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.     

A phytosociological study of the macrophytic vegetation of running waters in western lower saxony (Federal Republic of Germany)

A survey was carried out of the macrophytic vegetation in running waters of Western Lower Saxony. Three hundred and eighty-two phytosociological relevées were classified by common table work to 30 vegetation types of different phytosociological validity. The vegetation types are described and floristically characterized. Most of them belong to the complex Sparganium emersum community, which is characterized by the dominant occurrence of the nymphaeid species Sparganium emersum Rehm., Nuphar lutea (L.) Sm., Sagittaria sagittifolia L. and Potamogeton natans L., but magnopotamid-, parvopotamid-, batrachiid- and pleustophyte-dominated communities also occur. Similarity between the different vegetation types was checked by multivariate techniques, one classification technique (minimum variance clustering) and one ordination technique (principal components analysis). Additionally, syndynamical relations between the types were studied by observing the vegetation changes of 46 sample points within 2 years. The relations between different vegetation types are shown by combination of the different approaches. Finally, some more general statements are made regarding the handling of such data sets, as well as the consequences of the results for the classification of macrophytic vegetation in Central European rivers.     

The development of numerical classification and ordination

Summary The paper reviews the constraints and influences which have affected the development of numerical classification and ordination of vegetation.Initial development of ordination techniques and their reception by ecologists was hindered by the mistaken idea that ordination involved acceptance of variation in vegetation as a continuum, as well as by a general suspicion of mathematical approaches.Three distinct approaches to ordination, largely unrecognised at the time, are apparent in earlier work: direct gradient analysis, reduction in dimensionality and path-seeking (catenation) (Dale 1975).Modifications of simple initial techniques made them more efficient at the cost of increased computation. Acceptance of heavier computation as computers increased in capacity and speed turned attention to prineipal component analysis and the superficially similar factor analysis. These have been widely misunderstood largely because they were initially applied in the same way as in the analysis of psychological data, in which different constraints and objectives apply. The initial failure to recognise that principal component analysis involves a preliminary data transformation, the form of which depends on answers to biological, not mathematical, questions, was particularly unfortunate.Principal component analysis has limitations as a technique of ordination resulting from its assumptions of linearity and additivity of plant responses. Attempts to devise more effective techniques raise questions about the practical importance of non-linearity if the objective is data-exploration rather than elucidating the nature of species-response curves and about the adequacy of using simulated data as test data when we do not know how to simulate realistic data.Data-exploration has been more prominent in practical uses of ordination but many methodological developments have concentrated rather on species-response curves.Numerical classification also met obstacles to its acceptance additional to a general aversion to numerical techniques. The first numerical techniques were presented in the context of the relationships of a particular set of data, rather than of a generally valid system, which was the more familiar concept in non-numerical classification.Both numerical and non-numerical classification aim to produce as homogeneous groups as possible. The distinctive contribution of numerical methods is to allow the data to indicate the most efficient criteria of classification; this was an unfamiliar idea.The strategy of classification may be either divisive or agglomerative and either monothetic or polythetic. Choice of strategy in earlier work was not only constrained by computational limitation but may also have been influenced by an author's previous experience of non-numerical classification. As with ordination, the distinction between preliminary data transformation and subsequent analysis was at first not appreciated.Numerical classification has been influenced by parallel numerical developments in formal taxonomy. Because objectives and assumptions are not always the same, this influence has not been altogether helpful.The limitations of real data suggest that developments of technique are at risk of becoming too concerned with refinements of methodology. Increasingly complex methods and increasing availability of programmes for such methods carry the risk that they may be used without adequate understanding of what they do.     

Environmental conditions in relation to variability of deciduous forest vegetation — a study from volcanic mountain in central Slovakia

The deciduous forests represent dominant natural vegetation of Central European landscape and an important functional component for maintenance of biological diversity. However, their syntaxonomy and ecological gradients still remain unclear. The numerical classification was conducted to determine the main units of forest vegetation, while ordination techniques were used to explain the structure of vegetation-environmental data matrix consisting of 110 forest stands in the Štiavnické vrchy Mts (central Slovakia). Ten vegetation types within the phytosociological classes of deciduous forests Quercetea robori-petraeae and Querco-Fagetea were distinguished. The major environmental driver responsible for variation in forest species composition was interpreted as a response to soil moisture which also accounted for a large part of species variability (3.74%). Soil nutrient/acidity complex expressed by pH, Ca and Al concentration was also an important source of vegetation variability. Relevance of soil conditions in relation to plant survival and community distribution was discussed. Along the soil moisture gradient, vegetation types were arranged from the subxerophilous oak forests through the mesophilous beech and ravine forests to the hygrophilous alder ash vegetation.     

我国植被数量分析方法的研究概况和发展趋势

植被数量分析是现代植被研究的重要手段,数量分类和排序是现代植被生态学研究最重要的,也是应用最广泛的生态学技术。数量分析方法在20世纪50年代引入植被生态学研究领域,我国学者在70年代后期开始研究植被的数量分类和排序。本文主要从相关资料、应用研究、新方法研究三个方面论述了我国植被数量分析方法的发展,重点阐述了从20世纪70年代后期以来出现的并且被广泛应用的新方法及其应用研究概况,并在此基础上分析了植被数量分析方法未来的发展趋势。     

Data-processing in phytosociology,retrospect and anticipation

Summary The existence of the Working Group for Data-Processing in Phytosociology spans about six years, 1969–1975. Its activities are summarized and a series of papers to appear in Vegetatio, presenting results of various numerical treatments, mainly of salt marsh vegetation, is introduced. A survey is given of seven releve selections to which treatments were applied. Five main fields of activities are discussed: systematical data-processing, rearrangement of phytosociological tables, development of classification and ordination techniques, numerical syntaxonomy, and theoretical phytosociology.The establishment of the Working Group for Theoretical Phytosociology and its first symposium on theoretical aspects is announced.Contribution from the Working Group for Data-Processing in Phytosociology, International Society for Vegetation Science.     

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.     

Classification of vegetation: Past,present and future

Abstract. This paper is a report on the past, status-quo and perspectives of vegetation classification, still a major occupation of many vegetation scientists. The history of vegetation classification is discussed against a background of several controversial issues such as the problem of continuum vs. discontinuum, naturalness vs. arbitrariness of the nature of plant communities, universality vs. ad hoc character of syntaxonomic schemes, as well as classical versus numerical approaches to data analysis for classification purposes. The development of the methodology of vegetation science and the present image of vegetation classification is documented by a bibliometric analysis of the publication record of four majorjournals: Journal of Vegetation Science, Vegetatio, Phytocoenologia and Tuexenia. This analysis revealed a persisting controversy between traditional and numerical approaches to vegetation classification. A series of important changes in vegetation science (foundation of new journals, change of editorial policy by the established, important meetings) punctuate a period called the ‘Innovation period’. Several trends in the development of methods of vegetation systematics are summarized under the headings formalism, pluralism, functionalism, pragmatism and indeterminism. Some new features, such as the development and improvement of numerical tools, use of large data banks and attempts to summarize the theory of vegetation classification are discussed. The new growth-form system of Barkman initiated a revival of physiognomy-based vegetation classification. Within this framework the use of the character-type concept and the development of new numerical methods for studying the hierarchical structure of character-set types seems to be a promising approach. The achievements of population biology and ecophysiology have affected vegetation science by emphasizing the functionality of species within plant communities. The use of guilds and other functional groups has experienced an increasing interest from vegetation scientists. Applied in vegetation science, fuzzy-set theory has bridged the techniques of classification and ordination of plant communities.     

Effect of streaked shearwaterCalonectris leucomelas on species composition ofPersea thunbergii forest on Kanmurijima Island,Kyoto Prefecture,Japan

The effects of trampling and burrowing by streaked shearwaters were studied on thePersea thunbergii forest in Kanmurijima Is. The vegetation data were analysed using both the phytosociological tabulation method and principal component analysis (PCA). The vegetation ordination on the first axis reflected the environmental gradient from light to heavy disturbance by the streaked shearwater, and the vegetation ordination on the second axis represented the gradient from the latter phase of forest succession to retrogressive succession. The position of the three vegetation groups in the ordination diagram successfully explained the relation between the habitat and the species composition. The disturbance caused by activities of the streaked shearwater such as burrowing and trampling causes a decrease of species, especially character species of Camellietea japonicae, and also causes retrogressive succession: patches ofMallotus japonicus develop and many heliophytes occur.     

Numerical studies of compositional gradients in oreophilous vegetation (northern Apennines,Italy)

Summary A numerical study of a set of phytosociological data from oreophilous vegetation in the Apennines (Italy) has been performed. Ranking, classification and ordination methods have been used. The advantages of the joint use of the different methods is diseussed. The results confirm a compositional gradient related to degradation due to grazing, and another, related to base-richness in the soil. Compositional variation does not appear strongly disjointed.Nomenclature follows Zangheri (1976)This work was supported by a grant from the Italian CNR, Pfoject Environment Quality. We wish to thank Professor L. Orlóci for discussions and facllities, and Drs. D. Ubaldi and C. Ferrari for participation in the fieldwork.     

云南西畴县草果山常绿阔叶林的数量分类研究

一、引言 在植物生态学中,植物群落数量分类的研究始于50年代。Goodall(1953)首次把数量分类方法引入植物生态学的研究中。以后,Williams和Lambert(1959、1960、1961)作了大量的工作。Greig-Smith于1957年出版了专著《定量植物生态学》(1964年增订再版),系统     

Soft-water vegetation (Littorellion) in Spanish mountains

This paper presents a systematic review of Spanish high mountain soft water communities based on 46 phytosociological relevés. Using a numerical analysis with ordination (correspondence analysis—CA) and classification (Jaccard index and complete-linkage clustering) three groups corresponding to the phytosociological associationsIsoeto lacustris-Sparganietum borderei, Sparganio angustifolii-Isoetetum echinospori andSparganio angustifolii-Callitrichetum font-queri have been defined. SpanishLittorellion communities are characterized by boreal-alpine and endemic elements. The boreal-alpine elements probably spread via the Pyrenees-Iberian System through to the mountains of the western peninsula during a cold period, thus sharing the habitat with the Ibero-Atlantic flora which was able to adapt to the changing climatic conditions.     

Numerical analysis of vegetation complexes and community diversity of major coastal dinaric mountains

Abstract. This study includes 24 transects on slopes of the Velebit, Biokovo and Orjen mountains, the tallest coastal mountains along the Adriatic Sea, with a very high species richness according to European standards and having the largest regional concentrations of endemics. Within each transect the presence or absence of phytosociological alliances, according to the Zürich-Montpellier School, was interpreted from local studies. The alliance composition of the transects was compared with methods used in numerical syntaxonomy. Three types of vegetation complexes (N. Dinaric inland, transitional and S. Dinaric coastal) were derived from a numerical classification of the transects; they were defined on the basis of their characteristic alliances. The maritime impact affecting the mountains through the river canyons reduces the contrast in vegetation between inland and coastal slopes. The existence of a climatic macrogradient is indicated by seriation of the basic alliance/transect data matrix leading to a diagonal structure, by a gradual shift of the alliance composition within the same vegetational type, and by the distribution of the transects and alliances along the first axis of a Correspondence Analysis ordination. Transect scores on the second ordination axis are negatively correlated with transect vegetational diversity (number of alliances per transect); this axis may reflect a decrease in humidity and temperature range. This interpretation is supported by the result of a comparison of the total mountain vegetational diversity (Mann-Whitney test), which showed that Orjen is the most diverse mountain ridge. The expected correlation between plant community diversity and altitude of the transects is blurred, probably due to the impact of the cold Bora wind and subsequent compression of vegetational zones.     

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.     

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.     

Classification of basiphilous pine forests in Telemark, SE Norway: a numerical approach

The basiphilous pine forests in southern Telemark, SE Norway, have previously been classified according to the ordinary phytosociological methods of the Braun—Blanquet system. The same material was later treated by numerical analysis, i.e., TABORD—classification and the ordination technique 'reciprocal averaging'. No significant differences were found between the numerical and the non—numerical classification except for the classification of some transitional communities. Ordination by 'reciprocal averaging' gave a useful complement to the TABORD classification, and re—leves and species were distributed along interpretable gradients (i.e., along a dry—wet gradient and along a gradient from forest—rim conditions to forest conditions).     

应用间接排序识别盐沼植被的光谱特征:以崇明东滩为例

遥感技术已成为大尺度植被分类的重要手段,而地面植物群落特征与其光谱特征之间的关系是解译遥感影像的关键。该研究选择上海崇明东滩自然保护区的盐沼植物群落为对象,应用ASD地物光谱仪测定其植物群落的光谱反射率,并采用10个小型机载成像光谱仪(CASI)默认植被波段组,应用主分量分析法和相关分析分析了不同群落光谱特征与生态环境因子之间的关系。分析结果表明,间接排序法PCA能够识别盐沼植被中光滩、海三棱 草(Scirpus mariqueter)群落、芦苇(Phragmites australis)群落和互花米草(Spartina alterniflora)等群落的光谱特征,绝大多数盐沼湿地植物群落组成与光谱特征之间有显著的相关,识别效果最好的波段组是736~744 nm、746~753 nm、775~784 nm、815~824 nm和860~870 nm;对光谱反射率影响最大的生态环境因子分别是植物群落的高度和盖度,高程和其它环境因子的影响次之。研究成果可为遥感监测崇明东滩自然保护区内入侵种互花米草的空间分布和扩散规律提供技术支撑,为高光谱遥感影像的影像判读和解译分类以及盐沼湿地植被制图提供科学依据。     

Vegetation—environmental relationships in the bed of Wadi El-Sheikh of Southern Sinai

This study deterines the magnitudes of variation in the phytosociological structure in the bed of Wadi El-Sheikh of Southern Sinai. It evaluates the relationships between its vegetation and environment.Vegetation and soil were sampled in 68 stands representing physiographic variations in the study area. Relative frequency, density and coverage were determined for each perennial and were added to provide an estimate of its importance value. The stands were classified by the mutual information method. The principal components analysis was applied to achieve a two-dimentional ordination of stands.Determinations of the percentages of surface sediments of different size classes, soil pH, conductivity, moisture content, water-holding capacity, CaCO₃ and organic matter were carried out.The classification and ordination resulted in a clear demonstration of the vegetation pattern in the study area in quantitative terms. Other phytosociological groups were characterized in addition to those identified in previous studies. Vegetation analysis indicated that it is dominated by Artemisia judaica, Zilla spinosa, Fagonia tristis var. boveana and Lygos raetam. Factors influencing the distribution of vegetation fall into two important groups, one includes the moisture factors and the other includes factors of soil fertility.     

Multivariate analysis of vegetational variation in different habitats at Omayed,Egypt

The application of indicator species analysis has proved useful in classifying stands into groupings coinciding with topographic variations. 41 indicator pseudo-species are identified in the vegetation along a phytosociological gradient: some are characteristic of specific habitats, and the others indicate transition between groups of habitats.The use of qualitative estimations (transformed density records) in multivariate analysis in the present study indicates that they may be more preferable than quantitative estimations, because of their easy and fast recording in the field, and the less comprehensive computations, while yielding precise results.The X-axis of the reciprocal averaging ordination is related to the salinity and fertility gradients, while the Y-axis reflects soil texture.The phytosociological gradients of the canonical variate axes reflect essentially contrasts between groups of species, each correlated with one or the other of the environmental gradients.Comparing the results obtained by the two ordination methods (reciprocal averaging and canonical variates) we find that the first method detects the overall phytosociological changes along strong environmental gradients which would be helpful in studying large surveys. On the other hand, the second technique, which is mainly predictive, is more sensitive to changes within both the phytosociological and environmental gradients and can detect the impact of these changes on the overall variance of each gradient.Nomenclature follows Täckholm (1964).The authors are grateful to Dr J. Jeffers and Mr and Mrs P. Howard of the Institute of Terrestrial Ecology, England, for their help with applying the multivariate techniques to the vegetation data.     

基于SOM的野生秤锤树群落的空间分布和环境解释

本文采用自组织特征映射网络(self-organizing map, SOM)对南京老山野生秤锤树(Sinojackia xylocarpa)群落进行数量分类和排序, 分析了其与环境因子之间的关系。结果表明: (1) SOM将秤锤树野生群落的100个样方划分为5个群丛类型, 分类结果在空间上反映了秤锤树野生群落的演替变化趋势, 各群丛的群落结构和物种组成存在差异且群丛界限明显, 可较好地进行排序与分类的环境解释。(2)通过环境因子梯度的可视化方法, 确定了海拔、坡位和土壤厚度是影响该地区秤锤树生长和分布的主要因子, 同时也揭示了以不同优势种为代表的各群丛和环境因子的关系。(3) SOM可以摆脱许多定量技术的限制性假设, 使得神经网络对于群落生态特征及探索群落和环境相互关系具有良好展现力; SOM群落生态数据具有更高的映射能力, 进行群落分类以及较少程度的排序的潜力, 将有利于不同群落类型的分类和管理, 对于濒危植物保护具有重要意义。