ANNOTATION: This dissertation focus on spontaneously developed forests (SDFs) on mesic stands. The present tree layer was described and the possible changes in the tree layer was estimated. Influence of site- and context-dependent factors on the species composition and general character of the herb layer was studied in detail. An experiment was established to study seed and safe-site limitation of nine forest herb species.
Summary
The study area was situated into the southern part of the Czech Republic (40°35-38´ N, 14°11-17´ E), altitude 665-940 m (Fig. 1). Climatic conditions moderately oceanic between the warm and cold type (climatic regions MT 3 – Ch 7), average year temperature 6.8°C and average year precipitation 718-1 003 mm.
The SDFs on mesic stand (neither waterlogged nor desiccated, with vegetation neither ruderalised nor typical for poor soils) were studied. Forty eight plots (100 m2) were fixed in SDFs, which differed in age to cover up the chronosequence of successional sere directed toward forests in the study area. The species composition and the influence of the site- and context-dependent factors on the species composition and character of the herb layer were studied, and an estimation of the probable future development of the tree layer was calculated.
The studied SDFs were on average 7.4 ha large (ranging from 0.08 to 22 ha), the shape differed much between the studied SDFs (an average relative length of the SDF border was found to be 3.1, ranging from 1.1 to 15.6), and the nearest distance to the border of the SDF was found to be not long (on average 22 m, ranging from 10 to 80 m).
The SDFs were semi-open growths with an average relative irradiance of 20% of incoming radiation in the 120 cm above ground (ranging from 3 to 68%), with a relatively dense herb layer reducing the incoming radiation on average at 6% in the 5 cm above ground (ranging from 1 to 36%). Most of the SDF were on shallow soils (mean depth of the organic soil horizon was found to be 23 cm, ranging from 4 to 70 cm), with relatively high rock fragment content (mean value 42%, ranging from 6 to 86%). Several soil characteristics were found to be intermediate between the numbers typical for grassland and forest soils: soil reaction (mean value 4.3, ranging from 3.8 to 5.6), organic matter content (mean value 11%, ranging from 5 to 21%), basal respiration of the microbial community (mean value 2.7 g C CO2.g1.hod1, ranging from 0.7 to 5.8 g C CO2.g1.hod1).
Most of the SDFs were surrounded by SDFs (on average 38%), grassland (31%), and forests (23%), while fields (6%) and “other” land cover (2%) were low abundant. The distance to the nearest grassland was only 45±60 m, while the nearest forest was found to be 111±152 m away and the nearest field 743±912 m.
The average age of the tree layer was found to be 30 years (ranging from 10 to 46 years), but the age structure of the tree layer was not found to be homogeneous, because the maximum age of the tree layer was 46 years (ranging from 12 to 84 years). This finding corresponds with the data from the historical aerial photos: most of the SDFs were not covered by trees in 1952 (65% grasslands, 19% fields; 13% covered by scattered trees, and 4% by closed tree canopies), while half of the SDFs were covered by trees in 1966 (33% grassland, 13% fields, 25% scattered trees, 29% closed tree canopy) and most of the plots were covered by trees in 1983 (4% fields, 13% grassland, 23% scattered tree canopy, 60% closed tree canopy).
The tree layer changes from the dense growth of young thin trees (up to 179 trees.are-1) to open canopies of adult trees (down to 2 trees.are-1). The age structure of the older tree layer is less age uniform than in the young SDFs.
The tree layer was dominated by pioneer tree species. In total, 11 tree species were listed in the tree layer. The anemochorous species dominated the tree layer, but endozoochorous species were also able to colonise these growths. On average 2.6 species were found in a plot, and 83% of the trees were deciduous ones. The birch (Betula pendula Roth.) was found to be the most common and most abundant species (occupying 88% of plots, 60% of trees), but the European aspen (Populus tremula L.), Norway spruce (Picea abies (L.) Karsten), mountain ash (Sorbus aucuparia L.), and Scots pine (Pinus sylvestris L.) were able to dominate the tree layer as well. Of other species, the goat willow (Salix caprea L.) and common ash (Fraxinus excelsior L.) were more common incidental trees in these growths.
The species composition of the tree-layer was related from the studied factor to the Ellenberg T indicator value and shape of the SDF (14% of the total variability in the data explained). The Norway spruce and Scots pine preferred colder stands, while the sycamore maple, cherry-tree and common ash preferred warmer stands. The birch occupied sites with more irregular shapes, unlike the European aspen which preferred less complicated shapes of the SDF (it may be a result of vegetative reproduction of this species). The growths dominated by the European aspen were found to be more occupied by the endozoochorous cherry tree and mountain ash, while in the SDFs dominated by birch the anemochorous goat willow was more common. More tree species were found to be at higher altitudes and/or at colder stands, in younger growths and in the SDFs with more complicated shapes. The abundance of the two most common species (birch and Norway spruce) were influenced mostly by the soil conditions, history, age and proportion of SDF, and forests in the 300 m surroundings.
In total, fifteen tree-seedling species were listed. Most of the seedlings were 10-40 cm high. An average plot contained 145 seedlings of five species. The best colonisers were species non-specific to the stage of succession (59% of the seedlings, 96% of the plots), and anemochorous species (75% of the seedlings, 98% of the plots). Species typical for terminal stages of succession were able to colonise 65% of the plots, but they took only 4% of the total number of seedlings. Zoochorous species occupied 85% of the plots and took 25% of the seedlings. Species typical for the early stages of succession preferred plots more irradiated at the 20 cm above ground, lower altitudes, less productive sites, and sites near to the forest. Species typical for terminal stages of succession preferred lower altitudes, older growths, medium rich stands with more organic matter content, lower soil reaction, and sites near forests. Species non-specific to the stage of succession gained in medium-aged SDFs and under a dense tree layer. Anemochorous and zoochorous species were not found to be influenced by any studied factor.
The sycamore maple (Acer pseudoplatanus L.) was the most common seedling species (60%), while the European aspen (10%), mountain ash (10%), Norway spruce (9%), cherry tree (5%), and common ash (3%) were also abundant. The altitude and soil reaction significantly influenced the tree-seedling species composition. Sycamore maple seedlings were more common in younger SDFs, which grew faster. Norway spruce seedlings were negatively influenced by relative irradiance at the 20 cm above ground. The number of birch seedlings was negatively influenced by the relative irradiance at the 20 cm above ground and/or by the abundance of typical grassland species. More cherry tree seedlings were found at plots in lower altitudes, near forests and in older stands. The mountain ash was more common in the plots where the herb layer was more similar to the herb layer of the forests.
An estimation of the changes for the next generation of the trees was done. It was based on a comparison of the present and estimated species composition. The estimated tree layer species composition was derived from the number of tree seedlings according to their height. The estimation predicts big changes in the tree layer species composition. The estimated tree layer species composition was related to the time of succession, nutrients, and humidity (25% of the total variability explained). The shift was found to be directed toward the species composition typical for terminal stages of succession, but the species composition of the second generation of the tree layer was still estimated to be very different. The succession will probably be long, and several centuries can be expected to be necessary for the successional changes leading to the species composition near the natural species composition.
Most of the species common in the present tree layer are supposed to change their abundance in the estimated tree layer. The most important finding is the estimated decrease of the pioneer species the birch, and the predicted increase of the ubiquitous species the Norway spruce and the sycamore maple and the ability of species typical for terminal stages of succession (beech, pedunculate oak) to establish themselves in the next tree generation. The sycamore maple, European beech, and pedunculate oak (Quercus petraea (Mattuschka) Liebl.) were found to be typical species in the estimated tree layer, although present common species (birch, European aspen, Norway spruce) will be more common.
The results are relatively optimistic for the forest managers, because only 5% of the plots probably will not be colonised by species “valuable” for forest management, while 47% of plots will contain more than one of these species. Most of the plots (79%) exceeded 2,000 seedlings of “valuable” species per hectare, which is sufficient for natural regeneration. Almost half of the trees “valuable” for forest management were found to be deciduous.
The general character of the herb layer of the SDFs in the first tree generation could be described as follows: grassland character prevails (average G = 20.5, Gw = 45.1), while woodland character plays a minor role (average W = 8.7, Ww = 18.0). Most of the species listed in the herb layer were related to the grassland vegetation (in total, 86 species, mean abundance 33%), followed by a group of species growing both in grassland and wood (30 species, 17%), woodland species (26 species, 10%), antropochorous and/or ruderal species (18 species, 3%), shrub species (10 species, 1%) and species typically growing at clearings (4 species, 2%). Most of the species related to the grassland vegetation were exozoochorous species (average Gw_Zex = 22.6) and species non-specific to the type of distribution (average Gw_Ns = 11.5), while most of the species related to the forest vegetation were anemochorous (average Ww_Wd = 6.5), endozoochorous species (average Ww_Zend = 5.6) or species non-specific to the type of distribution (average Ww_Ns = 4.5).
Species grouped according to the biotope were in general more influenced by the context-dependent factors, especially by the historical vegetation cover and by the proportion of grassland and forests in the surroundings, while the proportion of “other” land cover and the character of the SDF (area, shape and position within the SDF) were of less importance. Of the site-dependent factors, the soil characteristics were more important for several groups of species, but the relationships were group dependent.
General trend observed was the preference of grassland species to the soil conditions closer to the soil conditions typical for grassland soils (i.e., lower organic matter content, higher soil reaction). The most important finding was that the abundance of the woodland species increased with the age of the tree layer and with the duration of the tree layer. However, a decrease in grassland species was not observed.
The increase in the woodland character of the herb layer with the duration of the tree layer (Aa, Amax, historical vegetation cover) was promoted using the (weighted) grasslandness and (weighted) woodlandness indeces. The relationships of the other factors were index dependent, and no general trend was observed, except for the woodlandness related to the soil conditions more similar to the forest soils (i.e., higher organic matter content, lower soil reaction and higher microbial respiration).
Almost all of the groups of species according to the type of distribution related to the woodland were found to be more abundant in the SDFs with the tree layer developed for a longer time period (Aa, Amax, historical vegetation cover), but the decrease in groups of species related to grassland vegetation was not observed. Many of the site- and context-dependent factors significantly influenced the abundance of the groups of species, but the relationships were group dependent and no general pattern could be observed.
In total, 161 vascular plant species were listed, on average 32±8.6 species per 100 m2. The herb layer of SDFs was found to be highly variable (23% of the species were found only at one plot, while only 18% were found at more than 33% of the plots). An average diversity value was found to be 2.6 (ranging from 0.4 to 3.5), measured as a Shannon Wiener index, which is relatively high.
All the common species in this study were also common species in the study area and species widespread in Central Europe. The most common species were the following: Agrostis capillaris L., Galium mollugo agg. L., Holcus mollis L., Veronica chamaedrys L. (present at more than 80% plots), while the most abundant were the following species: Agrostis capillaris, Avenella flexuosa (L.) Drejer, Holcus mollis and Vaccinium myrtillus L. (more than 7% abundance per occupied plot). Most of the diagnostic and constant species were species related to the grassland vegetation (33 out of 161 species), especially of the Arrhenatherion elatioris, Polygono bistortae Trisetion flavescentis and Violion caninae alliance. Only ten species were related to the forest vegetation, i.e., of the Dentario enneaphylli Fagetum association, Luzulo Fagion and Quercion petreae alliance. Only several typical forest species were able to establish in the SDFs, and they were usually low abundant. Ferns and Vaccinium myrtillus were the only more abundant typical forest species, and of the Oxalis acetosella L. was relatively frequent. However, some forest species typical for oak-beech or beech forests were also listed in the herb layer, e.g., Asarum europaeum L., Geranium robertianum L., Mercurialis perennis L. and Mycelis muralis L.
In general, the influence of the studied factors on the frequent species abundance was species specific, and only a few general trends could be observed. Both site- and context-dependent factors influenced the abundance of frequent species, and there was found to be no big difference or general trend true for these two types of factors.
The seed and safe-site limitation of forest herbs was studied in a factorial designed experiment in a 12-year abandoned grassland (48°41.468´ N, 14°17.382´ E, altitude 700 m). The influence of disturbance and shading on species composition, general characteristics of the vegetation, and on species abundances was studied, and seedling establishment of forest species was monitored. In total, 16 treatments in six randomised blocks were fixed and the following manipulations were performed: i) shading at the 40% level of incoming PhAR, ii) shading at the 5% level of incoming PhAR, iii) removal of above-ground plant biomass, iv) disturbance of the turf, v) seed addition of typical forest species (Actaea spicata L., Carex sylvatica Huds., Dentaria enneaphyllos L., D. bulbifera L., Galium odoratum (L.) Scop., Maianthemum bifolium (L.) F.W. Schmidt, Mercurialis perennis, Oxalis acetosella, Paris quadrifolia L.).
However, the between-year variability in species composition was higher than the changes in vegetation caused by disturbance (iii and iv) or shading treatments (i and ii), and the treatments were found to have had a significant effect. Disturbance treatments were found to be able to influence the species composition directly, and the changes in the species composition caused by the disturbance treatments were more quantitative than qualitative, unlike the shading treatments. Shading was found to slow down the changes in the species composition between years, and therefore this treatment was able to stabilise the species composition.
However, the turf disturbance was the most radical treatment causing the biggest changes in the vegetation one year after the treatment practice, and the following year the vegetation was able to close again. The reduction at the 40% level of the incoming radiation treatment was not able to produce new gaps in the vegetation, but the shading at 5% of the incoming radiation was able to increase the abundance of gaps.
The decrease in species diversity in control plots was observed, while the removal of above-ground vegetation was able to stabilise the diversity and the turf disturbance treatment increased the diversity, but only for a one-year period.
The annuals, rosettes and small perennial herbs were more abundant in the plots with the turf disturbance treatment.
Most of the forest species were not able to germinate, although the seed densities were high. The germination capacity in laboratory conditions was also found to be low for most of the species.
Only forest species sown directly into the study site were found. This finding indicates the seed limitation of the forest species. All the recruited species were found the following year after sowing. Dentaria enneaphyllos was the only well establishing species, but it was not able to germinate in 56% of the plots and its survival was found to be on average 3% in the plots, where it was able to germinate. None of the seedlings was able to develop into a further stage than to the seedling with cotyledons. Shading treatment was able to enhance the seedling recruitment of D. enneaphyllos, and there was also a similar trend (however insignificant) for disturbance of the turf treatment. The numbers of Oxalis acetosella seedlings indicate a similar pattern, although there was found to be an insufficient number of O. acetosella for the statistical analysis. These findings indicate the possible safe-site limitation of the forest species at a relatively early stage of succession.