Jan Roleček

LOCAL AND REGIONAL PATTERNS OF SPECIES RICHNESS IN CENTRAL EUROPEAN VEGETATION TYPES ALONG THE pH/CALCIUM GRADIENT

We investigated the relationship between soil pH/calcium content and species richness of vascular plants in seven broadly defined Central European vegetation types, using Ellenberg indicator values for soil reaction and a phytosociological data set of 11,041 vegetation sample plots from the Czech Republic. The vegetation types included (A) broad-leaved deciduous forests, (B) meadows, (C) dry grasslands, (D) reed-bed and tall-sedge vegetation, (E) fens and transitional mires, (F) perennial synanthropic vegetation and (G) annual synanthropic vegetation. Relationships between local species richness (alpha diversity) and pH/calcium were positive for vegetation types A and C, negative for D and G, unimodal for E, and insignificant for B and F. Ellenberg soil reaction values explained 37% of variation in local species richness for vegetation type E, 24% for A, 13% for D, but only less than 4% for the others. Species pool size, i.e., the number of species that can potentially occur in a given habitat, was calculated for each plot using Beals index of sociological favourability applied to a large phytosociological database. For most vegetation types, the relationships between species pool size and pH/calcium were similar to the relationships between local species richness and pH/calcium, with the exception of meadows (weak unimodal) and perennial synanthropic vegetation (weak negative).These patterns suggest that for those types of Central European vegetation that developed independently of human influence in the Pleistocene or early Holocene (dry grasslands, deciduous forests), there are larger pools of calcicole than calcifuge species. This pattern is also found at the level of local species richness, where it is, however, less clearly pronounced, possibly due to the predominance of a few widespread and generalist calcifuges in acidic habitats. The unimodal pattern found in mires may result from similar underlying mechanisms, but in high pH environments mineral-rich spring waters probably decrease species richness by having toxic effects on plant growth. By contrast, vegetation types developed under direct human influence (meadows, synathropic vegetation) show weak negative or no relationships of local species richness or species pool to pH/calcium gradient. These results support the hypothesis ofPärtel (Ecology 83: 2361–2366, 2002) andEwald (Folia Geobot. 38: 357–366, 2003), that the modern calcicole/calcifuge disparity in the species pool of Central European flora has resulted from historical and evolutionary processes that took place on high pH soils. In the Pleistocene, calcareous soils dominated both the dry continental landscapes of Central Europe and glacial refugia of temperate flora, which were mostly situated in southern European mountain ranges with abundant limestone and dolomite. The negative pattern of species richness along the pH/calcium gradient found in reed-bed and tall-sedge vegetation, however, is not consistent with this historical explanation.

Sampling design in large-scale vegetation studies: Do not sacrifice ecological thinking to statistical purism!

Most of the historical phytosociological data on vegetation composition have been sampled preferentially and thus belong to those ecological data that do not fulfill the statistical assumption of independence of observations, necessary for valid statistical testing and inference. Nevertheless, phytosociological data have been recently used for various ecological meta-analyses, especially in studies of large-scale vegetation patterns. For this reason, we focus on the comparison of preferential sampling with other sampling designs that have been recommended as more convenient alternatives from the point of view of statistical theory. We discuss that while simple random sampling, systematic sampling and stratified random sampling better meet some of the statistical assumptions, preferential sampling yields data sets that cover a broader range of vegetation variability. Moreover, today’s large phytosociological databases provide huge amounts of vegetation data with unrivalled geographic extent and density. We conclude that in the near future ecologists will not be able to replace the preferentially sampled phytosociological data in large-scale studies. At the same time, phytosociological databases have to be complemented with relevés of vegetation composed mostly of common and generalist species, which are under-represented in historical data. Stratified random sampling seems to be a suitable tool for doing this. Nevertheless, a methodology and input data for stratification have to be developed to make stratified random sampling an ecologically more relevant and practical method.

Experimental restoration of coppice-with-standards: Response of understorey vegetation from the conservation perspective

A substantial part of European lowland woodlands was managed as coppices or wood pastures for millennia. However, traditional management forms were almost completely abandoned in Central Europe by the middle of the 20th century. Combined with the effects of nitrogen deposition and herbivore pressure, shifts in management resulted in biodiversity loss affecting particularly light-demanding oligotrophic plant species. Experimental thinning was applied in a former oak coppice-with-standards in an attempt to restore vanishing understorey plant communities. Two levels of thinning intensity and zero management as control were used on 90 plots. Ten years after the treatment, significant changes in species composition and diversity were observed in heavily thinned plots, while moderate thinning had mostly insignificant effects. Light-demanding oligotrophic species significantly increased, indicating positive consequences of restoration. However, heavy thinning also brought about the expansion of native ruderal species. Alien species remained unchanged. We conclude that the restoration of coppice-with-standards can be an efficient tool to support vanishing light-demanding woodland species. Combined with biodiversity benefits, the increasing demand for biofuel may contribute to the renaissance of traditional management forms in forestry.

Habitat requirements of endangered species in a former coppice of high conservation value

Transformation of coppices to high forests has caused fundamental changes in site conditions and a decline of many species across Central Europe. Nevertheless, some formerly coppiced forests still harbour a number of the declining species and have become biodiversity hotspots in the changing landscape. We focused on the best-preserved remnant of formerly grazed and coppiced subcontinental oak forest in the Czech Republic – the Dúbrava forest near the town of Hodonín. To improve our understanding of the ecology of declining species, we studied local habitat requirements of vascular plants most endangered at the national level. We recorded vegetation composition and sampled important site variables in plots with the largest populations of endangered species and in additional plots placed randomly across all major forest habitats. We demonstrated that sites with endangered species have a highly uneven distribution in ecological space and that their species composition is often similar to open-canopy oak forests. Within this habitat, the endangered species are concentrated in places with a high light availability and high soil pH. Light-demanding species characteristic of subcontinental oak forests are the best indicators of these sites, while broadly distributed shade-tolerant and nutrient-demanding species avoid them. These results support the view that the occurrence of many endangered species in the Dúbrava forest is a legacy of the long history of traditional management that kept the canopies open. Light-demanding species are now threatened by ongoing successional changes. Therefore, active conservation measures are recommended, including opening up the canopies, early thinning of young stands, control of expansive and invasive species and understorey grazing or mowing.

Merging of Pedicularis exaltata and P. hacquetii in the Carpathians: from local history to regional phylogeography based on complex evidence

The disjunct occurrence of Pedicularis exaltata in the White Carpathians (Czech Republic), isolated by more than 500 km from the nearest populations in the Eastern Carpathians, has been considered one of the mysteries of the Western Carpathian flora. We used molecular methods (AFLP, ITS and cpDNA sequencing) to reconstruct a possible scenario of the evolution of P. exaltata and its closely related congener P. hacquetii and to evaluate their differentiation. We paid particular attention to the origin of the isolated population in the White Carpathians. We also analysed the vegetation composition at the sampling sites to characterize the habitat preferences of the Pedicularis species and compare different sites. The pattern of molecular variability does not support a species boundary between P. exaltata and P. hacquetii. These assumed species should be merged into one bearing the name Pedicularis hacquetii following the priority rule, as also proposed by several early morphology-based taxonomic studies. The cpDNA variability pattern supports an evolutionary scenario involving a distribution centre (refugium) in the Eastern Carpathians from which the species expanded to the more westerly parts of its extant geographical range. Low population differentiation in AFLPs, high gene diversity and high DW index in isolated populations indicate that this expansion was contiguous rather than based on long-distance dispersal. Under such a scenario, the White Carpathian population is considered one of isolated relict populations scattered throughout the Carpathians and the East European Plain. A complete phylogeographic reconstruction of P. hacquetii s.l., however, requires ancient DNA analysis of herbarium specimens of nowadays extinct populations of the East European Plain. Extant habitats of P. hacquetii s.l. consist of species-rich dry-mesic to mesic (sub)montane grasslands and subalpine tall-herb growths, which may be relics of vegetation widespread in the late Glacial/early Holocene, when the contiguous expansion of the species possibly proceeded. The analysis of species with high fidelity to P. hacquetii s.l. throughout its range indicated that middle altitudes of south-western White Carpathians belong to the regions most suitable for the taxon within the Czech Republic and Slovakia. We found no support for the late introduction/long-distance dispersal scenarios and consider the relic survival scenario most parsimonious for the White Carpathians.

Introduction to this special issue on the ecology and evolution of the Carpathian flora

The Carpathian Mountains, spanning from eastern Czechia through southern Poland, Slovakia and eastern Ukraine to Romania, are one of the least known mountainous diversity hotspots in Europe. Besides a unique diversity of endemic species and vegetation units, they also host numerous relicts from various time periods and had likely served as an important refugium for the boreal and temperate biota. For decades, the Carpathians have attracted botanists from surrounding countries, and their interest is not diminishing. This issue of Folia Geobotanica summarizes research presented at a joint conference of the Czech and Slovak Botanical Societies, held in the autumn of 2016 in Prague. The event was focused on describing Carpathian plant diversity from various angles, ranging from historical and floristic aspects to genetic variation and the structure of plant communities. The contributions made at this conference constitute the main body of the present issue.