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Part of the nature-protected area (NSG) "Griesheimer Düne und Eichwäldchen" in the inlanddune area of the northern upper Rhine valley with dominant Allio-Stipetum aspect and flowering Euphorbia cyparissias. In the background: former fields, now part of the NSG (May 2005. Photo: A. Schwabe).

Part of the nature-protected area (NSG) "Griesheimer Düne und Eichwäldchen" in the inlanddune area of the northern upper Rhine valley with dominant Allio-Stipetum aspect and flowering Euphorbia cyparissias. In the background: former fields, now part of the NSG (May 2005. Photo: A. Schwabe).

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To examine the necessity of livestock grazing for managing threatened sand vegetation so as to en-sure and develop its nature-conservation value, we investigated a grazing system in a model nature reserve (German upper Rhine valley) characterized by threatened steppic sandy grassland (Allio-Stipetum vegetation complex: Habitats Directive Annex I, p...

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... et Novák 1941stands (Koelerion glaucae Volk 1931: priority habitat 6120 "Xeric sand calcareous grasslands") and large areas of Allio-Stipetum Korneck 1974 steppic sandy grassland (Festuco-Brometea Br.-Bl. et Tx. in Br.-Bl. 1949, Festucetalia valesiacae Br.-Bl. et Tx. ex Br.-Bl. 1949; priority habitat 6240 "Sub-pannonic steppic grasslands", see Fig. 1); (European Habitats Directive 92/43/EEC, EUROPEAN COMMISSION 2007). This vegetation mosaic has extraordinary importance in a regional and whole European context ( ZIMMERMANN et al. 2010); it is an example of a "European High Nature Value (HNV)" farming system ( OPPERMANN et al. 2012). As in many other European eco- systems, the ...
Context 2
... the target species (Festuco-Brometea and Koelerio-Corynephoretea species) are con- sidered (Fig. 10), the Au plots are characterized by a constant decline in species numbers, while there was an increase in the Ag plots (with the exception of the last year). A highly significant grazing effect could be found in the mixed linear model, showing enhancement of target species by grazing from 2002 on. Fig. 6a, b, c. Proportion of species ...
Context 3
... species ratio". Die ausgewählten Covarianzstrukturen sind bei jeder Variable angeführt: ante = ante-dependence, arma (1,1) = autoregressive moving-average, hf = Huynh-Feldt, toep = Toeplitz. Nicht signifikante (P > 0.05) Effekte und Interaktionen wurden aus den endgültigen Modellen entfernt. Accordingly, there is a decrease of the TSR qual values (Fig. 11a) in the Au plots, whereas the grazed plots tend to increase in TSR qual values (with the exception of the last two years, where there was a slight decrease). There is a significant interaction of treatment and year, leading to significant grazing effects since 2007. Both treatments show low values in the abiotically extreme year 2003, ...
Context 4
... concerning the TSR quant (Fig. 11b) there is a significant interaction term. Beginning with the year 2007 there is a significant difference of the TSR quant depending on treatment. The ungrazed plots show first constant values, later decreases of the TSR quant values. On the other hand, the grazed plots show nearly a doubling of their TSR from 0.30 to ...

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... Allowing large herbivores to roam freely, within the carrying capacity of an area, can contribute, to some extent, towards biodiversity conservation, and it is sustainable economically (Schwabe et al., 2013;Timmermann et al., 2015) because forage supply is generally not needed. Moreover, it is sustainable environmentally as this practice limits the input of additional nutrients into the habitat. ...
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Traditional Mediterranean Mountain landscapes in Spain have suffered dramatic environmental and social changes over the last seven decades. The loss of these landscapes has had consequences for biodiversity, soil erosion, landscape quality and ecosystem services as croplands and pastures were mainly converted into forests and scrublands. Many animal breeds present in traditional land uses such as the Catalan donkey (Equus asinus var. catalana), are also at risk of extinction, but can provide environmental services while recovering traditional landscapes. In Puy de Cinca, a village in mountainous northeast Spain, we studied how grazing by Catalan donkeys reduces pasture biomass and the effects on plant diversity in pastures. We used Sentinel-2 satellite imagery to calculate the Normalized Difference Vegetation Index (NDVI), a biomass sensitive spectral index, throughout the grazing period to monitor pasture biomass and compared it to pastures without grazing. We also calculated several plant diversity indicators in pastures with and without grazing donkeys. Furthermore, we studied land use changes over the last seven decades using old (1956) and current (2018) aerial images, with forest, agricultural lands, and trails interconnecting the village being mapped to understand landscape changes. The results indicated a great increase in the forested area (348.3 ± 17.0 ha). Meanwhile, a severe decrease in cropland area (73 %) and trail length (62.5 %) was also observed. Concerning the effect of donkey grazing, biomass was lower in pastures with grazing donkeys, with NDVI values decreasing once donkeys started grazing. Nevertheless, plant diversity was higher in pastures with grazing donkeys than in abandoned pastures. This study demonstrated the capacity of low-to-moderate-intensity donkey grazing to improve plant diversity and reduce biomass in pastures. Furthermore, the study of land use changes allowed an understanding of landscape dynamics , which can help address the social and environmental recovery of the village.
... The few relics of the "Campagna Romana" landscape are an example of how a moderate human pressure can increase the number of available niches and decrease competition, allowing a great number of species to thrive. Among many other practices, grazing promotes a high environmental heterogeneity (Schwabe et al. 2013) and species diversity (Aavik et al. 2008;Dostálek and Frantík 2008) and sometimes can prevent the increase of ruderal species (Schwabe et al. 2013). ...
... The few relics of the "Campagna Romana" landscape are an example of how a moderate human pressure can increase the number of available niches and decrease competition, allowing a great number of species to thrive. Among many other practices, grazing promotes a high environmental heterogeneity (Schwabe et al. 2013) and species diversity (Aavik et al. 2008;Dostálek and Frantík 2008) and sometimes can prevent the increase of ruderal species (Schwabe et al. 2013). ...
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... In the last two decades, year-round grazing systems with large herbivores aiming to substitute extinct wild grazers (Vera, 2000;Bunzel-Drüke et al., 2008;Svenning et al., 2016;Pedersen et al., 2020) are increasingly applied to counteract further degradation of nutrient-rich (Gilhaus et al., 2014) and nutrient-poor acidic grasslands or heathlands (Schwabe et al., 2013;Rupprecht et al., 2016;Henning et al., 2017a,b). For these habitats, large herbivores are known to shape the pasture through their specific feeding behavior and utilization (Putfarken et al., 2008;Gilhaus et al., 2014) creating diverse vegetation structures, appropriate biomass removal and improved light conditions at the soil surface through reduced litter layer and woody plant cover (Bokdam and Gleichnam, 2000;Hejcman et al., 2005;Borer et al., 2014). ...
... Frontiers in Ecology and Evolution 08 frontiersin.org Schwabe, 2007;Schwabe et al., 2013;Henning et al., 2017a) assuming that a sufficient, adapted stocking rate is guaranteed as undergrazing can lead to ruderalization or tall species dominance (Sýkora et al., 2009;Moinardeau et al., 2021). Concerning faunistic diversity, forb enhancement of grassland is usually beneficial to insects as forbs provide both nesting and foraging sites (Bonari et al., 2017), and in the consequence beneficial to higher trophic animal species like birds (Lovász et al., 2021;Schmidt et al., 2022). ...
... teeth protecting from abrasion (coevolution of grasses and grazers beginning in the Tertiary; Stebbins, 1981;Bunzel-Drüke et al., 2008). These characteristics make horses well suited to reduce grass encroachment in dry grasslands (Süss and Schwabe, 2007;Schwabe et al., 2013). ...
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Introduction Dry calcareous grasslands are among the most species-rich habitats worldwide but strongly endangered by abandonment causing a severe decline of characteristic species such as orchids. To counteract further degradation, economically sustainable restoration tools such as megaherbivore grazing, that aim to substitute extinct wild grazers, should be considered. However, the long-term effects on target species of dry grasslands, and in particular for orchid populations, is still unclear. Methods To contribute to this knowledge gap, we applied vegetation surveys (5 m × 5 m), as well as large-scale census-based orchid observation and mapping of habitat structures (50 m × 50 m) in a year-round grazing scheme of a Natura 2000 site in Central Germany over 12 years. The horses and their grazing activity were observed via GPS telemetry. We fitted linear mixed models to evaluate whether Ophrys apifera density was affected by horse feeding frequency, habitat structure variables (bare soil patches, woody plant cover), grassland type or years. Results A The main results were that the target dry calcareous grassland vegetation significantly increased in species numbers and cover. The total abundance of O. apifera increased from 1,237 (2013) and 1,893 (2018), to 4,652 (2021) individuals. Ophrys apifera density was positively affected by horse feeding frequency as well as grazing-induced enhanced bare soil patches but underlying mechanisms varied between the three classified grassland types. Discussion Our results indicate that low-intensity year-round horse grazing as a relatively new restoration tool in dry calcareous grasslands has the potential to enhance floristic biodiversity in general, and particularly O. apifera density in the long run. Furthermore, we showed that highvalue xeric grasslands with outstanding orchid abundances can be integrated into the year-round grazing system and that there is no need of fencing off such sections.
... In contrast, vegetation scientists carried out plot surveys, so-called relevés, since the beginning of the 20 th century 6 , and some of these plots have been repeatedly surveyed. Such vegetation-plot time series have mainly been collected for particular habitats, such as forests [7][8][9][10][11][12][13][14][15][16][17][18][19] , hedgerows 20 , wet grasslands [21][22][23][24] , mesic grasslands [25][26][27][28][29][30][31] , dry grasslands 24,[32][33][34][35][36][37] , acid grasslands and heathlands [38][39][40] , alpine grasslands 41,42 , rivers 43 , riverbanks 44 , peatlands [45][46][47][48] , roadsides 49 or arable land [50][51][52] . Sometimes, they were recorded to assess the changes in species composition across all communities that occur in a certain area [53][54][55][56][57] . ...
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... Die signifikante Zunahme der offenen Bodenstellen (Henning et al. 2017) durch Tritt und Komfortverhalten der Weidetiere (Trittsiegel, Tierpfade und Wälzstellen) führt zur Aktivierung der Diasporenbank und zu einer Verbesserung der Keimungs-und Etablierungsbedingungen insbesondere für konkurrenzschwache, lichtliebende, kleinwüchsige Arten (vgl. auch Schwabe et al. 2013). Gleichzeitig wird die Ausbreitung dieser Arten über Hufe, Dung und Fell der Weidetiere gefördert (Schwabe et al. 2013). ...
... auch Schwabe et al. 2013). Gleichzeitig wird die Ausbreitung dieser Arten über Hufe, Dung und Fell der Weidetiere gefördert (Schwabe et al. 2013). Auf den unbeweideten Kontrollflächen blieben die absoluten Artenzahlen über den gesamten Untersuchungszeitraum hinweg mehr oder weniger konstant. ...
Article
The extent and speed of biodiversity loss in Germany, that has been continuing for decades, is unprecedented and has developed into a veritable biodiversity crisis. The long-term downward trend in species diversity is found not only in the intensively used agricultural landscape, but also in protected areas. Using the example of the 800 ha semi-open pasture landscape "Oranienbaumer Heide", we studied the development of vascular plant species richness as well as of breeding bird species populations that are characteristic of open and semi-open landscapes. Our study shows that the loss of species diversity as a result of lengthy succession processes can not only be halted, but species diversity can be increased enormously through broad-scale and long-term open land management which is nature conservation-compliant. The significant increase in mean plant species richness - in particular due to the increase in selected target species and indicator species of nutrient-poor conditions - is accompanied by a significant improvement in habitat structures. The marked increase in arthropods as well as the improved habitat structures also led to a verifiable promotion of target bird species.
... Among them, Festuco-Brometea grasslands are valuable for their high plant diversity and richness in rare plant species [11,12] and provide habitat for several animal species, such as endangered Agronomy 2021, 11, 6 2 of 12 birds and butterflies [13,14]. During the last few decades, several programs concerning the reintroduction of grazing in abandoned dry grasslands have been successfully implemented in Europe to support biodiversity and rare species conservation and to control invasive shrubs and highly competitive grass species [15][16][17][18]. Selective grazing, trampling, defecation, and heterogeneous distribution of the stocking rates enhance the sward's structural heterogeneity, thus supporting the ecological needs of a rich flora [19][20][21]. ...
... Most of the pioneer xerothermic species are annual and light-demanding and require bare ground microsites to germinate and grow. In penning areas, their competition was probably enhanced by the opening of gaps in the sward and the reduction in the litter layer induced by trampling, which provided suitable germination microsites [18,19,21]. In particular, Alyssum alyssoides (L.) L., Arenaria serpyllifolia L., H. salicifolium, O. reclinata, and Petrorhagia saxifraga (L.) Link were the most abundant pioneer xerothermic species in the penning areas at the end of the grazing period (Table S1). ...
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... We assume that in recent extremely dry years dry gassland species benefited from their capacity to gain ressources even under severe drought conditions. Generally, the reduced shading and competition improved light conditions on the ground through livestock grazing and trampling (ROSENTHAL et al. 2012, ELIAS et al. 2018a which increased germination probabilities, establishment and vegetative spread for low-competitive (calcareous) dry grassland species (SCHWABE 1997, SCHWABE et al. 2013, ELIAS et al. 2018a, b, SILVA et al. 2019 (Fig. 5). ...
... In den letzten Jahren mit extremer Trockenheit profitierten die Trockenrasenarten vermutlich von ihrer Fähigkeit, Wasser-und Nährstoffressourcen auch unter sehr trockenen Bedingungen erschliessen zu können. Generell erhöhte die verringerte Beschattung und Konkurrenz die Lichtverfügbarkeit am Boden (ROSENTHAL et al. 2012, ELIAS et al. 2018a, wodurch sich die Keimungswahrscheinlichkeit, Etablierung und vegetative Vermehrung für konkurrenzschwache (Kalk-) Trockenrasenarten verbesserte (SCHWABE 1997, SCHWABE et al. 2013, ELIAS et al. 2018aSILVA et al. 2019 Auf den unbeweideten Flächen blieb die Artenvielfalt über einen relativ langen Zeitraum erhalten (insb. auf den flachgründigen Steilhängen), im letzten Erfassungsjahr wurde jedoch schon eine Abnahme der Zielartendeckung beobachtet. ...
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Dry calcareous grasslands are one of the most species-rich habitats in the cultural landscape of Central Europe. The species richness largely resulted from the activities of humans and their livestock. Due to large-scale abandonment driven by socioeconomic changes during the last century, shrub encroachment is one of the main threats to many orchid-rich calcareous grasslands. Goat pasture has been reported to efficiently restore highly shrub encroached habitats. However, positive effects may be opposed by negative effects on target species, in particular, orchids, but long time data on goat-restored orchid sites are still rare. Over 10 years, we studied the development of large-scale woody plant cover and plant species on 12 permanent 25-m² plots in a spring, summer and autumn goat paddock and three unpastured control slopes. In the summer paddock and an adjacent unpastured site, where the largest population of the main orchid target species Ophrys sphegodes occurred, individual number, leaf index, number of blossoms and seed capsules, annual recruitment, mortality, and vegetative dormancy were observed on seven 1-m² plots. Woody plant cover was significantly reduced in all paddocks with best results in the spring and summer paddock, whereas the unpastured control showed a significant increase. The goat observations revealed a high browsing frequency for the most abundant shrubs Cornus sanguinea and Viburnum lantana. The number and cover of characteristic dry calcareous grassland species increased or remained stable in pastured sites, whereas they finally declined in unpastured sites. Ophrys sphegodes individual number significantly increased in the summer paddock and unpastured plots. Pasturing promoted a higher recruitment probably due to enhanced light availability and the creation of open soil. But we also observed a higher mortality and vegetative dormancy through stronger desiccation of these open soil patches, leading to higher population dynamics on the pastured sites. However, the main advantage of reduced shrub encroachment trades off possible individual losses in the long term. Other orchid species showed high year-to-year variations but were still abundant in the paddocks after eight years of regular pasture. We conclude, that goat pasturing is suitable to restore shrub encroached dry calcareous grasslands with a higher efficiency in spring and summer. However, spring pasturing must be adapted to the phenology of occurring orchid species, especially if they are locally rare, or should be alternately combined with summer or autumn pasture to ensure generative reproduction. Generally, the high year-to-year variations of orchid species call for a long-term scientific monitoring to evaluate the restoration outcomes.
... However, the cattle effect is limited by their preference for herbaceous plants [14] and by the quality of the food offered by the Mediterranean understory [15]. The equines are well suited to reduce grass encroachment as they can graze in low-quality fodder [16,17]. It has been observed that horses and goats grazing together in a Mediterranean scrub reduced plant phytovolume significantly [18]. ...
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During the second half of the 20th century, European countries experienced an increase in their forest area due to the global change. Consequently, there has been an increase in large forest fires, mainly in the Mediterranean basin, and this has forced the development of several types of prevention programs. One of them is the control of the understory by livestock. In this sense, browsing with a combination of donkeys and goats could be a good option, as both animals usually feed on forest species. However, little is known about their preferences for the key species of the Mediterranean forest. Using a cafeteria test, the preferences and consumption of both animals have been determined for five typical species of the Mediterranean forest, such as Quercus ilex, Pinus halepensis, Phillyrea latifolia, Rubus ulmifolius, and Brachypodium retusum. Results showed that donkeys and goats could act complementarily in the reduction of the fuel biomass of forests. Donkeys appear to act more on fine fuel, such as B. retusum, and goats on the more pyrophyte species, in this case P. halepensis. In addition, given that donkeys are at severe risk of extinction in Europe, this role of providing ecosystem services could contribute to their conservation. Despite this study only showing that goats and donkeys would consume all five presented plant species and that there are some differences in consumption during a short-term test, it constitutes a useful first step for conservation and fire prevention in the Mediterranean forests.
... Grasslands on acid inland dunes comprise an important habitat for the conservation of biodiversity in Europe and are protected as Natura 2000 habitats (EUROPEAN COMMISSION 2013). In nemoral and boreal vegetation zones, they are principally semi-natural habitat types, created as a result of man-made deforestation and long-lasting intensive humaninduced disturbance (including high grazing pressure) (SCHWABE et al. 2013, DENGLER et al. 2020a. Their area significantly decreased in the 20 th century, primarily due to the abandonment of traditional land-use practices and subsequent intentional afforestation as well as natural succession (WOLFF et al. 2017, DENGLER et al. 2020b). ...
Article
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Psammophilous grassland communities of inland dunes often occurring together with common juniper scrub are among the most threatened habitat types in Central Europe. Once they were related to disturbance caused by traditional agriculture and forest management (sheep grazing, burning, litter raking and overexploitation of forests). Currently, after the above-mentioned drivers have disappeared, those communities are subjected to secondary succession leading to loss of their typical biodiversity, an important component of which are cryptogams (bryophytes and lichens). In the presented study we ask whether the recently increasing off-road activity, concentrated in dune areas, can maintain such habitats and their biodiversity or instead leads to an even faster deterioration. To answer this question, we studied the cryptogam diversity in an off-road disturbed inland dune environment of a seldom used military training area located in Central Poland. A set of vegetation samples was collected, together with a range of measured or estimated variables including soil parameters , off-road disturbance intensity and factors related to ecological succession, like common juniper scrub cover. The obtained results suggest that highest off-road intensity led to a transformation of the habitat into its very initial stage with active dune processes, while within undisturbed patches the succession led to the development of initial pine forest. The highest number of lichen and bryophyte species occurred within not or only slightly disturbed habitats. However, lichens preferred less shrub cover than bryo-phytes. Nevertheless, pioneer, typical psammophilous grassland species, which also include some threatened lichen species, benefited from small to medium disturbance intensity. Diverse and spatially complex off-road activity increases the number of microhabitats and can thus be a positive factor promoting the coexistence of all ecological groups of cryptogams connected with the acid inland dune environment. In our opinion, off-road activity may to some extent be used as an inexpensive nature conservation tool in order to maintain open, cryptogam-rich dune habitats.
... Dominant species in degraded plots belonged to generalists represented by graminoids and competitively strong forbs. If such a site remains unmanaged, competitive graminoids such as Calamagrostis epigejos or Arrhenatherum elatius will become dominant (Stroh et al. 2002;Süss et al. 2010;Schwabe et al. 2013). On the other hand, specialists indicating sandy grasslands, drought-tolerant and nutrient-poor species, and threatened species had a high abundance in well-preserved vegetation (Beil et al. 2012), also at the restored sites. ...
Article
Natural open inland sandy grasslands are a highly threatened habitat in Europe. The originally nutrient‐poor habitats have been affected by increased aerial N‐deposition and the spread of competitively strong plants. We investigated the restoration of degraded psammophilous vegetation in the southeastern part of the Czech Republic, central Europe, where the competitively strong grass Calamagrostis epigejos had formed a compact cover over many hectares formerly dominated by valuable psammophilous grasslands. To expose nutrient‐poor sand, the upper nutrient‐rich layer was scraped off down to a depth of 10–30 cm. Vegetation was recorded by phytosociological relevés annually in a total of 15 permanent plots (5 × 5 m) established between 2012 and 2014 (five plots in each consecutive year). Reference sites with target psammophilous vegetation and degraded sites were also sampled. The target psammophilous species colonised the open sand areas immediately from the adjacent well‐preserved vegetation fragments. Out of 101 vascular plant species recorded in total in restored sites, 34 species were classified as target species, out of which 25 species were classified as threatened. After six years, the species richness and number of threatened species in the restored plots approached those of the reference sites. The results of multivariate analyses revealed that spontaneous development of restored plots run fast towards psammophilous vegetation. The optimal restoration strategy of eutrophicated open sandy grasslands should consist in creating a mosaic of topsoil removal patches followed by disturbance after some time and preservation of still existing fragments of target vegetation. This article is protected by copyright. All rights reserved.