Slash Pile Burning Effects on Soil Biotic and Chemical Properties and Plant Establishment: Recommendations for Amelioration

Ponderosa pine forest restoration consists of thinning trees and reintroducing prescribed fire to reduce unnaturally high tree densities and fuel loads to restore ecosystem structure and function. A current issue in ponderosa pine restoration is what to do with the large quantity of slash that is created from thinning dense forest stands. Slash piling burning is currently the preferred method of slash removal because it allows land managers to burn large quantities of slash in a more controlled environment in comparison with broadcast burning slash. However burning slash piles is known to have adverse effects such as soil sterilization and exotic species establishment. This study investigated the effects of slash pile burning on soil biotic and chemical variables and early herbaceous succession on burned slash pile areas. Slash piles were created following tree thinning in two adjacent approximately 20‐ha ponderosa pine (Pinus ponderosa) restoration treatments in the Coconino National Forest near Flagstaff, Arizona. We selected 30 burned slash pile areas and sampled across a gradient of the burned piles for arbuscular mycorrhizal (AM) propagule densities, the soil seed bank, and soil chemical properties. In addition, we established five 1‐m² plots in each burned pile to quantify the effect of living soil (AM inoculum) and seeding amendments on early herbaceous succession in burned slash pile areas. The five treatments consisted of a control (no treatment), living soil (AM inoculum) amendment, sterilized soil (no AM inoculum) amendment, seed amendment, and a seed/soil (AM inoculum) amendment. Slash pile burning nearly eliminated populations of viable seeds and AM propagules and altered soil chemical properties. Amending scars with native seeds increased the cover of native forbs and grasses. Furthermore adding both seed and living soil more than doubled total native plant cover and decreased ruderal and exotic plant cover. These results indicate that seed/soil amendments that increase native forbs and grasses may enhance the rate of succession in burned slash pile areas by allowing these species to outcompete exotic and ruderal species also establishing at the site through natural regeneration.     

Sustaining recovered grasslands is not likely without proper management: vegetation changes after cessation of mowing

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.     

Effects of litter on seedling establishment: an indoor experiment with short-lived Brassicaceae species

Recruitment by seeds is essential both in vegetation dynamics and in supporting biodiversity in grasslands. The recruitment by seeds is feasible in suitable vegetation gaps from the seed rain and/or by establishment from persistent soil seed banks. Cessation of grassland management results in litter accumulation, which leads to the decline of species diversity by the decreased availability of open patches. Low amounts of litter is often beneficial, while high amounts of litter is detrimental for seed germination and seedling establishment of short-lived species. In a designed indoor experiment, we explored the effect of litter on seedling establishment by germinating six short-lived Brassicaceae species with both increasing seed mass and litter cover. We found that both seed mass and litter had significant effect on germination and establishment of the sown species. Small-seeded species were significantly negatively affected by the 300 and/or 600 g/m² litter layers. No negative litter effect was detected for species with high seed masses (Lepidium spp.). No overall significant positive litter effect was found, although for most of the species cumulative seedling numbers were not the highest at the bare soil pots. Our results suggest that the negative effects of litter are less feasible on the large-seeded short-lived species than on that of small-seeded ones.     

Response of native and non-native ruderals to natural and human disturbance

The ruderal strategy is widely shared among non-native plants, providing a general explanation for the commonly observed positive effects of disturbance on invasions. How native ruderals respond to disturbance and how their abundance compares to that of non-native ruderals remains, however, poorly understood. Similarly, little is known about the role that disturbance type plays in the coexistence between native and non-native ruderals. We proposed that natural disturbance favors native over non-native ruderals, whereas novel anthropogenic disturbance favors non-natives over natives. To assess our general hypothesis, we conducted extensive field samplings in which we measured relative abundance, richness, and diversity of native and non-native ruderals in sites with natural and anthropogenic disturbance in central Argentina, a system where the ruderal strategy is common to a large number of native and non-native species. We found that natives dominated ruderal communities growing in recently burned grasslands, whereas non-natives dominated in roadsides. Additionally, the richness and diversity of native ruderal species were much greater than those of non-natives in sites with fire and in sites with grazing, but species richness and diversity did not differ between groups in roadsides. Because vegetation evolved with fire in our system and, in contrast, the construction and maintenance of roads is recent in it, these results support our hypothesis. Our work indicates that the ruderal strategy does not seem to suffice to explain why disturbance facilitates invasions. According to our data, species origin interacts with disturbance type to determine dominance in communities with coexisting native and non-native ruderals.     

Litter effects on seedling establishment interact with seed position and earthworm activity

Seedling establishment is influenced by litter cover and by seed predators, but little is known about interactions between these two factors. We tested their effects on emergence of five typical grassland species in a microcosm experiment. We manipulated the amounts of grass litter, seed sowing position and earthworm activity to determine whether: (i) the protective effect of litter against seed predation depends on cover amount and seed sowing position, i.e., on top or beneath litter; (ii) seed transport by earthworms changes the effect of seed sowing position on seedling emergence; and (iii) seeds transported into deeper soil layers by earthworms are still germinable. Litter cover and presence of earthworms lowered seedling emergence. The impact of seed position increased with seed size. Emergence of large-seeded species was reduced when sown on the surface. Additionally, we found an important seed position × earthworm interaction related to seed size. Emergence of large-seeded species sown on top of the litter was up to three times higher when earthworms were present than without earthworms. Earthworms also significantly altered the depth distribution of seeds in the soil and across treatments: on average 6% of seeds germinated after burial. In contrast to the seed position effect, we found no size effect on mobility and germinability of seeds after burial in the soil. Nevertheless, the fate of different-sized seeds may differ. While burial will remove large seeds from the regeneration pool, it may enhance seed bank build up in small-seeded species. Consequently, changes in the amount of litter cover and the invertebrate community play a significant role in plant community composition.     

Restoration of species‐rich flood‐plain meadows from abandoned arable fields in NE France

Abstract. Attempts to restore species‐rich flood‐plain meadows from abandoned arable fields in the valley of the river Meuse, NE France, were studied. The study area was sown with a commercial seed mixture, composed of Phleumpratense, Festuca pratensis, Lolium perenne and Trifolium repens. The above‐ground vegetation in the study area 1, 2 and 3 yr after restoration was compared to (1) the vegetation present during the previous 5‐yr fallow stage and (2) target flood‐plain meadows. Before restoration, the above‐ground fallow vegetation was dominated by ruderal and annual species, while only very few meadow species were present. Sowing led to tall, dense vegetation, mainly dominated by the sown species. Ruderal and annual species had decreased 3 yr after restoration, but target species were still poorly represented. Species richness was significantly lower in the sown site than in the semi‐natural target meadows and the vegetation had a different composition. Analysis of the soil seed bank of the restored meadow showed that only a few meadow species were present and that it was dominated by a few ruderal species. Three years after sowing, the vegetation of our experimental site is moving slowly towards the target communities but impoverished seed sources seem to limit the success of this restoration operation and will lead to under‐saturated communities.     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

死、活地被物对红松伴生树种天然更新影响的实验研究

1963年在小兴安岭丰林自然保护区红松林及有关的皆伐迹地上设置了61组,325个实验样方,定期观察到1965年,以研究死、活地被物对人工播种的红松林伴生树种种子发芽和幼苗初期存活、生长的影响。实验表明死、活地被物,特别是死地被物的干湿变化剧烈,影响幼苗的成活,甚至使种子或幼根无法接触土壤,因而种粒越小,或轻而带翅,受影响越大。此外本文还指出了桦树种子的强迫休眠和延迟发芽现象。讨论了实验结果在红松林演替和人工促进天然更新中的意义。     

Vegetation development after a large scale restoration of species-rich grasslands in a Central European floodplain

Large scale restoration using local high-diversity seed mixture combined with turf transfer was applied on ex-arable land in the Morava River floodplain in the western Slovakia in the years 1999–2012. The post-restoration vegetation development was recorded during 12 years after the restoration using floristic records per restored polygons with cover estimation in simple 3-degree scale. Temporal changes in species composition were evaluated by gradient analysis and number of characteristic grassland and ruderal species on restored sites was analysed by general linear models. Species composition changed gradually towards the species composition typical for species-rich floodplain grasslands, but the trajectory was not straightforward and several irregularities were observed. They were probably induced by extreme weather events (drought, floods). The decrease in ruderal species and increase in the number of typical floodplain grassland species were observed, when floodplain grassland species permanently outcompeted ruderal species since 8th year after the restoration. However the development in large scale was slower, than expected from previous small-scale experiments, it is evident, that combination of local seed mixture sowing with a turf transfer is a feasible method for the restoration of species-rich floodplain grasslands from arable land.     

Revegetation of Motorway Slopes Using Different Seed Mixtures

Motorway slopes cover large areas and could, if properly managed, partially compensate for the loss of natural habitats and connect remnant populations of target species. One of the main problems on such slopes is soil erosion; therefore, quick and stable revegetation is essential. Two widely used commercial seed mixtures were sown and tested to determine which one is better for restoration of motorway slopes. Both mixtures contained CSR strategists (“competitive stress‐tolerant ruderals”) but one included nurse species (Lolium perenne, L. multiflorum, Trifolium spp.), whereas the other did not. Both treatments were sampled every year over a period of 6 years and analyzed for species richness, diversity, similarity, and plant cover. The research took place in the sub‐Mediterranean part of Slovenia, where low precipitation during summer is characteristic. It was found that the seed mixture with stress‐tolerating species but without nurse species resulted in a smoother succession to a species‐rich grassland with stable cover. Succession was, by contrast, delayed in the mixture with nurse‐plants because they inhibited the establishment of the stress‐tolerating species and target species, respectively.     

Effects of small-scale animal disturbances on plant assemblages of set-aside land in Central Germany

Abstract. The distribution, density and cover of small-scale natural disturbances was surveyed in semi-natural and set-aside vegetation in a hilly landscape in Central Germany in the spring of 1995. More of the landscape was disturbed on set-aside agricultural land in valley s (1.02 %) and on warm south-facing slopes(1.33 %) than on rocky hilltops (0.3 %) and cool north-facing slopes (0.56 %). The major agents of disturbance on set-aside fields were moles, rodents (and their predators) and wild boars. In surrounding semi-natural grasslands, rabbit warrens were common on south-facing slopes and mound-building ants on north-facing slopes. Disturbances were significantly clustered and frequently superimposed. In order to investigate the effects of disturbance quality on plant assemblages in set-aside fields, two common types of disturbances were compared. These were grazing lawns (0.4–1.0 m²) maintained throughout the summer by common voles (Microtus arvalis), and excavations, 0.7–1.6 m² in area, where wild boars (Sus scrofd) grubbing for food had removed vegetation and top soil in early spring. Both types of disturbances increased plant species richness on the local scale. Wind-dispersed annual and pauciennial forbs of ruderal habitats (e.g. Carduus acanthoides, Cirsium vulgare, Matricaria maritima, Senecio vernalis) were most abundant on the superficial disturbances made by voles, whereas small, ephemeral field weeds (e.g. Polygonum aviculare, Anagallis arvensis, Chenopodium album, Fallopia convolvulus) predominated on patches grubbed by boars. This study confirms that small-scale disturbances by animals provide a variety of regeneration niches for field weeds and ruderals. The rate of decline in plant species richness on set-aside land is likely to be reduced where land is utilized by a variety of herbivorous and soil-moving mammals.     

Grassland restoration in practice: Do we achieve the targets? A case study from Saxony-Anhalt/Germany

Grassland restoration on arable land is the second most implemented compensation measure in Germany to counteract impacts of infrastructural projects on nature. Most grassland restoration has been carried out using standardized commercial seed mixtures with large amounts of perennial generalists, cultivars and seeds of non-local origin. To evaluate whether this current practice is appropriate for developing regional types of species-rich mesophile grasslands, we analyzed four widely used treatments in a real-world setting (48 plots): (1) sowing a non-site-specific herb-poor mixture; (2) sowing a non-site-specific herb-enriched mixture; (3) sowing a site-specific herb-enriched mixture; and (4) spontaneous regeneration. After up to nine years, restored sites differed from target grasslands in: (1) number of species; (2) abundance and dominance of target species; and (3) dominance structure. Sown fields were dominated by sown species from the beginning. Because most differences were due to increasing cover of a small number of sown species, we found little development toward regional types of species-rich mesophile grasslands. In contrast, species composition on spontaneously regenerated sites changed to a greater degree and showed gradual development toward target grasslands. The limiting factor for successful restoration on all sites was availability of propagules. On sown sites, dominance patterns - particularly of Festuca rubra cultivars - had a negative effect on immigration and development of target species. For future restoration practice, we strongly recommend avoiding standardized commercial non-local seed mixtures. In particular, highly competitive cultivars should never be used. Even spontaneous regeneration should be preferred over standardized mixtures. However, in species-poor environments enrichment with selected species is necessary to reach target state.     

The Use of Mixtures of Sulfur and Bracken Litter to Reduce pH of Former Arable Soils and Control Ruderal Species

High soil pH has been highlighted as a constraint to the restoration of heathland on ex‐arable land. Previous studies at the Minsmere Reserve of the Royal Society for the Protection of Birds (RSPB) in England have shown that it is possible to acidify ex‐arable soils using elemental sulfur and bracken litter, although sulfur (S) is more effective. Current recommendations suggest that 4 tS/ha need to be applied to reduce soil pH below pH 4, control vigorous ruderal species, and create conditions suitable for Calluna vulgaris (heather) establishment. However, S is relatively expensive, and as bracken litter is moderately abundant within the reserve, it made economic sense to evaluate the potential for mixing S with bracken to see if adequate pH reductions could be achieved at lower S rates. Accordingly an experiment was designed to test the effects of combining S (0–8 t S/ha) and bracken litter (0–10 cm depth layers) on (1) soil pH, (2) cover of ruderal species, and (3) the developing plant community. Significant interactions were detected, especially in the period immediately after application. Where bracken litter was applied the soil pH fell immediately; in contrast, S took at least six months to start reducing pH. Where mixtures were applied there was a synergistic effect, which produced a lower pH than the S or bracken litter applied alone. These effects were most marked at low S application rates, between 0.5–4 t S/ha. The effects of the bracken litter addition also reduced the growth of ruderal species in the period immediately after application, probably through a combination of acidification and physical smothering. There is, therefore, a clear potential to acidify ex‐arable soils using combinations of S and bracken litter in schemes designed to restore Calluna heathland.     

Effects of grazing on plant communities and successional processes vary along an aridity gradient at a northern temperate grassland

The intermountain grasslands of North America reach their most northern geographic extent in interior British Columbia’s Cariboo-Chilcotin region. Here, this study examined the long-term effects of livestock grazing exclusion and reductions in grazing severity on plant community characteristics including plant and litter cover, species richness and abundance of leading species of 33 grassland sites across a broad aridity and soil property gradient. Across the aridity gradient, grazing reduced species richness, plant cover, and litter cover. However, the effects of grazing on dominant species varied across the gradient. In more arid grasslands, historical grazing substantially reduced cover of late-seral native bunchgrass Psuedoroegnaria spicata, and repeated measurements indicate that very long time periods are necessary for successional processes associated with recovery of native bunchgrasses. At the cool-wet end of the aridity gradient, successional processes are more rapid but dominated by exotic species Poa pratensis and Tragopogon pratensis. Recent (past 20 years) light grazing and rest-rotation have favored Poa pratensis at the expense of native needlegrasses (Achnatherum spp. and Hesperostipa spp.). We suggest that absence of a dominant large-stature native bunchgrass for mesic grasslands was a key factor in the invasion and dominance of Poa pratensis.

     

Effects of the litter layer of Pteridium aquilinum on seed banks under experimental restoration

Questions: Does the litter layer of Pteridium aquilinum (bracken) act as a barrier to certain species in the seed bank? Does bracken control/restoration treatment affect seed transfer through the litter layer? Location: Five experiments at three sites across the UK covering two major vegetation types; acid‐grassland and heath‐land. Methods: At each experiment a range of bracken control and vegetation restoration treatments were applied for about ten years. The seed bank was sampled in both the bracken litter and the soil. The cover (%) of each species in the vegetation and the bracken litter abundance (cover and depth) was also estimated. Results: The bracken litter layer acts as an inert barrier as it contained a large proportion of seeds available in the litter‐soil profile (38%– 67% of the total). Bracken litter depth and cover also influenced significantly the seed bank composition in both the bracken litter and the soil. These effects were site‐specific, and species‐specific. The application of treatments changed significantly the balance between seed inputs and outputs in the bracken litter layer for some species. This was either a positive or negative response relative to the untreated control plots. Conclusion: For heathland and acid‐grassland restoration, the bracken litter layer may be an important seed source, but it must be disturbed particularly before seed addition.     

Native Alternatives for Non-Native Turfgrasses in Central Florida: Germination and Responses to Cultural Treatments

Little information exists about the establishment of native longleaf pine flatwoods species for use in restoration efforts and as buffers around natural areas in the southeastern United States. Composition of groundcover in these systems is dominated by perennial graminoid species. Vegetation in current buffers is generally non‐native turfgrass that can escape into natural areas, often reducing establishment and survival of native species. Where management objectives involve actively restoring native groundcover or reducing the probability of invasion by these non‐native turfgrasses, identification of native species and restoration methods is needed. We investigated seed germination and establishment of four species native to longleaf pine flatwoods in central Florida and one species native to the adjacent wetland communities. Paspalum setaceum, Panicum anceps, Eustachys petraea, and Eragrostis refracta were directly seeded, and P. distichum was planted as sprigs into three former P. notatum pastures. Irrigation, fertilization, weed control, and mowing treatments were assessed in terms of cover development of the sown species. Paspalum distichum developed the highest percent cover—over 80% in wet areas after 1 year. Mowing had mixed impacts depending on the species, and fertilization never significantly increased cover. Directly seeded species developed sparse cover (0–40%), probably as a result of drought conditions. However, E. petraea and E. refracta appeared more promising for use on rights‐of‐way when using high sowing rates. A second experiment conducted on a roadside included these two species and sprigged P. distichum. Both E. petraea and P. distichum developed more than 45% cover on the roadside. Establishment of these natives from seed or sprigs was significantly enhanced when site preparation effectively reduced the seedbank of other species present in the soil.     

The ecological significance of annuals on a Mediterranean grassland (Mt Ventoux,France)

A study was carried out on a Mediterranean limestone grassland to search for the ecological significance of the occurrence of annuals. In the studied grassland, most of annuals are concentrated in patches. Authors tested if annuals (1) correspond to the R-strategy (ruderal) according to Grime's theory, i. e. they are adapted to live in disturbed, productive habitats, or (2) if they present an adaptation to a xeric climate in keeping with Raunkiaer.The correspondence analyses and the examination of plants strategies reveal that:(1) annuals are grouped with the more xerophytic perennials in the ordination,(2) the cover of annuals is higher in plots with a high cover of S species (stress-tolerators) and a lower cover of C species (competitors) and CS species (intermediate strategy).These results constitute arguments to integrate annuals of Mediterranean grasslands into stress-tolerators. In the discussion are examined the importance of patches in which annuals are preferentially located. Although patches are generated by disturbances, the presence of annuals is explained by their life cycle fitting the microclimate variations, characterized by an extreme summer drought. Their cycle is not adapted here to the frequency of disturbances (unless the summer drought is considered as annual disturbance - this last point of view is discussed). In the studied grassland, annuals are then better thought of as highly developed stress-tolerators than as ruderals. A generalization is put forward by distinguishing two cases: in xeric habitats, annuals are stress-tolerant, and in productive habitats, they are ruderal.Nomenclature: follows Kerguélen (1993)     

Partial restoration of fungal and plant species diversity by removal of litter and humus layers in stands of Scots pine in the Netherlands

A field experiment was set up in five mature Scots pine (Pinus sylvestris) stands in the Netherlands in order to study the effects of removal of litter and humus on ectomycorrhizal fungi and vegetation of higher plants, bryophytes, and lichens. Lime was also applied as a countermeasure to the removal of surplus cations. Application of lime did not have significant effects on mycorrhizal fungi. Removal of ectorganic material increased species richness of ectomycorrhizal fungi. The number of fruit bodies was also higher in the sod-cut plots, but the difference compared with the controls was not significant in all years. A major effect of removal of the ectorganic layer was a reversion of ectomycorrhizal succession to earlier stages. Liming increased the number and cover of ruderal, nitrophilous plant species. Sod-cutting caused a reversion of the green vegetation to earlier successional stages.     

Effects of Nutrient Manipulations and Grass Removal on Cover,Species Composition,and Invasibility of a Novel Grassland in Colorado

Cover and richness of a 5‐year revegetation effort were studied with ,respect to small‐scale disturbance and nutrient manipulations. The site, originally a relict tallgrass prairie mined for gravel, was replanted to native grasses using a seed mixture of tall‐, mixed‐, and short‐grass species. Following one wet and three relatively dry years, a community emerged, dominated by species common in saline soils not found along the Colorado Front Range. A single species, Alkali sacaton (Sporobolus airoides), composed nearly 50% of relative vegetation cover in control plots exhibiting a negative relationship between cover and richness. Seeded species composed approximately 92% of vegetation cover. The remaining 8% was composed of weeds from nearby areas, seed bank survivors, or mix contaminants. Three years of soil nutrient amendments, which lowered plant‐available nitrogen and phosphorus, significantly increased relative cover of seeded species to 97.5%. Fertilizer additions of phosphate enhanced abundance of introduced annual grasses (Bromus spp.) but did not significantly alter cover in control plots. Unmanipulated 4‐m² plots contained an average of 4.7 planted species and 3.9 nonplanted species during the 5‐year period, whereas plots that received grass herbicide averaged 5.4 nonplanted species. Species richness ranged from an average 6.9 species in low‐nutrient, undisturbed plots to 10.9 species in the relatively high‐nutrient, disturbed plots. The use of stockpiled soils, applied sparingly, in conjunction with a native seed mix containing species uncommon to the preexisting community generated a species‐depauperate, novel plant community that appears resistant to invasion by ruderal species.     

Mass effects,clonality, and phenology but not seed traits predict species success in colonizing restored grasslands

Preserved grasslands commonly host species‐rich plant communities but a large part of the grasslands were plowed up in the past. Their restoration often requires a long time and initial restoration measures might trigger ecosystem recovery, which is then followed by spontaneous colonization. We evaluate the establishment success of target grassland species, which were not sown but established spontaneously in the restored grasslands of Bílé Karpaty Mts., SE Czech Republic. According to their key functional traits and incidence in the landscape (mass effect; acquired from the results of a grid mapping project in the region), we examined the frequency of species and their mean cover in 82 restored grasslands. The best predictor of species frequency in the grasslands was their mass effect, followed by a high capacity for clonal growth and late phenology. Seed dispersal traits (seed mass, terminal velocity, epizoochory ranking index) and plant height had no significant effect. Specific leaf area was positively correlated with mass effect. Species having a high cover in the restored grasslands had a high capacity for clonal growth. In the preparation of seed mixtures, we should therefore consider that nonclonal species relying on regeneration from seeds will be generally less able to reproduce and should be promoted by artificial sowing. At the same time, species common in the landscape, which spread well clonally, and those with a late phenology, might be expected to colonize restored meadows on their own, so that sowing them is not necessary.