Bias in Ellenberg indicator values — problems with detection of the effect of vegetation type

A recent analysis published in this journal found different relationships between mean Ellenberg indicator values and environmental measurements in different vegetation types. The cause was stated as bias in mean Ellenberg values between relevés which in turn suggested to reflect a bias in individual Ellenberg values. We discuss two phenomena that could explain these results without the need to invoke bias in either individual or mean Ellenberg values. Firstly, slopes of linear regression lines underestimate true relationships when analyses involve explanatory variables measured with error. Secondly, syntaxon‐specific distributions of Ellenberg values follow from the floristic definition of phytosociological units. Mean Ellenberg values per relevé therefore carry the stamp of their associated syntaxon even though associated abiotic conditions may vary between relevés. This will lead to variation in slopes and intercepts between vegetation types not because of bias in individual Ellenberg values but because of prescribed bias in the distribution of Ellenberg values between syntaxa. The residual variation in calibrations carried out across vegetation types is undoubtedly reduced by introducing vegetation type as a factor. However users should note that this is unlikely to reflect bias in individual Ellenberg values but is more likely to reflect error in environmental measurements as well the constraint imposed by phytosociological classification.     

Vegetation maps based on remote sensing are informative predictors of habitat selection of grassland birds across a wetness gradient

Vegetation is a major environmental factor influencing habitat selection in bird species. High resolution mapping of vegetation cover is essential to model the distribution of populations and improve the management of breeding habitats. However, the task is challenging for grassland birds because microhabitat variations relevant at the territory scale cannot be measured continuously over large areas to delineate areas of higher suitability. Remote sensing may help to circumvent this problem. We addressed this issue by using SPOT 5 imagery and phytosociological data. We mapped grassland vegetation in a floodplain using two methods. We (i) mapped the continuous Ellenberg index of moisture and (ii) identified 5 vegetation classes distributed across the wetness gradient. These two methods produced consistent output maps, but they also provided complementary results. Ellenberg index is a valuable proxy for soil moisture while the class approach provided more information about vegetation structure, and possibly trophic resources. In spite of the apparent uniformity of meadows, our data show that birds do not settle randomly along the moisture and vegetation gradients. Overall birds tend to avoid the driest vegetation classes, i.e. the highest grounds. Thus, vegetation maps based on remote sensing could be valuable tools to study habitat selection and niche partition in grassland bird communities. It is also a valuable tool for conservation and habitat management.     

Measurement errors and regression to the mean cannot explain bias in average Ellenberg indicator values

Smart & Scott (2004, this is sue) criticized our paper (Wamelink et al. 2002) about the bias in average Ellenberg indicator values. Their main criticism concerns the method we used, regression analysis. They state the bias can be mimicked by the construction of an artificial data set and that regression analysis is not a suited tool to investigate underlying phenomena. Moreover they claim that the present bias is caused by the distribution of Ellenberg indicator values between syntaxa, instead of a bias in average Ellenberg indicator values per species. We show that their criticism of the use of regression analysis does not hold. We selected average Ellenberg values per vegetation group for several pH classes and applied an F‐test to determine whether or not the vegetation groups within each pH class differed significantly from each other. This was the case for all tested classes (P < 0.001). Moreover we simulated an artificial data set, of which the F‐test for varying measurement error could not explain the magnitude of the F‐value we found earlier. This indicates that the bias we found in average Ellenberg indicator values cannot be explained by measurement errors or by regression to the mean. In the end, Smart & Scott, as we did, come to the conclusion that there is a bias present and that separate regression lines per vegetation type are necessary, but the debate remains open on whether or not this is caused by the bias in Ellenberg indicator values per species.     

Braun‐Blanquet's legacy and data analysis in vegetation science

Abstract. This article investigates whether the Braun‐Blanquet abundance/dominance (AD) scores that commonly appear in phytosociological tables can properly be analysed by conventional multivariate analysis methods such as Principal Components Analysis and Correspondence Analysis. The answer is a definite NO. The source of problems is that the AD values express species performance on a scale, namely the ordinal scale, on which differences are not interpretable. There are several arguments suggesting that no matter which methods have been preferred in contemporary numerical syntaxonomy and why, ordinal data should be treated in an ordinal way. In addition to the inadmissibility of arithmetic operations with the AD scores, these arguments include interpretability of dissimilarities derived from ordinal data, consistency of all steps throughout the analysis and universality of the method which enables simultaneous treatment of various measurement scales. All the ordination methods that are commonly used, for example, Principal Components Analysis and all variants of Correspondence Analysis as well as standard cluster analyses such as Ward's method and group average clustering, are inappropriate when using AD data. Therefore, the application of ordinal clustering and scaling methods to traditional phytosociological data is advocated. Dissimilarities between relevés should be calculated using ordinal measures of resemblance, and ordination and clustering algorithms should also be ordinal in nature. A good ordination example is Non‐metric Multidimensional Scaling (NMDS) as long as it is calculated from an ordinal dissimilarity measure such as the Goodman & Kruskal γ coefficient, and for clustering the new OrdClAn‐H and OrdClAn‐N methods.     

Plant communities and species distribution of low alpine Betula nana heaths in Northernmost Sweden

Summary Low alpine Betula nana heaths were studied with regard to plant communities, species distribution, and environmental conditions. It is clear from cluster analyses and PCA-ordinations that the distribution of winter snow is the most important factor for the composition of plant communities and the distribution of most plant populations. Under deep snow conditions soil instability is also important as a differentiating factor.The numerical treatment of data gives a phytosociological grouping in eoncordance with earlier opinions based on stand tabulations.The cluster analysis group the species into chionophobous, chionophilous, moderate snow bed species, and an additional group of wide snow depth amplitude. Only a few species remain unclassified. The ecological species groups are readily identified also in PCA-ordinations. They are independent of differences in species numbers and combinations.Nomenelature follows Lid (1963), Nyholm (1954–1969), Arnell (1956), and Dahl & Krog (1972).I am grateful to Dr. Eddy van der Maarel, University of Nijmegen, for critical comments on the manuseript, to Dr. Ewa Kvillner, University of Lund, and to my colleagues in Göteborg for generous help and encouragement throughout the study, which forms part of a Ph. D. thesis in the University of Göteborg. The work has been supported by grants from The Royal Swedish Academy of Sciences and from the Faculty of Natural Sciences, University of Göteborg.     

Decline ofPotentillo albae-Quercetum phytocoenoses associated with the invasion ofCarpinus betulus

The study area in the Bialowieza primeval forest-size 28000 m², divided into quadrats of 100 m², comprised two patches ofPotentillo albae-Quercetum as distinguished by phytosociological methods, and a fragment of an adjacent very largeTilio-Carpinetum stand. The directional variation in the distribution ofCarpinus betulus trunk diameter values in the southern part of the study area indicates invasion of this oak forest byCarpinus. This process probably started 1962–1968. The invasion ofCarpinus betulus caused considerable deterioration in light conditions of the ground layer, as deduced from a decrease in Ellenberg light index values. The phytosociological division of the study area correlates with variation in the spatial pattern ofCarpinus mean trunk diameter distribution.Quadrat groups were distinguished by association analysis. They were described phytosociologically using systematic group values for species, constituting the characteristic combination for thePotentillo albae-Quercetum. Areas occupied by still olderCarpinus individuals correspond to successive stages of oak forest regression. The deterioration of light conditions caused recession of thermo-and heliophilous oak-forest species and decline ofPotentillo albae-Quercetum phytocoenoses. However, this was not accompanied by the expansion pansion of species ofCarpinus forests.     

Validity of Ellenberg indicator values judged from physico‐chemical field measurements

Abstract. The relationship between mean Ellenberg indicator values (IV) per vegetation relevé and environmental parameters measured in the field usually shows a large variation. We tested the hypothesis that this variation is caused by bias dependent on the phytosociological class. For this purpose we collected data containing vegetation relevés and measured soil pH (3631 records) or mean spring groundwater level (MSL, 1600 records). The relevés were assigned to vegetation types by an automated procedure. Regression of the mean indicator values for acidity on soil pH and the mean indicator values for moisture on MSL gave percentages explained variance similar to values that were reported earlier in literature. When the phytosociological class was added as an explanatory factor the explained variance increased considerably. Regression lines per vegetation type were estimated, many of which were significantly different from each other. In most cases the intercepts were different, but in some cases their slopes differed as well. The results show that Ellenberg indicator values for acidity and moisture appear to be biased towards the values that experts expect for the various phytosociological classes. On the basis of the results, we advise to use Ellenberg IVs only for comparison within the same vegetation type.     

Optimum-transformation of plant species cover-abundance values

Most of the methods used in the multivariate analysis of data on vegetation and environment, or transformations implied in such methods, put disproportionate emphasis on species with a relatively wide ecological amplitude occurring with relatively high cover-abundance values, and/or rare species. This problem can be overcome to some extent by reducing the cover-abundance values to presence-absence data, but this means a severe loss of information. A standardization of values by species maxima as is done automatically in some programs, may lead to an undesirable emphasis on species represented with low values only.In this paper a method is presented, by which relatively low cover-abundance values of species are upweighted to an arbitrarily chosen higher value, if these low values are considered to indicate an optimum response of that particular species. The method has been tested on a selection of 40 phytosociological relevés from dune slacks in the Voorne dunes, as well as on the Dune Meadow data set used in the textbook of Jongman et al. (1987). The cluster structure obtained with the optimum-transformation appears to be clearer and the contribution of typical dune slack species to the cluster structure increased significantly. Canonical correspondence analysis of the transformed data gave slightly more important main axes.Abbreviations CA = Correspondence Analysis - CCA = Canonical Correspondence Analysis - DOL = Detection of Optimality Level - SWOM = Standardized Weighted Optimality Measure - WPGMA = Weighted Pair Group Method Average linking clustering     

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.     

Shift in Plant Species Composition Reveals Environmental Changes During the Last Decades: A Long-Term Study in Beech (Fagus sylvatica) Forests in Bavaria, Germany

Changes in vegetation composition due to the increasing temperatures in the past few decades have already been reported from several parts of Europe. It has been shown that single species move either northwards or to higher elevations. We expected that the species composition of forest stands should also have changed, i.e., an increase of thermophilous species. Another site factor changing for decades is nitrogen availability; we therefore also expected an increase of nitrophilous species, which was one main result in former long-term studies. We studied the species composition of beech (Fagus sylvatica) forests in southern Germany (Bavaria), comparing old (from 1949 to 1985) and young (2010) phytosociological relevés. Ellenberg indicator values representing plant species specific environmental factors combined with climatic data were used in a partial canonical correspondence analysis (pCCA) for vegetation comparisons. Changes in plant species composition were analyzed considering species frequency, distribution of Ellenberg indicator values, shares of i) non-native plants and ii) tree, shrub and herbaceous species. Contrary to our expectations, global warming in Bavaria during the past decades resulted only in an explained dispersion of 5 % in the species composition. On the species level, an overall increase of thermophilous walnut tree saplings (Juglans regia) was conspicuous in some study areas. Nitrophilous species, however, generally increased in frequency throughout the study areas. Throughout Bavaria the most significant change was a striking increase of juvenile tree species and a decrease of herbaceous species. Up to now the increased nitrogen input into forests had a stronger provable influence on species composition shift than the global warming of the last decades. Additionally general changes in forest management also had effects on forest species. Therefore community reorganization mirroring temperature factors in beech forests in central Europe seems to be only at its very beginning.     

Adaptive leaf structures in a Myrica-Erica stand on Tenerife (Canary Islands)

This study is based on a phytosociological table and a matrix of species and morphological characters. The relevés were taken along a transect in a Myrica faya-Erica arborea formation on Tenerife (Canary Islands). The analysis of the phytosociological table reveals a vegetational succession and a corresponding ecological gradient. The study of correlations between morphological and ecological data leads to the creation of groups of species with different combinations of characters, and to a discussion of the adaptive values of each character combination. The species of each group occupy similar niches.Abbreviations AOC= Analysis of Concentration - P/C= index ratio between the perimeter of a leaf and the circumference of a circle having the same area as the leaf     

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.     

Genetic patterns in the Lathyrus pannonicus complex (Fabaceae) reflect ecological differentiation rather than biogeography and traditional subspecific division

The endangered European relict species complex Lathyrus pannonicus shows distinct morphological variation, reflected by the number of subspecies recognized, and complicated patterns of genetic variation. The traditionally recognized subspecies appear to possess different ecological preferences and disjunct distributions, particularly in the western range of the species. In this study, L. pannonicus was investigated by the correlation of distance matrices based on phytosociological, ecological, molecular and morphological data. Ecological characteristics of selected stands of L. pannonicus throughout Europe were assessed using ‘Ellenberg values’ of all the constituent taxa in phytosociological relevés. Genetic distances were calculated using recently developed methods to analyse high degrees of intra‐individual nuclear‐encoded internal transcribed spacer variability. We found that the remarkable genetic (and morphological) diversity in L. pannonicus could not be explained solely by the fragmentation of the distributional range. Instead, patterns of morphological and genetic differentiation were a reflection of the moisture regime in the sampled stands. Two major lineages could be identified: (1) a lineage adapted to dry conditions (Ellenberg indicator F‐value ≤ 3.5) and (2) a lineage preferring moist conditions (F‐value ≥ 4.5). Although both lineages occurred in close proximity in the Pannonian area, they appeared to be reproductively isolated in general. Further data are needed to determine whether these genetically and ecologically defined lineages, or ecospecies, within the L. pannonicus species complex can be formalized as (Linnaean) species or subspecies. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 402–421.     

Phytosociological amplitudes of some Dutch coastal species and their ecological interpretation

A 4-axis Reciprocal Averaging ordination of phytosociological tables of Dutch plant communities revealed environmentally interpretable gradients: (1) overall moisture, (2) salinity, (3) soil fertility, and (4) structural complexity of vegetation. The quantitative occurrence of all species along these gradients has been computed. Some examples of the distribution of species from coastal communities are given, to show how these data may help to summarize and visualize existing knowledge and how they can form a basis for further study.Nomenclature of species follows Heukels-van Ooststroom (1975), that of syntaxonomical units Westhoff & den Held (1969).     

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.     

Vegetation of beech forests in the Rychlebské Mountains,Czech Republic,re-inspected after 60 years with assessment of environmental changes

From 1941–;1944 nearly 30 phytosociological relevés were completed by F. K. Hartmann in the Rychlebské Mountains, a typical mountainous area in northeastern Czech Republic. Of the original plots still covered with adult grown beech (Fagus sylvatica) forest, 22 were resampled in 1998 and 1999. In order to describe the recent vegetation variability of the sites 57 relevés were recorded. Changes in vegetation were estimated using relative changes in species density and ordinations (PCA, RDA). Environmental changes were assessed using Ellenberg indicator values when no direct measurements were available. A decline in species diversity has been documented, particularly, many species occurring frequently in deciduous forests with nutrient and moisture well-supplied soils around neutral have decreased. In contrast, several light-demanding, acid- and soil desiccation-tolerant species have increased. Natural succession, quantified as forest age, contributed slightly to these changes. In Ellenberg indicator values, a decline in F (soil moisture), R (soil calcium) and N (ecosystem productivity), and an increase in L (understorey light) were shown. This is interpreted as the influence of modified forestry management and of airborne pollutants. Intensified logging caused the canopy to open and soil conditions to worsen. The latter is most likely also due to acid leaching of soil cations (Ca, K, Na). This caused a decline in soil productivity, thus the effect of nitrification could not be detected. The original relevés may have differed in size influencing the results. This revised version was published online in August 2006 with corrections to the Cover Date.     

A probabilistic approach to estimating species pools from large compositional matrices

Abstract. Species pools are increasingly recognized as important controls of local plant community structure and diversity. While existing approaches to estimate their content and size either rely on phytosociological expert knowledge or on simple response models across environmental gradients, the proposed application of phytosociological smoothing according to Beals exploits the full information of plant co‐occurrence patterns statistically. Where numerous representative compositional data are available, the new method yields robust estimates of the potential of sites to harbour plant species. To test the new method, a large phytosociological databank covering the forested regions of Oregon (US) was subsampled randomly and evenly across strata defined by geographic regions and elevation belts. The resulting matrix of species presence/absence in 874 plots was smoothed by calculating Beals' index of sociological favourability, which estimates the probability of encountering each species at each site from the actual plot composition and the pattern of species co‐occurrence in the matrix. In a second step, the resulting lists of sociologically probable species were intersected with complete species lists for each of 14 geographical subregions. Species pools were compared to observed species composition and richness. Species pool size exhibited much clearer spatial trends than plot richness and could be modelled much better as a function of climatic factors. In this framework the goal of modelling species pools is not to test a hypothesis, but to bridge the gap between manageable scales of empirical observation and the spatio‐temporal hierarchy of diversity patterns.     

What makes a plant species specialist in mixed broad-leaved deciduous forests?

Knowledge about relationships between specialization degree of species, i.e. the width of their realized niche and functional traits, may have important implications for the assessment of future population developments under environmental change. In this study, we used a recently introduced method to calculate ecological niche widths of plant species in mixed broad-leaved deciduous forests and to investigate the dependence between niche widths of plants and their functional traits and Ellenberg indicator values. The research is based on a dataset of 4556 phytosociological relevés of mixed broad-leaved deciduous forests in Slovenia. We calculated theta indices for 326 species, which ranks them along a continuous gradient of habitat specialization. For 272 species, we compiled 26 functional traits and Ellenberg indicator values. We found some significant correlations between theta indices of species and their functional traits and Ellenberg indicator values; habitat specialists thrive primarily on the highest altitudes, on colder, dry sites and achieve the age of first flowering later than generalists. They also have smaller seed diameter, lower leaf dry matter content, lower mean canopy height and bigger specific leaf area than generalists. Two species groups, chamaephytes and spring green species, are particularly characterized as specialist species. The added value of our work is in complementing the knowledge about the niche differentiating along different environmental gradients and species coexistence in mixed broad-leaved deciduous forests.     

Diversity of subalpine and alpine vegetation of the eastern part of the Nízke Tatry Mts in Slovakia: major types and environmental gradients

Changes in composition and structure of alpine and subalpine plant communities in relation to ecological factors were analysed in the Nízke Tatry Mts, Slovakia. Species cover values of vascular and non-vascular plants in each vegetation plot were recorded on the nine-degree scale. A data set of 156 relevés of alpine and subalpine vegetation was sampled recently during one year in the eastern part of the Nízke Tatry National Park. The data set was analysed by cluster analysis and Detrended Correspondence Analysis. analyses were carried out on the entire data set, including the subset of short grassland and dwarf-shrub vegetation. Major gradients and clusters were ecologically interpreted using Ellenberg indicator values. In the entire data set, the major gradient in species composition was associated with nutrient availability and the second most important gradient with light. In the case of short grassland and dwarf-shrub vegetation, the gradients were different. The first one was associated with soil reaction and the second gradient was associated with moisture. Clusters proposed by numerical classification reproduced many traditional phytosociological associations, namely Seslerietum distichae, Sphagno capillifolii-Empetretum nigri, Junco trifidi-Callunetum vulgaris, Juncetum trifidi, Dryopterido dilatatae-Pinetum mugo, Luzuletum obscurae, Agrostio pyrenaiceae-Nardetum strictae, while some other associations were less clearly differentiated (communities of the alliances Calamagrostion villosae, Adenostylion alliariae, Trisetion fusci, Cratoneuro filicini-Calthion laetae or Salicion herbaceae). The next clusters included Vaccinium and Festuca supina dominated communities and artificial roadside grasslands sown 50 years ago. Bryophytes and lichens were highly represented among diagnostic species of particular associations. Distribution pattern of particular plant communities was strongly influenced by site position either on northern or southern slope of the mountains.