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Sporobolus cryptandrus is a C4 perennial bunchgrass native to extensive areas of North America. As a non-native species, it has been reported from several continents, and it has been described as a transformer species in sand steppes of Central and Eastern Europe. However, its spreading ability across the landscape and within habitats has not been quantified, and factors determining its success have not yet been assessed. In this study, we focused on the largest stronghold of S. cryptandrus invasion in Hungary, where the species was first recorded in 2016, and investigated its present distribution in the landscape by mapping along dirt roads. In a separate local study in a heavily infested sand dune site of 2 km2, we assessed the infestation level and factors affecting the species' establishment. Our landscape-scale mapping found that in April 2023, the distribution of S. cryptandrus encompassed a largely contiguous 600 km2, with documented presence from 282 1-km2 mapping units. The species occurred more than 5 m away from roads in 71 mapping units, mostly in the centre of its distribution area. Sporobolus cryptandrus presence was negatively related to soil organic matter content and positively related to sand content. At the local scale, we found the species in 39% of vegetation plots in a sand dune site originally covered by Pannonic sand steppes, a priority habitat in the EU Habitats Directive. Sporobolus cryptandrus presence at this site was negatively related to the total cover of resident grassland but, surprisingly, was unrelated to the distance from roads. Collectively, these results suggest that primary spread occurs mostly along roads; these roadside populations likely serve as sources for establishment in neighbouring grasslands, but subsequent mass invasion becomes independent from roads. Together with the species' broad macroclimatic tolerance in North America and reported mass invasion events in Ukraine and Russia, our results suggest that S. cryptandrus likely poses a broad-scale threat to Eurasian dry grasslands, in particular on coarse-textured sandy soils with low vegetation cover. [ABSTRACT FROM AUTHOR]

Background: Burrowing mammals are important ecosystem engineers, especially in open ecosystems where they create patches that differ from the surrounding matrix in their structure or ecosystem functions. Methods: We evaluated the fine-scale effects of a subterranean ecosystem engineer, the Lesser blind mole rat on the vegetation composition of sandy dry grasslands in Hungary. In this model system we tested whether the characteristics of the patch (mound size) and the matrix (total vegetation cover in the undisturbed grassland) influence the structural and functional contrasts between the mounds and the undisturbed grasslands. We sampled the vegetation of 80 mounds and 80 undisturbed grassland plots in four sites, where we recorded the total vegetation cover, and the occurrence and cover of each vascular plant species. We used two proxies to characterise the patches (mounds) and the matrix (undisturbed grassland): we measured the perimeter of the mounds and estimated the total vegetation cover of the undisturbed grasslands. First, we compared the vegetation characteristics of the mounds and the surrounding grasslands with general linear models. Second, we characterised the contrasts between the mounds and the undisturbed grassland by relative response indices (RRIs) of the vegetation characteristics studied in the first step. Results: Species composition of the vegetation of the mounds and undisturbed grasslands was well separated in three out of the four study sites. Mounds were characterised by lower vegetation cover, lower cover of perennial graminoids, and higher diversity, and evenness compared to undisturbed grasslands. The contrast in vegetation cover between mounds and undisturbed grasslands increased with decreasing patch size. Increasing vegetation cover in the matrix grasslands increased the contrasts between the mounds and undisturbed grasslands in terms of total cover, perennial graminoid cover, diversity, and evenness. Our results suggest that mole rat mounds provide improved establishment conditions for subordinate species, because they are larger than other types of natural gaps and are characterised by less intense belowground competition. The ecosystem engineering effect, i.e., the contrast between the patches and the matrix was the largest in the more closed grasslands. [ABSTRACT FROM AUTHOR]

Background: Soil seed banks play a central role in vegetation dynamics and may be an important source of ecological restoration. However, the vast majority of seed bank studies examined only the uppermost soil layers (0-10 cm); hence, our knowledge on the depth distribution of seed bank and the ecological significance of deeply buried seeds is limited. The aim of our study was to examine the fine-scale vertical distribution of soil seed bank to a depth of 80 cm, which is one of the largest studied depth gradients so far. Our model systems were alkaline grasslands in EastHungary, characterised by harsh environmental conditions, due to Solonetz soil reference group with Vertic horizon. We asked the following questions: (1) How do the seedling density and species richness of soil seed bank change along a vertical gradient and to what depth can germinable seeds be detected? (2) What is the relationship between the depth distribution of the germinable seeds and the species traits? Methods: In each of the five study sites, four soil cores (4 cm diameter) of 80 cm depth were collected with an auger for soil seed bank analysis. Each sample was divided into sixteen 5-cm segments by depth (320 segments in total). Samples were concentrated by washing over sieves and then germinated in an unheated greenhouse. Soil penetration resistance was measured in situ next to each core location (0-80 cm depth, 1-cm resolution). We tested the number and species richness of seedlings observed in the soil segments (N = 320), using negative binomial generalized linear regression models, in which sampling layer and penetration resistance were the predictor variables. We ran the models for morphological groups (graminoids/forbs), ecological groups (grassland species/weeds) and life-form categories (short-lived/perennial). We also tested whether seed shape index, seed mass, water requirement or salt tolerance of the species influence the vertical distribution of their seed bank. Results: Germinable seed density and species richness in the seed bank decreased with increasing soil depth and penetration resistance. However, we detected nine germinable seeds of six species even in the deepest soil layer. Forbs, grassland species and short-lived species occurred in large abundance in deep layers, from where graminoids, weeds and perennial species were missing. Round-shaped seeds were more abundant in deeper soil layers compared to elongated ones, but seed mass and ecological indicator values did not influence the vertical seed bank distribution. Our research draws attention to the potential ecological importance of the deeply buried seeds that may be a source of recovery after severe disturbance. As Vertisols cover 335 million hectares worldwide, these findings can be relevant for many regions and ecosystems globally. We highlight the need for similar studies in other soil and habitat types to test whether the presence of deep buried seeds is specific to soils with Vertic characteristics. [ABSTRACT FROM AUTHOR]

Aims: We tested the establishment success of grassland species in the presence or absence of zoochory by livestock and identified the traits associated with successful establishment. Location: Hortobágy National Park, Hungary. Methods: In six restored species-poor grasslands we established two species-rich 4 m × 4 m source plots per site in 2013. One source plot was managed by extensive cattle grazing (“zoochory on”), the other one was fenced and protected from grazers (“zoochory off”). We monitored the vegetation development in the source plots (2014–2018), and the establishment success of sown species (2016–2018) in adjacent dispersal plots in four, 20-m-long strips per source plot in a total of 960 dispersal plots. We calculated community-weighted means of thirteen plant traits related to regeneration and competitive ability. The effect of grazing, year, distance from source plots, grazing–year and grazing–distance interactions on population and trait dynamics in the source and dispersal plots were analysed with generalized linear mixed-effect models. Results: Although grazing reduced the cover and flowering success of sown species in the source plots, the number of successful establishment events was higher in dispersal plots next to grazed source plots. The sown species were the most abundant in plots adjacent to source plots, but occurred at all monitored distances. Zoochory favoured perennial plants with high specific leaf area (SLA), low leaf dry matter content (LDMC), short stature, light seeds, early flowering period, high clonal index and anemochory index. Conclusions: We showed that species that can colonize the matrix with and without zoochory have contrasting trait syndromes. In grazing-adapted ecosystems, fencing source populations of target species does not always seem to be a good option. It was a good tool for increasing the cover and flowering success of the sown species in the short run, but it significantly decreased their chance to colonize the surrounding area in the long run. [ABSTRACT FROM AUTHOR]

Sowing grasses supports the rapid development of a closed perennial vegetation, which makes the method universally suitable for fast and effective landscape‐scale restoration of grasslands. However, to increase their diversity and to create a natural‐like species‐rich grassland is a challenging task. Understanding the role of seed bank compositional changes and vegetation dynamics can help to design management regimes that support the establishment of target species and suppress unwanted weeds. Our aim was to reveal the effect of post‐restoration management on the vegetation and seed bank dynamics in grasslands restored in one of the largest European landscape‐scale restoration projects. Eight years after restoration, we sampled the vegetation and the seed bank in 96 quadrats located in 12 restored grasslands in the Great Hungarian Plain. In each grassland stand, we studied and compared a mown (mown from Year 1 to Year 8) and an abandoned plot (mown from Year 1 to Year 3 then abandoned from Year 4 to Year 8). Mown and abandoned plots showed divergent vegetation and seed bank development. Abandonment led to the decline of sown grasses and higher cover of weeds, especially in the alkaline grasslands. Our study underlined that the developing seed bank had a limited contribution to the maintenance of biodiversity in both grassland types. We found that 5 years of abandonment had a larger effect on the seed bank than on the vegetation. We stress that long‐term management is crucial for controlling the emergence of the weeds from their dense seed bank in restored grasslands. [ABSTRACT FROM AUTHOR]

Sowing of grass seed mixtures is a feasible and cost‐effective method for landscape‐scale grassland restoration. However, sowing only grasses usually leads to species‐poor and dense swards, where the establishment of target forbs is hampered by microsite and propagule limitation. To overcome these limitations and increase the diversity of species‐poor sown grasslands, we developed a novel method by creating "establishment gaps.". We used tillage to open gaps of 1‐, 4‐, and 16‐m2 size in the dense grass sward of six species‐poor restored grasslands in the Great Hungarian Plain. We sowed high‐diversity seed mixtures of 35 native species into all gaps. We analyzed vegetation development during the first 5 years after setting up the trial. We also studied the colonization dynamics of the sown species along four 20‐m transects around each gap, resulting in a total of 1440 plots of 1‐m2 size that were studied. Our results indicated that most of the sown species were able to establish permanently in the gaps. The total cover and the cover of perennial sown species increased and the cover of short‐lived sown species decreased independent of gap size. There was only a moderate level of weed abundance in the gaps, and weed cover decreased over the years. The sown target species started to colonize the species‐poor grasslands surrounding the gaps within 5 years. The highest number of species and individuals dispersed from the 4‐m2 gaps, as they had a more stable development than smaller gaps and were exposed to lower grazing pressure than large ones. [ABSTRACT FROM AUTHOR]

In intensively used landscapes biodiversity is often restricted to fragmented habitats. Exploring the biodiversity potential of habitat fragments is essential in order to reveal their complementary role in maintaining landscape-scale biodiversity. We investigated the conservation potential of dry grassland fragments in the Great Hungarian Plain, i.e. patch-like habitats on ancient burial mounds and linear-shaped habitats in verges, and compared them to continuous grasslands. We focused on plant taxonomic diversity, species richness of specialists, generalists and weeds, and the phylogenetic diversity conserved in the habitats. Verges meshing the landscape are characterised by a small core area and high level of disturbance. Their species pool was more similar to grasslands than mounds due to the lack of dispersal limitations. They held high species richness of weeds and generalists and only few specialists. Verges preserved only a small proportion of the evolutionary history of specialists, which were evenly distributed between the clades. Isolated mounds are characterised by a small area, a high level of environmental heterogeneity, and a low level of disturbance. Steep slopes of species accumulation curves suggest that high environmental heterogeneity likely contributes to the high species richness of specialists on mounds. Mounds preserved the same amount of phylogenetic diversity represented by the branch-lengths as grasslands. Abundance-weighted evolutionary distinctiveness of specialists was more clustered in these habitats due to the special habitat conditions. For the protection of specialists in transformed landscapes it is essential to focus efforts on preserving both patch-like and linear grassland fragments containing additional components of biodiversity. [ABSTRACT FROM AUTHOR]

Agricultural intensification has resulted in severe declines in the extent and diversity of seminatural habitats in Europe, whereas the extent of secondary habitats has increased considerably. River embankments have become one of the most extensive and widespread secondary habitats in former floodplains. We compared the diversity patterns of secondary dry and wet grasslands on river embankments with those of seminatural dry and wet grasslands in a Hungarian agricultural landscape using the following community descriptors: (a) species diversity, (b) phylogenetic diversity and (c) functional diversity. We also performed trait‐based analyses to evaluate the ecosystem services provided by these secondary grasslands. Both grassland types of the embankments showed significantly higher Shannon diversity compared with their seminatural counterparts. The cover of generalist species (i.e., cosmopolitan species, weeds and nonindigenous plant species) was high in the secondary grasslands. We found significant differences in phylogenetic diversity between the secondary and seminatural grasslands: secondary grasslands showed significantly lower mean nearest taxon distances than the seminatural grasslands. Functional diversity did not differ between the secondary and seminatural grasslands according to the Rao's quadratic entropy. However, we found higher community‐weighted means of specific leaf area, plant height and flowering period in the secondary grasslands, which are related to important ecosystem services (via biomass production and pollination). Well‐planned management actions and restoration activities could help further improve the ecological function and conservation value of secondary grasslands on river embankments, contributing to the maintenance of species diversity and sustaining the functionality of ecosystems in agricultural landscapes. [ABSTRACT FROM AUTHOR]

Questions: Sustainable rangeland utilization considering traditions and economic reasons is compulsory for harmonising the needs of the agricultural and nature conservation sectors. For proper rangeland management it is crucial to compare the grazing effects of traditional breeds and crossbred animals of the same species that might have different effects on the rangelands. To fill this knowledge gap, in a grazing experiment, we investigated the effect of cattle breeds on the vegetation to test the effects on nature conservation value and agricultural production value. We hypothesized that the effects of cattle grazing on habitat conservation values and forage quality depend on the grazing breed, because breeds differ in selectivity, body size and trampling effect. Location: Marshes and alkaline wet grasslands in Hortobágy National Park, Hungary. Methods: We recorded the percentage cover of vascular plants in three consecutive years in a total of 60 plots in 12 areas grazed by traditional (0.61 AU/ha) and large‐sized crossbred beef cattle (0.68 AU/ha). Results: We found that the effect of cattle breed on the habitat conservation values and forage quality is dependent on the habitat type. The traditional breed maintained a significantly higher species number and Shannon diversity in marshes than the crossbred beef cattle. Grazing of crossbred cattle led to decreasing moisture indicator values in marsh habitats. Conclusions: Our findings revealed that traditional breeds should be prioritized in the management of wet alkaline grasslands and marshes. Crossbred beef cattle might be a substitute but only in case traditional breeds are not available for the management of alkaline wet grasslands. In marshes, however, we recommend prioritizing the traditional breeds as they maintain higher diversity compared to crossbred beef cattle. [ABSTRACT FROM AUTHOR]

Question: Spontaneous succession is the most natural and cost‐effective solution for grassland restoration. However, little is known about the time required for the recovery of grassland functionality, i.e., for the recovery of reproductive and vegetative processes typical of pristine grasslands. Since these processes operate at different scales, we addressed the question: do reproductive and vegetative processes require different recovery times during spontaneous succession? Location: Kiskunság sand region (Central Hungary). Methods: As combinations of plant traits can be used to highlight general patterns in ecological processes, we compared reproductive (pollination‐ and dispersal‐related) and vegetative (growth form) traits between recovered grasslands of different age (<10 years old; 10–20 years old; 20–40 years old) and pristine grasslands. Results: During spontaneous succession, the reproductive trait spectra became similar to those of pristine grasslands earlier than the vegetative ones. In arable land abandoned for 10 years, pollination‐ and dispersal‐related trait spectra did not show significant difference to those of pristine grasslands; anemophily and anemochory were the prevailing strategies. Contrarily, significant differences in the growth form spectrum could be observed even after 40 years of abandonment; in recovered grasslands erect leafy species prevailed, while the fraction of dwarf shrubs and tussock‐forming species was significantly lower than in pristine grasslands. Conclusions: The recovery of the ecological processes of pristine grasslands might require different amounts of time, depending on the spatial scale at which they operate. The reproductive trait spectra recovered earlier than the vegetative one, since reproductive attributes first determine plant species sorting at the regional level towards their respective habitats. The recovery of the vegetative trait spectrum needs more time as vegetative‐based interactions operate on a smaller spatial scale. Thus, vegetative traits might be more effective in the long‐term assessment of restoration success than the reproductive ones. [ABSTRACT FROM AUTHOR]

Spontaneous succession is increasingly involved in grassland restoration, because it offers a cost‐effective solution compared to technical reclamation methods. This topic is especially important nowadays, as large areas of marginal croplands are being abandoned on poor soils of Central and Eastern Europe, which offers a possibility for the spontaneous recovery of typical target grasslands. Studying the vegetation composition, aboveground biomass, and seed bank in old fields of different age and target calcareous sand grasslands using the chronosequence method, we aimed to answer the following questions: (1) Which species contribute to the seed banks of old fields and reference grasslands? (2) Does the direction of vegetation and seed bank succession trend toward the reference grasslands? (3) How are the vegetation changes in spontaneous succession reflected by the soil seed banks of old fields? In reference grasslands on the dune tops only sporadic seed banks were detected, while several hygrophytes had dense seed banks in reference grasslands in dune slacks. Similarity between the species composition of vegetation and seed banks was low. The development of vegetation and seed banks in old fields progressed toward that of target grasslands and the proportion of weedy species (e.g. indigenous weeds and invasive species) also decreased with time. The cryptogam biomass correlated significantly negatively, while the soil phosphorus significantly positively with the weedy species seed bank density. Our results indicated that the role of persistent seed banks in the regeneration of calcareous sand grasslands from old fields is rather limited and promising vegetation changes are mostly driven by spatial dispersal. [ABSTRACT FROM AUTHOR]

Aims Cyanobacteria may have considerable effects on community functioning, mostly because they produce various metabolites that adversely affect other organisms. Here we synthesized existing knowledge about the effects of toxic cyanobacteria on the germination and growth of terrestrial plants. We also aimed to test the chemical effects of a Nostoc (Cyanobacteria) extract on the germination and growth of species of alkali habitats to investigate whether cyanobacteria can alter community structure and diversity via affecting the establishment success of plants. Location Cyanobacterium colonies from the Hortobágy National Park, east Hungary; indoor experiments at the University of Debrecen, Hungary. Methods To review the effects of toxic cyanobacteria on terrestrial plants, we conducted a literature search. To test these effects on native plants, field-collected Nostoc colonies were used to prepare a cell-free water extract, and treatments (watering with Nostoc extract and watering with tap water) were tested on 3 × 100 seeds of nine alkali grassland species. After 5 wk, seedling number, seedling length and fresh and dry weights were measured. Results We collected data on the effects of cyanobacteria on 27 species, but they were mostly focused on crops irrigated with cyanobacteria-containing water, not on floras native to natural ecosystems. In the germination experiment species identity and treatment had a significant effect on almost all variables, but their interaction only affected germination rate and fresh weight. Fresh weight decreased significantly only in the invasive Hordeum jubatum, but germination rate decreased significantly in five species. Conclusions Based on our findings, terrestrial cyanobacterium colonies can affect the establishment success of grassland plants, through which they may be important in determining which species can be incorporated into the community. Thus, cyanobacteria might play an important role in shaping diversity, species composition and the structure of natural plant communities. [ABSTRACT FROM AUTHOR]

Classical old-field succession studies focused on vegetation changes after the abandonment of annual croplands or on succession after the elimination of cultivated crops. Perennial-crop-mediated succession, where fields are initially covered by perennial crops, reveals alternative aspects of old-field succession theories. We tested the validity of classical theories of old-field succession for perennial-crop-mediated succession. We formulated the following hypotheses: (1) functional diversity increases with increasing field age; (2) resource acquisition versus conservation trade-off shifts toward conservation at community level during the succession; (3) the importance of spatial and temporal seed dispersal decreases during the succession; and (4) competitiveness and stress-tolerance increases and ruderality decreases at community level during the succession. We studied functional diversity, trait distributions and plant strategies in differently aged old-fields using chronosequence method. We found increasing functional richness and functional divergence, but also unchanged or decreasing functional evenness. We detected a shift from resource acquisition to resource conservation strategy of communities during the succession. The role of spatial and temporal seed dispersal was found to be important not only at the initial but also at latter successional stages. We found an increasing stress-tolerance and a decreasing ruderality during succession, while the competitiveness remained unchanged at the community level. Despite the markedly different starting conditions, we found that classical and perennial-crop-mediated old-field successions have some similarities regarding the changes of functional diversity, resource acquisition versus conservation trade-off, and seed dispersal strategies. However, we revealed also the subsequent differences. The competitive character of communities remained stable during the succession; hence, the initial stages of perennial-crop-mediated succession can be similar to the middle stages of classical old-field succession. Moreover, the occupied functional niche space and differentiation were larger in the older stages, but resources were not effectively utilized within this space, suggesting that the stabilization of the vegetation requires more time. [ABSTRACT FROM AUTHOR]

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Facilitation is an important driver of community assembly, and often overwhelms the effect of competition in stressed habitats. Thus, net effect of biotic interactions is often positive in stressed grasslands, where dominant species and litter can protect the subordinate species. Besides facilitation, niche partitioning can also support species coexistence leading to limiting similarity between subordinate species. Our aim was to provide a detailed analysis of fine-scale biotic interactions in stressed alkali grasslands. We supposed, that there are positive relationships between the main biomass fractions and species richness. We expected the expansion of trait ranges and the increase of trait dissimilarity with increasing biomass scores (total litter, green biomass of dominant species) and species richness. We studied the relationships between main biomass fractions, species richness, functional diversity and functional trait indices (ranges, weighted means and Rao indices). We used fine-scale biomass sampling in nine stands of dry alkali grasslands dominated by Festuca pseudovina . The detected relationships were always positive between the main biomass fractions (green biomass of dominant species, total litter and green biomass of subordinate species) and species richness. We found that the green biomass of dominant species and total litter increased ranges and dissimilarity of functional traits. Our results suggest that in dry alkali grasslands facilitation is crucial in shaping vegetation composition. The green biomass of dominant species and total litter increased the biomass production of subordinate species leading to overyielding. We found that mechanisms of facilitation and limiting similarity were jointly shaping the species coexistence in stressed grasslands, such as alkali grasslands. [ABSTRACT FROM AUTHOR]

Grasslands recovered by sowing low diversity seed mixtures of local provenance are usually managed by mowing. Besides restoration success only a few studies have focused on the direct effects of post-restoration mowing on recovered grassland vegetation. In this study we followed vegetation changes in 13 successfully recovered grasslands in 5 × 5-m-sized sites with continuous and ceased mowing at Hortobágy National Park, East-Hungary. We asked the following questions: (i) What are the effects of cessation of mowing on the vegetation structure and diversity of recovered grasslands? (ii) What are the effects of cessation of mowing on the abundance of sown grasses, target and undesirable species? (iii) Is yearly mowing an appropriate management tool for the maintenance of recovered grasslands? Our results showed that the cessation of mowing caused litter accumulation, while diversity, total vegetation cover and the cover of sown grasses decreased compared to the mown sites. The cover of undesirable perennial species was significantly higher in unmown sites than in mown ones. The species composition of mown sites remained more similar to near-natural grasslands than the unmown ones. Our results suggest that without regular post-restoration mowing the favourable status of recovered grasslands can rapidly decline due to litter accumulation and by the expansion of undesirable species, even in the short-run. We also stress that while yearly mowing is enough to maintain grasslands recovered by low-diversity seed sowing, it cannot be considered to be enough to recover target vegetation composition. [ABSTRACT FROM AUTHOR]

Question Explaining the biomass-species richness relationship is key to understanding vegetation dynamics. Several possible mechanisms have been suggested, but complex analysis of plant strategies, major biomass and species richness components along a long productivity gradient is still lacking. We provide a detailed analysis of the relationship between major biomass components (total above-ground biomass, green biomass and litter), plant strategies and species richness along a long gradient of alkali and loess grasslands in a steppe landscape in Central Europe. Location Hortobágy, Great Hungarian Plain, East Hungary. Methods Above-ground biomass of characteristic alkali and loess grassland stands was sampled along a gradient of increasing productivity. In each grassland stand, a 25-m2 sample site was randomly selected. Within each site, ten above-ground biomass samples (20 × 20 cm) were collected randomly in June 2009, at the peak of biomass production. We classified all species into mixed C-S-R strategy types. To obtain correlations between various biomass and species richness data, Spearman rank correlation was used. The relationship between plant strategies and species composition were displayed with a DCA ordination. Results The frequently detected humped-back relationship was valid for the relation of total biomass and species richness. With increasing amount of total biomass, we detected an increasing proportion of competitors, and a decreasing proportion of stress tolerators in green biomass. A low proportion of ruderals was detected at both low and high biomass levels. Species richness was affected positively by litter at low litter scores, but there was a negative litter effect from much lower scores than detected previously (from 400 g·m−2). There was a positive relationship between green biomass production and species richness. Conclusions The study revealed that at the initial part of a productivity gradient, stress is likely responsible for low species richness. Our results show that litter can shape changes in species richness along the whole biomass gradient, thus the litter effect is one of the major mechanisms structuring grassland diversity. [ABSTRACT FROM AUTHOR]

Spontaneous succession is often underappreciated in restoration after the cessation of intensive agricultural management. Spontaneous succession could improve the success of restoration programmes, and offers a cost-effective option with little active intervention. We studied the spontaneous recovery of loess grasslands in extensively managed lucerne Medicago sativa fields mown twice a year using space for time substitutions to highlight the importance of spontaneous processes in grassland restoration. With increasing field age a gradual replacement of lucerne by perennial native grasses and forbs and increase of mean species richness was detected. As the age of fields increased, lucerne decreased from 75% to 2% of total vegetation cover, whereas perennial graminoids increased from 0·5 to 50% cover. Mean total cover showed no significant differences between the age groups; weed cover was less than 10%. The phytomass of lucerne was negatively correlated with graminoid phytomass. As the age of the fields increased, lucerne phytomass decreased and grass phytomass increased. We found a negative correlation between litter and forb phytomass but there was no relationship with the age of the field. There was no litter accumulation and no increase of mean total phytomass as the age of fields increased. Synthesis and applications. Native grasses within loess grasslands recovered within 10 years, but characteristic native forbs remained rare. The advantages of spontaneous succession in lucerne fields compared to technical reclamation include: (i) no early stages dominated by weeds, (ii) minimal litter accumulation, (iii) a spontaneous decrease in lucerne over time, and (iv) negligible cost. In addition, the requirement for twice yearly mowing in the early years will guarantee farmer involvement because of the high forage value of lucerne. The complete restoration of species rich grasslands will require more active management such as propagule transfer by hay and/or moderate grazing to encourage the return of native forbs. [ABSTRACT FROM AUTHOR]

Regulation of plant biomass accumulation is a key issue in effective grassland conservation in Europe. Burning is an alternative tool to regulate biomass dynamics in semi-natural grasslands even in the absence of grazing or mowing. We tested the effects of regular spring burning on the biomass fractions and fine-scale plant species composition of species-rich foothill grasslands in Aggtelek National Park, North-Hungary. There were five regularly burned and five control grasslands in the study; we collected twenty 20×20-cm sized biomass samples from each. We analyzed the main fractions (litter, graminoid and forb biomass), and the species-level biomass scores, and flowering success in the control and burned grasslands. We revealed that fire increased the amount of forb biomass and decreased the amount of litter, which suggested that regular burning might be feasible for regulating biomass dynamics. The non-metric multi-dimensional scaling (NMDS) showed a high similarity of the control and burned grasslands in species composition. However, plant diversity, and the number of flowering shoots decreased significantly in the burned grasslands. In regularly burned sites we found a significant decline of specialist species, as well as of steppic flora elements. Our results showed that besides its positive effect on biomass dynamics, high-frequency burning threatens the overall diversity and specialist plant species in semi-natural grasslands. We recommend that proper fire regimes should be first studied experimentally, to provide a scientific basis for the application of prescribed burning management in such habitats. [ABSTRACT FROM AUTHOR]

Highlights: [•] Seed banks play an important role sustaining overall grassland biodiversity. [•] We studied vegetation and seed banks in differently elevated alkali grasslands. [•] The lowest seed density was found in the most stressed grassland type. [•] Characteristic grasses and forbs possessed at most sparse seed banks. [•] Rapid vegetation shifts are not likely driven by persistent seed banks. [ABSTRACT FROM AUTHOR]

Open landscapes in many parts of Europe have been negatively affected by large-scale drainage and amelioration to support agricultural production. In continental alkali grasslands, amelioration and establishment of drainage ditch systems were typical in the 1950s and 60s. Drainage ditches caused a considerable fragmentation and degradation of natural grasslands; thus several projects aimed at eliminating these linear landscape elements. In a multi-site study, we explored the drivers of grassland recovery after soil-filling of drainage ditches in landscape-scale restoration projects in Hortobágy National Park, East-Hungary. Ditch embankments, formerly built from the excavated soil, were used to fill the 8-m wide ditches and grazing was applied to facilitate the recovery of grasslands similar to the surrounding matrix. Three age classes were selected for the study: 1-, 6- and 8-year-old filled ditches; with nine sites per age group, surrounded by three grassland types (27 ditches in total). We recorded the percentage cover of vascular plant species in 18 plots per ditch, 486 plots in total. We found that the species pool of the filled ditches became more similar to the reference grasslands with increasing successional age and increasing distance to the central zone of the ditches regardless of grassland type. Species richness of the filled ditches became more similar to that of the reference grasslands with increasing successional age. However, we found that several target species, especially salt-tolerant pioneers, could establish even in the first year. Grassland recovery was most successful in sites adjacent to dry grasslands characterised by soils with high salt content, which favoured specialist species and suppressed non-target species. Cover of non-target species was higher in wet meadows with moist, nutrient-rich soils which favoured generalists and non-target species. Our study revealed that passive restoration after soil filling of disused drainage ditches can effectively support grassland recovery even within less than ten years, when restoration sites are surrounded by natural grasslands. [ABSTRACT FROM AUTHOR]

Abstract: Technical reclamation of grasslands is a powerful tool for conserving biodiversity in fragmented landscapes. Plant material transfer and sowing seeds of local provenance are used most frequently to recover grassland vegetation in former croplands. The joint application of these methods is rarely used, although it has the potential to gain a predictable and directed vegetation development with effective early weed suppression. We studied the effectiveness of combining low diversity seed sowing and hay-transfer in weed suppression and recovery of perennial grassland vegetation in Hortobágy Puszta, Central-Europe, by testing the following hypotheses: (i) lower weed cover and biomass is expected in vegetation recovered by the joint method of sowing and hay transfer than by seed sowing only. (ii) With sowing and additional hay transfer a higher rate of establishment of Festuca species is expected than with sowing only. Our results supported both hypotheses. We found that the additional application of hay significantly accelerated the development of perennial grassland vegetation and provided a higher weed suppression rate in the first year and onwards in most plots than seed sowing only. A higher establishment rate was detected both in the cover and the biomass of perennial grasses including Festuca species in all plots with hay addition than with sowing only. Our results suggest that a combination of hay transfer and low diversity sowing may provide a cost-effective alternative to the more costly high-density sowing and if proper sources for high-diversity hay are available, it may replace high-diversity seed mixtures. [Copyright &y& Elsevier]

A decrease in habitat amount and connectivity causes immediate or delayed species extinctions in transformed landscapes due to reduced functional connections among populations and altered environmental conditions. We assessed the effects of present and historical grassland amount and connectivity as well as local habitat factors typical of the present landscape on the current species richness of grassland specialist and generalist plants and arthropods in grassland fragments. We surveyed herbaceous plants, ants, orthopterans, true bugs and rove beetles in 60 dry grassland fragments in Hungary. We recorded the area of the focal grassland, the slope and the cover of woody vegetation. By using habitat maps of the present and historical landscape, we calculated grassland amount and connectivity for four time periods covering 158 years (1858–2016). We found evidence for an unpaid extinction debt in specialist plants, suggesting that they have not come to equilibrium with the grassland amount and connectivity of the present landscape yet. This localised and typically long-lived group responded slowly to the landscape changes. Specialist arthropod taxa with short generation times responded much faster to habitat loss than plants and did not show an extinction debt. Generalist plants and animals adapted to a wide range of habitats were affected by the landscape-scale decline of grassland habitats. Despite decreased habitat connectivity, grassland fragments with dry environmental conditions and high environmental heterogeneity can sustain specialist plants in transformed landscapes. Unpaid extinction debt should be considered an early warning signal: Restoration of grassland connectivity is necessary to halt ongoing extinction processes. [ABSTRACT FROM AUTHOR]