The robustness of Correspondence,Detrended Correspondence,and TWINSPAN Analysis

Abstract. Simulated vegetation data of known structure and varying complexity were analyzed with Correspondence Analysis (CA), Detrended Correspondence Analysis (DCA), and a polythetic divisive method of classification as incorporated in the computer program TWINSPAN. The CA and DCA axes were statistically compared with the original gradients. In the case of TWINSPAN, the results were visually evaluated. The analyses showed that both CA and DCA will recover the first vegetation gradient in an acceptable manner only if the variation in the vegetation due to second or higher order gradients is small. Second gradients were never recovered in a meaningful manner. The sample points along these axes were so far displaced from their original positions along the gradients that the axes could not be used to represent the gradients. In particular, when the two gradients have the same length, CA and DCA gave very poor results for both the first and the second axes. This is thought to be one of the main reasons why TWINSPAN does not perform well, especially after the first division.     

瑞典河漫滩草甸植被的数量分类和排序

本文用目前各国学者广泛使用的一些数量分类和排序方法对瑞典河漫滩草甸样地资料进行了分类和排序。所用的方法包括多元等级聚合分类(TABORD程序),多元等级分划分类(TWINSPAN程序),PCA排序(ORDINA程序),RA和DCA排序(DECORANA程序)。研究结果表明可以把28个样地分为6个群落类型,它们的分布格局是与土壤水分梯度密切相关的。此外本文还对数量分类和排序方法在植物群落学研究中的应用以及所用方法的比较进行了讨论。     

Trends in structural compositional attributes of dune-interdune vegetation and their edaphic relations in the Indian desert

Vegetation of 127 sites on different aspects of dune-interdunes in the Indian Thar Desert was classified using TWINSPAN. TWINSPAN groupings of sites separated better vegetated dunes of the northeast form the poorly vegetated dunes of the northwest and the southwest. Of the different ordinations using noncentred, centred and centred and standardized principal component analysis, reciprocal averaging and detrended correspondence analysis (DCA), the site and species classes in DCA correlated well with ten edaphic and ten vegetational attributes of each site. Strong correlation of vegetation groupings with soil texture, moisture holding capacity and low correlation with pH and electrical conductivity revealed the possible importance of soil physical properties in affecting vegetation composition.The 11 species classes in TWINSPAN were regrouped into 18 species classes in DCA, which separated highly frequent species from those of less and least frequent species. Based on dominance-diversity attributes, Calligonum polygonoides-Lasiurus sindicus was brought out as bioedaphic climax stage. Correlation of ordination scores in different site groupings with vegetational attributes showed specific trends: From the zero of x, y and z axes to their maximum, the ordination scores of grasses and browse species declined while score of spinous species increased. The sites near the origin of the x, y and z axes were therefore least degraded and those at or near the maximum of x, y & z axes were most disturbed as was confirmed by the dominance diversity trends. Thus trends of compositional and functional attributes of vegetation of sites in different groupings helped in inferring a site's degradation status.Abbreviations RIV Relative importance value - TWINSPAN Two Way Indicator Species Analysis - PCA Principal Component Analysis - RA Reciprocal Averaging - DCA Detrended Correspondence Analysis - EC Electrical Conductivity - WHC Water Holding Capacity - BD Beta Diversity - DC Dominance Concentration - WWS Windward slope of the dune - LWS Leeward slope of the dune - ID Interdune Nomenclature: Bhandari, (1990)     

Effects of sample distribution along gradients on eigenvector ordination

In general, disproportionately heavy sampling of the ends of a gradient increases the interpretability of eigenvector ordinations. More specifically, correspondence analysis (CA) and detrended correspondence analysis (DCA) best reproduce the original positions of samples in simulated coenoclines when samples are clustered toward the ends of the axis. Principal components analysis (PCA) reproduces the original sample positions less well than either CA or DCA and shows no improvement as samples are increasingly clustered toward the ends of the axis. PCA and CA show less curvature of one dimensional data into the second axis when sampling favors the ends of the axis.     

Bootstrapped ordination: a method for estimating sampling effects in indirect gradient analysis

Indirect gradient analysis, or ordination, is primarily a method of exploratory data analysis. However, to support biological interpretations of resulting axes as vegetation gradients, or later confirmatory analyses and statistical tests, these axes need to be stable or at least robust into minor sampling effects. We develop a computer-intensive bootstrap (resampling) approach to estimate sampling effects on solutions from nonlinear ordination.We apply this approach to simulated data and to three forest data sets from North Carolina, USA and examine the resulting patterns of local and global instability in detrended correspondence analysis (DCA) solutions. We propose a bootstrap coefficient, scaled rank variance (SRV), to estimate remaining instability in species ranks after rotating axes to a common global orientation. In analysis of simulated data, bootstrap SRV was generally consistent with an equivalent estimate from repeated sampling. In an example using field data SRV, bootstrapped DCA showed good recovery of the order of common species along the first two axes, but poor recovery of later axes. We also suggest some criteria to use with the SRV to decide how many axes to retain and attempt to interpret.Abbreviations DCA= detrended correspondence analysis - SRV= scaled rank variance     

长白山高山冻原植物群落的数量分类和排序

本文用目前在国际上比较先进的双向指示种分析(TWINSPAN)多元等级分划分类方法和无趋势对应分析(DCA)排序方法对长自山高山冻原植物群落进行了研究。研究结果表明:DCA的第2轴与长白山高山冻原59个植物群落类型的土壤水分梯度紧密相关, TWINSPAN将长白山高山冻原59个植物群落分为12组。此外, 本文还对长白山高山冻原植物群落的TWINSPAN和DCA分类结果与经典分类方法对其分类的结果进行了比较。     

Plant-environment relationships on the Montlake wildlife area,Seattle, Washington,USA

Grassland vegetation on the Montlake fill was analyzed using TWINSPAN. Eight herb communities were recognized. Moisture, proximity to gas vents, and disturbance are the main factors that control species and community distributions. Binary discriminant analysis (BDA) and detrended correspondence analysis (DCA) were used to study species-environment relationships. BDA revealed complex species response patterns and the resultant indicator values were used to interpret the ordination axes. Species distributions are controlled primarily by moisture, but also influenced by soil pH. Multiple regressions revealed little about plant-environment relationships not discovered by BDA. Before robust nonlinear methods are available, BDA, metric ordination with data stratification and nonmetric ordination are methods that can yield satisfactory results in exploratory plant-environment studies. BDA alone is an efficient, useful first approach where response patterns of species are initially unknown.Abbreviations BDA Binary Discriminant Analysis - DCA Detrended Correspondence Analysis     

子午岭地区植被恢复演替过程与环境因子的分类与排序

在野外实地调查基础上,采用双向指示种分析法(TWINSPAN)和除趋势对应分析(DCA)对子午岭地区植物群落进行了数量分类和排序。结果显示:(1)TWINSPAN将50个样方划分为14个群系;DCA排序结果与TWINSPAN分类结果相互印证。(2)第一排序轴上显示出植被由草地、灌丛到森林的恢复演替系列,在此演替过程中土壤中的速效磷逐渐减少,全氮和有机质逐渐增加;第二排序轴反映环境因子的梯度不明显,但植被的演替过程会受到海拔、坡度的影响。研究表明,随着植被恢复演替的进展群落结构在变化,乔木层的重要值增加,所在群落中的全氮、速效磷、有机质等亦呈现规律性变化。     

Urban plant ecology patterns and processes: a case study of the flora of the City of Plymouth, Devon, U. K.

Aim Using a large database that has been created over the past 5 years with the RECORDER package, presence/absence data for 829 species of vascular plants in the 103 1-km² squares that cover the city of Plymouth (pop. 243,373) have been analysed by two-way indicator species analysis (TWINSPAN ) and canonical correspondence analysis (CCA) to establish the major species assemblages and to examine their spatial distribution across the city in relation to variation in land use. Location The City of Plymouth. Methods Nine groupings of squares emerged and their distribution was mapped across the city. Interpretation of those groups and their variation in relation to the land use ordination axes showed that TWINSPAN groups lying along the first axis of variation correlated floristic variation with the process of urban development and the historical evolution of urban structure. The second axis appeared to be related to particular remnant semi-natural habitats within the city that could be regarded as ‘hot-spots’ for survival of many plant species. Species were categorized into four types on the basis of their recency and mode of arrival in the city and by using the historical flora of Plymouth produced by T. R. Archer-Briggs in 1880. Variations in the sources of species in relation to the TWINSPAN classification groups were then examined. A second ordination was also computed; this time with geological, altitudinal and distance variables. TWINSPAN groups were then superimposed on the ordinations using polygons to assist with biogeographical interpretation. Results and conclusions The results of both the multivariate analyses and distribution of species by source are then discussed in the context of previous research into urban plant ecology, particularly in Central Europe. The problems of inferring process from pattern in this meso-scale study are reviewed and suggestions are made for further research into urban ecology and biogeography in the city.     

对应分析和去趋势分析在有蹄类取食模式研究中的应用

本研究将2009年1月和2010年1月小兴安岭大沾河湿地自然保护区二可河林场内驼鹿冬季食性作为原始数据,分别以对应分析（CA）、去趋势分析（DCA）,并将数据以样本为单位进行标准化后,再进行去趋势分析（DCA_std）3种排序方法,对驼鹿冬季取食模式进行了研究,后通过普鲁克分析,比较了不同排序方法对大型有蹄类取食模式研究的效果。结果表明,3种排序法的1轴和2轴均能涵盖绝大多数信息量,CA涵盖79.27%,DCA涵盖66.65%,DCA_std涵盖68.22%;3种方法均能够在1轴上区分针叶树和落叶乔木类食物,在2轴上,3种方法主要能够达到针叶树种与除落叶乔木外的其他植物类别的区分。虽三者均能够展现有蹄类取食模式,但在图形可视化后,仅DCA_std无明显的弓形效应。普鲁克分析结果表明,DCA_std样本位移平方和与CA和DCA均有很大差异,即将数据先进行标准化再进行DCA分析能够有效去除弓形效应。因此,在由多度组成的食性数据在进行标准DCA分析前,应对数据进行前期处理会得到更好的效果。同时,以样本为单位的标准化将使排序分析结果生态学意义更明确。     

Species and plant community distribution in a Mojave-Great Basin desert transition

Multivariate analyses were used to describe the vegetation characteristics of a transition from lowelevation Mojave desert to higher-elevation Great Basin desert. Vegetation data used were from Plutonium Valley in the Nevada Test Site. Data from forty nine releves were analyzed with two classifications (two-day indicator analysis or TWINSPAN and unweighted paried group cluster analysis or CLUSTER). Three ordinations, reciprocal averaging (RA), detrended reciprocal averaging (DCA) and non-metric multidimensional scaling (MNDS), were also used. A rotational correlation analysis was used to determine the vector direction of environmental gradients that correlate best with ordination results. Only token correspondence was found between multivariate classes generated by TWINSPAN and CLUSTER, and seven classes (plant communities) identified from field reconnaissance. The latter seven communities were based on differences in dominant species. Distribution of the vegetation was related more to beta diversity than alpha diversity. Individual species were much less diagnostic than the amount of plant cover, groups or guilds of species or differences in elevation and steepness of slope. Because of the high beta diversity the NMDS ordination gave results with the greatest ease of interpretation.     

Geographical and ecological differentiation in Greek Fagus forest vegetation

Question: Which are the gradients of floristic differentiation in Greek beech (Fagus sylvatica) forests? Which is the role of geographical and ecological factors in this differentiation? Location: Beech forests of the plant geographical regions Northeast, North Central and East Central Greece. Methods: A total of 1404 published and unpublished phytoso‐ciological relevés were used in the analyses. TWINSPAN and DCA were applied to classify and ordinate the relevés. Altitude, Indicator Values of relevés and their X and Y coordinates were used in a posteriori interpretation of the ordination axes. Kendall's correlation coefficients were calculated between DCA relevé scores and explanatory variables. Multiple linear regression was used to partition the variation explained by the first two DCA axes, between the geographical and the ecological variables. Results: Classification resulted in 14 vegetation units defined by species composition. Two types of gradients, ecological and geographical, were revealed by the DCA of all releves. The partition of the variation accounted for by the first and second DCA axis was attributed mainly to ecological and geographical variables, respectively. Conclusions: Beech forests of northeast and Central Greece show phytogeographical differences, while ecologically similar vegetation units occur in both regions. A west‐east gradient is revealed in Greek beech forest vegetation. The extent of the study area, its position along regional gradients and the comprehensiveness of the data set that is analysed determine the types of the gradients which can be revealed in a vegetation study.     

The impact on the Senjogahara ecosystem of extreme run-off events from the river Sakasagawa,Nikko National Park

Vegetational data from Senjogahara moor, Nikko National Park, Japan have been analysed using two contrasting but parallel multivariate approaches as a basis for future studies. The plant communities objectively defined using a divisive polythetic classificatory approach (TWINSPAN) have been successfully overlayed on an ordination of the same data using detrended correspondence analysis (DCA). The results reveal a vegetational mosaic reflecting differential levels of flood damage to the vegetation and its subsequent rates of recovery, in a space-time continuum. The results from the TWINSPAN classification show that 13 plant communities are present in the sample area and these correlate closely with complete absence, slight, or extremely heavy disturbance. The form of the disturbance as either channel erosion or alluvial depositional features is shown to also correspond quite closely with the vegetational types and is reflected in the ordination axes derived from DCA.     

芦芽山亚高山草甸的数量分类与排序研究

芦芽山亚高山草甸是在华北地区最典型的山地草甸之一,该草甸在水土保持、生态旅游资源开发和生物多样性保护等方面发挥着重要作用。应用双向指示种分析法（TWINSPAN）和除趋势对应分析（DCA）相结合的方法,对山西芦芽山亚高山草甸进行数越分类和排序,将该草甸划分为13个群丛类型,分别位于不同的海拔高度。分类结果很好地反映了植物群丛类型与环境梯度之间的关系,并在DCA二维排序图上得到较好的验证。DCA排序轴反映了海拔、土壤湿度和坡向的梯度变化,表明海拔高度、水热条件和地形是影响植物群丛变化的主要生态因子。     

Quantitative classification and ordination of forest communities in Pangquangou National Nature Reserve

Quantitative analysis of ecological relationships between vegetation and the environment has become an essential means in the field of research of modern vegetation ecology. In this article, based on data from 84 quadrates, forest communities in this reserve were investigated using TWINSPAN, DCA and DCCA. The results will be helpful in the construction and development of Pangquangou National Nature Reserve. Using TWINSPAN, the forest communities were classified into seven types. The distribution pattern of vegetation reflects the comprehensive influence of environments. The results of DCA and DCCA clearly reflect the relationship between the pattern of forest communities and environmental gradients. The ordination result of DCCA indicates that altitude is more important than other environmental factors because the change of altitude gradient will lead to changes in the temperature and humidity gradients. The first of the DCA ordination axes indicates the humidity gradient, and the second indicates the temperature gradient. All these results show that the main factors restricting the distribution of communities in this reserve are temperature and humidity. The ecological meaning of the ordination axis in DCCA is much clearer than that in DCA, and the species-environment correlation of DCCA is more obvious than DCA. The first DCCA axis indicates the altitude gradient among the communities, while the second is the gradient in aspect and slope among the communities. DCCA ordination can simultaneously express similarities of species and environment. Therefore, the quadrat location in the DCCA ordination figure is much closer than in the DCA.     

Quantitative classification and ordination of forest communities in Pangquangou National Nature Reserve

Quantitative analysis of ecological relationships between vegetation and the environment has become an essential means in the field of research of modern vegetation ecology. In this article, based on data from 84 quadrates, forest communities in this reserve were investigated using TWINSPAN, DCA and DCCA. The results will be helpful in the construction and development of Pangquangou National Nature Reserve. Using TWINSPAN, the forest communities were classified into seven types. The distribution pattern of vegetation reflects the comprehensive influence of environments. The results of DCA and DCCA clearly reflect the relationship between the pattern of forest communities and environmental gradients. The ordination result of DCCA indicates that altitude is more important than other environmental factors because the change of altitude gradient will lead to changes in the temperature and humidity gradients. The first of the DCA ordination axes indicates the humidity gradient, and the second indicates the temperature gradient. All these results show that the main factors restricting the distribution of communities in this reserve are temperature and humidity. The ecological meaning of the ordination axis in DCCA is much clearer than that in DCA, and the species-environment correlation of DCCA is more obvious than DCA. The first DCCA axis indicates the altitude gradient among the communities, while the second is the gradient in aspect and slope among the communities. DCCA ordination can simultaneously express similarities of species and environment. Therefore, the quadrat location in the DCCA ordination figure is much closer than in the DCA.     

Data manipulation and gradient length estimation in DCA ordination

Abstract. The effects of different kinds of data manipulation on gradient length estimation by non-linear rescal-ing (as in DCA ordination) are evaluated by considering the first axis inDCA ordinations of 11 field data sets from four investigations. Gradient length estimates are dependent on the range of the abundance scale; the more the scale favours the quantitative aspect (abundance) of the data over the qualitative aspect (presence), the longer the DCA axes. The gradient length estimate decreases when infrequent species are deleted. A new formula is proposed to replace the option for downweighting of rare species in DCA, as the option presently available has some undesirable properties. Some implications for interpretation of gradient length estimates by non-linear rescaling in general (and in DCA in particular) and for comparison of gradient length estimates between studies, are discussed. The potential of non-linear rescaling of gradients for estimation of β diversity is emphasized.     

On the relationship between sample plot size and beta diversity in boreal coniferous forests

Nested sample plots of three sizes (16, 1, and 1/16 sq. m) from three different studies of Norwegian coniferous forests have been subjected to DCA ordination using the same choice of options. At each sample plot size, species quantities are recorded as frequency in 16 subplots. Beta diversity, measured as length of the first DCA axis, invariably increased upon lowering of sample plot size. The same applied to the eigenvalues of the axes. This is explained as a consequence of the weakening of structure in the data matrices when the fine-grained patterns of the vegetation are emphasized.     

毛乌素沙化草地景观生态分类与排序的研究

 本文通过对毛乌素沙地景观生态数据的处理与分析,以广泛应用于植被科学的数量分类方法TWINSPAN为工具,对毛乌素沙地景观生态类型进行了自上而下的等级式的数量分类研究,并与实际情况相结合,提出了毛乌素沙化草地景观生态分类系统。然后又利用算法上与TWINSPAN相似的DCA对沙化草地景观生态类型进行了排序分析,并与数量分类结果相结合,辅以逐步回归分析等多元分析方法,对所划分的各景观生态类型之间的关系进行了分析与探讨,指出毛乌素沙化草地具有较为丰富的景观生态类型,地下水位,覆沙厚度,基质类型控制着沙化草地景观生态类型的发生与演化;提出了毛乌素沙化草地景观生态的演化模式。本文还表明,TWINSPAN和DCA是景观生态研究比较有效的数量分析方法。     

林区火灾后植物群落与冻土环境的多元分析

本文运用国际较为先进的双向指示种分析(TWINSPAN)多元等级分划分类方法和无趋势对应分析(DCA)排序方法对大兴安岭阿木尔林区1987年特大森林火灾后的植物群落进行了研究。通过排序结果和冻土环境的多元分析,得出：DCA排序轴1代表了土壤岩性与地形的梯度而决定植物群落的分布;DCA排序轴2则反映了森林火烧的程度梯度,并通过冻土融化深度和浅层地温值的变化而影响植物群落的分布,运用相关回归分析方法,定量地找出了它们之间相互关系的回归方程。 据此,进而探讨了该地区森林火灾后冻土环境的变化对植物次生演替方向和进程的影响。