zbornik gozdarstva in lesarstva 79 (2006), s. 37 - 52 gdK: 182.51+181.1+165:176.1:(497.12)(045) Prispelo / Recived: 06. 05. 2005 Sprejeto / Accepted: 11. 04. 2006 Izvirni znanstveni članek Original scientifc paper PLANT DIVERSITy OF SELECTED Quercus robur l. AND Quercus petraea (matt.) liebl. FORESTS IN SLOVENIA Lado KUTNAr1 Abstract In Slovenia, the plant species diversity on 225 research plots dominated by pedunculate oak (Quercus robur L.) and by sessile oak (Quercus petraea (matt.) Liebl.) has been analysed. Plots of Q. robur are located in fve, and plots of Q. petraea in four semi-natural managed forest complexes. In the tree layer, 28 species were found beside the dominant two oak species, with Carpinus betulus L., Picea abies (L.) Karst, Quercus cerris L. and Fagus sylvatica L. having signifcant shares of growing stock. Based on the understorey vegetation (shrub and herb layer, terricolous mosses), the detrended Correspondence Analysis (dCA) made a clear distinction between plots with dominant Q. robur and those with Q. petraea. The understorey vegetation also proved to be a valuable indicator of the site conditions and of forest management in the past. Based on ordination, lowland pedunculate oak forests of relatively long standing near to natural management have been separated from the pedunculate oak forests where spruce was favoured by the forest management, and from the man-made pedunculate oak stands on primary sites of Q. petraea. dCA clearly differentiated the sessile oak forests in warmer climate of Sub-mediterranean region, and in warmer meso-sites of Pre-Pannonian region from other sessile oak forests. The main gradients of vegetation structure and of species diversity, as well main ecological gradients in different oak forests were obtained by ordination technique. Key words: foristic composition, vegetation structure, biodiversity, growing stock, Quercus robur, Quercus petraea, dCA, Slovenia RASTLInSKA VRSTnA DIVERZITETA IZBRAnIh DOB OVIh In gRADnOVIh gOZDOV V SLOVEnIJI Izvleček na 22 raziskovalnih ploskvah v dobovih (Quercus robur L.) in gradnovih (Quercus petraea (matt.) Liebl.) gozdovih smo analizirali rastlinsko vrstno pestrost. Dobove ploskve so postavljene v petih, gradnove pa v štirih gospodarjenih gozdnih kompleksih. V drevesni plasti preučevanih gozdnih ekosistemov se poleg dominantnih vrst hrasta pojavlja še 28 vrst, med katerimi so v lesni zalogi bolj zastopani beli gaber (Carpinus betulus L), smreka (Picea abies (L.) Karst.), cer (Quercus cerris L.) in bukev (Fagus sylvatica L.). Ordinacija DCA (Detrended Correspondence Analysis) je samo na osnovi vrst v pritalnih plasteh vegetacije (grmovna in zeliščna plast, na tleh rastoči mahovi) jasno ločila ploskve s prevladujočim dobom od ploskev z gradnom. Vrste pritalnih plasti so dober indikator rastiščnih razmer kot tudi gospodarjenja z gozdovi v preteklosti. na tej osnovi je ordinacija ločila nižinske dobove gozdove z razmeroma sonaravnim gospodarjenjem v daljšem obdobju od dobovih gozdov, v katerih so v preteklosti pospeševali smreko, in od umetno zasnovanega dobovega gozda na primarnih rastiščih gradna. Analiza je ločila tudi gradnov gozd, ki uspeva v toplejšem podnebju v submediteranskem območju, in gradnov gozd na toplejših rastiščih v predpanonskem območju od preostalih dveh gradnovih gozdov. Ordinacija raziskovalnih ploskev dobro odseva razlike v foristični sestavi hrastovih gozdov, nakazuje gradiente pestrosti pritalnih plasti vegetacije in glavne ekološke gradiente. Ključne besede: foristična sestava, struktura vegetacije, biodiverziteta, lesna zaloga, Quercus robur, Quercus petraea, DCA, Slovenia INTRODUCTION UVOD Biodiversity has become an increasingly popular topic within the discussion of sustainability in the last decade, though the maintenance of diversity of forest ecosystems has been required for many years (SWINdEL / CoNdE / SmITH 1984, SCHULEr 1998). Biodiversity and heterogeneous vegetation structure play an important role in stable forests. Biodiversity changes could be a valuable indicator of global warming too. global warming is altering the distribution and abundance of plants. Application of a basic law of ecology predicts that many species will vanish if temperatures continue to rise (PoUNdS / PUSCHENdorF 2004). many forest understorey plants are sensitive indicators of environmental changes (OKLANd et al. 2004). dr. L. K., gozdarski inštitut Slovenije, oddelek za gozdno ekologijo, Večna pot 2, 1000 Ljubljana, SLo 1 38 Zbornik gozdarstva in lesarstva, 79 High biodiversity based on high variability of forest sites is one of the main characteristics of the European lowland oak forests. The lowland oak forests and other foodplain forest ecosystems could be considered as being biodiversity hotspots in Europe. However, due to human impact, which results from their vicinity to settlements, these forests are one of the most altered forest communities in Europe. Additionally, strong sources of air pollutants or changes of groundwater level and other causes diminish the stability and resistance of lowland forests. The foodplain forest ecosystems, especially oak forests throughout Europe, have always been under heavy anthropogenic impact (KLImo / HAgEr 2001). In Slovenia, as a main or admixed tree species fve autochthonous oak species (Quercus robur agg., Q. petraea agg., Q. cerris, Q. pubescens, and Q. ilex) represent less than 8 % of the total growing stock (SmoLEj / HAgEr 1995). The oak forests grow mostly in the lowland areas and their hilly margins, where the abundance of forest cover is low, but human population density is high. The greatest threat to native biodiversity of lowland oak forests is the loss and fragmentation of natural habitats. generally speaking, the loss and degradation of remnant native vegetation results in the loss of biodiversity. one of the signifcant threats to native biological diversity of foodplain forests and other lowland forests are invasions caused by alien species (KLImo / HAgEr 2001). In Slovenian lowlands, there has been a noticeable decline and physiological weakening of pedunculate oak (Quercus robur), connected with dry climate, unfavourable precipitation patterns and human infuences through hydromeliorations and changes in the water table (ČATEr / LEVANIČ 2004). many studies of oaks in Slovenia from different aspects have been done, such as phytosociological (e.g. ACCETTo 1974, PUNCEr / zUPANČIČ 1979, šUgAr et al. 1995, dAKSKoBLEr 1997, zUPANČIČ 1997), from phytoindi-cation aspects (SmoLE 1993, SmoLE / KUTNAr 1994a, 1994b, KUTNAr 1997), from ecophysiological aspects (ČATEr / SImoNČIČ / BATIČ 1999, ČATEr / BATIČ 1999, 2000), from morphological aspect (SmoLE / BATIČ 1992, BATIČ et al. 1997, TrAjBEr et al. 2001), from morphological and genetic aspects (BATIČ / SINKoVIČ / jAVorNIK 1995, BrEzNIKAr et al. 2000), and from many others (e.g. SmoLEj / HAgEr 1995, rogL et al. 1996, ČATEr / KUT- NAr / ACCETTo 2001, ČATEr / LEVANIČ 2004). These studies did not focus much on the evaluation of oak forest biodiversity in a narrow sense, and related vegetation-stand structure as a crucial component of stable forests. The aim of our study was to determine biodiversity and the main gradients of the vegetation structure of selected pedunculate oak (Quercus robur L.) and sessile oak (Quercus petraea (matt.) Liebl.) forests in Slovenia. MATERIAL AND METHODS MaTeRIal IN MeTODe STUDY AREA AND RESEARCH PLOTS štuDijsko območje in raziskovalne MeTODe A total of nine oak forest complexes were selected in different parts of Slovenia (SmoLEj 1995). All nine oak complexes are semi-natural managed forests. Five forest complexes of pedunculate oak (Quercus robur L.) are located mostly in the eastern part of Slovenia (Fig. 1): I. Polom, II. Krakovski gozd, III. dobrava, IV. Cigonca, V. Hraščica. Four forest complexes of sessile oak (Quercus petraea (matt.) Liebl.) are more dispersed (Fig. 1): 1. Panovec, 2. Bo-janci, 3. Pišece, 4. Bukovnica. most of the Q. robur complexes are located in the food-plains of rivers on the margins of Pannonian basin (SmoLEj 1995). on the Q. robur complexes deep hydromorphic soil prevails (KALAN 1995), which developed under the infu-ence of either water-logging above less permeable soil layers (pseudogley soils) or of a high groundwater-table (gley soils – amphigleys and hypogleys). The exception is the Polom complex, which differs from the others essentially. It lies in the rolly permeable lime hill country near Kočevje, where some surface waterfow can appear occasionally, but there is no direct groundwater infuence. The selected Q. petraea complexes are placed on more or less permeable hilly terrain with very different types of forest soils out of direct ground-water infuences. due to ecological regions in Slovenia (KUTNAr et al. 2002), all complexes of Q. robur except Polom are located in different sub-regions of the Pre-Pannonian region. The Polom complex lies in the Pre-dinaric region. The three of four Q. Kutnar, L.: Plant diversity of selected Quercus robur L. and Quercus petraea (Matt.) Liebl. forests in Slovenia 39 Fig. 1: Location of the oak forest research complexes in Slovenia Slika 1: Položaj hrastovih raziskovalnih kompleksov v Sloveniji petraea complexes are located in different parts of the Pre-Pannonian region. The exception is Panovec complex that is situated in the Sub-mediterranean region. The potential natural forest vegetation of complexes has been described (SmoLE 1993, 1995, SmoLE / KUTNAr 1994a, 1994b). The Polom complex was described as a potential site of sessile oak-beech forest hedero-Fagetum Koš. (62, 79) 94 nom.nov. (syn.: Querco-Fagetum Koš.62). The vegetation of the dobrava, Cigonca and Hraščica complexes were described as forests of pedunculate oak and hornbeam Querco roboris-Carpinetum SoÓ 40. The Krakovski gozd complex belongs to the Pseudostellario-Quercetum roboris ACC.73 association. The Panovec complex belongs to the Sub-mediterranean oak forests Carici umbrosae-Quercetum petraeae PoLd. 82 var. geogr. Sesleria autumnalis dAKS.87. The forest of the Bojanci complex was described as an association of Epi-medio-Carpinetum (HT.38) BorH.63. The Pišece complex lies in a region of potential beech forest (hacquetio-Fagetum Koš.62 var. geogr. Ruscus hypoglossum (mAr.& zUP.78) Koš.79), and the Bukovnica in region of potential hornbeam forest (Pruno padi-Carpinetum betuli (mAr. & zUP.84) mAr.94). Each quadratic (100×100 metres) complex (Fig. 1) was divided in 25 square (20×20 metres) plots (SmoLEj 1995). fIELD SAMPLINg TeReNskO DelO Trees with diameter at breast high (dBH) exceeding 10 centimetres were taken into account (AzAroV 1995). The understorey vegetation of the research plots was surveyed according to the standard Central European method (BrAUN-BLANQUET 1964). The cover of all species in shrub, herb and moss layer (only terricolous mosses) was estimated. The cover of the young trees (dBH <10 cm) was estimated in the same way as other plants in understorey layers. The sources of the nomenclature were mArTINČIČ et al. (1999) for vascular plants; mArTINČIČ (2003) for mosses. DATA ANALYSIS aNalIZa PODaTkOV The vegetation of 225 research plots was analysed. on all research plots, based on phytosociological relevés of the 0 Zbornik gozdarstva in lesarstva, 79 understorey vegetation, the following parameters were calculated: a) species richness (S), b) sum of cover estimations of all species per plot, c) mean species cover, d) Shannon [H’ = - S (pi log (pi))] diversity index, e) Simpson [d = 1 - E pi2] diversity index, and f) Evenness [E = H’/ln (S)]; pi - share of plant species of total. Based on feld measurement, the growing stock of trees was analysed by AzAroV (1995). We also calculated the number of trees per plot and the number of different tree species. Based on the results of AzAroV (1995), we estimated the total growing stock (gS) and growing stock of the most common tree species: Quercus robur L., Quercus petraea (matt.) Liebl, Carpinus betulus L., Fagus sylvaticaL., Picea abies (L.) Karst, Acer campestre L., Quercus cerris L., Alnus glutinosa (L.) gaertn., and Tilia cordata mill. The main structure gradients of vegetation were extracted by the detrended correspondence analysis (dCA) (HILL / gA-UCH 1980). The foristic composition was used as a criterion for ordination of the research plots. The dCA was carried out with the PC-ord program (mcCUNE / mEFFord 1999). The Spearman correlation coeffcients were calculated between the dCA axes (plot scores) and: a) tree layer parameters; b) understorey (shrub, herb, moss) vegetation parameters; RESULTS AND DISCUSSION ReZUlTaTI IN RaZPRaVa on 225 research plots, a total of 30 tree and shrub species with a diameter at breast high over 10 centimetres were found. The most common tree species are following (in parentheses are numbers of research plots with their presence): Quercus robur (124 plots), Quercus petraea (100), Carpinus betulus (90), Fagus sylvatica (56), Picea abies (50), Acer campestre (42), Quercus cerris (18), Alnus glutinosa (16), and Tilia cordata (16). other less common tree and shrub species with a diameter at breast high over 10 centimetres are following (order of precedence is due to frequency of species): Pinus nigra, Pyrus pyraster, Sorbus torminalis, Prunus avium, Crataegus monogyna, Pinus sylvestris, Ostrya carpinifolia, Fraxinus or- nus, Larix decidua, Populus tremula, Acer pseudoplatanus, Abies alba, Sorbus aria, Fraxinus excelsior, Betula pendula, Acer platanoides, ulmus glabra, Crataegus laevigata, ulmus laevis, Corylus avellana and Ilex aquifolium. The total number of trees and shrubs with a diameter at breast high over 10 centimetres was 4,003. on average, almost 18 individual trees (shrubs) (dBH>10 cm) per 20×20 metres plot were found. The number of individual trees (shrubs) per plot ranges between 4 and 35. The estimation of total growing stock per plot varies from 5.1 to 48.8 m3. Based on the foristic composition (presence/absence), nine oak-forest complexes are well-separated in the plot ordination space (Figs. 2, 3, 4). dCA analysis clearly differentiates between the Quercus robur and the Q. petraea dominated plots respectively. In dCA1 vs. dCA2 ordination (Figs. 2 and 3), plots of Quercus robur are below the diagonal of graph (start from point dCA1=0, dCA2=0), and plots of Q. petraea are above this diagonal. As a result of similarity of foristic composition (Figs. 2 and 3), in 3-d ordination space, the dobrava and Cigonca plots are grouped close together. There are swamped, gleyed soils resulting from impact of high groundwater-table. The Hraščica and Krakovski gozd plots are quite similar in foristic composition (Fig. 2). However, in 3-d ordination space (Fig. 3), signifcant differences between these plots are shown (Krakovski gozd plots obtained higher dCA3 scores than Hraščica plots). It is mainly a result of different soil conditions of these plots (KALAN 1995). on the Hraščica plots, eutric brown soils were found, and gleyed swampy soils were found on the Krakovski gozd plots. As a result of intensive groundwater infuence on the Krakovski gozd, Cigonca and dobrava plots, the similarity in Figure 3 is shown. due to differences in site and climate conditions, the third dCA axis segregates the Bojanci plots from the Bukovnica plots. The Panovec and Pišece plots obtain low scores along the dCA1 axis (Figs. 2 and 3). resulting from similar local climate conditions, they lie close together in ordination space. Although they originate from different parts, their foristic composition shows quite similar site conditions. due to relative humidity and cold soils on the fysch of Panovec plots, as well due to moderate northern exposure of plots, the effect of the warm mediterranean climate is not so pronounced. Ho- Kutnar, L.: Plant diversity of selected Quercus robur L. and Quercus petraea (Matt.) Liebl. forests in Slovenia 1 wever, the Pišece plots are situated on very warm southern slopes in the Pre-Pannonian region. In the Polom region, the Q. robur acorns were an important nutrition source for domestic pigs in the past (SmoLEj 1995). Thus, in this region the forest stands dominated by the Q. robur are man-made. The oak trees are growing under conditions in which groundwater has no direct infuence. Potentially, the Polom region is site of mixed forest of Q. petraea and Fagus sylvatica. Consequently, the understorey vegetation of the Polom plots is more similar to that on plots overgrown with Q. petraea forests than with Q. robur forests. Therefore, the Polom plots lie quite closer to Q. petraea plots in ordination space (Figs. 2 and 3). Vectors of Q. robur and of Q. petraea show an increase in the growing stock of these two tree species (Figs. 2 and 3). Based only on the foristic composition of understorey layers, the differentiation between Q. robur and Q. petraea dominated plots has been confrmed. The understorey vegetation has indicated very specifc site conditions of plots. Besides similar foristic composition, the Cigonca and dobrava plots have signifcant shares of Picea abies in the tree layer (Figs. 2 and 3). In last decades, the Norway spruce has been favoured by forest management. The signifcant positive correlation was found between the frst dCA axis and the growing stock of Picea abies (Table 1). Apart from these tree species, in dCA analysis (Fig. 3) only the vector of the growing stock of Fagus sylvatica is shown. The growing stock of beech was found to increase towards the Bukovnica plots. Among the nine common tree species that have been analysed (Table 1), the weak negative correlation between the dCA2 axis and Carpinus betulus growing stock, and between the dCA2 and Acer campestre growing stock is also shown. The number of different tree species increases slightly towards plots in lower parts of the dCA ordination space (Fig. 2, Table 1). In the understorey vegetation layers, a total of 256 species was recorded. on average, 31 plant species per plot were n""5 A. \ ' Do,mci IIUDO I-PO fj :Vec/^TncG Axis 1 Fig. 2: dCA analysis of research plots and vectors of tree layer parameters (Axis 1 vs. Axis 2) Slika 2: DCA-analiza raziskovalnih ploskev in vektorji parametrov drevesne plasti (os 1: os 2) Legend. Q. robur complexes : Po-Polom (I), Kg-Krakovski gozd (II), do-dobrava (III), CI-Cigonca (IV), Hr-Hraščica (V); Q. petraea complexes: PA-Panovec (1), Bo-Bojanci (2), PI-Pišece (3), BU-Bukovnica (4); gS-Q. robur (-Q. petraea, -Picea abies) - growing stock of Q. robur (Q. petraea, Picea abies) Legenda. Kompleks doba Q. robur: PO-Polom (I), Kg-Krakovski gozd (II), DO-Dobrava (III), CI-Cigonca (IV), hR-hraščica (V); Kompleks gradna Q. petraea: PA-Panovec (1), BO-Bojanci (2), PI-Pišece (3), Bu-Bukovnica (); gS-Q. robur (-Q. petraea, -Picea abies) – lesna zaloga Q. robur (Q. petraea, Picea abies) 2 Zbornik gozdarstva in lesarstva, 79 found. The number of species per plot ranges between 4 and 70. on all 225 research plots, Carpinus betulus is the most frequent plant species of understorey vegetation that is present on 173 research plots (Appendix 1). other very common plant species are Acer campestre (129 plots) and Prunus avium (128 plots). In the understorey vegetation, the dominant tree species Quercus robur (113 plots) and Quercus petraea (94 plots) are also frequent. other common woody plant species are Corylus avellana (109), Ligustrum vulgare L. (90), Crataegus monogyna jacq. (86), and Euonymus europaea L. (83). The herb species with high frequencies are Anemone nemorosa L. (124), Athyrium flix-femina (L.) roth (116), Polygonatummultiforum (L.) All. (111), Violareichenbachi-ana jord. ex Boreau (94), Carex brizoides L. (90), galium sylvaticum L. (85), and Ajuga reptans L. (80). Species richness was found to increase towards the Polom plots (Fig. 4), where the mean number of species in the understorey layers is 62 (Appendix 2). on average, a high number of species has been found on the Pišece plots (49 species/ plot), and on the Krakovski gozd plots (45 species/plot). on the dobrava and on Cigonca plots, we found very few species (9 and 12 species/plot). due to the high number of species, and due to the relative balance of their cover, the Polom, Pišece, and Krakovski gozd plots have high diversity indexes (H’ and d), and the evenness index (E) is also high on these plots (Appendix 2). due to the predominance of Corylus avellana in the shrub layer of the Polom plots, evenness is slightly lower on these plots than on the plots of other two complexes. The mean species cover increases towards the dobrava and Cigonca plots (Fig. 4), with positive correlation with the frst dCA axis (Table 1). Besides the low number of species, the majority of these plots are completely overgrown by Carex brizoides. The sum of all species cover is high on the Polom and Krakovski gozd plots. These are plots with a high number of species in the herb layer, and in the shrub layer as well. CONCLUSIONS zaključki research of oak forest ecosystems in Slovenia (SmoLEj / HAgEr 1995) was supposed to explain most of the predominating factors (e.g. drought, frost, air pollution) in the process of oak decline, and was a contribution to the wider 200 Axis 3 QS-Fagus sylvatica EL> cwJJI-DO *aQ21 DOD02 DOS\B0ri Axis 1 Fig. 3: dCA analysis of research plots and vectors of tree layer parameters (Axis 1 vs. Axis 3) Slika 3: DCA-analiza raziskovalnih ploskev in vektorji parametrov drevesne plasti (os 1: os 3) Kutnar, L.: Plant diversity of selected Quercus robur L. and Quercus petraea (Matt.) Liebl. forests in Slovenia Axis 2 ,-K#aS ~+BU 2-BQ p024 ' |,n. BjgjSjauZil BUI? BUlJ\ buaB017B02l y BUS x Of08223.»^ BUI« ,B025 )f fa2BJ23 BU19 V-Tffl / *1 HB15 ..-.JX . *ds8 cc5 mean species covets -A ^9 III-DO Axis 1 Fig. 4: dCA analysis of research plots and parameters of understorey vegetation (Axis 1 vs. Axis 2) Slika : DCA-analiza raziskovalnih ploskev in parametri pritalnih plasti vegetacije (os 1: os 2) Legend. Q. robur complexes: Po-Polom (I), Kg-Krakovski gozd (II), do-dobrava (III), CI-Cigonca (IV), Hr-Hraščica (V); Q. petraea complexes: PA-Panovec (1), Bo-Bojanci (2), PI-Pišece (3), BU-Bukovnica (4); E - Evenness index, H’ - Shannon diversity index, d - Simpson diversity index. Legenda. Kompleks doba Q. robur: PO-Polom (I), Kg-Krakovski gozd (II), DO-Dobrava (III), CI-Cigonca (IV), hR-hraščica (V); Kompleks gradna Q. petraea: PA-Panovec (1), BO-Bojanci (2), PI-Pišece (3), Bu-Bukovnica (); E – indeks poravnanosti; h’ – Shannonov indeks pestrosti, D – Simpsonov indeks pestrosti. ecological understanding of oaks and oak sites in Slovenia as a part of the wider area of oak forest communities in Europe. In the present study, the biodiversity and vegetation structure of oak forests in Slovenia have been analysed. The importance of conserving biodiversity for a sustainable future is widely accepted. Areas of remnant native vegetation support a range of biodiversity. our attention has been focused on species diversity in relation to forest stand structure. With the ordination technique, the diversity of the vegetation structure of different oak forests was obtained. The ordination has differentiated clearly between plots dominated by Q. robur, and those with Q. petraea. due to specifc site conditions and human infuences, the differences in biodiversity of understorey vegetation and tree layer parameters have been shown as well. With the ordination of oak forest based on the foristic composition, the main ecological gradients were also indicated (e.g. soil-moisture gradient). Warmer oak sites (Panovec and Pišece plots) are clearly separate from the other oak forests of mesic conditions. The plant species diversity proved to be a valuable indicator of the site conditions and of forest management in the past as well. Although heavy anthropogenic impacts have been present in the past, a variety of oak forests still exists in Slovenia. The Quercus robur and Quercus petraea dominated forests create a wide range of different types with heterogeneous vegetation structure. In the studied oak forests, a high number of tree-layer species and high species richness of understorey layers have been recorded. But in some cases, such as dobrava and Cigonca plots, forest management practices in the past could be one of the reasons for signifcant biodiversity impoverishment. Forest management practices often have a major impact on biodiversity, causing changes to site conditions, tree species composition and forest structure (mITCHELL / KIr-BY 1989). 3 44 Zbornik gozdarstva in lesarstva, 79 Table 1: Spearman rank correlations between dCA scores and 1) tree layer parameters, 2) understorey vegetation para- meters. Preglednica 1: Spearmanova korelacija rangov med DCA-koordinatami in 1) parametri drevesne plasti, 2) parametri pritalnih plasti vegetacije Spearman r dCA1 -0,259 *** dCA2 0,006 / dC -0,024 A3 1) NUmBEr oF TrEES (STEmS) / ŠTEVILO DREVES / NUmBEr oF TrEE SPECIES / ŠTEVILO DREVESnIh VRST 0,010 / -0,487 *** 0,298 *** ToTAL groWINg SToCK (gS) / CELOTnA LESnA ZALOgA (LZ) 0,367 *** -0,026 / -0,047 / gS / Lz - Quercus robur 0,770 *** -0,408 *** -0,133 * gS / LZ - Quercus petraea -0,694 *** 0,547 *** 0,069 / gS / LZ - Quercus cerris -0,367 *** -0,221 *** 0,122 / gS / Lz - Carpinus betulus 0,264 *** -0,516 *** -0,106 / gS / LZ - Acer campestre 0,001 / -0,421 *** 0,014 / gS / LZ - Alnus glutinosa 0,307 *** -0,167 * 0,132 * gS / LZ - Fagus sylvatica -0,295 *** 0,342 *** 0,443 *** gS / LZ - Picea abies 0,707 *** 0,112 / 0,022 / gS / LZ - Tilia cordata -0,154 -0,617 * *** -0,323 -0,602 *** *** 0,082 0,060 / 2) SPECIES rICHNESS (S) / VRSTnO BOgASTVO (S) / SUm oF SPECIES CoVEr / SEŠTEVEK ZASTIRAnJA VSEh VRST -0,299 *** -0,461 *** -0,154 * mEAN SPECIES CoVEr /POVPREČnA STOPnJA ZASTIRAnJA nA VRSTO 0,614 *** 0,143 * -0,395 *** EVENNESS (E) /InDEKS PORAVnAnOSTI (E) -0,571 *** -0,164 * 0,276 *** SHANNoN dIVErSITY INdEX (H’) / ShAnnOnOV DIVERZITETnI InDEKS ( h’) -0,627 *** -0,506 *** 0,130 / SImPSoN dIVErSITY INdEX (d) / SIMPSOnOV DIVERZITETnI InDEKS (D) -0,617 *** -0,422 *** 0,142 * SUMMARY POVZeTek Velika biodiverziteta v povezavi z raznolikostjo gozdnih rastišč je ena izmed značilnosti nižinskih hrastovih gozdov v Evropi. Po drugi strani pa so ti gozdovi zaradi neposrednih človekovih vplivov tudi med najbolj ogroženimi in spremenjenimi. Nižinski poplavni gozdovi in še posebej hrastovi gozdovi so bili vseskozi pod močnimi antropogenimi pritiski (KLImo / HAgEr 2001). Pet avtohtonih vrst hrasta (Quercus robur agg., Q. petra-ea agg., Q. cerris, Q. pubescens in Q. ilex) v Sloveniji sestavlja manj kot 8 % celotne lesne zaloge (SmoLEj / HAgEr, 1995). Večina hrastov uspeva v nižinah in gričevnatem obrobju, kjer je delež gozdov razmeroma majhen, gostota prebivalstva pa največja. Biodiverziteta ima poseben pomen pri trajnostnemu gospodarjenju z gozdom. Vrstna diverziteta in heterogena struktura vegetacije sta pomembni za stabilno funkcioniranje gozdnih ekosistemov. Namen raziskave je bil ugotoviti vrstno diverziteto in glavne strukturne gradiente vegetacije izbranih hrastovih gozdov v Sloveniji. V okviru raziskave smo analizirali sedanje stanje vegetacije, ki je rezultat dolgotrajne prilagoditve vegetacije na rastišče (vegetacija kot indikator rastiščnih razmer) in hkrati rezultat sprememb zaradi človekovega delovanja (vegetacija kot indikator gospodarjenja z gozdom). V različnih delih Slovenije smo izbrali devet gozdnih kompleksov (SmoLEj 1995). Vsi so v kategoriji gospodarskih gozdov. Večina od petih izbranih dobovih kompleksov (I. Polom, II. Krakovski gozd, III. dobrava, IV. Cigonca, V. Hraščica) leži v vzhodnem delu Slovenije (slika 1). štirje gra- Kutnar, L.: Plant diversity of selected Quercus robur L. and Quercus petraea (Matt.) Liebl. forests in Slovenia dnovi kompleksi (1. Panovec, 2. Bojanci, 3. Pišece, 4. Bukov-nica) pa so bolj raztreseni. raziskovalni kompleksi velikosti 100×100 metrov so bili razdeljeni na 25 kvadratnih (20×20 m) ploskev (SmoLEj 1995). V študiji smo posebej obravnavali drevesno plast na eni strani in pritalne plasti vegetacije (grmovna, zeliščna in mahovna) na drugi strani. V drevesni plasti so bili zajeti vsi osebki, katerih prsni premer je presegal 10 centimetrov. Na osnovi predhodne raziskave (AzAroV 1995) smo ugotovili lesno zalogo (Lz) po ploskvah in ocenili Lz najpogostejših drevesnih vrst Quercus robur, Quercus petraea, Carpinus be-tulus, Fagus sylvatica, Picea abies, Acer campestre, Quercus cerris, Alnus glutinosa in Tilia cordata. Poleg tega smo ugotovili število dreves (osebkov) in število različnih drevesnih vrst na vseh 225 ploskvah. Vegetacijo pritalnih plasti smo popisali po standardni srednjeevropski metodi (BrAUN-BLANQUET 1964). Na tej osnovi smo ocenili stopnjo zastiranja vrst v grmovni, zeliščni in mahovni plasti (vključeni samo na tleh rastoči mahovi). V popis in oceno pritalnih plasti vegetacije smo vključili tudi drevesa, katerih prsni premer še ne presega 10 centimetrov. Na osnovi popisa pritalnih plasti vegetacije smo za vsako ploskev izračunali vrstno bogastvo, seštevek zastiranja vseh vrst, povprečno stopnjo zastiranja na vrsto, indeks poravna-nosti (E), Shannonov diverzitetni indeks (H’) in Simpsonov diverzitetni indeks (d). glavne strukturne gradiente smo analizirali z ordinacijo dCA (Detrended Correspondence Analysis) (HILL / gAU-CH 1980), ki je temeljila na foristični sestavi raziskovalnih ploskev. Analiza je bila napravljena z računalniškim paketom PC-ord (mcCUNE / mEFFord 1999). Izračunali smo tudi Spearmanovo korelacijo rangov med dCA-koordinatami in: a) parametri drevesne plasti; b) parametri pritalnih plasti vegetacije. Na 225 raziskovalnih ploskvah smo ugotovili 30 drevesnih in grmovnih vrst s prsnim premerom nad 10 centimetrov. Najpogostejše vrste v drevesni plasti (v oklepaju je število ploskev, na katerih se pojavljajo) so Quercus robur (124 ploskev), Quercus petraea (100), Carpinus betulus (90), Fagus sylvatica (56), Picea abies (50), Acer campestre (42), Quercus cerris (18), Alnus glutinosa (16) in Tilia cordata (16). druge manj pogoste drevesne in grmovne vrste s prsnim premerom nad 10 centimetrov so (vrstni red v skladu s frekvenco pojavljanja dreves po ploskvah): Pinus nigra, Pyrus pyraster, Sorbus torminalis, Prunus avium, Crataegus mono-gyna, Pinus sylvestris, Ostrya carpinifolia, Fraxinus ornus, Larix decidua, Populus tremula, Acer pseudoplatanus, Abies alba, Sorbus aria, Fraxinus excelsior, Betula pendula, Acer platanoides, ulmus glabra, Crataegus laevigata, ulmus la-evis, Corylus avellana in Ilex aquifolium. Na vseh ploskvah smo evidentirali 4003 dreves (grmov), ki so presegali merski prag. število dreves (grmov) na ploskev se giblje med 4 in 35 (povprečno skoraj 18). ocena Lz na ploskev se giblje med 5 in 49 m3. ordinacija dCA je samo na osnovi foristične sestave (prisotnost/odsotnost) pritalnih plasti vegetacije jasno ločila komplekse s prevladujočim dobom (pod diagonalo na slikah 2 in 3) od kompleksov s prevladujočim gradnom (nad diagonalo). zaradi foristične podobnosti se ploskve dobovih kompleksov dobrava in Cigonca, ki se pojavljajo na močvirnih, oglejenih tleh kot rezultat vpliva visoke podtalnice, v ordinaciji nahajajo skupaj (slika 2 in 3). razmeroma podobna sta si tudi dobova kompleksa Hraščica in Krakovski gozd (slika 2), vendar pa se določene razlike pokažejo s tretjo ordinacij-sko osjo (slika 3). razlike v foristični sestavi lahko v veliki meri povežemo z razlikami v tleh. V kompleksu Hraščica prevladujejo evtrična rjava tla, medtem ko so tla v Krakovskem gozdu neprimerno bolj izpostavljena procesom oglejevanja zaradi vpliva podtalnice (KALAN 1995). gradnova kompleksa Panovec in Pišece sta si v primerjavi z drugimi gradnovi kompleksi najbolj podobna, čeprav ležita v povsem različnih delih Slovenije (sliki 2 in 3). Podobnost je posledica razmeroma podobnih mezo-rastiščnih razmer. Pri prvem vpliv toplega mediteranskega podnebja ne prihaja do izraza zaradi razmeroma hladnih in vlažnejših fišnih tal ter zmerne severne ekspozicije. drugi pa leži na toplih južnih legah v predpanonskem območju. razlike v foristični sestavi med gradnovima kompleksoma Bojanci in Bukovnica, ki so posledica rastiščno-podnebnih razlik, se pokažejo s tretjo or-dinacijsko osjo (slika 3). dobov kompleks Polom je bil osnovan na potencialnih rastiščih mešanih gozdov gradna in bukve. dob so pospeševali zaradi želoda, ki je bil v preteklosti pomemben vir prehrane 6 Zbornik gozdarstva in lesarstva, 79 domačih prašičev (SmoLEj 1995). Pritalne plasti vegetacije kompleksa Polom kažejo na večjo foristično podobnost z gradnovimi kot dobovimi kompleksi. delitev med gradnovimi in dobovimi kompleksi samo na osnovi foristične sestave (prisotnost/odsotnost) pritalnih plasti vegetacije potrjujejo tudi vektorji naraščanja lesne zaloge glavnih dveh drevesnih vrst, doba in gradna. Vegetacija nakazuje specifčne razlike v rastiščnih razmerah dobovih in gradnovih gozdov. Podobnost v rastiščnih razmerah ploskev Cigonca in dobrava se kaže v foristični sestavi. Hkrati pa je podobna sestava vegetacije lahko deloma tudi posledica podobnega načina gospodarjenja v preteklosti, ki je močno pospeševal smreko, kar nakazuje tudi vektor naraščanja lesne zaloge smreke proti tema kompleksoma. V pritalnih plasteh vegetacije smo našli 256 rastlinskih vrst. število vrst na ploskev se giblje med 4 in 70 (povprečno 31 vrst/ploskev). V pritalnih plasteh smo na največjem številu ploskev našli navadni gaber Carpinus betulus (173 ploskev). druge pogoste lesnate rastline v pritalnih plasteh so maklen Acer campestre (129 ploskev), češnja Prunus avium (128), dob Quercus robur (113), navadna leska Corylus avellana (109) in graden Quercus petraea (94). zeliščne vrste z veliko frekvenco pojavljanja so podlesna vetrnica Anemone nemorosa (124), navadna podborka Athyri-um flix-femina (116), mnogocvetni salomonov pečat Polygo-natum multiforum (111), gozdna vijolica Viola reichenbachi-ana (94), migalični šaš Carex brizoides (90), gozdna lakota galium sylvaticum (85) in plazeči skrečnik Ajuga reptans (80). Vektor vrstnega bogastva je usmerjen proti kompleksu Polom, kjer smo v pritalnih plasteh vegetacije povprečno popisali 62 vrst na ploskev. Veliko število vrst smo našli tudi na ploskvah kompleksov Pišece (49 vrst/ploskev) in Krakovski gozd (45 vrst/ploskev). število vrst je razmeroma majhno na ploskvah kompleksov Cigonca (12 vrst/ploskev) in dobrava (9 vrst/ploskev). glede na veliko število vrst in njihovo medsebojno uravnoteženost smo za ploskve kompleksov Polom, Pišece in Krakovski gozd izračunali tudi velike vrednosti diverzitetnih indeksov (H’ in d) ter indeksa poravnanosti (E). Slednji je sicer nekoliko manjši na ploskvah Poloma, kjer v grmovni plasti močno prevladuje leska. Povprečna stopnja zastiranja vrst na ploskev izrazito narašča proti kompleksoma dobrava in Cigonca z značilnim majhnim številom vrst in močno prevladujočim migaličnim šašem. V raziskavi smo dali poudarek biodiverziteti in vegetacijski strukturi izbranih hrastovih gozdov, ki sta pomembna elementa trajnostnega razvoja gozdnih ekosistemov S pomočjo ordinacijskih tehnik smo nakazali razlike med gozdovi doba in gradna ter nakazali glavne ekološke gradiente. Pestrost pritalne vegetacije v povezavi s sestojnimi parametri (vertikalna struktura) se je pokazala kot dober indikator specifčnih rastiščnih razmer in tudi gospodarjenja z gozdom v preteklosti. Kljub močnim antropogenim pritiskom na hrastove gozdove v preteklosti in sedanjosti lahko pri nas še vedno najdemo razmeroma pestre gozdove doba in gradna z razgibano vegetacijsko strukturo. zanje je na splošno značilno veliko število drevesnih vrst in veliko bogastvo vrst v pritalnih plasteh vegetacije. ACKNOwLEDgEMENTS ZahVala Very special thanks must go to my colleague Ivan Smole who performed the basic work on the feld vegetation survey and the determination of plant species. Especially at the initial stages of this research, he has given me important lessons and assistance. Thanks are due to Professor Franc Batič who provided the determination of some problematic plant species. Thanks also go to Igor Smolej, Evgenij Azarov-Stjopa and janko Kalan for all their useful comments and help at the beginning of this study. I wish to thank to all other collaborators from the Slovenian Forestry Institute for their feld and technical assistance. Thanks also to the referees whose criticism has considerably improved an earlier version of the manuscript. The English language of the manuscript was kindly checked by Terry T. jackson. REfERENCES VIRI ACCETTo, m., 1974. združbi gabra in evropske gomoljčnice (Pseudostellario-Carpinetum) ter doba in evropske gomoljčnice (Pseudostellario-Quercetum) v Krakovskem gozdu = Communities of hornbeam and Pseudostellaria europaea (Pseudostellario-Carpinetum), and of pedunculate oak and Pseudostellaria europaea (Pseudostellario-Quercetum) in Krakovski gozd.- gozdV 32, 10: 357-369. Kutnar, L.: Plant diversity of selected Quercus robur L. and Quercus petraea (Matt.) Liebl. forests in Slovenia 7 AzAroV, E., 1995. Qualitative and quantitative characteristics of oaks on permanent research plots.- In: SmoLEj, I. / HAgEr, H. (eds.). oak decline in Slovenia: Endbericht über die Arbeiten 1995. gozdarski inštitut Slovenije, Ljubljana, Institut für Waldökologie, Wien: 26-34. BATIČ, F. / SINKoVIČ, T. / jAVorNIK, B., 1995. morphological and genetic variability of pedunculate oak (Quercus robur L.) populations in Slovenia.- zb. gozd. lesar. 46: 75-96. BATIČ, F. / mAVSAr, r. / SINKoVIČ, T. / KrALj, A., 1997. morfološka variabilnost populacij doba (Quercus robur L.) v Sloveniji.- Acta biol. Slov 41, 2-3: 127-140. BrAUN-BLANQUET, j., 1964. Pfanzensoziologie. grundzüge der Vegetationskunde.- Springer, Wien, New York. 865 s. BrEzNIKAr, A. / KUmP, B. / CSAIKL, U. / BATIČ, F. / KrAIgHEr, H., 2000. Taxonomy and genetics of chosen oak populations in Slovenia.-glas. šumske pokuse 37: 361-373. ČATEr, m. / SImoNČIČ, P. / BATIČ, F., 1999. Pre-dawn water potential and nutritional status of pedunculate oak (Quercus robur L.) in the northeast of Slovenia.- Phyton (Horn) 39, fasc. 4 (Spec. issue: “Eurosilva”): 13-22. ČATEr, m. / BATIČ, F., 1999. Nekateri ekofziološki kazalci stresa pri dobu (Quercus robur L.) v severovzhodni Sloveniji = Some ecophysiological stress indicators of pedunculate oak (Quercus robur L.) in the north easteren of Slovenia.- zb. gozd. lesar. 58: 47-83. ČATEr, m. / BATIČ, F., 2000. Ecophysiological parameters as a possible tool for the assessment of natural and artifcial regeneration in pedunculate oak in lowland parts of Slovenia.- glas. šumske pokuse 37: 201-213. ČATEr, m. / KUTNAr, L. / ACCETTo, m., 2001. Slovenian lowland and foodplain forests.- In: KLImo, E. / HAgEr, H. (eds.). The foodplain forests in Europe: current situation and perspectives. European Forest Institute research report, 10. Leiden; Boston; Köln: Brill.: 233-248. ČATEr, m. / LEVANIČ, T, 2004. Increment and environmental conditions in two Slovenian pedunculate oak forest complexes.- Ekológia, Bratislava 23: 353-365. dAKSKoBLEr, I., 1997. Fitocenološka oznaka sestojev črnega hrasta Quercus ilex L. na Sabotinu in nad izvirom Lijaka (zahodna Slovenija) = Phytosociological characteristics of holm oak Quercus ilex L. stands on mount Sabotin and above the source of the Lijak river (Western Slovenia).- Acta biol. Slov. 41, 2-3: 19-42. HILL, m. o. / gAUCH, H. g., 1980. detrended correspondence analysis, an improved ordination technique.- Vegetatio 42: 47-58. KALAN, j., 1995. Basic soil analyses.- In: SmoLEj, I. / HAgEr, H. (eds.). oak decline in Slovenia: Endbericht über die Arbeiten 1995. gozdarski inštitut Slovenije, Ljubljana, Institut für Waldökologie, Wien.: 43-46. KLImo, E. / HAgEr, H, 2001. Executive Summary.- In: KLImo, E. / HAgEr, H. (eds.) The Floodplain Forests in Europe: Current Situation and Perspectives. European Forest Institute research report 10, Brill, Leiden, Boston, Köln: VI-XI. KUTNAr, L., 1997. Primerjava metod vrednotenja okoljskih razmer gozdnih ekosistemov na osnovi ftoindikacije = A comparison of evaluation methods regarding environmental conditions of forest ecosystems on the basis of phytoindication.- zb. gozd. lesar. 54: 5-44. KUTNAr, L. / zUPANČIČ, m. / roBIČ, d. / zUPANČIČ, N / žITNIK, S. / KrALj, T. / TAVČAr, I. / doLINAr, m. / zrNEC, C. / KrAIgHEr, H., 2002. razmejitev provenienčnih območij gozdnih drevesnih vrst v Sloveniji na osnovi ekoloških regij = The delimitation of the regions of provenance of forest tree species in Slovenia based on ecological regions.- zb. gozd. lesar. 67: 73-117. mArTINČIČ, A. / WrABEr, T. / jogAN, N. / rAVNIK, V. / PodoBNIK, A. / TUrK, B. / VrEš, B., 1999. mala fora Slovenije - Ključ za določanje praprotnic in semenk = Flora of Slovenia - key for determination of ferns and vascular plants.- Tehniška založba Slovenije, Ljubljana, 845 s. mArTINČIČ, A., 2003. Seznam listnatih mahov (Bryopsida) Slovenije.-Hacquetia 2: 91-166. mcCUNE, B. / mEFFord, m. j., 1999. PC-ord: multivariate Analysis of Ecological data, Version 4.0.- mjm Software design,glenden Beach, oregon, 237 s. mITCHELL, P. L. / KIrBY, K. j., 1989. Ecological effects of forestry practices in long established woodland and their implications for nature conservation.- oxford Forestry Institute, oxford Forestry Institute occasional Papers 39, 172 s. OKLANd, T. / BAKKESTUEN, V. / OKLANd, r.H. / EILErTSEN, o., 2004. Changes in forest understorey vegetation in Norway related to long-term soil acidifcation and climatic change.- journal of Vegetation Science15: 437-448. PoUNdS, j. A. / PUSCHENdorF, r., 2004. Ecology: Clouded futures.-Nature, 427: 107-109. PUNCEr, I. / zUPANČIČ, m., 1979. Novi združbi gradna v Sloveniji (melampyro vulgati-Quercetum petraeae ass. nova s. lat.) = Two new associations of durmast oak in Slovenia (Melampyro vulgati-Quercetum petraeae ass. nova s. lat.).- Scopolia 2: 1-47. rogL, S. / jAVorNIK, B. / SINKoVIČ, T. / BATIČ, F., 1996. Characterization of oak (Quercus L.) seed proteins by electrophoresis.-V: KrAIgHEr, H / BATIČ, F. / gUTTENBErgEr, H. / grILL, d. / AgErEr, r. / HANKE, d. (eds), Bioindication of stress in forest trees and forest ecosystems, Phyton (Horn) 36, fasc. 3: 159-162. SCHULEr, A., 1998. Sustainability and biodiversity - forest historical notes on two main concerns of environmental utilisation.- In: BACHmANN, P. / KÖHL, m. / PÄIVINEN, r. (eds.), Assessment of Biodiversity for Improved Forest Planning. Kluwer Academic Publishers, dordrecht: 353-360. SmoLE, I. / BATIČ, F., 1992. The importance of morphological characteristics for identifcation of oak species.- zb. gozd. lesar. 39: 133-172. SmoLE, I., 1993. Vegetacijske in rastiščne razmere na trajnih raziskovalnih ploskvah hrasta v Sloveniji : I. del: Krakovski gozd, Cigonca, Hraščica, Bojanci, Polom = Vegetation and site conditions of the permanent research plots of oak in Slovenia, part I: Krakovski gozd, Cigonca, Hraščica, Bojanci, Polom.- gozdarski inštitut Slovenije, Ljubljana, 86 s. SmoLE, I., 1995. Vegetations- und Standortsverhältnisse der Ständigen Versuchs-fächen in den Eichenwäldern Sloweniens.- In: SmoLEj, I. / HAgEr, H. (eds.). oak decline in Slovenia: Endbericht über die Arbeiten 1995. gozdarski inštitut Slovenije, Ljubljana, Institut für Waldökologie, Wien.: 47-59 SmoLE, I. / KUTNAr, L., 1994a. Vegetacijske in rastiščne razmere na trajnih raziskovalnih ploskvah hrasta v Sloveniji : II. del: Panovec, dobrava, Bukovnica, Pišece) = Vegetation and site conditions of the permanent research plots of oak in Slovenia, part II: Panovec, dobrava, Bukovnica, Pišece.- gozdarski inštitut Slovenije, Ljubljana, 56 s. SmoLE, I. / KUTNAr, L., 1994b. Vegetacijske in rastiščne razmere na trajnih raziskovalnih ploskvah : (III. del: Povzetek I. in II. dela naloge) = Vegetation and site conditions of the permanent research plots of oak in Slovenia, part III: summaries of parts I and II.- gozdarski inštitut Slovenije, Ljubljana, 50 s. SmoLEj, I., 1995. Permanent research plots.- In: SmoLEj, I. / HAgEr, H. (eds.). oak decline in Slovenia: Endbericht über die Arbeiten 1995. gozdarski inštitut Slovenije, Ljubljana, Institut für Waldökologie, Wien.: 11-14 SmoLEj, I. / HAgEr, H. (eds.) 1995. oak decline in Slovenia: Endbericht über die Arbeiten 1995. gozdarski inštitut Slovenije, Ljubljana, Institut für Waldökologie, Wien, 99 s. SWINdEL, B. F. / CoNdE, L. F. / SmITH j. E., 1984. Species diversity: concept, measurement, and response to clearcutting and site preparation.-Forest Ecology and management 8: 11-22. šUgAr, I. / zUPANČIČ, m. / TrINAjSTIĆ, I. / PUNCEr, I., 1995. Forets thermophiles de chene pubescent et de molinie (Molinio-Quercetum pubescentis šugar 1981) dans la zone limitrophe de Croatie et de Slovénie = Termoflni gozd puhovca in navadne stožke (Molinio- 8 Zbornik gozdarstva in lesarstva, 79 Quercetum pubescentis šugar 1981) na obmejnem območju Hrvaške in Slovenije.- Biol. vestn. 40, 3-4: 113-124. TrAjBEr, d. / BrEzNIKAr, A. / SINKoVIČ, T. / BATIČ, F., 2001. Ugotavljanje križancev doba (Quercus robur L.) in gradna (Quercus petraea (matt.) Liebl.) z morfološko analizo listov = Assessment of common (Quercus robur L.) and sessile oak (Quercus petraea (matt.) Liebl.) hybrids by morphological leaf analyses.- Hladnikia 12-13: 167-175. zUPANČIČ, m., 1997. (Sub)mediteranski forni element v gozdni vegetaciji submediteranskega fornega območja Slovenije = (Sub)mediterranean foral element in the forest vegetation of the Submediterranean foric area of Slovenia.- razpr.- Slov. akad. znan. umet., razr. naravosl. vede, Ljubljana 38: 257-298. Appendix 1: Species of understorey vegetation (shrub, herb and moss layer) - presence per complexes (in the table only 163 species with presence on 10 or more research plots are shown; the total number of plant species is 256). Priloga 1: Vrste pritalnihplasti vegetacije (grmovna, zeliščna in mahovnaplast) -pojavljanje vrste po kompleksih (vpreglednici je samo 163 vrst, ki se pojavljajo na 10 ali več raziskovalnih ploskvah; celotno število vrst je 26). Legend: Q. robur complexes: Po-Polom (I), Kg-Krakovski gozd (II), do-dobrava (III), CI-Cigonca (IV), Hr-Hraščica (V); Q. petraea complexes: PA-Panovec (1), Bo-Bojanci (2), PI-Pišece (3), BU-Bukovnica (4). Legenda: komplex doba Q. robur: Po-Polom (I), Kg-Krakovski gozd (II), do-dobrava (III), CI-Cigonca (IV), Hr-Hraščica (V); komplex gradna Q. petraea complexes: PA-Panovec (1), Bo-Bojanci (2), PI-Pišece (3), BU-Bukovnica (4). PO kG Do cI hr PA Bo PI BU sum sum sum I II III IV V 1 2 3 4 I-V 1-4 All N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=125 N=100 N=225 1 Carpinus betulus L. 25 22 2 22 25 24 12 22 19 96 77 173 2 Acer campestre L. 24 24 0 0 25 24 7 25 0 73 56 129 3 Prunus avium L. 18 11 1 0 16 24 25 25 8 46 82 128 4 Anemone nemorosa L. 16 25 1 10 25 17 17 1 12 77 47 124 5 Athyrium flix-femina (L.) roth 19 25 15 23 11 0 17 0 6 93 23 116 6 Quercus robur L. 25 22 16 25 25 0 0 0 0 113 0 113 7 Polygonatum multiforum (L.) All. 25 25 1 7 17 2 14 0 20 75 36 111 8 Corylus avellana L. 25 25 5 0 13 2 25 14 0 68 41 109 9 Quercus petraea (matt.) Liebl. 0 0 0 0 0 25 25 21 23 0 94 94 10 Viola reichenbachiana jordan ex Boreau 21 12 3 0 24 20 2 10 2 60 34 94 11 Carex brizoides L. 0 25 25 25 15 0 0 0 0 90 0 90 12 Ligustrum vulgare L. 25 0 0 0 18 13 8 25 1 43 47 90 13 Crataegus monogyna jacq. 25 0 0 0 0 25 11 25 0 25 61 86 14 galium sylvaticum L. 11 0 0 0 0 25 25 21 3 11 74 85 15 Euonymus europaea L. 25 20 1 0 23 8 0 6 0 69 14 83 16 Ajuga reptans L. 22 10 5 0 22 8 1 6 6 59 21 80 17 Polytrichum formosum Hedw. 13 0 8 11 0 6 6 7 22 32 41 73 18 Sorbus torminalis (L.) Crantz 12 0 0 0 0 19 17 25 0 12 61 73 19 Fagus sylvatica L. 9 0 1 0 0 1 12 24 25 10 62 72 20 rosa arvensis Huds. 19 4 0 0 0 24 0 25 0 23 49 72 21 Brachypodium sylvaticum (Huds.) P.Beauv. 13 6 0 0 0 21 5 25 0 19 51 70 22 Pulmonaria offcinalis L. 21 0 0 0 24 3 0 22 0 45 25 70 23 Hieracium umbellatum L. 17 0 0 0 1 18 0 16 17 18 51 69 24 galium odoratum (L.) Scop. 0 21 0 0 0 0 9 19 17 21 45 66 25 Carex sylvatica Huds. 11 9 0 0 14 9 7 11 4 34 31 65 26 Sanicula europaea L. 25 0 0 0 5 1 9 24 1 30 35 65 27 Cruciata glabra (L.) Ehrend. 25 3 0 0 0 15 11 10 0 28 36 64 28 Fraxinus ornus L. 0 0 0 0 0 25 13 25 0 0 63 63 29 galeopsis speciosa mill. 1 1 9 12 21 0 17 0 2 44 19 63 30 Cornus mas L. 21 22 0 0 0 0 19 0 0 43 19 62 31 Euphorbia dulcis L. 9 11 0 0 0 21 0 20 0 20 41 61 32 Viburnum opulus L. 21 8 0 0 3 5 13 11 0 32 29 61 33 dryopteris carthusiana (Vill.) H.P.Fuchs 2 3 24 23 1 1 0 0 6 53 7 60 34 rubus hirtus W. & K. 0 0 0 1 3 14 25 17 0 4 56 60 Kutnar, L.: Plant diversity of selected Quercus robur L. and Quercus petraea (Matt.) Liebl. forests in Slovenia PO kG Do cI hr PA Bo PI BU sum sum sum I II III IV V 1 2 3 4 I-V 1-4 All N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=125 N=100 N=225 35 Symphytum tuberosum L. 15 17 0 0 0 3 9 14 2 32 28 60 36 Fragaria moschata (duchesne) Weston 25 0 0 0 0 10 0 23 0 25 33 58 37 Lonicera caprifolium L. 0 0 2 0 20 25 0 11 0 22 36 58 38 Primula vulgaris Huds. 25 0 0 0 0 6 6 21 0 25 33 58 39 Tilia cordata mill. 25 0 0 0 0 1 24 7 0 25 32 57 40 Pyrus pyraster (L.) Burgsd. 21 2 0 0 1 5 2 25 0 24 32 56 41 Serratula tinctoria L. 0 0 0 0 1 25 5 25 0 1 55 56 42 Acer pseudoplatanus L. 8 0 0 0 5 3 14 25 0 13 42 55 43 deschampsia cespitosa (L.) P.Beauv. 0 3 22 19 9 1 0 0 1 53 2 55 44 Salvia glutinosa L. 25 0 0 0 0 4 2 24 0 25 30 55 45 Solidago virgaurea L. 24 0 0 0 0 10 0 21 0 24 31 55 46 galeobdolon montanum (Pers.) Pers. ex rchb. 21 25 0 0 8 0 0 0 0 54 0 54 47 Festuca heterophylla Lam. 24 0 1 0 0 10 1 17 0 25 28 53 48 Pteridium aquilinum (L.) Kuhn 8 4 0 0 0 2 25 0 14 12 41 53 49 Circaea lutetiana L. 0 25 0 0 20 0 7 0 0 45 7 52 50 Anemone ranuculoides L. 2 18 0 0 25 1 4 0 0 45 5 50 51 Carex facca Schreber 12 0 0 0 0 13 0 25 0 12 38 50 52 Scrophularia nodosa L. 10 1 7 2 0 0 21 0 7 20 28 48 53 Frangula alnus mill. 4 0 10 14 0 3 9 0 7 28 19 47 54 gentiana asclepiadea L. 15 6 3 0 0 0 23 0 0 24 23 47 55 Picea abies (L.) Karsten 1 0 25 21 0 0 0 0 0 47 0 47 56 ranunculus fcaria L. 0 21 0 0 25 0 0 0 0 46 0 46 57 Prunus spinosa L. 14 0 0 0 0 7 4 21 0 14 32 46 58 Hedera helix L. 0 0 0 0 0 24 0 21 0 0 45 45 59 Lamium orvala L. 18 0 0 0 0 0 18 8 0 18 26 44 60 mercurialis perennis L. 23 20 0 0 0 0 0 0 0 43 0 43 61 molinia arundinacea Schrank 0 0 0 4 0 18 18 1 0 4 37 41 62 Cardamine bulbifera (L.) Crantz 0 25 0 0 0 0 2 13 0 25 15 40 63 melampyrum pratense L. 0 0 0 0 0 2 23 15 0 0 40 40 64 Viburnum lantana L. 15 0 0 0 0 2 0 22 0 15 24 39 65 Crataegus laevigata (Poir.) dC. 0 25 0 0 13 0 0 0 0 38 0 38 66 Lathyrus niger (L.) Bernh. 0 0 0 0 0 21 0 17 0 0 38 38 67 Aegopodium podagraria L. 3 24 0 0 6 0 4 0 0 33 4 37 68 Tamus communis L. 0 0 0 0 0 1 12 24 0 0 37 37 69 Luzula pilosa (L.) Willd. 10 0 1 0 0 18 4 0 3 11 25 36 70 Clematis vitalba L. 7 0 0 0 0 0 3 25 0 7 28 35 71 Heracleum sphondylium L. 21 4 0 0 6 1 0 3 0 31 4 35 72 Hieracium sylvaticum (L.) L. 5 0 0 0 4 9 1 13 3 9 26 35 73 rubus caesius L. 19 16 0 0 0 0 0 0 0 35 0 35 74 Aposeris foetida (L.) Less. 0 10 0 0 0 0 0 24 0 10 24 34 75 Calamagrostis arundinacea (L.) roth 0 0 0 0 0 25 0 9 0 0 34 34 76 daphne mezereum L. 23 10 0 0 0 0 0 0 1 33 1 34 77 dryopteris flix-mas (L.) Schott 9 8 0 0 0 1 12 1 3 17 17 34 78 melittis melissophyllum L. 0 0 0 0 0 8 5 21 0 0 34 34 79 Paris quadrifolia L. 22 12 0 0 0 0 0 0 0 34 0 34 80 Aruncus dioicus (Walter) Fernald 25 0 0 0 0 0 7 0 0 25 7 32 81 Cornus sanguinea L. 0 0 0 0 0 7 0 25 0 0 32 32 82 Cyclamen purpurascens mill. 25 0 0 0 0 0 0 6 0 25 6 31 83 Campanula trachelium L. 24 0 0 0 0 0 0 6 0 24 6 30 84 Cephalanthera longifolia (L.) Fritsch 6 0 0 0 0 11 0 2 11 6 24 30 85 Convallaria majalis L. 4 0 0 0 0 0 21 5 0 4 26 30 86 dactylis glomerata L. 1 0 0 0 5 13 0 10 0 6 23 29 87 melampyrum nemorosum L. 24 5 0 0 0 0 0 0 0 29 0 29 9 Zbornik gozdarstva in lesarstva, 79 PO kG Do cI hr PA Bo PI BU sum sum sum I II III IV V 1 2 3 4 I-V 1-4 All N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=125 N=100 N=225 88 oxalis acetosella L. 11 3 4 8 3 0 0 0 0 29 0 29 89 Aremonia agrimonoides (L.) dC. 20 0 0 0 0 0 8 0 0 20 8 28 90 rubus fruticosus agg. 0 20 0 0 0 0 0 0 8 20 8 28 91 Carex montana L. 25 0 0 0 0 0 0 2 0 25 2 27 92 gagea lutea (L.) Ker-gawl. 0 21 0 0 6 0 0 0 0 27 0 27 93 Scilla bifolia L. 0 21 0 0 6 0 0 0 0 27 0 27 94 Vicia oroboides Wulf. 0 0 0 0 0 3 0 21 3 0 27 27 95 Betonica offcinalis L. 25 0 1 0 0 0 0 0 0 26 0 26 96 Ilex aquifolium L. 0 0 0 0 1 25 0 0 0 1 25 26 97 Luzula luzuloides (Lam.) dandy & Wilm. 0 0 0 0 0 24 0 1 1 0 26 26 98 Vinca minor L. 0 0 0 0 0 25 0 0 1 0 26 26 99 dryopteris dilatata (Hoffm.) A.gray 0 0 0 25 0 0 0 0 0 25 0 25 100 Epimedium alpinum L. 0 0 0 0 0 0 25 0 0 0 25 25 101 omphalodes verna moench 25 0 0 0 0 0 0 0 0 25 0 25 102 ruscus aculeatus L. 0 0 0 0 0 25 0 0 0 0 25 25 103 Sesleria autumnalis (Scop.) F.W.Schultz 0 0 0 0 0 25 0 0 0 0 25 25 104 Asarum europaeum L. 0 12 0 0 0 0 0 12 0 12 12 24 105 Berberis vulgaris L. 24 0 0 0 0 0 0 0 0 24 0 24 106 Crocus napolitanus mord. & Loisel. 0 20 1 3 0 0 0 0 0 24 0 24 107 Impatiens noli-tangere L. 0 24 0 0 0 0 0 0 0 24 0 24 108 Leucojum vernum L. 0 22 0 0 2 0 0 0 0 24 0 24 109 Pulmonaria dacica Simonk. 0 24 0 0 0 0 0 0 0 24 0 24 110 Carex pilosa Scop. 0 11 0 0 0 12 0 0 0 11 12 23 111 glechoma hederacea L. 0 17 0 0 6 0 0 0 0 23 0 23 112 Brachypodium rupestre (Host) roem. & Schult. 22 0 0 0 0 0 0 0 0 22 0 22 113 Carex digitata L. 0 0 0 0 0 20 0 0 2 0 22 22 114 Lonicera xylosteum L. 22 0 0 0 0 0 0 0 0 22 0 22 115 Ulmus glabra Huds. 0 0 0 0 0 0 0 22 0 0 22 22 116 Cardamine trifolia L. 0 21 0 0 0 0 0 0 0 21 0 21 117 Lamium maculatum L. 0 19 0 0 1 0 1 0 0 20 1 21 118 Alliaria petiolata (mB.) Cav. & grande 0 20 0 0 0 0 0 0 0 20 0 20 119 mycelis muralis (L.) dum. 8 0 0 0 1 0 5 0 6 9 11 20 120 Asplenium trichomanes L. 17 0 0 0 0 0 0 2 0 17 2 19 121 Stellaria holostea L. 0 7 0 5 6 0 0 0 1 18 1 19 122 Festuca gigantea (L.) Vill. 4 2 0 0 0 0 12 0 0 6 12 18 123 Peucedanum venetum (Sprengel) Koch 18 0 0 0 0 0 0 0 0 18 0 18 124 ranunculus nemorosus dC. 18 0 0 0 0 0 0 0 0 18 0 18 125 Arum maculatum L. 0 17 0 0 0 0 0 0 0 17 0 17 126 Cardamine impatiens L. 0 17 0 0 0 0 0 0 0 17 0 17 127 galium aparine L. 0 14 0 0 3 0 0 0 0 17 0 17 128 Inula spiraeifolia L. 17 0 0 0 0 0 0 0 0 17 0 17 129 Pulmonaria mollissima Kerner 0 0 4 0 0 0 0 13 0 4 13 17 130 Veronica hederifolia L. 0 9 0 0 8 0 0 0 0 17 0 17 131 Cardamine amara L. 0 16 0 0 0 0 0 0 0 16 0 16 132 Fragaria vesca L. 0 0 0 0 0 0 16 0 0 0 16 16 133 geum urbanum L. 0 16 0 0 0 0 0 0 0 16 0 16 134 Isopyrum thalictroides L. 1 15 0 0 0 0 0 0 0 16 0 16 135 melica unifora retz. 7 1 0 0 0 5 3 0 0 8 8 16 136 Peucedanum oreoselinum (L.) moench 15 0 0 0 0 0 0 1 0 15 1 16 137 Senecio fuchsii C.C.gmelin 12 0 0 0 0 0 0 4 0 12 4 16 138 Solanum dulcamara L. 14 0 0 2 0 0 0 0 0 16 0 16 139 Urtica dioica L. 0 13 0 0 3 0 0 0 0 16 0 16 140 Veronica persica Poir. 0 8 0 0 8 0 0 0 0 16 0 16 0 Kutnar, L.: Plant diversity of selected Quercus robur L. and Quercus petraea (Matt.) Liebl. forests in Slovenia 1 PO kG Do cI hr PA Bo PI BU sum sum sum I II III IV V 1 2 3 4 I-V 1-4 All N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=25 N=125 N=100 N=225 141 Abies alba mill. 1 0 2 0 0 12 0 0 0 3 12 15 142 Actaea spicata L. 8 0 0 0 0 0 7 0 0 8 7 15 143 Buglossoides purpurocaerulea (L.) I.m.johnston 0 0 0 0 0 0 0 15 0 0 15 15 144 Carex umbrosa Host 0 0 0 0 0 0 0 0 15 0 15 15 145 Hacquetia epipactis (Scop.) dC. 3 11 0 0 0 0 0 1 0 14 1 15 146 Knautia drymeia Heuff. 3 0 0 0 11 0 0 1 0 14 1 15 147 moehringia trinervia (L.) Clairv. 0 0 0 0 11 0 4 0 0 11 4 15 148 Potentilla erecta (L.) räuschel 13 0 0 0 0 2 0 0 0 13 2 15 149 Pseudostellaria europaea Schaeftlein 0 14 0 1 0 0 0 0 0 15 0 15 150 ranunculus lanuginosus L. 7 7 0 0 0 0 0 0 0 14 0 14 151 Campanula persicifolia L. 12 0 0 0 0 0 0 1 0 12 1 13 152 Sorbus aria (L.) Crantz 0 0 0 0 0 1 0 12 0 0 13 13 153 Taraxacum offcinale F.Weber in Wiggers 9 0 0 0 0 0 0 2 2 9 4 13 154 Carex pendula Huds. 0 12 0 0 0 0 0 0 0 12 0 12 155 juniperus communis L. 0 0 0 0 0 6 0 5 1 0 12 12 156 Angelica sylvestris L. 8 2 0 1 0 0 0 0 0 11 0 11 157 Chamaecytisus hirsutus (L.) Link 1 0 0 0 0 2 1 7 0 1 10 11 158 Chrysosplenium alternifolium L. 0 11 0 0 0 0 0 0 0 11 0 11 159 Iris graminea L. 0 0 0 0 0 0 0 11 0 0 11 11 160 Agrostis canina L. 0 0 0 10 0 0 0 0 0 10 0 10 161 Clinopodium vulgare L. 0 0 0 0 0 0 0 10 0 0 10 10 162 Platanthera bifolia (L.) L.C.rich. 1 0 0 0 0 1 2 4 2 1 9 10 163 ranunculus auricomus L. 0 9 0 0 1 0 0 0 0 10 0 10 2 Zbornik gozdarstva in lesarstva, 79 Appendix 2: mean values per plot of nine oak dominated forest complexes in Slovenia Priloga 2: Povprečne vrednosti po devetih kompleksih hrasta v Sloveniji krAkoVSkI „ POLOM „ DoBrAVA cIGoNcA HRAŠČICA GozD PANOVEC BoJANcI PIŠECE BUKOVNICA LABEL OF COMPLEX / OZNAKA KOMPLEKSA ABBrEVIAtIoN / OKRAJŠAVA NUMBER OF PLOTS / ŠTEVILO PLOSKEV I II III IV V PO kG Do cI hr N=25 N=25 N=25 N=25 N=25 123 4 PA Bo PI BU N=25 N=25 N=25 N=25 ELEVATIoN (m) / nADMORSKA VIŠInA (m) INCLINATIoN (°) / POVP. nAKLOn( °) LoNgITUdE / gEOg. DOLŽInA LATITUdE / gEOg. ŠIRInA 370 150 160 260 180 10 0 0 0 0 5489640 5532690 5550925 5545165 5598240 5067090 5082090 5088981 5135670 5167490 140 280 470 230 5 15 25 10 5397650 5521290 5551040 5601740 5090244 5038915 5096765 5173740 TREE LAyER / DREVESNA PLAST NUmBEr oF TrEES (STEmS) / ŠTEVILO DREVES NUmBEr oF TrEE SPECIES / ŠTEVILO DREVESnIh VRST ToTAL groWINg SToCK (in m3) / CELOTnA LESnA ZALOgA (LZ) GroWING Stock (in m3) OF / LESnA ZALOgA (LZ): Quercus robur Quercus petraea Quercus cerris Carpinus betulus Acer campestre Alnus glutinosa Fagus sylvatica Picea abies Tilia cordata 17 12 25 14 17 3.2 2.9 2.0 2.8 2.4 12 22 27 23 22 9.3 16.4 23.8 18.3 18.5 / / /// / / /// 0.8 4.4 / 0.6 3.4 / 0.4 / / 0.1 / 0.6 / 0.2 / / / /// / / 3.4 3.4 / 1.5 / / / / 26 12 21 17 2.0 1.6 3.8 2.4 12 19 24 16 /// / 11.7 18.9 18.3 13.5 / / 3.9 / / / / 0.1 / / 0.1 / /// / / 0.2 1.1 2.3 /// / / 0.1 / / UNDERSTOREy VEGETATION LAyERS / PRITALNE PLASTI VEGETACIJE SPECIES rICHNESS / VRSTnA PESTROST (BOgASTVO) SUm oF SPECIES CoVEr (%) / SEŠTEVEK ZASTIRAnJA VSEh VRST mEAN SPECIES CoVEr (%) / POVPREČnA STOPnJA ZASTIRAnJA nA VRSTO EVENNESS(E) / InDEKS PORA VnAnOSTI SHANNoN (H’) / ShAnnOnOV InDEKS PESTROSTI SImPSoN (d) / SIMPSOnOV InDEKS PESTROSTI 62 45 9 12 23 182 178 74 105 114 3 4 10 9 5 0.79 0.81 0.46 0.37 0.70 3.26 3.07 0.99 0.94 2.17 0.92 0.92 0.43 0.37 0.79 37 31 49 12 130 154 124 27 453 3 0.80 0.72 0.82 0.77 2.89 2.47 3.18 1.89 0.90 0.84 0.92 0.72