The steppes of Tierra del Fuego: floristic and growthform patterns controlled by soil fertility and moisture

Northern Tierra del Fuego is the insular extreme of the Magellanic steppe, a unique example of a cold-temperate oceanic grassland in South America. We surveyed this poorly known region with a landscape-based procedure to classify plant communities and investigate environmental gradients. Sampling sites were selected with the aid of aerial photos, and floristic composition, soil, topography and landform type were recorded in each site. Classification was achieved with Two-Way Indicator Species Analysis and gradients were analyzed with Detrended Canonical Correspondence Analysis. Two broad groups of communities were separated by the presence of typical acidophilous dwarf shrubs: acidophilous/mesotrophic and neutrophilous vegetation. Along the main floristic gradient, ericoid dwarf shrubs were replaced by forbs and grasses as soil acidity decreased. The acidic habitats were on Pleistocene glacial and alluvial deposits, and had coarse texture, poorly decomposed organic matter, and low base status. The eutrophic habitats were on Tertiary rocks of medium to fine texture and had well-decomposed organic matter and high base status. A second axis of variation was associated to latitudinal position and slope aspect, revealing a precipitation/evaporation gradient. Scrubs dominated by midheight shrubs, with species from the southern forests, were at the humid extreme (higher latitudes/south aspects); and short grass steppes, with taxa commonly found further North in the continent, at the xeric end (lower latitudes/north aspects). Although fertility and moisture seem to control the overall floristic gradient, the extreme vegetation types of the fertility gradient, both having a very simple structure, could be attributed to grazing disturbance. Floristics and ecology of the Fuegian steppe resemble to a certain degree some boreal oceanic systems.     

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.     

山西高原植被与气候的关系分析及植被数量区划的研究

本文利用去势典范对应分析和数量区划的方法,研究了山西高原植被与气候之间的关系,并进行了数量区划。排序的结果表明：DCCA的第—轴代表山西高原植被和气候梯度的纬向性,热量梯度是决定植被分布最主要的气候因子,水分梯度中的年降水量也对第—轴有较大的影响,由于山西高原南北跨度大,植被与气候因子表现出明显的纬向性;DCCA第二轴代表山西高原植被和气候梯度的经向性,与DCCA第二轴相关性较大的是水分因子中的年降水量、年蒸发量,由于山西高原东西跨度不大,而且大部分地区处在吕梁山脉和太行山脉之间,东西向的气候变化幅度不大,所以植被与气候梯度的经向性不明显。植被数量区划的结果表明：山西高原可划分为17个植被区,用图示的方法确定山西高原大致有三个极点和—个中心。     

中条山混沟地区森林乔木种的数量分类与环境解释

通过对中条山混沟地区森林植被调查资料的多元分析——TWINSPAN分类、CCA排序与环境解释,划分了该地区的植被类型,给出了植被与环境因子的定量关系。结果如下:1)40个样地可划分为青檀林(Form.Pteroceltis tatarinowii)、栾树林(Form. Koelreuteria paniculata)、槲栎林(Form. Quercus aliena)、栓皮栎林(Form. Quercus variabilis)、鹅耳枥+葛萝槭林(Form. Carpinus turczaninowii+Acer grosseri)、元宝槭+千金榆林(Form. Acer truncatum+Carpinus cordata)和辽东栎林(Form. Quercus liaotungensis)7种群落类型,体现了中条山地带性植被类型为暖温带落叶阔叶林的特点。2)群落类型的划分主要反映了CCA排序第一、二轴的环境梯度,CCA排序第一轴突出反映了海拔与土壤养分梯度,第二轴与土壤pH值、湿度指数和坡度显著相关。总体来说,海拔和土壤因子是影响混沟地区乔木物种分布分异的最主要环境因子。3)环境因子和空间因子解释了物种格局变化的46.14%,其中环境因子占30.79%,空间因子占8.48%,空间因子和环境因子交互作用解释的部分占6.87%。良好的环境解释反映了调查取样和环境因子选取的合理性,同时也体现了混沟地区植被的原始性。     

Altitudinal pattern of vegetation variation on Tenerife

Abstract. The variation in structure and floristic composition of the vegetation of Tenerife is related to different environmental factors. 200 sample plots, stratified according to altitude, exposure to wind, kind and age of parent rock, inclination and degree of disturbance were analysed throughout the island. The resulting data were analysed using univariate statistics (altitudinal distribution profiles of the more frequent species) and multivariate methods (Correspondence, Cluster and Pattern Analysis). Altitude, exposure to wind and, to a lesser extent, disturbance were found to be the major determinants of vegetational variation, whereas parent rock and inclination were less important. Ordination results show a continuum in structural and floristic variation of the vegetation with species distributions varying continually along the altitudinal gradient. Nevertheless, altitudinally bound plant community types were recognized with Cluster Analysis. Vegetational discontinuities, which were also detected by pattern analysis, are more evident on the windward slope of the island; they have been attributed to the existence of a major underlying environmental discontinuity the ‘cloud sea’-which is associated with a climatic inversion. On the leeward slope, beyond the influence of the cloud sea, discontinuities are less evident and vegetational variation seems to be closer to a coenocline, i.e. a vegetational continuum.     

拉萨河流域植物群落的数量分类与排序

青藏高原植物群落空间分异格局是异质生境条件下物种性状、种间相互作用等生态学过程共同作用的结果,对其分析有助于深入理解群落形成与环境因子之间的关系。基于拉萨河流域自然植被样带调查,采用双向指示种分析(TWINSPAN)和典范对应分析(CCA)等方法,探讨了群落的结构组成及影响其结构分异的主导环境因子。结果表明:(1) TWINSPAN数量分类将拉萨河流域草地系统划分成12个群系类型,即圆叶合头菊+唐古拉翠雀花群系;紫花针茅群系;青藏臺草群系;雪层杜鹃+鲜卑花-西藏嵩草群系;高山嵩草群系;小叶金露梅群系;硬叶柳+杯腺柳群系;水栒子+拱枝绣线菊-高山嵩草群系;绢毛蔷薇-冷蒿+白草群系;大果圆柏-垂穗披碱草群系;铺地柏-藏橐吾+高原荨麻群系;醉鱼草+砂生槐群系。12种群系类型包含了较多的植被类型,包括高寒灌丛草甸、高寒灌丛草原、稀树草原、高寒草甸和高寒草原等。(2) CCA排序表明:影响拉萨河流域植物群系分布的主要环境因子是年均温度、海拔和经度和纬度,其次是年均降雨量。(3) TWINSPAN分类与CCA排序结合反映了群系分布格局变异与环境因子之间的关系,可为拉萨河流域草地的保护和可持续利用,以及相关的植被群落研究提供参考。     

雅鲁藏布江河岸植物物种丰富度分布格局及其环境解释

沿环境梯度物种多样性的分布格局及其环境影响因子的研究是生态学研究的重点内容之一。雅鲁藏布江是我国海拔最高的大河, 目前还没有针对其流域的河岸植物群落物种丰富度分布格局的系统研究。作者沿海拔梯度从雅鲁藏布江上游源头区至下游共设置了15个海拔样带, 在每个海拔样带随机取6个5 m×5 m样方, 共对90个样方的河岸植物群落物种组成和植被类型进行了调查, 共记录到238种维管束植物。DCA排序结果表明物种组成变化主要受海拔和经度控制。CCA排序区分出5种大的植被类型, 即高寒草甸或高寒草原、高山草原、两种亚高山灌丛草原和林缘灌丛植被。各植被类型的分布和变化主要受海拔和经度影响。雅鲁藏布江河岸植物物种丰富度沿海拔和经度梯度呈浅“U”形分布格局, 物种丰富度与年均降水量和植被盖度成显著正相关。     

Vegetation ecology of phytogenic hillocks (nabkhas) in coastal habitats of Jal Az-Zor National Park,Kuwait: Role of patches and edaphic factors

Phytogenic sandy hillocks (Arab. ‘nabkha’) are very frequently occurring aeolian deposits along the coastal plain of Kuwait. We investigated the vegetation of 42 nabkhas in a coastal habitat of Jal Az-Zor National Park, Kuwait. Sixty-two species were recorded (47 annuals and 15 perennials) in the studied nabkhas. Four vegetation types were recognized after classifying the vegetation of the nabkhas by TWINSPAN. They were named after their dominating host species which are Nitraria retusa, Zygophyllum qatarense, Haloxylon salicornicum and Panicum turgidum. Using Detrended Correspondence Analysis (DCA) and Canonical Correspondence Analysis (CCA), we assessed the relationships between environmental gradients, floristic composition, species diversity, and geomorphology aspects of the studied habitats. Notable environmental variables affecting the distribution of the vegetation types in the study area were: geomorphological aspect, size of plants forming the core of the nabkha, moisture and nutrients contents, salinity, sand and silt components, and pH.     

长白山北坡植物群落间物种共有度的海拔梯度变化

群落同各生活植物及种所有植物种的共有度,一定程度上反映了群落间的相关性及沿环境梯度的物种替代关系。应用梯度格局法在长白山北坡海拔700－2600m之间,海拔每上升100m设置一块样地,共计20块样地,应用Jaccard指数,对各海拔植物群落间物种共有度的海拔梯度变化进行了分析,不同海拔群落间物种的共有度,无论以哪一层次的植物来表达,基本都以与其最相邻海拔的群落之间为最高;如相邻海拔的两个群落属于不同植被类型,其共有度较低。群落间物种共有度的峰值与谷点反映了群落类型随海拔具有间断性变化,乔、灌、草各生活型物种的共有度以及所有植物物的共有度,在同一海拔差群落间的数值都非常接近,并均随着海拔的增加呈明显的下降趋势,表明群落各层次物种的构成有很大的相关性以及物种之间的共存性。     

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.     

东灵山华北落叶松人工林的分布格局及环境解释

应用TWINSPAN、CCA和物种多样性分析,从植物群落、植物种与环境的生态关系等方面研究了东灵山华北落叶松人工林的植被分布格局,并给予合理的环境解释。TWINSPAN分类将华北落叶松人工林分为4个类型,分类结果在CCA二维排序图上得到了较好的验证。样方与物种的CCA排序较好地揭示了该区华北落叶松人工林的分布格局与环境梯度的关系：坡位、坡度、土壤厚度和海拔是影响该区华北落叶松人工林的主导因子。结合物种多样性分析,结果表明东灵山绝大多数华北落叶松人工林生长良好,并根据华北落叶松喜凉湿习性,建议在该区自然分布海拔以下造林时,应优先选择阴坡和半阴坡的中、下位坡。     

Floristic composition in relation to environmental gradients across KwaZulu‐Natal,South Africa

Conservation planning in the face of global change is still in its infancy. A suggested approach is to incorporate environmental gradients into conservation planning as they reflect the ecological and evolutionary processes generating and maintaining diversity. Our study provides a framework to identify the dominant environmental gradients determining floristic composition and pattern. Nonmetric multidimensional scaling was used on 2155 sampling plots in savanna and grassland habitat located across the province of KwaZulu‐Natal, South Africa (94 697 km²), a floristically rich region having steep environmental gradients, to determine the dominant gradients. Hierarchical cluster analysis was used to group similar plots which were then used in a Classification and Regression Tree analysis to determine the environmental delimiters of the identified vegetation clusters. Temperature‐related variables were the strongest delimiters of floristic composition across the province, in particular mean annual temperature. Frost duration was the primary variable in the Classification and Regression Tree analysis with important implications for savanna/grassland dynamics. Soil properties (base, pH status) and moisture variables accounted for most of the variation for the second and third axes of floristic variation. Given that climatic and edaphic variables were well correlated with floristic composition, it is anticipated that a changing climate will have a marked influence on floristic composition. We predict warmer temperatures may facilitate the spread of frost sensitive savanna species into previously cooler, grassland areas. Species associated with specific soil types will not easily be able to move up the altitudinal gradient to cooler climes because geology is aligned in an approximately north‐south direction compared with increasing altitude from east‐west. Future conservation planning should take cognisance of these gradients which are surrogates for ecological and evolutionary processes promoting persistence.     

Relationships between vegetation and climate on the Loess Plateau in China

The Loess Plateau is one of the most environmentally sensitive regions in China. This study addresses the relationships between vegetation and climate of this area quantitatively at a large-scale, in order to determine the factors that control vegetation distribution. The Loess Plateau, located at 101°01′–155°10′ E and 34°02′–40°40′ N, covers an area of 52 million hectares. Vegetation data were collected from the vegetation map (1:500,000) and the Landsat Thematic Mapper scenes of the Loess Plateau. The Loess Plateau was divided into small districts of 30′ latitude by 30′ longitude on the vegetation map. In each district, areas with different vegetation were measured and used as vegetation data. The climatic data were average values of county meteorological records in each district in the past 25 years. GIS, TWINSPAN and canonical correspondence analysis (CCA) were employed for analysis. 257 small districts were clustered into 7 groups using TWINSPAN, representing 7 vegetation regions or subregions. The first three CCA axes had significant correlations with climate. The first CCA axis represented the variation of vegetation and climate along the latitude gradient, while the second CCA axis the variation along the longitude gradient. The distribution pattern of 171 vegetation formations on the CCA plot is identical to that of vegetation regions (districts). The spatial distribution of vegetation is closely related to climate variables on the Loess Plateau. Water variables and temperature are important in both latitude and longitude gradients, while the sunshine hours, accumulated temperature and wind speed are more important than water variables and temperature in longitude gradients.     

Altitudinal zonation of montane Quercus forests along two transects in Chirripó National Park,Costa Rica

Abiotic and vegetation data were collected along two altitudinal transects through mature montane Quercus forests on the Pacific and Atlantic slopes of Costa Rica's Chirripó Massif. Between 2000 and 3200 m asl twenty-four 0.05 ha forest plots were selected at altitudinal intervals of 100 m, and eight soil profiles were described at intervals of 200 m. A TWINSPAN classification aided in the determination of eight zonal forest communities on the basis of their floristic composition. They are grouped in two sets of four: (i) the palm-rich lauraceous-fagaceous Lower Montane Mollinedia-Quercus Forests (2000–2600 m asl) and (ii) the bamboo-rich myrsinaceous-fagaceous Upper Montane Schefflera-Quercus Forests (2500–3200 m asl), respectively. Vegetation changes seem correlated with two major climatic gradients: (i) a temperature gradient (altitude), and (ii) a moisture gradient (wet Atlantic vs. moist Pacific slope). Most soils are Andepts, and residual, colluvial or derived from volcanic material. Humus layers are thicker on the wetter Atlantic slope. A total of 431 vascular plant species consisted of 86 pteridophytes, 1 gymnosperm, 296 dicots and 48 monocots. Species richness, canopy height and stem diameter decrease with increasing altitude, while the canopy surface becomes more flattend. A comparison with other studies shows that Chirripó's montane Quercus forests fit within the environmental ranges known from altitudinal zonations elsewhere in the Tropics.Abbreviations asl above sea level - dbh diameter at breast height - LM Lower Montane - Mt. Mountain - TWINSPAN two way indicator species analysis - UM Upper Montane - VU code referring to soil profiles as presented in Van Uffelen (1991) This paper is dedicated to the memory of Alwyn H. Gentry, an outstanding and inspiring tropical botanist who tragically died in a plane crash in the mountains of Ecuador on August 3 1993, when surveying possible boundaries for a new tropical cloud forest reserve.     

河北小五台山国家级自然保护区森林群落与环境的关系

植物与环境之间的关系是一个复杂的演变过程,运用数量生态学方法探讨森林群落的物种组成、种群的生态特征、不同植物群落与环境之间的关系,有助于保护该区森林群落的稳定性和生物多样性。根据148个森林群落样方数据,选用双向指示种分析(TWINSPAN)和典范对应分析(CCA)方法,对河北小五台山国家级自然保护区森林群落进行分类和排序研究。结果表明:(1)TWINSPAN将该区的森林群落分为20个类型;(2)CCA排序结果较好地反映出群落分布格局与环境梯度的关系,各个森林群落类型在前两轴分异明显,在11个环境因子中,海拔、坡位、凋落层厚度、土壤导电率、土壤温度、土壤湿度、土壤厚度和干扰程度这8个环境因子对森林群落的分布起较大的作用,影响森林群落的分布格局,形成不同的植被类型。(3)乔木层优势种的CCA二维排序图所揭示的环境梯度与群落类型的分布有很大的相似性;t值双序图阐明了海拔、凋落层厚度、土壤温度、干扰程度等环境因子对森林群落乔木层优势种有着重要影响。采用TWINSPAN分类与CCA排序的方法,较好地解释了森林群落与环境因子的关系,为小五台山地区森林生态系统的科学管理和保护提供了理论依据,研究结果也为同类地区森林生态系统研究及保护提供参考和借鉴。     

Classification of dry grassland vegetation in Denmark

Abstract. A hierarchic classification of Danish semi‐natural grassland vegetation on well‐drained soils is presented. TWINSPAN was used for clustering of 614 samples of grassland vegetation showing floristic gradients and turnover in species composition in more dimensions. The optimal hierarchic level of clustering was determined by indicator species analysis. The classification was interpreted in terms of variables relating to abiotic environment and vegetation structure and to major ecoclines previously identified by gradient analysis. The 12 final clusters were compared to syntaxa of formal phytosociology and to communities in the British Vegetation Classification. Criteria for achieving floristically homogeneous clusters without sacrificing the ecological interpretability and validity of the clusters in time and along geographical gradients are discussed.     

Spatial distribution pattern of the riparian vegetation in a basin in the NW Spain

The riparian vegetation of a basin in the NW Spain was studied to establish its spatial variation pattern and to relate floristic and structural differences in the community to environmental factors. Eighty-seven sampling units in 43 sampling stations were used. Samples were classified in 5 groups using Two Way Indicator Species Analysis (TWINSPAN). Three groups represented reaches with riparian wood along their banks: Mediterranean alderwoods and shrubby willow woods. The remaining two corresponded to floodplains with vegetation colonizing moderately eutrophicated deposits of gravel. Structural characteristics of richness and diversity differentiated the alder woods. In these, the shaded environment created by the woody species limited herbaceous vegetation development. This was dominated by Carex acuta subsp. broteriana. Classification and CCA ordination results were compared. The TWINSPAN groups could be recognized in the CCA graph. The ordination was related to a pollution gradient associated with altitude. This pollution gradient involved bank ruderalization, incorporation of nitrophilous species and a decrease in the vegetation quality. The influence of the lithological features on vegetation was also evident.     

Topo-edaphic, floristic and physiognomic gradients of woody plants in a semi-arid African savanna woodland

The variation in topo-edaphic conditions, woodland composition and physiognomic structure of a semi-arid savanna woodland, namely the 59 400 ha Klaserie Nature Reserve, was studied with multivariate techniques. Spatial variation in the texture and chemistry of both the A-and B-horizons was substantial. One gradient of the A-horizon was of increasing clay, silt, organic matter, pH and conductivity, while a second gradient was of increasing rockiness, slope and gravel content, associated in part with upper landscape positions. Physiognomic structure of the woodland also displayed substantial spatial variation. Variation in tree density had the greatest effect on total canopy volume, and the density of coppice and dead individuals were closely correlated. Topo-edaphic variation, after the effect of vegetation composition had been accounted for, had a direct effect on woodland structure, as clay content of soils or rockiness influenced shrub density, and catenal position together with silt-or gravel-content of soils influenced coppice density. Ordination analyses identified that the included topo-edaphic variables accounted for a significant amount of floristic variation, but also showed that important environmental variables were omitted. Floristic variation of woody elements was of the nature of a large number of relatively equivalent, independent gradients rather than of a few simple primary gradients. Ten woodland types were identified by TWINSPAN which differed in their topo-edaphic and physiognomic character. Distinctive woodland assemblages were found on rocky outcrops or alluvial areas, on deep, sandy soils or on clay-or silt-rich soils. Colophospermum mopane formed the densest woodlands. Characteristics of the B-horizon were often dissimilar to those of the A-horizon, and seemed important for woodland composition.     

云南大围山种子植物区系海拔梯度格局分析

 海拔梯度包含了多种环境因子的梯度效应,因而研究山地植物区系的海拔梯度格局对揭示植物区系的环境梯度变化规律、了解生物适应性和生物多样性沿海拔梯度的变化趋势等具有重要意义。为了探讨山地植物区系构成特征及其海拔梯度的生态意义,该文根据对大围山国家级自然保护区植被线路调查和垂直样带调查,并结合文献研究等获得的植物区系资料,分析了该保护区种子植物区系构成的基本特征及其随海拔梯度的变化趋势;利用系统聚类的方法寻找和研究大围山植物区系沿海拔梯度变化的断点位置。研究结果表明：1)大围山大多数热带成分分布的上限位于海拔1 500 m左右,以此为界划分热带雨林和常绿阔叶林是合理的。 2)湿润雨林分布于海拔700 m以下;山地雨林分布于海拔700～1 500 m;季风常绿阔叶林分布于海拔1 300～1 800 m;山地苔藓常绿阔叶林分布于海拔1 800 m以上;在海拔2 100 m以上的迎风坡面、土层瘠薄的地段分布有不甚典型的山地苔藓矮林。