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IZVORNI I ZNANSTVENI ČLANCI – ORIGINAL SCIENTIFIC PAPERS Šumarski list br. 11–12, CXXXIV (2010), 581-591
UDK 630* 182 (001)


SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS
ON ABANDONED AGRICULTURAL LAND IN HALOZE


VRSTE DRVEĆA I GRMLJA TE STRATEGIJA ZARASTANJA NAPUŠTENOG
POLJOPRIVREDNOG ZEMLJIŠTA NA PODRUČJU HALOZA U SLOVENIJI


Mateja COJZER*, Robert BRUS**


SUMMARY: In Slovenia, as well as in others parts of Europe, the share of
abandoned agricultural land overgrown by forest has been increasing every
year. This article deals with this process of succession in Haloze, in the northeastern
part of Slovenia. The main aim of this research was to find out how
much of the abandoned agricultural land on the studied area has succeeded to
forest in the last 20 years, to examine differences in species composition and the
density of individuals of tree and shrub species between abandoned areas and
younger developmental phases of forest, as well as to point out the strategies of
succession on abandoned areas and compare them with the vegetation process
of younger developmental phases in forest. Forest area increased by 7 % in the
period from 1985 to 2005 in the study region. The results show that the successional
process on abandoned land starts with shrub species, while in younger
phases of forest, tree species prevail entirely.


Key words: abandoned agricultural land, successional pathways on
abandoned land, old-field succession, vegetation process in forest, species
composition, density of individuals, younger developmental phases of forest,
the Haloze region.


1. INTRODUCTION – Uvod
Dense forest cover is one of the basic features in conditions; and socio-economic and political circum-
Slovenia, where forests cover more than half (58.4 %) stances (Golob et al., 1994). This process has been
of the total territory (ZGS, 2007). This is in large part going on since the beginning of the last century, and at
because Slovenia is a mountainous country. More than an accelerated rate after World War II (Hudoklin,
one third of the land area is above the altitude of 600 m, 2004). During the second half of the 20th century, the
and two thirds of this area is forested. Approximately mountainous and hilly areas in Slovenia were abandohalf
of the land has a slope incline greater than 20 % ned and suffered high population emigration (Mlinand
a good fifth more than 35 % (Perko,2004). Fo-šek,1968; Cunder,1998; Kob ler,2001; Hočevar
rest has been mainly preserved at higher and steeper lo-et al., 2004; Kozorog, 2004; Kobler et al., 2005).
cations, which are less suitable for agriculture. The changes in land use during the last 60 years have led


to extensive re-vegetation with an increase in shrubs and


The amount of forest area in Slovenia is still increaforests.
According to ZGS (2007), the forest area has in-


sing. The main reasons for this are: the abandonment of
creased by 60.5 % in the last 130 years and by 34.6 %


farmland and agricultural land (grasslands, pastures, vi-
since 1947. Similar to other parts of Slovenia, the share


neyards, meadows and orchards) (Perpar, 2002); a de-
of forest area and abandoned agricultural land have in


crease in livestock pressure; inconvenient natural
creased in the Haloze region, for the same aforementio


* ned reasons.
Mateja Cojzer, univ. dipl. inž. gozd., ZGS, OE Maribor,


Tyrševa 15, 2000 Maribor, Slovenia, mateja.cojzer@zgs.gov.si


Succession on abandoned agricultural land has be


** Prof. dr. Robert Brus, univ. dipl. inž. gozd., Biotechnical Faculty,


come a serious problem not only in Slovenia, but also


Department of Forestry and Renewable Forest Resources,
Večna pot 83, 1000 Ljubljana, Slovenia, robert.brus@bf.uni-lj.si elsewhere in Europe (Borec et al., 2004). In studies




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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591
conducted throughout Europe, most studies have examined
natural and socio-economical factors associated
with old-field succession (Walther, 1986; Baldock
et al., 1996; MacDonald et al.; 2000; Mather,
2001; Kozak,2003; Kobler etal., 2005; DLG, 2005;
Lasanta et al., 2006; Gellrich, 2007; Pueyo and
Begueria,2007; Rickebusch 2007; ). There is, however,
a lack of research on tree and shrub composition
on abandoned land and a lack of research on planned
management of successive changes of stands, which occurred
in these areas. Even rarer are studies that investigate
the hilly Sub-Pannonian district, such as Haloze.
Consequently, our research in these abandoned areas fo


cused on the species composition, which is extremely
rich in the north-eastern part of Slovenia.


Specifically, our objectives were to:


Determine how much of the Haloze area has become
overgrown in the period from 1985 to 2005,
and what the estimated area of forests will be by the
year 2015;


Study the differences in composition and density of
shrub and tree species between abandoned land and
forest;


Study the strategies of succession on the abandoned
land and compare them to the vegetation dynamics
in forest.
2 MATERIAL AND METHODS – Materijal i metode


2.1 Research area – Područje istraživanja
This research was carried out in Haloze (at 46°19´N,
15°52´E), in the north-eastern part of Slovenia.


The hilly relief of Haloze is very steep with numerous
valleys and ditches (Belec, 1961). The absolute altitudes
on the study area range from 220 to 458 m a.s.l.
The climate is Sub-Pannonian and is characterised by
hot summers, dry and sunny early autumns, and cold
winters. The annual mean air temperature is 9.7°C (Klimatografija
Slovenije, 1995), and the average tempera


ture during the vegetation period is 15.3°C. The average
annual precipitation varies from 928 up to 1.075 mm/m2
(Bračič, 1982). Geology in the region consists of
Miocene sediments, mainly covered by dystric soil on
marl and marl sandstone (Belec, 1961). The Haloze
region is on the Sub-Pannonian margin of the Predinaric
phytoclimatic territory (Košir, 1994). The study area
is dominated by beech forest sites. Neutrophilic beech
forests (Hedero-Fagetum Košir /62, 79/ 94) grow in rich


Figure 1 Locations of the research plots in the study area in north-eastern Slovenia (Slovenia Forest Service)


Slika 1. Lokacije istraživačkih ploha na području istraživanja, koje leži na sjeveroistočnom djelu Slovenije


and moist habitats, while acidophilic beech forests (Ca-regular water supply. The syntaxonomic nomenclature
staneo-Fagetum sylvaticae /Mar. & Zup. 79/ Mar. & in this study follows Robič and Accetto (1999).
Zup. 95) grow on well-drained and dry sites with an ir


2.2 Data collection – Prikupljanje podataka
Abandoned land areas were recorded in the latest was used as a basis to determine the plot site locations.
forest management plans for the Vzhodne Haloze and Plot locations were chosen on the basis of digital ort-
Rodni vrh units (ZGS 2005a, 2005b). This database hophotos (scale 1:5.000). Important criteria for choo




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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591
sing plots were the size, the shape and the developmental
stage of the abandoned land. All plots were square
shaped and north-east oriented.


Vegetation records were carried out on 52 plots in
total. 37 plots (20x20 m) were set on abandoned agricultural
land, of which 5 were in young growth developmental
stages, 15 in a thicket phase, and 17 in a pole
stand phase. 15 plots (20x10 m), 5 for each developmental
phase, were analysed in forest areas. The survey was
carried out from the beginning of August until the end of
October 2006, and in September 2007. Each plot was set
according to its developmental phase (one plot – one developmental
phase), and its floristic composition was recorded
by counting the individuals of tree and shrub


species. The source of the plant nomenclature was the
Mala flora Slovenije (Martinčič et al., 2007).


To analyse the vegetation composition for every
single developmental phase, an appropriate treatment
was assigned to each study plot. Six equivalent treatments
were: A – abandoned land in young growth
phase, B – abandoned land in thicket phase, C – abandoned
land in pole stand phase, D – forest in young
growth phase, E – forest in thicket phase, F – forest in
pole stand phase. All vegetation samples which were
carried out on abandoned land, together created Z treatment,
and all vegetation samples which were carried
out on forest plots, formed G treatment.


2. 3 Statistical analyses – Statističke analize
MapInfo v. 8.5 software was used for spatial data
processing, and Excel for data analysis. Statgraphics
Plus for Windows software was used for statistical data
processing. Data were analysed by analysis of variance
(One-way ANOVA), where parametric (t-test) and nonparametric
tests (LSD test, Duncan variance homogeneity
test) were used. Since the number of subjects per
ha is not a normally distributed variable, a preliminary
»square root« transformation was performed for the
ANOVA. Species diversity was calculated by the Shannon’s
diversity index (H´; H´= -.(pi ln pi)) (Shannon,
1948). CANOCO 4.5 for Windows (ter Braak and


Šmilauer, 2002) was used for calculation. The number
of species per plot (species composition), their stability,
and species density (the number of species
individuals per ha) were calculated for each plot separately.
Species stability was calculated according to the
share of individual species occurrence in the previous
samples. We analysed the plot similarity vegetation
composition by using the detrended correspondence
analysis (DCA), considering the number of individuals
per ha. DCA analysis was carried out with the help of
CANOCO 4.5 for Windows (ter Braak and Šmilauer,
2002).


3 RESULTS – Rezultati


3. 1 The share of forest area and abandoned land –
Udio šumskih površina i zemljišta u zarastanju


In twenty years (1985-2005) the forest area of Haloze
increased by 6.9 % (Tab. 1). The share of forest
cover in the Haloze region is thereby 42.2 %, but
twenty years ago it used to be 39.4 %. And, if (by the
next forest management plan renewal) all presently recorded
areas of abandoned land are classified as forest,
in 2015 (comparably to the year 1985) the forest area
will have increased by 13.0 %, and the total forest area
of Haloze will total 44.5 %.


Table 1
Changes in the forest area in Haloze in the period
from 1985 to 2005 (ZGS, 2005)


Tablica 1. Promjena šumskih površina u Halozama od 1985
do 2005 godine


Year
Godina
Forest area
Površina šuma
(ha)
Index
Indeks
(%)
1985 6916.70 100.0
2005 7396.70 106.9
2015 7803.49 112.8


3. 2 Species composition and density of individuals (on abandoned land and forest) –
Vrste drveća i grmlja te gustoća jedinki (na napuštenom zemljištu te u šumi)


Abandoned land was more species diverse than forest;
47 species were identified on abandoned land (33
tree and 14 shrub species), and 36 species in forest (25
tree and 11 shrub species) (Tab. 4).


With regard to the treatment, the ordination shows
(Fig. 2), that by the first axis, which explains most of
the variability, we can distinguish between two groups
of samples: abandoned land (Z – treatment) and forest
(G - treatment). The total inertion of DCA is 6.281; the
eigenvalues of the first four axes were 0.728, 0.470,


0.322, 0.201, which cumulatively explained 11.6 %,


19.1 %, 24.2 %, 27.4 % of variability in species data.
Gradient length of the first axis is 4.663, which justifies
the use of uni-modal ordination methods (ter Braak and
Šmilauer, 2002). The second axis shows a relatively
high variability in vegetation composition per plot on
abandoned land and a little lower variability in forest.
Table 2 shows the average number of species per
plot. There was no difference in average number of all
species per plot, and in the average number of tree spe




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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591
Figure 2 DCA ordination plot of 52 vegetation samples. Samples are marked with the sym-
bols denoting the treatment (Z – abandoned land, G – forest). First and second
axes explain 11.6 % and 7.5 % of variability in species data, respectively.


Slika 2.DCA ordinacija popisa označenih po tretmanima (Z – zemljište u zarastanju,
G – šuma). Prva os pojašnjava 11,5 % a druga 7,5 % varijabilnosti podataka


o distribuciji vrsta po popisima.
cies per plot between forest and
abandoned land at p<0.05. But
there was a significant difference in
the average number of shrub species
per plot.


In forest there were mainly


(91.5 %) tree species individuals,
while shrub species represent only
8.5 %. On abandoned land the proportion
was more balanced, where
tree species represent 53.0 %, and
shrub species 47.0 % of the share.
In the forest area, Fagus sylvatica
was the most abundant tree species,
followed by Carpinus betulus and
Quercus petraea. For shrubs, Sambucus
nigra was the dominant species,
followed by Corylus avellana
and Cornus sanguinea. On abandoned
land, Carpinus betulus was the
dominant tree species, followed by
Alnus glutinosa and Acer pseudoplatanus.
Among the shrub species,
Cornus sanguinea was the most
abundant, followed by Corylus
avellana and Prunus spinosa.
Table 2 The average number of species per plot separately according to treatments (Z - abandoned land, G - forest)


Tablica 2. Prosječan broj vrsta po jedinici popisa (plohi) s obzirom na tretmane (Z - napušteno zemljište, G - šuma)


All species Tree species Shrub species
Sve vrste Vrste drveća Vrste grmlja
Average F p Average F p Average F p
Treatments Count number number number
Tretmani Broj ploha per plot
Prosječan broj
na plohi
per plot
Prosječan broj
na plohi
per plot
Prosječan broj
na plohi
G 15 12.3 9.7 2.7
Z 37 13.7 9.2 4.4
G x Z 1.01 0.312 0.16 0.691 8.74 0.005


Table 3 The density of individuals per ha separately according to treatments (Z - abandoned land, G - forest)


Tablica 3. Gustoća jedinki na hektar, prikazano po tretmanima (Z - napušteno zemljište, G - šuma)


Treatments
Tretmani
Count
Broj
ploha
All species
Sve vrste
Tree species
Vrste drveća
Shrub species
Vrste grmlja
Average
number of
individuals
Prosječan
broj jedinki
(ha-1)
F p Average
number of
individuals
Prosječan
broj jedinki
(ha-1)
F p Average
number of
individuals
Prosječan
broj jedinki
(ha-1)
F p
G 15 23906.3 21884.0 2022.3
Z 37 19447.2 10301.2 9146.0
G x Z 1.17 0.203 12.02 0.001 17.32 <0.001
Transformation
Transformacija
Sqrt (number of individuals
all species ha-1)
Korijen(broj jedinki
svih vrsta na ha)
Sqrt (number of individuals
tree species ha-1)
Korijen (broj jedinki
vrste drveča na ha)
Sqrt (number of individuals
shrub species ha-1)
Korijen (broj jedinki
vrste grmlja na ha)




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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591
586
There was no difference in the Shannon’s diversity
index between the forest and abandoned land treat-
ments (ANOVA at p<0.05; F=0.06; p=0.802) (Fig. 3).
On average a little higher H’ value was typical for the
abandoned land treatment (1.48) and the forest treat-
ment (1.43) (Fig. 3).
Figure 3 Box-plot of plant species richness (S) and the Shannon’s diversity index (H’) of treatments (Z - abandoned land, G - forest).
Slika 3. Box-plot za prosječan broj vrsta drveća i grmlja (S) za Shannonov indeks (H’) po tretmanima (Z - napušteno zemljište, G - šuma).
3.3 Species composition and the density of plant species individuals between
the developmental phases of forest and abandoned land –
Vrste drveća i grmlja te gustoća jedinki vrsta drveća i grmlja
po razvojnim stadijima (tretmanima)
Treatments differ in average number of all species
per plot (p<0.05; F=3.09; p=0.017), in average number
of tree species (F=4.95; p=0.001) and in average number
of shrub species per plot (F=2.62; p=0.036) (Tab. 5).
Table 5 Mean values of the number of species between the treatments. Homogeneous groups are marked with small letters
(Duncan test at p<0.05).
Tablica 5. Prosječne vrijednosti broja vrsta drveća i grmlja po tretmanima. Homogene grupe (skupine) su označene malim
slovima (Duncanov test kod p<0.05)
All species Tree species Shrub species
Sve vrste Vrste drveća Vrste grmlja
Count Average Homogeneous Average Homogeneous Average Homogeneous
Treatments Broj number groups number groups number groups
Tretmani ploha per plot Homogene per plot Homogene per plot Homogene
Prosječan broj grupe Prosječan broj grupe Prosječan broj Homogene
na plohi na plohi na plohi
A 5 7.8 a 3.8 a 4.0 abc
B 15 14.5 b 9.7 b 4.7 c
C 17 14.7 b 10.4 b 4.3 bc
D 5 11.2 ab 9.4 b 1.8 a
E 5 13.8 b 11.6 b 2.2 ab
F 5 12.0 ab 8.0 b 4.0 abc
The average number of species per plot increased
with developmental phase aging on abandoned land.
The number of tree species increased as well, whilst the
number of shrub species did not change significantly
by developmental phase aging. The average number of
species in forest did not change essentially by develop-
mental phase aging, and the average number of tree
species did not change significantly as well, but the
average number of shrub species increased with deve-
lopmental phase aging.
There was a difference in the density of tree species
individuals (F=7.46; p=0.000) and in the density of
shrub species (F=6.34; p=0.000) (Tab.6).


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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591
It is evident (Tab. 3) that the species density of spe-the density of shrub species individuals. In forest the
cies individuals was a little higher in forest than on density of tree species individuals was twice as high as
abandoned land. There was no difference in the density the density of tree species individuals on abandoned
of all species individuals at p<0.05 between the forest land. And the density of shrub species individuals was
and abandoned land treatments. But there was a diffe-considerably higher on abandoned land than in forest.
rence in the density of tree species individuals and in


Table 4
The average number of individuals per plant species, separately for forest and abandoned land and according to
treatments (developmental phases). Data are shown as a number of individuals ha-1.


Tablica 4. Prosječan broj jedinki pojedinih vrsta drveća i grmlja, posebno za šumu i napuštena zemljišta te odvojeno/posebno
po tretmanima (stadiji razvoja šuma). Podaci su izraženi u brojevima jedinki po hektaru.


Species
Vrste
Tree/
shrub
Drveće/
grmlje
Forest
Šuma
Abandoned land
Napuštena zemljišta
Treatments
Tretmani
Average
number of
individuals
Prosječan
broj
jedinki (ha-1)
Presence
number of
plot
Frekvencija
pojavljanja
po plohama
Average
number of
individuals
Prosječan
broj
jedinki (ha-1)
Presence
number of
plot
Frekvencija
pojavljanja
po plohama
A B C D E F
Picea abies tree -drveće 750 4 125 4 0 50 75 50 550 150
Abies alba tree -drveće 11650 6 500 2 0 0 500 8600 3050 0
Pinus sylvestris tree -drveće 130 2 475 2 0 200 275 50 80 0
Pinus strobus tree -drveće 160 1 0 0 0 0 0 0 160 0
Larix decidua tree -drveće 50 1 0 0 0 0 0 0 0 50
Fagus sylvatica tree -drveće 145115 13 5450 14 0 475 4975 102015 37350 5750
Malus sylvestris tree -drveće 0 0 250 3 0 150 100 0 0 0
Pyrus pyraster tree -drveće 50 1 5910 10 0 870 5040 50 0 0
Quercus petraea tree -drveće 35030 13 18465 26 200 4455 13810 22330 5850 6850
Castanea sativa tree -drveće 13685 9 5950 17 0 1050 4900 1055 11130 1500
Aesculus hippocastanum tree -drveće 0 0 50 1 0 50 0 0 0 0
Robinia pseudacacia tree -drveće 0 0 725 2 125 600 0 0 0 0
Juglans regia tree -drveće 1210 4 3240 17 0 1985 1255 0 160 1050
Acer pseudoplatanus tree -drveće 8880 11 34735 29 400 12480 21855 810 5620 2450
Acer platanoides tree -drveće 50 1 1195 6 0 150 1045 0 50 0
Fraxinus excelsior tree -drveće 50 1 2125 2 0 25 2100 0 0 50
Ulmus glabra tree -drveće 800 3 200 2 0 25 175 0 800 0
Ulmus carpinifolia tree -drveće 0 0 1375 2 0 500 875 0 0 0
Tilia cordata tree -drveće 3695 9 1830 15 0 900 930 1315 1130 1250
Carpinus betulus tree -drveće 52185 13 141360 30 50 29140 112170 7135 11750 33300
Sorbus domestica tree -drveće 0 0 50 1 0 0 50 0 0 0
Prunus avium tree -drveće 17415 13 8825 22 30 2955 5840 8795 4720 3900
Prunus domestica tree -drveće 100 2 17855 13 1350 2140 14365 0 0 100
Prunus cerasus tree -drveće 0 0 275 2 0 175 100 0 0 0
Acer campestre tree -drveće 12920 14 32580 30 1490 17300 13790 2680 5890 4350
Sorbus torminalis tree -drveće 7480 9 2230 10 100 100 2030 4850 2130 500
Fraxinus ornus tree -drveće 8435 8 17025 17 315 6570 10140 615 2470 5350
Quercus cerris tree -drveće 0 0 4555 7 0 2850 1705 0 0 0
Populus tremula tree -drveće 1050 2 17370 12 0 12210 5160 0 0 1050
Populus alba tree -drveće 0 0 2295 2 0 150 2145 0 0 0
Alnus glutinosa tree -drveće 6300 1 39610 12 400 26825 12385 0 0 6300
Alnus incana tree -drveće 0 0 500 1 0 0 500 0 0 0
Betula pendula tree -drveće 580 2 8910 16 1485 3100 4325 0 80 500
Salix caprea tree -drveće 490 2 4805 12 0 4000 805 0 490 0
Salix sp. tree -drveće 0 0 300 1 0 0 300 0 0 0
Cornus sanguinea shrub - grmlje 5550 7 194550 34 54870 100615 39065 2850 650 1850
Crataegus monogyna shrub - grmlje 1200 6 11545 15 2475 840 8230 250 650 300
Prunus spinosa shrub - grmlje 350 1 34700 19 11825 19400 3475 0 0 350
Ligustrum vulgare shrub - grmlje 4000 4 18695 18 1005 9185 8505 750 700 2550
Corylus avellana shrub - grmlje 7675 10 41375 18 0 13000 28375 225 4850 2800
Sambucus nigra shrub - grmlje 9910 6 11650 10 0 7225 4425 1010 7600 1300
Viburnum lantana shrub - grmlje 1350 2 3365 5 685 1280 1400 0 1300 50
Viburnum opulus shrub - grmlje 0 0 765 2 0 365 400 0 0 0
Berberis vulgaris shrub - grmlje 50 1 475 2 0 425 50 0 0 50
Lonicera xylosteum shrub - grmlje 150 1 100 1 0 0 100 0 0 150
Juniperus communis shrub - grmlje 50 1 4140 3 0 1840 2300 0 0 50
Rosa sp. shrub - grmlje 50 1 10685 23 2580 5850 2255 0 0 50
Frangula alnus shrub - grmlje 0 0 2605 7 0 975 1630 0 0 0
Euonymus europaea shrub - grmlje 0 0 3750 7 2000 1700 50 0 0 0




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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591
Table 6
Mean values of the density of individuals per ha between treatments. Homogeneous groups are marked with small
letters (Duncan test at p<0.05).


Tablica 6. Prosječne vrijednosti gustoće jedinki na ha, posebno po tretmanima. Homogene grupe (skupine) su označene
malim slovima (Duncanov test kod p<0.05)


Treatments
Tretmani
Count
Broj
ploha
All species
Sve vrste
Tree species
Vrste drveća
Shrub species
Vrste grmlja
Average
number
per plot
Prosječan broj
jedinki (ha-1)
Homogeneous
groups
Homogene
grupe
Average
number of
individuals
Prosječan broj
jedinki (ha-1)
Homogeneous
groups
Homogene
grupe
Average
number of
individuals
Prosječan broj
jedinki (ha-1)
Homogeneous
groups
Homogene
grupe
AB
C
D
E
F
5
15
17
5
5
5
16277.0
19612.0
20234.1
33087.0
21842.0
16790.0
a
a
a
a
a
a
1189.0
8765.3
14336.5
32070.0
18692.0
14890.0
a
b
b
c
bc
b
15088.0
10864.7
5897.6
1017.0
3150.0
1900.0
c
bc
ab
a
a
a


The density of all species individuals
on abandoned land increased
with developmental phase aging
(Fig. 4). This pattern is reversed in
forest, where the density of all species
individuals decreased with developmental
phase aging (Fig. 4).
The findings for the density of tree
species individuals were similar.
On abandoned land the density of
shrub species individuals decreased
with developmental phase aging.
There was no significant difference


shrub species - vrsta grmlja
tree species - vrsta drveća
Figure 4 The density of tree and shrub species individuals per ha between treatments
in the density of shrub species indi


(A - abandoned land in young growth phase, B - abandoned land in thicket phase,
viduals in forest. C - abandoned land in pole stand phase, D - forest in young growth phase,


E - forest in thicket phase, F - forest in pole stand phase).


Slika 4.
Gustoća jedinki vrsta drveća i grmlja po hektaru, posebno po tretmanima ( A - zemljište
u zarastanju u razvojom stadiju mladika, B - zemljište u zarastanju u razvojom
stadiju koljika, C - zemljište u zarastanju u razvojom stadiju letvika, D - šuma u razvojom
stadiju mladika, E - šuma u fazi koljika, F - šuma u razvojnom stadiju letvika)


3.4 Overgrowing strategies on abandoned land and vegetation process in forest –
Strategija zarastanja na zemljištima u zarastanju te vegetacijski procesi u šumi


Treatments A, B and C deal with developmental “Cornus sanguinea” stage in this phase was replaced
phases on abandoned land. In treatment A there was the by the “Carpinus betulus” stage.
most shrub species (Fig. 4). Thus, in young growth The share of the most numerous species, Cornus
Cornus sanguinea was dominant (Tab. 4). Among tree sanguinea, strongly decreased in treatment A with de-
species Acer campestre and Betula pendula were the velopmental phase aging, whilst the share of Carpinus
most abundant. This is the first initial stage of shrub ve-betulus increased. Shrub species were replaced by tree
getation, which can be defined as a »Cornus species in later developmental phases.
sanguinea« stage. Shrub species still prevailed in treat-Treatments D, E and F denote initial developmental
ment B, but their share decreased in favour of tree spe-phases in forest. In treatment D there was an abundance
cies. Cornus sanguinea thus still prevailed in the of tree species (Fig. 4). Among the young growth pre-
thicket stage, and was followed by Carpinus betulus. vailed Fagus sylvatica. Cornus sanguinea was the do-
We can still talk about the »Cornus sanguinea« stage. minant shrub species (Tab. 4). In treatment E tree
In treatment C tree species already prevailed over species still prevailed; their share was the least in this
shrub species. Carpinus betulus was abundant, while phase, but the share of shrub species increased a bit. In
Cornus sanguinea was abundant in the shrub layer. The the thicket phase Fagus sylvatica still prevailed, alt




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hough its share decreased by almost a half. Sambucus
nigra had a significant share among shrub species.
Treatment F was in the pole stand phase. The tree species,
whose share again increased a bit, still strongly
prevailed over shrub species in this phase. The most
abundant was Carpinus betulus. Among shrub species
Corylus avellana had a significant share.


The share of Fagus sylvatica, as the most numerous
species in treatment D, strongly decreased with developmental
phase aging, whereas the share of Carpinus
betulus increased. On abandoned land as well as in forest
the share of Carpinus betulus increased with developmental
phase aging.


4 DISCUSSION – Rasprava


The Haloze region has a similar trend of forest
growth as Slovenia. Succession on abandoned agricultural
land is influenced by numerous natural, socio-economic,
infrastructural and accidental factors (Hočevar
et al., 2004). The successinal process on farmland immediately
starts after mowing or grazing is ceased, and
in Haloze also after abandonment of vineyards and orchards.
Vegetation on abandoned land develops through
different stages (pioneer, middle and final stages), and
proceeds in the direction of a late successional forest.
Mature forest stage restoration is a long-term process
that is connected with distinctive changes in species
(Benabdellah et al., 2003). On the studied area
Beech sites prevail, species composition changes
through stages and is oriented toward beech associations
ZGS, 2005a, 2005b). Only pioneer (early) stages were
studied on abandoned land.


47 different species were recorded on abandoned
land in the Haloze study area, of which 33 tree and 14
shrub species were found. A similar study was done in
the Kočevje region (Mlinšek, 1968), where 38 species
were recorded on fir-beech sites, of which 20 tree and 18
shrub species were found. If we compare the leading
species, the most numerous shrub species on the Haloze
hilly area is Cornus sanguinea, and among tree species
Carpinus betulus prevails. In the Kočevje region the
successive development inolves Corylus avellana, and
among tree species Populus tremula prevails (Mlinšek,
1968). In Slovenia, as well as elsewhere in Europe
and around the world, early stages on abandoned land
involve different species. The succession of species immigration
is different, depending mostly on the site and
also on coincidence, and the immigration rate depends
on the seed bank presence, soil state of preparation for
germination and on the pioneer competition (Mlin šek,
1968). In the Pyrenees succession occurs with
Pinus sylvestris (Pueyo and Begueria,2007), but on
Slovenian karst it occurs with Pinus nigra (Eler,
2007). The results of other studies are hardly comparable
with our findings, since most of the studies deal with


successive changes of herbaceous species, which were
not investigated in our case, and some other studies deal
only with tree species without shrub species.


In this study on abandoned land in Haloze, it was demonstrated
that shrub species first appear, among which
Cornus sanguinea is the most frequent. Shrub species
represent an important ecological middle stage in forest
development. In the next stage the share of shrub species
decreases, and the share of tree species increases. At this
developmental stage some economically significant tree
species are already present. In the pole stand phase its
structure and species composition are a bit more formed.
At this stage tree species strongly prevail over shrub
species, which have successfully completed their task
and are slowly disappearing. On overgrowing land pioneer
forest developed, by which site conditions are improved,
since under its protection gradually other
managed, shade tolerant and semi-shade tolerant species
develop, and form managed forest. In later stages the
share of economically significant species increases quickly
(Mlinšek, 1968).


It was found that in rejuvenated gaps of forest,
Fagus sylvatica regenerates abundantly, which is otherwise
a shade tolerant species. In very good conditions
it germinates in abundance, but after a few years
the number of individuals decreases quickly. This is accordance
with the statement by Marinček (1987),
who ascertained that elimination of individuals is particularly
severe in the first few years and that the number
of individuals slows down later. We discovered that its
share almost halves in the thicket phase (Tab. 4), yet it
still prevails over other tree species. In the pole stand
phase the share of Fagus sylvatica diminishes again. In
the race for territory Carpinus betulus won, and Quercus
petraea is also competitive.


The final vegetation stage is beech forest. It is thereby
expected that the share of Fagus sylvatica will increase
again in older developmental phases, and the
share of Carpinus betulus will decrease.


5. CONCLUSION – Zaključak
Abandoned land area in the Haloze region is increa-We also found that abandoned land is a bit more di-
sing every year. It was found that forest area in the period verse than forest with regard to species composition.
from 1985 to 2005 increased by 6.9 %, and it is expected Abandoned land differs from forest in the number of
to increase by another 5.5 % in the next decade. shrub species, but it does not differ in the number of




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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591
tree species. The biggest distinction in species composition
was found between developmental phases within
abandoned land, since pole stands are the most diverse,
and the least diverse is young growth. There are also
differences in the density of tree and shrub species between
abandoned land and forest. In forest the density
of tree individuals is higher, whilst on abandoned land
the density of shrub species individuals is higher. The
density of tree species individuals on abandoned land
increases with developmental phase aging, but the density
of shrub species individuals decreases. The oppo


site pattern occurs in forest, where the density of tree
species individuals decreases with developmental
phase aging, and the density of shrub species individuals
does not change significantly with developmental
phase aging.


The successinal process on abandoned farmland
goes from initial “Cornus sanguinea” – “Cornus sanguinea”
– ”Carpinus betulus” stages, and vegetation
process in forest from “Fagus sylvatica” – ”Fagus sylvatica”
– ”Carpinus betulus” stages.


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stevilkah-2007/index.htm


SAŽETAK: Slovenija je jedna od najšumovitijih država u Europi, jer – po
podacima Slovenske službe za šumarstvo (Zavod za gozdove Slovenije, 2007)


– šume pokrivaju više od polovice državnog teritorija (58,5 %). Zarastanje
napuštenog poljoprivrednog zemljišta šumom i dalje napreduje te tako postaje
ozbiljan problem. Glavni razlog zarastanja je napuštanje poljoprivrednih
imanja, ponajprije zbog teških prirodnih uvjeta te socioekonomske politike.
Taj proces traje već od početka 20. stoljeća, a posebno se ubrzao nakon kraja
drugog svjetskog rata.
U istraživanju smo se ograničili na područje Haloza, koje leži na području
sjeveroistočne Slovenije i sastavni je dio rubnog dijela panonske nizine. Istraživanjem
smo željeli proučiti koliko se poljoprivrednih površina u zadnjih dvadeset
godina pretvorilo u šumu, utvrditi razlike u sastavu te gustoći jedinki
pojedinih vrsta drveća i grmlja na područjima u zarastanju i u mlađim razvojnih
stadijima šuma, te naznačiti strategiju zarastanja napuštenih poljoprivrednih
površina uz istovremenu usporedbu sa stanjem vegetacijskih procesa
u mlađim razvojnim stadijima šuma.




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Istraživanjem smo utvrdili da se na području Haloza u zadnjih dvadeset
godina (u razdoblju od 1985. do 2005. godine) površina šuma povećala za
6,9 %, dok u sljedećih deset godina očekujemo povećanje od 5,5 %. Vegetacijske
smo popise, koje smo napravili na 52 plohe (37 ploha na poljoprivrednom
zemljištu i 15 ploha unutar šume), analizirali pomoću DCA analize. Ordinacija
je pokazala da pomoću prve osi, koja pojašnjava najveći dio varijabilnosti
(11,6 %), možemo razlikovati dvije izrazite skupine/grupe: u prvoj grupi su
plohe koje smo snimali u šumi, a u drugoj one koje smo snimali na zemljištu u
zarastanju (bivšem poljoprivrednom zemljištu). Na zemljištima u zarastanju
prosječno smo po plohi evidentirali 13,7 vrste, od toga je bilo 9,2 vrsta drveća
te 4,4 vrsta grmlja, dok smo po plohi u šumi evidentirali 12,3 vrste, od toga
9,7 vrsta drveća te 2,7 vrsta grmlja. Na zemljištima u zarastanju broj se vrsta
drveća sa starošću povećava, dok se broj vrsta grmlja bitno ne mijenja. U
šumi je situacija obrnuta: staranjem razvojnih stadija povećava se broj vrsta
grmlja, dok broj jedinki vrsta drveća ostaje više ili manje jednak. Veću smo
gustoća jedinki zabilježili u šumi (23.906,3 jedinki po hektaru), od toga vrste
drveća predstavljaju 91,5 %, a vrste grmlja samo 8,5 % svih jedinica po hektaru.
Najbrojnija vrsta drveća u šumi je Fagus sylvatica, a među vrstama grmlja
Sambucus nigra. Broj jedinki vrsta drveća u šumi uz staranje razvojnih
stadija pada, dok se broj jedinki vrsta grmlja bitno ne mijenja. U stadiju razvoja
mladika glavna je vrsta Fagus sylvatica, u fazi koljika je najčešće pojavljuje
Carpinus betulus. Na zemljištima u zarastanju zabilježili smo 19.447,2
jedinki po hektaru, od toga je udio vrsta drveća 53,0 % a vrsta grmlja 47,0 %
svih jedinica po hektaru. Između vrsta drveća na zemljištima u zarastanju je
najčešća vrsta Carpinus betulus, a među vrstama grmlja Cornus sanguinea.
Na zemljištima u zarastanju udio jedinki vrsta drveća uz staranje stadija razvoja
raste, a udio vrsta grmlja pada. Tako u mladiku dominiraju vrste grmlja,
među kojima u cjelini dominira Cornus sanguinea. U razvojnom stadiju koljika
njihov se udio polako smanjuje u korist vrsta drveća, no još uvijek je dominirajuća
vrsta Cornus sanguinea, dok je na drugem mjestu Carpinus
betulus. U fazi letvika vrste drveća već dominiraju nad vrstama grmlja. Najčešće
se pojavljuje Carpinus betulus.


Proces zarastanja na bivšim poljoprivrednim zemljištima prolazi preko inicijalnih
stadija “Cornus sanguinea”-”Cornus sanguinea”-”Carpinus betulus”,
a vegetacijski procesi u šumi preko stadija “Fagus sylvatica”-”Fagus
sylvatica”-”Carpinus betulus”. Konačni vegetacijski stupanj/stadiji je u oba
primjera šuma bukve. U Halozama ima Carpinus betulus na staništima bukve
(koji niže prelaze u šume bijelog graba) ulogu pionirske vrste. U starijim stadijima
ponovo dominira Fagus sylvatica.