GDK: 23:174.7 Abies alba Mill.(045)=111 Prispelo / Recived: 17. 08. 2009 Sprejeto / Accepted: 30. 11. 2009 Izvirni znanstveni članek Original scientific paper STAND DIVERSITY IN THE DINARIC FIR-BEECH FORESTS Milan KOBAL1, David HLADNIK2 Abstract Data from forest research and permanent sampling plots were used to quantify stand structure and forest diversity in uneven-aged Dinaric silver fir and beech forest at the level of forest management unit. The study examined the indicators that are expected to be used in the monitoring of biodiversity and the favourable conservation status of forest habitat types in Natura 2000 sites. By comparing the present and past stand structures, it was possible to establish the developmental dynamics of the stands and the changes of structure indices of the forest stands on the 2-hectare research plots in the last 50 years. We adopted the indicators for the assessment of biotic and stand structure diversity based on tree species composition, stand densities, mean DBH and coefficient of variation, species and DBH-based Shannon's index, spatial mingling and size differentiation of trees on the permanent sampling plots. The indicators confirmed stand structural diversity in four forest management classes of uneven-aged Dinaric fir-beech forests in the Leskova dolina (valley) of the Sneznik-Javorniki high Karst region. Continuous forest inventory and the presented indicators of stand structure and diversity may potentially represent the framework for the monitoring of the favourable conservation status of forest habitat types. Key words: stand structure, stand diversity monitoring, indicators, uneven-aged fir and beech stands, Abies alba Mill., Fagus sylvatica L. SESTOJNA RAZNOVRSTNOST V DINARSKIH JELOVO-BUKOVIH GOZDOVIH Izvleček Na podlagi podatkov gozdarskih raziskovalnih ploskev in stalnih vzorčnih ploskev smo ocenili zgradbo in pestrost v dinarskih jelovo-bukovih gozdovih na ravni gozdnogospodarske enote. Predstavljeni so kazalniki, ki bi jih lahko uporabili za ocenjevanje raznovrstnosti in ugodnega ohranitvenega stanja gozdnih habitatnih tipov na območjih Natura 2000. Na podlagi primerjave današnje in nekdanje sestojne zgradbe na 2 hektarskih raziskovalnih ploskvah je bilo mogoče oceniti razvojno dinamiko in spremembe strukturnih znakov gozdnih sestojev v zadnjih 50 letih. Na stalnih vzorčnih ploskvah smo uporabili kazalnike za ocenjevanje biotske in strukturne raznovrstnosti gozdnih sestojev na podlagi sestave drevesnih vrst, sestojnih gostot, povprečnih premerov in koeficientov variacije za premere dreves, Shannonovega indeksa za vrstno pestrost in debelinsko strukturo dreves, prostorsko razmestitev mešanosti in dominantnosti drevesnih vrst. S kazalniki smo potrdili razlike v zgradbi gozdnih sestojev štirih gozdnogospodarskih razredov, ki obsegajo raznodobne dinarske jelovo-bukove gozdove v Leskovi dolini na območju Snežniško-Javorniškega masiva. Stalne vzorčne ploskve in predstavljeni kazalniki sestojne zgradbe in raznovrstnosti bi lahko postali izhodišče za monitoring ugodnega ohranitvenega stanja gozdnih habitatnih tipov. Ključne besede: sestojna zgradba, monitoring raznovrstnosti gozdnih sestojev, kazalniki, raznodobni jelovo-bukovi gozdovi, Abies alba Mill., Fagus sylvatica L. UVOD INTRODUCTION The assessment of structural diversity of forest stands has been important in forest management for a long time - initially for predicting future stand growth, the resistance against storms, snow loads of stands and the effectiveness as a protection against other natural hazards. Stand diversity has become an important topic within the discussion of sustaina-bility, biological diversity, namely alpha diversity that refers to the ecosystem diversity - the diversity within forest stands in forest ecosystems. Many authors have suggested and tested different measures of forest structural diversity, which can be characterized with respect to tree species diversity, tree size diversity and spatial diversity (Pommerening, 2002; Aguir-re et al., 2003; Varga et al., 2005; McRoberts et al., 2008). Cantarello and Newton (2008) suggested that the indicators of forest biodiversity, developed in order to support the assessment of sustainable forest management, may be of value for the assessment of the conservation status of forested habitats in the Natura 2000 sites. Forests represent more than a half of the Natura 2000 sites in the network representing one of the main actions for biodiversity conservation in Europe. In Slovenia, the Natura 2000 forest habitat types represent more than 260,000 ha of forests and it was estimated that the Natura 2000 sites encompass approximately one half of all the forests (Golob, 2006). Although the general guidance on the conservation status assessment has been provided (EC, 2005), 1 M. K., univ. dipl. inž. gozd., Gozdarski inštitut Slovenije, Večna pot 2, 1000 Ljubljana, milan.kobal@gozdis.si 2 doc. dr. D. H., UL, Biotehniška fakulteta, Oddelek za gozdarstvo in obnovljive gozdne vire, Večna pot 2, 1000 Ljubljana, david.hladnik@bf.uni-lj.si a common standard for the implementation of monitoring of the Natura 2000 sites and forest habitat types has not yet been created (Cantarello and Newton, 2008). In the large-area forest structural biodiversity assessments, the national forest inventories or other permanent monitoring plots (for example EU/ICP Forests programme) provide the most comprehensive and geographically extensive tree species and size diversity data. Unfortunately, the sampling intensity in regional inventories is typically too low and the plots are too small to adequately characterise individual stands (Koehl et al., 1995). These data are not sufficient to determine the habitat and species diversity within landscape and forest compartment structures, or to investigate the causal factors such as land use change or forest management that affect the patterns of structural diversity and contribute to them. As proposed by Pommerening (2002), the quantification of stand structure has the advantage of being easier to survey by means of common forest inventories than more direct measures of diversity. Apart from using algorithms and concepts to relate the forest stand structure to the habitat functions, they can also be used to inform forest managers about the consequences of silvicultural activities. Even though the results presented in forest management plans provide us with important information on the effectiveness of silvicultural treatments, long term permanent observation is very important in order to understand the forest dynamics and to establish a monitoring strategy, appropriate for the assessment of the effectiveness of measures maintaining or improving a favourable conservation status of the forests. In order to characterise the stand structure of forests, several parameters are needed. Forest scientists have developed a number of indices for quantification of different aspects of stand structure including small scale differences within stands. As has been shown by Fischer and Pommerening (2003), these indices require different data input, which can practically be provided by different inventory types. McElhinny et al. (2005) reviewed literature concerning forest and woodland structure at the scale of individual stands. As a group, the attributes identified describe stand structure in terms of foliage arrangement, canopy cover, tree diameter, tree height, tree spacing, tree species, stand biomass, understorey vegetation, and deadwood. The review indicates that there is no definite suite of structural attributes; different authors emphasised the subset of different attributes, and relatively few studies provided quantitative evidence linking attributes to the provision of faunal habitat or other measures of biodiversity, although a number of studies identified attributes that distinguish between successive stages. Neumann and Starlinger (2001) used the data from permanent observation plots, established within the frame of the Pan-European Programme for Intensive and Continuous Monitoring of Forest Ecosystems, to quantify stand structure and species diversity by means of different indices (species richness, evenness, diversity, complexity, nonrandomness, clumping, aggregation, neighbouring pattern). Each of the plots where the mapping of the stands has been carried out comprised an area of 0.25 ha. A comparison of different indices has shown that there exist only few weak correlations between species diversity of different vegetation layers and between plant species diversity and the diversity of stand structure, while among all the indices of stand structure a correlation exists. Bachofen and Zingg (2001) have shown that even for the assessment of silvicultural treatment several parameters are needed. From the results of the first survey of subalpine Norway spruce forests they concluded that the variables h/d ratio, spatial distribution and the stand density index (SDI) were the best indicators of silvicultural treatments. They stressed that one survey is not sufficient to understand the forest dynamics; long-term permanent observation is very important and long-term commitments are needed. Based on the review of scientific literature and recommendations of international research projects for monitoring forest biodiversity and sustainable forest management, Canta-rello and Newton (2008) selected 17 indicators for monitoring the favourable conservation status of the forested habitats in the Natura 2000 sites. Most of these indicators were also harmonized by the representatives of the national forest inventories of 27 European countries and the USA participating in COST Action E43 »Harmonization of the National forest inventories in Europe« (Winter et al., 2008). In this project, the biodiversity variables were ranked relative to their utility for the national forest inventory based biodiversity assessment. The most important and feasible variables were deadwood, stand-level development phases and forest types, horizontal and vertical structures, microsites, vegetation naturalness, tree age and diameter, abundance of old trees, species of trees and shrubs, variables related to forest management systems and disturbances. Similar to biodiversity and to sustainable forest management monitoring in the National forest inventories, the first reports of monitoring the conservation status of the habitats have been prepared at the national scale. The common standard for implementation of monitoring the Na- tura 2000 sites and forest habitat types has not been created and the current results suggest that flexibility may be required to select the most appropriate indicators for individual sites (Cantarello and Newton, 2008). During the argumentation of financially acceptable indicators for the evaluation of the favourable conservation status of forest habitats it was estimated that the scientific literature does not yet provide information on the indicators usable in the monitoring of forest habitat types at the level of site or stand (Cantarello in Newton, 2008). Thus, the researchers determine the biodiversity and the condition of forest habitat types indirectly. During the process, the most frequently used key indicators at the level of forest stands were the following: number of trees (ha-1), basal area of stands (m2/ha), mean diameter of trees (cm), standard deviation of diameters and coefficient of variation (CV), Shannon's index for the assessment of tree species diversity and stand structural diversity, percentage of big trees in forest stands, mean height of trees (m), volume of snags and downed dead wood (m3/ha). The objective of this study was to test the proposed variables for describing forest structural diversity at the level of forest stands, based on the continuous forest inventory in forest management unit. We adopted the indicators for the assessment of stand structure diversity, developed within the framework of national forest inventories and forest monitoring projects in the European countries. The methods for characterisation of stand structure in the Dinaric fir-beech uneven-aged forests were used to assess to what extend were the data available within the Slovenian control sampling method suitable for calculating measures of stand diversity and differences between stand structures. We hypothesized that: • permanent sample plots can be used for assessing the stands characteristics of forest habitat type at the local scale, • the measurements of the stand diversity indicators vary between different forest management classes, which encompass forest management unit in the Dinaric fir-beech forests. The objective did not include investigations of causal factors such as silvicultural measures or disturbances that affect stand structural diversity. The study examined the indicators that are expected to be used in the monitoring of biodiversity and the favourable conservation status of forest habitat types in Natura 2000 sites. Although it was estimated that most of the indicators relevant for the assessment of the conservation status and monitoring are already in use in the forest management planning system in Slovenia (Golob, 2006), the reference values of stand structures and the variation within and between different indicators of forest stand structure were not tested in different forest habitat types as a part of proposed integral monitoring, incorporated into the planning process. The reference values from the investigations of stand structure performed up to now on research plots (Kotar, 2006; Hladnik and Skvarča, 2008) proved to be a good starting point for the assessment of forest habitat types and for the comparison with data and information gained through the continuous forest inventory in Slovenia. The research plots, established 60 years ago in the uneven-aged Dinaric beech-fir forest stands to study the optimal values of growing stock levels and the structure of forests, were used to estimate changes of stand diversity in periods of time, comparable with 10-year periods in forest management planning. The aim of this part of the study was to compare the indicators of stand structure diversity on the research plots in the past 50 years, during which the structure of the forests stands in the former selection forests has changed gradually. MATERIALS AND METHODS MATERIAL IN METODE STUDY AREA AND DATA SOURCES RAZISKOVALNO OBMOČJE IN VIRI PODATkOV The indicators for the assessment of stand structures were tested in the Leskova valley, which represents a forest management unit and a part of Special Protection Area Sne-žnik-Pivka, included also in the habitat types of Illyrian beech forests. The forest management unit encompasses 3,011 ha of forests on the northeast slopes of the Dinaric mountain massive Snežnik. In Dinaric silver fir and beech forests on the high Karst with high diversity of natural conditions, uneven-aged forest stands are prevalent. They developed from a mature forest with old beech and silver fir trees and rich fir regeneration in the middle of the 19th century (Bončina et al., 2003). After the period of unsuppressed growth of fir and selection forest treatment, favouring selective harvesting of single trees or small groups of trees, the natural regeneration of fir was disturbed by red deer. These processes, the fir decline and uneven-aged forest management in the last 40 years have created a great variability of forest structures. Table 1: General characteristics of forest management classes in Dinaric fir-beech forests in Leskova valley (Forest management plan; Gozdnogospodarski načrt ..., 2007) Preglednica 1: Osnovne značilnosti gozdnogospodarskih razredovjelovo-bukovih gozdov v Leskovi dolini (Gozdnogospodarski načrt..., 2007) Characteristics Omphalodo-Fagetum lycopodietosum homogynetosum mercurialetosum typicum Area (ha, percentage) 300.98 (10.0 %) 769.43 (25.6 %) 252.23 (8.4 %) 1120.82 (37.2 %) Development phase or stand structure Large timber 48.3 % Regeneration 41.9% Uneven-aged 97.2 % Uneven-aged 96.1 % Uneven-aged 96.0 % Average growing stock (m3/ha) 524 422 412 463 Percentage of growing stock by 20 cm DBH classes (%) 4 / 8 / 8 / 17 / 63 4 / 8 / 15 / 24 / 49 4 / 10 / 15 / 21 / 50 4 / 7 / 13 / 19 / 57 Annual volume increment (m3/ha) 11.2 8.4 8.3 9.8 Number of tree species 6 9 11 8 Number of sample plots 56 151 49 215 Stand diversity was assessed in four forest management classes, which encompass fir and beech forests in the altitude range between 750 and 1,250 m. These classes were named after the prevalent vegetation syntaxa of this part of Dinaric fir-beech forests: Omphalodo- Fagetum (Treg. 57) Mar et al. 93 (syn. Abieti-Fagetum dinaricum (Treg. 57) em. Punc. 79); OF-typicum, OF-homogynetosum, OF-mercurialetosum, OF-Lycopodietosum (Gozdnogospodarski načrt ..., 2007). METHODS METODE The study includes 471 permanent sample plots on 250 x 200 m large systematic sampling grid, established by Slovenian Forest Service (Table 1). A permanent sample plot consists of two concentric plots of 200 m2 and 500 m2. All trees with the DBH of more than 10 cm were measured within the inner circular plot, whereas those with the DBH of more than 30 cm were measured within a circular plot of 500 m2. The position of each tree on a sample plot was determined by the azimuth and the distance from the plot centre that enabled the calculation of indices of the spatial structure of trees in forest stands. Based on the concept of the tree factor or expansion factor, stand densities and stand diversity indices were calculated for sample plots, measured in 2003. In the same year, the measurement of trees on the research plots in Leskova valley and in a neighbouring forest management unit Mašun was finally repeated after 50 years of observations (Hladnik, 2004b). In 1950, 2-hectare-research plots were set up in silver fir and beech forests to study the optimal values of growing stock levels and the structure of selection forests on those plots. The assessment of stand densities (the number of trees, the basal area, the stand volume) is one of the key elements in forest inventories and the data gathered on the sampling plots represent the basis for estimation of several indicators and proxy variables. On the permanent sampling plots, the statistic parameters are calculated based on the concept of the tree factor. Measurements on sample plots are summarized and expressed on a per unit area basis - per hectare for the metric system. The expansion factor referred to as the tree factor (Husch et al., 2003) was also suggested to harmonize the data from different sampling units in the COST Action E43. The Shannon index (H') was intended for the assessment of species diversity, but it was used by many authors for the assessment of structural diversity of forest stands as well (Varga et al., 2005; McRoberts et al., 2008), with p. representing the proportion of the basal area of trees in the DBH class compared with the total basal area: H' = - Sp^ ln(p) The coefficient of variation (CV) as a measure of differences between individual trees as regards their mean diameters was used to assess tree-size diversity in forest inventories or to estimate the structure of even-aged stands on the research plots (Kotar 1991, 2006). The reference values from previous researches of the stand structure in Slovenia are a good starting point for the assessment of forest habitat types and for the comparison with the data and information on the control sampling method in Slovenia. The methods for the assessment of the spatial structure of forest stands based on the coefficient of variation for the assessment of the spacing between the trees have already been presented in the previous researches (Puhek, 1998; Hladnik, 2004a). However, on the permanent sampling plots it is possible to estimate the differences in the species structure and in the position of trees regarding the size of the neighbouring trees as well (Pommerening, 2002; Agui-rre et al., 2003). Aguirre et al. (2003) presented a method for comparing the spatial structure of different natural forest sites using a new approach for describing complex forest structures in a straightforward manner. They described the structure of forest stands by the spatial distribution of tree positions, by the spatial mingling of different tree species and by the spatial arrangement of tree dimensions, size differentiation and contagion. The spatial characteristics of stands have been established merely by evaluating the immediate neighbourhood of the given number of reference trees. M. = iLvj v, = 1, neighbour j belongs o the same species a reference tree i 0, otherwise 0 < M.^1 The parameters M (mingling) or U. (differentiation) can have 5 values: 0.00 - all the neighbouring trees are of other tree species or are thicker than the reference tree, 0.25 - one of the neighbouring trees is of the same tree species or is thinner than the reference tree, 0.50 - two neighbours are of the same tree species or are thinner than the reference tree, 0.75 - three neighbours are of the same tree species as the reference tree, 1.00 - all five trees (the reference tree and the four neighbours) are of the same tree species or all four neighbouring trees are thinner than the reference tree. On the sampling plots, the relations between the neighbouring trees with mean diameters of less than 30 cm, measured only on 200 m2 subplots, could not be estimated in the same way as those between thicker trees, measured on the 500 m2 plots. The radius of 7.98 m on the sampling plots was too small to enable the analysis of spatial arrangement for reference fir, beech and spruce trees and their 4 neighbouring trees within a 200 m2 plot. The estimation of mingling distribution of thinner firs would be especially useful, since according to the forest management plan (Gozdnogospodarski načrt ..., 2007) it is necessary to support these firs and to enable them the in-growth into the mature stands. However, a representative assessment could not be carried out, because the calculation could comprise only the data for the fir trees growing on less than a half of the sampling plots. Statistical analyses and computing were performed using R (R Foundation for Statistical Computing). Descriptive statistics was employed in computing mean values, variances and standard errors. ANOVA was used in establishing differences between indices of species and DBH-based diversity for forest management classes. Frequency distributions of cut and survival trees by 20 cm DBH classes were compared using Brandt-Snedecor test for the 10- year periods of measurements on research plots. The present structure of forest stands in the area of four forest management classes was compared to the data of forest management plan from the year 1954. In this forest management plan, the data of total tallies of trees in forest compartments for individual tree species were shown for the first time. In the environment of geographic information systems (GIS), we were able to align the spatial data from old forest maps with present strata of forest management classes, owing the fact that the borders of forest departments remained unchanged and that at the level of forest compartments, only small changes were observed in the past as well. RESULTS REZULTATI The three research plots represent a part of development processes taking place in fir and beech forests for the last 50 years and the share of starting points for the comparison with other fir and beech forests in this area. In the beginning of the observation period 55 years ago, the fir was prevalent in the majority of forest compartments encompassing four forest management classes of fir and beech forests in the area of the Leskova valley nowadays. Only in 4 of the 123 forest compartments, measured at that time with full callipering, the beech or the spruce was prevalent. In the group of compartments ranged nowadays in the forest management class Omphalodo-Fagetum typicum, the share of the fir on the basal area was 76%, in OF-lycopodietosum group 71%, in OF-mer-curialetosum 96% and in OF homogynetosum 64%. The values of the basal area of stands on the research plots were on the increase from the beginning of the 1970s onwards. In the ensuing period, characterized as the period of silver fir decline, however, the basal areas diminished until in the end of the 1980s the amount of firs in the basal area came to a stan- Table 2: Stand densities and diversity indices for three research plots in uneven-aged forest stands of fir and beech in 2003 Preglednica 2: Sestojne gostote in kazalniki sestojne raznovrstnosti za tri raziskovalne ploskve v raznodobnih jelovo-bukovih gozdovih v letu 2003 Characteristics Plot 97 Plot 98 Plot 99 Number of trees (ha-1) 382 217 413 Basal area (m2/ha) 49.4 38.1 48.9 Growing stock (m3/ha) 718 571 700 Big trees DBH>70 cm (% BA) 5 33 22 Shannon index (H') based on 5 cm classes 2.37 2.64 2.70 20 cm classes 1.07 1.23 1.30 species diversity 0.63 0.89 0.73 Mean diameter Fir Spruce Beech Fir Spruce Beech Fir Spruce Beech Quadratic (cm) 37.2 45.0 19.8 54.8 37.6 42.2 44.6 40.3 24.6 Arithmetic (cm) 32.2 42.3 18.8 49.5 33.0 39.0 39.7 34.6 22.0 Coefficient of variation (DBH) 58 36 33 48 55 41 51 59 50 dstill or even increased (Fig. 1). In the course of development of stands, illustrated by quadratic mean changes, the beech gained significance on the plot 98. In 50 years, its share on the basal area has increased from 30% to 45%, whereas the share of the fir declined from 67% to 50%. On this research plot, the beech grew even into thicker DBH classes. In the year 1954, the share of the beech in the basal area of the DBH class of more than 50 cm was 8.6 %, while 50 years later it was 36.2%. After 1971, the cutting of thinner trees in the lowest DBH class of 10-30 cm contributed to more significant changes of quadratic means of the fir, the beech and the spruce (Fig. 1). Especially noticeable are the changes of quadratic means of the beech and the spruce on the plot 98 after 1988. Using the Brandt-Snedecor test, we were able to confirm that after 1971 the frequency distributions of the cut trees typically differed from the distributions of other trees regarding the DBH classes. Until 1988, the cuttings of firs in the lowest DBH class (Plot 99, P<0.001; plot 97, P<0.05), spruce (Plot 97, P<0.001) and beech (Plot 98, P<0.05) were prevalent. During the regeneration of fir and beech forests, thinner trees did not preserve the role prescribed to them in the former selection forest treatment. In last decades, the recruitment rate of these trees into higher DBH classes has declined. Being of bad quality, the beeches in the lowest DBH class that used to be prevalent in the structure of the cutting on the plot 98 even after 1988 (P<0.001) were cut before the regeneration of stands. 86% of the beeches cut at that time were in the 10-30 cm DBH class. On other research plots, the beech remained in the lower layer. On the plot dominated by the spruce (Plot 97), the be- ech did not overgrow the quadratic mean of 50 cm, while on the plot 99 its share in this DBH class declined from 4.5% in the year 1951 to 2.3% fifty years later. On both research plots, the share of the fir in the stand basal area declined by 5% in a period of 50 years. After the last measurements in 2003, the regeneration of stands has begun in the area of the research plots as well. On the permanent sampling plots measured in the year 2003, the prevalent share of the fir could still be estimated in the stand basal area of four forest management classes (Fig. 2). Its share declined most visibly in the forest management class OF-lycopodietosum (24%) on account of the share of the spruce in the stand basal area that was estimated to be two times bigger than 50 years ago. In other forest management classes, the average value of fir basal area was greater than the basal area of other tree species (P<0,001), although until the year 2003, the share of the spruce increased also in OF-homogynetosum class (by 9%), in OF- mercurialetosum (7%) and in OF- typicum (6%). The share of the beech in the stand basal area of uneven-aged stands increased most significantly (by 9%) in the forest management class OF- typicum. The differences between the stands were estimated indirectly, based on the quadratic means of individual tree species and on the coefficients of variation for these means. Our estimations have shown that while the average diameters of the fir are at the biggest, its coefficients of variation are at the smallest, which confirms its dominant position in the present stands in four forest management classes of fir and beech forests. The fir prevails in the stand basal area of the thickest 70 60 50 40 30 20 10 Plot 98 • Spmce • Fir o Beech Other iiuyy 1950 1960 1971 1979 Year 1988 2003 1951 1960 1970 1979 Year 1988 2003 60 50 40 I 30 20 10 0 Plot 97 ^^ Spruce ^ Fir ^ Beech ^^ Sycamore 1950 1960 1970 1980 1990 2000 Year 60 50 40 I 30 ■o" 20 10 0 Plot 98 Spruce Fir Beech Sycamore 1950 1960 1970 1980 Year 1990 2000 60 - Plot 99 50 - 40 - •i- 30 - ■s H ___ „ ____ 20 - H S--- H- - El -- 10 -0 - ^^ Spruce ^ Fir ^ Beech ^^ Sycamore 1950 1960 1970 1980 1990 Year 2000 Fig. 1: The changes in the structure of basal area and the development of the quadratic means of trees on three research plots in Dinaric fir-beech forests from 1950 to 2003 Slika 1: Spremembe v strukturi temeljnice in razvoj srednjetemeljničnih premerov dreves na treh raziskovalnih ploskvah v dinarskih jelovo-bukovih gozdovih od leta 1950 do 2003 trees, however, its share in the DBH class to 30 cm is smaller compared to the beech and the spruce (Fig. 3). On the basis of 500 m2 sampling plots we estimated the diversity of DBH-based structure in the fir and beech forest stands. Since the sampling plots are too small to enable such estimations, we confirmed a high level of variability in the stand structure indirectly, based on the coefficients of varia- tion arithmetic mean diameters of the spruce, the fir and the beech. On individual sampling plots, the groups of tree species representing uniform stand structures were present rather rarely. Only on one fifth of the sampling plots, the coefficients of variation for arithmetic diameters of the fir, the spruce and the beech were estimated to be smaller than 20%. Even in the forest management class OF-lycopodietosum, dominated BA mVha dq (cm) cv (%) 0 1 ■C I 50 40 30 - 'Iii Spruce Fir Beech Total 50 40 30 20 10 0 50 40 30 - Spruce Fir Beech Totai Spruce Fir Beech Totai 50 40 30 -20 10 -0 Li Spruce Fir Beech Totai Spruce Spruce Spruce Fir Beech Spruce Spruce Spruce Spruce Fir Beech Spruce Fig. 2: The estimations of the basal area, quadratic means and coefficients of variation for average diameters on the permanent sampling plots in the fir-beech forests of the Leskova valley in 2003 Slika 2: Ocene temeljnice, srednjetemeljničnih premerov in koeficientov variacije za povprečne premere na stalnih vzorčnih ploskvah v jelovo-bukovih gozdovih Leskove doline v letu 2003 according to data in the forest management plan (Gozdnogospodarski načrt 2007) by uniform fir and spruce stands with mixed deciduous trees, the coefficients of variation for arithmetic mean diameters were estimated to be smaller than 20% merely on 17% of the sampling plots. In the distribution of the coefficients of variation in Figure 4, the medians of the spruce and the beech were equal (CV = 37%), while the median of the fir was at 38%. The diversity of stand structure in the fir-beech forests was confirmed on the basis of the DBH-based Shannon index as well (Fig. 5). On 2-hectare-research plots, the DBH-based Shannon indices have increased and reached the highest value when the trees were represented in all 4 DBH classes. In DBH 70 cm DBH 30 cm • Spruce • Fir O Beech Other OF-lyco OF-homo OF-merc OF-typc OF-lyco OF-homo OF-merc OF-typc Fig. 3: The basal areas of tree species for the highest and the lowest DBH class in the fir- beech forest of the Leskova valley, estimated on the permanent sampling plots in 2003 Slika 3: Temeljnice drevesnih vrst za največji in najmanjši debelinski razred v jelovo-bukovih gozdovih Leskove doline, ocenjene na stalnih vzorčnih ploskvah leta 2003 80 i2 60 a. f o. E SS 40 0 1 E 20 M M • Spruce O Fir O Beech M k dHi 10 20 30 40 50 60 70 80 90 CV [%] Fig. 4: Frequency distribution of permanent sampling plots according to CV classes, calculated for arithmetic means of silver fir, beech and Norway spruce in fir-beech forest in the Leskova valley in 2003 Slika 4: Frekvenčna porazdelitev števila vzorčnih ploskev po posameznih razredih koeficientov variacije, ki so bili izračunani za aritmetično srednje premere treh drevesnih vrst v jelovo-bukovih gozdovih Leskove doline leta 2003 the year 2003, the Shannon index reached 94% of maximum values for 5 cm and 20 cm DBH classes on the plot with prevalent fir (Plot 99). The index was the lowest on the plot with prevalent spruce (Plot 97), where it had reached 90% of the highest value for 5 cm classes and 70% for 20 cm classes. The latest measurements on the research plots have estimated lower Shannon's indices for 5 cm and 20 cm DBH classes on the plot 98, since the trees have grown into higher DBH 3.0 -2.5 -2.0 1.5 1.0 -0.5 - 0.0 - -----t-:::: 5 cm DBH classes - 0 ________^ o--- El H----- KI----- -----BI---------- __ __0--- __- + ___ 0 - -+- --- o- -BI-KI- -H 20 cm DBH classes H - I2J .............+ + --M- ............ + tree speciess ° Plot 97 + Plot 98 ® Plot 99 1950 1960 1970 1980 1990 2000 Year Fig 5: Changes of DBH-based Shannon's index for 5 and 20 cm classes and species based index on tree research plots in Dinaric fir-beech forest between 1950 and 2003 Slika 5: Spremembe Shannonovih indeksov za debelinsko pestrost po 5 cm in 20 cm debelinskih razredih ter indeks vrstne pestrosti na treh raziskovalnih ploskvah v dinarskih jelovo-bukovih gozdovih v med letoma 1950 in 2003 classes and the share of the basal area in lower DBH classes has declined. This estimation is confirmed also by the coefficient of variation for mean diameter of trees that has declined from 58% to 46% since the year 1988. As regards the fir, the coefficient of variation remained the same as in the previous period, while that of the spruce diminished by 7% and of the beech by 12% on this research plot. The Shannon index of species diversity on the research and permanent sampling plots was the lowest among three comparative indices, because the major part of the basal area was dominated by two tree species, whereas on the plot with prevailing spruce, the Shannon index of species diversity grew even smaller after the year 1970. The shares of the fir, beech and sycamore Acer pseudoplatanus in the stand basal 5 cm DBH classes 20 cm DBH classes tree species i i -i- OF-lyco OF-homo OF-merc OF-typc OF-fyco OF-homo OF-merc OF-typc OF-lyco OF-homo OF-merc OF-typc Fig. 6: Shannon's indices for DBH and species diversity, estimated on the permanent sampling plots in 4 forest management classes in 2003 Slika 6: Shannonovi indeksi za debelinsko in vrstno pestrost, ocenjeni na stalnih vzorčnih ploskvah v 4 gozdnogospodarskih razredih leta 2003 D t^