Ilona Knollová

Separating habitat invasibility by alien plants from the actual level of invasion.

Habitats vary considerably in the level of invasion (number or proportion of alien plant species they contain), which depends on local habitat properties, propagule pressure, and climate. To determine the invasibility (susceptibility to invasions) of different habitats, it is necessary to factor out the effects of any confounding variables such as propagule pressure and climate on the level of invasion. We used 20 468 vegetation plots from 32 habitats in the Czech Republic to compare the invasibility of different habitats. Using regression trees, the proportion of alien plants, including archaeophytes (prehistoric to medieval invaders) and neophytes (recent invaders), was related to variables representing habitat properties, propagule pressure, and climate. The propagule pressure was expressed as the proportion of surrounding urban and industrial or agricultural land, human population density, distance from a river, and history of human colonization in the region. Urban and industrial land use had a positive effect on the proportion of both archaeophytes and neophytes. Agricultural land use, higher population density, and longer history of human impact positively affected the proportion of archaeophytes. Disturbed human-made habitats with herbaceous vegetation were most invaded by both groups of aliens. Neophytes were also relatively common in disturbed woody vegetation, such as broad-leaved plantations, forest clearings, and riverine scrub. These habitats also had the highest proportion of aliens after removing the effect of propagule pressure and climate, indicating that they are not only the most invaded, but also most invasible. These habitats experience recurrent disturbances and are rich, at least temporarily, in available nutrients, which supports the hypothesis that fluctuating resources are the major cause of habitat invasibility. The least invaded habitats were mires and alpine-subalpine grasslands and scrub. After removing the effect of propagule pressure and climate, some habitats actually invaded at an intermediate level had very low proportions of aliens. This indicates that these habitats (e.g., dry, wet, and saline grasslands, base-rich fens, and broad-leaved deciduous woodlands) are resistant to invasion.

Stratified resampling of phytosociological databases: some strategies for obtaining more representative data sets for classification studies

Abstract Question: The heterogeneous origin of the data in large phytosociological databases may seriously influence the results of their analysis. Therefore we propose some strategies for stratified resampling of such databases, which may improve the representativeness of the data. We also explore the effects of different resampling options on vegetation classification. Methods: We used 6050 plot samples (relevés) of mesic grasslands from the Czech Republic. We stratified this database using (1) geographical stratification in a grid; (2) habitat stratification created by an overlay of digital maps in GIS; (3) habitat stratification with strata defined by traditional phytosociological associations; (4) habitat stratification by numerical classification and (5) habitat stratification by Ellenberg indicator values. Each time we resampled the database, taking equal numbers of relevés per stratum. We then carried out cluster analyses for the resampled data sets and compared the resulting classifications using a newly developed procedure. Results: Random resampling of the initial data set and geographically stratified resampling resulted in similar classifications. By contrast, classifications of the resampled data sets that were based on habitat stratifications (2–5) differed from each other and from the initial data set. Stratification 2 resulted in classifications that strongly reflected environmental factors with a coarse grain of spatial heterogeneity (e.g. macroclimate), whereas stratification 5 resulted in classifications emphasizing fine-grained factors (e.g. soil nutrient status). Stratification 3 led to the most deviating results, possibly due to the subjective nature of the traditional phytosociological classifications. Conclusions: Stratified resampling may increase the representativeness of phytosociological data sets, but different types of stratification may result in different classifications. No single resampling strategy is optimal or superior: the appropriate stratification method must be selected according to the objectives of specific studies. Abbreviations: ASS = Phytosociological association; ELL = Ellenberg indicator values; GEO = Geographical stratification; GIS = Geographical information system; NUM = Numerical classification; RAN = Random resampling.

Local ranges of phytosociological associations: are they reflected in numerical classification?

In the tradition of European phytosociology, delimitations of vegetation units such as associations are mostly based on data from small areas where more detailed vegetation sampling has been carried out. Such locally delimited vegetation units are often accepted in large-scale synthetic classifications, e.g. national vegetation monographs, and tentatively assigned to a small geographical range, forming groups of similar (vicarious) vegetation units in different small areas. These vicarious units, however, often overlap in species composition and are difficult to recognize from each other. We demonstrate this issue using an example of the classification of dry grasslands (Festuco-Brometea) in the Czech Republic. The standard vegetation classification of the Czech Republic supposes that the majority of accepted associations (66 out of 68) have a restricted distribution in one of the two major regions, Bohemia or Moravia. We compared the classification into traditional associations with the numerical classification of 1440 phytosociological relevés from the Czech Republic, in order to test whether the traditionally recognized associations with small geographical ranges are reflected in numerical classification. In various comparisons, the groups of relevés identified by numerical analysis occupied larger areas than the traditional associations. This suggests that with consistent use of total species composition as the vegetation classification criterion, the resulting classification will usually include more vegetation units with larger geographical ranges, while many of the traditional local associations will disappear.

Classification of European and Mediterranean coastal dune vegetation

Aims: Although many phytosociological studies have provided detailed local and regional descriptions of coastal dune vegetation, a unified classification of this vegetation in Europe and the Mediterranean Basin has been missing. Our aim is to produce a formalized classification of this vegetation and to identify the main factors driving its plant species composition at a continental scale. LocationAtlantic and Baltic coasts of Europe, Mediterranean Basin and the Black Sea region. - Methods: We compiled a database of 30,759 plots of coastal vegetation, which were resampled to reduce unbalanced sampling effort, obtaining a data set of 11,769 plots. We classified these plots with TWINSPAN, interpreted the resulting clusters and used them for developing formal definitions of phytosociological alliances of coastal dune vegetation, which were included in an expert system for automatic vegetation classification. We related the alliances to climatic factors and described their biogeographic features and their position in the coastal vegetation zonation. We examined and visualized the floristic relationships among these alliances by means of DCA ordination. - Results: We defined 18 alliances of coastal dune vegetation, including the newly described Centaureo cuneifoliae-Verbascion pinnatifidi from the Aegean region. The main factors underlying the differentiation of these alliances were biogeographic and macroclimatic contrasts between the Atlantic-Baltic, Mediterranean and Black Sea regions, along with ecological differences between shifting and stable dunes. The main difference in species composition was between the Atlantic-Baltic and Mediterranean-Black Sea regions. Within the former region, the main difference was driven by the different ecological conditions between shifting and stable dunes, whereas within the latter, the main difference was biogeographic between the Mediterranean and the Black Sea. - Conclusions: The first formal classification of the European coastal dune vegetation was established, accompanied by an expert system containing the formal definitions of alliances, which can be applied to new data sets. The new classification system critically revised the previous concepts and integrated them into a consistent framework, which reflects the main gradients in species composition driven by biogeographic influences, macroclimate and the position of the sites in the coast-inland zonation of the dune systems. A revision of the class concept used in EuroVegChecklist is also proposed.