Jaanus Paal

Global sampling of plant roots expands the described molecular diversity of arbuscular mycorrhizal fungi.

We aimed to enhance understanding of the molecular diversity of arbuscular mycorrhizal fungi (AMF) by building a new global dataset targeting previously unstudied geographical areas. In total, we sampled 96 plant species from 25 sites that encompassed all continents except Antarctica. AMF in plant roots were detected by sequencing the nuclear SSU rRNA gene fragment using either cloning followed by Sanger sequencing or 454-sequencing. A total of 204 AMF phylogroups (virtual taxa, VT) were recorded, increasing the described number of Glomeromycota VT from 308 to 341 globally. Novel VT were detected from 21 sites; three novel but nevertheless widespread VT (Glomus spp. MO-G52, MO-G53, MO-G57) were recorded from six continents. The largest increases in regional VT number were recorded in previously little-studied Oceania and in the boreal and polar climatic zones — this study providing the first molecular data from the latter. Ordination revealed differences in AM fungal communities between different continents and climatic zones, suggesting that both biogeographic history and environmental conditions underlie the global variation of those communities. Our results show that a considerable proportion of Glomeromycota diversity has been recorded in many regions, though further large increases in richness can be expected in remaining unstudied areas.

Diversity of bryophyte vegetation in some forest types in Estonia: a comparison of old unmanaged and managed forests

The bryophyte vegetation of 3 pairs of unmanaged and managed forest stands, representing Oxalis drained peatland, Aegopodium and Oxalis forest site type, were compared. The total number of bryophyte species in unmanaged stands was 74 and that in managed stands 54. Out of the 20 species occurring only in unmanaged forests, 9 grow on decaying wood, and 3 on trunks or bases of big trees; 13 of them were hepatics. In unmanaged forests 11 hemerophobic species were recorded altogether. Although the difference in substrate and species diversity between unmanaged and managed stands is not statistically significant, in unmanaged forests more substrates characteristic for an old-growth stand are available, and the percentage of species preferring dryer habitats or prolonged humidity is a bit higher than in managed forests; the percentage of species associated with better illuminated habitats is higher in managed forests. Analysis of classification structure of the bryophyte layer synusia shows that the number of societies is also higher in unmanaged forests. This is associated with more numerous microhabitats; the average light and humidity indices calculated for every society, confirm this conclusion. The large discrepancy in bryophyte layer classification structure in old-growth and managed forests of the same forest site type is manifested not so much by species content in synusia as by their abundance proportions. The larger diversity of classification units in unmanaged forests is also seen at the synusia facies level; four of nine facies are confined exclusively to unmanaged stands. This is a strong argument for the informativeness of bryophyte layer classification structure for purposes of indication and monitoring as well.

Epiphytic and epixylic lichen species diversity in Estonian natural forests

The epiphytic and epixylic lichen flora of natural forests was recorded in different parts of Estonia. Altogether 232 taxa of lichens, lichenicolous fungi, or non-lichenized fungi were recorded, 10 of them listed in the Estonian Red Data Book. We found regional differences in lichen species composition and diversity caused by differences in the forest types. The tree-species-rich boreo-nemoral forests had the most diverse lichen flora, while the boreal forest dominated by coniferous trees or birch had the lowest diversity. The stand age proved to be significant in regard to the number of lichen species in a forest. The most remarkable effect on the diversity of forest lichen species was caused by the presence of Populus tremula. Aspen had the highest number of lichen species on the basal trunk and twigs, and also the highest number of host-specific lichen species.

Rare and threatened plant communitiesof Estonia

This paper introduces the criteria for identifying rare and threatened plant communities in Estonia. An overview of such communities is given and the general problems associated with their protection are briefly discussed. Of forest communities, alvar forests and boreo-nemoral forests must be preserved very carefully. Once widely distributed minerotrophic mobile-water swamp forests and flood plain forests nowadays cover only a small area. Numerous rare communities are associated with coastal regions and islands. Wooded meadows, the most species rich vascular plant communities in northern Europe, as well as flooded meadows and many other communities require continuation of traditional management for their preservation.

Influence of tree stand age and management on the species diversity in Estonian eutrophic alvar and boreo-nemoral Pinus sylvestris forests

Abstract The influence of tree stand age, silvicultural management and several habitat variables on species diversity was analysed in Calamagrostis site type alvar forests and Hepatica site type boreo-nemoral forests. On the community scale, the herb-layer species richness appeared to be determined mostly by forest soil and light conditions. However, the number of bryophyte species growing on tree stem bases and on decaying wood was dependent on management intensity and forest site quality class. On the same substrate types the number of lichen species was promoted only by tree species diversity. On the 1 m2 quadrat scale, diversity and density of tree layer, species richness of bush layer and soil pH have significant layer-specific effects on species richness. Bryophyte diversity increases together with increasing shade and moisture in forest ground vegetation, while an increase of herb-layer diversity is connected with lightness. Therefore, maintenance of the small-scale species richness of forest ground vegetation cannot be achieved by uniform management practice.

Diversity of epiphytic lichens in boreo-nemoral forests on the North-Estonian limestone escarpment: the effect of tree level factors and local environmental conditions

The species richness and composition of lichens was quantified for four temperate broad-leaved tree species ( Acer platanoides, Fraxinus excelsior, Tilia cordata and Ulmus glabra ) in boreo-nemoral forests on the talus slope of the North-Estonian limestone escarpment (North-Estonian Klint). Thirteen study sites were distributed along the klint on a west to east gradient. The effects of tree and stand characteristics and geographical location of a stand on composition and diversity of epiphytic lichens were evaluated by multivariate analyses (DCA, CCA, pCCA) and by general linear mixed models (GLMM). Tree level variables (e.g. bark pH, bryophytes cover and host tree species) explained the largest fraction of the variation in lichen species composition. However, species richness and composition were significantly influenced also by the unique habitat conditions of klint forest (length of the forest fragment, proximity of the stand to the sea and height of the escarpment). A significant correlation between stand locality and lichen diversity on the tree bole was found which is most likely related to local air pollution gradients caused by alkaline cement dust and acid pollutants in the north-eastern part of Estonia.

Forest land-use history affects the species composition and soil properties of old-aged hillock forests in Estonia

Decisions regarding forest typology, management and protection are often based on the structures of present-day forests, ignoring their successional history. Forests growing on kames, eskers and various moraine hillocks common in regions with Holocene glaciation are good examples of this approach. In Estonia, these forests locally persist as fragments of continuous primary forest, but usually they are situated on former slash-and-burn areas (bushlands) or reforested agricultural land. Our aim was to elucidate the strength of the effect of long-term land-use history on the present-day vegetation compositions of mature hillock forests and their soil chemistry. It appeared that even the mature secondary hillock forests are still distinct from historically continuous stands in terms of species composition. We discovered connections between stand history and species content in hillock forests as well as transformed soil properties. The carbon and nitrogen contents in the humus horizons of secondary forests are lower while their carbon–nitrogen ratios are higher than in continuous forests. The relationship between vegetation and stand history is demonstrated by the higher proportions of anthropophytic and apophytic species in the herb layer of the secondary forests. The presence of species that are tolerant of anthropogenic impact on the secondary hillock forests floor can also be partly explained by the effect of different species in the tree and shrub layers, gaps in the tree canopy, and the boundary effect caused by the small areas of forest patches, neighboring grasslands or fields. The extinction debt in secondary communities should also be considered.

FLOODPLAIN FORESTS IN ESTONIA: TYPOLOGICAL DIVERSITY AND GROWTH CONDITIONS

Flooded forests in 79 sample areas were studied throughout the country. Using multivariate methods, six community types were established: (i)Tilia cordata-Mercurialis perennis, (ii)Ulmus laevis-Allium ursinum, (iii)Populus tremula-Convallaria majalis, (iv)Alnus incana-Cirsium oleraceum, (v)Alnus glutinosa-Filipendula ulmaria and, (vi)Alnus glutinosa-Carex acutiformis. The species composition of these types is partly overlapping but the abundance proportions of species are clearly different and all types have several significant (P < 0.05) indicator species. Communities of the first type grow mainly on temporarily overmoistened Dystri-Gleyic Arenosols on floodplain terraces but sometimes also in lower areas close to a riverbed on Eutric Gleysols or Gleyic Fluvisols. The second and third types embrace communities on levees where mainly Eutric Gleysols or Gleyic Fluvisols are represented. Communities of the fourth type are characteristic of low levees or flat areas with Mollic Fluvisols or Eutric Gleysols. The communities of the fifth and sixth types represent floodplain backswamp black alder forests on Eutric Gleysols, Molli-Histic and Histic Fluvisols. The convergence of communities, i.e., a situation where plant communities growing in various hydrological conditions and on different soils can have similar structure and belong to one community type is rather conspicuous for floodplain forests. In the conditions of Estonia, like in the Central European and Scandinavian countries, it is reasonable to consider as floodplain forest not only those growing on Fluvisolssensu stricto but all forests that are almost annually regularly covered with flood-water at least for a couple of weeks; the amount of alluvial sediments is not a decisive criterion in this case.

Cryptogams in Estonian Alvar Forests: Species Composition and their Substrata in Stands of Different Age and Management Intensity

Species-rich alvar forests represent a rare vegetation type in northern Europe. They grow locally on shallow soils formed on calcareous parent material. The aim of this study was to compare cryptogam species richness and composition on various substrata in alvar forests of different age and management rate. Altogether 101 bryophyte and 54 lichen species were recorded. Substratum explained species variance more than did the stand age or management intensity. Younger and older forests had a different bryophyte and lichen species composition. Decaying wood was the richest as cryptogam species substratum. It had quite a unique bryophyte composition, including a high amount of hepatics. Juniperus communis appeared to be an important substratum for cryptogam species diversity in alvar forests. Bryophyte species composition appeared to be similar on the bases of Pinus sylvestris, Betula pendula and Picea abies and different on J. communis. Most of the recorded threatened bryophyte species grew in old subnatural forests on stones or decaying wood.

MOSS SYNUSIAE IN SOUTH ESTONIAN FORESTS

On the Karula Upland, South Estonia, the forest moss layer was analysed using a transect of 726 contiguous 0.2×0.2 m plots. The sample plots were classified according to a multistage clustering procedure based on the sequential use of algorithms with different criteria. Several obtained clusters are rather similar in species composition, but abundance relationships among dominant species are distinctly different. For detailed analysis of mutual relations among societies a formal definition of adjacency is proposed, and two aspects of the cluster continuum — transitionality and distinctness — are estimated. It appears that almost all resulting societies are very distinct (P<0.05), but at the same time can be continual in the sense of transitionality. Spatial changes in vegetation along transects are also discontinuous. The null hypothesis assuming the independency of neighbouring sample plots type was refuted withP=0.01. The spatial extent of different synusiae is typically several times larger than it should be if there were no structure in moss vegetation.