Numerical classification and ordination of ruderal plant communities (Sisymbrietalia,Onopordetalia) in the western part of Slovakia

The ruderal communities of the orders Sisymbrietalia and Onopordetalia from the western part of Slovakia have been subjected to numerical classification and ordination. The ordination techniques proved to be a useful tool in the elucidation of the cluster pattern as well as in the detection of the main environmental variation underlying the floristic variation within the data. Results obtained with numerical techniques and traditional syntaxonomical classification have been compared. The similarity between these results is low at the level of the orders. This incompatability is explained by the differences in the weighting of the species in the course of the classification process and by the addition of non-floristical criteria that often occurs in syntaxonomical classification according to Braun-Blanquet. The highest value has been observed at the 3-clusters level (both orders and the Malvion neglectae). High similarity among the results of the numerical techniques have been observed, particularly in the group of space-dilating clusterings (Ward's method, Complete linkage clustering and MeQuitty's similarity analysis). Average linkage clustering produces the most diverse result. The Malvion neglectae appeared as a separate group in all numerical techniques adopted. This suggests the upranking of its syntaxonomical position. The Bromo-Hordeion murini turned out to be a very heterotoneous syntaxon.     

A numerical-syntaxonomical study of the Calthion palustris Tx. 37 in the Netherlands

Summary After some remarks on ambiguities in the conventions of the Braun-Blanquet approach of phytosociology, a syntaxonomical study is presented of theCalthion palustris (wet grassland vegetation) of the Netherlands and surroundings. A numerical analysis of the non-optimal vegetation yielded a classification well comparable with the one established before. One new association is described. The pattern of variation in this dataset is compared in several ways with the one described elsewhere. Synoptic tables of different associations from several authors were compared by numerical analyses. The results suggested a large geographical variation within theCalthion of Western Europe.The authors wish to thank Mr H.M. van der Steeg (Nijmegen) and Mr J.C. Smittenberg (Assen) for kindly placing some unpublished relevés at their disposal, as well as several members of the plant sociology working group of the Nederlandse Jeugdbond voor Natuurstudie.     

Pre-climax epiphyte communities of bryophytes and lichens in Mediterranean forests from the Central Plateau (Spain)

The pre-climax epiphytic communities of forests from the Spanish Mediterranean region (Central Plateau) were studied by numerical and traditional floristic methods. One hundred phytosociological relevés were analyzed through a classical numerical approach based on hard partitions improved by PCoA — Principal Coordinates Analysis — ordinations. Two groups easily related to two different suballiances included in Frullanion dilatatae Lecointe 1975, Ulotenion crispae (Barkman 1958) Lecointe 1975 and Fabronienion pusillae Barkman 1958 were detected. Two new syntaxa within the ass. Ortotrichetum lyellii (Allorge 1922) Lecointe 1975 are proposed. Phytogeography, syntaxonomy and ecology of these communities are discussed.     

A phytosociological survey of the boreal forest (shape Vaccinio-Piceetea) in North America

A survey of syntaxa of vegetation of North American boreal forests (class Vaccinio-Piceetea) is presented. This phytosociological survey, carried out combining the Braun-Blanquet method with numerical syntaxonomical analyses (cluster and correspondence analysis), describes the associations of the North American boreal forests, which have several species, varieties or vicariant species in common with their Eurasian counterparts, and can be placed in the class Vaccinio-Piceetea. By means of tabular and multivariate analyses, 2084 North American relevés were compared with 3273 relevés from European, Japanese and Korean boreal forest, to describe and typify 4 orders, 10 alliances and 37 associations. Diagnostic tables, ordination, clustering, and climatic, edaphic and biogeographical data were used to show floristic affinities among these syntaxa and interpret their distribution areas. Syntaxa were briefly characterized by their floristic composition, physiognomy, succession, zonation, and biogeographical distribution.     

Numerical phytosociology of the subalpine belt of the Kazbegi region,Caucasus, USSR

A vegetational transect across a glacial valley was investigated. Structure and species composition, as well as influence of disturbance by biotic factors (grazing of sheep and cattle) and abiotic factors (topography, wind, water) are discussed.According to species composition and community structure, 4 main vegetation types were distinguished by comparing field data from phytosociological relevés with results of numerical treatments (3 methods of numerical classification and DCA ordination): 1) Chamaesciadium acaule-Polygonum viviparum, 2) Festuca cf. woronowii-Carex tristis, 3) Veratrum lobelianum-Taraxacum stevenii and 4) Rhododendron caucasicum-Vaccinium myrtillus communities.Species diversity (evenness, and number of species) was low in the ungrazed dwarf-shrub alpine communities, which were dominated by Rhododendron caucasicum, in comparison with the grazed meadow. The highest values of these characteristics were found in the transitions among the types.     

New similarity indices for the assignment of relevés to the vegetation units of an existing phytosociological classification

Every proposed vegetation classification is sooner or later confronted with an accumulation of new data, which has to be assigned to existing vegetation units. Calculation of similarity indices between new relevés (vegetation plots) and constancy columns of established vegetation units is a suitable method for computerised assignment of relevés to these units. This paper compares several similarity indices using simulated data set where either randomly distributed or diagnostic species prevail in the species composition of the tested relevé. Traditional indices, based only on species composition, produce different results than similarity indices that consider species fidelity. However, both types of indices failed in some situations and thus cannot be widely accepted as suitable methods of additional relevé assignment. Therefore a combined Frequency-Positive Fidelity Index (FPFI) is proposed. This new index includes compositional similarity of an assigned relevé with vegetation unit and retains the advantages and lacks the disadvantages of tested indices. The calculation of all these indices is available in the JUICE program (http://www.sci.muni.cz/botany/juice.htm).     

Interspecific associations in phytosociological data sets: how do they change between local and regional scale?

Interspecific associations detected in phytosociological data sets sampled in local areas can reflect locally specific combinations of environmental factors and may thus differ from the interspecific associations existing on a regional scale. As a result, vegetation units derived from numerical classifications of local data sets can accurately reflect local environmental gradients, but their boundaries or spectra of diagnostic species must be frequently adjusted when transferred to the regional scale. Local vegetation classifications can be useful for some purposes, but regional classifications are superior, as they facilitate communication among the researchers from different areas. We demonstrated changes in interspecific associations between regional and local scale, using a data set of 14 589 relevés of herbaceous vegetation of the Czech Republic, and 16 local subsets of this national data set. We focused on sociological species groups, derived statistically in the national data set. Changes in coherence of these groups when applied to the local data sets were described on the basis of statistical association between the relevés containing some species of these groups and the species belonging vs. not belonging to these groups. The results were summarized using the principal components analysis (PCA). In addition, relevé data sets were compared with respect to presence/absence of sociological species groups, using the principal coordinate analysis (PCoA). The results of PCA and PCoA were compared by Procrustean analysis. Local data sets differed from the national data set to different extent. The national data set was more remote to the local data sets if the analysis focused on the coherence of species group rather than on presence/absence. The species groups from the national data set retained most of their coherence in low-altitude hilly landscapes with thermophilous flora, i.e., the most diverse landscape type of the Czech Republic. On the other hand, many species groups from the national data set could not be recognized in mountainous areas or flat lowlands. These results suggest that interspecific associations existing on regional scale are best reproduced in those local areas which have a high habitat heterogeneity or which have a central position along the major gradients existing on regional scale.     

Classification of South Swedish Isoetid vegetation with the help of numerical methods

Relevés of Isoetid vegetation from 60 lakes in southern Sweden have been classified with the help on numerical methods. A community system is constructed, at the variant level by clustering with the TABORD program, and at the community and subcommunity levels by reference to traditional floristic characteristics. The diagnostic species were selected with the help of PCA. Isoëtes lacustris, Lobelia dortmanna and Littorella uniflora determined the community composition together with Eleocharis acicularis. The differentation of the syntaxa along the first three ordination axes of a PCA was clear. The complex water depth factor determines clearly the position of the small syntaxa (variants) in the PCA ordination space. At high levels of similarity the relevés were grouped effectively with the TABORD program, so that the clusters are floristically homogeneous and easy to identify on the basis of floristicsociological criteria. At higher syntaxon levels a selective use of diagnostic species was made.     

Clusla: A computer program for the clustering of large phytosociological data sets

Summary CLUSLA, a computer program for the clustering of very large phytosociological data sets is described. It is an elaboration of Janssen's (1975) simple procedure. The essence of the program is the creation of clusters, each starting with one relevé, as the relevés are entered in the program. Each new relevé that is sufficiently distinct from already existing clusters is considered a new cluster. The fusion criterion is the attainment of a certain level of (dis-) similarity between relevé and cluster. Bray and Curtis' dissimilarity measure with presence-absence data was used.The program, written in FORTRAN for an IBM 370–158 system, can deal with practically unlimited numbers of relevés, provided the product of the number of primary clusters and the number of species does not exceed 140.000. We adopted maxima of 100 and 1400 respectively.After the primary clustering round a reallocation is performed. Then a simple table is printed with information on the significance of occurrence of species in clusters according to a chi-square approach. The primary clusters can be treated again with a higher fusion threshold; or approached with more elaborate methods, in our case particularly the TABORD program.The program is demonstrated with a collection of 6072 relevés with 889 species of salt marsh vegetation from the Working-Group for Data-Processing.Contribution from the Working Group for Data-Processing in Phytosociology, International Society for Vegetation Science. Nomenclature follows the Trieste system, which will be published later.The authors are very grateful to Drs. Jan Janssen, Mike Dale, László Orlóci and Mike Austin for their comments on drafts of the program, and to Wil Kortekaas for her help in the interpretation of the tables.     

Mediterranean hay meadow communities: diversity and dynamics in mountain areas throughout the Iberian Central Range (Spain)

This paper reports a phytosociological study on managed hay meadow communities in the main mountains of the Spanish Central Range. Multivariate techniques (Cluster analysis and CA-ordination) were applied in the classification of the relevés and ordination of vegetation units and species. Clusters were related to the following associations: Agrostio castellanae–Arrhenatheretum bulbosi, Festuco amplae–Cynosuretum cristati, and F. amplae–Agrostietum castellanae. The scattergram suggested that the dynamic patterns between communities are associated with soil nutrient content and moisture. We propose a new subassociation: F. amplae–A. castellanae gaudinietosum fragilis that includes Salmantincensean Agrostis castellana communities growing in regions with lower supramediterranean thermotype.     

A method for the analysis of temporal changes in vegetation pattern at the landscape level

A new approach to study temporary changes in small landscape units is proposed, based on vegetation-complex relevés (sigma relevés). The method is introduced using extensively grazed or abandoned heathland and Nardus-sward landscapes in the Black Forest as examples. Thirty-six sigma relevés of the years 1979–80 are compared with those of 1989–90. Each of the compared plot areas covers on the average 2–4 hectares, and has a diversity of about 20 different vegetation types. The vegetation complex relevés can be ordered to 3 different types which characterize landscape ecological zones dependant on former rural management.As a result of the comparison 2 main developments which have a spatial differentiation in the Black Forest are confirmed and documentated:

Permanent plot research in wet meadows and cutting experiment

Vegetation mowing is usually recommended for preserving wet meadows, in particular for the prevention of land warping and brushwood encroachment. The concept and the methods for the study of the influence of different cutting rhythms on the vegetation of two wet meadows, as well as some short term results are presented. Cutting experiments with permanent plots have been undertaken in vegetation communities such as Orchio-Schoenetum nigricantis and Caricetum elatae, abandoned for many years. The strategy of the study was to have diverse but complementary methodological approaches in order to assess changes in the vegetation and species abundance, especially concerning the botanical composition, the vegetation structure and the species vitality. The methods include counting of plants or part of plants, morphometry of Phragmites communis, biomass measurements, phenological records, structure relevés with the point-intercept technique as well as classical vegetation relevés with coverage estimation.On a short term base we can conclude, on the whole, that the yearly winter cut treatment induces a better vitality (number of individuals, flowering, biomass) of the studied species, while in the control plots the vitality is low, due notably to the important layer of accumulating litter and the lack of light on the ground, which hamper soil warming. The yearly summer cut treatment and the winter cut made every three years show in general intermediate characteristics. Furthermore, the vegetation structure is noticeably different according to whether the vegetation is cut (with hay removal) or not. Concerning the methodology, it is concluded that a diverse approach is a good way to assess changes in the vegetation with a minimum loss of information. If the rather subjective method of vegetation relevés with cover estimation may be suitable for long term survey on permanent plots, more quantitative and objective techniques have to be set if short term or precise changes in the botanical composition, the species vitality and the biomass, as well as structural changes, are sought.This paper was presented at the meeting of the working Group for Succession on Permanent Plots, held in Bern (Switzerland), 4th–7th September 1991.     

Variation in species composition and species richness within Phragmites australis dominated riparian zones

In riparian wetlands total standing crop often fails to account for a significant part of the observed variation in species richness and species composition within communities. In this study, we used abundance of the dominant species instead of total standing crop as the biotic predictor variable and investigated its relationships with species composition and species richness in communities dominated by Phragmites australis (Cav.) Trin. ex Steudel. This was done by measuring soil organic matter content, litter cover and elevation, Phragmites abundance (standing crop and stem density) and species composition in 78 relevés. In addition, we tried to identify the environmental boundaries of Phragmites communities by sampling relevés in neighbouring communities.Two gradients were related to a decline in Phragmites abundance: one gradient, perpendicular to the shoreline, was mainly related to increased elevation and the second gradient ran parallel to the shoreline and was related to increased amounts of soil organic matter. Within the relevés dominated by Phragmites, stem density of Phragmites and litter cover were the only factors significantly related to species composition in the RDA solution. Litter cover and standing crop of the dominant accounted for 64% of the variation in species richness within the Phragmites-dominated community. These results show that dead and living biomass of the dominant species may account for a substantial part of the variation in species composition and species richness within a single community.     

Variability and ecology of the alpine dwarf shrub community Loiseleurio-Cetrarietum

Summary Variation in the dwarf shrub community Loiseleurio-Cetrarietum from Mt. Patscherkofel, Austria, was investigated using polar ordination, principal component analysis, hierarchical cluster analysis, and multiple correlation analysis. Comparison was made with existing data (relevé tables) from the Alps. In addition to phytosociological methods the canopy structure and the morphological variability of the dominant species Loiseleuria procumbens were studied. The result of the namerical phytosociological analyses yield 3 clusters representing: 1) A dense Loiseleuria carpet, 2) a Loiseleuria carpet damaged by erosion and 3) an open Loiseleuria community on skeletal soils. The first two communities can be combined as one type because of their canopy structure in the form of dense intertwined individual plants and because of the morphological uniformity of Loiseleuria, although in the second group there are species growing on erosion gaps and showing close floristic connections to the third group. The comparison of 140 relevés from the Alps proves that the Loiseleurio-Cetrarietum is characterized only by the dominance of Loiseleuria procumbens while the abundantly occurring lichens change in their species composition along the gradient of wind, and elevation. The open Loiseleuria community at the most extreme habitats (cluster 3) is regarded as a new association. Not only floristic but also physiognomical, morphological and ecophysiological evidence led to this conclusion. The name Gymnomitrio-Loiseleurietum (ass. prov.) is proposed. This plant community shows floristic and ecological correlations to the Diapensia-Loiseleuria-Cesiolichen community known from Scandinavia.Nomenclature of species follows Ehrendorfer (1973) (vascular plants), Poelt (1974) (lichens) and Gams (1973) (mosses).For PCA, hierarchical cluster analysis, and multiple correlation analysis, computer programs from the data bank system (AFO-data) of the Alpine Forschungsstelle Obergurgl, Austria were used. I am indebted to Dr. S. Pamer and P. Moser for their invaluable help. For polar ordination a separate program was developed (language: Simula, author: Dr. G. Knecht). I am also indebted to Dr. H. Hilscher for the translation of the text.     

Influences of transformation of cover values on classification and ordination of lake vegetation

Summary Numerical ordination (Program ORDINA) and classification (Program TABORD) have been applied to vegetation data from a lake in south Sweden.Full percentage values and different transformations of the estimated percented cover scale have been tested on relevés with nymphaeids (+ elodeids) and on relevés containing obly helophytes. The used scales have been: percentage (P), Hult-Sernander-Du Rietz (HSD), Log (%+1) (LP), 1+ log% (L), 100+ % (HP) and Presence/Absence (P/A). The main aim has been to find if there is an optimal transformation regarding both quantitative and qualitative information in vegetation data.In the classification of helophytes a dataset with 87 relevés and 20 species have been used applying P-and L-scales and P/A. The similarity between the different scales has been compared in relation to common relevés between every two corresponding clusters. L and P/A give very similar results, more similar than L and P or PA and P. It is also interesting that the result with P-scale is more similar to L than to P/A. The P-scale neglects with the species with a low cover value, whilst the L-scale takes notice of both quantity and quality.In the classification of 90 nymphaeid relevés containing only 6 species P, HSD, L, HP and P/A were applied. The L-scale (and partly PH-scale) gave a result similar to the most quantitative P-and HSD-scales. P/A-values gave a very deviating result. In the ordination of helophytes only 38 relevés with 18 species, taken along a transcect, were used and P, P/A and 4 different transformations were applied. The similarity between order of relevés in the graphs and real order of relevés along the transect was used as an objective test of the effectiveness of the scale applied. The P-scale gave a bad result, whilst P/A gave the best result. The intermediate scales L and HP gave acceptable results.In the ordination of nymphaeids (+ elodeids) 90 relevés, where only pattern was studied, the L-scale gave the best result most similar to P-and HSD, but applying the two extremes, P-scale and P/A, resulted in large reduction of information.In the submitted ordinations and classifications the applied L-scale (and HP-scale) always gave ecologically interpretable results, but the extreme transformation were less reliable.This work has been supported by grants from the University of Lund, for which I am duty grateful.Nomenclature follows Lid (1974).I am grateful to Professor Nils Malmer, head of the Department of Plant Ecology, Lund, Sweden, Dr. Eddy van der Maarel, University of Nijmegen, Holland, and my colleague Fil. Kand. Stefan Persson for valuable advice and discussions. I also thank my wife Fil. Kand. Eva Waldemarsson Jensén, Institute of Ecological Botany, Uppsala, for valuable discussions, Mr. Tommy Olsson for valuable technical advice, Dr. R.S. Clymo for advice with DIVINF and PCA and correction of the language, Mrs. Mimmi Varga for drawing the figures and Mrs. Brita Billstein for typewriting the text and the tables.     

A phytosociological survey of the Irish Molinio-Arrhenatheretea using computer techniques

Summary The Irish approach to the computer processing of grassland relevés is introduced. A suite of programs in Fortran IV has been developed for the purpose. The preparation and analysis of the relevés using these programs is explained. The cost and ecological effectiveness is discussed. It is concluded that the approach is not radically different to the familiar Braun-Blanquet technique of tabular comparison of relevés.Contribution from the Working Group for Data-Processing in Phytosociology, International Society for Vegetation Science.     

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 snow-bed vegetation in the Northern Apennines

The snow-bed vegetation in the Northern Apennines was studied by the Braun-Blanquet method. The phytosociological relevés were classified by numerical methods. Four main vegetation types were identified: Salicetum herbaceae, community of Carex foetida, Poo-Cerastietum cerastioidis and Oligotricho-Gnaphalietum supini. The latter was described as a new association. The vegetation types were ecologically characterized by an indirect gradient analysis based on principal component analysis. The results were interpreted by means of: i) geomorphological observations, ii) soil profiles, iii) measurements of the length of the snow-free period.     

Computerized matching of relevés and association tables,with an application to the British National Vegetation Classification

When a new relevé is to be assigned to a pre-existing type, its composition is compared with an association table. Bayesian inference may seem a good way to make the comparison, but presents difficulties. In an alternative approach, three indices of goodness-of-fit are proposed. Compositional satisfaction is a measure of how well the species composition of the relevé fits the constancy classes in the table; it is a minor modification of the Czekanowski coefficient of similarity between observed and expected numbers of species in each constancy class. Dominance satisfaction is a modification of the Czekanowski similarity between the relevé and cover values that might be expected from the association table. Dominance constancy is a weighted mean of the constancy class of the four most abundant species in the relevé. A computer program, TABLEFIT, combines them into a single index. It has been tested on British mire vegetation.     

Syntaxonomical analysis of beech woods in the Apennines (Italy) using the program package IAHOPA

IAHOPA, an overlay program package performing intersection analysis and information analysis has been applied to a large data set of relevés of beech woods in the Apennines (Italy) completed by several authors following the Braun Blanquet approach. The results have been treated by several numerical methods testing classification efficiency and predictivity. Ecological indicator values have been used to test for predictivity. The classification proposed by Gentile has been confirmed in its main lines. However 2 new associations (Polysticho-Fagetum and Digitali-Fagetum) and 12 new subassociations are described. Furthermore the Veronico-Fagetum Montacchini 1972 has been recognized also for the Apennines. The clusters corresponding to the association level could be classified in two main alliances: Geranio nodosi-Fagion and Geranio striati-Fagion as suggested by Gentile, however their syntaxonomical justification should be based on numerical comparisons of the data from the entire area of European beech woods.