The research was conducted in three old−growth forests consisted of silver fir Abies alba Mill., European beech Fagus sylvatica L., and Norway spruce Picea abies Karst. located in the southern part of Poland (Żarnówka and Oszast) and the south−eastern Bosnia and Hercegovina (Perućica). The aim of the study was to compare basic stand characteristics and spatial heterogeneity in terms of variation in stand basal area and vertical structure. In each research area, small sample plots (0.015 ha) were localised in a regular 20×20 m grid covering approximately 10 ha. In each sample plot the diameter at breast height (d1.3 ≥ 7 cm) and species of all live trees were recorded. For each plot the basal area of live trees and an index of structural diversification were determined. As a measure of structural diversification, the simple variance in tree height was computed and scaled through comparison with a hypothetical variance of the uniform distribution. The spatial patterns of basal area and structural diversification were tested with paired−plot approach. In addition, simulation techniques were used to model variation in the basal area of live trees dependent on spatial scale. The Carpathian and Dinaric stands considerably differed in basal area (ranging from 36.1 to 65.2 m²/ha) and volume (varying from 522 to 1176 m³/ha), but all of them had diameter distributions proximate to a negative exponential model. The basal area recorded on the 0.015 ha plots had a very similar pattern of variation, which could be generalised as a truncated normal distribution. The distribution of the structural diversification index was different and resembled an uniform (Perućica) or a bimodal distribution with modal values at its extremes (Oszast and Żarnówka). However, in the spatial scales above 1,000 m² the index distribution become similar to a normal (Oszast) or a truncated normal (Perućica and Żarnówka) ones with high mean values, indicating the predominance of complex vertical structures. In general, the spatial variability in basal area and structural diversification of live trees tended to be random. These results suggest that the patch−mosaic assumption being fundamental for the developmental cycle hypothesis is inapplicable to the studied primeval forests. Regardless of differences in geographic location and site conditions, the studied stands show a similar spatial pattern of structural heterogeneity, suggesting a close resemblance of disturbance regimes driving its dynamics.
The aim of this study was to characterize the texture of a primeval forest composed of Fagus sylvatica (L.), Abies alba (Mill.) and Picea abies ((L.) H. Karst). Empirical data were collected in the Babia Góra National Park (southern Poland) in the stand being under strict protection since 1934. 259 circular plots with a radius of 7.0 m and an area of 154 m2 each were established in nodes of 20×20 m grid. For individual plots and blocks of the combined plots representing gradient of spatial scales between 0.015 and 0.640 ha, the number of trees, diameter at breast height (dbh) distributions, basal areas and the values of structural diversity indices of Gini (GI), Shannon (SH) and Staudhammer−LeMay (STVI) were determined. The indices were also calculated for several types of theoretical distributions. Based on the values obtained for the theoretical distributions, the individual plots and the blocks of the combined plots were classified as representing simple (GI0.30; STVI0.10), moderately diversified (0.30GI0.45 and 0.10 STVI0.30) or complex (GI0.45; STVI0.30) dbh structure. For all the spatial scales analyzed the average values of GI and STVI indices reached the level typical for populations of a high structural diversity (i.e. exceeded the values of 0.45 and 0.30 respectively). According to the GI and STVI values, the portion of stand patches with complex dbh structure ranged from 70.9% and 68.2% at the individual plots scale, respectively, up to 100% in the blocks of 16 plots (4×4). In general, in all the spatial scales analyzed the spatial diversification of the dbh distributions and basal area levels was higher than in managed selection forests and much higher than in managed single−storied stands. The dominant frequency of highly diversified dbh distributions found in the analysed stand was not concordant with the predictions of the forest dynamics theory based on developmental stages, according to which in primeval forests with a significant partition of Abies alba and Picea abies should prevail stand patches of rather simple dbh structure, characteristic for the long−lasting optimum stage.
The objective of the study was to compare the seven tree species in terms of their role in the formation of a complex vertical structure in mixed broadleaved stands. The data came from 18 multi−storey old−growth forests representing different communities of lowland deciduous forests in the Niepołomice Forest. In each stand, squared sample plot (80×80 m) was established. The height of trees of dbh≥7 cm was measured. Then, for each studied species, values of the two structural indices were determined. The structural diversity index (ZS) quantifies tree height variation within individual population, whereas the index of structure−forming role (RS) expresses the contribution of the given population into vertical structural diversity of the whole stand. The greatest tendency to form a population with a large height diversity belonged to Tilia cordata, Fraxinus excelsior and Ulmus laevis. Significant opportunities in this regard, however, were exhibited by other tree species, including shade−intolerant Pinus sylvestris and Alnus glutinosa. This phenomenon in the conditions of eutrophic habitats, with the constantly present competitive pressure from shade−tolerant species, should be considered as an exceptional. A major contribution to vertical structural diversity of the stand was often an attribute of the populations with a low internal height diversity, which concerned especially Quercus robur. This means that an important ecological mechanism responsible for the formation of a multi−layer canopy is complementary filling the space above ground by trees of the species with different light requirements. However, the lack of correlation between RS and ZS indices suggests that the ability of particular species to form the population with a high internal diversity is equally important. The results prove the possibility of forming stands with a very complex vertical structure in the conditions of lowland mixed deciduous forests. However, in terms of sustainable maintenance of such stand structure, the issue of the growth rate of trees at different competitive pressure and the nature of their response to the release at different stages of ontogeny remains open.