Next Article in Journal
Habitat Destruction Alters the Mechanisms of Species Coexistence by Modifying Competitive Structure
Previous Article in Journal
Postglacial Origin and Regional Differentiation of Microtus arvalis in the Baltic Region
Previous Article in Special Issue
Six New Records of Cortinarius and Russula from Northeastern China
 
 
Search for Articles:
Title / Keyword
Author / Affiliation / Email
Journal
Article Type
 
 
Advanced Search
 
Section
Special Issue
Volume
Issue
Number
Page
 
Journals
Diversity
Volume 18
Issue 4
10.3390/d18040216
diversity-logo
Submit to this Journal Review for this Journal Propose a Special Issue
► Article Menu

Article Menu

share Share announcement Help format_quote Cite question_answer Discuss in SciProfiles
first_page
settings
Order Article Reprints
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Review

Semi-Natural Dry Grasslands in Decline: A Review of Characteristics, Threats and Conservation Challenges

by
Justyna Wielgos
and
Mariusz Kulik
*
Department of Grassland and Landscape Planning, University of Life Sciences in Lublin, Akademicka 15, 20-950 Lublin, Poland
*
Author to whom correspondence should be addressed.
Diversity 2026, 18(4), 216; https://doi.org/10.3390/d18040216
Submission received: 2 March 2026 / Revised: 3 April 2026 / Accepted: 6 April 2026 / Published: 8 April 2026
(This article belongs to the Special Issue Biodiversity, Community Structure and Ecology of Terrestrial Ecosystems Under Global Change)
Browse Figures
Versions Notes

Abstract

In Europe, the most valuable grasslands are semi-natural ecosystems maintained by long-term extensive human management, particularly pastoralism, and therefore do not represent climax vegetation. According to the Natura 2000 habitat interpretation manual (EUR-28), key habitats include xerothermic grasslands of Festuco-Brometalia (code 6210*) on calcareous soils and sandy grasslands of Koelerion glaucae (code 6120*) on poor substrates. Only 10–15% of their area in the EU has favorable conservation status. The main threat is secondary succession and encroachment (83.94%), caused by abandonment of traditional management (81.75%). Without mowing or grazing, dominant grasses replace rare species, followed by shrubs and trees. Other pressures include intensive agriculture (75.18%), habitat loss and fragmentation (69.34%), climate change (37.96%), invasive species (23.36%) and urbanization (14.60%). Multiple threats often co-occur, so cumulative percentages exceed 100%. The most effective conservation method is restoring or maintaining extensive grazing, particularly with local sheep and goat breeds. Grazing limits succession, increases structural diversity and promotes seed dispersal, creating a mosaic of microhabitats that enhances biodiversity. Effective protection requires landscape-scale actions, limiting urban development, and long-term support for farmers under the Common Agricultural Policy. Increasing public awareness of the ecological and cultural value of these ecosystems is also essential.

1. Introduction

Grasslands, defined as plant formations dominated by grasses and other graminoids such as Cyperaceae and Juncaceae, constitute one of the largest terrestrial biomes, covering from 25% to 40% of the Earth’s land surface [1,2]. They occur on all continents except Antarctica and perform key ecosystem functions, such as forage production, soil carbon sequestration, protection of water and soil resources and supporting high biodiversity [3,4,5,6,7]. In the context of grassland management, this refers to the traditional, economic function of these ecosystems, which involved obtaining fodder. Today, however, management has a crucial ecological dimension, biomass removal halts succession through the combined effects of nutrient depletion, physical disturbance and the prevention of litter accumulation. While moderate management can provide forage for livestock, it is important to note that improper grazing intensity may lead to competitive exclusion or reduced food availability for wild herbivores [8].
In the European context, most ecologically valuable grasslands are of a semi-natural character, reflecting a unique synergy between post-glacial climate and long-term traditional land use. These ecosystems do not constitute climax communities, instead, they represent stable anthropogenic stages maintained by centuries of human activity dating back to the Neolithic period [9]. Deforestation, followed by regular, extensive use such as grazing or mowing, halted the natural process of ecological succession towards forest communities. This process allowed for the development and survival of unique phytocoenoses characterized by exceptional alpha-diversity. These communities maintain high vascular plant species density because regular management prevents the competitive exclusion of smaller plants by dominant species, thus acting as critical strongholds for heliophilous flora. While the transition towards forest communities is a natural successional process, this study focuses on the immediate conservation challenges of maintaining the anthropogenic biodiversity of open grasslands. Therefore, the aim of this paper is to provide a comprehensive review of the current threats to 6210 and 6120 habitats in Europe, evaluate the effectiveness of active management practices and propose evidence-based recommendations for their long-term preservation [10,11,12,13,14,15,16,17,18,19,20] .
Unfortunately, in recent decades, a drastic decline in the area and degradation of these ecosystems has been observed throughout Europe [9,17,21]. Socio-economic changes in agriculture, leading, on the one hand, to the abandonment of traditional, unprofitable forms of farming and, on the other hand, to the intensification of production on more fertile lands, have become the main cause of this phenomenon [4,22,23].
The aim of this publication is to review and synthesize current knowledge on the ecological characteristics of xerothermic grasslands (6210*) and thermophilous sandy grasslands (6120*), to identify the main factors threatening these habitats across the European Union and to evaluate the effectiveness of active conservation measures. Particular attention is given to examples and data from Poland and Central Europe, where these habitat types have been extensively studied. Codes 6210* and 6120* designate priority habitats within the Natura 2000 network, which are subject to strict legal protection across the European Union. They identify the most valuable ecosystems threatened with extinction that require urgent conservation measures for their survival.

2. Literature Search

This paper presents a literature review and analysis based on scientific publications, examining the characteristics, threats and current status of thermophilous grasslands in Europe. To identify and classify the main threats to these habitats, a systematic literature review was conducted across the Web of Science and Scopus databases on February 9, 2026.
To guarantee a robust and exhaustive overview of the subject matter, the search utilized a combination of specific keywords: (“xerothermic grassland*” OR “sandy grassland*” OR “calcareous grassland*” OR “acidic grassland*” OR “semi-dry grassland*” OR “Festuco-Brometalia” OR “Koelerio-Corynephoretea”) AND (“biodiversity” OR “characteristic*” OR “composition” OR “species richness”) AND (“threat*” OR “conservation” OR “management” OR “restoration” OR “succession”). This strategy was designed to capture the most relevant publications.
The publication selection followed a multi-stage process according to the PRISMA model [24,25]. The procedure included an initial assessment of titles and abstracts, followed by a detailed full-text analysis of eligible papers. The structure and workflow of the literature review are presented in the flow diagram (Figure 1). An initial search of the Web of Science and Scopus databases yielded a total of 727 records, which were supplemented by 6 additional items identified through the “snowball” method (citation tracking). After removing 146 duplicates, a total of 581 unique publications remained for screening. In the first stage of selection (screening), 389 items were excluded based on title and abstract analysis as they did not meet the research scope. Subsequently, 198 reports were assessed for full-text eligibility. Within this pool, 6 documents were unavailable, and a further 61 were excluded after detailed reading due to a lack of direct relevance to threats facing grassland habitats. Out of the total records identified, 6 documents were excluded from the analysis as their full-text versions were unavailable. To ensure data consistency and reliability, only publications with accessible full content were included in the synthesis. In addition to the systematic database search, six further records were identified through citation searching. Ultimately, 137 publications met the inclusion criteria for the final analysis. The analysis covered peer-reviewed articles, reports and official documents published between 2000 and 2025. This 25-year period was selected to encompass the most relevant research following the implementation of the Natura 2000 network and to include the latest available data on habitat conservation status.
The literature review focused on identifying threats to xerothermic and sandy grassland habitats. Data regarding specific threat factors—including abandonment of land use, negative agricultural activity, habitat loss and fragmentation, natural succession, invasive alien species, climate change and urbanization—were extracted from each source. The distribution of threat categories was recorded in a binary matrix (Table 1), where ‘1’ signifies that a publication identified or discussed a specific threat, and ‘0’ indicates its absence. The percentages presented in the results section represent the proportion of the total analyzed publications (N = 137) that identified a given threat factor. A threat was recorded if it was either the primary subject of the study or was explicitly identified by the authors as a significant factor affecting the studied habitats. Based on this matrix, a bar chart (Figure 2) was generated to illustrate the frequency of each category. A total of 529 mentions were identified across 137 qualified publications; this discrepancy arises from the fact that many works address multiple threat factors simultaneously.
Due to the descriptive and narrative nature of this review, a formal risk of bias assessment of the included studies was not conducted. The review is based on a thematic synthesis of data from all included studies, rather than a detailed summary of each individual study.

3. Characteristics of Selected Grassland Types

3.1. Semi-Natural Dry Grasslands and Scrubland Facies on Calcareous Substrates (Festuco-Brometalia) (* Important Orchid Sites) 6210*

Xerothermic grasslands (Figure 2) are among the most floristically rich and, simultaneously, most endangered plant communities in Poland and Europe [160]. Their uniqueness stems from their formation and functioning in specific, often extreme, habitat conditions. Due to their high natural value, they have been included in the Natura 2000 network as the priority habitat 6210* “Semi-natural dry grasslands and scrubland facies on calcareous substrates (Festuco-Brometalia)” [16,17,19,20,161,162]. Those sites featuring important orchid populations are considered to be of particular importance within this habitat [16,18,162].
The existence of xerothermic grasslands is inextricably linked to human activity, making them a classic example of a semi-natural ecosystem [163,164,165]. Their origin dates to the Neolithic period, when the clearing and burning of forests began to acquire land for cultivation and pastoralism [9,10]. It was precisely the centuries-long, extensive grazing of animals that became the key factor preventing the return of the forest and allowing for the survival and development of heliophilous steppe vegetation [10,11,12,14,162]. Many species that build these communities are post-glacial relics that arrived in Central Europe during a period of climate warming and survived precisely thanks to humans maintaining the open character of the area. In countries such as Poland, these habitats represent a significant portion of the regional biodiversity, acting as a bridge between Atlantic and Continental flora [10].
The occurrence of xerothermic grasslands is strictly restricted to sites with a specific combination of abiotic factors. A fundamental condition is a substrate rich in calcium carbonate. They develop on soils such as rendzinas, pararendzinas and chernozems, formed on limestone, chalk or gypsum rocks, or on loess [10,16,17,19,161,162,166,167]. These soils are characterized by an alkaline pH, which leads to the low availability of certain micronutrients, including iron and manganese [167]. They are typically shallow, skeletal, highly permeable and nutrient-poor soils with a low water retention capacity [14,138,167]. They inhabit almost exclusively steep, sun-exposed slopes with a southern, southwestern or southeastern aspect [12,16,167,168,169]. Such a location, in combination with low soil moisture, creates an extremely warm and dry microclimate. On sunny summer days, the temperature at the ground surface can reach as high as 50 °C, and evaporation often exceeds precipitation [16].
Xerothermic grasslands are among the most species-rich habitats in Central Europe. Nearly one hundred species of vascular plants can be found on a single square meter [9,10,166]. In Poland, they are divided into three main types [10]:
•
Rock grasslands: pioneer communities on steep limestone rock faces;
•
Stipa grasslands: highly thermophilous grasslands resembling steppes, formed by tussock grasses, mainly feather grasses (Stipa sp.);
•
Flowery grasslands: the most diverse and lush group, growing on gentler slopes, dominated by large, dicotyledonous perennials.
They are a habitat for many rare and protected plant species, including those covered by the Habitats Directive, e.g., typical and representative of the Central European area. These include, among others, Carlina onopordifolia, Echium russicum, Cypripedium calceolus, Rosa gallica, Gentiana cruciata, Stipa pennata, Inula ensifolia, Anthericum liliago, as well as numerous species Salvia sp., Centaurea sp. and Campanula sp. [9,10,19,167,169].
At the same time, a problem in these habitats can be the expansion of certain grass species, such as Brachypodium pinnatum and Bromus erectus, which, by forming dense swards, outcompete other, rarer plant species. The expansion of Bromus erectus is particularly well-documented in certain dry and semi-dry grasslands in Central Europe. While Bromus erectus is a characteristic species of xerothermic grasslands, an uncontrolled increase in its dominance leads to the displacement of other typical species; therefore, it is regarded as a highly competitive dominant in this context [160,170,171].
The specific conditions and floral richness make these grasslands a key habitat for many specialized animal species, especially invertebrates. They are a stronghold for rare butterflies, e.g., from the genus Jordanita [172] and beetles, e.g., Onthophagus grossepunctatus or Cheilotoma musciformis, which is endangered in Poland [17,138,168]. They also constitute the only known habitat for the relict, predatory Saga pedo, which is an indicator species for well-preserved, open xerothermic habitats [164]. They are also important hunting grounds for birds of prey [166].
On a European scale, xerothermic grasslands have an island-like and azonal character, being a remnant of former steppe formations [17]. Their largest share in the Natura 2000 network is found in the Mediterranean region [10,173]. In the European Union, this habitat is protected in 3930 sites (Figure 3).

3.2. Xeric Sand Calcareous Grasslands (Koelerion glaucae) 6120*

Thermophilous sandy grasslands (Figure 4), also referred to as psammophilous (from Greek: psammos—sand, philos—loving), are thermophilous communities occurring in very poor, sandy habitats [89,160]. Similar to xerothermic grasslands, they are semi-natural habitats whose existence depends on appropriate, extensive use [89,160,162,174]. Within the Natura 2000 network, they constitute a priority habitat with the code 6120* [20,162].
Thermophilous sandy grasslands in Central Europe were formed mainly as a result of human activity. Their genesis is linked to medieval deforestation of areas with a sandy substrate (often inland dunes in river valleys) and the maintenance of their open character through centuries of intensive grazing [11,14,171]. In some cases, fires and military activities, which led to the transformation of forest areas into open heathlands and grasslands, also contributed to their formation [171].
These habitats are characterized by extreme edaphic (soil) conditions. They develop on sandy soils that are extremely poor in nutrients and calcium carbonate (mainly on podsols) [14,20]. They have an acidic pH, which causes high solubility and availability of aluminum, iron and manganese ions, which in higher concentrations can be toxic to many plants [20]. These soils are characterized by low organic matter content and a low water retention capacity [14]. The vegetation forms a loose, often discontinuous turf, with a large share of exposed soil patches and a well-developed moss-lichen layer [89,172,175]. The species composition of these grasslands is fully adapted to the acidic substrate and the deficit of water and nutrients. Typical species include xeromorphic grasses such as Corynephorus canescens and Koeleria glauca, as well as Thymus serpyllum, Dianthus deltoides, Dianthus carthusianorum, Helichrysum arenarium and numerous lichens, e.g., Cladina mitis [89,162,171,175,176,177,178]. The main threat to the species diversity of thermophilous sandy grasslands is the expansion of wood Calamagrostis epigejos. Thanks to its competitive abilities, dense rhizomes and high biomass production, this species limits the development of lower, heliophilous plants and leads to the homogenization of the habitat [160].
Open, sandy and warm habitats are extremely important for many animal groups. They constitute a key habitat for wild bees (Apiformes) and other pollinating insects that nest in the exposed, sandy soil [21,172]. They are also home to rare butterfly species, e.g., Jordanita chloros [172,179]. They are also an important nesting and foraging site for many bird species, including waders [89].
The largest number of protected thermophilous sandy grasslands occurs in Central Europe [20,173]. The analyzed literature mentioned their presence in, among other places, the Lower Odra Valley and the Podlasie Bug Gorge in Poland [21,175,177] and in eastern Germany [172]. Many of these sites are located at the limit of their range, which makes them communities of a relict character [178]. Within the European Union, the habitat is protected in 446 sites (Figure 5).

4. Habitat Loss

Analysis of data from the last fully completed reporting cycle under Article 17 according to the 2020 national report on the implementation of the Habitats Directive (covering the period 2013–2018) provides quantitative evidence of the alarming situation of two priority grassland habitats. It should be emphasized that more recent data, covering the period 2019–2024, are currently being collected and analyzed by Member States, and their publication in aggregate form for the entire EU is expected in the future [180].
In the case of semi-natural dry grasslands and scrubland facies on calcareous substrates (Festuco-Brometalia) 6210*, the data revealed a striking disparity between conservation objectives and reality. Only 15% of the area of this habitat in the European Union were rated “favorable” (FV). This suggests that the overwhelming majority did not meet the criteria for good conservation status. A total of 80% of the habitat was in “Unsatisfactory” condition, with 50% rated as “unfavorable–inadequate” (U1) and 30% as “unfavorable–bad” (U2). Furthermore, the status of 5% of the habitat area was used as “Unknown”, meaning that monitoring was available at these locations after the advice was issued [180].
The situation of protected xeric sand calcareous grasslands (Koelerion glaucae) 6120* is, from a statistical perspective, even more dramatic. The share of habitats in “favorable” (FV) condition was marginal, at just 10%. The dominant category in the assessment of these unique ecosystems was the worst possible—“unfavorable–bad” (U2), which covered as much as 55% of the total habitat area. A further 25% was classified as “unfavorable–inadequate” (U1). This suggests that a total of 80–90% of sandy grasslands in the EU were in unsatisfactory condition, with over half of this being critically degraded. In the case of this habitat, the share of the “Unknown” rating was even higher and amounted to 10% of the total habitat area [180].
Although these official EU-wide statistics reflect the state as of 2018, recent national monitoring results and expert assessments confirm that these negative trends persist. In Poland, the latest data from the Chief Inspectorate for Environmental Protection [181] indicate that the conservation status of xerothermic and sandy grasslands shows no significant improvement, with secondary succession remaining the primary threat. This is further corroborated by the “Shadow List of Natura 2000 habitats in Poland” [182], which highlights ongoing habitat degradation and critical gaps in management. Similar trends are observed across Europe; for instance, the latest UK Habitats Regulations Reporting [183] confirms that most dry grassland habitats remain in an “unfavorable–bad” status with a declining trend due to atmospheric nitrogen deposition and insufficient grazing.
The data presented, although published as of 2018, provides a current, official summary that quantitatively records the ongoing process of biodiversity loss. The predominance of U1 and U2 ratings, supplemented by additional areas of unknown status, constitutes statistical evidence of the failure of current protection measures. These findings highlight the need for a fundamental reassessment of current conservation strategies. This requires an immediate shift in conservation priorities and resource allocation, as well as the implementation of active management and comprehensive monitoring systems. Without such actions, future reporting cycles may continue to document further declines in these habitats.

5. Threats to Dry Grassland Habitats

Below are a detailed description of the chart and the individual categories of threats to grasslands (Figure 6). Importantly, from all the literature analyzed for this publication, 137 key items in which the discussed topic was addressed were selected to create this summary. Based on them, a picture of the most serious factors threatening the existence and proper condition of these habitats was formed. The distribution presented in the pie chart illustrates that these threats are not uniform, specifically, those associated with human activity and the cessation of traditional management practices represent a significantly greater proportion.
Succession and encroachment by trees/shrubs (83.94%) are the most frequently indicated threat. This is a natural ecological process that poses a significant threat to grassland persistence. When grazing or mowing is absent, the process of succession begins. Initially, tall grasses and forbs enter the grassland, shading and outcompeting the light-loving, low-growing species typical of these habitats. Subsequently, seedlings of trees (e.g., pine, birch) and shrubs (e.g., blackthorn, hawthorn) appear. Over time, the open space of the grassland transforms into scrubland and ultimately into a forest, which means the complete loss of the habitat and its unique flora and fauna [8,11,17,27,28,29,30,31,32,33,34,35,36,37,38,39,43,44,46,47,48,49,50,51,52,53,54,55,56,58,59,60,61,62,63,64,66,67,68,69,71,72,73,74,75,77,79,80,81,82,83,84,85,86,87,88,89,90,91,92,93,94,95,96,97,98,99,101,103,104,106,107,108,110,111,112,113,114,115,116,117,119,120,121,122,123,124,125,126,127,128,130,131,132,133,134,136,138,140,141,142,143,145,146,147,148,149,150,151,152,153,154,155,157,159].
Abandonment of use and lack of management (81.75%). Grasslands, especially xerothermic and sandy types, are semi-natural habitats. This suggests that their existence and species richness depend on the continuation of traditional, extensive human activities, such as the grazing of animals like sheep, goats and cattle, which prevents the excessive development of tall grasses such as Festuca ovina or Koeleria glauca and the expansion of shrubs. It also includes regular or one-off mowing with hay collection, which removes biomass and prevents overgrowth. The abandonment of these practices, caused, for example, by economic unprofitability, leads directly to the next threat, which is succession [8,11,17,26,27,28,29,30,31,32,33,34,35,36,37,38,39,41,43,44,46,47,48,49,51,52,54,55,56,57,58,59,60,61,62,63,64,66,68,69,71,72,74,75,77,79,80,81,82,83,85,86,87,88,89,90,91,93,94,95,96,97,98,99,100,101,102,103,104,106,107,108,109,110,111,113,114,115,116,117,119,120,121,122,123,124,125,126,127,128,129,132,133,134,136,137,138,139,140,141,142,145,146,147,148,149,150,151,152,153,154,157].
Negative agricultural activities (75.18%). This category includes agricultural activities that are too intensive for delicate grassland ecosystems, namely plowing, fertilization, and re-seeding of grasses. The literature mentions the complete and irreversible liquidation of grasslands to convert them into arable fields. Additionally, the use of fertilizers (especially nitrogen-based) leads to eutrophication. In the context of dry grasslands, eutrophication does not refer to the typical phenomenon of algal blooms found in aquatic ecosystems. Instead, in terrestrial ecosystems, it signifies the accumulation of excess nutrients in the soil, primarily nitrogen and phosphorus. This surplus, originating from atmospheric deposition (e.g., pollution) or runoff from nearby agricultural fields, leads to a drastic change in habitat conditions. This allows for the dominance of fast-growing and competitive species, which outcompete the rare, light-demanding species characteristic of grasslands. Consequently, this results in the loss of the unique biodiversity of these habitats. This changes the soil conditions, promoting the growth of a few nitrophilous (nitrogen-loving) species that quickly dominate and displace hundreds of others adapted to nutrient-poor soils. Furthermore, re-seeding with grasses introduces productive but alien forage grass species, which consequently destroys the natural species composition [11,17,26,27,30,31,32,33,34,35,36,37,38,40,41,42,43,44,45,47,48,49,51,53,54,57,58,59,60,61,62,63,64,65,66,68,69,70,71,72,74,75,76,77,78,79,80,81,82,83,85,86,87,91,92,93,94,95,96,97,98,100,102,103,106,109,110,111,112,113,115,116,118,119,121,122,123,124,126,127,128,129,130,131,132,133,134,135,136,137,139,141,142,143,144,148,151,152,153,155,156,157,158].
Habitat loss and fragmentation (69.34%). The development of infrastructure, agriculture and urbanization leads to the division of once-extensive grassland complexes into small, isolated patches. This limits gene flow and increases the risk of local population extinction, especially for species with limited mobility [8,27,29,31,32,33,34,36,37,38,40,41,42,43,44,45,47,54,56,57,59,60,61,62,63,64,65,66,68,69,70,75,76,77,79,80,82,83,84,85,86,87,88,90,91,92,93,94,96,97,98,99,101,102,103,104,107,108,110,112,113,114,116,119,120,122,123,124,125,126,127,128,129,130,131,132,133,134,135,136,139,140,142,143,144,147,148,149,151,152,153,154,155,157,158].
Climate change (37.96%) is the least frequently mentioned threat in the analyzed works, although its significance is growing. Climate change can affect grasslands in many ways. Extreme droughts can lead to the weakening and extinction of species not adapted to long periods without water. In turn, milder winters may favor the expansion of alien and invasive species. Furthermore, a change in the timing of growing seasons can disrupt the life cycles of plants and the insects associated with them [17,29,33,34,35,36,38,39,41,44,47,48,50,53,55,59,60,62,63,68,69,72,74,77,83,84,85,91,94,97,101,102,103,105,107,109,116,117,118,119,121,122,132,135,136,138,139,142,143,153].
Invasive and alien species (23.36%). Among the biological threats, invasive plant species play a dominant role in the degradation of dry grasslands. While the term “invasive species” can encompass various taxa, this review focuses on vascular plants as they directly compete with native grassland species for light and nutrients. These are species with no natural enemies in a given area, characterized by enormous expansiveness. A serious threat is posed by invasive alien plants, e.g., Robinia pseudoacacia, Solidago sp. and Ailanthus altissima, especially in Mediterranean areas. They form dense, monospecific stands, displacing valuable grassland flora. Additionally, the black locust, through its symbiosis with nitrogen-fixing bacteria, drastically changes the soil chemistry (eutrophication), which eliminates species adapted to nutrient-poor conditions [26,28,34,35,36,42,47,50,53,59,64,65,66,69,75,82,83,85,88,90,91,92,94,96,113,125,126,134,135,142,143,157].
Urbanization and infrastructure development (14.60%) entails the direct, physical liquidation of habitats. Grasslands, often located on attractive, sun-exposed slopes, are destroyed for residential and recreational construction, the building of roads, railway lines and other infrastructure, as well as the development of industry and mines [26,33,36,41,59,63,66,83,84,88,92,97,123,132,135,136,140,142,143,159].

6. Methods of Conservation and Management of Dry Grassland Habitats

The conservation of xerothermic grasslands, as semi-natural ecosystems, must be based on active measures that mimic traditional forms of land use. Passive (strict) conservation is ineffective in their case and leads to their decline as a result of succession [12,13,15]. Effective conservation and management of these valuable ecosystems involve a range of integrated actions, including extensive animal grazing, mowing, removal of tree and shrub encroachment, controlled burning, restoration of degraded habitats, active protection of rare species, monitoring and adaptive management.

6.1. Extensive Animal Grazing

The systematic review and thematic synthesis of the selected publications clearly indicates that extensive animal grazing is the primary, most effective and historically justified method of grassland conservation [10,11,17,19,160,161,163,165,166,177]. Its positive impact is multifaceted. Grazing prevents secondary succession. By browse on seedlings and young shoots of trees and shrubs (e.g., blackthorn, Prunus spinosa), animals effectively inhibit the process of encroachment, which is the main threat to heliophilous grassland species [12,13,15,19,160]. Through trampling, animals create gaps in the dense sward, exposing patches of bare soil. These are so-called regeneration niches, ideal for the germination of seeds of many rare species [17,19,161,174]. Furthermore, animals disperse seeds on their fur, in their hooves and through their digestive tracts. This mechanism is crucial for maintaining genetic connectivity between isolated grassland patches. Studies have shown that a single flock of sheep can transport millions of seeds over a single season [12,89]. Extensive grazing removes dead organic matter (so-called thatch), which prevents the matting of the sward and facilitates plant development [16,19,161]. Studies have also shown higher soil enzymatic activity on grazed sites [17,19].
It is crucial to apply extensive grazing with a stocking density not exceeding 0.5–1.0 LSU/ha [184] (Livestock Unit per hectare) to avoid habitat degradation [12,89]. Native, local breeds of sheep and goats are most suitable, as they are perfectly adapted to the harsh conditions and the low nutritional value of the grassland biomass. This applies to breeds such as Olkuska Sheep, Polish Heath Sheep and Świniarka Sheep in Poland, but also to numerous local races in southern Europe, such as the Lacaune sheep in France or the Murciana goat in Spain [9,11,151,164,173,176]. Mixed-species grazing (e.g., sheep with goats or horses) is also beneficial, as different species complement each other in their feeding preferences [10,12,160,166].

6.2. Mowing and Removal of Tree and Shrub Encroachment

Woody and shrubby vegetation management involves mowing and mechanical removal. Mowing, used as an alternative to grazing, is considered less effective because it does not create a diverse turf microstructure and can promote grass expansion. Mowing late enough is crucial, allowing valuable species to set seed [9,20]. Mechanized removal of successional vegetation, involving manual or machine cutting of trees and shrubs, is often a necessary preparatory step before reintroducing grazing. However, as a standalone method, it only yields short-term results, as plants regrow very quickly [13,15,17,19,21,161,167].

6.3. Controlled Burning

Fire is one of the most fundamental, yet most complex, factors shaping grassland ecosystems. Its role is ambivalent—on a global and historical scale, it was a natural element of grassland dynamics, which are susceptible to it due to the accumulation of dry biomass [1]. Fire effectively limited the succession of trees and shrubs, stimulated the growth of grasses and influenced the nutrient cycle [1,2,162]. Historically, it was also a human tool used to create land for pasture by burning forests, thus maintaining an open agricultural landscape [10,171]. Even in more recent times, fires, for example, along railway lines, have contributed to the preservation of valuable habitats [14].
However, the action of fire is multifaceted and carries serious ecological consequences. On the one hand, it poses a mortal threat to fauna, especially to organisms with limited mobility, such as insects, amphibians, and reptiles [12]. It also acts as a powerful selection factor for flora, destroying sensitive species while favoring resistant, expansive grasses such as Calamagrostis epigejos and Brachypodium pinnatum, which leads to the homogenization of the community and a decline in biodiversity [11,12]. Furthermore, fire destroys the layer of organic matter that is crucial for dry habitats, which increases susceptibility to drought and can degrade the soil in the long term, creating conditions for the invasion of alien species. On the other hand, for some plants, e.g., from the legume family (Fabaceae), thermal shock can stimulate seed germination [162].
In a scientific context, the concept of prescribed burning emerges as a specialized conservation tool. This is a deliberate procedure, carried out under strictly defined conditions, which can serve to maintain large, open areas where other methods are impossible to implement [162]. Studies suggest that infrequent, controlled fires (e.g., on a 15–25 year cycle) can inhibit succession without negatively affecting soil carbon stocks [162]. However, it must be emphasized that this is an experimental method that requires vast knowledge and experience [10]. In the Polish context, a clear distinction must be made. The theoretical concept of prescribed burning has nothing in common with the widespread, illegal practice of spring grass burning. According to Polish law, the burning of meadows and wastelands is strictly prohibited [177]. It is perceived exclusively as a destructive threat that destroys biodiversity and degrades valuable habitats [160]. Therefore, the effective conservation of grasslands in Poland must be based on proven and ecologically sound methods, such as extensive grazing and mowing.

6.4. Restoration and Adaptive Managment

As part of active restoration on highly degraded habitats, e.g., by invasive species, radical measures can be effective, such as the removal of the topsoil layer (topsoil stripping) combined with the transfer of fresh hay from well-preserved patches to introduce a seed bank of desired species [6,14].
This method involves targeted actions aimed at supporting species that are particularly threatened or have specific requirements. An example includes creating and maintaining the aforementioned regeneration niches—small patches of bare soil created by animal trampling, which are ideal for the germination of seeds of many rare species. Another action can be the transfer of hay from well-preserved grassland patches to enrich the seed bank in degraded areas [185,186].
This is a key element of a long-term conservation strategy, based on the principle of “learning by doing.” It includes the systematic observation of the effects of applied conservation measures, e.g., assessing the impact of grazing on species composition or the effectiveness of shrub removal and adjusting actions based on the collected data. A broader aspect of adaptive management also includes integrating socio-economic factors that can support conservation goals. One such example is wisely managed tourism. Extensive grazing, being a key method of grassland conservation [6,19,89,160,161], simultaneously has significant potential for creating a tourism offer. Traditional pastoralism, especially using native breeds, creates a picturesque landscape that becomes a living tourist attraction and a basis for agritourism [12,89,165,177]. However, it must be remembered that uncontrolled tourist traffic poses a threat, e.g., through trampling, which is why it is crucial to promote responsible tourism, which involves sticking to designated trails and supporting local initiatives [12].
In summary, wisely managed tourism creates an economic justification for maintaining pastoralism, which is key to preserving grassland biodiversity. In this model, nature conservation generates a tourism product, and the income from it finances further conservation activities, creating a system beneficial to the environment and local communities [3,12].

6.5. Financial and Legal Support

The practical implementation of the aforementioned conservation methods requires robust financial and legal frameworks. European dry grasslands are recognized as threatened biodiversity hotspots, yet their maintenance is strictly dependent on continued extensive management [142]. The Common Agricultural Policy (CAP), particularly through its agri-environmental–climate schemes, remains the primary instrument for providing systemic support to farmers and landowners. These subsidies compensate for the lower productivity and higher labor costs associated with extensive grazing and mowing in sensitive habitats. However, as emphasized by Pe’er et al. (2020) [187], for the CAP to effectively address sustainability challenges, it must move toward more targeted, results-oriented support that treats biodiversity as a high-value public good. Furthermore, large-scale restoration efforts are frequently funded through LIFE projects, which are essential for coordinating actions at a landscape scale, such as restoring ecological corridors and strengthening the conservation impact of the Natura 2000 network [188]. As demonstrated by successful European initiatives, such as “LIFE to alvars” or “Trockenrasen Saar,” long-term success is contingent upon the continuity of funding that extends beyond short-term grants, ensuring that active management remains economically viable for local communities.

7. Conclusions and Recommendations

The synthesis of the reviewed literature confirms a dramatic decline and degradation of valuable, semi-natural dry grasslands across Europe. Their existence and species richness are inextricably linked to centuries-old, extensive human activities, such as grazing and mowing. However, in recent decades, these habitats have been significantly reduced and transformed due to socio-economic changes in agriculture. Our analysis highlights that the preservation of Festuco-Brometalia and Koelerion glaucae depends on a fundamental transition from passive protection to consistent, active management. This shift implies a necessary redefinition of conservation policy from static preservation of areas to process-based conservation that supports the socio-economic foundations of pastoralism. Analysis reveals that the most critical challenges for these ecosystems are secondary succession and encroachment, driven by the abandonment of land use. Crucially, these threats do not operate in isolation but form a degradative feedback loop. For instance, atmospheric nitrogen deposition acts as a catalyst, significantly accelerating the rate of succession and making the habitats more susceptible to invasive alien species. The lack of appropriate management and the cessation of traditional agricultural practices initiate a successional process, manifested by the expansion of dominant grass species that displace rarer plants, eventually giving way to shrubs and trees. Other significant factors degrading these habitats include detrimental agricultural activities, habitat loss and fragmentation, and climate change. The interaction between habitat fragmentation and climate change is particularly dangerous, as isolated patches lack the genetic resilience and connectivity needed for species to migrate or adapt to shifting climatic conditions.
The list of threats is completed by invasive alien species and progressing urbanization and infrastructure development, which directly destroy natural habitats. The current conservation status of these ecosystems is alarming, as confirmed by data from the report under the Habitats Directive (2013–2018). In the case of xerothermic grasslands (6210*), only 15% of the EU’s habitat area was assessed as being in a favorable condition, while as many as 80% was in an unfavorable condition. The situation of sandy grasslands (6120*) is even more dramatic—only 10% of habitats were in a favorable condition, while as many as 80–90% was in an unfavorable condition, with over half in a state of critical degradation. Previous conservation efforts for Festuco-Brometalia and Koelerion glaucae habitats have often relied on establishing legally protected areas (e.g., reserves) without ensuring long-term funding for active management. These measures frequently failed because they focused on passive protection, which in the case of semi-natural grasslands leads to secondary succession and the loss of biodiversity. Furthermore, many conservation projects were fragmented and short-term, lacking the continuity necessary to maintain the results after the initial funding ended. Without constant grazing or mowing, the effects of invasive species removal or shrub clearing were quickly reversed. To address these failures, a clear ranking of priority conservation strategies is required. The restoration of extensive grazing must be considered the primary strategic priority due to its multi-functional impact on vegetation structure, while mechanical clearing and seed transfer serve as essential secondary measures for sites already heavily degraded.
The findings of this study underscore that the primary ecological characteristic of the 6210* and 6120* habitats is the dominance of specialized heliophilous species. These phytocoenoses are characterized by high light requirements and a low tolerance for competitive shading. Our analysis demonstrates that the structural stability of these grasslands is directly contingent upon regular disturbance regimes, such as grazing or mowing, which prevent successional infilling. Without such interventions, the expansion of woody vegetation creates a closed-canopy environment, leading to the rapid loss of photophilous biodiversity. Therefore, maintaining the specific light availability niche is an absolute ecological necessity for the long-term preservation of these anthropogenic ecosystems.
Based on the above conclusions, the following recommendations have been formulated:
•
For managers of protected areas and nature conservation practitioners: The implementation of active conservation programs based on extensive grazing should be prioritized. All conservation actions should be preceded by a detailed natural inventory and conducted in a way that ensures the maintenance of a habitat mosaic. It is necessary to conduct constant monitoring of the effectiveness of the actions undertaken and to flexibly adapt conservation plans. Habitat mosaic is key to maintaining diversity, as it provides a variety of microhabitats, thus supporting species richness. Effective monitoring and adaptive management are essential to actively adjust conservation efforts to changing conditions. Monitoring should assess specific indicators, and management should respond flexibly to the results to ensure the durability of ecosystems;
•
The decline in profitability of traditional management practices driven by rural to urban migration, falling prices of agricultural products (e.g., wool and milk) and rising labor costs has led to widespread abandonment. As a result, many grasslands have been abandoned, which has led to secondary succession, woody species encroachment and the loss of their functional and structural integrity as open grassland habitats. Therefore, to ensure the sustainability of these ecosystems, it is necessary to create new, attractive support mechanisms. They must compensate for financial losses and make extensive grassland management profitable again for farmers and landowners. For decision-makers and institutions, this means moving beyond temporary grants toward long-term systemic support within the Common Agricultural Policy that treats biodiversity as a high-value agricultural product.
•
Furthermore, long-term conservation success depends on fostering social awareness and education. It is essential to develop communication programs that highlight the ecological significance and cultural value of dry grasslands. Educational initiatives should target local communities, landowners and tourists to promote an understanding of active management, e.g., the role of grazing and to build public support for the necessary socio-economic shifts in agricultural policy.
It is crucial to simplify bureaucratic procedures and increase the effectiveness of the implementation of the Natura 2000 network. Landscape-scale conservation should be supported by promoting the maintenance of ecological corridors and implementing stricter spatial planning to limit urban and industrial development pressure on ecologically valuable areas. This includes preventing habitat fragmentation by ensuring that infrastructure projects do not disrupt the connectivity between isolated grassland patches. In the context of grassland conservation, implementing measures at a landscape scale is critical. The conservation of small, isolated fragments, despite their value, is insufficient to ensure the long-term survival of species. Habitat fragmentation and population isolation are major drivers of biodiversity decline. Therefore, it is necessary to implement strategies that connect isolated habitats into ecological networks. The successful implementation of these measures requires the cooperation of multiple stakeholders (farmers, local authorities, NGOs) and the inclusion of ecological corridors. Case studies from successful projects (e.g., “LIFE to alvars” in Estonia [189] or “Trockenrasen Saar” in Germany [190]) demonstrate that large-scale, coordinated actions can significantly increase the effectiveness of conservation efforts.

Author Contributions

J.W.; software, J.W.; formal analysis, J.W. and M.K.; writing—original draft preparation, J.W.; writing—review and editing, J.W. and M.K. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

No new data were created or analyzed in this study. Data sharing is not applicable to this article.

Conflicts of Interest

The authors declare no conflicts of interest.

References

  1. Blair, J.; Nippert, J.; Briggs, J. Grassland ecology. In Ecology and the Environment; Monson, R.K., Ed.; The Plant Sciences 8; Springer: New York, NY, USA, 2014; pp. 389–423. [Google Scholar] [CrossRef]
  2. Briggs, J.; Knapp, A.; Collins, S. Steppes and Prairies. In Encyclopedia of Ecology, 2nd ed.; Elsevier: Amsterdam, The Netherlands, 2008; Volume 2, pp. 634–642. [Google Scholar] [CrossRef]
  3. Burczyk, P.; Gamrat, R.; Gałczyńska, M.; Saran, E. The role of grasslands in providing ecological sustainability of the natural environment. Water-Environ. Rural Areas 2018, 18, 21–37. [Google Scholar]
  4. Emadodin, I.; Corral, D.E.F.; Reinsch, T.; Kluß, C.; Taube, F. Climate Change Effects on Temperate Grassland and Its Implication for Forage Production: A Case Study from Northern Germany. Agriculture 2021, 11, 232. [Google Scholar] [CrossRef]
  5. Srivastava, P.; Raghubanshi, A.S. Impact of Parthenium hysterophorus L. invasion on soil nitrogen dynamics of grassland vegetation of Indo-Gangetic plains, India. Environ. Monit. Assess. 2021, 193, 286. [Google Scholar] [CrossRef] [PubMed]
  6. Szymura, M.; Świerszcz, S.; Szymura, T.H. Restoration of ecologically valuable grassland on sites degraded by invasive Solidago: Lessons from a 6-year experiment. Land Degrad. Dev. 2022, 33, 1985–1998. [Google Scholar] [CrossRef]
  7. Varga, K.; Halász, A.; Kovács, G.P.; Csízi, I. Investigation of Carbon-Dioxide-Emissions from Underutilized Grassland between 2019 and 2020. Agronomy 2022, 12, 931. [Google Scholar] [CrossRef]
  8. Tälle, M.; Deák, B.; Poschlod, P.; Valkó, O.; Westerberg, L.; Milberg, P. Grazing vs. mowing: A meta-analysis of biodiversity benefits for grassland management. Agric. Ecosyst. Environ. 2016, 222, 200–212. [Google Scholar] [CrossRef]
  9. Klichowska, E.; Szczecińska, M.; Ślipiko, M.; Nobis, M. Different but valuable: Anthropogenic habitats as genetic diversity reservoirs for endangered dry grassland species—A case study of Stipa pennata. Ecol. Indic. 2020, 111, 105998. [Google Scholar] [CrossRef]
  10. Barańska, K.; Żmihorski, M.; Pluciński, P. Report on the Project: Conservation of Xerothermic Grasslands in Poland—Theory and Practice LIFE08 NAT/PL/513; Naturalists’ Club Publishing House: Perth, Australia, 2013; ISBN 978-83-63426-09-5. [Google Scholar]
  11. Bornkamm, R. Fifty years vegetation development of a xerothermic calcareous grassland in Central Europe after heavy disturbance. Flora 2006, 201, 249–267. [Google Scholar] [CrossRef]
  12. Hanczaruk, R.; Bąba, W. Threats and protection of valuable natural habitats in Poland on the example of calcareous grasslands. Kosmos 2019, 68, 599–611. [Google Scholar] [CrossRef]
  13. Michalska-Hejduk, D.; Kopeć, D. Dynamics of Semi-Natural Vegetation with a Focus on Molinion Meadows after 50 Years of Strict Protection. Pol. J. Environ. Stud. 2012, 21, 1731–1741. [Google Scholar]
  14. Řehounková, K.; Jongepierová, I.; Šebelíková, L.; Vítovcová, K.; Prach, K. Topsoil removal in degraded open sandy grasslands: Can we restore threatened vegetation fast? Restor. Ecol. 2020, 29, e13188. [Google Scholar] [CrossRef]
  15. Rosenthal, G. Secondary succession in a fallow central European wet grassland. Flora 2010, 205, 153–160. [Google Scholar] [CrossRef]
  16. Banach, M. Maintenance of biodiversity of xerothermic habitats in Małopolska. Stud. Mater. CEPL 2010, 12, 248–255. [Google Scholar]
  17. Futa, B.; Kulik, M.; Kajtoch, Ł.; Mazur, M.A.; Jaźwa, M.; Ścibior, R.; Wielgos, J. Enzymatic Activity of Soil on the Occurrence of the Endangered Beetle Cheilotoma musciformis (Coleoptera: Chrysomelidae) in Xerothermic Grasslands. Insects 2024, 15, 307. [Google Scholar] [CrossRef] [PubMed]
  18. General Directorate for Environmental Protection. Natura 2000 Standard Data Form: PLH240009 Ostoja Środkowojurajska. Available online: https://natura2000.egov.bg/EsriBg.Natura.Public.Web.App/PublicDownloads/SDF/Site_BG0000396.pdf (accessed on 5 April 2026).
  19. Sowa, S.; Kulik, M.; Koroluk, A.; Toporowska, J.; Marek, E.; Szewczyk, W.; Szewczyk, M.; Krzysztof, K.; Paczos-GrzęDa, E. Genetic structure of Carlina acanthifolia subsp. utzka populations on the north-western margins of the species range. Glob. Ecol. Conserv. 2020, 24, e01225. [Google Scholar] [CrossRef]
  20. Wala, M.; Kołodziejek, J.; Wilk, T. Acidity and availability of aluminum, iron and manganese as factors affecting germination in European acidic dry and alkaline xerothermic grasslands. PeerJ 2022, 10, e13255. [Google Scholar] [CrossRef]
  21. Banaszak, J.; Twerd, L. Importance of thermophilous habitats for protection of wild bees (Apiformes). Community Ecol. 2018, 19, 239–247. [Google Scholar] [CrossRef]
  22. Gelabert, P.; Rodrigues, M.; de la Riva, J.; Ameztegui, A.; Sebastià, M.; Vega-Garcia, C. LandTrendr smoothed spectral profiles enhance woody encroachment monitoring. Remote. Sens. Environ. 2021, 262, 112521. [Google Scholar] [CrossRef]
  23. Tiebel, M.; Mölder, A.; Plieninger, T. Small-scale private forest owners and the European Natura 2000 conservation network: Perceived ecosystem services, management practices, and nature conservation attitudes. Eur. J. For. Res. 2021, 140, 1515–1531. [Google Scholar] [CrossRef]
  24. Pent, G.J. Over-yielding in temperate silvopastures: A meta-analysis. Agrofor. Syst. 2020, 94, 1741–1758. [Google Scholar] [CrossRef]
  25. Page, M.J.; Moher, D.; Bossuyt, P.M.; Boutron, I.; Hoffmann, T.C.; Mulrow, C.D.; Shamseer, L.; Tetzlaff, J.M.; Akl, E.A.; Brennan, S.E.; et al. PRISMA 2020 explanation and elaboration: Updated guidance and exemplars for reporting systematic reviews. BMJ 2021, 372, n160. [Google Scholar] [PubMed]
  26. Apostolova, I.; Dengler, J.; Di Pietro, R.; Gavilán, R.G.; Tsiripidis, I. Dry Grasslands of Southern Europe: Syntaxonomy, Management and Conservation. Hacquetia 2014, 13, 5–18. [Google Scholar] [CrossRef]
  27. Huber, S.; Huber, B.; Stahl, S.; Schmid, C.; Reisch, C. Species diversity of remnant calcareous grasslands in south eastern Germany depends on litter cover and landscape structure. Acta Oecol 2017, 83, 48–55. [Google Scholar] [CrossRef]
  28. Rendeková, A.; Miškovic, J.; Mičieta, K.; Michalová, M.; Eliašová, M.; Ballová, D. Changes in the Diversity of Dry Calcareous Grasslands After Abandonment of Traditional Management in Devínska Kobyla National Nature Reserve (Southwestern Slovakia, Central Europe). Ekológia 2020, 39, 101–115. [Google Scholar] [CrossRef]
  29. Ballesteros, M.; Řehounková, K.; Šebelíková, L.; Müllerová, A.; Vítovcová, K.; Prach, K. Participation of grassland species in various successional series in a temperate European region and implications for habitat management. Glob. Ecol. Conserv. 2023, 49, e02761. [Google Scholar] [CrossRef]
  30. Jacquemyn, H.; Van Mechelen, C.; Brys, R.; Honnay, O. Management effects on the vegetation and soil seed bank of calcareous grasslands: An 11-year experiment. Biol. Conserv. 2011, 144, 416–422. [Google Scholar] [CrossRef]
  31. Rico, Y.; Boehmer, H.J.; Wagner, H.H. Effect of Rotational Shepherding on Demographic and Genetic Connectivity of Calcareous Grassland Plants. Conserv. Biol. 2014, 28, 467–477. [Google Scholar] [CrossRef]
  32. Balmer, O.; Erhardt, A. Consequences of Succession on Extensively Grazed Grasslands for Central European Butterfly Communities: Rethinking Conservation Practices. Conserv. Biol. 2000, 14, 746–757. [Google Scholar] [CrossRef]
  33. Janišová, M.; Bartha, S.; Kiehl, K.; Dengler, J. Advances in the conservation of dry grasslands: Introduction to contributions from the seventh European Dry Grassland Meeting. Plant Biosyst. 2011, 145, 507–513. [Google Scholar] [CrossRef]
  34. Ridding, L.E.; Redhead, J.W.; Upcott, E.V.; Hulmes, L.; Hulmes, S.; Mitschunas, N.; Howell, K.; Christelow, J.; Wickenden, A.R.; Fussell, M.; et al. Practical methods for the control of tor-grass (Brachypodium pinnatum s.l.) and the restoration of calcareous grassland. J. Nat. Conserv. 2024, 78, 126566. [Google Scholar] [CrossRef]
  35. Barbaro, L.; Dutoit, T.; Cozic, P. A six-year experimental restoration of biodiversity by shrub-clearing and grazing in calcareous grasslands of the French Prealps. Biodivers. Conserv. 2001, 10, 119–135. [Google Scholar] [CrossRef]
  36. Janišová, M.; Wellstein, C.; Willner, W.; Dengler, J. Succession, restoration, and management of dry grasslands: Special Feature with contributions from the 7th European Dry Grassland Meeting 2010 in Smolenice. Tuexenia 2011, 31, 227–234. [Google Scholar]
  37. Roleček, J.; Čornej, I.I.; Tokarjuk, A.I. Understanding the extreme species richness of semi-dry grasslands in east-central Europe: A comparative approach. Preslia 2014, 86, 13–34. [Google Scholar]
  38. Barbaro, L.; Dutoit, T.; Anthelme, F.; Corcket, E. Respective influence of habitat conditions and management regimes on prealpine calcareous grasslands. J. Environ. Manag. 2004, 72, 261–275. [Google Scholar] [CrossRef]
  39. Janišová, M.; Magnes, M.; Devánová, K.; Dúbravková, D. Mid-term changes of dry grasslands with conservation management in the Hainburger Berge Mountains. Biologia 2024, 79, 1091–1112. [Google Scholar] [CrossRef]
  40. Römermann, C.; Tackenberg, O.; Jackel, A.-K.; Poschlod, P. Eutrophication and fragmentation are related to species’ rate of decline but not to species rarity: Results from a functional approach. Biodivers. Conserv. 2008, 17, 591–604. [Google Scholar] [CrossRef]
  41. Bauer, M.; Albrecht, H. Vegetation monitoring in a 100-year-old calcareous grassland reserve in Germany. Basic Appl. Ecol. 2019, 42, 15–26. [Google Scholar] [CrossRef]
  42. Jaunatre, R.; Buisson, E.; Dutoit, T. First-year results of a multi-treatment steppe restoration experiment in La Crau (Provence, France). Plant Ecol. Evol. 2012, 145, 13–23. [Google Scholar] [CrossRef]
  43. Rosenthal, G.; Schrautzer, J.; Eichberg, C. Low-intensity grazing with domestic herbivores: A tool for maintaining and restoring plant diversity in temperate Europe. Tuexenia 2012, 32, 167–205. [Google Scholar]
  44. Bennie, J.; Hill, M.O.; Baxter, R.; Huntley, B. Influence of slope and aspect on long-term vegetation change in British chalk grasslands. J. Ecol. 2006, 94, 355–368. [Google Scholar] [CrossRef]
  45. Joshi, J.; Stoll, P.; Rusterholz, H.-P.; Schmid, B.; Dolt, C.; Baur, B. Small-scale experimental habitat fragmentation reduces colonization rates in species-rich grasslands. Oecologia 2006, 148, 144–152. [Google Scholar] [CrossRef] [PubMed]
  46. Ruprecht, E.; Enyedi, M.Z.; Szabó, A.; Fenesi, A. Biomass removal by clipping and raking vs burning for the restoration of abandoned Stipa-dominated European steppe-like grassland. Appl. Veg. Sci. 2016, 19, 78–88. [Google Scholar] [CrossRef]
  47. Berki, B.; Botta-Dukát, Z.; Csákvári, E.; Gyalus, A.; Halassy, M.; Mártonffy, A.; Rédei, T.; Csecserits, A. Short-term effects of the control of the invasive plant Asclepias syriaca: Secondary invasion of other neophytes instead of recovery of the native species. Appl. Veg. Sci. 2023, 26, e12707. [Google Scholar] [CrossRef]
  48. Kahmen, S.; Poschlod, P. Plant functional trait responses to grassland succession over 25 years. J. Veg. Sci. 2004, 15, 21–32. [Google Scholar] [CrossRef]
  49. Rupprecht, D.; Gilhaus, K.; Hölzel, N. Effects of year-round grazing on the vegetation of nutrient-poor grass- and heathlands—Evidence from a large-scale survey. Agric. Ecosyst. Environ. 2016, 234, 16–22. [Google Scholar] [CrossRef]
  50. Berry, P.M.; Dawson, T.P.; Harrison, P.A.; Pearson, R.G. Modelling potential impacts of climate change on the bio-climatic envelope of species in Britain and Ireland. Glob. Ecol. Biogeogr. 2002, 11, 453–462. [Google Scholar] [CrossRef]
  51. Kahmen, S.; Poschlod, P. Effects of grassland management on plant functional trait composition. Agric. Ecosyst. Environ. 2008, 128, 137–145. [Google Scholar] [CrossRef]
  52. Rysiak, A.; Chabuz, W.; Sawicka-Zugaj, W.; Zdulski, J.; Grzywaczewski, G.; Kulik, M. Comparative impacts of grazing and mowing on the floristics of grasslands in the buffer zone of Polesie National Park, eastern Poland. Glob. Ecol. Conserv. 2021, 27, e01612. [Google Scholar] [CrossRef]
  53. Bobbink, R.; Hicks, K.; Galloway, J.; Spranger, T.; Alkemade, R.; Ashmore, M.; Bustamante, M.; Cinderby, S.; Davidson, E.; Dentener, F.; et al. Global assessment of nitrogen deposition effects on terrestrial plant diversity: A synthesis. Ecol. Appl. 2010, 20, 30–59. [Google Scholar] [CrossRef]
  54. Kaligarič, M.; Brecl, J.; Škornik, S. High potential of sub-Mediterranean dry grasslands for sheep epizoochory. Open Life Sci. 2016, 11, 177–184. [Google Scholar] [CrossRef][Green Version]
  55. Sărățeanu, V.; Cotuna, O.; Paraschivu, M.; Cojocariu, L.L.; Horablaga, N.M.; Rechițean, D.; Mircov, V.D.; Sălceanu, C.; Urlică, A.A.; Copăcean, L. Features of Natural Succession of Ex-Arable Forest Steppe Grassland (from Western Romania) under the Influence of Climate. Plants 2023, 12, 1204. [Google Scholar] [CrossRef] [PubMed]
  56. Kapás, R.E.; Plue, J.; Kimberley, A.; Cousins, S.A.O. Grazing livestock increases both vegetation and seed bank diversity in remnant and restored grasslands. J. Veg. Sci. 2020, 31, 1053–1065. [Google Scholar] [CrossRef]
  57. Schmid, B.C.; Poschlod, P.; Prentice, H.C. The contribution of successional grasslands to the conservation of semi-natural grasslands species—A landscape perspective. Biol. Conserv. 2017, 206, 112–119. [Google Scholar] [CrossRef]
  58. Boschi, C.; Baur, B. The effect of horse, cattle and sheep grazing on the diversity and abundance of land snails in nutrient-poor calcareous grasslands. Basic Appl. Ecol. 2007, 8, 55–65. [Google Scholar] [CrossRef]
  59. Kasari, L.; Saar, L.; de Bello, F.; Takkis, K.; Helm, A. Hybrid ecosystems can contribute to local biodiversity conservation. Biodivers. Conserv. 2016, 25, 3023–3041. [Google Scholar] [CrossRef]
  60. Schmitt, T.; Rákosy, L. Changes of traditional agrarian landscapes and their conservation implications: A case study of butterflies in Romania. Divers. Distrib. 2007, 13, 855–862. [Google Scholar] [CrossRef]
  61. Boschi, C.; Baur, B. Past pasture management affects the land snail diversity in nutrient-poor calcareous grasslands. Basic Appl. Ecol. 2008, 9, 752–761. [Google Scholar] [CrossRef]
  62. Kettermann, M.; Fartmann, T.; Poniatowski, D. Restoration of abandoned calcareous grasslands promotes habitat diversity and overall wild bee beta diversity. Glob. Ecol. Conserv. 2025, 64, e03977. [Google Scholar] [CrossRef]
  63. Schwarz, C.; Fartmann, T. Conservation of a strongly declining butterfly species depends on traditionally managed grasslands. J. Insect Conserv. 2021, 25, 255–271. [Google Scholar] [CrossRef]
  64. Bossuyt, B.; Hermy, M. The potential of soil seedbanks in the ecological restoration of grassland and heathland communities. Belg. J. Bot. 2003, 136, 23–34. [Google Scholar]
  65. Kiehl, K.; Kirmer, A.; Donath, T.W.; Rasran, L.; Hölzel, N. Species introduction in restoration projects—Evaluation of different techniques for the establishment of semi-natural grasslands in Central and Northwestern Europe. Basic Appl. Ecol. 2010, 11, 285–299. [Google Scholar] [CrossRef]
  66. Shyriaieva, D. Classification, ecological differentiation, and conservation value of Pontic sandy grasslands in the Southern Buh River Basin (Ukraine). Tuexenia 2022, 42, 57–94. [Google Scholar] [CrossRef]
  67. Bossuyt, B.; Honnay, O. Can the seed bank be used for ecological restoration? An overview of seed bank characteristics in European communities. J. Veg. Sci. 2008, 19, 875–884. [Google Scholar] [CrossRef]
  68. Klinkovská, K.; Sperandii, M.G.; Trávníček, B.; Chytrý, M. Significant decline in habitat specialists in semi-dry grasslands over four decades. Biodivers. Conserv. 2024, 33, 161–178. [Google Scholar] [CrossRef]
  69. Silva, V.; Catry, F.X.; Fernandes, P.M.; Rego, F.C.; Paes, P.; Nunes, L.; Caperta, A.D.; Sérgio, C.; Bugalho, M.N. Effects of grazing on plant composition, conservation status and ecosystem services of Natura 2000 shrub-grassland habitat types. Biodivers. Conserv. 2019, 28, 1205–1224. [Google Scholar] [CrossRef]
  70. Braschler, B.; Marini, L.; Thommen, G.H.; Baur, B. Effects of small-scale grassland fragmentation and frequent mowing on population density and species diversity of orthopterans: A long-term study. Ecol. Entomol 2009, 34, 321–329. [Google Scholar] [CrossRef]
  71. Koch, M.A.; Scheriau, C.; Schupfner, M.; Bernhardt, K.-G. Long-term monitoring of the restoration and development of limestone grasslands in north western Germany: Vegetation screening and soil seed bank analysis. Flora 2011, 206, 52–65. [Google Scholar] [CrossRef]
  72. Škornik, S. Plant Trait Composition in Lowland Calcareous and Acidic Semi-Dry Grasslands. Diversity 2025, 17, 22. [Google Scholar] [CrossRef]
  73. Bulokhov, A.D.; Petrenko, A.M. Communities of the class Koelerio-Corynephoretea Klika in Klika et Novák 1941 in the Bryansk region of Russia. Veg. Russ. 2017, 30, 29–34. [Google Scholar] [CrossRef]
  74. Köhler, B.; Gigon, A.; Edwards, P.J.; Krüsi, B.; Langenauer, R.; Lüscher, A.; Ryser, P. Changes in the species composition and conservation value of limestone grasslands in Northern Switzerland after 22 years of contrasting managements. Perspect. Plant Ecol. Evol. Syst. 2005, 7, 51–67. [Google Scholar] [CrossRef]
  75. Škornik, S.; Pipenbaher, N. Patterns in clonal traits in semi-dry calcareous grasslands in Slovenia. Preslia 2025, 97, 567–588. [Google Scholar] [CrossRef]
  76. Busch, V.; Klaus, V.H.; Schäfer, D.; Prati, D.; Boch, S.; Müller, J.; Chisté, M.; Mody, K.; Blüthgen, N.; Fischer, M.; et al. Will I stay or will I go? Plant species-specific response and tolerance to high land-use intensity in temperate grassland ecosystems. J. Veg. Sci. 2019, 30, 674–686. [Google Scholar] [CrossRef]
  77. Köhler, M.; Elias, D.; Hiller, G.; Hölzel, N.; Tischew, S. Restoration of orchid-rich dry calcareous grasslands by rotational goat pasturing. Tuexenia 2020, 40, 201–223. [Google Scholar] [CrossRef]
  78. Socher, S.A.; Prati, D.; Boch, S.; Müller, J.; Baumbach, H.; Gockel, S.; Hemp, A.; Schöning, I.; Wells, K.; Buscot, F.; et al. Interacting effects of fertilization, mowing and grazing on plant species diversity of 1500 grasslands in Germany differ between regions. Basic Appl. Ecol. 2013, 14, 126–136. [Google Scholar] [CrossRef]
  79. Butaye, J.; Adriaens, D.; Honnay, O. Conservation and restoration of calcareous grasslands: A concise review of the effects of fragmentation and management on plant species. Biotechnol. Agron. Soc. Environ. 2005, 9, 111–118. [Google Scholar]
  80. Kormann, U.; Rösch, V.; Batáry, P.; Tscharntke, T.; Orci, K.M.; Samu, F.; Scherber, C. Local and landscape management drive trait-mediated biodiversity of nine taxa on small grassland fragments. Divers. Distrib. 2015, 21, 1204–1217. [Google Scholar] [CrossRef]
  81. Stadler, J.; Trefflich, A.; Brandl, R.; Klotz, S. Spontaneous regeneration of dry grasslands on set-aside fields. Biodivers. Conserv. 2007, 16, 621–630. [Google Scholar] [CrossRef]
  82. Cremene, C.; Groza, G.; Rakosy, L.; Schileyko, A.A.; Baur, A.; Erhardt, A.; Baur, B. Alterations of Steppe-Like Grasslands in Eastern Europe: A Threat to Regional Biodiversity Hotspots. Conserv. Biol. 2005, 19, 1606–1618. [Google Scholar] [CrossRef]
  83. Krauss, J.; Bommarco, R.; Guardiola, M.; Heikkinen, R.K.; Helm, A.; Kuussaari, M.; Lindborg, R.; Öckinger, E.; Pärtel, M.; Pino, J.; et al. Habitat fragmentation causes immediate and time-delayed biodiversity loss at different trophic levels. Ecol. Lett. 2010, 13, 597–605. [Google Scholar] [CrossRef]
  84. Storm, C.; Cezanne, R.; Eichler, M.; Schwabe, A. Multi-stress-affected sandy grasslands: Livestock-grazing as a tool for nature conservation under the impact of drought events and rabbit population fluctuations? Tuexenia 2019, 39, 215–248. [Google Scholar] [CrossRef]
  85. Danko, H.; Gärtner, J.; Kasper, F.; Heinken, T.; Dengler, J.; Karrenberg, S. Vegetation change in dry grasslands in Northeast Germany over two decades: A resurvey. Biol. Conserv. 2026, 313, 111494. [Google Scholar] [CrossRef]
  86. Kuhn, T.; Domokos, P.; Kiss, R.; Ruprecht, E. Grassland management and land use history shape species composition and diversity in Transylvanian semi-natural grasslands. Appl. Veg. Sci. 2021, 24, e12585. [Google Scholar] [CrossRef]
  87. Storm, C.; Stroh, M.; Schwabe, A. Restoration of Koelerion glaucae vegetation in the Upper Rhine valley (Hesse, Germany): An update on the development of a model site after 19 years. Tuexenia 2024, 44, 67–101. [Google Scholar] [CrossRef]
  88. Dekoninck, W.; De Koninck, H.; Baugnée, J.-Y.; Maelfait, J.-P. Ant biodiversity conservation in Belgian calcareous grasslands: Active management is vital. Belg. J. Zool. 2007, 137, 137–146. [Google Scholar]
  89. Kulik, M.; Patkowski, K.; Warda, M.; Lipiec, A.; Bojar, W.; Gruszecki, T.M. Assessment of biomass nutritive value in the context of animal welfare and conservation of selected Natura 2000 habitats (4030, 6120 and 6210) in eastern Poland. Glob. Ecol. Conserv. 2019, 19, e00675. [Google Scholar] [CrossRef]
  90. Szabó, A.; Ruprecht, E. Restoration possibilities of dry grasslands afforested by pine: The role of seed bank and remnant vegetation. Tuexenia 2018, 38, 405–418. [Google Scholar] [CrossRef]
  91. Dengler, J.; Biurrun, I.; Boch, S.; Dembicz, I.; Török, P. Grasslands of the palaearctic biogeographic realm: Introduction and synthesis. In The Encyclopedia of the World’s Biomes; Elsevier: Amsterdam, The Netherlands, 2020; pp. 617–637. [Google Scholar]
  92. Kun, R.; Babai, D.; Csathó, A.I.; Erdélyi, A.; Hartdégen, J.; Lengyel, A.; Kálmán, N.; Mártonffy, A.; Hábenczyus, A.A.; Szegleti, Z.; et al. Effects of management complexity on the composition, plant functional dominance relationships and physiognomy of high nature value grasslands. Nat. Conserv. 2024, 55, 1–19. [Google Scholar] [CrossRef]
  93. Szabó, Á.R.; Ernst, L.M.; Gallé, R.; Batáry, P. Grassland type and presence of management shape butterfly functional diversity in agricultural and forested landscapes. Glob. Ecol. Conserv. 2022, 35, e02096. [Google Scholar] [CrossRef]
  94. Dengler, J.; Janišová, M.; Török, P.; Wellstein, C. Biodiversity of Palaearctic grasslands: A synthesis. Agric. Ecosyst. Environ. 2014, 182, 1–14. [Google Scholar] [CrossRef]
  95. Lanta, V.; Doležal, J.; Lantová, P.; Kelíšek, J.; Mudrák, O. Effects of pasture management and fertilizer regimes on botanical changes in species-rich mountain calcareous grassland in Central Europe. Grass Forage Sci. 2009, 64, 443–453. [Google Scholar] [CrossRef]
  96. Dmytrash-Vatseba, I.I.; Shumska, N.V. Dynamics of plant cover of meadow steppes after the cessation of traditional management in Opillia. Biosyst. Divers. 2020, 28, 224–229. [Google Scholar] [CrossRef]
  97. Lehmair, T.A.; Pagel, E.; Poschlod, P.; Reisch, C. Surrounding landscape structures, rather than habitat age, drive genetic variation of typical calcareous grassland plant species. Landsc. Ecol. 2020, 35, 2881–2893. [Google Scholar] [CrossRef]
  98. Teleki, B.; Sonkoly, J.; Erdős, L.; Tóthmérész, B.; Prommer, M.; Török, P. High resistance of plant biodiversity to moderate native woody encroachment in loess steppe grassland fragments. Appl. Veg. Sci. 2020, 23, 175–184. [Google Scholar] [CrossRef]
  99. Dolek, M.; Geyer, A. Conserving biodiversity on calcareous grasslands in the Franconian Jura by grazing: A comprehensive approach. Biol. Conserv. 2002, 104, 351–360. [Google Scholar] [CrossRef]
  100. Löffler, F.; Poniatowski, D.; Fartmann, T. Orthoptera community shifts in response to land-use and climate change—Lessons from a long-term study across different grassland habitats. Biol. Conserv. 2019, 236, 315–323. [Google Scholar] [CrossRef]
  101. Timmermann, A.; Damgaard, C.; Strandberg, M.; Svenning, J. Pervasive early 21st-century vegetation changes across Danish semi-natural ecosystems: More losers than winners and a shift towards competitive, tall-growing species. J. Appl. Ecol. 2015, 52, 21–30. [Google Scholar] [CrossRef]
  102. Duckworth, J.C.; Bunce, R.G.H.; Malloch, A.J.C. Modelling the potential effects of climate change on calcareous grasslands in Atlantic Europe. J. Biogeogr. 2000, 27, 347–358. [Google Scholar] [CrossRef]
  103. Loos, J.; Krauss, J.; Lyons, A.; Föst, S.; Ohlendorf, C.; Racky, S.; Röder, M.; Hudel, L.; Herfert, V.; Tscharntke, T. Local and landscape responses of biodiversity in calcareous grasslands. Biodivers. Conserv. 2021, 30, 2415–2432. [Google Scholar] [CrossRef]
  104. Peciña, M.V.; Ward, R.D.; Bunce, R.G.; Sepp, K.; Kuusemets, V.; Luuk, O. Country-scale mapping of ecosystem services provided by semi-natural grasslands. Sci. Total. Environ. 2019, 661, 212–225. [Google Scholar] [CrossRef]
  105. Duckworth, J.; Bunce, R.; Malloch, A. Vegetation-environment relationships in Atlantic European calcareous grasslands. J. Veg. Sci. 2000, 11, 15–22. [Google Scholar] [CrossRef]
  106. Lyons, A.; Ashton, P.A.; Powell, I.; Oxbrough, A. Impacts of contrasting conservation grazing management on plants and carabid beetles in upland calcareous grasslands. Agric. Ecosyst. Environ. 2017, 244, 22–31. [Google Scholar] [CrossRef]
  107. Vološčuk, I.; Uhliarová, E.; Sabo, P.; Škodová, M.; Švajda, J. Succession dynamics of vegetation in the Slovak Karst Biosphere Reserve Landscape (Western Carpathians). Ekologia 2016, 35, 13–31. [Google Scholar] [CrossRef]
  108. Dullinger, S.; Dirnböck, T.; Greimler, J.; Grabherr, G. A resampling approach for evaluating effects of pasture abandonment on subalpine plant species diversity. J. Veg. Sci. 2003, 14, 243–252. [Google Scholar] [CrossRef]
  109. Maalouf, J.; Le Bagousse-Pinguet, Y.; Marchand, L.; Bâchelier, E.; Touzard, B.; Michalet, R. Integrating climate change into calcareous grassland management. J. Appl. Ecol. 2012, 49, 795–802. [Google Scholar] [CrossRef]
  110. Wagner, H.H.; Rico, Y.; Lehnert, H.; Boehmer, H.J. Process-based long-term evaluation of an ecological network of calcareous grasslands connected by sheep herding. Ecography 2013, 36, 374–382. [Google Scholar] [CrossRef]
  111. Dutoit, T.; Forey, E.; Römermann, C.; Buisson, E.; Fadda, S.; Saatkamp, A.; Gaignard, P.; Trivelly, E. Rémanence des utilisations anciennes et gestion conservatoire des pelouses calcicoles en France. Biotechnol. Agron. Soc. Environ. 2005, 9, 125–132. [Google Scholar]
  112. Maccherini, S.; Bacaro, G.; Favilli, L.; Piazzini, S.; Santi, E.; Marignani, M. Congruence among vascular plants and butterflies in the evaluation of grassland restoration success. Acta Oecol 2009, 35, 311–317. [Google Scholar] [CrossRef]
  113. Walker, K.J.; Stevens, P.A.; Stevens, D.P.; Mountford, J.; Manchester, S.J.; Pywell, R.F. The restoration and re-creation of species-rich lowland grassland on land formerly managed for intensive agriculture in the UK. Biol. Conserv. 2004, 119, 1–18. [Google Scholar] [CrossRef]
  114. Dzwonko, Z.; Loster, S. Changes in plant species composition in abandoned and restored limestone grassiands—the effects of tree and shrub cutting. Acta Soc. Bot. Pol. 2008, 77, 67–75. [Google Scholar] [CrossRef][Green Version]
  115. Marriott, C.A.; Fothergill, M.; Jeangros, B.; Scotton, M.; Louault, F. Long-term impacts of extensification of grassland management on biodiversity and productivity in upland areas. A review. Agron. Sustain. Dev. 2004, 24, 447–462. [Google Scholar] [CrossRef]
  116. WallisDeVries, M.F.; Poschlod, P.; Willems, J.H. Challenges for the conservation of calcareous grasslands in northwestern Europe: Integrating the requirements of flora and fauna. Biol. Conserv. 2002, 104, 265–273. [Google Scholar] [CrossRef]
  117. Dzwonko, Z.; Loster, S. Changes in species composition of an abandoned limestone grassland in relation to climatic conditions. Pol. J. Ecol. 2001, 59, 687–698. [Google Scholar]
  118. Mazalla, L.; Diekmann, M.; Duprè, C. Microclimate shapes vegetation response to drought in calcareous grasslands. Appl. Veg. Sci. 2022, 25, e12672. [Google Scholar] [CrossRef]
  119. Wenzel, M.; Schmitt, T.; Weitzel, M.; Seitz, A. The severe decline of butterflies on western German calcareous grasslands during the last 30 years: A conservation problem. Biol. Conserv. 2006, 128, 542–552. [Google Scholar] [CrossRef]
  120. Elias, D.; Hölzel, N.; Tischew, S. Goat paddock grazing improves the conservation status of shrub-encroached dry grasslands. Tuexenia 2018, 38, 215–233. [Google Scholar] [CrossRef]
  121. Meier, T.; Hensen, I.; Partzsch, M. Floristic changes of xerothermic grasslands in Central Germany: A resurvey study based on quasi-permanent plots. Tuexenia 2021, 41, 203–226. [Google Scholar] [CrossRef]
  122. Wesche, K.; Ambarlı, D.; Kamp, J.; Török, P.; Treiber, J.; Dengler, J. The Palaearctic steppe biome: A new synthesis. Biodivers. Conserv. 2016, 25, 2197–2231. [Google Scholar] [CrossRef]
  123. Elliott, T.; Thompson, A.; Klein, A.; Albert, C.; Eisenhauer, N.; Jansen, F.; Schneider, A.; Sommer, M.; Straka, T.; Settele, J.; et al. Abandoning grassland management negatively influences plant but not bird or insect biodiversity in Europe. Conserv. Sci. Pr. 2023, 5, e13008. [Google Scholar] [CrossRef]
  124. Milberg, P.; Tälle, M. Maintaining an open landscape: Comparison of management methods for semi-natural grasslands: A Swedish multi-site study. Glob. Ecol. Conserv. 2023, 48, e02721. [Google Scholar] [CrossRef]
  125. Wiezik, M.; Svitok, M.; Wieziková, A.; Dovčiak, M. Shrub encroachment alters composition and diversity of ant communities in abandoned grasslands of western Carpathians. Biodivers. Conserv. 2013, 22, 2305–2320. [Google Scholar] [CrossRef]
  126. Fabšičová, M.; Vymyslický, T.; Frei, I.; Zdražílková, M.; Smetanová, S.; Winkler, J.; Jiroušek, M. The importance of soil seed banks for biodiversity restoration in degraded grasslands. Folia Geobot. 2024, 59, 17–37. [Google Scholar] [CrossRef]
  127. Milić, D.; Rat, M.; Bokić, B.; Mudri-Stojnić, S.; Milošević, N.; Sukur, N.; Jakovetić, D.; Radak, B.; Tot, T.; Vujanović, D.; et al. Exploring the effects of habitat management on grassland biodiversity: A case study from northern Serbia. PLoS ONE 2024, 19, e0301391. [Google Scholar] [CrossRef]
  128. Willems, J.H. Problems, Approaches, and Results in Restoration of Dutch Calcareous Grassland During the Last 30 Years. Restor. Ecol. 2001, 9, 147–154. [Google Scholar] [CrossRef]
  129. Fagan, K.C.; Pywell, R.F.; Bullock, J.M.; Marrs, R.H. Do restored calcareous grasslands on former arable fields resemble ancient targets? The effect of time, methods and environment on outcomes. J. Appl. Ecol. 2008, 45, 1293–1303. [Google Scholar] [CrossRef]
  130. Mitchley, J.; Xofis, P. Landscape structure and management regime as indicators of calcareous grassland habitat condition and species diversity. J. Nat. Conserv. 2005, 13, 171–183. [Google Scholar] [CrossRef]
  131. Woodcock, B.; Pywell, R.; Roy, D.; Rose, R.; Bell, D. Grazing management of calcareous grasslands and its implications for the conservation of beetle communities. Biol. Conserv. 2005, 125, 193–202. [Google Scholar] [CrossRef]
  132. Valentin, D.N.; Adamo, M.; Richiardi, C.; Mammola, S.; Kull, T. Impact of climate and land use change on the distribution of orchids in Estonia. Plant Ecol. 2025, 226, 831–844. [Google Scholar] [CrossRef]
  133. Zlinszky, A.; Schroiff, A.; Kania, A.; Deák, B.; Mücke, W.; Vári, Á.; Székely, B.; Pfeifer, N. Categorizing Grassland Vegetation with Full-Waveform Airborne Laser Scanning: A Feasibility Study for Detecting Natura 2000 Habitat Types. Remote. Sens. 2014, 6, 8056–8087. [Google Scholar] [CrossRef]
  134. Godefroid, S.; Le Pajolec, S.; Hechelski, M.; Van Rossum, F. Can we rely on the soil seed bank for restoring xeric sandy calcareous grasslands? Restor. Ecol. 2018, 26, 1084–1094. [Google Scholar] [CrossRef]
  135. Newton, A.C.; Walls, R.M.; Golicher, D.; Keith, S.A.; Diaz, A.; Bullock, J.M. Structure, composition and dynamics of a calcareous grassland metacommunity over a 70-year interval. J. Ecol. 2012, 100, 196–209. [Google Scholar] [CrossRef]
  136. Gorris, P.; Bodin, Ö.; Giralt, D.; Hass, A.L.; Reitalu, T.; Cabodevilla, X.; Hannappel, I.; Helm, A.; Prangel, E.; Westphal, C. Social-ecological perspective on European semi-natural grassland conservation and restoration: Key challenges and future pathways. Biol. Conserv. 2025, 304, 111038. [Google Scholar] [CrossRef]
  137. Pedashenko, H.; Apostolova, I.; Boch, S.; Ganeva, A.; Janišová, M.; Sopotlieva, D.; Todorova, S.; Ünal, A.; Vassilev, K.; Velev, N.; et al. Dry grasslands of NW Bulgarian mountains: First insights into diversity, ecology and syntaxonomy. Tuexenia 2013, 33, 309–346. [Google Scholar]
  138. Greguła-Kania, M.; Nazar, P.; Kulik, M.; Patkowski, K.; Hahaj-Siembida, A.; Junkuszew, A. Some Physiological Responses of Native Sheep Breeds to Environmental Conditions during Grazing in Natura 2000 Habitats. Agriculture 2023, 13, 982. [Google Scholar] [CrossRef]
  139. Petit, S.; Elbersen, B. Assessing the Risk of Impact of Farming Intensification on Calcareous Grasslands in Europe: A Quantitative Implementation of the MIRABEL Framework. AMBIO 2006, 35, 297–303. [Google Scholar] [CrossRef] [PubMed]
  140. Grzędzicka, E.; Vahed, K. Habitat requirements of the endangered heath bush-cricket Gampsocleis glabra (Orthoptera, Tettigoniidae) in an isolated population. J. Insect Conserv. 2020, 24, 935–945. [Google Scholar] [CrossRef]
  141. Pietzsch, D.; Ochsner, S.; Mantilla-Contreras, J.; Hampicke, U. Low-intensity Husbandry as a Cost-efficient Way to Preserve Dry Grasslands. Landsc. Res. 2013, 38, 523–539. [Google Scholar] [CrossRef]
  142. Habel, J.C.; Dengler, J.; Janišová, M.; Török, P.; Wellstein, C.; Wiezik, M. European grassland ecosystems: Threatened hotspots of biodiversity. Biodivers. Conserv. 2013, 22, 2131–2138. [Google Scholar] [CrossRef]
  143. Piqueray, J.; Mahy, G. Revue bibliographique sur la restauration des pelouses calcicoles en Europe: Contraintes rencontrées et solutions proposées. Biotechnol. Agron. Soc. Environ. 2010, 14, 471–484. [Google Scholar]
  144. Habel, J.C.; Ulrich, W.; Biburger, N.; Seibold, S.; Schmitt, T. Agricultural intensification drives butterfly decline. Insect Conserv. Divers. 2019, 12, 289–299. [Google Scholar] [CrossRef]
  145. Podgórska, M.; Łazarski, G. Impact of Secondary Succession in the Xerothermic Grassland on the Population of the Eastern Pasque Flower (Pulsatilla patens)—Preliminary Studies. Sustainability 2021, 13, 12575. [Google Scholar] [CrossRef]
  146. Harásek, M.; Klinkovská, K.; Chytrý, M. Vegetation change in acidic dry grasslands in Moravia (Czech Republic) over three decades: Slow decrease in habitat quality after grazing cessation. Appl. Veg. Sci. 2023, 26, e12726. [Google Scholar] [CrossRef]
  147. Polus, E.; Vandewoestijne, S.; Choutt, J.; Baguette, M. Tracking the effects of one century of habitat loss and fragmentation on calcareous grassland butterfly communities. Biodivers. Conserv. 2007, 16, 3423–3436. [Google Scholar] [CrossRef]
  148. Hejcman, M.; Hejcmanová, P.; Pavlů, V.; Beneš, J. Origin and history of grasslands in Central Europe—A review. Grass Forage Sci. 2013, 68, 345–363. [Google Scholar] [CrossRef]
  149. Poniatowski, D.; Stuhldreher, G.; Helbing, F.; Hamer, U.; Fartmann, T. Restoration of calcareous grasslands: The early successional stage promotes biodiversity. Ecol. Eng. 2020, 151, 105858. [Google Scholar] [CrossRef]
  150. Helbing, F.; Fartmann, T.; Poniatowski, D. Restoration measures foster biodiversity of important primary consumers within calcareous grasslands. Biol. Conserv. 2021, 256, 109058. [Google Scholar] [CrossRef]
  151. Poschlod, P.; WallisDeVries, M.F. The historical and socioeconomic perspective of calcareous grasslands—Lessons from the distant and recent past. Biol. Conserv. 2002, 104, 361–376. [Google Scholar] [CrossRef]
  152. Helm, A.; Hanski, I.; Pärtel, M. Slow response of plant species richness to habitat loss and fragmentation. Ecol. Lett. 2006, 9, 72–77. [Google Scholar] [CrossRef]
  153. Prangel, E.; Reitalu, T.; Kasari-Toussaint, L.; Marja, R.; Jüriado, I.; Kupper, T.; Ingerpuu, N.; Oja, E.; Tiitsaar, A.; Karise, R.; et al. Grassland Restoration Drives Strong Multitrophic Biodiversity Recovery, but Climate Extremes Jeopardize Drought-Sensitive Species. Glob. Chang. Biol. 2025, 31, e70496. [Google Scholar] [CrossRef] [PubMed]
  154. Henning, K.; Lorenz, A.; von Oheimb, G.; Härdtle, W.; Tischew, S. Year-round cattle and horse grazing supports the restoration of abandoned, dry sandy grassland and heathland communities by supressing Calamagrostis epigejos and enhancing species richness. J. Nat. Conserv. 2017, 40, 120–130. [Google Scholar] [CrossRef]
  155. Pywell, R.F.; Bullock, J.M.; Hopkins, A.; Walker, K.J.; Sparks, T.H.; Burke, M.J.; Peel, S. Restoration of species-rich grassland on arable land: Assessing the limiting processes using a multi-site experiment. J. Appl. Ecol. 2002, 39, 294–309. [Google Scholar] [CrossRef]
  156. Hirst, R.A.; Pywell, R.F.; Marrs, R.H.; Putwain, P.D. The resistance of a chalk grassland to disturbance. J. Appl. Ecol. 2003, 40, 368–379. [Google Scholar] [CrossRef]
  157. Raduła, M.W.; Szymura, T.H.; Szymura, M.; Swacha, G.; Kącki, Z. Effect of environmental gradients, habitat continuity and spatial structure on vascular plant species richness in semi-natural grasslands. Agric. Ecosyst. Environ. 2020, 300, 106974. [Google Scholar] [CrossRef]
  158. Hooftman, D.; Kimberley, A.; Cousins, S.A.; Escribano-Avila, G.; Honnay, O.; Krickl, P.; Plue, J.; Poschlod, P.; Traveset, A.; Bullock, J.M. Dispersal limitation, eutrophication and propagule pressure constrain the conservation value of Grassland Green Infrastructure. Biol. Conserv. 2021, 258, 109152. [Google Scholar] [CrossRef]
  159. Rahmonov, O.; Oles, W. Vegetation succession over an area of a medieval ecological disaster. The case of the Bledów Desert, Poland. Erdkunde 2010, 64, 241–255. [Google Scholar] [CrossRef]
  160. Gruszecki, T.M.; Warda, M.; Kulik, M.; Junkuszew, A.; Patkowski, K.; Bojar, W.; Tomczuk, K.; Gregula-Kania, M.; Dudko, P.; Bielińska, E.J.; et al. Sheep grazing as a way to protect the diversity of plant communities in valuable natural habitats. Zootechn. News 2017, 55, 177–184. [Google Scholar]
  161. Cieślak, E.; Drobniak, S. Phylogeography of xerothermic Carlina acanthifolia subsp. utzka in Central Europe. Flora 2019, 253, 76–86. [Google Scholar] [CrossRef]
  162. Lednev, S.; Kasymov, D.; Semenkov, I.; Klink, G.; Agafontsev, M.; Koroleva, T. Fire as a Factor in the Dynamics of Meadow Vegetation: A Model Experiment in Western Siberia. Fire 2024, 7, 115. [Google Scholar] [CrossRef]
  163. Benthien, O.; Braun, M.; Riemann, J.C.; Stolter, C. Long-term effect of sheep and goat grazing on plant diversity in a semi-natural dry grassland habitat. Heliyon 2018, 4, e00556. [Google Scholar] [CrossRef]
  164. Della Rocca, F.; Repetto, E.; De Caria, L.; Milanesi, P. Identifying Ecological Corridors of the Bush Cricket Saga pedo in Fragmented Landscapes. Insects 2025, 16, 279. [Google Scholar] [CrossRef] [PubMed]
  165. Morabito, A.; Musarella, C.M.; Spampinato, G. Diversity and Ecological Assessment of Grasslands Habitat Types: A Case Study in the Calabria Region (Southern Italy). Land 2024, 13, 719. [Google Scholar] [CrossRef]
  166. Calaciura, B.; Spinelli, O. Management of Natura 2000 habitats: 6210 Semi-Natural Dry Grasslands and Scrubland Facies on Calcareous Substrates (Festuco-Brometalia) (Important Orchid Sites). European Commission. ISBN 978-92-79-08335-8. 2008. Available online: https://www.life-heliantheme.eu/fileadmin/Life/Heliantheme/Management_Festuco_Brometea.pdf (accessed on 5 April 2026).
  167. Lawless, P.J.; Baskin, J.M.; Baskin, C.C. Xeric Limestone Prairies of Eastern United States: Review and Synthesis. Bot. Rev. 2006, 72, 235–272. [Google Scholar] [CrossRef]
  168. Byk, A.; Cieślak, R.; Piskorek, W. Persistent isolated population of Onthophagus grossepunctatus REITTER, 1905 (Coleoptera: Scarabaeidae) near Kazimierz Dolny (Poland). Balt. J. Coleopterol. 2020, 20, 101–108. [Google Scholar]
  169. Wójcik, T.; Ziaja, M. Abundance and conservation status of Rosa gallica in Strzyżów Foothills (SE Poland). Ecol. Quest. 2022, 33, 83–90. [Google Scholar] [CrossRef]
  170. Meier, T.; Hensen, I.; Kühn, I. Functional approach to xerothermic grasslands in Central Germany. Preslia 2024, 96, 183–208. [Google Scholar] [CrossRef]
  171. Nienartowicz, A.; Kunz, M.; Adamska, E.; Boińska, U.; Deptula, M.; Gugnacka-Fiedor, W.; Kamiński, D.; Rutkowski, L. Relief and changes in the vegetation cover and the flora of the Zadroże Dune near the city of Toruń: Comparison of the conditions in 1948 and 2009. Ecol. Quest. 2010, 12, 17–51. [Google Scholar] [CrossRef][Green Version]
  172. Markl, G.; Hinneberg, H.; Tarmann, G. Drastic decline of extensive grassland species in Central Europe since 1950: Forester moths of the genus Jordanita (Lepidoptera, Zygaenidae) as a type example. Ecol. Evol. 2022, 12, e9291. [Google Scholar] [CrossRef] [PubMed]
  173. European Environment Agency. Natura 2000 Data—The European Network of Protected Sites. 2024. Available online: https://natura2000.eea.europa.eu/?views=Habitats_View (accessed on 11 July 2025).
  174. Süss, K.; Schwabe, A. Sheep versus donkey grazing or mixed treatment: Results from a 4-year field experiment in Armerio-Festucetum trachyphyllae sand vegetation. Phytocoenologia 2007, 37, 135–160. [Google Scholar] [CrossRef]
  175. Warda, M.; Kulik, M.; Gruszecki, T.; Lipiec, A.; Zubel, R. The natural values of selected dry grasslands under sheep grazing in the Kózki Nature Reserve. Grassl. Sci. Pol. 2015, 18, 229–239. [Google Scholar]
  176. Horka, S.; Jandt, U.; Bruelheide, H. Diversity Increases but Community Specialists Decline Over Three Decades in Dry Grassland Communities of Central Germany. J. Veg. Sci. 2025, 36, e70036. [Google Scholar] [CrossRef]
  177. Kulik, M.; Oleszek, M.; Lipiec, A.; Warda, M.; Gruszecki, T.; Bojar, W.; Król, A. Total protein and macroelement content in selected psammophilous grassland species under free-range sheep grazing in Kózki Nature Reserve. J. Elem. 2017, 22, 183–193. [Google Scholar] [CrossRef]
  178. Mezule, L.; Strazdina, B.; Dalecka, B.; Skripsts, E.; Juhna, T. Natural Grasslands as Lignocellulosic Biofuel Resources: Factors Affecting Fermentable Sugar Production. Energies 2021, 14, 1312. [Google Scholar] [CrossRef]
  179. Demarchi, L.; Kania, A.; Ciężkowski, W.; Piórkowski, H.; Oświecimska-Piasko, Z.; Chormański, J. Recursive Feature Elimination and Random Forest Classification of Natura 2000 Grasslands in Lowland River Valleys of Poland Based on Airborne Hyperspectral and LiDAR Data Fusion. Remote. Sens. 2020, 12, 1842. [Google Scholar] [CrossRef]
  180. EEA Report No 10/2020: State of Nature in the EU. Available online: https://www.eea.europa.eu/en/analysis/publications/state-of-nature-in-the-eu-2020 (accessed on 4 July 2025).
  181. Chief Inspector of Environmental Protection. Nature Monitoring Bulletin No. 31; GIOŚ: Warsaw, Poland, 2025. [Google Scholar]
  182. Naturalists’Club. Shadow List of Natura 2000 Habitats in Poland; Naturalists’ Club: Świebodzin, Poland, 2024. [Google Scholar]
  183. JNCC. Habitats Regulations Reporting 2019–2024: Summary of Conservation Status Assessments for UK Terrestrial and Freshwater Habitats. Joint Nature Conservation Committee, Peterborough. 2024. Available online: https://jncc.gov.uk/our-work/habitats-regulations-reporting-2019-to-2024/ (accessed on 26 March 2026).
  184. Stalenga, J.; Brzezińska, K.; Stańska, M.; Berbeć, A.; Błaszkowska, B.; Czekała, W.; Dach, J.; Dzierża, P.; Ebertowska, B.; Feledyn-Szewczyk, B.; et al. Code of Good Agricultural Practices Promoting Biodiversity. Monograph, 2nd ed.; Publishing House of the Institute of Soil Science and Plant Cultivation—State Research Institute: Puławy, Poland, 2016; ISBN 978-83-7562-238-6. [Google Scholar]
  185. Bremer, L.L.; Farley, K.A.; Chadwick, O.A.; Harden, C.P. Changes in carbon storage with land management promoted by payment for ecosystem services. Environ. Conserv. 2016, 43, 397–406. [Google Scholar] [CrossRef]
  186. Grubb, P.J. The Maintenance of Species-richness in Plant Communities: The Importance of the Regeneration Niche. Biol. Rev. 1977, 52, 107–145. [Google Scholar] [CrossRef]
  187. Pe’Er, G.; Bonn, A.; Bruelheide, H.; Dieker, P.; Eisenhauer, N.; Feindt, P.H.; Hagedorn, G.; Hansjürgens, B.; Herzon, I.; Lomba, Â.; et al. Action needed for the EU Common Agricultural Policy to address sustainability challenges. People Nat. 2020, 2, 305–316. [Google Scholar] [CrossRef]
  188. Hermoso, V.; Villero, D.; Clavero, M.; Brotons, L. Spatial prioritisation of EU’s LIFE-Nature programme to strengthen the conservation impact of Natura 2000. J. Appl. Ecol. 2018, 55, 1502–1512. [Google Scholar] [CrossRef]
  189. European Commission. Restoration of Estonian Alvar Grasslands. Available online: https://webgate.ec.europa.eu/life/publicWebsite/project/LIFE13-NAT-EE-000082/restoration-of-estonian-alvar-grasslands (accessed on 23 September 2025).
  190. European Commission. Regeneration and Preservation of Dry Grassland in Germany. Available online: https://webgate.ec.europa.eu/life/publicWebsite/project/LIFE00-NAT-D-007058/regeneration-and-preservation-of-dry-grassland-in-germany (accessed on 23 September 2025).
Diversity 18 00216 g001
Figure 1. PRISMA flow diagram illustrating the process of selecting articles for a systematic literature review. Source: Page MJ et al. BMJ 2021;372:n71. doi: 10.1136/bmj.n71 [25]. This work is licensed under CC BY 4.0. To view a copy of this license, visit https://creativecommons.org/licenses/by/4.0/, accessed on 16 February 2026.
Figure 1. PRISMA flow diagram illustrating the process of selecting articles for a systematic literature review. Source: Page MJ et al. BMJ 2021;372:n71. doi: 10.1136/bmj.n71 [25]. This work is licensed under CC BY 4.0. To view a copy of this license, visit https://creativecommons.org/licenses/by/4.0/, accessed on 16 February 2026.
Diversity 18 00216 g001
Diversity 18 00216 g002
Figure 2. Xerothermic grassland—Natura 2000 area Kąty PLH060010, Poland (Fot. M. Kulik).
Figure 2. Xerothermic grassland—Natura 2000 area Kąty PLH060010, Poland (Fot. M. Kulik).
Diversity 18 00216 g002
Diversity 18 00216 g003
Figure 3. Semi-natural dry grasslands and scrubland facies on calcareous substrates (Festuco-Brometalia) (* important orchid sites) 6210*. Source: Own study based on data from: Data on Natura 2000 areas—European Environment Agency (EEA) [173], downloaded [11 July 2025], administrative boundaries of Europe—GIS-Support.pl, downloaded [11 July 2025]. Illustration based on free and open-source software QGIS 2.18 (https://www.qgis.org/en/site/), accessed 13 July 2025. Coordinate system (EPSG:3035).
Figure 3. Semi-natural dry grasslands and scrubland facies on calcareous substrates (Festuco-Brometalia) (* important orchid sites) 6210*. Source: Own study based on data from: Data on Natura 2000 areas—European Environment Agency (EEA) [173], downloaded [11 July 2025], administrative boundaries of Europe—GIS-Support.pl, downloaded [11 July 2025]. Illustration based on free and open-source software QGIS 2.18 (https://www.qgis.org/en/site/), accessed 13 July 2025. Coordinate system (EPSG:3035).
Diversity 18 00216 g003
Diversity 18 00216 g004
Figure 4. Xeric sand calcareous grasslands (Koelerion glaucae) 6120* (Fot. J. Wielgos).
Figure 4. Xeric sand calcareous grasslands (Koelerion glaucae) 6120* (Fot. J. Wielgos).
Diversity 18 00216 g004
Diversity 18 00216 g005
Figure 5. Xeric sand calcareous grasslands (Koelerion glaucae) 6120*. Source: Own study based on data from: Data on Natura 2000 areas—European Environment Agency (EEA) [173], downloaded [11 July 2025], administrative boundaries of Europe—GIS-Support.pl, downloaded [11 July 2025]. Illustration based on free and open-source software QGIS 2.18 (https://www.qgis.org/en/site/). Coordinate system (EPSG:3035).
Figure 5. Xeric sand calcareous grasslands (Koelerion glaucae) 6120*. Source: Own study based on data from: Data on Natura 2000 areas—European Environment Agency (EEA) [173], downloaded [11 July 2025], administrative boundaries of Europe—GIS-Support.pl, downloaded [11 July 2025]. Illustration based on free and open-source software QGIS 2.18 (https://www.qgis.org/en/site/). Coordinate system (EPSG:3035).
Diversity 18 00216 g005
Diversity 18 00216 g006
Figure 6. Frequency of identified threats to dry grasslands based on publications in the Web of Science and Scopus database.
Figure 6. Frequency of identified threats to dry grasslands based on publications in the Web of Science and Scopus database.
Diversity 18 00216 g006
Table 1. Matrix of threat category occurrences in publications identified in Web of Science and Scopus databases. A1—Succession and encroachment by trees/shrubs; A2—Abandonment of use and lack of management; A3—Negative agricultural activities; A4—Habitat loss and fragmentation; A5—Climate change; A6—Invasive and alien species; A7—Urbanization and infrastructure development.
Table 1. Matrix of threat category occurrences in publications identified in Web of Science and Scopus databases. A1—Succession and encroachment by trees/shrubs; A2—Abandonment of use and lack of management; A3—Negative agricultural activities; A4—Habitat loss and fragmentation; A5—Climate change; A6—Invasive and alien species; A7—Urbanization and infrastructure development.
PublicationA1A2A3A4A5A6A7PublicationA1A2A3A4A5A6A7PublicationA1A2A3A4A5A6A7
[26]0110011[27]1111000[28]1100010
[29]1101100[30]1110000[31]1111000
[32]1111000[33]1111101[34]1111110
[35]1100010[36]1111111[37]1111000
[38]1111100[39]1100100[40]0011000
[41]0111101[42]0011010[43]1111000
[44]1111100[45]0011000[46]1100000
[47]1111110[48]1110100[49]1110000
[50]1000110[51]1110000[52]1100000
[53]1010110[54]1111000[55]1100100
[11]1110100[56]1101000[57]0111000
[58]1110000[59]1111111[60]1111100
[61]1111000[62]1111100[63]1111101
[64]1111010[65]0011010[66]1111011
[67]1000000[68]1111100[69]1111110
[70]0011000[71]1110000[72]1110100
[73]1000000[74]1110100[75]1111010
[76]0011000[77]1111100[78]0010000
[79]1111000[80]1111000[81]1110000
[82]1111010[83]1111111[84]1000100
[85]1111110[86]1111000[87]1111001
[88]1101011[89]1100000[90]1101010
[91]1111110[92]1011011[93]1111000
[94]1111110[95]1110000[8]1101000
[96]1111010[97]1111101[98]1111000
[99]1101000[100]0110100[101]1101100
[102]0111100[103]1111100[104]1101000
[105]0010100[106]1110000[107]1101100
[108]1101100[109]0110100[110]1111000
[111]1110000[112]1011000[113]1111010
[114]1101000[115]1110000[116]1111100
[117]1100100[118]0010100[119]1111100
[120]1101000[121]1110100[122]1111100
[123]1111001[124]1111000[125]1101010
[126]1111010[127]1111000[128]1111000
[129]0111000[130]1011000[131]1011000
[17]1110000[132]1111101[133]1111000
[134]1111010[135]0011111∑ 137 11511210395523220
[136]1111101[137]0110000
[138]1100100[139]0111100
[140]1101001[141]1110000
[142]1111111[143]1011111
[144]0011000[145]1100000
[146]1100000[147]1101000
[148]1111000[149]1101000
[150]1100000[151]1111000
[152]1111000[153]1111100
[154]1101000[155]1011000
[156]0010000[157]1111010
[158]0011000[159]1000001
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Share and Cite

MDPI and ACS Style

Wielgos, J.; Kulik, M. Semi-Natural Dry Grasslands in Decline: A Review of Characteristics, Threats and Conservation Challenges. Diversity 2026, 18, 216. https://doi.org/10.3390/d18040216

AMA Style

Wielgos J, Kulik M. Semi-Natural Dry Grasslands in Decline: A Review of Characteristics, Threats and Conservation Challenges. Diversity. 2026; 18(4):216. https://doi.org/10.3390/d18040216

Chicago/Turabian Style

Wielgos, Justyna, and Mariusz Kulik. 2026. "Semi-Natural Dry Grasslands in Decline: A Review of Characteristics, Threats and Conservation Challenges" Diversity 18, no. 4: 216. https://doi.org/10.3390/d18040216

APA Style

Wielgos, J., & Kulik, M. (2026). Semi-Natural Dry Grasslands in Decline: A Review of Characteristics, Threats and Conservation Challenges. Diversity, 18(4), 216. https://doi.org/10.3390/d18040216

Note that from the first issue of 2016, this journal uses article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop