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Bryophyte vegetation on volcanic rock outcrops and dead wood is studied in a near-natural montane beech stand in northern Hungary. Substrate specificity of the species and  the existing interspecific relationships are described. The most important species combinations and their diversity are evaluated using information theoretical functions and Monte-Carlo simulations. All analyses are based on presence/absence data of 33 species in 1508 100 cm2 microplots. Most species exhibit strong substrate specificity. Of the species that occurred with frequencies higher than 10, 8 are associated to rock, 5 to dead wood and 5 to both substrate types. Analyses of interspecific associations and agglomerative classification reveal that frequent species of species-poor bare rocks are separated from species-rich assemblages of humus-rich outcrops and coarse woody debris.  Monte Carlo simulations reveal that many species combinations are significantly more frequent than expected under the assumption of random combining of species. Observed number, diversity and evenness of species combinations are significantly lower, whereas interspecific constraint (expressed as associatum) is significantly higher than under the neutral models even when data are stratified according to substrate type. The presence of coarse woody debris, not only provides habitat for wood inhabiting bryophytes, but also results in diverse rupicolous bryophyte assemblages on humus-rich outcrops.

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Different types of forest use significantly changed the structure and species composition of European temperate forests. Herbaceous species and seedlings are important parts of the forest ecosystem, thus it is necessary to understand the effects of stand characteristics on the species composition of the understory. In our study we assessed the main factors that affect the species composition of herb and tree seedling assemblages in Quercus petraea and Q. cerris dominated stands (age 50–150 years) in the Bükk Mountains, Hungary. The relationship between the studied assemblages and explanatory variables (tree species composition, stand structure, canopy closure and topography) were explored by Redundancy Analysis (RDA). The occurrence of herbaceous species was affected by canopy closure, stand structure (mean DBH and DBHcv of trees), topography and the density and diversity of shrub layers. Oak forest species were associated with more open stands with sparsely distributed large trees, while mesic forest species were positively associated with heterogeneous stand structure, low shrub density, and western exposure. Seedlings of trees and shrubs showed a dispersal limited phenomenon. The composition of seedlings was significantly influenced by the mean DBH of trees, the structural heterogeneity of the overstory, the tree species diversity and the density of shrub layers. However the seedlings of both dominant oak species required the same stand structure, sessile oak was able to regenerate almost exclusively in those stands where it was dominant in the overstory, which is significant for the management of the species. Generally, forest management affects species composition and structure of the overstory, accordingly it had direct and indirect effects on the understory community as well.

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The effects of stand structure, tree species composition, proportion of habitat types and land use history on breeding bird assemblages in temperate mixed forests in Western Hungary were studied. The species richness, the abundance and the composition of the whole breeding bird assemblage and of some groups formed on the basis of nesting site and rarity were examined. Stand structural variables had the highest impact on the breeding bird assemblage, while tree species composition, the varying proportion of vegetation types and land use history had no significant effect. In the case of the species richness, the abundance and the composition of the whole assemblage, the most important variables were the mean diameter of trees, the vegetation cover of the forest floor and the dead wood volume. The explained variance in the linear models of different groups varied between 20% and 60%, and the relative importance of these three variables also differed considerably. These results indicate that forest management may considerably influence the diversity and the composition of birds, as all the structural elements affecting birds deeply depend on it. Within the shelterwood management system, the elongation of the rotation and regeneration periods, and the relatively high proportion of retention tree groups after harvest could contribute to the conservation of forest birds. Our results also showed that for the forest bird communities, both the prevalence of big trees and the presence of a dense understory layer are important. Management regimes which apply continuous forest cover might be more appropriate for providing these structural elements simultaneously on small spatial scales, and for the maintenance of a more diverse bird community, thus healthier forest ecosystems.

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Different diversity measures of forest floor assemblages were evaluated in order to check if they can be used as indicators of forest naturalness. We compared vascular and bryophyte vegetation of two habitat types in an unmanaged beech-dominated reserve and five managed stands of different ages. We used systematically collected data characterizing four spatial scales obtained by successively aggregating neighbouring quadrats. Species richness did not always differentiate near natural sites from managed sites, and the observed difference depended very much on the spatial scale used. The behaviour of Shannon-Wiener diversity function can only be understood if both the species richness and the evenness components are considered. Near natural plots had high Shannon-Wiener diversity values even at the finest spatial scale not only because of high number of species, but also because of high evenness. We found that a simple measure of pattern diversity — spatial variation of species importance — was the most effective in differentiating the diversity of plots with different levels of naturalness. The absolute values of pattern diversity in the forest floor vegetation were the highest in those plots where the characteristics of important limiting ecological factors were generated by natural disturbance. Vascular and bryophyte species responded differently to tree stand structural characteristics. The diversity of vascular vegetation was determined mainly by the spatial variation of light availability, whereas that of bryophyte vegetation responded to the amount and spatial heterogeneity of appropriate substrates (dead wood, rock). The use of pattern sensitive diversity measures is necessary to reveal diversity-naturalness relationships. We suggest that all diversity descriptors should be calculated for different spatial scales, since their change with spatial scale was as informative as their actual values.

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Three indirect light measurement methods were compared in mixed deciduous and coniferous forests with heterogeneous stand structure: tRAYci — a spatially explicit light model calculating percentage of above canopy light (PACL); LAI-2000 Plant Canopy Analyzer measuring diffuse non-interceptance (D1FN); and spherical densiometer estimating canopy openness (CO). Correlations between the different light variables were analyzed at several spatial scales (at 5 × 5, 10 × 10, 15 × 15, 20 × 20 and 30 × 30 m 2 ). Relationships between light variables and the cover of alight flexible plant, blackberry ( Rubus fruticosus agg.), as a potentially sensitive response variable for light conditions were also investigated. LAI-2000 (D1FN) and tRAYci (PACL) seemed the most appropriate for the description of the light environment in the investigated stands. D1FN and PACL had stronger correlations with each other and with blackberry cover than CO. Spatial heterogeneity of light (expressed with coefficient of variation) showed much stronger correlations than mean values both between the methods and between light intensity and Rubus cover. The correlation values between the methods increased towards coarser scales (from 5 × 5 to 30 × 30 m 2 ), while the correlation between light intensity and blackberry cover had a maximal response at the scale of 20 × 20 m 2 if a lower resolution of light estimation was used, and had also a maximum at smaller scales if the light was calculated for more points per plot by tRAYci. LAI-2000 can be recommended for the comparison of different stands, however, for fine scale description of light conditions of a stand tRAYci seems to be more appropriate.

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Dry oak forests have one of the richest understory vegetation in Europe, but the environmental drivers of this community have been scarcely revealed. In this study, we assessed whether the amount of light, soil pH or stand heterogeneity affect primarily the species composition of this community. We investigaed 332 sampling plots in 40-165 year old managed and abandoned Quercus cerris and Q. petraea dominated forests in North Hungary. Presence-absence data of herbaceous species and seedlings of woody species were recorded in 28 subplots within each sampling plot. Stand structure, canopy openness and soil pH were also measured in each plot. The relationships between stand characteristics and the species assemblage were explored by redundancy analysis, while the individual responses of species and species groups were studied by generalized linear mixed models. Multivariate methods and individual species response analyses provided similar results, the amount of light and soil pH were equally important variables (both of them explained 2.8% of species variance), while stand heterogeneity had a bit lower, albeit still significant role in determining understory species composition (1.9% of species variance explained). Seedlings of woody species preferred shaded (half-shaded) conditions, while many herbaceous species were positively related to light. The effect of the three explanatory variables was hard to separate, since they influenced each other as well. Sessile oak seedlings and herbs typical of dry forests, forest edges, grasslands and acidic soil habitats preferred light rich habitats with homogeneous stand structure and low soil pH. Mesic forest herbs and seedlings of other woody species were related to relatively high soil pH, heterogeneous stand structure and closed canopy. These two understory types were clearly separated regarding composition. This study emphasizes the importance of heterogenous light conditions and mosaic, diverse forest structure (presence of homogeneous and heterogeneous forest patches) during forest management for the maintenance of understory biodiversity.

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Community Ecology
Authors: G. Ónodi, Gy. Kröel-Dulay, E. Kovács-Láng, P. Ódor, Z. Botta-Dukat, B. Lhotsky, S. Barabás, J. Garadnai, and M. Kertész

Aboveground plant biomass is one of the most important features of ecosystems, and it is widely used in ecosystem research. Non-destructive biomass estimation methods provide an important toolkit, because the destructive harvesting method is in many cases not feasible. However, only few studies have compared the accuracy of these methods in grassland communities to date. We studied the accuracy of three widely used methods for estimation of aboveground biomass: the visual cover estimation method, the point intercept method, and field spectroscopy. We applied them in three independent series of field samplings in semi-arid sand grasslands in Central Hungary. For each sampling method, we applied linear regression to assess the strength of the relationship between biomass proxies and actual aboveground biomass, and used coefficient of determination to evaluate accuracy. We found no evidence that the visual cover estimation, which is generally considered as a subjective method, was less accurate than point intercept method or field spectroscopy in estimating biomass. Based on our three datasets, we found that accuracy was lower for the point intercept method compared to the other two methods, while field spectroscopy and visual cover estimation were similar to each other in the semi-arid sand grassland community. We conclude that visual cover estimation can be as accurate for estimating aboveground biomass as other approaches, thus the choice amongst the methods should be based on additional pros and cons associated with each of the method and related to the specific research objective.

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