rich habitats in Estonia. Land-use change and cessation of traditional agricultural practices have led to a decrease of the area of these valuable grasslands during the past century. Therefore, their conservation and restoration are becoming increasingly important. Efforts to restore these habitats have already been made in recent years. Land suitability analysis for potential restoration sites, using the machine learning technique Random Forest (RF), was performed for the first time in this study, which aimed to assess the use of RF for a suitability analysis of alvar grassland. RF predicted 610.91 km2 of areas suitable for restoring alvar grasslands or for creating alvar- like habitats in Estonia. These areas include all existing alvar areas as well an additional 140.91 km2 suitable for establishing new habitat similar to calcareous alvar grasslands. We discuss suitability analysis to help with restoration planning and find it to be a reasonable and efficient tool that has potential to provide relevant information. The quality of the prediction could be improved by including additional data relevant for alvar grasslands, such as soil depth, but such data was unfortunately unavailable.

Recent habitat change in semi-natural grasslands due to a lack of management has been shown to affect the genetic diversity of grassland plants. However, it is unknown how a change in local environment affects genetic diversity at adaptive loci. We applied RADseq (restriction-site associated DNA sequencing) to extract > 3,000 SNPs across 568 individuals from 32 Estonian populations of Primula veris, a plant species common to semi-natural grasslands. We evaluated the effect of recent grassland overgrowth following management abandonment on the genetic diversity at putatively neutral and adaptive loci, which we distinguished by applying three methods, i.e., linear and categorical environmental association analyses, and an FST outlier test. For validation, we randomised the genotype to sample assignments. Effects of recent habitat change on genetic diversity differed between neutral and adaptive SNP sets. Genetic diversity assessed at putatively neutral loci was similar in open and overgrown habitats but showed a significant difference between these habitat types at putatively adaptive loci: overgrown (i.e. newly established) habitats exhibited higher genetic diversity at putatively adaptive loci than open (i.e. old) habitats, likely due to the exertion of novel selection pressures imposed by new habitat conditions. This increase in genetic diversity at putatively adaptive loci in the new environment points to currently ongoing selection processes where genetic adaptation to the old habitat is potentially lost through altered allele frequencies. Our study suggests that a recent change in local habitat conditions may not be reflected in neutral loci whereas putatively adaptive loci can inform about potential selection processes.

© 2021 The Authors., Aims: Understanding fine-grain diversity patterns across large spatial extents is fundamental for macroecological research and biodiversity conservation. Using the GrassPlot database, we provide benchmarks of fine-grain richness values of Palaearctic open habitats for vascular plants, bryophytes, lichens and complete vegetation (i.e., the sum of the former three groups). Location: Palaearctic biogeographic realm. Methods: We used 126,524 plots of eight standard grain sizes from the GrassPlot database: 0.0001, 0.001, 0.01, 0.1, 1, 10, 100 and 1,000 m and calculated the mean richness and standard deviations, as well as maximum, minimum, median, and first and third quartiles for each combination of grain size, taxonomic group, biome, region, vegetation type and phytosociological class. Results: Patterns of plant diversity in vegetation types and biomes differ across grain sizes and taxonomic groups. Overall, secondary (mostly semi-natural) grasslands and natural grasslands are the richest vegetation type. The open-access file ”GrassPlot Diversity Benchmarks” and the web tool “GrassPlot Diversity Explorer” are now available online (https://edgg.org/databases/GrasslandDiversityExplorer) and provide more insights into species richness patterns in the Palaearctic open habitats. Conclusions: The GrassPlot Diversity Benchmarks provide high-quality data on species richness in open habitat types across the Palaearctic. These benchmark data can be used in vegetation ecology, macroecology, biodiversity conservation and data quality checking. While the amount of data in the underlying GrassPlot database and their spatial coverage are smaller than in other extensive vegetation-plot databases, species recordings in GrassPlot are on average more complete, making it a valuable complementary data source in macroecology., GrassPlot development has been supported by the Bavarian Research Alliance (BayIntAn_UBT_2017_58), the Eurasian Dry Grassland Group (EDGG) and the International Association for Vegetation Science (IAVS); IB, CorM, JAC, IGM, DGM, MHe, DL and MTo were supported by the Basque Government (IT936‐16); CorM, IAx, MCh, JDa, PD, MHá, ZL, ZPr, EŠ and LT were supported by the Czech Science Foundation (19‐28491X); TR was supported by the Estonian Research Council (PUT1173); RJP was funded by the Strategic Research Programme of the Scottish Government’s Rural and Environmental Science and Analytical Services Division”; SBa was supported by the GINOP‐2.3.2‐15‐2016‐00019 project; GFi was partially supported by the MIUR initiative “Department of excellence” (Law 232/2016)"; BJA was funded by the Spanish Research Agency (grant AEI/ 10.13039/501100011033); AK, VB, IM, DS, IV and DV were supported by the National Research Foundation of Ukraine (2020.01/0140); MP and AH were supported by the Estonian Research Council (PRG874, PRG609), and the European Regional Development Fund (Centre of Excellence EcolChange); Data collection of HCP was funded by FORMAS (Swedish Research Council for Environment, Agricultural Science and Spatial Planning) and The Swedish Institute; JR was supported by the Czech Science Foundation (grant No. 20‐09895S) and the long‐term developmental project of the Czech Academy of Sciences (RVO 67985939); ATRA was funded by the Grant of Excellence Departments, MIUR‐Italy (ARTICOLO 1, COMMI 314 – 337 LEGGE 232/2016); JMA was supported by Carl Tryggers stiftelse för vetenskaplig forskning and Qatar Petroleum; AAli was supported by the Jiangsu Science and Technology Special Project (Grant No. BX2019084), and Metasequoia Faculty Research Startup Funding at Nanjing Forestry University (Grant No. 163010230), and he is currently supported by Hebei University through Faculty Research Startup Funding Program; ZB was supported by the NKFI K 124796 grant; The GLORIA‐ Aragón project of JLBA was funded by the Dirección General de Cambio Climático del Gobierno de Aragón (Spain); MCs and LDem were supported by DG Environment through the European Forum on Nature Conservation and Pastoralism and Barbara Knowles Fund, in collaboration with Pogány‐havas Association, Romania; JDa was partially supported by long‐term research development project no. RVO 67985939 of the Czech Academy of Sciences; BD and OV were supported by the NKFI KH 126476, NKFI KH 130338, NKFI FK 124404 and NKFI FK 135329 grants; BD, OV and AKe were supported by the Bolyai János Scholarship of the Hungarian Academy of Sciences; BE was funded by the Environmental Department of the Tyrolean Federal State Government, the MAB Programme of the Austrian Academy of Science, the Mountain Agriculture Research Unit and the Alpine Research Centre Obergurgl of Innsbruck University. The GLORIA projects of BE were funded by the EU project no. EVK2‐CT‐2000‐00056, the Earth System Sciences Program of the Austrian Academy of Sciences (project MEDIALPS), the Amt für Naturparke, Autonome Provinz Bozen‐Südtirol, the Südtiroler Wissenschaftsfonds and the Tiroler Wissenschaftsfonds; RGG was supported by the Spanish Ministry of Research to sample GLORIA sites in central Spain (CGL 2008‐00901/BOS) and present works by the Autonomous Region of Madrid (REMEDINAL TE‐CM, S2018/EMT‐4338); MJ was supporteLatviaed by Latvia Grant No. 194051; NP and SŠ were partly supported by the Slovenian Research Agency, core fundings P1‐0403 and J7‐1822.

Highlights • New empirical studies are repeatedly demonstrating the importance of rare species. • However, very little is still known about species rarity in the aquatic realm. • Continental maps suggest variable latitudinal trends in freshwater plants. • Human footprint, past and present-day climates predict rarity hotspots. • Current centers of freshwater plant rarity might shrink within a century. Patterns of species rarity have long fascinated ecologists, yet most of what we know about the natural world stems from studies of common species. A large proportion of freshwater plant species has small range sizes and are therefore considered rare. However, little is known about the mechanisms and geographical distribution of rarity in the aquatic realm and to what extent diversity of rare species in freshwater plants follows their terrestrial counterparts. Here, we present the first in–depth analysis of geographical patterns, potential deterministic ecogeographical factors and projected scenarios of freshwater vascular plant rarity using 50 × 50 km grid cells across Europe (41°N–71°N) and North America (25°N–78°N). Our results suggest that diversity of rare species shows different patterns in relation to latitude on the two continents, and that hotspots of rarity concentrate in a relatively small proportion of the European and North American land surface, especially in mountainous as well as in climatically rare and stable areas. Interestingly, we found no differences among alternative rarity definitions and measures when delineating areas with notably high diversity of rare species. Our findings also indicate that few variables, namely a combination of current climate, Late Quaternary climate–change velocity and human footprint, are able to accurately predict the location of continental centers of rare species diversity. However, these relationships are not geographically homogeneous, and the underlying factors likely act synergistically. Perhaps more importantly, we provide empirical evidence that current centers of rare species diversity are characterized by higher anthropogenic impacts and might shrink disproportionately within this century as the climate changes. Our reported distributional patterns of species rarity align with the known trends in species richness of other freshwater organisms and may help conservation planners make informed decisions mitigating the effects of climate change and other anthropogenic impacts on biodiversity.

The prevailing nature-based solution to tackle climate change is tree planting. However, there is growing evidence that it has serious contraindications in many regions. The main shortcoming of global tree planting is its awareness disparity to alternative ecosystem types, mainly grasslands. Grasslands, where they constitute the natural vegetation, may support higher biodiversity and a safer, soil-locked carbon stock than plantations and other forests. We suggest replacing “tree planting” by “restore native vegetation.” This improved action terminology reduces the risks of inappropriate afforestation and, by diversifying target ecosystem types, does not reduce but increases potential land area for nature-based climate mitigation.

Species dispersal, establishment, and assembly are crucial stages of the life history of plants, and clear understanding of the governing forces and rules that shape species composition in a particular community is vital for successful ecological restoration. In this article, we focus on five aspects of seed dispersal and plant establishment, which should be considered during habitat restoration actions. In the first two sections, we discuss the success of spontaneous dispersal and establishment on restoration based on either spatial dispersal or local seed banks. In the third section, we assess the possibilities of species introduction and assisted dispersal. In the fourth section, we introduce some possibilities for the improvement of establishment success of spontaneously dispersed or introduced species. Finally, we highlight issues influencing long‐term persistence and sustainability of restored habitats, related to the alteration of management type and intensity, climate change, and spread of non‐native species. With the present article, we introduce the special issue entitled “Seed dispersal and soil seed banks – promising sources for ecological restoration” containing 15 papers by 62 authors from 10 countries arranged in the abovementioned five topics.

1. The latitudinal diversity gradient hypothesis suggests that species richness should be highest at low latitudes, whereas Rapoport’s rule states that largest ranges ought to be found for species at high latitudes. However, there is no consensus over these patterns and their underlying drivers in the freshwater realm. 2. We investigated species richness and mean range size of freshwater plants in 50 × 50 km grid cells across Europe (40°N–71°N) and North America (25°N–78°N), supplemented with data based on 1° latitudinal bands for mean range size. We were especially interested to find out whether there are similarities and differences in these ecogeographical patterns and their underlying drivers between the continents due to their contrasting historical characteristics, spatial extent and topography. 3. First, we used partial regression to reveal whether species richness and mean range size of freshwater plants have a linear or quadratic relationship with latitude. Second, we employed variation partitioning based on partial regression to model relationships between plant species richness and mean range size and four explanatory variable groups (i.e. environmental features, current climate, historical climate and geographical location). Third, we utilized boosted regression tree analysis to further investigate species richness and mean range size of freshwater plants in relation to a set of explanatory variables. 4. Our results revealed that species richness showed relatively similar patterns in relation to latitude between the continents. Similarly, mean range size trends were alike in North America whether we used 50 × 50 km grid cell data or 1° latitudinal bands. Instead, different patterns in mean range size emerged between the used datasets in Europe. For both species richness and mean range size, current climate (with different individual predictor variables) was the main driver in both the continents, but historical effects had a small influence on the response variables. 5. Synthesis. Our findings indicated that major ecogeographical rules can strongly vary for the same taxonomic group across broad scales between continents. It is also premature to rely solely on well‐known terrestrial taxonomic groups when drawing generalizations about ecogeographical rules.

During the past century, grasslands in Europe have undergone marked changes in land-use, leading to a decline in plant diversity both at local and regional scales, thus possibly also affecting the mechanisms of species sorting into local communities. We studied plant species assembly in grasslands with differing habitat history and hypothesised that trait divergence prevails in historical grasslands due to niche differentiation and trait convergence prevails in more dynamic grasslands due to competitive exclusion and dispersal limitation. We tested these hypotheses in 35 grassland complexes in Estonia, containing neighbouring grassland habitats with different land-use histories: continuously managed open historical grassland, currently overgrown former grassland and young developing grassland. We assessed species assembly patterns in each grassland type for finer scale-a 2 × 2 m plot scale from a local community pool and for broader scale-a local community from the habitat species pool for that grassland stage and observed changes in trait means at finer scale. We found that grasslands with long management history are assembled differently from former grasslands or young developing grasslands. In historical grasslands, divergence or random patterns prevailed at finer scale species assembly while in former or developing grasslands, mostly convergence patterns prevailed. With increasing scale convergence patterns become more prevalent in all grassland types. We conclude that land-use history is an important factor to consider when assessing grassland functional trait assembly, particularly at small scales. Understanding the mechanisms behind species assembly and their relationship with land-use history is vital for habitat conservation and restoration.

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.

Traditionally managed and unregulated floodplain meadows are characterised by high and habitat-specific biodiversity. In most parts of Europe, however, these habitats are almost completely lost. Despite possessing high importance in conservation, little is known about the dynamics of important species groups in those valuable communities. Here, we studied the diversity and composition of ground-dwelling spider fauna in ten sites of temporarily flooded semi-natural grasslands in the Matsalu Bay, Western Estonia. As fluctuation of aquatic and terrestrial phases is the strongest factor determining the environmental conditions in flooded coastal and floodplain meadows, we estimated the effect of floods on spider diversity and composition. Studied grasslands were characterized by opportunistic, eurytopic and hydrophilic spider species that are also abundant in other disturbed habitats. Most species belonged to families characterised by species with good dispersal ability (Linyphiidae and Lycosidae). Average ecological flexibility of spiders (i.e. number of preferred habitats) increased, while activity density of spiders declined towards the sea, where floods are more frequent and more saline. Our results indicate that the spider fauna of floodplain meadows is dependent on flooding regime and is likely re-structured after major flooding events via colonisation from nearby habitats.

GrassPlot is a collaborative vegetation-plot database organised by the Eurasian Dry Grassland Group (EDGG) and listed in the Global Index of Vegetation-Plot Databases (GIVD ID EU-00-003). GrassPlot collects plot records (relevés) from grasslands and other open habitats of the Palaearctic biogeographic realm. It focuses on precisely delimited plots of eight standard grain sizes (0.0001; 0.001; ... 1,000 m²) and on nested-plot series with at least four different grain sizes. The usage of GrassPlot is regulated through Bylaws that intend to balance the interests of data contributors and data users. The current version (v. 1.00) contains data for approximately 170,000 plots of different sizes and 2,800 nested-plot series. The key components are richness data and metadata. However, most included datasets also encompass compositional data. About 14,000 plots have near-complete records of terricolous bryophytes and lichens in addition to vascular plants. At present, GrassPlot contains data from 36 countries throughout the Palaearctic, spread across elevational gradients and major grassland types. GrassPlot with its multi-scale and multi-taxon focus complements the larger international vegetation-plot databases, such as the European Vegetation Archive (EVA) and the global database “sPlot”. Its main aim is to facilitate studies on the scale- and taxon-dependency of biodiversity patterns and drivers along macroecological gradients. GrassPlot is a dynamic database and will expand through new data collection coordinated by the elected Governing Board. We invite researchers with suitable data to join GrassPlot. Researchers with project ideas addressable with GrassPlot data are welcome to submit proposals to the Governing Board. Abbreviations: EDGG = Eurasian Dry Grassland Group; EVA = European Vegetation Archive; GrassPlot = Database of Scale-Dependent Phytodiversity Patterns in Palaearctic Grasslands; SAR = species-area relationship.

Temperate calcareous grasslands are characterized by high levels of species richness at small spatial scales. Nevertheless, many species from a habitat-specific regional species pool may be absent from local communities and represent the dark diversity' of these sites. Here we investigate dry calcareous grasslands in northern Europe to determine what proportion of the habitat-specific species pool is realized at small scales (i.e. how the community completeness varies) and which mechanisms may be contributing to the relative sizes of the observed and dark diversity. We test whether the absence of particular species in potentially suitable grassland sites is a consequence of dispersal limitation and/or a low ability to tolerate stress (e.g. drought and grazing). We analysed a total of 1223 vegetation plots (1 x 1 m) from dry calcareous grasslands in Sweden, Estonia and western Russia. The species co-occurrence approach was used to estimate the dark diversity for each plot. We calculated the maximum dispersal distance for each of the 291 species in our dataset by using simple plant traits (dispersal syndrome, growth form and seed characteristics). Large seed size was used as proxy for small seed number; tall plant height and low S-strategy type scores were used to characterise low stress-tolerance. Levels of small-scale community completeness were relatively low (more species were absent than present) and varied between the grasslands in different geographic areas. Species in the dark diversity were generally characterized by shorter dispersal distances and greater seed weight (fewer seeds) than species in the observed diversity. Species within the dark diversity were generally taller and had a lower tolerance of stressful conditions. We conclude that, even if temperate grasslands have high levels of small-scale plant diversity, the majority of potentially suitable species in the regional species pool may be absent as a result of dispersal limitation and low stress-tolerance. (Less)

This Special Feature of the Journal of Vegetation Science contains ten contributions from the 56th Symposium of the International Association for Vegetation Science (IAVS), which was held in Tartu, Estonia, 26–30 Jun 2013, and focused on processes underlying vegetation patterns. These contributions give an overview of recent advances in vegetation science, with topics ranging from the introduction of new methods and ideas to comprehensive analysis of vegetation patterns on a global scale. The focus of these Special Feature contributions indicates that vegetation science is becoming more global, more complex and more interdisciplinary. It is likely that progress in methods and accumulation of data have led vegetation science into a ‘new era of discovery’.

Species richness is infl uenced both by mechanisms occurring at landscape scales, such as habitat availability, and local-scale processes, that are related to abiotic conditions and plant – plant interactions. However, it is rarely tested to what extent local species richness can be explained by the combined eff ect of factors measured at multiple spatial scales. In this study, we quantifi ed the simultaneous infl uence of historical landscape-scale factors (past human population density, and past habitat availability – an index combining area and connectivity) and small-scale environmental conditions (shrub cover, and heterogeneity of light, soil depth, and other soil environmental variables) on plant species richness in dry calcareous grasslands (alvars). By applying structural equation modelling (SEM) we found that both landscape conditions and local environmental factors had signifi cant direct and indirect (i.e. through the modifi cation of another factor), eff ects on species richness. At the landscape scale, we found a direct positive infl uence of historical habitat availability on species richness, and indirect positive infl uence of past human population (via its eff ects on historical habitat availability). At small scales, we found a positive direct infl uence of light heterogeneity and shrub cover on species richness. Conversely, we found that small-scale soil environmental heterogeneity, which was mainly determined by soil depth heterogeneity, had a negative eff ect on species richness. Our study indicates that patterns of species richness in alvar grasslands are positively infl uenced by the anthropogenic management regime that maintained the landscape habitat conditions in the past. However, the abandonment of management, leading to shrub invasion and increased competition for light resources also infl uenced species richness. In contrast to the positive heterogeneity – diversity relationship we found that soil heterogeneity reduced species richness. Environmental heterogeneity, occurring at the plant neighbourhood scale (i.e. centimetres), can increase the isolation among suitable soil patches and thus hinder the normal functioning of populations. Th e combination of previous knowledge of the system with new ecological theories facilitates disentangling how species richness responds to complex relationships among factors operating at multiple scales.

A number of studies show contrasting results in how plant species with specific life-history strategies respond to fragmentation, but a general analysis on whether traits affect plant species occurrences in relation to habitat area and isolation has not been performed. We used published data from forests and grasslands in north-central Europe to analyse if there are general patterns of sensitivity to isolation and dependency of area for species using three traits: life-span, clonality, and seed weight. We show that a larger share of all forest species was affected by habitat isolation and area as compared to grassland species. Persistence-related traits, life-span and clonality, were associated to habitat area and the dispersal and recruitment related trait, seed weight, to isolation in both forest and grassland patches. Occurrence of clonal plant species decreased with habitat area, opposite to non-clonal plant species, and long-lived plant species decreased with grassland area. The directions of these responses partly challenge some earlier views, suggesting that further decrease in habitat area will lead to a change in plant species community composition, towards relatively fewer clonal and long-lived plants with large seeds in small forest patches and fewer clonal plants with small seeds in small grassland patches. It is likely that this altered community has been reached in many fragmented European landscapes consisting of small and isolated natural and semi-natural patches, where many non-clonal and short-lived species have already disappeared. Our study based on a large-scale dataset reveals general and useful insights concerning area and isolation effects on plant species composition that can improve the outcome of conservation and restoration efforts of plant communities in rural landscapes.

Intensification or abandonment of agricultural land use has led to a severe decline of semi-natural habitats across Europe. This can cause immediate loss of species but also time-delayed extinctions, known as the extinction debt. In a pan-European study of 147 fragmented grassland remnants, we found differences in the extinction debt of species from different trophic levels. Present-day species richness of long-lived vascular plant specialists was better explained by past than current landscape patterns, indicating an extinction debt. In contrast, short-lived butterfly specialists showed no evidence for an extinction debt at a time scale of c. 40 years. Our results indicate that management strategies maintaining the status quo of fragmented habitats are insufficient, as time-delayed extinctions and associated co-extinctions will lead to further biodiversity loss in the future.

1 Species-rich semi-natural grasslands in Europe developed during prehistoric times and have endured due to human activity. At the same time, intensive grassland management or changes in land use may result in species extinction. As a consequence, plant diversity in semi-natural calcareous grasslands may be related to both historical and current human population density. 2 We hypothesize that current vascular plant diversity in semi-natural calcareous grasslands is positively correlated with the Late Iron Age (c. 800-1000 years ago) density of human settlements (indicated by Late Iron Age fortresses and villages) due to enhancement of grassland extent and species dispersal, and negatively correlated with current human population density due to habitat loss and deterioration. 3 We described the size of the community vascular plant species pool, species richness per 1 m(2) and the relative richness (richness divided by the size of the species pool) in 45 thin soil, calcareous (alvar) grasslands in Estonia. In addition to historical and current human population density we considered simultaneously the effects of grassland area, connectivity to other alvar grasslands, elevation above sea level (indicating grassland age), soil pH, soil N, soil P, soil depth, soil depth heterogeneity, geographical east-west gradient, precipitation and spatial autocorrelation. 4 Both the size of the community species pool and the species richness are significantly correlated with the Late Iron Age human population density. In addition, species richness was unimodally related to the current human population density. The relative richness (species 'packing density') was highest in the intermediate current human population densities, indicative of moderate land-use intensity. 5 Community species pool size decreased non-linearly with increasing soil N, and was highest at intermediate elevation. Small-scale richness was greater when sites were well connected and when the elevation was intermediate. Spatial autocorrelation was also significant for both species pool size and small-scale richness. 6 In summary, human land-use legacy from prehistoric times is an important aspect in plant ecology, which could be an important contributor to the current variation in biodiversity. (Less)

Alvar grasslands are thin-soil calcareous grasslands on Silurian and Ordovician limestone bedrock. They are considered unique plant communities for northern Europe and America with a high conservation value. Until now, European alvar grasslands have only been described in Sweden and Estonia. We described species-rich alvar grasslands on the Izhora plateau, NW Russia and compared this vegetation with that of Estonian alvar grasslands. Floristic composition and environmental conditions of Russian alvars were similar to Estonian alvar grasslands on thicker dry soil. Russian alvars are isolated from similar communities in Estonia, but extensive transport of domestic animals from Estonia to NW Russia in past centuries has probably enhanced species dispersal. Some characteristic Estonian alvar species were missing in NW Russia because it lies outside their natural ranges. Other differences in vegetation were explained by higher atmospheric nitrogen deposition and younger age of Russian sites. In NW Russia the alvar grasslands were small, but several red-listed plant species were present. In order to preserve European alvar vegetation effectively, Russian alvar grasslands need further study and conservation as an outpost of this threatened vegetation type.

Changes in species richness along the ecological succession gradient may be strongly determined by coexisting extinction debts of species from the original habitats and colonization credits of those from the replacing habitats. The magnitude of these processes and their causes remain largely unknown. We explored the extinction debt and colonization credit for grassland and forest specialist plants, respectively, and the local and landscape factors associated to the richness of these species groups in a 50-year process of forest encroachment into semi-natural Mediterranean grasslands. A set of sampling plots of persistent grasslands and forests and their transitional habitat (wooded grasslands) was selected within fixed-area sites distributed across the landscape. Our results confirm the extinction debt and suggest colonization credit (according to observed trends and model predictions) in wooded grasslands when compared to persistent forests, despite wooded grasslands and persistent forests having similar tree cover. Grassland connectivity and solar radiation had opposing effects on the richness of both grassland and forest specialists, and it is possible that the availability of seed sources from old forests may have accelerate the payment of colonization credit in the wooded grasslands. These results suggest that extinction debt and colonization credit have driven species turnover during the 50 years of forest encroachment, but at different rates, and that local and landscape factors have opposing effects on these two phenomena. They also highlight the importance of documenting biodiversity time lags following habitat change when they are still in progress in order to timely and adequately manage habitats of high conservation value such as the grasslands studied here.

Local extinction of species can occur with a substantial delay following habitat loss or degradation. Accumulating evidence suggests that such extinction debts pose a significant but often unrecognized challenge for biodiversity conservation across a wide range of taxa and ecosystems. Species with long generation times and populations near their extinction threshold are most likely to have an extinction debt. However, as long as a species that is predicted to become extinct still persists, there is time for conservation measures such as habitat restoration and landscape management. Standardized long-term monitoring, more high-quality empirical studies on different taxa and ecosystems and further development of analytical methods will help to better quantify extinction debt and protect biodiversity.