52
1  Agricultural Institute of Slovenia,  
Hacquetova ulica 17, 1000 Ljubljana,  
Ljubljana, Slovenia
* Corresponding author:  
E-mail address: mateja.grasic@kis.si
Citation: Grašič, M., Šabić, A., & Lukač, B. 
(2023). A review of methodology for grassland 
restoration and management with practical 
examples. Acta Biologica Slovenica, 66(1).
https://doi.org/10.14720/abs.66.1.13230
This article is an open access article distributed 
under the terms and conditions of the Creative 
Commons Attribution (CC BY SA) license
Review Article
A review of methodology  
for grassland restoration 
with practical examples
Mateja Grašič 1,*, Azra Šabić 1, Branko Lukač 1
Abstract
Currently, the majority of high nature value Slovenian grasslands have an 
unfavourable conservation status. Based on the available data from habitat type 
mappings, the surface of high nature value grasslands (6210(*) – semi-natural 
dry grasslands and scrubland facies on calcareous substrates, 6410 – Molinia 
meadows, and 6510 – lowland hay meadows) at Natura 2000 sites is decreasing. 
The existing agri-environment measures have been only partly effective in 
promoting grassland biodiversity. The main threats to biodiversity are driven 
by various anthropogenic activities, which result in a continuous change in 
landscape identity, habitat fragmentation, and ecosystem degradation. Therefore, 
biodiversity restoration became an urgent step in the conservation of high nature 
value grasslands. Multiple pathways may lead to the ecological restoration of 
grasslands with an altered, uncharacteristic floristic composition, or grasslands 
with an unfavourable conservation status. In this paper, we present an overview of 
the methods used in the restoration of grasslands from different parts of Europe. 
In an adapted form, these methods could also be used for the restoration of 
Slovenian grasslands. Grasslands may be left to spontaneous succession, which 
is mainly suitable for small-scale areas located in the proximity of grasslands 
with well-preserved biodiversity. However, to re-establish plant communities 
with specialist species, a more proactive approach is usually required, such as 
sowing of regional or commercial seed preservation mixtures, transfer of mature 
plant material, or topsoil transfer from donor sites with appropriate botanical 
composition. Grassland restoration methods should be carefully thought-out 
and carried out before the habitat or species is endangered. We conclude that 
optimally chosen post-restoration management may have an impact that is 
comparable to or even greater than the impact of a suitable restoration method. 
Nevertheless, the maintenance of well-preserved grasslands is still much more 
cost-effective than the restoration of degraded grasslands.
Keywords 
grassland restoration, management, methodology, Natura 2000 grassland 
habitat types, agri-environment measures
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Acta Biologica Slovenica, 2023, 66 (1)
Pregled metodologije obnove travišč s primeri iz prakse
Izvleček
Večina slovenskih travišč z visoko naravno vrednostjo ima trenutno neugoden ohranitveni status. Glede na 
razpoložljive podatke s kartiranj habitatnih tipov se površine travinja z visoko naravno vrednostjo (6210(*) – polnaravna 
suha travišča in grmiščne faze, 6410 – travniki s prevladujočo modro stožko in 6510 – nižinski gojeni travniki) na 
območjih Natura 2000 zmanjšujejo. Predlagani ukrepi v preteklih kmetijsko-okoljskih programskih obdobjih so bili 
le delno učinkoviti pri ohranjanju biotske raznovrstnosti travišč. Glavno grožnjo biotski raznovrstnosti predstavljajo 
različne antropogene aktivnosti, ki vodijo v stalno spreminjanje krajine, razdrobljenost habitatov in degradacijo 
ekosistemov. Obnova biotske raznovrstnosti je zato postala nujen ukrep za ohranjanje travišč z visoko naravno 
vrednostjo. Do ekološke obnove travišč s spremenjeno, neznačilno floristično sestavo, ali pa travišč v neugodnem 
naravovarstvenem stanju, vodi več poti. V tem prispevku predstavljamo pregled metod, uporabljenih pri obnovi 
travišč iz različnih predelov Evrope, ki bi jih lahko ustrezno prilagojene uporabili tudi za obnovo slovenskih travišč. 
Travnike lahko prepustimo spontani sukcesiji, ki je primerna predvsem za manjše degradirane površine v bližini 
travišč z dobro ohranjeno biotsko raznovrstnostjo. Praviloma pa vnovična vzpostavitev rastlinskih združb z značilnimi 
vrstami zahteva bolj proaktiven pristop, kot je setev regionalnih ali komercialnih ohranjevalnih semenskih mešanic, 
prenos dozorelega rastlinskega materiala ali prenos vrhnje plasti tal z donorske površine z ustrezno vrstno sestavo. 
Metode obnove travinja je potrebno natančno pretehtati in z obnovo vrstne pestrosti začeti še preden je habitat ali 
določena vrsta ogrožena. Ugotavljamo, da ima lahko optimalen način upravljanja travinja po obnovi primerljiv ali celo 
večji učinek glede na tistega, ki ga lahko dosežemo z ustrezno metodo obnove. Kljub vsemu pa še vedno velja, da je 
vzdrževanje dobro ohranjenih travišč z ustreznim načinom rabe cenejše od obnove degradiranih travišč.
Ključne besede 
obnova travišč, upravljanje, metodologija, Natura 2000 travniški habitatni tipi, kmetijsko-okoljski ukrepi
Introduction
Grasslands are areas dominated by naturally occurring 
grasses and other herbaceous species, used mainly for 
grazing by livestock and wildlife (Allen et al. 2011). They 
comprise approximately 40.5% of the Earth’s surface 
(excluding Greenland and Antarctica), making them one 
of the world’s largest ecosystems (Suttie et al. 2005).
In general, grasslands can be divided into two types: 
primary (natural) and secondary (semi-natural). The first 
type is represented by sites that are unfavourable for 
the establishment of trees, while the second type is of 
anthropogenic origin, sustained by regular mowing and/
or livestock grazing. Despite their anthropogenic origin, 
secondary grasslands represent some of the most spe-
cies-rich and also some of the most vulnerable habitats 
in Europe since they have been extensively managed for 
several hundred years (Törok et al. 2020). In contrast to 
intensively managed systems with high productivity and 
thus high input requirements per unit area, extensively 
managed systems use small amounts of labour and capital 
per unit area and primarily rely on natural soil fertility, water 
availability, and climate (Ashton et al. 2012). Therefore, 
extensively managed grasslands have higher biodiversity 
value (Lesschen et al. 2014). The most species-rich sec-
ondary grasslands are dry and semi-dry grasslands. They 
are found on shallow to deep soils and are predominantly 
formed on calcareous or volcanic bedrocks from lowlands 
to mountains. Calcareous grassland types are especially 
species-rich. However, they are increasingly threatened by 
woody encroachment (Elias et al. 2018).
Grasslands may also be divided according to their age, 
namely into temporary and permanent grasslands. Accord-
ing to the European Commission, permanent grasslands 
are defined as “land used to grow grasses or other herba-
ceous forage naturally (self-seeded) or through cultivation 
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Acta Biologica Slovenica, 2023, 66 (1)
(sown), and that is not included in the crop rotation of the 
holding for five years or longer” (Commission Regulation 
(EC) No 796/2004). Grasslands that are less than five years 
of age (including in a crop rotation) are therefore defined as 
temporary grasslands (Reheul et al. 2007).
Natural and semi-natural grasslands are usually 
defined by climatic, pedological, or topological factors. 
However, in natural grasslands, plant species richness is 
further increased by fires and grazing, which also prevent 
litter accumulation and thus limit woody encroachment 
in addition to climate and extreme habitat conditions 
(Kuzemko et al. 2016, Török et al. 2020). Both natural and 
semi-natural grasslands depend on microclimatic, soil, and 
bedrock gradients (Sutcliffe et al. 2016). Among these, the 
main drivers of species composition are soil depth, soil 
texture, and soil pH. The relationship between pH and plant 
diversity of a certain region is generally hump-shaped, as 
the highest plant species richness is found under neutral 
or slightly basic conditions (Palpurina et al. 2017). A hump-
shaped relationship can also be found between plant 
species richness and primary productivity (Fraser et al. 
2015). The productivity of grassland communities is mostly 
driven by the availability of water and nutrients, which 
affects the biodiversity of these communities. The amount 
of moisture and its seasonal variation influences the pro-
ductivity of natural grasslands, whereas the productivity 
of semi-natural grasslands is controlled by the fertility 
gradient (Török et al. 2020). Limiting nutrients play an 
important role in shaping the richness-productivity pattern 
(Palpurina et al. 2017) since various species are adapted 
to nitrogen (N) and phosphorus (P) limitation (Roeling et 
al. 2018). Generally, grasslands rich in N, P, and potassium 
(K) have low biodiversity (Merunková and Chytrý 2012). 
Although fire is not as crucial for the formation of grass-
land plant communities as climate, its contribution to this 
process in terms of seasonality, intensity, and return rate is 
still significant (Ewing and Engle 1988, Biondini et al. 1989). 
Moreover, areas that have recently undergone fires are 
usually richer in nutrients, attracting large ungulates that 
may further change species composition (Milchunas et al. 
1988, Hartnett et al. 1996).
Natural and semi-natural extensively managed grass-
lands provide many ecosystem services that cannot be 
provided by other land uses. Their ecosystem services are 
of higher value compared to those of sown and intensively 
managed grasslands (Wick et al. 2016). For example, 
natural and semi-natural extensively managed grasslands 
harbour rich and unique flora and fauna, they produce 
biomass, serve as forage for herbivores, provide natural 
medicines, present habitat for pollinators and birds, 
ensure resources for water infiltration, flood reduction, 
purification, and storage, they prevent erosion, balance 
local climate, and play an important role in nutrient cycling 
and nutrient retention (Wick et al. 2016, Török et al. 2020). 
They also act as carbon dioxide (CO2) sinks and thus have 
a great carbon (C) sequestration potential (Ammann et al. 
2007) since they contain up to 12% of the soil organic C 
global pool (Schlesinger 1997). Converting grasslands into 
arable land leads to a decline in soil C due to C loss by 
tillage and lower C input from litter (Jones and Donnelly 
2004). Grasslands also offer a variety of intangible 
aesthetic, cultural, and recreational services (Török et 
al. 2017) and also have a good potential for biogas and 
biofuel production (Heinsoo et al. 2010).
Despite their ecological, cultural, and agricultural 
importance, grasslands are threatened globally due to 
numerous anthropogenic factors and climate change. 
Their degradation results in a reduced provision of eco-
system goods and services (Wick et al. 2016). The most 
common reasons for grassland degradation include, on the 
one hand, their fragmentation and isolation caused by the 
intensification of agriculture and increased development 
of the secondary and tertiary economic sector, especially 
in lowlands, along with the abandonment of extensive 
management by mowing or grazing in less accessible 
areas, resulting in overgrowth and encroachment by 
trees and shrubs. Other frequently mentioned causes 
include the absence of naturally occurring fires, biological 
invasions caused by introductions of non-native species, 
overgrazing, eutrophication, and the already mentioned 
climate change (Wick et al. 2016, Török et al. 2020).
In Slovenia, permanent grasslands cover 58% of the 
total agricultural land (Factsheet on the 2014-2022 Rural 
Development Programme for Slovenia 2022). About 20% 
of the Slovenian Natura 2000 network is represented 
by utilised agricultural land, among which extensive 
meadows are the most important (Prioritised action 
framework (PAF) for Natura 2000 in Slovenia 2019). The 
latest report by the European Environment Agency on 
the state of nature in the European Union from the year 
2020 claims that 50% of the assessments for grasslands 
in Slovenia under the Habitats Directive are showing a 
bad conservation status (U2) and about 22% are showing 
an inadequate (U1) conservation status. Therefore, alto-
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Acta Biologica Slovenica, 2023, 66 (1)
gether, no less than 72% of Slovenian grassland areas 
have an unfavourable conservation status (EEA 2020). 
Among grassland areas, semi-natural dry grasslands and 
scrubland facies on calcareous substrates (Natura 2000 
habitat type 6210(*)) are particularly affected. The main 
reason for such a negative trend is their dependency 
on particular sustainable management measures (EEA 
2020). Kaligarič et al. (2019) studied four grassland-spe-
cific agri-environment measures (AEM) in Slovenia in the 
period from 2007 to 2013 and found that these failed to 
preserve high nature value (HNV) grasslands in Slovenia 
since AEM did not reach most of the HNV grasslands in 
Slovenia and since this limited interest in the AEM targeted 
any permanent grassland, irrespective of its conserva-
tion value. As a result, many grasslands included in the 
AEM were not HNV grasslands. The authors concluded 
that this was due to the lack of pre-selection criteria for 
grasslands and the lack of monitoring of the efficiency of 
the measures, and due to low interest of farmers in the 
subsidies (Kaligarič et al. 2019). Ivajnšič et al. (2019) also 
reported negligible integration of agricultural holdings 
in the AEM in Slovenia. Unfortunately, the AEM in the 
period from 2014 to 2020 retained more or less the same 
concept as in the previous period (Kaligarič et al. 2019). 
Therefore, Kaligarič et al. (2019) suggested that future 
schemes should be prepared on a completely different 
basis, wherein HNV grasslands should be prioritised, and 
monitoring of biodiversity should be the most important 
requirement. The proposed Common Agricultural Policy 
(CAP) for the period from 2021 to 2027 was said to be 
more flexible and effective (Lovec et al. 2020). However, 
Lovec et al. (2020) argue that the actual long-term impact 
was not thoroughly considered in the new CAP. Therefore, 
no significant improvements in biodiversity conservation 
can be expected (Lovec et al. 2020). Šumrada et al. (2021) 
conducted a study where they explored the potential 
of a payment-by-results approach as an alternative to 
management-based schemes (MBS) in Slovenia. MBSs 
are schemes that provide payments for farming practices, 
which are believed to secure certain services instead of 
being tied to their actual provision (Burton and Schwarz 
2013). Alternatively, result-based schemes (RBS) remu-
nerate farmers for ecological results, demonstrated by 
certain indicators (e.g., presence of certain plant species, 
breeding success of farmland birds, etc.) (Herzon et 
al. 2018). Šumrada et al. (2021, 2022) found that most 
farmers and experts were in favour of the introduction of 
such RBSs for grassland conservation. Farmers knew the 
selected plant indicators and preferred monitoring of their 
presence over the current system (Šumrada et al. 2021). 
However, for the successful conservation of HNV grass-
lands, institutional capacity is also needed to implement 
RBSs on a larger scale (Šumrada et al. 2021). In addition, 
regardless of their many advantages in comparison to 
MBSs (Pe’er et al. 2022), RBSs do not seem to be better 
suited than the current schemes in addressing the specific 
needs of small and (semi-)subsistence farmers (Davidova 
2011) and in the cases where land ownership is highly frag-
mented (Hartvigsen 2014). For this reason, older farmers 
and those who manage semi-subsistent and small farms 
are mostly not in favour of RBSs (Šumrada et al. 2022). 
The findings of Šumrada et al. (2021) indicate that there is 
an institutional gap in the understanding of agroecology 
and of the importance of integration of biodiversity policy, 
which needs to be bridged to be able to enforce changes 
into the current system and achieve better conservation 
outcomes. Pe’er et al. (2022) argue that a combination of 
both result-based and action-oriented payments might be 
optimal.
According to the various ecosystem services that 
grasslands provide, it is vital to prevent their further deg-
radation in the future. In this review article, we present an 
overview of the techniques for grassland restoration used 
in experiments carried out in various European countries 
(but not in Slovenia), evaluate their practical use in the 
past, and form recommendations for their future applica-
tion. All findings and recommendations are drawn from 
these studies.
Grassland restoration  
in general
The best way to maintain the biodiversity of grasslands is 
to prevent both their abandonment and overexploitation 
by using extensive traditional agricultural practices, mainly 
grazing and/or mowing. These two practices have been 
shown to exert many positive effects on grassland biodi-
versity and are often recommended as tools for both the 
restoration and maintenance of grasslands (Galvánek and 
Lepš 2008, Török et al. 2016). Among these, wild herbivore 
grazing on open areas (by wild horses and cattle) and 
low-intensity grazing by herded livestock (local breeds) are 
often suggested (Török et al. 2016, Tóth et al. 2018).
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Acta Biologica Slovenica, 2023, 66 (1)
To retain grassland biodiversity in (near) optimum 
conditions, not only one single type but a whole scheme 
of traditional management practices is required (Babai 
and Molnár 2014). In addition, mosaic management 
should also be taken into account to ensure the highest 
biodiversity (i.e., temporally and spatially dynamic com-
bination of abandoned and mown grassland patches) 
(Török et al. 2017). Nonetheless, it is often the case that 
traditional practices are not possible or economically 
sustainable. For this reason, conservation authorities 
and site managers are looking for alternative practices, 
such as prescribed burning during the dormant season. 
Prescribed burning with long fire-return periods could be 
a suitable and economically feasible way of eliminating 
accumulated litter and sustaining grassland biodiversity 
(Valkó et al. 2014). However, in some cases, soil biota 
may be negatively affected by fires (McLeod and Gates 
1998, Vasconcelos et al. 2017, Zaitsev et al. 2017). Annual 
prescribed fires were found to be inappropriate for 
maintaining the desired species richness and structure, 
whereas periodic prescribed fires every two to six years 
may have a positive impact even in terms of soil biological 
properties (Valkó et al. 2014).
For partly degraded grasslands, a decrease in manage-
ment intensity is often sufficient for their recovery, while for 
fully degraded grasslands that have been converted into 
other land uses, spontaneous succession or technical rec-
lamation is required for a successful restoration (Prach and 
Hobbs 2008). Spontaneous succession has been reported 
to be a promising restoration tool in several different 
Central European grassland habitats, especially where the 
proportion of target grassland communities is high (Albert 
et al. 2014, Prach et al. 2015). However, for large-scale res-
toration projects to be successful, well-preserved donor 
grasslands acting as spontaneous sources of propagules 
must be available nearby (Török et al. 2020). Regarding 
technical reclamation, the most common methods that 
have been successfully used in some extensive grassland 
restoration projects (Lengyel et al. 2012, Prach et al. 2015) 
are sowing of regional seed mixtures and transfer of plant 
material (Török et al. 2011).
The main goal of many restoration actions is not to 
directly increase biodiversity on chosen plots per se, 
but rather to (re-)establish and maintain the presence of 
characteristic indicator species of target grassland com-
munities, with an additional aim of reducing cover and 
impact of non-target weeds (Lepš et al. 2007).
Restoration of species-rich grasslands can be 
time-consuming, and additionally, it usually requires plot 
preparation (Kiehl et al. 2010, Krautzer et al. 2011) and 
appropriate post-restoration management (Kiehl et al. 
2010, Török et al. 2011). Many sources claim that seeding 
on bare soil improves the chances of vegetation estab-
lishment (Kiehl et al. 2010, Krautzer et al. 2011, Török et 
al. 2011). Soil preparation is mostly done by ploughing or 
disking, followed by raking for seedbed preparation. After 
sowing, the covering of seeds is carried out by raking 
or ring rolling (Török et al. 2011). If an existing sward is 
present, it should be cut to the height of 3-5 cm, when 
necessary, and then opened (Krautzer et al. 2011). Grass-
land restoration should first be carefully thought-out and 
then carried out according to the plan shown in Fig. 1.
Figure 1. The stages of grassland restoration.
Slika 1. Zaporedje korakov pri obnovi travnikov.Collection of information
on previous and current state of chosen plots
Choice of methodology 
according to target vegetation and conditions at 
target and donor sites 
STAGE 1 
STAGE 2
Restoration action
according to chosen methodologySTAGE 3
Evaluation of restoration
according to target vegetationSTAGE 5
Post-restoration managementSTAGE 4
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Acta Biologica Slovenica, 2023, 66 (1)
Most commonly used methods 
in grassland restoration with 
practical examples
When restoring degraded permanent grasslands, we can 
choose between different methods. Each of them has 
its advantages and disadvantages, which are described 
below and summarised in Tab. 1.
Table 1. A summary of grassland restoration techniques used in different European countries.
Tabela 1. Povzetek tehnik obnove travnikov, ki so bile uporabljene v različnih evropskih državah.
Grassland restoration technique Suitable for Advantages Limitations Reference /  
An example of use
A decrease in management intensity degraded grasslands 
of various habitat 
types (mesophilic, 
calcareous, and  
acidic grasslands)
reduction and 
prevention of further 
nutrient accumulation 
in the soil; general 
improvement of soil 
properties; an increase 
of taxonomical and 
functional biodiversity; 
increased landscape 
heterogeneity
drastic decrease 
in management 
intensity can lead to 
abandonment and 
biodiversity loss, 
causing overgrowth 
of weeds, forbs, and 
shrub encroachment; 
positive impacts of 
fertilisation reduction 
are not noticeable 
immediately; might 
negatively impact 
grass coverage in 
mesic grasslands  
and pastures
Marriott et al. 2004, 
Milberg et al. 2017, 
Mayel et al. 2021, 
Resch et al. 2021
Spontaneous succession ruined grasslands; 
small-scale areas 
located in the proximity 
of well-preserved 
grasslands (mesic 
Arrhenatherion 
grasslands, dry 
Festuco-Brometea 
grasslands)
natural process;  
no action is needed
requires a high 
proportion of target 
grassland communities; 
high probability for an 
increased presence 
of weeds; long-term 
method; most suitable 
for recently degraded 
grasslands
Bossuyt and Honay 
2008, Fagan et al. 
2008, Lencová and 
Prach 2011, Knappová 
et al. 2012, Albert et al. 
2014, Prach et al. 2014, 
Sojneková and Chytrý 
2015, Prach et al. 2021a
Sowing of 
regional 
seed 
mixtures
low-
diversity 
seed 
mixtures
manual 
collection
degraded grasslands; 
ruined grasslands; 
plots that are at risk of 
erosion; larger plots 
(mesic Arrhenatherion 
grasslands, dry 
Festuco-Brometea 
grasslands)
quick results requires  
well-preserved donor 
grasslands; manual 
collection is time-
consuming, requires 
good seed recognition; 
proper harvesting time
Lepš et al. 2007, Kiehl 
et al. 2010, Hofmann et 
al. 2020
vacuum 
harvesting
combine 
harvesting
high-
diversity 
seed 
mixtures
manual 
collection
ruined grasslands; 
smaller plots (mesic 
Arrhenatherion 
grasslands, dry 
Festuco-Brometea 
grasslands)
greater restoration 
success; a useful 
first step towards 
revegetation
requires  
well-preserved donor 
grasslands; limited 
availability; larger 
collecting effort; 
manual collection 
is time-consuming, 
requires good seed 
recognition; proper 
harvesting time
Jongepierová et al. 
2007, Lepš et al. 2007, 
Kiehl et al. 2010, Török 
et al. 2011, Mitchley et 
al. 2012, Haslgrübler 
et al. 2013, Prach et al. 
2013, 2014, Baasch et 
al. 2016, Hofmann et 
al. 2020, Prach et al. 
2021a
vacuum 
harvesting
combine 
harvesting
Decrease in management intensity
Grassland management (especially traditional) is a crucial 
tool in the maintenance of semi-natural grasslands 
(Bischoff et al. 2009, Milberg et al. 2017, Goret et al. 2021). 
Cessation of management intensity (extensification) on 
previously intensively used grasslands has recently been 
discussed as a useful grassland restoration tool in many 
conservation policies and attempts. It might, theoretically, 
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Acta Biologica Slovenica, 2023, 66 (1)
Grassland restoration technique Suitable for Advantages Limitations Reference /  
An example of use
Transfer 
of plant 
material
hay 
threshing
plant 
clipping or 
green hay
ruined grasslands;  
all types of habitats
greater restoration 
success; genetic 
diversity preserved; 
low-cost; applicable  
on a larger scale
requires  
well-preserved  
donor grasslands; 
lower seed content
Kiehl et al. 2006, 
Donath et al. 2007, 
Edwards et al. 2007, 
Galvánek and Janák 
2008, Kiehl et al. 2010, 
Rydgren et al. 2010, 
Krautzer et al. 2011, 
Albert et al. 2019, 
Hofmann et al. 2020
dry hay/ 
‘hay sensu 
stricto’
seed stripping does not require 
vegetation cutting
requires  
well-preserved  
donor grasslands
brush harvester  
(most common)
ruined grasslands;  
tall meadows  
(Molinion, Cnidion)
successful and widely 
used restoration 
method; does not 
require vegetation 
cutting; greater 
restoration success
requires well-
preserved donor 
grasslands; seeds 
of shorter species 
underrepresented; 
specific equipment
Topsoil transfer ruined grasslands greater restoration 
success
destructive for donor 
site; laborious; costly
Kiehl and Pfidenhauer 
2007, Török et al. 2011
Mechanical disturbance, applied  
prior to restoration techniques
variously managed 
temperate grasslands; 
severely degraded 
grasslands; wet 
meadows; Nardus 
grasslands
increased soil seed 
bank potential and 
seedling recruitment
low specificity 
and selectivity; 
might improve the 
development of weeds
Hofmann and Isselstein 
2004, Donath et al. 
2007, Edwards et al. 
2007, Mitchell et al. 
2009, Krautzer et al. 
2011, Klaus et al. 2018
Planting of entire plant individuals degraded grasslands; 
individual/specific 
species; for stimulating 
succession to later 
stages
better establishment  
of endangered  
species
laborious; costly Török et al. 2011
Cessation of fertilisation grasslands degraded 
by over-fertilisation 
(mesic semi-natural 
grasslands containing 
Molinio-Arrhenatherea 
and Festuco-Brometea 
species; Nardus 
grasslands)
low-cost; no active 
measurements  
needed
unpredictable species 
shift; depends on 
abiotic parameters 
(climate); highly 
dependent on initial 
state; time-consuming
Marriott et al. 2004, 
Hejcman et al. 2007, 
Královec et al. 2009, 
Korzeniak 2016, Van 
Daele et al. 2017, Prach 
et al. 2021b
increase biodiversity on chosen plots by reducing work 
intensity, but at the same time increasing work effective-
ness on larger grassland areas (Marriott et al. 2004, Milberg 
et al. 2017). However, results are context-dependent and 
differ according to the methodology used and the initial 
state at chosen sites. A study on the impact of grassland 
extensification by Marriott et al. (2004) showed that man-
agement cessation had a moderately negative impact on 
the diversity of calcareous and mesic grasslands, wherein 
changes were primarily visible in the long term. On the 
other hand, management abandonment had a significant 
negative impact primarily on mesic grasslands, which 
were showing signs of succession to forest vegetation 
in a 13-year period and were overgrown by shrubs and 
forest species in 30 years. Acidic grasslands were over-
grown by non-palatable grasses (such as Molinia caerulea 
and Deschampsia flexuosa) and ericoid shrubs, whereas 
calcareous grasslands were overgrown by Brachypodium 
pinnatum (Marriott et al. 2004). Milberg et al. (2017) found 
that cessation of mowing intensity (from annually to once 
every three years) had a weak impact on diversity of 
wet grasslands, whereas the diversity of dry grasslands 
decreased significantly. Significant biodiversity loss was 
noticeable one decade into the experiment, more spe-
cifically after 11-14 years, which is somehow similar to the 
results of the study by Marriott et al. (2004). Rare mowing 
has a particularly negative impact on short-growing 
species, which can be impaired by higher and/or woody 
species that out-compete them and benefit from sporadic 
management. A combination of annual mowing of grasses 
and forbs in swards and occasional (once every few years) 
mowing of shrubby, overgrown parts of vegetation might 
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be a solution that would combine the benefits of the afore-
mentioned policy of reduced, but more efficient work, and 
conservation of grassland habitats (Milberg et al. 2017). 
The benefits of grassland management are discussed in 
more detail in the chapter Post-restoration management 
with grazing and mowing.
Spontaneous succession
Spontaneous secondary succession is a natural process 
of changes in species’ structure and interaction dynamics 
in an already established ecosystem following natural or 
anthropogenic disturbance events. Particularly in the case 
of grassland restoration actions, secondary succession 
often occurs following the abandonment of previously 
used arable land. It is the most natural way of grassland 
restoration, relying on naturally (spontaneously) occurring 
processes in an ecosystem (Török et al. 2011). It is strongly 
dependent on the local availability of seeds (propagules) 
of the target grassland species and their efficient disper-
sal by different vectors, deriving from natural vegetation 
near the restoration target sites (Hölzel and Otte 2003, 
Donath et al. 2007, Kardol et al. 2008, Kiehl et al. 2010, 
Krautzer et al. 2011, Török et al. 2011, Hofmann et al. 2020). 
Natural succession of ex-arable fields usually begins with 
the establishment of annual and perennial ruderal plants, 
followed by perennial grasses and forbs – species of 
later successional stages, which are better adapted to 
specific grassland conditions (such as resource limitation). 
However, the transition to more complex plant commu-
nities during later stages of ecological succession rarely 
occurs. The structure of these plant communities stag-
nates with an increased presence of weeds, ruderals, and 
grasses, due to the low germination potential of grass-
land species’ seeds in the seed banks and their limited 
dispersal potential (Kleijn 2003, Lawson et al. 2004). As 
stable ecosystems, grasslands do not typically rely on 
soil seed banks since characteristic grassland species do 
not produce long-lived or numerous seeds (Bossuyt and 
Honnay 2008). They rather rely on the dispersal of seeds 
(Edwards et al. 2007, Bossuyt and Honnay 2008). A study 
by Knappová et al. (2012) reported on the ability of dry 
grassland species (Festuco-Brometea) to colonise grass-
lands in their proximity as an alternative to restoration 
actions of a particular grassland type. While about 68% 
of the species present showed dispersal potential, some 
of the ‘grassland specialists’, such as Helianthemum num-
mularium subsp. grandiflorum, Carex humilis, Anthericum 
ramosum, Filipendula vulgaris, Melampyrum nemorosum, 
Gymnadenia conopsea, and Campanula glomerata, did 
not colonise proximal area. Overall, the authors defined 
the distance of 0.5 kilometres as the limit for successful 
propagule spreading, pointing out that in addition to 
distance and plot area, biotic and abiotic factors of the 
selected grasslands also need to be considered when 
discussing this topic (Knappová et al. 2012).
A review study by Bossuyt and Honnay (2008) showed 
that grassland soil seed banks were amongst the ones 
with the highest diversity, richness, and evenness, in 
comparison to those of forests, marshes, and heathlands. 
However, their generally low seed density and common 
absence of target species’ seeds (especially in calcareous 
grasslands) made them an unreliable source for resto-
ration. Exceptionally, seed banks might be useful at sites 
that were recently degraded (up to 5 years ago) (Bossuyt 
and Honnay 2008). Intensive use of agricultural fields and 
increased soil fertility negatively impact grassland soil 
seed banks (Edwards et al. 2007), favouring the devel-
opment of weeds as remnants of previous successional 
stages (Török et al. 2011), along with invasive and non-tar-
get species that are commonly found at earlier stages of 
succession (Bossuyt and Honnay 2008). Another common 
issue with spontaneous succession is landscape fragmen-
tation, resulting in the isolation of grassland plots and the 
absence of suitable propagule sources (Kleijn 2003, Kiehl 
et al. 2010, Lawson et al. 2004, Török et al. 2011), along 
with their dispersal limitation (Donath et al. 2007). All 
these issues generally cause delayed restoration (Kleijn 
2003, Török et al. 2011). In addition, natural regeneration is 
often limited by inadequate germination conditions found 
on plots where vegetation is already established (mainly 
regarding light availability and soil). These differ between 
species of early and later successional stages. Species of 
early successional stages are usually taller and are con-
sequently better competitors for light. Furthermore, they 
have a better capability for vegetative reproduction and 
thus out-compete species of later successional stages 
(Kleijn 2003).
Proactive measures of grassland restoration are 
usually needed since restoration by spontaneous succes-
sion is mainly suitable for small-scale areas located in the 
proximity of well-preserved grasslands, which serve as an 
adequate source of propagules. It is a good restoration 
method where no urgent results are expected due to 
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slow and often unpredictable development of vegetation 
(Török et al. 2011). For example, Fagan et al. (2008) showed 
that they preferred the regeneration of dry grasslands by 
natural succession, combined with moderate manage-
ment, defined by local agro-environmental schemes, 
rather than regeneration by sowing seed mixtures. 
Another study favouring natural succession over sowing 
of seed mixtures as a form of dry grassland restoration 
was conducted by Lencová and Prach (2011). They claim 
that if target sites are in the proximity of adequate prop-
agule sources, properly managed, and not (extremely) 
loaded with nutrients, no sowing is required unless urgent 
results for grassland production are needed.
Sowing of seed mixtures
Many grassland restoration attempts rely on different 
methods of plant material introduction (Hofmann et al. 
2020). One of the most commonly used methods is seed 
addition by sowing seed mixtures (Török et al. 2011, 
Hofmann et al. 2020). Seed mixture composition depends 
on numerous factors, such as target vegetation, condi-
tions at the target site, and seed availability (Török et al. 
2011).
Török et al. (2011) mention two main types of seed 
mixtures used in restoration: (1) low-diversity (LD) seed 
mixtures, usually consisting of seeds of 2-8 species, 
mostly dominant grasses and/or forbs of the target veg-
etation, and (2) high-diversity (HD) seed mixtures, which 
contain seeds of 10 or more target species. LD seed 
mixtures are often applied in restoration experiments that 
aim for quicker results – for instance, on plots that are 
at high risk of erosion. On larger plots, where HD seed 
mixtures might not be quantitatively sufficient due to their 
limited availability and greater collection effort, mixtures 
may be combined and/or applied in patches. Seeds can 
be either commercially sourced or locally harvested. 
Commercial seed mixtures are acceptable if they contain 
seeds of target species from local populations. It is, 
however, highly suggested to use seeds of plants grown 
or harvested locally, as this way the chances of successful 
restoration are higher. Non-native ecotypes in commercial 
seed mixtures namely have lower genetic and ecological 
compatibility with local conditions (Kiehl et al. 2010, Török 
et al. 2011). Additionally, foreign ecotypes may hybridise 
with local ecotypes, causing genetic biodiversity loss and 
reduced fitness in new hybrid populations (Kiehl et al. 
2010). Seed collection, as opposed to the application of 
commercially sourced propagules, may help in the rein-
troduction of certain rare species, whose seeds are not 
readily available (Kiehl et al. 2010). Seeds can be collected 
manually or by using specific equipment, such as vacuum 
or combine harvesters (Edwards et al. 2007), which require 
special attention in the choice of proper harvesting time 
(Krautzer et al. 2011). An exception is vacuum harvesting, 
which can be easily carried out with machines used for 
leaf vacuuming and blowing even after seed shedding. 
Due to its complexity, this method should be carried out 
primarily at locations where other methods are not readily 
applicable (Kiehl et al. 2010). Application of different seed 
densities results in varying success rates of grassland 
restoration. Generally, the application of a larger number 
of seeds results in greater restoration success. However, 
it could also cause greater resource competition among 
sown species, which might result in decreased diversity 
of target species (Lepš et al. 2007, Török et al. 2011). As 
a potentially good alternative to spontaneous succession, 
Török et al. (2011) recommend sowing densities from 4000 
to 13000 seeds/m2 on areas of up to a few hundred square 
meters, whereas larger areas (measured in hectares) might 
require between 20 and 45 kg of seeds per hectare (for 
more details, see Török et al. 2011). Sowing of regional 
seed mixtures has been repeatedly proven as a successful 
restoration method for dry grasslands (Lepš et al. 2007, 
Jongepierová et al. 2007, Mitchley et al. 2012, Joha-
nidesová et al. 2014, Prach et al. 2021a). Jongepierová et 
al. (2007) reported that sowing of regional seed mixtures 
is a very good method for the restoration of Bromion 
grasslands, indicating the use of a specific methodology 
in the form of strip sowing. Regional seed mixtures may be 
expensive and not readily accessible. In this case, strips 
of regional seed mixtures could be applied inside fields 
designated for natural regeneration, which should theo-
retically be colonised by previously sown target species. 
Mitchley et al. (2012) studied vegetation at the same plots 
in 2009, following the first part of the survey by Jonge-
pierová et al. (2007), further confirming the positive effects 
of sowing regional seed mixtures, with the highest success 
achieved by sowing regional grasses, which covered 
around 50% of the studied plots. All experimental plots 
showed divergence towards ancient grasslands in the 
10-year period, but donor and recipient sites still showed 
significant differences in species composition, suggesting 
that the 10-year period might not be sufficient to achieve 
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the intended results and prove the importance of time 
dimension in grassland restoration experiments. Regard-
ing different types of seed mixtures, Lepš et al. (2007) 
confirmed the positive impact of sowing both HD and LD 
seed mixtures on grassland vegetation succession. Both 
LD and HD seed mixtures were proven as suitable, even 
though HD seed mixtures were generally more successful 
and had higher productivity. An additional benefit that 
they provided was a higher probability of compensation 
in the case of species establishment failure, expressing 
the ‘insurance effect’ on diversity. Grasses were generally 
more successful than forbs. However, non-sown control 
plots subjected to natural succession had higher species 
richness and diversity, which did not result from dispersal 
from sown plots. Namely, sown species never achieved 
dominance on control plots, even though they spread rela-
tively successfully over short distances into adjacent plots. 
Sowing of species is therefore defined as better suited for 
species introduction and not necessarily for biodiversity 
increase. An ideal combination might be planned to sow 
seed mixtures in strips, with some space left for natural 
colonisation between the strips. When seed mixtures are 
used in restoration actions, the species composition of 
both applied mixtures and recipient sites should be taken 
into consideration, focusing on the colonisation potential 
and competitiveness of present species. The presence of 
highly competitive species in mixtures might negatively 
impact the development and survival of other species, 
which were either previously sown or initially present at 
the site. Due to the competitiveness or invasiveness of 
various plant species (particularly weeds), the presence of 
those species in plant communities that are located near 
the restored grasslands should also be taken into account. 
By doing that, we might prevent their spreading into newly 
sown sites, where they might cause negative shifts in target 
species composition. Johanidesová et al. (2014) addition-
ally reported that restoration by seed addition could be 
a successful first step towards revegetation and that its 
continuation in the form of natural succession depends 
on the vicinity of appropriate grasslands that would serve 
as a natural source of propagules. The latter is important 
due to limitations in the sowing of some grassland species’ 
seeds for many reasons, either biological or technical. If 
natural spreading does not fulfil expectations in the set 
time frame, repeated sowing of selected target species 
may be required. Prach et al. (2014) studied the differences 
between restoration attempts of two grassland types of 
the Arrhenatherion and Bromion alliances in the White 
Carpathian Mountains, pointing out that spontaneous 
succession and use of commercially prepared mixtures led 
to the establishment of mesic vegetation, whereas local 
mixtures favoured dry grasslands. A study by Prach et al. 
(2013) on the same area gave the same results, favouring 
the use of local seed mixtures. 98% of sown species grew 
successfully, along with unsown species that spread from 
nearby reference ancient grasslands.
Transfer of plant material
Other commonly used seed collection techniques include 
threshing, seed stripping, and the use of a brush harvester. 
Brush harvesting is a generally successful and widely 
used restoration method that does not require vegetation 
cutting. It is overall better suited for tall meadows since 
seeds of shorter species are often underrepresented in 
the harvested material. Shortgrass meadows could there-
fore benefit from collecting plant material by cutting or 
even raking, followed by transfer of raked material, which 
would additionally help transfer propagules of bryophytes 
and lichens. Hay threshing and brush harvesting are more 
expensive methods due to the use of specific equipment 
(Kiehl et al. 2010).
Restoration of species-rich grasslands usually requires 
transplantation of plant material from donor sites since the 
collection of adequate HD seed mixtures demands a lot of 
time and resources (Török et al. 2011). The application of 
plant material containing seeds is a cost-effective method 
that is applicable on a large scale and in a wide range 
of habitats. In addition, this method might be beneficial 
due to the introduction of an entire gene pool of the 
donor community, which might also include some rare or 
endangered species that cannot be sown. Therefore, the 
genetic diversity of locally-adapted ecotypes is preserved, 
along with the provision of microsites required for seed 
germination. Like other restoration methods, it is highly 
context-dependent, and it requires the proper selection 
of quality donor plant material applied at corresponding, 
ecologically compatible sites. Although the seed content 
of the transferred plant material is an important variable in 
the evaluation of restoration success, colonisation ability 
is also very important (Hölzel and Otte 2003).
When plant material with propagules is collected by 
mowing (cutting), it can be applied either as fresh or dry 
hay (Kiehl et al. 2010, Krautzer et al. 2011). Application 
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of fresh hay is known as ‘plant clippings’ or ‘green hay’, 
whereas dry hay is known as ‘hay sensu stricto’ (Kiehl et 
al. 2010). Application of fresh plant material increases the 
chances of species establishment (Kiehl et al. 2010, Török 
et al. 2011), while dry hay requires additional manipulation 
of the material and therefore increases the costs of the 
process (Krautzer et al. 2011), with a limited restoration 
potential due to lower seed content. Low-productivity 
grasslands, such as calcareous grasslands, typically 
require around 300-600 g/m2 of freshly cut hay with a 2:1 
to 3:1 donor-to-recipient plot ratio, whereas other habitat 
types, such as fens, require a 1:1 to 2:1 plot ratio (Kiehl et al. 
2010). Immediate transfer of freshly cut plant material with 
ripe seeds is known as hay strewing (Edwards et al. 2007). 
Plant material can be applied in a 5-15 cm layer, which is 
advised for mesotrophic to eutrophic grasslands (Kiehl et 
al. 2010). Thicker layers are not recommended since they 
can suppress the colonisation of target species (Kiehl 
et al. 2010, Török et al. 2011). Transfer of plant material 
containing seeds is best combined with prior preparation 
of recipient sites with various soil disturbance methods, 
which are chosen according to the grassland type and its 
nutrient status. The importance of mechanical disturbance 
for grassland restoration based on seed bank potential 
was studied by Klaus et al. (2018). Results showed that 
mechanical disturbance could potentially increase the 
diversity of severely degraded grasslands. The sole impact 
of seed banks on partly developed grasslands is, however, 
questionable. The similarity between seed banks and veg-
etation stands did not increase with disturbance frequency. 
Thus, other measures of propagule introduction, such as 
sowing, were required (Klaus et al. 2018). Apart from the 
quality of applied seed mixtures, another crucial factor is 
the availability of microsites that would provide adequate 
conditions for seedling germination and development, 
preventing seed desiccation. This can be achieved by 
providing an additional mulch layer that also prevents soil 
erosion (Kiehl et al. 2010).
Diaspore transfer was proved as a successful method 
in the restoration of wet meadows of the Molinion and 
Cnidion alliances (Hölzel and Otte 2003, Donath et al. 
2007). Previous disturbance of existing grass swards 
in the form of rotavating before the application of 
seed-containing plant material was proved beneficial for 
the establishment of target vegetation of wet grasslands, 
although the authors noted that competitive relationships 
within the stand (with a specific focus on grasses) severely 
dictated the dynamics of restoration. They also noted 
that heavier disturbance (such as ploughing of existing 
sward) might be needed to expand niches for introduced 
species (Donath et al. 2007). Both studies reported low 
establishment rates for sedges (Carex spp.) (Hölzel and 
Otte 2003, Donath et al. 2007). This can be explained 
by their specific phenology and ecology. Namely, their 
seeds ripen early in the vegetation season and are absent 
at the time of seed collection in late summer or autumn. 
They also have specific germination requirements, such 
as prolonged incubation in warm and moist conditions, 
and due to their successful vegetative reproduction, they 
are better suited for other methods of propagule intro-
duction, such as turf transfer (Donath et al. 2007). Donath 
et al. (2007) do not recommend simultaneous sowing of 
grasses when applying seed mixtures due to the potential 
of grasses to out-compete target vegetation and their 
ability to successfully colonise sites of stands in the prox-
imity, even though simultaneously sown grasses do not 
disturb the development of newly introduced vegetation 
(Donath et al. 2006, 2007). Already established grass 
stands presented a bigger obstacle to the recruitment of 
new vegetation (Donath et al. 2007).
Green hay transfer was proved by Kiehl et al. (2006) 
to be a successful method of restoration of ancient 
grasslands, although it is also not suitable for the reintro-
duction of sedges (Carex spp.). Albert et al. (2019) found 
that green hay transfer was the most efficient restoration 
method for Bromion erecti grasslands, followed by triple 
brush harvesting and single brush harvesting. Donor 
sites were located in the Protected Landscape Area and 
Biosphere Reserve of the White Carpathian Mountains in 
the Czech Republic. Thirty-five species were recorded at 
the recipient site, on which green hay consisting of 112 
species collected at donor plots was applied, whereas 
triple harvesting showed a similar success rate with 33 
newly established species. The success of this method 
may be due to the fact that plants were transferred as 
specimens, providing a possibility for the seeds to ripen 
on parent plants, whereas harvesting methods require 
effort to remove fruits and seeds from plants. Harvesting 
proved to be more successful when repeated during the 
season. However, all of the tested options had a relatively 
low success rate, with minimal differences between green 
hay and triple harvesting. Albert et al. (2019) noted that, 
due to the higher practicality of seed mixtures collected 
by harvesting, which can be stored and transferred, brush 
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harvesting has a significant potential for restoration, 
whereas hay transfer needs to be applied as soon as 
possible after mowing and is usually limited to smaller 
areas. Hofmann et al. (2020) tested different methods of 
grassland restoration on various vegetation types (Fes-
tuco-Brometea, Nardo-Callunetea, Sedo-Scleranthetea, 
and Trifolio-Geranietea). Their results confirmed the 
efficiency of hay transfer. However, they noted the impor-
tance of choosing the right time for the application of this 
method, which should be before the seeds are ripe, and 
should be repeated if necessary.
Edwards et al. (2007) studied the influence of prepa-
ratory disturbance practices (power harrowing and turf 
stripping) on the efficiency of two commonly used grass-
land restoration methods (brush harvesting and green 
hay strewing) applied to lowland hay meadows and chalk 
grasslands as two target vegetation types. Both methods 
of seed addition proved successful. Hay strewing was 
more efficient for seed collection of low-growing plant 
species due to the greater height at which the brush 
harvester collected seeds. In addition, it sampled a wider 
phenological range of different plants, a trait specific 
to mesic chalk grasslands. Early cutting might favour 
perennial grasses, whereas later hay cut includes a wider 
range of different forbs, of which many have a conserva-
tion value. This confirms the importance of phenology 
knowledge in choosing the adequate restoration method. 
Disturbance generally had a positive impact on grassland 
establishment, except for power harrowing in chalk 
grasslands. They did, however, benefit from turf stripping, 
even though the positive impact might have been related 
to a decrease in nutrients in the soil caused by the given 
disturbance. This study confirmed the importance of back-
ground knowledge and context dependence. The choice 
should be in line with phenology, and management tech-
niques should be considered.
Other methods of grassland restoration
Other, not commonly used methods of grassland resto-
ration include topsoil transfer, turf transplantation, and 
community translocation directly from donor sites to 
recipient plots (Török et al. 2011). Potential advantages 
of chosen methods include the transfer of diaspores with 
their associated soil fauna and microbiota, which could 
increase the chances of vegetation establishment in its 
original form and dynamics (as close as possible) (Török 
et al. 2011). Kiehl and Pfidenhauer (2007) confirmed a 
positive impact of topsoil removal on the re-establish-
ment of dry Bromus grasslands by an increased cover of 
various target species, such as Thymus praecox, Hippo-
crepis comosa, and Dorycnium germanicum, following 
the experiment. However, transplantation methods are 
still not widely used due to their destructive nature, the 
high effort that they require, and the costs they cause 
(Török et al. 2011). This was also noted by the authors 
of the original research (Kiehl and Pfidenhauer 2007). 
In addition to sowing or hay transfer, planting of entire 
plant individuals or their belowground parts is sometimes 
carried out in restored areas. Due to higher costs, it is 
only recommended in specific cases, for example, for a 
better establishment of endangered species or for stimu-
lating succession to later stages (Török et al. 2011). Other 
potential issues in soil transfer are eutrophication, caused 
by increased nutrient mineralisation in the applied soil, 
and ruderalisation, which require additional management 
measures afterwards (Kiehl et al. 2010).
As an option for the restoration of grasslands dom-
inated by N. stricta, Mitchell et al. (2009) suggested 
the creation of gaps in turf for other plant seedlings by 
rotavation. Nonetheless, regular cutting and chopping of 
biomass using a flail or rotary mower is not advised since 
it may cause changes in species composition and dom-
inance of some grasses (Krahulec et al. 2007, Galvánek 
and Janák 2008). Too regular burning is also not rec-
ommended, as it can promote the spreading of invasive 
species (Bensettiti et al. 2005).
For long-term maintenance of Nardus grasslands 
in Serbia, Dajić Stevanović et al. (2008) recommend 
practices such as juniper burning or roller chopping, or 
the introduction of horse grazing to control the spreading 
of some undesired low-quality grasses, and mechanical 
clearance of woody species, along with the reintroduction 
of cattle and sheep grazing.
Parolo et al. (2011) defined a few management practices 
necessary for the preservation of Nardus grasslands. They 
suggested mechanical removal of woody pioneer species 
to prevent encroachment at pasture edges, establishing 
electric paddocks for more intensive grazing at the periph-
ery of pastures, and using electric fences on lower pastures 
that are grazed twice to ensure the concentration of cows 
at pasture edges. It is advisable to distribute animals 
along the peripheral parts of pastures so that nitrophilous 
species have less potential for spreading (Bensettiti et al. 
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2005). Moreover, Parolo et al. (2011) recommended turf 
stripping in the centre and fertilisation at the periphery 
of pastures, and combining cattle with small herbivores, 
such as sheep and goats, for a more efficient limitation 
of woody encroachment following restoration. However, 
fertilisation in these habitats is usually forbidden or at least 
very restricted, as it can lead to eutrophication (Galvánek 
and Janák 2008). Some countries encourage liming for the 
improvement of Nardus grasslands, as higher calcium (Ca) 
levels may positively affect species diversity (Common et 
al. 1991). Nevertheless, a thorough assessment is required 
before application since liming, like P, has a long-term 
effect on species composition (Hejcman et al. 2007). Turf 
stripping is otherwise mainly used in the case of eutrophi-
cation of the upper soil horizons. This technique removes 
nutrients from the upper soil layers, and thus restoration of 
such oligotrophic habitats is possible (Galvánek and Janák 
2008).
Sometimes it is necessary to apply several restoration 
techniques at once. However, this correspondingly also 
requires more effort. Since grasslands that are to be 
restored are often abandoned and, therefore, densely 
overgrown, a technique that is often used is the removal 
of trees and shrubs by hand or with machinery. If a 
habitat is not densely overgrown, cutting and chopping of 
biomass with a flail or rotary mower may be used. When, 
however, trees and shrubs are too lush, a cultivator has 
to be used. Another possible, but costly method is also 
manual cutting with a brush cutter (Galvánek and Janák 
2008). All these methods for scrub cutting are effective, 
but only if regular management is assured afterwards 
(Bensettiti et al. 2005), as scrub encroachment could turn 
out to be even more intense after cutting (Galvánek and 
Janák 2008).
Restoration of completely destroyed habitats is very 
expensive, yet feasible. There were some attempts of 
restoration of severely damaged habitats in Belgium by 
turf transplantation using sod-cutting techniques and 
by utilisation of hay or mulch from species-rich donor 
grasslands (Galvánek and Janák 2008). The turf should 
be placed on open land in a chessboard layout. Such an 
arrangement prompts seed dispersal and recruitment, and 
thus enables faster rehabilitation of disturbed habitats. 
In addition, turf also prevents soil erosion (Stanová et 
al. 2007). If farmers are not interested in maintaining 
these habitats, or when these habitats are threatened by 
various economic activities, land acquisition is of great 
importance in order to ensure their proper management. 
In any case, no matter which restoration measure is used, 
regular active management of these habitats is still much 
more cost-effective than their restoration (Galvánek and 
Janák 2008).
Timeline of grassland 
restoration
When designing restoration studies, it is important to take 
into account the conditions and area of both donor and 
recipient grassland plots, along with the phenology of the 
community and target grassland species (Edwards et al. 
2007, Török et al. 2011). Seed collection should take place 
when most of the seeds in the donor plant community 
are ripe (Edwards et al. 2007, Török et al. 2011). The most 
appropriate times for seed collection in different habitat 
types are shown in Tab. 2. Nevertheless, it should be 
noted that these data were collected in different parts of 
Europe and, therefore, cannot be directly applied to the 
conditions in Slovenia.
For example, to maximise the yield of grass seeds, 
mowing of European dry grasslands should be carried out 
in June, whereas mesic grasslands should be mown later 
during the vegetation season (between June and July) 
(Török et al. 2011). To promote better biomass decomposi-
tion at the recipient sites in Nardus grasslands, propagule 
collection for restoration purposes in these grasslands 
should be carried out as early as May (Van Daele et al. 
2017), in the period from June to July (Rūsiņa et al. 2017), 
in mid-July (Galvánek and Janák 2008, Kurtogullari et 
al. 2020), or by the end of July at the latest (Háková et 
al. 2004). Actions taken later in the vegetation season 
are generally more suitable for Nardus grasslands and 
pastures at higher altitudes (Kurtogullari et al. 2020). Wet 
grasslands are usually mown in late summer, only in August 
(Török et al. 2011). Generally, earlier cuts of grasslands 
favour grasses, whilst later or repeated cuts favour forbs 
(Krautzer et al. 2011, Haslgrübler et al. 2013). Late propagule 
collection can severely decrease the chances of successful 
grassland restoration. To avoid this, it is recommended to 
carry out multiple collections throughout the vegetation 
season (Török et al. 2011). The area ratio between recipient 
and donor site size varies between 1:2 and 1:10, and mostly 
depends on the state of vegetation at donor sites (Edwards 
et al. 2007, Török et al. 2011).
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Table 2. An approximate timeline of plant material collection (primarily by mowing), which is used for the restoration of different European 
grassland habitat types. Differently coloured cells in the following table indicate theoretically defined optimal timing for mowing at different 
grassland habitat types. Individual restoration actions should, however, consider climate conditions and the state of donor grassland vegetation 
at a given time. Following works by Háková et al. (2004), Galvánek and Janák (2008), Krautzer et al. (2011),Török et al. (2011), Haslgrübler et al. 
(2013), Rūsiņa et al. (2017), Van Daele et al. (2017), Kurtogullari et al. (2020).
Tabela 2. Časovni okvir nabiranja rastlinskega materiala (predvsem s košnjo) za obnovo različnih evropskih habitatnih tipov travišč. Različno 
obarvana polja v spodnji preglednici označujejo teoretično optimalen čas za košnjo in nabiranje rastlinskega materiala v različnih travniških 
habitatnih tipih. Kljub temu pa je pri akcijah obnavljanja travnikov vedno potrebno upoštevati klimatske razmere in stanje vegetacije območja 
izbranih donorskih površin. Prirejeno po Háková in sod. (2004), Galvánek in Janák (2008), Krautzer in sod. (2011), Török in sod. (2011), Haslgrübler 
in sod. (2013), Rūsiņa in sod. (2017), Van Daele in sod. (2017), Kurtogullari in sod. (2020).
Ja
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Ju
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Dry calcareous grasslands (habitat type 6210(*))
Mesic grasslands (habitat types 6510, 6520)
Oligotrophic Nardus grasslands (habitat type 6230) Dependent on area
Wet grasslands (habitat type 6410)
Choice of donor sites for 
grassland restoration
When choosing adequate donor sites for grassland res-
toration, one should opt for donor sites that fulfil specific 
criteria. Primarily, we should choose donor sites that are 
biogeographically and phytocoenologically suitable 
for recipient sites, with representative target plant com-
munity structure and low presence of neophytes. Some 
databases for grassland sites in Germany also cite the 
need for adequate management status and naturalness 
of the site (not sown with commercial seeds), without 
incoming/planned changes in land use (Krautzer et al. 
2011). Additionally, donor sites must be compatible with 
recipient plots regarding their nutrient status, hydrology, 
and substrate, with special attention paid to differences in 
water and nutrient status of dry, nutrient-poor grasslands 
(Bromion), mesic and mesotrophic grasslands (Arrhen-
atherion), and wet grasslands (Molinion and Deschamp-
sion) (Krautzer et al. 2011).
Soil fertility as a crucial factor  
in grassland restoration
On land formerly used for cropland, the establishment 
of target grassland species can be limited by increased 
nutrient levels in topsoil, which stimulate the growth 
of highly competitive annuals and weeds (Kardol et al. 
2008). The removal of the topsoil layer might be applied 
to lower nutrient concentrations (Hölzel and Otte 2003, 
Kardol et al. 2008, Kiehl et al. 2010, Török et al. 2011) and 
to remove propagules of weeds (Hölzel and Otte 2003, 
Kiehl et al. 2010, Török et al. 2011). To prevent soil erosion, 
the removal of 25-50 cm of soil is recommended, whereas 
soil removal on a larger scale is not advised. Soils of 
ex-arable fields are often saturated with inorganic N, 
which stimulates the development of non-target species 
in the initial phases of vegetation succession. To prevent 
overgrowth by non-target species (namely weeds), a 
decrease in soil fertility is required and carried out by 
different actions: topsoil removal, offtake optimisation, 
or the currently popular C addition (for more details, see 
Török et al. 2011).
Two different studies, one from Switzerland and one 
from the Czech Republic, reported a long-term impact of 
fertilisation on Nardus grasslands. In the latter study, the 
impact of fertilisation was evident even 37 years after the 
last nutrient application, especially on behalf of P and Ca 
(Hegg et al. 1992, Hejcman et al. 2007). Therefore, due to 
their preference for oligotrophic soils, restoration of exten-
sive Nardus grasslands where fertilisers were applied in 
the past is difficult (Dähler 1992, Hejcman et al. 2007). 
Van Daele et al. (2017) claim that to restore species-rich 
Nardus grasslands, it is crucial to reduce bioavailable 
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Acta Biologica Slovenica, 2023, 66 (1)
P below 10 mg/kg or to select sites with bioavailable P 
contents below this threshold. Similarly, Korzeniak (2016) 
found that, especially for mesic Nardus grasslands in the 
lower montane zone, nutrient levels should be kept low to 
successfully control the expansion of nitrophilous species. 
For the restoration of species-rich Nardus grasslands, 
Van Daele et al. (2017) suggest inoculation of soil with a 
native soil community since native soil communities are 
known to promote restoration management (Middleton 
and Bever 2012, Wubs et al. 2016). Another criterion for 
selecting suitable sites for restoration could be a pH of 
approximately 4.5 to lift seed limitation. Last but not least, 
they recommend mowing in May, as this measure could 
reduce the competitive disadvantage of the slower-germi-
nating Nardus species. They emphasised that to increase 
the success of restoration management, knowledge 
about establishment limitation should be considered (Van 
Daele et al. 2017).
According to Schelfhout et al. (2017), a prerequisite for 
successful restoration is that the requirements regarding 
the abiotic conditions are met. These authors conducted 
a study where they tried to restore Nardus grasslands 
on formerly intensively managed agricultural land. They 
discovered that traditional mowing and grazing did not 
change community composition in such a way that it would 
resemble Nardus grasslands. They concluded that when 
threshold values for abiotic conditions are exceeded, 
abiotic restoration should be performed before biotic res-
toration. Therefore, it is crucial to perform measurements 
of important initial soil characteristics (e.g., pH, nutrients, 
etc.) before restoration. For abiotic restoration, the authors 
suggest P-mining or topsoil removal. Nevertheless, topsoil 
removal is a costly procedure from both the perspectives 
of time and money. Topsoil removal is, therefore, more 
feasible in larger restoration projects with more funding. 
Restoration projects with less funding should focus on 
sites that were previously managed less intensively 
(Schelfhout et al. 2017).
Post-restoration management 
with grazing and mowing
The most commonly applied methods in post-restoration 
management, used in the studies analysed, are listed in 
Tab. 3. It is very important to use an appropriate type of 
post-restoration management since optimally chosen 
post-restoration management may have an impact that 
is comparable to or even greater than the impact of a 
suitable restoration method (Paolinelli Reis et al. 2022).
Grasslands in Europe are semi-natural ecosystems 
that greatly depend on how they are managed (Butaye et 
al. 2005). This fact confirms the importance of disturbance 
in grasslands (Edwards et al. 2007), primarily mowing 
and/or grazing in restored grasslands, once basic vege-
tation is established (Butaye et al. 2005, Kiehl et al. 2010, 
Török et al. 2011). Disturbance in the form of grazing and 
mowing enhances the colonisation of target species by 
creating better germination and establishment conditions 
at microsites by reducing the cover of highly competitive 
species (Edwards et al. 2007). Cutting taller plants is 
especially effective, as it stimulates the establishment of 
sown forbs (Lawson et al. 2004) and improves plot diver-
sity (Török et al. 2011). One of the many positive impacts 
that grazing and mowing as forms of management also 
have on grasslands is the reduction of accumulated abo-
veground plant (litter) biomass since they open space and 
offer new niches for new plants to emerge and establish 
in the community (Török et al. 2011). This is a specificity of 
mesic grasslands due to their higher biomass production 
in comparison to xeric grasslands, which do not benefit 
from grazing and/or mowing as much (Hayes and Holl 
2003). Huhta et al. (2001) indicated that different mowing 
timeline has a different impact on grasslands: late mowing 
as a form of grassland upkeep and litter removal, whereas 
early mowing might shift species structure due to dis-
turbed seed production.
Turtureanu et al. (2014) claim that the highest plant 
species richness in semi-natural grasslands is provided 
by mowing. However, grazing and minor disturbance 
by animals may also be important for a higher diversity 
of plant species in semi-natural grasslands (Enyedi et al. 
2008). Mowing may affect species composition since, for 
example, early mowing facilitates early-flowering species. 
As opposed to mowing, grazing usually inhibits graminoids 
and promotes the development and replenishment of forb 
species. However, grazing has a positive impact on the 
development and germination of many important grass-
land species, primarily by providing open soil surfaces 
and niches by livestock trampling while grazing (Török 
et al. 2020). Nevertheless, grazing animals may exert a 
large force on the soil surface due to their heavy weight 
and, at the same time, relatively small hoof area (Bilotta 
et al. 2007). As a result, trampling may reduce both bio-
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Acta Biologica Slovenica, 2023, 66 (1)
Grassland management 
technique
Suitable for Advantages Limitations Reference /  
An example of use
Grazing mesic and semi-dry 
semi-natural grasslands, 
e.g., habitat types 
6210(*) and 6230*; 
not optimal for hay 
meadows and xeric 
grasslands
relatively frequent, but 
low-intensity disturbance; 
when carried out 
adequately (in line with 
species phenology), it can 
prevent shrub overgrowth 
and encroachment; 
enhancement of target 
species colonisation; 
removal of accumulated 
biomass and litter; grazing 
animals might serve as 
vectors for propagule 
transfer; decreases fire 
hazard; provision of open 
areas (niches) for the 
development of grassland 
species by animal trampling; 
increase and maintenance of 
microhabitat heterogeneity
selective (might cause an 
overgrowth of woody and 
thorny species, not palatable 
for certain types of livestock 
– primarily cows, so grazing 
by goats is recommended 
in this case); favours 
grasses over forbs due to 
their biology; dependent 
on species phenology; not 
suitable for xeric grasslands
Halada et al. 2001, 
Muller 2002, Hayes 
and Holl 2003, Pykälä 
2004, Edwards et al. 
2007, Galvánek and 
Lepš 2008, Török et al. 
2011, Turtureanu et al. 
2014, Török et al. 2016, 
Bonari et al. 2017, Tóth 
et al. 2018, Silva et al. 
2019, Köhler et al. 2020, 
Kurtogullari et al. 2020, 
Mrázková-Štýbnarová 
et al. 2020, Török et al. 
2020, Tölgyesi et al. 
2022, Zarzycki et al. 
2022
Mowing various semi-natural 
grasslands, including 
habitat types 6210(*), 
6230*, 6410, 6510, 6520
generally more accessible 
than grazing; early mowing 
facilitates the development 
of early-flowering species; 
enhancement of target 
species colonisation; 
improvement of the 
development of sown 
forbs by increasing 
competitiveness of higher 
plants; elimination of 
accumulated biomass and 
litter
beneficial only if carried 
out at an appropriate time 
in the season (when most 
diagnostic grassland forb 
species are ripe) and not 
too frequently (regime 
dependent on grassland 
type); when carried out 
with machinery, it is a 
non-selective and high-
intensity disturbance 
single event (compared to 
grazing) – hand mowing 
has more advantages 
that are similar to grazing, 
but it is costly and time-
consuming; might cause 
vegetation homogenisation; 
heavy machinery threatens 
invertebrate diversity
Halada et al. 2001, 
Muller 2002, Lawson 
et al. 2004, Edwards et 
al. 2007, Enyedi et al. 
2008, Galvánek and 
Lepš 2008, Královec et 
al. 2009, Halada et al. 
2011, Török et al. 2011, 
Valkó et al. 2012, Török 
et al. 2016, Bonari et al. 
2017, Milberg et al. 2017, 
Van Daele et al. 2017, 
Tälle et al. 2018, Török 
et al. 2020, Zarzycki et 
al. 2022
Prescribed burning (with 
long fire-return periods)
various types of 
grasslands
elimination of accumulated 
biomass and litter
might cause drastic species 
shift; might increase chances 
of biological invasions
Bensettiti et al. 2005, 
Galvánek and Janák 
2008, Valkó et al. 2014
Mulching nutrient-poor  
grasslands
prevents overgrowth by 
woody species; promotes 
the development of target 
species found at nutrient-
poor sites
Moog et al. 2002
Table 3. A list of post-restoration grassland management techniques used in the studies reviewed.
Tabela 3. Seznam tehnik vzdrževanja travnikov po njihovi obnovi, ki so bile uporabljene v pregledanih raziskavah.
diversity and vegetation cover at grazed sites (Matches 
1992). Thus, the load imposed on the soil by grazing 
animals should always be taken into account. The amount 
of pressure exerted on the soil differs according to the 
species and age of grazing animals (Bilotta et al. 2007). 
Furthermore, there are differences regarding grazing 
preference according to the type of grazing livestock. 
Cattle and horses feed on taller grasses, whereas sheep 
prefer short grasses and forbs (Tóth et al. 2018). Moreover, 
browsers, such as goats, can reduce shrub encroachment 
into grasslands (Elias et al. 2018). On Nardus grasslands, 
late onset of grazing, badly organised grazing, or too low 
grazing intensity might lead to the spreading of small 
shrubs, such as Vaccinium myrtillus or V. uliginosum, 
which results in a lower pasturing value of these habitats 
(Bensettiti et al. 2005).
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Acta Biologica Slovenica, 2023, 66 (1)
Grassland management 
technique
Suitable for Advantages Limitations Reference /  
An example of use
Cessation of fertilisation suitable for oligotrophic 
grasslands (such as 
Nardus grasslands); 
previously used arable 
fields
retains naturally occurring 
nutrient status of 
oligotrophic habitats
revegetation might be  
time-consuming
Critchley et al. 2007, 
Galvánek and Janák 
2008, Královec et al. 
2009, Korzeniak 2016, 
Van Daele et al. 2017, 
Kurtogullari et al. 2020
Mechanical elimination 
of woody vegetation
Nardus grasslands prevention of overgrowth at 
pasture edges
if not followed by regular 
management, it might cause 
worse shrub overgrowth 
compared to initial 
conditions
Galvánek and Janák 
2008, Parolo et al. 2011
Mixed management various semi-natural 
(including restored) 
grassland types
favours plant and butterfly 
diversity; takes into 
consideration the fact that 
different habitats require 
different approaches and 
that there is no ultimate 
solution that could be 
applied to all habitats
time- and resource-
challenging
Bonari et al. 2017
Grazing is a more selective form of management that 
favours rosette and prostrate plant life forms, so its aban-
donment favours the development of taller plant stands. 
Pykälä (2004) also found that grazing improves the 
diversity of different plant functional types and life forms 
in mesic grasslands more than in wet or arid habitats. The 
latter are additionally limited by natural limiting factors, 
such as drought or flooding. In this study, grazing partic-
ularly benefitted annual, biennial, and perennial plants, 
whereas geophytes had a negative response to grazing. 
Cattle grazing might increase the coexistence of different 
species with different life strategies and functional types, 
if given species are not under selective pressure by cows 
as a food source. Apart from species’ traits and habitat 
type, impacts of this specific disturbance are also geo-
graphically defined and, therefore, context-dependent. 
Török et al. (2011) noted that grazing could potentially 
have more positive impacts on grasslands in comparison 
to mowing since grazing animals can serve as vectors for 
propagule transfer. Furthermore, through their selective 
grazing, they can form more heterogeneous landscapes 
with different microclimatic and microstructural condi-
tions. However, selective grazing can negatively affect 
the diversity and establishment of target species since 
grazers (especially cows) usually avoid woody and thorny 
species. Consequently, these can overgrow the target 
area. Such common field weeds should be avoided in 
forage production and may be suppressed with occasional 
clean cuts. Otherwise, goat grazing is also recommended 
in areas under pressure by shrub vegetation (Török et al. 
2011). This was confirmed by Köhler et al. (2020), whose 
study showed that grazing by goats (browsing) had a 
positive effect on orchid-rich dry calcareous grasslands in 
Germany at a Natura 2000 site classified as habitat type 
6210(*), due to the presence of Gymnadenia conopsea, 
Orchis purpurea, O. militaris, Ophrys sphegodes, O. 
apifera, and O. insectifera. Browsing should be conducted 
in early spring in accordance with the phenology of 
orchids and the onset of shrub development, when young 
shrub plants are still palatable. The results of this study are 
promising. However, they should be further researched 
since this study was conducted on a relatively small scale 
and with orchid cover varying throughout the eight-year 
study. Grazing and trampling probably enhance light avail-
ability in pastures, promoting the annual recruitment of 
orchids (Köhler et al. 2020). The positive effects of grazing 
are also reflected in epizoochory (Tölgyesi et al. 2022).
Mowing is a more accessible form of management. 
However, it does not offer the same probability of possible 
improvement in grassland and landscape diversity as 
grazing does, often causing homogenisation of vege-
tation and negatively impacting invertebrate diversity 
in grasslands, particularly when carried out by heavy 
machinery. When possible, mowing by hand is recom-
mended, although this method itself is not very useful at a 
larger scale (Török et al. 2011).
Muller (2002) compared different management prac-
tices between different Natura 2000 grassland habitat 
types and denoted mowing without fertilisation as an 
adequate form of management for dry grasslands (6210(*)) 
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Acta Biologica Slovenica, 2023, 66 (1)
since fertilisation led to structurally- and diversity-de-
graded forms of habitats dominated by grasses. On the 
other hand, abandonment led to overgrowth by shrubs. 
Low-intensity mowing without fertilisation is also recom-
mended for Molinia grasslands (6410), which are very 
sensitive to eutrophication. Grazing without fertilisation 
turned out to be the best method for species-rich Nardus 
grasslands (6230*) since grazing prevents regressive 
succession to the previous form (heathlands), whereas 
fertilisation would disturb the oligotrophic nature of this 
habitat. Hay meadows (Natura 2000 habitat types 6510 
and 6520) require extensive management with low fertil-
isation and late cutting. Moog et al. (2002) noted that for 
Arrhenatherum elatius and Bromus erectus grasslands, 
the best management regimes for grassland conservation 
include regular grazing, mowing, and mulching since 
these practices promote the development of species that 
are typical of nutrient-poor habitats. On the other hand, 
irregular mulching was proved to have a very similar 
impact as natural succession. Succession favoured the 
development of seedlings of woody species, such as 
Fraxinus excelsior, Acer platanoides, A. pseudoplata-
nus, and species of nutrient-rich habitats. Therefore, to 
prevent grassland degradation, it is highly discouraged to 
cease their management. Královec et al. (2009) reported 
on the possibility of natural regeneration of agriculturally 
used fields following cessation of fertilisation and regular 
management in the form of multiple cuttings per year. 
However, this experiment was relatively small-scale and 
was probably highly dependent on the colonisation of 
target species in the proximity and regular management 
rather than on the seed bank itself. A positive impact of 
post-restoration management on dry grasslands, evident 
from the suppression of ruderal perennial vegetation, was 
experimentally proved by Kiehl and Pfidenhauer (2007), 
particularly in the later phases of experiments.
The importance of regular management of grassland 
habitat types 6210(*), 6410, and 6510 was confirmed by 
Milberg et al. (2017), whose results showed that in com-
parison to annual mowing, mowing every three years had 
a negative impact on plant diversity after 11-14 years from 
cessation of management. Cessation of management had 
the highest impact on short plants, so the authors recom-
mended extensive management (for example, mowing 
once every two years) on taller grasslands, whereas short-
grown swards are suited for annual mowing. Even though 
irregular mowing might not be the ideal solution for the 
conservation of all grassland plants, it is still more reason-
able and sustainable regarding diversity in comparison to 
abandonment, with additional benefits for pollinators.
One of the largest attempts at grassland restoration 
was carried out as part of the project LIFE04 NAT/HU/119 
titled ‘Grassland restoration and marsh protection in 
Egyek-Pusztakócs’ in the Hortobágy National Park in 
the Great Hungarian Plain (Valkó et al. 2021). Following 
restoration actions in the form of seed mixture sowing, 
the authors studied the importance of post-restoration 
management and the impact of seed bank on grassland 
development. Their results showed that cessation of man-
agement had a negative impact on both seed bank and 
vegetation stands, favouring weed development. Further-
more, seed bank itself had limited potential in grassland 
restoration since it mainly consisted of weed species and 
not target species. The similarity between vegetation 
stands and seed bank was low (Valkó et al. 2021).
When trying to reduce N. stricta cover in the case of 
N. stricta dominance on Nardus grasslands, it is generally 
better to choose cattle than sheep, as sheep usually avoid 
N. stricta (Grant et al. 1996), and plant species diversity 
increases under cattle grazing (Armstrong et al. 1997). 
Similarly, in a study in the Hrubý Jeseník Mountains in the 
Czech Republic, overall species richness increased on a 
previously long-term unmanaged pasture after six years 
of revived cattle grazing. In addition, rare and endangered 
species became more dominant. The authors concluded 
that this was probably the result of cattle trampling, which 
formed small open habitats that enabled the germination 
and survival of new species (Mrázková-Štýbnarová et 
al. 2020). However, according to Hejcman et al. (2008), 
sheep grazing proved to be a suitable management 
practice for degraded meadows in the Giant Mts. in the 
Czech Republic, as it reduced the extent of undesirable 
species typical of long-term abandoned swards. Crawley 
(1983) even promotes mixed stocking with sheep and 
cattle (or goats), as one species improves the environment 
for the other, and thus facilitation is enabled. Accordingly, 
Holland et al. (2008) observed that the combined use of 
cattle and sheep was more effective in creating struc-
tural change on N. stricta grasslands compared to when 
sheep alone were used. Next, Kurtogullari et al. (2020) 
believe that pastures at lower altitudes should have a 
lower grazing intensity. Velev and Apostolova (2008) also 
found in their study in Bulgaria that the abundance of N. 
stricta was lower at lower grazing intensity. This could 
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Acta Biologica Slovenica, 2023, 66 (1)
be achieved by moving cattle to higher altitudes earlier 
during the season. At higher altitudes, cattle should first 
be sent and enclosed in areas where N. stricta is dominant 
in order to reduce its cover as long as it is still palatable 
(Kurtogullari et al. 2020).
According to Bedia and Busqué (2013), the most 
suitable management practice for maintaining spe-
cies-rich Nardus grasslands is grazing by large herbivores, 
as it reduces not only N. stricta dominance through defo-
liation, but also shrub encroachment through trampling 
and fertilisation (Hartley and Mitchell 2005). In general, 
low-intensity management by mowing, grazing, or a 
combination of both is crucial to retain Nardus grasslands 
(Krahulec et al. 2001, Dullinger et al. 2003). A combination 
is especially beneficial, as it enables the occurrence of 
a greater number of different species (Galvánek and 
Janák 2008). Most of all, a spatially variable disturbance 
regime is needed (Dullinger et al. 2003). Parolo et al. 
(2011) showed that heterogeneous grazing management 
had a positive impact on plant diversity in the alpine spe-
cies-rich Nardus pastures in Italia. Likewise, Zarzycki et al. 
(2022) claim that to preserve the biodiversity of mountain 
grasslands, it is vital to preserve a mosaic spatial structure 
and retain extensive management practices with various 
forms of human impact, such as mowing and grazing. On 
the contrary, Vassiliev et al. (2011) suggested the forma-
tion of sheep pens for a more spatially uniform grassland 
management within the mountain in less accessible sites 
with abandoned pastures in the Western Balkan Mts. in 
Bulgaria. However, they also advocated extensive man-
agement with a zonation regime (Vassiliev et al. 2011). 
Yearly rotational grazing is also recommended to limit 
woody encroachment in remote areas on the one hand 
and also overgrazing near populated areas on the other 
hand (Başnou et al. 2009). In a study by Lüth et al. (2011), 
the Sieversio montanae-Nardetum strictae grasslands 
were most species-rich in the case of mowing, slight 
fertilisation, and grazing in autumn. Thus, they suggested 
that traditional hay management is the most appropriate 
practice (Lüth et al. 2011). Some use of manure or leaving 
cut grass on the ground every now and then is advisable 
when the only management practice on Nardus grass-
lands is mowing (Háková et al. 2004), as regular removal of 
biomass might gradually lead to oligotrophisation of these 
grasslands, especially on very poor soils, which reflects in 
a lower number of species (Krahulec et al. 1996, Halada 
et al. 2001). However, several soil parameters have to be 
considered first to prevent eutrophication (Galvánek and 
Janák 2008). In the case when Nardus grasslands are only 
mown, it is also highly recommended to introduce artificial 
disturbance to ensure space for those plant species that 
are less competitive (Háková et al. 2004). Fischer and 
Wipf (2002) advised to continue with traditional mowing of 
subalpine meadows and to switch back to mowing in the 
case of recently grazed meadows, as grazing negatively 
affected plant species richness in their study. However, 
this traditional management practice presents high costs 
and is, therefore, often no longer feasible (Galvánek and 
Janák 2008). Considering this, traditional mowing may 
also be alternated with sheep grazing. In fact, the alter-
nation between mowing and grazing is advisable not only 
from the perspective of lowering the costs, but also to 
suppress the spreading of invasive species, as utilisation 
of only one restoration technique is often not enough to 
curb the expansion of invasive species (Pecháčková and 
Krahulec 1995, Krahulec et al. 2001). Pecháčková and 
Krahulec (1995) reported that by using multiple restoration 
techniques at once, it is possible to restore species-rich 
grasslands in three to five years. Nonetheless, Halada 
et al. (2001) found that the vitality of invasive species 
was also suppressed by applying regular mowing only. 
Along with the position of grasslands along the altitudinal 
gradient, management practices should also always take 
into consideration the sensitivity of grasslands to isolation 
and fragmentation (Reitalu et al. 2012, Janišová et al. 
2014). Korzeniak (2016) stated that the above-mentioned 
suggestions are necessary especially for thermophilous 
grasslands with Nardus in the lower montane zone.
Evaluation of grassland 
restoration techniques
Evaluation of grassland restoration techniques and their 
comparison is a very demanding task since every resto-
ration action has specific initial conditions at both target 
and donor sites (Török et al. 2011). Every restoration 
action depends on interspecies’ interactions on the field, 
particularly competition (Hölzel and Otte 2003), and there-
fore requires individual approaches (Török et al. 2011). 
Additionally, technical details differ for each restoration 
action. Different methods are generally harder to compare. 
Therefore, only similar actions should be compared (e.g., 
sowing of seeds and addition of plant material). In general, 
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any restoration planning effort should start with the col-
lection of reliable information regarding the previous and 
current state of chosen plots. The choice of methodology 
should be carefully considered and planned according to 
the target vegetation and conditions at both target and 
donor sites, which should be ecologically similar (Török 
et al. 2011). Sengl et al. (2017) noted that commonly used 
variables, such as similarity and dissimilarity indices, 
biodiversity indices, number and coverage of chosen func-
tional plant groups, and species number, are applicable 
only if sites are similar or located nearby, and not when 
they are distant and/or degraded. For instance, sites with 
high diversity could be degraded by invasive species and 
weeds, and thus their ecosystem services could be dis-
rupted. Therefore, Sengl et al. (2017) recommend the eval-
uation of grassland naturalness, more specifically, the use 
of ecological indicator values, which explain the state of 
grasslands through the presence of specific plant species, 
level of invasiveness, grassland functional diversity, and 
resilience of grasslands to disturbance and degradation.
The evaluation of success is highly context-depen-
dent. According to Prach et al. (2021a), varying abiotic and 
biotic factors at chosen sites affect restoration success 
and are also influenced by the chosen restoration method. 
Furthermore, soil and landscape characteristics are also 
highly important. Their experiment showed that the use 
of regional seed mixtures benefitted the establishment of 
dry grasslands, whereas methods like natural succession 
and commercial seed mixtures favoured the establish-
ment of mesic grassland species (Prach et al. 2021a). They 
also noted that researchers should evaluate measures of 
post-restoration management, which would primarily help 
in the establishment of characteristic target species. In 
addition, restoration must also be thought out from the 
technical and economic points of view. When possible, the 
desired restoration actions should be tested. Restoration 
progress should be monitored and sampled using an 
appropriate methodology, which should be able to show 
the differences and results of revegetation attempts. In 
addition, these should be documented for future refer-
ence (Török et al. 2011). The survival of newly established 
plant populations is often limited by ecosystem services, 
such as seed production and dispersal, but also by other 
factors, such as lack of suitable pollinators, limited genetic 
diversity in sexual mating, vegetative reproduction by 
dominant species, herbivory, and inadequately planned 
management (Albert et al. 2021). Albert et al. (2021) found 
that donor sites provide more ecosystem services in com-
parison to recipient sites. These show signs of pressure 
by previous arable use and dominance of ruderals 
and weeds, supporting mostly herbivory, whereas the 
presence of late-flowering meadow generalists supports 
pollination and pollinator diversity on donor grasslands.
Conclusions
Today, grasslands are under threat all over the world, 
mainly due to human activity and climate change, which is 
why their ability to provide ecosystem services is rapidly 
decreasing.
The agri-environment measures for the preservation 
of high nature value grasslands in Slovenia are not suc-
cessful for various reasons. Therefore, they should be 
reconsidered and based on a different basis.
Given the many benefits that grasslands provide, it 
is vital to prevent their further decline. The best way to 
maintain grassland biodiversity is to ensure traditional 
extensive agricultural practices, such as mowing and/
or grazing. For this reason, these two practices are also 
often recommended for grassland restoration. In order to 
successfully maintain grassland biodiversity, not only one 
type of appropriate practice but rather the introduction of 
a wider scheme of traditional management practices is 
necessary. In addition, mosaic management should also 
be taken into account.
In the case of partly degraded grasslands, a reduction 
in management intensity is usually sufficient for their 
successful restoration. On the other hand, for entirely 
degraded grasslands, spontaneous succession or tech-
nical reclamation (for example, sowing of regional seed 
mixtures or transfer of plant material) is necessary for their 
recovery. In any case, the proximity of well-preserved 
donor grasslands, which serve as a natural source of repro-
ductive units (propagules) for degraded grasslands, is key 
to the success of larger grassland restoration projects.
Restoration of species-rich grasslands can be a very 
time-consuming process and usually requires a certain 
amount of soil preparation on the land that will be subject to 
restoration, as well as further management after restoration. 
It was found that optimally chosen post-restoration man-
agement may have an impact that is comparable to or even 
greater than the impact of a suitable restoration method.
When designing restoration studies, it is always neces-
sary to take into account the conditions and area of donor 
72
Acta Biologica Slovenica, 2023, 66 (1)
and recipient grassland plots, as well as the phenology of 
the community and target grassland species.
When selecting donor plots for grassland restoration, 
attention should be paid to the fact that donor plots should 
be compatible with recipient plots in terms of nutrient 
status, hydrological conditions, and substrate. In addition, 
donor plots must also demonstrate a biogeographically and 
phytocoenologically representative plant community com-
position, with as little presence of neophytes as possible.
In any case, active maintenance of well-preserved 
grasslands that have not yet been degraded is more 
cost-effective than the restoration of grasslands that have 
already been degraded.
Funding
This review was written within the project LIFE20 NAT/
SI/000253 titled ‘Conservation of priority grassland 
habitats in Slovenia through the establishment of the 
seed bank and in situ restoration’ (acronym: LIFE FOR 
SEEDS), which is co-financed by the European Union 
from the LIFE programme, Sigrid Rausing Trust, and 
the Ministry of Public Administration of the Republic of 
Slovenia. The authors would also like to acknowledge the 
programme P4-0072 and the CRP project V4-2021. The 
review reflects the authors’ view, and financers are not 
responsible for any use that may be made of the informa-
tion it contains.
References
Albert, Á.-J., Kelemen, A., Valkó, O., Miglécz, T., Csecserits, A., Rédei, T., Deák, B., Tóthmérész, B., Török, P., 2014. Secondary succession in sandy old fields: 
a promising example of spontaneous grassland recovery. Applied Vegetation Science, 17, 214-224.
Albert, Á.-J., Mudrák, O., Jongepierová, I., Fajmon, K., Frei, I., Ševčíková, M., Klimešová, J., Doležal, J., 2019. Grassland restoration on ex-arable land by 
transfer of brush-harvested propagules and green hay. Agriculture, Ecosystems and Environment, 272, 74-82.
Albert, Á.-J., Götzenberger, L., Jongepierová, I., Konečná, M., Lőkkösné-Kelbert, B., Májeková, M., Mudrák, O., Klimešová, J., 2021. Restoration of ecosystem 
functions: seed production in restored and ancient grasslands. Applied Vegetation Science, 24, 202107.
Allen, V.G., Batello, C., Berretta, E.J., Hodgson, J., Kothmann, M., Li, X., McIvor, J., Milne, J., Morris, C., Peeters, A., Sanderson, M., 2011. An international 
terminology for grazing lands and grazing animals. Grass and Forage Science, 66, 2-28.
Ammann, C., Flechard, C.R., Leifeld, J., Neftel, A., Fuhrer, J., 2007. The carbon budget of newly established temperate grassland depends on management 
intensity. Agriculture, Ecosystems & Environment, 121, 5-20.
Armstrong, R.H., Grant, S.A., Common, T.G., Beattie, M.M., 1997. Controlled grazing studies on Nardus grassland: effects of between tussock sward height 
and species of grazer on diet selection and intake. Grass and Forage Science, 52, 219-231.
Ashton, M.S., Tyrrell, M.L., Spalding, D., Gentry, B. (eds.), 2012. Managing Forest Carbon in a Changing Climate. Springer Science+Business Media B.V., 
Dordrecht.
Babai, D., Molnár, Z., 2014. Small-scale traditional management of highly species-rich grasslands in the Carpathians. Agriculture, Ecosystems & Environment, 
182, 123-130.
Baasch, A., Engst, K., Schmiede, R., May, K., Tischew, S., 2016. Enhancing success in grassland restoration by adding regionally propagated target species. 
Ecological Engineering, 94, 583-591.
Başnou, C., Pino, J., Šmilauer, P., 2009. Effect of grazing on grasslands in the Western Romanian Carpathians depends on the bedrock type. Preslia, 81, 
91-104.
Bedia, J., Busqué, J., 2013. Productivity, grazing utilization, forage quality and primary production controls of species-rich alpine grasslands with Nardus 
stricta in northern Spain. Grass and Forage Science, 68, 297-312.
Bensettiti, F., Boullet, V., Chavaudret-Laborie, C., Deniaud, J. (eds.), 2005. “Habitats manuals” Natura 2000. Knowledge and management of habitats and 
species of community interest. Tome 4. Agropastoral habitats. Volumes 1 and 2, 445 pp. and 487 pp.
Bilotta, G.S., Brazier, R.E., Haygarth, P.M., 2007. The impacts of grazing animals on the quality of soils, vegetation, and surface waters in intensively 
managed grasslands. Advances in Agronomy, 94, 237-280.
Biondini, M.E., Steuter, A.A., Grygiel, C.E., 1989. Seasonal fire effects on the diversity patterns, spatial distribution and community structure of forbs in the 
Northern Mixed Prairie, USA. Vegetatio, 85, 21-31.
73
Acta Biologica Slovenica, 2023, 66 (1)
Bischoff, A., Warthemann, G., Klotz, S., 2009. Succession of floodplain grasslands following reduction in land use intensity: the importance of environmental 
conditions, management and dispersal. Journal of Applied Ecology, 46, 241-249.
Bonari, G., Fajmon, K., Malenovský, I., Zelený, D., Holuša, J., Jongepierová, I., Kočárek, P., Konvička, O., Uřičár, J., Chytrý, M., 2017. Management of semi-
natural grasslands benefiting both plant and insect diversity: the importance of heterogeneity and tradition. Agriculture, Ecosystems and Environment, 
246, 243-252.
Bossuyt, B., Honnay, O., 2008. Can the seed bank be used for ecological restoration? An overview of seed bank characteristics in European communities. 
Journal of Vegetation Science, 19, 875-884.
Burton, R.J.F., Schwarz, G., 2013. Result-oriented agri-environmental schemes in Europe and their potential for promoting behavioural change. Land Use 
Policy, 30, 628-641.
Butaye, J., Adriaens, D., Honnay, O., 2005. Conservation and restoration of calcareous grasslands: a concise review of the effects of fragmentation and 
management on plant species. Biotechnologie, Agronomie, Société et Environnement, 9, 111-118.
Common, T.G., Hunter, E.A., Floate, M.J.S., Eadie, J., Hodgson, J., 1991. The long-term effects of a range of pasture treatments applied to three semi-natural 
hill grassland communities. 1. Pasture production and botanical composition. Grass and Forage Science, 46, 239-251.
Crawley, M.J., 1983. Herbivory: The Dynamics of Animal-Plant Interactions. Studies in Ecology, Volume 10. Blackwell Scientific Publications, Oxford, 437 pp.
Critchley, C.N.R., Fowbert, J.A., Wright, B., 2007. Dynamics of species-rich upland hay meadows over 15 years and their relation with agricultural 
management practices. Applied Vegetation Science, 10, 307-314.
Dajić Stevanović, Z., Peeters, A., Vrbničanin, S., Šoštarić, I., Aćić, S., 2008. Long term grassland vegetation changes: case study Nature Park Stara Planina 
(Serbia). Community Ecology, 9, 23-31.
Davidova, S., 2011. Semi-subsistence farming: an elusive concept posing thorny policy questions. Journal of Agricultural Economics, 62, 503-524.Dähler, 
W., 1992. Long term influence of fertilization in a Nardetum. Vegetatio, 103, 141-150.
Donath, T.W., Hölzel, N., Otte, A., 2006. Influence of competition by sown grass, disturbance and litter on recruitment of rare flood-meadow species. 
Biological Conservation, 130, 315-323.
Donath, T.W., Bissels, S., Hölzel, N., Otte, A., 2007. Large scale application of diaspore transfer with plant material in restoration practice – impact of seed 
and microsite limitation. Biological Conservation, 138, 224-234.
Dullinger, S., Dirnböck, T., Greimler, J., Grabherr, G., 2003. A resampling approach for evaluating effects of pasture abandonment on subalpine plant 
species diversity. Journal of Vegetation Science, 14, 243-251.
Edwards, A.R., Mortimer, S.R., Lawson, C.S., Westbury, D.B., Harris, S.J., Woodcock, B.A., Brown, V.K., 2007. Hay strewing, brush harvesting of seed and 
soil disturbance as tools for the enhancement of botanical diversity in grasslands. Biological Conservation, 134, 372-382.
EEA (European Environment Agency), 2020. State of nature in the EU. Results from reporting under the nature directives 2013–2018. EEA Report No 
10/2020.
Elias, D., Hölzel, N., Tischew, S., 2018. Goat paddock grazing improves the conservation status of shrub-encroached dry grasslands. Tuexenia, 38, 215-233.
Enyedi, Z.M., Ruprecht, E., Deák, M., 2008. Long-term effects of the abandonment of grazing on steppe-like grasslands. Applied Vegetation Science, 11, 
55-62.
Ewing, A.L., Engle, D.M., 1988. Effects of late summer fire on tallgrass prairie microclimate and community composition. The American Midland Naturalist, 
120, 212-223.
Factsheet on the 2014-2022 Rural Development Programme for Slovenia, 2022. European Commission. https://agriculture.ec.europa.eu/system/
files/2022-10/rdp-factsheet-slovenia_en.pdf (16. 3. 2023).
Fagan, K.C., Pywell, R.C., Bullock, J.M., Marrs, R.H., 2008. Do restored calcareous grasslands on former arable fields resemble ancient targets? The effect 
of time, methods and environment on outcomes. Journal of Applied Ecology, 45, 1293-1303.
Fischer, M., Wipf, S., 2002. Effect of low-intensity grazing on the species-rich vegetation of traditionally mown subalpine meadows. Biological Conservation, 
104, 1-11.
Fraser, L.H., Pither, J., Jentsch, A., Sternberg, M., Zobel, M., Askarizadeh, D., … Zupo, T., 2015. Worldwide evidence of a unimodal relationship between 
productivity and plant species richness. Science, 349, 302-305.
Galvánek, D., Janák, M., 2008. Management of Natura 2000 habitats. 6230 *Species-rich Nardus grasslands. Technical Report 2008 14/24. European 
Commission, Brussels, 24 pp.
Galvánek, M., Lepš, J., 2008. Changes of species richness pattern in mountain grasslands: abandonment vs. restoration. Biodiversity and Conservation, 
17, 3241-3253.
Goret, T., Janssens, X., Godefroid, S., 2021. A decision-making tool for restoring lowland grasslands in Europe. Journal for Nature Conservation, 63, 
126046.
Grant, S.A., Torvell, L., Sim, E.M., Small, J.L., Armstrong, R.H., 1996. Controlled grazing studies on Nardus grassland: effects of between-tussock sward 
height and species of grazer on Nardus utilization and floristic composition in two fields in Scotland. Journal of Applied Ecology, 33, 1053-1064.
Halada, Ľ., Ružičková, H., David, S., 2001. Monitoring of the grassland communities in the East Carpathians biosphere reserve – basis for the future ILTER 
site. Ekológia, 20, 107-114.
74
Acta Biologica Slovenica, 2023, 66 (1)
Halada, Ľ., Evans, D., Romão, C., Petersen, J.-E., 2011. Which habitats of European importance depend on agricultural practices?. Biodiversity Conservation, 
20, 2365-2378.
Háková, A., Klaudisová, A., Sádlo, J. (eds.), 2004. Zásady péče o nelesní biotopy v rámci soustavy Natura 2000. Planeta XII, 8/2004, druhá část. Ministerstvo 
životního prostředí, Praha, 144 pp.
Hartley, S.E., Mitchell, R.J., 2005. Manipulation of nutrients and grazing levels on heather moorland: changes in Calluna dominance and consequences for 
community composition. Journal of Ecology, 93, 990-1004.
Hartnett, D.C., Hickman, K.R., Walter, L.E.F., 1996. Effects of bison grazing, fire, and topography on floristic diversity in tallgrass prairie. Journal of Range 
Management, 49, 413-420.
Hartvigsen, M., 2014. Land reform and land fragmentation in Central and Eastern Europe. Land Use Policy, 36, 330-341.
Haslgrübler, P., Krautzer, B., Blaschka, A., Graiss, W., Pötsch, E.M., 2013. Quality and germination capacity of seed material harvested from an Arrhenatherion 
meadow. Grass and Forage Science, 69, 454-461.
Hayes, G.F., Holl, K.D., 2003. Cattle grazing impacts on annual forbs and vegetation composition of mesic grasslands in California. Conservation Biology, 
17, 1694-1702.
Hegg, O., Feller, U., Dähler, W., Scherrer, C., 1992. Long term influence of fertilization in a Nardetum: phytosociology of the pasture and nutrient contents 
in leaves. Vegetatio, 103, 151-158.
Heinsoo, K., Melts, I., Sammul, M., Holm, B., 2010. The potential of Estonian semi-natural grasslands for bioenergy production. Agriculture, Ecosystems & 
Environment, 137, 86-92.
Hejcman, M., Klaudisová, M., Štursa, J., Pavlů, V., Schellberg, J., Hejcmanová, P., Hakl, J., Rauch, O., Vacek, S., 2007. Revisiting a 37 years abandoned 
fertilizer experiment on Nardus grassland in the Czech Republic. Agriculture, Ecosystems & Environment, 118, 231-236.
Hejcman, M., Žáková, I., Bílek, M., Bendová, P., Hejcmanová, P., Pavlů, V., Stránská, M., 2008. Sward structure and diet selection after sheep introduction 
on abandoned grassland in the Giant Mts, Czech Republic. Biologia, 63, 506-514.
Herzon, I., Birge, T., Allen, B., Povellato, A., Vanni, F., Hart, K., Radley, G., Tucker, G., Keenleyside, C., Oppermann, R., Underwood, E., Poux, X., Beaufoy, 
G., Pražan, J., 2018. Time to look for evidence: results-based approach to biodiversity conservation on farmland in Europe. Land Use Policy, 71, 347-354.
Hofmann, M., Isselstein, J., 2004. Seedling recruitment on agriculturally improved mesic grassland: the influence of disturbance and management schemes. 
Applied Vegetation Science, 7, 193-200.
Hofmann, S., Conradi, T., Kiehl, K., Albrecht, H., 2020. Effects of different restoration treatments on long-term development of plant diversity and functional 
trait composition in calcareous grasslands. Tuexenia, 40, 175-200.
Holland, J.P., Waterhouse, A., Robertson, D., Pollock, M.L., 2008. Effect of different grazing management systems on the herbage mass and pasture height 
of a Nardus stricta grassland in western Scotland, United Kingdom. Grass and Forage Science, 63, 48-59.
Hölzel, N., Otte, A., 2003. Restoration of a species-rich flood meadow by topsoil removal and diaspore transfer with plant material. Applied Vegetation 
Science, 6, 131-140.
Huhta, A.-P., Rautio, P., Tuomi, J., Laine, K., 2001. Restorative mowing on an abandoned semi-natural meadow: short-term and predicted long-term effects. 
Journal of Vegetation Science, 12, 677-686.
Ivajnšič, D., Pintarič, D., Škornik, S., Kaligarič, M., Pipenbaher, N., 2019. SOSKOPOP Haloze: podporni sistem potencialnim uveljaviteljem ukrepov KOPOP 
na nivoju travišč. Revija za geografijo, 14, 49-64.
Janišová, M., Michalcová, D., Bacaro, G., Ghisla, A., 2014. Landscape effects on diversity of semi-natural grasslands. Agriculture, Ecosystems & Environment, 
182, 47-58.
Johanidesová, E., Fajmon, K., Jongepierová, I., Prach, K., 2014. Spontaneous colonization of restored dry grasslands by target species: restoration proceeds 
beyond sowing regional seed mixtures. Grass and Forage Science, 70, 631-638.
Jones, M.B., Donnelly, A., 2004. Carbon sequestration in temperate grassland ecosystems and the influence of management, climate and elevated CO2. 
New Phytologist, 164, 423-439.
Jongepierová, I., Mitchley, J., Tzanopoulous, J., 2007. A field experiment to recreate species rich hay meadows using regional seed mixtures. Biological 
Conservation, 139, 297-305.
Kaligarič, M., Čuš, J., Škornik, S., Ivajnšič, D., 2019. The failure of agri-environment measures to promote and conserve grassland biodiversity in Slovenia. 
Land Use Policy, 80, 127-134.
Kardol, P., Van der Wal, A., Bezemer, T.M., de Boer, W., Duyts, H., Holtkamp, R., Van der Putten, W.H., 2008. Restoration of species-rich grasslands on ex-
arable land: seed addition outweighs soil fertility reduction. Biological Conservation, 141, 2208-2217.
Kiehl, K., Thormann, A., Pfadenhauer, J., 2006. Evaluation of initial restoration measures during the restoration of calcareous grasslands on former arable 
fields. Restoration Ecology, 14, 148-156.
Kiehl, K., Pfidenhauer, J., 2007. Establishment and persistence of target species in newly created calcareous grasslands on former arable fields. Plant 
Ecology, 189, 31-48.
Kiehl, K., Kirmer, A., Donath, T.W., Rasran, L., Hölzel, N., 2010. Species introduction in restoration projects – evaluation of different techniques for the 
establishment of semi-natural grasslands in Central and Northwestern Europe. Basic and Applied Ecology, 11, 285-299.
75
Acta Biologica Slovenica, 2023, 66 (1)
Klaus, V.H., Hoever, C.J., Fischer, M., Hamer, U., Kleinebecker , T., Mertens, D., Schäfer, D., Prati, D., Hölzel, N., 2018. Contribution of the soil seed bank to 
the restoration of temperate grasslands by mechanical sward disturbance. Restoration Ecology, 26, S114-S122.
Kleijn, D., 2003. Can establishment characteristics explain the poor colonization success of late successional grassland species on ex-arable land?. 
Restoration Ecology, 11, 131-138.
Klimeš, L., Hájek, M., Mudrák, O., Dančák, M., Preislerová, Z., Hájková, P., Jongepierová, I., Klimešová, J., 2013. Effects of changes in management on 
resistance and resilience in three grassland communities. Applied Vegetation Science, 16, 640-649.
Knappová, J., Hemrová, L., Münzbergová, Z., 2012. Colonization of central European abandoned fields by dry grassland species depends on the species 
richness of the source habitats: a new approach for measuring habitat isolation. Landscape Ecology, 27, 97-108.
Korzeniak, J., 2016. Mountain Nardus stricta grasslands as a relic of past farming – the effects of grazing abandonment in relation to elevation and spatial 
scale. Folia Geobotanica, 51, 93-113.
Köhler, M., Elias, D., Hiller, G., Hölzel, N., Tischew, S., 2020. Restoration of orchid-rich dry calcareous grasslands by rotational goat pasturing. Tuexenia, 
40, 201-223.
Krahulec, F., Blažková, D., Balátová-Tuláčková, E., Štursa, J., Pecháčková, S., Fabšičová, M., 1996. Louky Krkonoš: rostlinná společenstva a jejich dynamika. 
Opera Corcontica, 33. Správa Krkonošského národního parku, Vrchlabí, 252 pp.
Krahulec, F., Skálová, H., Herben, T., Hadincová, V., Wildová, R., Pecháčková, S., 2001. Vegetation changes following sheep grazing in abandoned mountain 
meadows. Applied Vegetation Science, 4, 97-102.
Krahulec, F., Chytrý, M., Härtel, H., 2007. Nardus grasslands and heathlands. In: Chytrý, M. (ed.): Vegetation of the Czech Republic. 1. Grassland and 
Heathland Vegetation. Academia, Prague, pp. 281-306.
Královec, J., Pocová, L., Jonášová, M., Macek, P., Prach, K., 2009. Spontaneous recovery of an intensively used grassland after cessation of fertilizing. 
Applied Vegetation Science, 12, 391-397.
Krautzer, B., Bartel, A., Kirmer, A., Tischew, S., Feucht, B., Wieden, M., Haslgrübler, P., Rieger, E., Pötsch, E.M., 2011. Establishment and use of high nature 
value farmland. In: Proceedings of the 16th EGF Symposium on “Grassland farming and land management systems in mountainous regions”. Gumpenstein, 
Austria, August 29-31, pp. 457-469.
Kurtogullari, Y., Rieder, N.S., Arlettaz, R., Humbert, J.-Y., 2020. Conservation and restoration of Nardus grasslands in the Swiss northern Alps. Applied 
Vegetation Science, 23, 26-38.
Kuzemko, A.A., Steinbauer, M.J., Becker, T., Didukh, Y.P., Dolnik, C., Jeschke, M., Naqinezhad, A., Uğurlu, E., Vassilev, K., Dengler, J., 2016. Patterns and 
drivers of phytodiversity in steppe grasslands of Central Podolia (Ukraine). Biodiversity and Conservation, 25, 2233-2250.
Lawson, C.S., Ford, M.A., Mitchley, J., 2004. The influence of seed addition and cutting regime on the success of grassland restoration on former arable 
land. Applied Vegetation Science, 7, 259-266.
Lencová, K., Prach, K., 2011. Restoration of hay meadows on ex-arable land: commercial seed mixtures vs. spontaneous succession. Grass and Forage 
Science, 66, 265-271.
Lengyel, S., Varga, K., Kosztyi, B., Lontay, L., Déri, E., Török, P., Tóthmérész, B., 2012. Grassland restoration to conserve landscape-level biodiversity: a 
synthesis of early results from a large-scale project. Applied Vegetation Science, 15, 264-276.
Lepš, J., Doležal, J., Bezemer, T.M., Brown, V.K., Hedlund, K., Igual Arroyo, M., Jörgensen, H.B., Lawson, C.S., Mortimer, S.R., Peix Geldart, A., Rodríguez 
Barrueco, C., Santa Regina, I., Šmilauer, P., van der Putten, W.H., 2007. Long-term effectiveness of sowing high and low diversity seed mixtures to enhance 
plant community development on ex-arable fields. Applied Vegetation Science, 10, 97-110.
Lesschen, J.P., Elbersen, B., Hazeu, G., van Doorn, A., Mucher, S., Velthof, G., 2014. Defining and Classifying Grasslands in Europe. Wageningen University 
and Research, Wageningen.
Lovec, M., Šumrada, T., Erjavec, E., 2020. New CAP delivery model, old issues. Intereconomics, 2, 112-119.
Lüth, C., Tasser, E., Niedrist, G., Dalla Via, J., Tappeiner, U., 2011. Classification of the Sieversio montanae-Nardetum strictae in a cross-section of the 
Eastern Alps. Plant Ecology, 212, 105-126.
Marriott, C.A., Fothergill, M., Jeangros, B., Scotton, M., Louault, F., 2004. Long-term impacts of extensification of grassland management on biodiversity 
and productivity in upland areas. A review. Agronomie, 24, 447-462.
Matches, A.G., 1992. Plant response to grazing: a review. Journal of Production Agriculture, 5, 1-7.
Mayel, S., Jarrah, M., Kuka, K., 2021. How does grassland management affect physical and biochemical properties of temperate grassland soils? A review 
study. Grass Forage Science, 76, 215-244.
McLeod, R.F., Gates, J.E., 1998. Response of herpetofaunal communities to forest cutting and burning at Chesapeake Farms, Maryland. The American 
Midland Naturalist, 139, 164-177.
Merunková, K., Chytrý, M., 2012. Environmental control of species richness and composition in upland grasslands of the southern Czech Republic. Plant 
Ecology, 213, 591-602.
Middleton, E.L., Bever, J.D., 2012. Inoculation with a native soil community advances succession in a grassland restoration. Restoration Ecology, 20, 218-226.
Milberg, P., Tälle, M., Fogelfors, H., Westerberg, L., 2017. The biodiversity cost of reducing management intensity in species-rich grasslands: mowing 
annually vs. every third year. Basic and Applied Ecology, 22, 61-74.
76
Acta Biologica Slovenica, 2023, 66 (1)
Milchunas, D.G., Sala, O.E., Lauenroth, W.K., 1988. A generalized model of the effects of grazing by large herbivores on grassland community structure. 
The American Naturalist, 132, 87-106.
Mitchell, R.J., Rose, R.J., Palmer, S.C.F., 2009. The effect of restoration techniques on non-target species: case studies in moorland ecosystems. Applied 
Vegetation Science, 12, 81-91.
Mitchley, J., Jongepierová, I., Fajmon, K., 2012. Regional seed mixtures for the re-creation of species-rich meadows in the White Carpathian Mountains: 
result of a 10-yr experiment. Applied Vegetation Science, 15, 253-263.
Moog, D., Poschlod, P., Kahmen, S., Schreiber, K.-F., 2002. Comparison of species composition between different grassland management treatments after 
25 years. Applied Vegetation Science, 5, 99-106.
Mrázková-Štýbnarová, M., Holec, J., Štencl, R., Kolářová, M., Tyšer, L., Oldřich, L. 2020., Impact of renewed cattle grazing on floristic composition in the 
Hrubý Jeseník Mountains. Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis, 68, 335-342.
Muller, S., 2002. Appropriate agricultural management practices required to ensure conservation and biodiversity of environmentally sensitive grassland 
sites designated under Natura 2000. Agriculture, Ecosystems & Environment, 89, 261-266.
Palpurina, S., Wagner, V., von Wehrden, H., Hájek, M., Horsák, M., Brinkert, A., … Chytrý, M., 2017. The relationship between plant species richness and soil 
pH vanishes with increasing aridity across Eurasian dry grasslands. Global Ecology and Biogeography, 26, 425-434.
Paolinelli Reis, B., Szitár, K., Kövendi-Jakó, A., Török, K., Sáradi, N., Csákvári, E., Halassy, M., 2022. The long-term effect of initial restoration intervention, 
landscape composition, and time on the progress of Pannonic sand grassland restoration. Landscape and Ecological Engineering, 18, 429-440.
Parolo, G., Abeli, T., Gusmeroli, F., Rossi, G., 2011. Large-scale heterogeneous cattle grazing affects plant diversity and forage value of Alpine species-rich 
Nardus pastures. Grass and Forage Science, 66, 541-550.
Pecháčková, S., Krahulec, F., 1995. Efficient nitrogen economy: key to the success of Polygonum bistorta in an abandoned mountain meadow. Folia 
Geobotanica et Phytotaxonomica, 30, 211-222.
Pe’er, G., Finn, J.A., Díaz, M., Birkenstock, M., Lakner, S., Röder, N., Kazakova, Y., Šumrada, T., Bezäk, P., Concepción, E.D., Dänhardt, J., Morales, M.B., Rac, 
I., Špulerová, J., Schindler, S., Stavrinides, M., Targetti, S., Viaggi, D., Vogiatzakis, I.N., Guyomard, H., 2022. How can the European Common Agricultural 
Policy help halt biodiversity loss? Recommendations by over 300 experts. Conservation Letters, 2022, e12901.
Prach, K., Hobbs, R.J., 2008. Spontaneous succession versus technical reclamation in the restoration of disturbed sites. Restoration Ecology, 16, 363-366.
Prach, K., Jongepierová, I., Řehounková, K., 2013. Large-scale restoration of dry grasslands on ex-arable land using a regional seed mixture: establishment 
of target species. Restoration Ecology, 21, 33-39.
Prach, K., Jongepierová, I., Řehounková, K., Fajmon, K., 2014. Restoration of grasslands on ex-arable land using regional and commercial seed mixtures and 
spontaneous succession: successional trajectories and changes in species richness. Agriculture, Ecosystems and Environment, 182, 131-136.
Prach, K., Fajmon, K., Jongepierová, I., Řehounková, K., 2015. Landscape context in colonization of restored dry grasslands by target species. Applied 
Vegetation Science, 18, 181-189.
Prach, K., Fajmon, K., Řehounková, K., Jongepierová, I., 2021a. Hierarchy of environmental factors driving restoration of dry grasslands: a multi-site analysis. 
Applied Vegetation Science, 24, 202104.
Prach, K., Vítovcová, K., Řehounková, K., Královec, J., 2021b. Three decades of vegetation changes in a submontane grassland after the cessation of 
intensive fertilization. Preslia, 93, 169-179.
Prioritised action framework (PAF) for Natura 2000 in Slovenia pursuant to Article 8 of Council Directive 92/43/EEC on the conservation of natural habitats 
and of wild fauna and flora (the Habitats Directive) for the Multiannual Financial Framework period 2021 – 2027, 2019. Ministry of the Environment and 
Spatial Planning. https://natura2000.gov.si/fileadmin/user_upload/IP_PAF_Slovenia_final.pdf (17. 10. 2022).
Pykälä, J., 2004. Cattle grazing increases plant species richness of most species trait groups in mesic semi-natural grasslands. Plant Ecology, 175, 217-226.
Reheul, D., De Vliegher, A., Bommelé, L., Carlier, L., 2007. The comparison between temporary and permanent grassland. In: Proceedings of the 14th EGF 
Symposium on “Permanent and temporary grassland: plant, environment, economy”. Ghent, Belgium, September 3-5, pp. 1-13.
Reitalu, T., Purschke, O., Johansson, L.J., Hall, K., Sykes, M.T., Prentice, H.C., 2012. Responses of grassland species richness to local and landscape factors 
depend on spatial scale and habitat specialization. Journal of Vegetation Science, 23, 41-51.
Resch, M.C., Schütz, M., Buchmann, N., Frey, B., Graf, U., van der Putten, W.H., Zimmerman, S., Risch, A.C., 2021. Evaluating long-term success in grassland 
restoration: an ecosystem multifunctionality approach. Ecological Applications, 31, e02271.
Roeling, I.S., Ozinga, W.A., van Dijk, J., Eppinga, M.B., Wassen, M.J., 2018. Plant species occurrence patterns in Eurasian grasslands reflect adaptation to 
nutrient ratios. Oecologia, 186, 1055-1067.
Rūsiņa, S., Auniņš, A., Spuņģis, V., 2017. Chapter 13. 6230* Species-rich Nardus grasslands, on siliceous substrates in mountain areas. In: Rūsiņa, S. (ed.): 
Protected Habitat Management Guidelines for Latvia. Nature Conservation Agency, Sigulda, pp. 155-163.
Rydgren, K., Nordbakken, J.-F., Austad, I., Auestad, I., Heegaad, E., 2010. Recreating semi-natural grasslands: a comparison of four methods. Ecological 
Engineering, 36, 1672-1679.
Schelfhout, S., Mertens, S., Perring, M.P., Raman, M., Baeten, L., Demey, A., Reubens, B., Oosterlynck, S., Gibson-Roy, P., Verheyen, K., De Schrijver, A., 2017. 
P-removal for restoration of Nardus grasslands on former agricultural land: cutting traditions. Restoration Ecology, 25, S178-S187.
Schlesinger, W.H., 1997. Carbon balance in terrestrial detritus. Annual Review of Ecology and Systematics, 8, 51-81.
77
Acta Biologica Slovenica, 2023, 66 (1)
Sengl, P., Magnes, M., Erdős, L., Berg, C., 2017. A test of naturalness indicator values to evaluate success in grassland restoration. Community Ecology, 18, 184-192.
Silva, V., Catry, F.X., Fernandes, P.M., Rego, F.C., Paes, P., Nunes, L., Caperta, A.D., Sérgio, C., Bugalho, M.N., 2019. Effects of grazing on plant composition, 
conservation status and ecosystem services of Natura 2000 shrub-grassland habitat types. Biodiversity and Conservation, 28, 1205-1224.
Sojneková, M., Chytrý, M., 2015. From arable land to species-rich semi-natural grasslands: succession in abandoned fields in a dry region of Central Europe. 
Ecological Engineering, 77, 373-381.
Stanová, V., Šeffer, J., Ripka, J., 2007. Hodnotenie vegetácie a návrh monitoringu zjazdoviek lyžiarskeho strediska Veľká Rača. Daphne - Inštitút aplikovanej 
ekológie, Bratislava.
Suttie, J.M., Reynolds, S.G., Batello, C. (eds.), 2005. Grasslands of the World. Plant Production and Protection Series, No. 34. Food and Agricultural 
Organization of the United Nations, Rome.
Sutcliffe, L.M.E., Germany, M., Becker, U., Becker, T., 2016. How does size and isolation affect patches of steppe-like vegetation on slumping hills in 
Transylvania, Romania?. Biodiversity and Conservation, 25, 2275-2288.
Šumrada, T., Vreš, B., Čelik, T., Šilc, U., Rac, I., Udovč, A., Erjavec, E., 2021. Are result-based schemes a superior approach to the conservation of High 
Nature Value grasslands? Evidence from Slovenia. Land Use Policy, 111, 105749.
Šumrada, T., Japelj, A., Verbič, M., Erjavec, E., 2022. Farmers’ preferences for result-based schemes for grassland conservation in Slovenia. Journal for 
Nature Conservation, 66, 126143.
Tälle, M., Deák, B., Poschlod, P., Valkó, O., Westerberg, L., Milberg, P., 2018. Similar effects of different mowing frequencies on the conservation value of 
semi-natural grasslands in Europe. Biodiversity and Conservation, 27, 2451-2475.
Tóth, E., Deák, B., Valkó, O., Kelemen, A., Miglécz, T., Tóthmérész, B., Török, P., 2018. Livestock type is more crucial than grazing intensity: traditional cattle 
and sheep grazing in short-grass steppes. Land Degradation & Development, 29, 231-239.
Tölgyesi, C., Vadász, C., Kun, R., Csathó, A.I., Bátori, Z., Hábenczyus, A., Erdős, L., Török, P., 2022. Post-restoration grassland management overrides the 
effects of restoration methods in propagule-rich landscapes. Ecological Applications, 32, e02463.
Török, P., Vida, E., Deák, B., Lengyel, S., Tóthmérész, B., 2011. Grassland restoration on former croplands in Europe: an assessment of applicability of 
techniques and costs. Biodiversity and Conservation, 20, 2311-2332.
Török, P., Miglécz, T., Valkó, O., Kelemen, A., Tóth, K., Lengye,l S., Tóthmérész, B., 2012. Fast restoration of grassland vegetation by a combination of seed 
mixture sowing and low-diversity hay transfer. Ecological Engineering, 44, 133-138.
Török, P., Hölzel, N., van Diggelen, R., Tischew, S., 2016. Grazing in European open landscapes: how to reconcile sustainable land management and 
biodiversity conservation?. Agriculture, Ecosystems & Environment, 234, 1-4.
Török, P., Janišová, M., Kuzemko, A., Rūsiņa, S., Dajić Stevanovic, Z., 2017. Grasslands, their threats and management in Eastern Europe. In: Squires, V.R., 
Dengler, J., Hua, L., Feng, H. (eds.): Grasslands of the World: Diversity, Management and Conservation, 1st Edition. CRC Press, Boca Raton, FL, pp. 64-88.
Török, P., Dembicz, I., Dajić-Stevanović, Z., Kuzemko, A., 2020. Grasslands of Eastern Europe. In: Goldstein, M.I., DellaSala, D.A. (eds.): Encyclopedia of the 
World’s Biomes. Elsevier, Amsterdam, pp. 703-713.
Turtureanu, P.D., Palpurina, S., Becker, T., Dolnik, C., Ruprecht, E., Sutcliffe, L.M.E., Szabó, A., Dengler, J., 2014. Scale- and taxon-dependent biodiversity 
patterns of dry grassland vegetation in Transylvania. Agriculture, Ecosystems & Environment, 182, 15-24.
Valkó, O., Török, P., Matus, G., Tóthmérész, B., 2012. Is regular mowing the most appropriate and cost-effective management maintaining diversity and 
biomass of target forbs in mountain hay meadows?. Flora, 207, 303-309.
Valkó, O., Török, P., Deák, B., Tóthmérész, B., 2014. Review: prospects and limitations of prescribed burning as a management tool in European grasslands. 
Basic and Applied Ecology, 15, 26-33.
Valkó, O., Deák, B., Török, P., Tóth, K., Kiss, R., Kelemen, A., Miglécz, T., Sonkoly, J., Tóthmérész, B., 2021. Dynamics in vegetation and seed bank composition 
highlight the importance of post-restoration management in sown grasslands. Restoration Ecology, 29, e13192.
Van Daele, F., Wasof, S., Demey, A., Schelfhout, S., De Schrivjer, A., Baeten, L., van Ruijven, J., Mertens, J., Verheyen, K., 2017. Quantifying establishment 
limitations during the ecological restoration of species-rich Nardus grassland. Applied Vegetation Science, 20, 594-607.
Vasconcelos, H.L., Maravalhas, J.B., Cornelissen, T., 2017. Effects of fire disturbance on ant abundance and diversity: a global meta-analysis. Biodiversity 
and Conservation, 26, 177-188.
Vassiliev, K., Pedashenko, H., Nikolov, S.C., Apostolova, I., Dengler, J., 2011. Effect of land abandonment on the vegetation of upland semi-natural grasslands 
in the Western Balkan Mts., Bulgaria. Plant Biosystems, 145, 654-665.
Velev, N.I., Apostolova, I.I., 2008. Successional changes of Nardus stricta communities in the Central Balkan Range (Bulgaria). Phytologia Balcanica, 14, 75-84.
Wick, A.F., Geaumont, B.A., Sedivec, K., Hendrickson, J.R., 2016. Grassland degradation. In: Shroder, J.F., Sivanpillai, R. (eds.): Biological and Environmental 
Hazards, Risks, and Disasters. Elsevier, Amsterdam, pp. 257-276.
Wubs, E.R.J., van der Putten, W.H., Bosch, M., Bezemer, T.M., 2016. Soil inoculation steers restoration of terrestrial ecosystems. Nature Plants, 2, 16107.
Zaitsev, A.S., Gongalsky, K.B., Korobushkin, D.I., Butenko, K.O., Gorshkova, I.A., Rakhleeva, A.A., Saifutdinov, R.A., Kostina, N.V., Shakhab, S.V., Yazrikova, 
T.E., 2017. Reduced functionality of soil food webs in burnt boreal forests: a case study in Central Russia. Contemporary Problems of Ecology, 10, 277-285.
Zarzycki, J., Korzeniak, J., Perzanowska, J., 2022. Impact of land use changes on the diversity and conservation status of the vegetation of mountain 
grasslands (Polish Carpathians). Land, 11, 252.