Ecological preferences of alien plant species in North-Eastern Germany

The large, comprehensive vegetation database of Mecklenburg-Vorpommern/NE Germany with 51,328 relevés allowed us to study an entire regional flora of 133 non-native plants (NNP, immigration after 1492 AD) with regard to their preferences to all kinds of habitats and along different ecological gradients. For each relevé, we computed average Ellenberg indicator values (EIV) for temperature, light, moisture, reaction, nutrients and salt as well as plant strategy type weights. We partitioned the dataset into relevés with and without occurrences of NNP and compared them with respect to the relative frequencies of EIVs and strategy type weights. We identified deviations from random differences by testing against permuted indicator values. To account for bias in EIV between community types, NNP preferences were differentiated for 34 phytosociological classes. We tested significance of preferences for the group of NNP as a whole, as well as for single NNP species within the entire dataset, as well as differentiated by phytosociological classes and formations. NNP as a group prefer communities with high EIVs for temperature and nutrients and low EIVs for moisture. They avoid communities with low EIV for reaction and high EIV for salt. NNP prefer communities with high proportions of ruderal and low proportion of stress strategists. The differentiation by phytosociological classes reinforces the general trends for temperature, nutrients, moisture, R and S strategy types. Nevertheless, preferences of single species reveal that NNP are not a congruent group but show individualistic ecological preferences.     

Phytosociological data give biased estimates of species richness

Abstract. Large phytosociological data sets of three types of grassland and three types of forest vegetation from the Czech Republic were analysed with a focus on plot size used in phytosociological sampling and on the species‐area relationship. The data sets included 12975 relevés, sampled by different authors in different parts of the country between 1922 and 1999. It was shown that in the grassland data sets, the relevés sampled before the 1960s tended to have a larger plot size than the relevés made later on. No temporal variation in plot sizes used was detected in forest relevés. Species‐area curves fitted to the data showed unnatural shapes, with levelling‐off or even decrease in plot sizes higher than average. This distortion is explained by the subjective, preferential method of field sampling used in phytosociology. When making relevés in species‐poor vegetation, researchers probably tend to use larger plots in order to include more species. The reason for this may be that a higher number of species gives a higher probability of including presumed diagnostic species, so that the relevé can be more easily classified in the Braun‐Blanquet classification system. This attitude of phytosociologists has at least two consequences: (1) in phytosociological data bases species‐poor vegetation types are underrepresented or relevés are artificially biased towards higher species richness; (2) the suitability of phytosociological data for species richness estimation is severely limited.     

Bayesian classification of vegetation types with Gaussian mixture density fitting to indicator values

Question: Is it possible to mathematically classify relevés into vegetation types on the basis of their average indicator values, including the uncertainty of the classification? Location: The Netherlands. Method: A large relevé database was used to develop a method for predicting vegetation types based on indicator values. First, each relevé was classified into a phytosociological association on the basis of its species composition. Additionally, mean indicator values for moisture, nutrients and acidity were computed for each relevé. Thus, the position of each classified relevé was obtained in a three‐dimensional space of indicator values. Fitting the data to so called Gaussian Mixture Models yielded densities of associations as a function of indicator values. Finally, these density functions were used to predict the Bayesian occurrence probabilities of associations for known indicator values. Validation of predictions was performed by using a randomly chosen half of the database for the calibration of densities and the other half for the validation of predicted associations. Results and Conclusions: With indicator values, most reléves were classified correctly into vegetation types at the association level. This was shown using confusion matrices that relate (1) the number of relevés classified into associations based on species composition to (2) those based on indicator values. Misclassified relevés belonged to ecologically similar associations. The method seems very suitable for predictive vegetation models.     

Vegetation and environmental factors during primary succession on glacier forelands: Some outlines from the Italian Alps

ABSTRACT

Relationships between plant communities and the physical environment during primary succession on recently deglaciated glacier forelands were studied in 3 areas of the Italian Alps. The aim of the research was to relate traditional phytosociological data with environmental variables. Twenty-eight phytosociological relevés were performed, each associated with twenty-six environmental variables; quantitative parameters of richness and diversity were also calculated. Species/relevés, environmental variables/relevés and species/environmental variables matrices were analyzed by cluster analysis, PCA and Spearman correlation coefficient. Three main stages of succession were identified by floristic composition and confirmed by environmental parameter evaluation. A complex of environmental variables seems to be closely correlated with terrain age and richness/diversity parameters, even though diversity decreases in late successional stages. The phytosociological significance of species is in accordance with their position in the context of succession.     

A phytosociological classification of Swiss mire vegetation

The mapping and monitoring of Swiss mires has so far relied on a classification system based on expert judgement, which was not supported by a quantitative vegetation analysis and which did not include all wetland vegetation types described in the country. Based on a spatially representative sample of 17,608 relevés from 112 Swiss mires, we address the following questions: (1) How abundant are wetland vegetation types (phytosociological alliances) in Swiss mires? (2) How are they distributed across the country––is there a regional pattern? (3) How clearly are they separated from each other? (4) How clear and reliable is their ecological interpretation? Using published wetland vegetation relevés and lists of diagnostic species for phytosociological units (associations and alliances) established by experts, we developed a numerical method for assigning relevés to units through the calculation of similarity indices. We applied this method to our sample of 17,608 relevés and estimated the total area covered by each vegetation type in Switzerland. We found that vegetation types not included in previous mapping were either rare in Switzerland (partly due to mire drainage) or poorly distinguished from other vegetation units. In an ordination, the Swiss mire vegetation formed a triangular gradient system with the Sphagnion medii, the Caricion davallianae and the Phragmition australis as extreme types. Phytosociological alliances were clearly separated in a subset of 2,265 relevés, which had a strong similarity to one particular association, but poorly separated across all relevés, of which many could not be unequivocally assigned to one association. However, ecological gradients were reflected equally well by the vegetation types in either case. Overall, phytosociological alliances distinguished until now proved suitable schemes to describe and interpret vegetation gradients. Nevertheless, we see the urgent need to establish a data base of Swiss wetland relevés for a more reliable definition of some vegetation units.     

Neophyte Invasion in Moldavia (Eastern Romania) in Different Habitat Types

The actual state of neophyte invasion in Moldavia (Eastern Romania) is described in this paper on the basis of 11,055 phytosociological relevés. We analyzed the i) proportion of relevés with neophytes, ii) mean proportion of neophytes per relevés, and iii) mean coverage of neophytes per relevé for 36 EUNIS habitat types to identify general plant invasion patterns. The level of invasion differed considerably between habitats. The invasion of neophytes especially affected habitats strongly determined or influenced by man, such as anthropogenic woodlands, ruderal habitats, arable lands or trampled areas. Most natural habitats are either slightly invaded or entirely free of neophytes. Only riverine willow stands and wet tall-herb stands are relatively highly invaded. However, the absence of neophytes in some natural habitats less represented in the phytosociological dataset could be of artifactual nature. No significant relationship between the number of neophytes and non-neophytes was found in the analysis across different habitats. When the analyses were made within-habitats, both negative and positive relationships were found, which confirm that the relationship between alien and native species richness depends on the habitats. A total number of 105 neophyte species were recorded in the phytosociological relevés used in this study. Among these, 13 species that are currently considered invasive in Moldavia occur in at least 10 types of habitats.     

Life-history traits and habitat preferences of colonizing plant species in long-term spontaneous succession in abandoned gravel–sand pits

The following question was asked: Species of what traits and ecological demands are the best and the least successful colonizers in the course of succession in disused gravel–sand pits?Abandoned gravel–sand pits (36) were studied in the Czech Republic. Together 224 phytosociological relevés were recorded in 5 m×5 m plots in all available seral stages ranging in age from 1 to 75 years. Complete lists of species occurring in (semi-)natural habitats were surveyed up to a distance of 100 m from each relevé. The colonization success of each species was expressed by an index between 0 and 1 which was obtained as the ratio of the number of relevés with species present in a pit/the number of relevés with the species occurrence in their surroundings. Species characteristics were taken from available databases. Data were elaborated by ordination analysis and regression tree analysis. Generally, the best colonizers of gravel–sand pits were hydrophytes with the ability to vegetatively reproduce. In particular, at the beginning of succession, anemochorous species played the most important role. Stress tolerant species, possessing light diaspores, were typical of nitrogen poor and acidic habitats. Later, sciophilous and nitrophilous species of mesic habitats increased in importance. They have heavier diaspores, and are mostly phanerophytes or geophytes, often with the ability to vegetatively reproduce as well as exhibiting higher demand for pH. Plant functional traits were recognized as powerful tools to predict the colonization success of plants available in the local species pool. They may help predict vegetation succession in various human-disturbed sites and thus be used in various restoration programmes.     

A simple method for estimating homotoneity of sets of phytosociological relevés

The homogeneity of sets of phytosociological relevés or syntaxa—the homotoneity—is tested by means of a homotoneity-coefficient. This coefficient consists of the basic homotoneity-coefficient expressing the proportion of highly constant species (61–100%) in the species composition of the average relevé and the correcting factor which is proportional to the difference of species number between the richest and poorest relevé. Successive analysis of heterotoneity of sets of relevés for detecting sources of heterotoneity is proposed.     

Arranging phytosociological tables by species-relevé groups

Abstract. A newmethod of arranging phytosociological tables by species-relevé groups (blocks) on the basis of the density of rows and columns (relative species frequency and relative species number) is presented. Each block fulfils a given minimum criterion regarding its density. With a density threshold of e.g. 50 %, blocks are formed with only those species that are present in at least 50 % of the block's relevés, while at the same time the block's relevés comprise at least 50 % of the block's species. The procedure starts with the complete table and iteratively masks species and relevés with the smallest densities. This is performed step by step with species and relevés and continues until the given density threshold for all species and relevés is exceeded. Without any further parameters other than the minimum density, species-relevé groups of gradually decreasing size are formed. Data processing is performed with a computer program (ESPRESSO). The main application of the method is found in an effective pre-sorting of relevé data. The blocks formed can be arranged in the final table as desired.     

Random sampling does not exclude spatial dependence: The importance of neutral models for ecological hypothesis testing

Lájer (2007) notes that, to investigate phytosociological and ecological relationships, many authors apply traditional inferential tests to sets of relevés obtained by non-random methods. Unfortunately, this procedure does not provide reliable support for hypothesis testing because non-random sampling violates the assumptions of independence required by many parametric inferential tests. Instead, a random sampling scheme is recommended. Nonetheless, random sampling will not eliminate spatial autocorrelation. For instance, a classical law of geography holds that everything in a piece of (biotic) space is interrelated, but near objects are more related than distant ones. Because most ecological processes that shape community structure and species coexistence are spatially explicit, spatial autocorrelation is a vital part of almost all ecological data. This means that, independently from the underlying sampling design, ecological data are generally spatially autocorrelated, violating the assumption of independence that is generally required by traditional inferential tests. To overcome this drawback, randomization tests may be used. Such tests evaluate statistical significance based on empirical distributions generated from the sample and do not necessarily require data independence. However, as concerns hypothesis testing, randomization tests are not the universal remedy for ecologists, because the choice of inadequate null models can have significant effects on the ecological hypotheses tested. In this paper, I emphasize the need of developing null models for which the statistical assumptions match the underlying biological mechanisms.     

SaudiVeg ecoinformatics: Aims,current status and perspectives

During the last decade many electronic databases of vegetation plots were established in many countries around the world. These databases contain valuable phytosociological information assisting both governmental and NGO (Non-governmental organizations) agencies to formulate strategies and on-ground plans to manage and protect nature resources. This paper provides an account on aims, current status and perspectives of building of a vegetation database for the Central Region (Najd) of Saudi Arabia – the founding element of the Saudi Vegetation Database (SVD). The data stored by the database are sample plots (vegetation relevés) collected according to the field techniques of the Braun-Blanquet approach (lists of taxa accompanied by semi-quantitative cover assessment), and are accompanied by general vegetation characteristics such as vegetation layering and cover, information on life-form of the recorded species, geographical coordinates, altitude, soil typology, topography and many more. More than 2900 vegetation-plot records (relevés) have so far been collected in the Najd region; of these more than 2000 have already been stored using the Turboveg database platform. These field records cover many habitats such as depressions, wadis (dry river beds), agricultural lands, sand dunes, sabkhas, and ruderal habitats. The ecological information collected in the database is currently the largest set of vegetation data collated into a database in the Middle East. These data are of great importance for biodiversity studies in Saudi Arabia, since the region is recording a loss of biodiversity at a fast rate due to environmental problems such as global warming and land-use changes. We envisage that this database would catalyze further data collection on vegetation of the entire Arabian Peninsula, and shall serve as one of the most important datasets for classification and mapping of the vegetation of the Kingdom of Saudi Arabia.     

Computer assisted floristic similarity analysis

A program for polythetic numerical evaluation of phytosociological material is described. Using Sörensen's coefficient of floristic similarity it computes the homogeneity of subjectively chosen sets of relevés, the affinity of each constituent relevé to the set as well as the similarity between any pair of sets. It is also able to plot a dendrogram of the hierarchic system obtained by agglomerating individual relevés into complex groupings. The program is suitable for processing phytosociological data arranged in any conventional table.     

Re-assessing the ecology of rare flood-meadow violets (<Emphasis Type="Italic">Viola elatior,V. pumila</Emphasis> and<Emphasis Type="Italic">V. persicifolia</Emphasis>) with large phytosociological data sets

This study demonstrates how conventional ecological knowledge on species together with models resulting from functional traits can be tested and refined by tapping large data sources that have been made available through recent electronic compilations. The study is based on the comparison of three rare, closely related flood-meadow violet species, which have been supposed to have similar ecological behaviour and niche occupation. In contrast, the analysis of 335 Central European relevés using different methods of numerical ordination, classification and calibration revealed distinct differences in habitat preferences between the three species. Detrended correspondence analysis ordination, and multiple-response permutation procedures and TWINSPAN classification displayed the separation ofViola persicifolia fromV. elatior andV. pumila along a moisture and base-richness gradient, while the latter two differed mostly in terms of mowing compatibility. Although the three violets are considered to be weak competitors they may be found under nutrient-poor as well as nutrient-rich site conditions. The distribution of C-S-R strategy types in relevés supported the hypothesis that at more fertile sites the violets crucially depend on disturbance by management or flooding events that create gaps and weaken strong competitors, while at less productive sites they may persist for a long time even under fallow conditions. The S/R strategy is shared by all three violets. Problems and perspectives arising with the use of phytosociological databanks as a source of ecological information are discussed.     

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.     

Quantitative prediction of the distribution and abundance of Vaccinium myrtillus with climatic and edaphic factors

Question: Can the distribution and abundance of Vaccinium myrtillus be reasonably predicted with soil nutritional and climatic factors? Location: Forests of France. Methods: We used Braun‐Blanquet abundance/dominance information for Vaccinium myrtillus on 2905 forest sites extracted from the phyto‐ecological database EcoPlant, to characterize the species ecological response to climatic and edaphic factors and to predict its cover/abundance at the national scale. The link between cover/abundance of the species and climatic (65 monthly and annual predictors concerning temperature, precipitation, radiation, potential evapotranspiration, water balance) and edaphic (two predictors: soil pH and C:N ratio) factors was investigated with proportional odds models. We evaluated the quality of our model with 9830 independent relevés extracted from Sophy, a large phytosociological database for France. Results: In France, Vaccinium myrtillus is at the southern limit of its European geographic range and three environmental factors (mean annual temperature, soil pH and C:N ratio) allow prediction of its distribution and abundance in forests with high success rates. The species reveals a preference for colder sites (especially mountains) and nutritionally poor soils (low pH and high C:N ratio). A predictive map of its geographic range reveals that the main potential habitats are mountains and northwestern France. The potential habitats with maximal expected abundance are the Vosges and the Massif central mountains, which are both acidic mountains. Conclusions: Complete niche models including climate and soil nutritional conditions allow an improvement of the spatial prediction of plant species abundance at a broad scale. The use of soil nutritional variables in distribution models further leads to an improvement in the prediction of plant species habitats within their geographical range.     

Malcolmia littorea: The isolated Italian population in the European context

We compared the coenological information of the only Italian population of Malcolmia littorea (L.) R. Br. with published phytosociological relevés, including ones of this species, throughout its European range. With the aim of highlighting the main climatic features influencing the distribution patterns of M. littorea, we integrated coenological data with some climatic variables and considered major drivers for plant distribution at the European scale. Finally, we analysed the population extent of M. littorea in Italy, in order to assess its conservation status at regional level. The DCA analysis, performed on a matrix 139 relevés × 183 species, separated the relevés according to their floristic composition, showing a geographic gradient from Portugal to Mediterranean coasts, until Italy; with Mediterranean relevés clearly separated from Atlantic ones as well. Along the beach-inland gradient, the analysis highlights that the species is typical of fixed dunes habitats (more inland, mainly stabilised dunes), although in the Atlantic it can also be found in mobile dunes. The analysis of climatic variables in relation to M. littorea distribution, suggested that the species is sensitive to low winter temperature and to summer drought. The only Italian population of M. littorea is subjected to many threats, due to its small dimensions (<1 ha), isolation from the rest of its distribution area, that ranges from Portugal to France (until the Camargue region) and intensive human disturbances. Using both field and remote sensing information, we showed a considerable decrease of the occupied surface in Italy, leading us to suggest that the IUCN threat category of M. littorea in Italy should be reassessed from endangered to critically endangered.     

JUICE,software for vegetation classification

Abstract. The program JUICE was designed as a Microsoft® WINDOWS® application for editing, classification and analysis of large phytosociological tables and databases. This software, with a current maximum capacity of 30 000 relevés in one table, includes many functions for easy manipulation of table and header data. Various options include classification using COCKTAIL and TWINSPAN methods, calculation of interspecific associations, fidelity measures, average Ellenberg indicator values, preparation of synoptic tables, automatic sorting of relevé tables, and export of table data into other applications (word processors, spreadsheet programs or mapping packages). JUICE is optimized for use in association with TURBOVEG which is the most widespread database program for storing phytosociological data in Europe.     

The Quercus petraea-dominated communities in Italy: Floristic,coenological and chorological diversity in an European perspective

In Italy, woods dominated by Quercus petraea are well documented within local and regional phytosociological studies but a critical revision of their principal ecological-floristic components based on a comprehensive Italian data set was missing. We gathered 209 published and unpublished Italian phytosociological relevés, where Q. petraea was dominant, that were investigated by means of multivariate analysis. The ecological requirements of the resulting groups were indirectly calculated by means of Ellenberg indicator value, and a chorological analysis was performed. The fidelity coefficient (phi) for the diagnostic species of each group was calculated. According to our analysis, five different types of Q. petraea woods were found to be present in Italy. Each group was characterized by means of its floristic, ecological and chorological components. These resulting five groups were further compared with similar Q. petraea-dominated woods recorded in other European locations, as to the floristic and chorological components. We collected 57 synoptic tables of Q. petraea communities coming from all over Europe. This data set, together with the Italian relevés, was submitted to exploratory multivariate analysis using also the chorological information. According to these analyses, the Italian woods dominated by Q. petraea can be subdivided in two main clusters: the communities belonging to It-1 group (Alps) fell within the central European communities, distinguished by the predominance of large distribution and European chorotypes. All the other Italian groups of relevés resulted to be positioned in the cluster which includes central southern, central eastern and eastern European Q. petraea communities. From the syntaxonomical point of view, the results of this analysis allow to attribute the Italian Q. petraea woods to seven alliances of three different orders. The ecological similarities between Erythronio-Quercion petraeae and Erythronio-Carpinion betuli are highlighted and the distribution of Erythronio-Quercion petraeae is limited to northern Apennines, in particular, to slopes facing the Po Valley.     

The influence of <Emphasis Type="Italic">Ceratophyllum demersum</Emphasis> L. and <Emphasis Type="Italic">Stratiotes aloides</Emphasis> L. on richness and diversity of aquatic vegetation in the lakes of mid-eastern Poland

The aim of this study was to estimate the influence of Ceratophyllum demersum L. and Stratiotes aloides L. on the species richness and phytocoenotic diversity of aquatic vegetation in lakes. The study was based on field investigations in the Łęczna-Włodawa Lake District located in mid-eastern Poland. The studied lakes (32) included both polymictic and dimictic reservoirs. Both in the polymictic and dimictic lakes, statistically significant correlation coefficients were obtained between the parameters that characterize Stratiotes aloides (the frequency in the phytosociological relevés, the percent share of Stratiotetum aloidis association in the phytolittoral) and those characteristic for species richness (the total number of hydromacrophytes, the number of charophytes, elodeids, nymphaeids and lemnids), phytocoenotic richness (the mean number of species in the phytosociological relevés) and phytocoenotic diversity (the Shannon–Wiener Index). The communities with a share of S. aloides are characterized by higher species richness. However, not a single case of statistically significant dependency was reported between the parameters that describe C. demersum (the frequency in the phytosociological relevés, the percent share Ceratophylletum demersi association in the phytolittoral) and the vegetation traits in both lake groups. Common occurrence of Stratiotes aloides might be considered to be evidence for good status of an aquatic ecosystem and a very good indicator of species richness and diversity of aquatic vegetation.