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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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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591 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. 6. REFERENCES – Literatura Baldock, D., G. Beaufoy, F. Brouwer, F. Godeschalk, 1996. Farming at the Margins: Abandonment or Redeployment of Agricultural Land in Europe. Institute for European Environmental Policy, London/The Hague. Belec, B., 1961. Morfologija Haloz, Geografski zbornik VI, 163–190, Ljubljana. Benabdellah, B., K. F. Albrecht,V. L. Pomaz, W. A. Denisenko, D. O. Logofet, 2003. Markov chain models for forest successions in the Erzgebirge, Germany. Ecological Modelling 159: 14–160. Borec, A., A. Flambard, K. Pažek, 2004. Relationships between production system of Slovenian mountain farms and dynamics of overgrowing areas. Agricultura 3: 32–36. Bračič, V., 1982. Gozdnate Haloze, Založba Obzorja, 154, Maribor. Cunder, T., 1998. Zaraščanje kmetijskih zemljišč v Slovenskem Alpskem svetu. Sonaravni razvoj v slovenskih Alpah in sosedstvu. 1. Melikovi geografski dnevi, Kranjska gora, 5. –7. november 1998. Filozofska fakulteta, Oddelek za geografijo, 165–175, Ljubljana. DLG, 2005. Land Abandonment, Biodiversity and the CAP. Government Service for Land and Water Management of the Netherlands (DLG), Utrecht. Eler, K., 2007. Dinamika vegetacije travišč v slovenskem submediteranu: vzorci in procesi ob spremem bah rabe tal: doktorska disertacija. Biotehniška fakulteta, Oddelek za agronomijo, 228 str., Ljubljana. Gellrich, M., P. Baur, B. Koch, N. E. Zimmer mann, 2007. Agricultural land abandonment and natural forest re-growth in the Swiss mountains: A spatially explicit economic analysis. Agriculture, Ecosystems and Environment 118: 93–108. Golob, S., M. Hrustel-Majcen, T. Cunder, 1994. Raba zemljišč v zaraščanju v Sloveniji. Kako izboljšati posestno strukturo v Sloveniji. 9. tradicionalni posvet kmetijske svetovalne službe. Ministrstvo RS za kmetijstvo in gozdarstvo, 89–98, Bled. Hočevar, M., G. Kušar, T. Cunder, 2004. Monitoring in analiza zaraščanja kraške krajine v GIS okolju. Zbornik gozdarstva in lesarstva 75: 21–52. Hudoklin, J., 2004. Regional Distribution of Landscape Types in Slovenia and Outstanding Landscapes of Slovenia. http://www.mop.gov.si/fileadmin/mop.gov.si/pageuploads/ podrocja/prostor/pdf/studije/s3krajina.pdf Klimatografija Slovenije za obdobje 1961–1990. 1995, Hidrometeorološki zavod republike Slovenije. Kobler, A., 2001. Sprejemljivost zaraščanja kot funkcija kakovosti kulturne krajine: prostorski model: magistrsko delo. Biotehniška fakulteta, Oddelek za gozdarstvo in obnovljive gozdne vire, 187 str., Ljubljana. Kobler, A., T. Cunder, J., Pirnat, 2005. Modelling spontaneous afforestation in Postojna area, Slovenia. J. Nat. Conserv. 13: 127–135. Košir, Ž., 1994. Ekološke in fitocenološke razmere v gorskem in hribovitem jugozahodnem obrobju Panonije. Zveza gozdarskih društev, 149 str., Ljubljana. Kozak, J., 2003. Forest cover change in the Western Carpathians in the past 180 years. Mt. Res. Dev. 23: 369–375. Kozorog, E., 2004. Ali je zaraščanje kmetijskih površin problem? Gozdarski vestnik 62 (9): 407–408. Lasanta, T., J.C. Gonzalez-Hidalgo, S.M. Vicente- Serrano, E. Sferi, 2006. Using landscape ecology to evaluate an alternative management scenario in abandoned Mediterranean mountain areas. Landscape Urban Plan. 78: 101–114. MacDonald, D., J. R. Crabtree, G. Wiesinger, T. Dax, N. Stamou, P. Fleury, J. G. Lazpita, A. Gibon, 2000. Agricultural abandonment in mountain areas of Europe: Environmental consequences and policy response, Journal 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 Environmental Management 5: 47–69. Marinček, L., 1987. Bukovi gozdovi na Slovenskem. Delavska enotnost, 153 str., Ljubljana. Martinčič,A., T. Wraber, N. Jogan,A. Podobnik, B. Turk, B. Vreš,V. Ravnik, S. Frajman, B. Strgulc-Krajšek, B. Trčak, T. Bačič, M.A. Fischer, K. Eler, B. Surina, 2007. Mala flora Slovenije, Ključ za določevanje praprotnic in semenk. Četrta, dopolnjena in spremenjena izdaja, Tehniška založba, 968 str., Ljubljana. Mather, A.S., 2001. The transition from deforestation to reforestation in Europe. In: Angelsen, A., Kaimowitz, D. (Eds.), Agricultural Technologies and Tropical Deforestation. Center for International Forestry Research (CIFOR). CABI, Oxon/New York, 35–52. Mlinšek, D., 1968. Premena grmišč v Sloveniji, 39 str., Ljubljana. Perko, F., 2004. Gozd in gozdarstvo Slovenije. Zveza gozdarskih društev Slovenije v sodelovanju: Ministrstvo za kmetijstvo, gozdarstvo in prehrano RS: Zavod za gozdove Slovenije, 39 str., Ljubljana. Perpar, A., 2002. Stanje in procesi v kmetijstvu v različnih tipih podeželskih območij v Sloveniji. Zbornik Biotehniške fakultete Univerze v Ljubljani. Kmetijstvo 97, 1: 281–300. Pueyo, Y., S. Begueria, 2007. Modelling the rate of secondary succession after farmland abandonment in a Mediterranean mountain area. Landscape and Urban Planning 83 (4): 245–254. Rickebusch, S., M. Gellrich, H. Lischke, A. Guisan, N. E. Zimmermann, 2007. Combining probabilistic land-use change and tree po pulation dynamics modelling to simulate responses in mountain forests. Ecological Modelling 209 (2–4): 157–168. Robič, D., M. Accetto, 1999. Pregled sintaksonomskega sistema gozdnega in obgozdnega rastlinja Slovenije. Študijsko gradivo iz fitocenologije. Biotehniška fakulteta, Oddelek za gozdarstvo in obnovljive gozdne vire, 18 str., Ljubljana. Robič, D., 2000. Različno razumevanje in pomen biodiverzitete v ekologiji, posebno v fitocenologiji. Zbornik gozdarstva in lesarstva, 63: 47–93, Ljubljana. Shannon, C. E., 1948. A mathematical theory of communication. Bell. System. Technol. J. 27: 379–423, 623–653. terBraak, C. J. F., P. Šmilauer, 2002. CANOCO 4.5 for Windows – Software for canonical community ordination. Walther, P., 1986. Land Abandonment in the Swiss Alps: a new understanding of a land use problem. Mountain Research and Development 6: 305–314. ZGS, 2005a. Gozdnogospodarski načrt za gozdnogospodarsko eno to Rodni vrh za obdobje 2004– 2015. 2005, Za vod za gozdove Slovenije, OE Maribor, 117 str., Maribor. ZGS, 2005b. Gozdnogospodarski načrt za gozdnogospodarsko enoto Vzhodne Haloze za obdobje 2005–2014. 2005, Zavod za gozdove Slovenije, OE Maribor, 150 str., Maribor. ZGS, 2007. http://www.zgs.gov.si/slo/gozdovi-slovenije/ o-gozdovih-slovenije/slovenski-gozdv- 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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M. Cojzer, R. Brus: SPECIES COMPOSITION AND SUCCESSIONAL PATHWAYS ON ABANDONED ... Šumarski list br. 11–12, CXXXIV (2010), 581-591 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. |