20/1 • 2021, 177–188
DOI: 10.2478/hacq-2020-0017
An example of fast old field succession in
a traditionally managed rural landscape
on the Slovenian Karst
Andraž Čarni1, 2,* , Zita Zimmermann3, 4 , Nina Juvan5 , Andrej Paušič6 ,
Gábor Szabó4 & Sándor Bartha3, 4
Key words: ex-arable land, grassland
restoration, land use legacy,
mowing, secondary succession,
spatial heterogeneity, species pool,
target species.
Ključne besede: ciljne vrste, košnja,
obnova travišč, opuščene površine,
prostorska heterogenost, sekundarna
sukcesija, sledi pretekle rabe
zemljišč, zaloga vrst.
Nomenclature of plant species
follow Martinčič (2007)
Received: 14. 3. 2020
Revision received: 25. 6. 2020
Accepted: 21. 7. 2020
Co-ordinating Editor:
Orsolya Valkó
Abstract
We report an exceptionally fast grassland recovery process from a karst plateau
in SW Slovenia. Vegetation of old fields with different ages was sampled using
a chronosequence of fields abandoned 1, 3, 6, 9, 13, 15 and 100 years ago. We
prepared dendrogram dividing the data set into 9 clusters that were further
analyzed: diagnostic species, ecological conditions and life forms were evaluated.
The initial stage of succession was characterized by segetal weeds and indicated
high levels of soil nutrients. The second stage was dominated by dense patches of
perennial forbs (most of them ruderal species) preferring also high levels of soil
nutrients. The third stage was dominated by caespitose hemicryptophyte grasses,
many of them of sub-Mediterranean origin. The first two stages took 13 years
and both could be considered as early successional stages developing on nutrient
rich soils. These stages were switched to late successional stage characterized by
seminatural grassland species. The quick succession can probably be attributed to
the rich species pool of natural grassland flora, the small size and annual mowing
of abandoned agricultural fields and the close proximity of seed sources.
Izvleček
Prispevek obravnava hitro zaraščanje opuščenih kmetijskih površin na kraški
planoti v jugovzhodni Sloveniji. Na podlagi kronosekvece smo vzorčili površine,
ki so bile opuščene pred 1, 3, 6, 9, 13, 15 in 100 leti. Na podlagi dendrograma
smo pridobljene podatke razdelili v 9 snopov, ki smo jih nadalje analizirali:
diagnostične vrste, ekološke razmere in življenjske oblike. Za prvi stadij zaraščanja
(sukcesije) je značilno pojavljanje plevelnih vrst in visoka vsebnost hranil v tleh.
V drugem stadiju prevladujejo goste zaplate trajnih zelišč (večinoma ruderalnih
vrst), ki dobro uspevajo na tleh z visoko vsebnostjo hranil. V tretjem stadiju
prevladujejo rušnate, trajne trave, od katerih mnoge izvirajo iz submediteranskega
območja. Prva dva stadija trajata 13 let in jih lahko oba obravnavamo kot zgodnja
sukcesijska, ki se razvijeta na s hranili bogatih tleh. Potem se razvije pozen
sukcesijski stadij, kjer prevladujejo travnate vrste. Hiter potek sukcesije povzročijo:
velika zaloga vrst polnaravnih travišč, majhna površina opuščenih površin,
vsakoletna košnja in neposredna bližina virov semen.
1 Research Centre of the Slovenian Academy of Sciences and Arts, Institute of Biology, Novi trg 2, SI-1001 Ljubljana, Slovenia
2 University of Nova Gorica, Faculty for Viticulture and Enology, Vipavska 13, Rožna Dolina, SI-5000 Nova Gorica, Slovenia
3 Centre for Ecological Research, GINOP Sustainable Ecosystems Group, H-8237 Tihany, Klebelsberg str. 3., H-8237 Hungary
4 Centre for Ecological Research, Institute of Ecology and Botany, Alkotmány str. 2-4., H-2163 Vácrátót, Hungary
5 Research Centre of the Slovenian Academy of Sciences and Arts, Anton Melik Geographical Institute, Novi trg 2, SI-1000 Ljubljana, Slovenia
6 University of Maribor, Faculty of Agriculture and Life Sciences, Pivola 10, SI-2311 Hoče, Slovenia
* Corresponding author. E-mail: carni@zrc-sazu.si
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Introduction
Old field succession is a process of natural regeneration
of vegetation after the cessation of cultivation (Cramer et
al. 2008). Secondary grasslands recovering after the abandonment of agricultural fields provide habitats for many
grassland species and partially compensate for the general
loss of grasslands. However, the rate of spontaneous old
field succession might vary greatly between habitats due
to various factors (Pywell et al. 2002, Walker et al. 2004,
Török et al. 2011a, Halassy et al. 2016). In early stages of
succession, high concentrations of soil nutrients (as legacy
from cultivation) and the dominance of invasive aliens are
considered as major factors limiting spontaneous regeneration (Cramer & Hobbs 2007, Cramer et al. 2008). In
middle stages of succession, several species (both native
and invasive alien) form nearly monodominant patches
suppressing other species and preventing the colonization
of native species (Bartha et al. 2003, 2014, Házi et al.
2011, Szentes et al. 2012). Spontaneous succession might
be limited also by the lack of available propagules (Donath et al. 2003, Öster et al. 2009, Halassy et al. 2016).
Patterns learnt about vegetation development in old
field succession (Pickett et al. 1987, Osbornová et al. 1989,
Cramer & Hobbs 2007) contributed to ecological theory
and supported restoration practice (Török & Helm 2017).
Case studies describing patterns of spontaneous successions
in different habitats provide indispensable background for
ecological restoration (Prach et al. 1999, 2001). Despite to
the large number of related studies (for bibliography see
Rejmánek & van Katwyk 2005, http://botanika.bf.jcu.cz/
suspa/pdf/BiblioOF.pdf ), generalizations about old field
succession are still limited due to the paucity of comparable
data (Prach & Walker 2019). Following recommendations
and the related framework of Prach & Walker (2019),
we intend to collect data comparable to other studies.
Therefore, we have estimated the ecological circumstances
by Ellenberg bioindicator values and assessed the presence
of invasive alien species and the spatial heterogeneity of
dominant species. We have also evaluated the success of
succession (i.e. the time required for recovering some
desirable target community with natural or semi-natural
species composition) and the temporal pattern of species
richness. For assessing the patchwork of midsuccessional
dominants (Bartha et al. 2014) we have sampled long
transects that able to cross several vegetation patches
within a particular stand estimating their diversity and
heterogeneity (Bartha et al. 2004).
Cultivated fields have been abandoned due to various
reasons (e.g. droughts, natural disasters, political and
economic crises or agricultural mismanagement). These
problems usually affect large areas, therefore abandoned
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Andraž Čarni, Zita Zimmermann, Nina Juvan, Andrej Paušič, Gábor Szabó &
Sándor Bartha
An example of fast old field succession in a traditionally managed rural landscape
on the Slovenian Karst
fields are typically aggregated in space and time. Therefore, in related studies the abandoned fields occupied
large part of the landscape after some drastic collapse of
local economy followed by drastic change in land use (e.g.
Osbornová et al. 1989, Molnár & Botta-Dukát 1998,
Palang et al. 2006, Csecserits et al. 2007). In contrast,
information about old field succession in traditionally
managed landscape with stable land use is still lacking.
In this study, we intend to fill this gap. Our study area is
located on a Karst plateau in SW Slovenia near the coast
of Adriatic Sea. This is a traditional rural landscape with
continuous low intensity agriculture since the Roman Era
(Vitasović et al. 2012, Kaligarič & Ivanjšič 2014, Batalha
et al. 2015, Breg Valjavec et al. 2018). Fields appropriate for cultivation are small and are surrounded by seminatural forests and grasslands. Field abandonment is rare.
Ex-arable fields are small and they are utilized as pastures
or meadows.
The aim of the research was to explore some characteristics of secondary succession in this specific traditional
rural landscape. We explored temporal and spatial differentiation of vegetation and described patterns of life
forms and ecological indicator values. We addressed the
following questions: How do diagnostic species, ecological conditions, life-form spectrum and spatial heterogeneity change along the succession? How long does the
development of semi-natural grasslands take after land
abandonment? How does our research fit with similar research in this field?
Materials and methods
Study site
The study area is located on a karst plateau in the northwest edge of the Dinaric Alps, Slovenia, in the area that
is under the influence of the Adriatic Sea. The climate is
sub-Mediterranean (Köppen-Type Cfa) with average January temperatures above 0 oC and annual precipitation
between 1400 and 1500 mm. The karst plateau consists
of karstified Mesozoic limestone, covered predominantly
by rendzinas and cambisols (Vrščaj et al. 2017). The zonal vegetation is forest, dominated by Ostrya carpinifolia
and Quercus pubescens (Čarni et al. 2002, 2009). After
intensive deforestation that occurred centuries ago (Gams
1993), karstic pastures (Carex humilis-Centaurea rupestris
community) and karstic meadows (Danthonia calycinaScorzonera villosa community) were established (Kaligarič
1997, Kaligarič et al. 2006). As a consequence of largescale socio-economic changes, afforestation began and
grasslands declined from 82 to 20% and forests pro-
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gressed from 17 to 73% in the last 250 years (Kaligarič &
Ivanjšič 2014).
The cultivated surfaces are mainly located in “cultural
dolinas”, i.e. in enclosed karstic depressions that are filled
with soil (Breg Valjavec et al. 2018) and only rarely appear on flat surfaces. Such a flat area was found above
the village Podpeč near Črni Kal (45.536° N, 13.900° E,
elevation 385 m a.s.l.), where we could find fields of different age after abandonment. These areas are still mown
once a year and occasionally grazed (Škornik et al. 2010,
Pipenbaher et al. 2011).
Sampling of vegetation
Using the method of improved space for time substitution (Pickett 1989, Molnár & Botta-Dukát 1998, Csecserits et al. 2007), we built a chronosequence representing a succession series in the region (Čarni et al. 2007).
Abandoned fields were selected close to each other in
order to have the same regional species pool and habitat
conditions. We sampled in the area of Podpeč near Črni
Kal, where we can find nearly all stages of succession on
a small area of 5 ha, only one plot (the 1-year-old-stand)
was 3 km apart. The size of individual abandoned fields
was small, ranged between 0.03 and 1 ha. We could find
fields abandoned 1, 3, 6, 9, 13, 15 and 100 years ago.
Age of the fields was determined using a time series of
aerial photos and information from local people. Specific
transect sampling was used for assessing within-stand heterogeneity of vegetation (Bartha et al. 2004, Supplement
S, Figure 1). In each field, the presence of rooted vascular plant species was recorded in 5 cm × 5 cm contiguous
microquadrats arranged along a 52 m long belt transect.
Transects were positioned in the middle of the individual
field to avoid the edge effect. Vegetation patchwork (spatially heterogeneous vegetation mosaic) is typical to these
transitional vegetation types (Pickett et al. 2001, Bartha
et al. 2004, Szentes et al. 2012). The 52 m long transects
were long enough to estimate properly the related spatial
heterogeneity at each field. Base-line transect data were
resampled by computer (with the method of computerized sampling, Podani 1987, Bartha et al. 2004). During
computerized resampling, each transect was subdivided
into 1 m long segments and species abundances were
calculated in each segment (by summarizing presences
of species in the particular segment) (for details, see S,
Figure 1). Based on previous analyses of spatial patterns
in succession (Bartha et al. 2004, Bartha 2007, Ruprecht
et al. 2007, Szentes et al. 2012), we chose the 1 m scale
to provide relatively homogeneous segments for further
analyses. The transects were sampled in 2012 in late May
– early June during the optimal growing period of veg-
Andraž Čarni, Zita Zimmermann, Nina Juvan, Andrej Paušič, Gábor Szabó &
Sándor Bartha
An example of fast old field succession in a traditionally managed rural landscape
on the Slovenian Karst
etation, before the fields were mown. The high resolution transect-based sampling design was chosen because
it was effective in several other old field studies (Bartha et
al. 2004, Jongepierová et al. 2004, Ruprecht et al. 2007,
Szentes et al. 2012).
Data analysis
Spatiotemporal differentiation of species composition
at the scale of 1 m long segments was analysed by classification and ordination of the full data set. For finding the most appropriate classification method (distance
measures between samples and method for construction
of clusters), the OptimClass program was applied (Tichý
et al. 2010). Classification was then carried out by the
PC-ORD software (McCune & Mefford 1999), run in
the JUICE 7.0 program. Non-metric multidimensional
scaling (NMDS) was also performed by the vegan program package run in R environment (Oksanen et al. 2017,
R Core Team 2018). Diagnostic species of each cluster
were determined in JUICE 7.0 program by calculating
the fidelity, using the phi-coefficient as fidelity measure
on basis of presence of species within and beyond each
cluster (Sokal & Rohlf 1995). In these calculations, each
cluster was compared with the other samples in the data
set, which were taken as single, undivided group. Species
with phi > 0.4 were considered as diagnostic for individual clusters, but species whose occurrence concentration
in the plots of a particular cluster was not significant at
p < 0.05 (Fisher´s exact test) were excluded (Chytrý et al.
2002, Tichý & Chytrý 2006).
Patterns of life forms and ecological indicator values
(EIV, called also Ellenberg bioindicator values) were
assessed according to Pignatti et al. (2005). Dry grasslands specialist species have been determined by a local
monograph (Kaligarič 1997) and a wider European list
(Mucina et al. 2016), all species considered as diagnostic for Festuco-Brometea dry grasslands were considered
as target species. Soil nutrient content was estimated by
abundance-weighted mean ecological indicator values at
the scale of 1 m segments of transects. Ecological indicator values were tested for significant differences between
coenostate clusters (vegetation patch types) found by classification. For this analysis we used Zelený’s methods (Zelený 2018) to account for potential biases.
We tested temporal trends in succession using years
since abandonment as predictor variable. We used nitrogen indicator values, number of all species and number of
specialist species as response variables. Years since abandonment was fitted with log transformation. We used a
linear model with polynomial terms in case of nitrogen
indicator values and number of species, while we used
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GLM ('glm' function with Gaussian family) in case of
specialist species. We fitted the models with polynomials
because of the non-linearity of our variables. We chose
cubic model because it had lower AIC values and higher
R2 values than a linear model. Figures were created using
'ggplot2' package in R (Wickham 2016).
Variables with different age (years since abandonment)
were compared with Kruskal–Wallis-H-test, followed by
Dunn-test as post hoc test. Significant differences between years were plotted with 'FSA' (Ogle 2018) and
'rcompanion' (Mangiafico 2000) packages in R.
Results
Classification of plots revealed nine clusters representing
different vegetation patch types, i.e. plots with similar
species composition (Appendix, Figure 2). Each cluster
had specific dominant and diagnostic species (Table 1).
NMDS ordination showed further structure: three main
distinct groups with overlapping subgroups (Figure 1).
The main groups represented temporal differentiation in
Andraž Čarni, Zita Zimmermann, Nina Juvan, Andrej Paušič, Gábor Szabó &
Sándor Bartha
An example of fast old field succession in a traditionally managed rural landscape
on the Slovenian Karst
vegetation (three stages in succession) while clusters within the main groups represented spatial heterogeneity of
stands. High compositional similarity was found between
clusters of 15 years old and 100 years old fields (clusters 8
and 9). Plantago major, Trifolium repens and weeds typical
to cultivated fields dominated in the first main group of
clusters (clusters 1–2): Cirsium arvense and many annual
species (e.g. Anagallis arvensis, Cerastium brachypetalum,
Chenopodium album) (Table 2). Within this stage, some
small differences were found between two clusters: with
more Chenopodium album in cluster 1 and more Cynodon dactylon in cluster 2. The second main group of clusters (clusters 3–7) represented the period in succession
with fields of ages between 2 and 13 years. These fields
had ruderal weeds (e.g. Bromus sterilis, Pastinaca sativa,
Melilotus officinalis, Elymus repens, Taraxacum officinale)
and species typical of mesic and eutrophic meadows (e.g.
Dactylis glomerata, Festuca pratensis, Verbena officinalis,
Leucanthemum liburnicum, Brachypodium rupestre). The
terminal main group (clusters 8 and 9) was characterized
by Bromus erectus, Chrysopogon gryllus, Scorzonera villosa,
Thymus longicaulis, Festuca rupicola and many other spe-
Figure 1: Bray-Curtis based non-metric multidimensional scaling (NMDS) plot of all samples. NMDS plot (stress 0.14) shows that samples were divided into three main groups: initial (clusters 1 and 2), early successional (clusters 3, 4, 5, 6 and 7) and late successional main group (clusters 8 and 9).
Slika 1: Nemetrično multidimenzionalno lestvičenje (NMDS) vseh vzorčnih ploskev na podlagi Bray-Curtisove podobnosti (stres 0,14) kaže, da so
vzorci razdeljeni v tri glavne skupine: inicialno (snopa 1 in 2), zgodnje sukcesijsko (snopi 3, 4, 5, 6 in 7) in pozno skucesijsko skupino (snopa 8 in 9).
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Andraž Čarni, Zita Zimmermann, Nina Juvan, Andrej Paušič, Gábor Szabó &
Sándor Bartha
An example of fast old field succession in a traditionally managed rural landscape
on the Slovenian Karst
Table 1: Diagnostic and dominant species of different successional stages according to nine clusters (coenostate clusters). Species
with phi > 0.4 were considered as diagnostic (associated) for individual clusters. Among them species with >50% abundance in the
1 m quadrats were considered as dominant.
Tabela 1: Diagnostične in dominantne vrste različnih sukcesijskih stadijev glede na devet snopov (cenostatski snopi). Vrste z
phi > 40 smo opredelili kot pridružene (diagnostične) za posamezen snop. Med njimi smo vrste z abundanco večjo od 50% na
ploskvah 1 m smatrali kot dominantne.
Successional Cluster Age of
stage
old fields
1
year 1
Initial stage
Early stage
Terminal
stage
Diagnostic species
Dominant species
Anagallis arvensis, Chenopodium album, Cirsium
arvense, Crepis taraxacoides, Plantago major,
Polygonum aviculare, Veronica persica
Cerastium brachypetalum, Cynodon dactylon,
Lolium perenne, Plantago major, Poa annua,
Trifolium repens
Lotus corniculatus, Medicago sativa, Poa bulbosa,
Poa sylvatica
Anagallis arvensis, Chenopodium album, Cirsium
arvense, Plantago major, Polygonum aviculare
2
year 1
3
year 6
year 9
year 13
4
year 9
Pastinaca sativa, Trifolium pratense
5
year 3
year 13
Festuca pratensis, Medicago lupulina,
Setaria pumila
6
year 3
Filipendula hexapetala, Leucanthemum
platylepis, Melilotus officinalis, Setaria pumila
7
year 3
year 13
8
year 15
year 100
9
year 15
year 100
Cerastium brachypetalum, Convolvulus arvensis,
Cynodon dactylon, Lolium perenne, Plantago major,
Poa annua, Taraxacum officinale, Trifolium repens
Brachypodium rupestre, Convolvulus arvensis,
Dactylis glomerata, Elytrigia repens, Lotus corniculatus, Medicago sativa, Pastinaca sativa, Plantago
lanceolata, Poa sylvicola, Taraxacum officinale,
Trifolium pratense
Convolvulus arvensis, Elytrigia repens, Linaria
vulgaris, Medicago lupulina, Medicago sativa,
Pastinaca sativa, Plantago lanceolata, Taraxacum
officinale, Trifolium pratense
Convolvulus arvensis, Elytrigia repens, Medicago
lupulina, Plantago lanceolata, Setaria pumila,
Taraxacum officinale, Trifolium pratense
Brachypodium rupestre, Convolvulus arvensis,
Dactylis glomerata, Elytrigia repens, Filipendula hexapetala, Melilotus officinalis, Plantago lanceolata,
Poa sylvicola, Setaria pumila, Taraxacum officinale
Brachypodium rupestre, Dactylis glomerata, Elytrigia repens, Medicago lupulina, Medicago sativa,
Plantago lanceolata, Trifolium pratense
Betonica serotina, Brachypodium rupestre,
Bromopsis erecta, Plantago media, Scorzonera
villosa, Thymus longicaulis
Betonica serotina, Bromopsis erecta, Festuca
rupicola, Filipendula hexapetala, Galium
lucidum, Salvia pratensis, Scorzonera villosa,
Thymus longicaulis
Anthyllis vulneraria, Bromopsis erecta, Carex flac- Anthyllis vulneraria, Bromopsis erecta, Chrysopogon
ca, Carex montana, Chrysopogon gryllus, Festuca gryllus, Festuca rupicola, Rhinanthus major,
rupicola, Hippocrepis comosa, Onobrychis tom- Scorzonera villosa, Thymus longicaulis
masiniana, Potentilla tommasiniana, Rhinanthus
major, Scorzonera villosa, Thymus longicaulis
cies typical for dry and oligotrophic grasslands. Clusters
represented vegetation differentiation both in space (spatial heterogeneity) and time (asynchrony). Spatial heterogeneity was obvious as different clusters appeared within a
particular field. Some clusters appeared in different fields,
i.e. these vegetation patch types were present over several
years (Table 2).
Table 2: Number of individual plots of each age in clusters
divided according to OPTIMCLASS proposal.
Tabela 2: Število posameznih ploskev določenega časovnega
obdobja razdeljenih v snope s postopkom OPTIMCLASS.
Cluster
1
2
3
4
5
6
7
8
9
A
1
41
11
3
6
52
36
14
2
g
9
15
37
e
13
15 100
6
2
44
51
1
3
49
Dominant age
1
1
6
9
3
3
13
15
100
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Andraž Čarni, Zita Zimmermann, Nina Juvan, Andrej Paušič, Gábor Szabó &
Sándor Bartha
An example of fast old field succession in a traditionally managed rural landscape
on the Slovenian Karst
Figure 2: Changes of the main life form
categories (annuals, perennial forbs,
perennial graminoids and woody species)
during succession.
Slika 2: Spremembe glavnih življenskih
oblik (enoletnice, trajna zelišča, trajne
trave in lesnate vrste) med sukcesijo.
Perennial forbs dominated the first stages in succession
then perennial grasses became dominant from the 13th
year (Figure 2). Annuals were abundant only in the initial
stage. Woody species (Pinus nigra, Prunus mahaleb and
Rosa gallica) appeared as early as in the 3rd year of succession with ca. 3% relative abundance. However, abundance of woody species decreased and remained negligible in subsequent years due to annual mowing.
Ecological indicator values for nutrients showed nonlinear decrease over time (Figure 3, Appendix, Table 3)
Figure 3: Relationship between nitrogen
indicator values and years since abandonment in old fields in SW Slovenia (p <
2.2e-16, adj R2 = 0.572, y = 5.56 + 0.55x
– 0.36x2 – 0.03x3). The 95% confidence
level interval for predictions from the
linear model is marked with gray envelop.
Individual points represent nitrogen
indicator values estimated in 1 m long
segments. Note the strong spatial heterogeneity represented by variability within
particular years.
Slika 3: Odnos med indikatorsko
vrednostjo za dušik in leti od opustitve
starih polj v JZ Sloveniji (p < 2,2e-16,
adj R2=0,572, y = 5.56 + 0.55x – 0,36x2
– 0,03x3). Interval zaupanja (95%) za
napovedi z lineranim modelom je prikazan s sivo. Posamezne točke predstavljajo
indikatorske vrednosti za dušik ocenjene
na 1 m odsekih. Opazna je močna
prostrska heterogenost zaradi variabilnosti
med posameznimi leti.
182
indicating mesotrophic conditions in early succession and
oligotrophic conditions in the 15-year-old and 100-yearold stands. In our study, the ecological indicator values
for nutrients were higher in initial and early successional
stages than in the 15-year-old and the 100-year-old stands
(Appendix, Figure 3a). Total species richness reached a local maximum early in succession in the 3rd year and the
fitted cubic relationship predicted a minimum between
the 15 and 100-years (Figure 4a, Appendix, Table 3). Due
to the high within-stand variation at 1 m plot scales (cf.
20/1 • 2021, 177–188
A
Andraž Čarni, Zita Zimmermann, Nina Juvan, Andrej Paušič, Gábor Szabó &
Sándor Bartha
An example of fast old field succession in a traditionally managed rural landscape
on the Slovenian Karst
B
Figure 4: Relationship (A) between the number of all species and years since abandonment (p < 2.2e-16, adj R2 = 0.22, y = 7.63 + 4.77x – 1.89x2 +
0.17x3) and (B) between the number of specialist species and years since abandonment (p < 2.2e-16, adj R2 = 0.616, y = 0.28 – 1.06x + 0.79x2
– 0.06x3) in old fields in SW Slovenia. The 95% confidence level interval for predictions from the linear model is marked with gray envelop.
Individual points represent species richness estimated in 1 m long segments. Note the strong spatial heterogeneity represented by variability within
particular years.
Slika 4: Odnos med (A) številom vseh vrst in leti od opustitve (p < 2,2e-16, adj R2=0,22, y = 7,63 + 4,77x – 1,89x2 + 0,17x3) in (B) številom
specialistov in leti od opustitve (p < 2,2e-16, adj R2=0,616, y = 0,28 – 1,06x + 0,79x2 - 0,06x3) starih polj v JZ Sloveniji. ). Interval zaupanja (95%)
za napovedi z lineranim modelom je prikazan s sivo. Posamezne točke predstavljajo indikatorske vrednosti za dušik ocenjene na 1 m odsekih.
Opazna je močna prostrska heterogenost zaradi variabilnosti med posameznimi leti.
Figure 4a), the total number of species did not differ between the initial and early successional (years 1–13) and
the other stands (year 15 and year 100) (Appendix, Figure 3b). Dry grassland specialist species were present in
all stands including the initial stage (Appendix, Table 1)
with increasing number of specialists along the successional gradient (Figure 4b). Number of grassland specialist species at 1 m plot scale did not differ between the
15-year-old and the 100-year-old stands. However, these
stands had higher number of specialists than earlier stages
(Appendix, Figure 3c).
Besides overall trends of temporal vegetation differentiation, using the method suggested by Zelený we
found significant spatial heterogeneity of ecological indicator values within-fields (between different clusters;
F = 227.965, adjusted p < 0.05).
Discussion
Fast and successful secondary
succession
In our study we found an example of fast and successful
secondary succession on abandoned agricultural fields in
a traditional rural landscape. Species richness and composition become similar to target grasslands on a field that
was abandoned only 15 years ago.
According to general models of succession (Cramer &
Hobbs 2007) and previous studies in the region (Čarni
& Kaligarič 1991, Kaligarič 1997, Kaligarič et al. 2006,
Kaligarič & Ivanjšič 2014) local spontaneous succession
should terminate in forest after ca. 50 years. Successional
stages on unmanaged abandoned fields consist of ruderal
communities, mesic perennial grasslands and shrublands
with increasing cover of woody components. In contrast,
annual mowing may alter the successional pathway to
the direction of dry semi-natural grasslands (Čarni &
Kaligarič 1991, Kaligarič & Ivanjšič 2014). This type of
dry semi-natural grasslands (dominated by Bromus erectus and Chrysopogon gryllus) can be considered as target
vegetation in our study area (Čarni & Kaligarič 1991,
Kaligarič & Ivanjšič 2014). We found low abundance of
woody species. Although some seedlings and saplings of
woody species appeared early after 3 years, their colonization remained unsuccessful with negligible contribution
of woody species in subsequent years.
After 15 years, old fields could develop into semi-natural grasslands in our study area. Note that spontaneous
succession would reach forest stage only after 50 years
(Čarni & Kaligarič 1991, Kaligarič 1997). In contrast
to other studies that reported the lack or minor development of weedy stages in fast successions (Jongepierová et
al. 2004, Török et al. 2011b), we found distinct early successional ruderal stages. These early stages with the dominance of ruderal species reflect land use legacy, i.e. the
183
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presence of specific seed bank composition and increased
nutrient content of soil (Cramer et al. 2008, Halassy et
al. 2016, Török et al. 2018). In our study, the ecological indicator values for nutrients were higher in initial
and early successional stages than in the 15-year-old and
the 100-year-old stands. In vegetation patches (clusters)
dominated by ruderal species, the mean Ellenberg N indicator values varied between 4 and 6 indicating medium
levels of nutrients. This medium level of soil nutrients
most probably is a legacy of the cultivation. Our results
suggest that nutrient stocks have been depleted after ca.
15 years. Annual mowing probably enhanced this process.
Invasive alien species often dominate early stages in old
field succession and might slow down or prevent further
regeneration (Prach & Pyšek 1999, Matus et al. 2003,
Csecserits et al. 2011, Bartha et al. 2014). It is important
to note that invasive alien species did not appear in this
succession. The lack of invasive aliens is probably typical
in landscapes with traditional extensive land use and with
high cover of natural and semi-natural vegetation types
(Jongepierová et al. 2004, Ruprecht 2006).
Rate of succession is often limited by the lack of available propagules (Molnár & Botta-Dukát 1998, Pywell et
al. 2002, Török et al. 2011a) and diversity of developing semi-natural grasslands is under the influence of surrounding habitats (Janišová et al. 2014). In our study, the
surrounding species pool was rich in mesic and xeric grassland species. Hedgerows, dry semi-natural grasslands, forests and shrublands providing these species were present
in the close vicinity (within 50 m) or often adjacent to
abandoned fields. As a consequence, the total number of
species increased fast in succession and reached maximum
already in the third year. Specialists appeared in the first
year and their number increased continuously along the
successional gradient. Therefore, we can conclude that
dispersal limitation had minor role here.
Species of mesic grasslands also colonized and some of
them became abundant in the early stage of succession (e.g.
Dactylis glomerata, Festuca pratensis). However, after 15
years, these species were replaced by dry grassland species.
Temporal patterns of life forms in secondary succession
at the karst plateau of SW Slovenia are similar to patterns
described in other studies from Central Europe (Prach et
al. 2007). Succession starts with the dominance of perennial forbs then perennial grasses become dominant after
ca. 10 years. This early period is very similar to spontaneous succession in other mesic habitats (Prach et al. 2007,
2014). However, annual mowing modifies the subsequent
patterns preventing the development of woody stages and
promoting the emergence of semi-natural dry grasslands.
Contribution of perennial forbs is high in all stages while
annuals are abundant only in the initial stage.
184
Andraž Čarni, Zita Zimmermann, Nina Juvan, Andrej Paušič, Gábor Szabó &
Sándor Bartha
An example of fast old field succession in a traditionally managed rural landscape
on the Slovenian Karst
Considering potential mechanisms, we suggest that
initial and early stages of succession are driven by the
moderately increased level of soil nutrients (a minor agricultural legacy effect). The subsequent fast succession to
target semi-natural dry grasslands can be explained by the
interaction of rich species pool, the good condition for
dispersal and the appropriate disturbance regime (annual
mowing). The small size of fields, the low ratio of ruderal
elements in the surrounding landscape and the lack of
invasive aliens contribute also to the success of succession.
Tölgyesi et al. (2019) proposed an alternative explanation
that the rate of recovery might depend on the plot-scale
functional redundancy of the target community, as redundant species come back slower due to a self-organized
establishment limitation. Species density is relatively low
in our case in the target grasslands in this area. Low species density implies low functional redundancy. Therefore, community assembly can proceed quickly and unhindered by internal establishment limitation.
Succession without middle stage?
In this study, we found evidence that early successional
stages can develop quickly to late successional grasslands, i.e. the expected midsuccessional stage(s) described by other studies (Čarni & Kaligarič 1991, Bartha
et al. 2014, Prach et al. 2014, Sojneková & Chytrý 2015,
Schmid et al. 2017) did not appear. The missing midsuccessional stage is an interesting aspect of this successional series (see also Jongepierová et al. 2004, Török et al.
2011b). One may argue against our interpretation saying
that our target community is not a terminal stage but it
is the “missing midsuccessional” stage due to deflected
(or arrested) succession (sensu Godwin 1929). However,
we must reject this alternative. First, because no “later
successional” grasslands are known from the region
(Kaligarič et al. 2006, Vitasović et al. 2012, Kaligarič &
Ivanjšič 2014). Second, because the fine-scale community organization of these fields (the 15 years old and the
100 years old stands in our study) is similar to natural
and semi-natural grasslands but differ from typical midsuccessional grasslands. In midsuccessional grasslands
low diversity patches are characteristic, formed by species
that are subordinate in the terminal community (Bartha
et al. 2014). These species become locally and temporally
dominant due to stochastic priority effects. The related
vegetation structure differs considerably from the structure of terminal community (Bartha 2007) and it arrests
or delays colonization of other target species (Prach &
Pyšek 1999, Házi et al. 2011, Szentes et al. 2012, Bartha
et al. 2014). The grasslands developed after 15 years in
secondary succession at the Karst plateau of SW Slovenia
20/1 • 2021, 177–188
did not show these features. Bromus erectus and Chrysopogon gryllus formed vegetation matrix (i.e. the same
dominant species as in target semi-natural grasslands),
and this matrix did not inhibit the colonization and persistence of other dry grassland species.
Spatial heterogeneity in succession
We identified nine vegetation patch types (called ‘coenostate clusters’) in this study. Patches occurring in early
succession were dominated by ruderal and mesic grassland
species (mainly legumes and grasses, e.g. Lotus corniculatus, Taraxacum officinale, Dactylis glomerata and Elymus
repens). On the 15-year-old abandoned field we found
that perennial grasses of target community (Bromus erectus, Chrysopogon gryllus) became dominant and formed
additional patch types. To some degree, these structures
were repetitive in space and time as we could find indicator species with significant associations (i.e. preference) to
these patch types. Our results are consistent with previous
studies reporting spatiotemporal contingencies of vegetation patterns in old field succession (Pickett et al. 2001).
At each field (with one exception) two or three patch
types were present and formed spatially heterogeneous
vegetation mosaic. Patch types (coenostate clusters) that
co-occurred on a field often had significant differences in
ecological indicator values or in species richness. These
patterns suggest the existence of fine-scale environmental
and functional heterogeneity within-stands. Spatial heterogeneity of soil characteristics (Robertson et al. 1988),
biomass (Symonides 1985) and vegetation structure (Symonides 1985, Lepš 1989, Ruprecht et al. 2007) has been
reported in previous old field studies. However, the number of related studies is limited and generalizations about
spatial patterns in succession (similar to generalizations
made about temporal patterns) are still missing.
Our results highlight the potential role of spatial patterns in controlling the rate and direction of spontaneous
regeneration processes (McCallum et al. 2018), and call
for further studies on fine scale spatial heterogeneity in
succession.
Conclusion
We report an example of fast vegetation succession on
abandoned agricultural fields from a traditionally managed “living rural landscape” where land abandonment is
a rare event and restricted to small areas. We found here
initial and early successional stages similar to other series.
However, we did not found typical midsuccessional stage.
In contrast, we found a 15-year-old stand distinct from
younger abandoned fields but similar to late-successional
Andraž Čarni, Zita Zimmermann, Nina Juvan, Andrej Paušič, Gábor Szabó &
Sándor Bartha
An example of fast old field succession in a traditionally managed rural landscape
on the Slovenian Karst
fields (represented by a 100-year-old stand in our data).
Our study provides evidence that dry semi-natural grasslands can regenerate with high rate within 15 years in this
specific rural landscape. Multiple factors (low intensity
agriculture with low fertilizer input, small size of agricultural fields, close proximity of seed sources and the annual
mowing of ex-arable lands) are probably generating and
controlling this specific pattern of fast and successful old
field succession. Assessing the generality of these patterns
needs further studies based on permanent plots and experiments with monitoring both vegetation characteristics and environmental factors.
Acknowledgements
We owe thanks to Iztok Sajko who kindly prepared the
graphics. The research was financially supported by Slovenian Research Agency (ARRS P1-0236), the bilateral
project between Hungarian Academy of Sciences and
Slovenian Academy of Sciences and Arts and by the GINOP-2.3.2-15-2016-00019 project.
Conflict of interest
The authors declare that there is no conflict of interests
regarding the publication of the manuscript.
Andraž Čarni , https://orcid.org/0000-0002-8909-4298
Zita Zimmermann , https://orcid.org/0000-0001-8592-3925
Andrej Paušič , https://orcid.org/0000-0003-3457-7097
Sándor Bartha , https://orcid.org/0000-0001-6331-7521
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