The ecology of a beech forest on Mt. Sanpoiwadake,Hakusan National Park,Japan I. Braun-Blanquet,TWINSPAN and DCA analysis

The Lindero membranaceae-Fagetum crenatae association on Mt. Sanpoiwadake, Hakusan National Park, Japan, has been classified using traditional Braun-Blanquet methods. The association was floristically sub-divided into six communities, all of which had been included as four subassociations by Hukusima (1982). The resulting table was compared with an analogous community table derived using computer-based TWINSPAN analysis. This dual approach was shown to provide a fully objective classification by method, the subjective element in the Braun-Blanquet method thus being substantiated TWINSPAN. Subsequent ordination of the data using DCA then provided a further degree of objective evaluation allowing any misclassified stand or species to be re-examined, and at the same time displaying the detailed inter-relationships between all species or stands.     

南亚热带常绿阔叶林取样技术研究

 本文研究了样地取样技术和无样地取样技术在南亚热带常绿阔叶林的应用。认为中心点四分法无样地取样技术具有较高的精度和效率,完全适用于南亚热带常绿阔叶林。 应用中心点四分法无样地取样技术所获得的能反映群落真实特征的中心点数目,称为“最小点数”。评定最小点数,可应用种类数目与中心点数目的相关曲线,即“种一点数曲线”;以及优势种重要值与中心点数的相关曲线,即“重要值一点数曲线’。据研究南亚热带常绿阔叶林的最小点数为20—22点。 “重要值—面积曲线”与“重要值一点数曲线”是评定最小面积与最小点数的测度新方法,在某种意义上较“种一面积曲线”更为理想。     

广东中西部桉树人工林植物多样性与林龄和土壤因子的关系

以位于广东省中西部的高要市桉树人工林林下植被群落为研究对象,对不同龄级林分物种多样性进行比较分析,采用典范对应分析(CCA)方法进行排序并通过相关分析,对物种分布和多样性与土壤因子的关系进行了研究。结果表明,每龄级24块样地充足,从第9块样地开始不同龄级的物种数目出现一定程度的差异。桉树林分样地中共有136种植物,隶属54科107属,灌木层的种类最多,但以草本层物种为优势种。不同龄级林分之间物种多样性差异不显著,但1—2年生林分低于2—4年生和5—6年生两个龄级;3—4年生的Shannon指数、Simpson指数和丰富度最高。对植被物种的分布,土壤中有机质、全磷、全钾和容重的影响明显,pH值也有一定程度作用,而对物种多样性,pH值和全氮是最为稳定的影响因子。     

Long‐term effects of the abandonment of grazing on steppe‐like grasslands

Question: What are the consequences of grazing abandonment on the Stipa lessingiana dominated steppe‐like grasslands? What is the relative importance of management and environmental factors in causing variation in species composition and abundance in the continuously grazed and abandoned grassland stands? Location: Transylvanian Lowland, Romania. Methods: Repeated vegetation mapping of a grassland stand, where grazing was abandoned 35 years ago; re‐sampling six grassland stands surveyed 29–57 years ago. For revealing long‐term changes in species composition and rank abundance PCoA ordination was applied. The relative importance of management and environmental factors in structuring vegetation were explored by CCA ordination. Diversity, evenness and the relative number and abundance of red‐listed species were compared between managed and abandoned stands. Results: Our results pointed out that grasslands which were formerly grazed and dominated by S. lessingiana, in the long‐term absence of grazing, have been transformed into a S. pulcherrima dominated type. Management, probably by creating bare surfaces and preventing litter accumulation, had the strongest effect on the species composition and abundance in the grasslands. Abandoned grassland stands had lower diversity and evenness compared to continuously grazed stands. While at the same time, the relative number of threatened, rare species did not differ between managed and abandoned sites. Conclusion: Maintaining extensively grazed, as well as un‐managed, Stipa dominated grasslands would be important in order to create various habitat conditions for plant species, especially threatened and rare species, and promote diversity on the landscape scale.     

An application of ordination to the identification of forest types

Summary The problems of allocating new vegetation stands to an existing classification are outlined. As one solution, the vegetation samples from an area may be ordinated by reciprocal averaging and classified by dividing the ordination into community-types. The ordination scores for a short list of species present in a new sample may then be averaged to locate that sample on the ordination axes and in one of the community-types. The method is tested by comparing estimates from incomplete species lists with that from a complete list of species in an forest stand in southern Ghana.We gratefully acknowledge the assistance of Mr. M.O. Hill in the correct formulation of the coordinate estimation procedure.     

Correlation between stand structure and ground vegetation: an analytical approach

The effect of stand structure on the diversity of the forest's ground vegetation was examined based on data on permanent sample plots collected in the northern parts of North-Carelia, eastern Finland. Different ordination methods (DCA, GNMDS, LNMDS, and HMDS) were used together with the TWINSPAN classification method. The aim was to construct a basis for classifying forests with respect to the biodiversity of the forest vegetation. Fertility and stand age showed the strongest correlation with the variation in ground vegetation. Other important factors were basal area, tree species composition, and crown cover. These variables were important in the division of the sample plots into different classes. According to the diversity indices, species diversity was at its highest in young stands on fertile forest sites. As a result, twenty-one different classes were formed based on the relative abundance of understorey species. The variables with the highest correlation were used to describe the stand structure in these classes.     

Forest community analysis and the point-centered quarter method

The point-centered-quarter (PCQ) method has been applied in community analysis since the publication of the method nearly 50 years ago. This and other distance methods offer increased sampling efficiency over fixed-area plots (FAP), but have long been known to produce biased density estimates when plant distribution deviates from random spatial patterns. Spatial indices have been developed to quantify the direction of this bias when plant distributions are aggregated or evenly distributed. Its continued use, especially in community analysis, requires additional scrutiny in measurements of community structure. We measured 14 forest stands of varying age, elevation and disturbance regime using FAP and PCQ methods. Density estimates were biased, with the point-centered quarter method lower than fixed-area plot estimates when stems were aggregated and higher when stems were evenly spaced. In general the PCQ method underestimated species richness. The efficiency of the PCQ method makes it popular for ordination studies, although comparison of community structure varied from 18% to 90% similarity between the measurements of species basal area in the same stands using the two different methods. The bias observed in calculations of stem density, species abundance and community similarity indicate that use of the PCQ method should be approached with caution when used in community level analysis.     

Dynamics of the horizontal structure of the tree stand in a pristine herb–bilberry forest of the subzone of the middle tiaga in the Komi Republic

According to observations from 1987 to 2010 on a constant sampling plot, using spatial statistics, we have shown dynamics of the horizontal structure of all-aged stands of a pristine spruce herb–bilberry forest. We note that the trees forming it provide a continual change of generations and, therefore, random community structure and its spatial stability. We determined that small-leaved arboreal plants, which represent part of the tree stand, tend to grouping at distances from 4 m. It was discovered that the mutual arrangement of trees of different sizes and species do not differ from a random occurrence.     

Permanent seed banks in chalk grassland under various management regimes: their role in the restoration of species-rich plant communities

A survey was carried out on four stands of a chalk grassland (open, intensively grazed, tall and scrub) in order to monitor the permanent soil seed bank under various management regimes responsible for the decline of the ecological interest of this ecosystem. There was no real similarity between the seed banks and the associated vegetation, except in the intensively grazed stand. When vegetation changes, seed banks appear to be quite stable and are mostly dominated by three species (Centaurium erythraea, Origanum vulgare, Hypericum perforatum). The seed bank seems to be quite useless in order to restore a species-rich community after both grazing intensification or abandonment. Because recolonization processes are the critical factor, the management of species-rich grassland has to be designed with a view to preserving a given plant community as well as to improving dispersion processes from a source to target areas.     

Disparate distribution patterns between canopy and subcanopy life-forms in two temperate North American forests

Quantitative vegetational data of canopy and woody subcanopy species (two life-forms adapted to occupy different strata at maturity) were compared with data collected in two temperate forest ecosystems to determine whether they exhibit a similar pattern of distribution. Tidal freshwater swamps (21 stands) and southern Appalachian forests (19 stands) were examined from data obtained using identical sampling methods. Separate structural analyses of the canopy, sapling, and subcanopy species were compared using the indirect ordination algorithm Detrended Correspondence Analysis. Environmental measurements collected in each stand were assessed for their relationship to the distribution of stands depicted by the ordination diagrams.Canopy trees and saplings showed a similar pattern of distribution, suggesting that the resource requirements of saplings and canopy-statured adults are similar. In contrast, the subcanopy species (species genetically adapted to an understory existence, i.e., shrubs and small understory trees) of neither ecosystem showed any discernable distributional relationship to the canopy or sapling layers (in tidal swamps, there was no clear way to even segregate subcanopy stands into communities). Environmental gradients associated with the subcanopy ordinations differed from those of the canopy and sapling strata in both forest systems, suggesting that subcanopy species partition different resources than do canopy species.If a lack of similarity in distribution patterns between canopy and subcanopy species is universal in temperate forests, then the common practice of combining sapling and subcanopy species in structural analyses may hinder our understanding of subcanopy structural patterns in forests.     

Plant communities,diversity and endemism of the Kula Volcano,Manisa, Turkey

Volcanoes often harbour specialized plant communities and shelter endemic plant species. Kula Volcano is one of 14 volcanoes in Turkey. Although this volcano is clearly a landmark of the Aegean region, only few botanical studies analysed the vegetation pattern at the Kula Volcano. None performed a phytosociological classification to delimit different plant communities. We applied a stratified random sampling design according to altitude and aspect and sampled 112 vegetation plots. We classified plant community types using a modified TWINSPAN analysis followed by the determination of diagnostic species based on φ coefficient fidelity values. Floristic relationships between plant community types were interpreted by ordination and ANOSIM analyses. Further, we used partial correlations of the ordination axes and environmental parameters in order to identify relationships between vegetation zonation and environment. We identified five major plant community types based on 85 diagnostic species. These plant community types were significantly correlated with altitude and aspect. Further, 13 endemic plant species were found from which one was endangered and one was classified as vulnerable according to International Union for Conservation of Nature (IUCN).     

Understorey vegetation development in North Finnish Picea forests after disturbance: re‐analysis of Sirén's data

Abstract. Sirén (1955) studied understorey species composition, tree stand properties and humus‐layer thickness in 64 unlogged forest stands on topographically and pedologically comparable sites. The stands were of even age (6 – 300 yr), stocked with the first or second tree generation after wildfire. The view of Sirén and several authors after him, that the vegetation of old‐growth boreal Picea forests is homogeneous on a broad scale, was examined by applying, in parallel, the partial variants of two ordination methods (DCA and PCA) to Sirén's vegetation data. Two main vegetation gradients were found: a major gradient running from recently burnt plots with prominence of pioneer species to plots with stand age > 100 yr, a well stocked tree layer and a thick humus layer, dominance of feather‐mosses and ample occurrence of shade‐tolerant as well as light‐preferring vascular plant species, and a second gradient along which first‐ and second‐generation plots segregate. The more prominent element of Betula trees in first‐ than in second‐generation stands < 100 yr contributed to the latter. A minor third gradient related to humus‐layer thickness was recovered by partial DCA only. The main vegetation gradient reappeared in separate ordinations of data from 47 mature forest stands (> 100 yr), but without being correlated with forest age. Variation among mature‐forest stands in the importance of pioneer species is considered mainly to be brought about by fine‐scale disturbance processes such as tree uprooting. Increasing importance of factors operating on within‐stand scales [development of a varied gap structure and stronger gradients in tree influence (radiation at ground level), soil moisture, soil depth and nutrient availability] with time is also reflected in the second and third mature‐forest ordination axes. Possible implications of the results for conservation of biological diversity and monitoring of changes in boreal forests are discussed.     

广东森林群落排序分析

本文应用极点排序和位置向量排序的方法对广东13个森林群落进行排序分析,并分别用极点排序图和位置向量排序的二维图和三维图表示排序的结果。同时对排序图的生态学意义及排序方法的优缺点进行讨论。结果表明,三维位置向量排序图能较好地把性质相近的群落类型聚在一起,可作为植被分类的辅助方法;积点排序图从一定程度上反映了植被的连续变化;极点排序与位置向量排序虽然取得一定结果,但由于同属线性排序,损失的信息量较多,寻求非线性排序方法是研究的方向。     

Stand characteristics,understorey associates and environmental correlates ofEucalyptus tetrodonta F. Muell. forests on Gunn Point,northern Australia

An ordination study of 20Eucalyptus tetrodonta forest stands growing on deep earths in monsoonal Australia revealed two major gradients in understorey vegetation type. The first axis reflected both floristic and structural understorey variation, where litter and shrub cover were inversely related to grass cover. This axis is thought to reflect a complex fire-vegetation type interaction, where vegetation is primarily determined by the saturation of the soil profile in the wet season, as measured by the colour of the iron rich soils. On the second axis of the ordination, floristic composition but not vegetation structure, and stand height were found to vary with the intercorrelated measures of soil gravel and moisture supply.E. tetrodonta is able to regenerate in the absence of fire, but firing appears to stimulate regeneration. All stands contained some advance growth, which occurs in distinct clumps, probably reflecting these plants clonal origin. Sapling presence in the stands is variable and the recruitment of advance growth into this size class appears to be related to over-wood competition. The size class distribution of trees was found to be similar amongst the stands, therefore stand structure appeared to be independent of understorey type. In comparison to general models of temperate eucalypt regeneration processes the tropical eucalypts have evolved different regeneration strategies, possibly in response to the severe annual drought.Nomenclature: Chippendale (1971), unless otherwise indicated. Structural classification: Specht (1970).I acknowledge the help of Mr. Clyde Dunlop who identified plant specimens and provided stimulating discussion, and Dr. Peter Minchin for permission to use the computer package ECOPACK to prepare and handle the floristic data, and his invaluable assistance with the ordination analysis. Mr. Lee Belbin and CSIRO Division of Water and Land Resources kindly gave their permission to use the Numerical Taxonomy Package (NTP).     

Soil seed bank in different habitats of the Eastern Desert of Egypt

The floristic composition and species diversity of the germinable soil seed bank were studied in three different habitats (desert salinized land, desert wadi, and reclaimed land) in the Eastern Desert of Egypt. Moreover, the degree of similarity between the seed bank and the above-ground vegetation was determined. The seed bank was studied in 40 stands representing the three habitats. Ten soil samples (each 25 × 20 cm and 5 cm depth) were randomly taken per stand. The seed bank was investigated by the seedling emergence method. Some 61 species belonging to 21 families and 54 genera were identified in the germinable seed bank. The recorded species include 43 annuals and 18 perennials. Ordination of stands by Detrended Correspondence Analysis (DCA) indicates that the stands of the three habitats are markedly distinguishable and show a clear pattern of segregation on the ordination planes. This indicates variations in the species composition among habitats. The results also demonstrate significant associations between the floristic composition of the seed bank and edaphic factors such as CaCO₃, electrical conductivity, organic carbon and soil texture. The reclaimed land has the highest values of species richness, Shannon-index of diversity and the density of the germinable seed bank followed by the habitats of desert wadi and desert salinized land. Motyka’s similarity index between the seed bank and the above-ground vegetation is significantly higher in reclaimed land (75.1%) compared to desert wadi (38.4%) and desert salinized land (36.5%).     

Gradient structure of forest vegetation in the central Washington cascades

Summary Forest vegetation located in three areas of the central Washington Cascades, arrayed along a gradient of increasing continentality and decreasing rainfall, were compared using ordination methods. Within each region, lower and upper elevation sites were analyzed separately and for each set of sites, trees and ground story vegetation were analyzed independently. Principal components analysis, reciprocal averaging, weighted averaging, and polar ordination were applied to each set. The characteristics of the data determined which method gave the most readily interpretable results, but RA and WA usually best ordered the stands along a complex, combined coenocline, while PO often decomposed the gradient into moisture and temperature components. PCA was of little use, even with relatively low beta diversity.Results are presented in the form of stand by species tables for each data set and the most appropriate two dimensional ordination. Both are correlated to the classification. A mosaic diagram for each region is synthesized from these analyses and habitat data. These diagrams indicate that community types occupy a smaller portion of the habitat as continentality increases.The ordination results were in close agreement with our earlier classification of these stands. The maritime (west) region contains stands of low richness and with relatively little coenocline differentiation. In contrast, the continental (east) region has high alpha and beta diversity. Species in the west tend to occupy a broad portion of the available habitat range, while species in the east do not. The analyses reveal that ordinations by different strata may produce similar stand sequences if both strata are responding directly to the same factors, but that the correspondence degenerates where the understory responds primarily to the nature of the canopy dominant species. Thus stand sequence correlations are highest in the lowland eastern region and lowest in the lowland western region.This study demonstrates that none of these metric ordination methods is fool-proof and that none should be used exclusively or in isolation. RA and PO are demonstrated to be useful general methods; WA gave results similar to those of RA. PCA never produced uniquely superior results. Analyses with too few species and moderate beta diversity often produce distortions as pronounced as those with many species and high beta diversity.Nomenclature follows Hitchcock & Conquist (1973).Funds provided by the Graduate School Research Fund, University of Washington and by grants GB-20963 and GB-36810X to the Coniferous Forest Biome, U.S.I.B.P. This is contribution # 318 to the Coniferous Forest Biome. M.F. Denton, R.S. Fleming, A.R. Kruckeberg, and R.H. Whittaker each made significant suggestions. We thank R.S. Fleming, S.G. Fleming, C. Brewer, B.C. Cannon, K.E. Wade, K. Loughney, and J.E. Canfield for their assistance with field work and data analysis.     

Long-term growth dynamics of natural forests in Hokkaido,northern Japan

Abstract. The long-term growth dynamics of natural forest stands on the island of Hokkaido were described on the basis of an analysis of data from 38 permanent plots spanning 15–22 yr. Stand structure was characterized by basal area, stem density and tree size variability. To detect trends in stand structure, regression models for recruitment rate (per ha per yr), mortality rate and the rate of change in stem density and tree size variability were developed by a stepwise method using initial basal area, stem density, tree size variability, species composition summarized by LNMDS ordination, altitude, annual mean temperature, annual precipitation, type of understorey vegetation, topography and slope aspect as candidates for predictor variables. The same analyses were conducted for basal area increment (net growth) and its components: survivor growth = basal area gain by growth of surviving individuals and mortality = basal area loss by death of individuals. Stem density remained generally unchanged; recruitment was relatively low even in very sparse stands. Stand basal area generally increased as survivor growth was approximately double the mortality. Recruitment rate was strongly affected by the presence of dwarf bamboo (Sasa spp.) vegetation on the forest floor which inhibited tree regeneration. Mortality rate was density-dependent; dense stands had higher mortality than sparse stands. Density change rate (recruitment rate - mortality rate) was, therefore, determined by both the type of understorey vegetation and stem density. Survivor growth was high in stands with high stem density and basal area. Mortality was dependent on basal area and altitude. Net basal area increment (net growth) was dependent only on stem density with other factors that influenced survivor growth and mortality omitted. Tree size variability decreased in stands with high tree size variability whereas it increased in stands with low size variability. Based on the obtained models for density change rate and net basal area increment, trajectories of stands were illustrated on a log-log diagram of stem density and basal area. The predicted differences in trajectories as affected by the understorey vegetation type indicated the importance of dwarf bamboo vegetation for forest dynamics on Hokkaido.     

The Finnish forest site type approach: ordination and classification studies of mesic forest sites in southern Finland

A.K. Cajander's forest site type classification system is based on definition of plant communities typical to certain climatical and edaphical site conditions, but the structure and composition of the tree stands in Finland are considered sensitive to random variation and are therefore not used as primary classification criteria. The system has often received criticism, usually that the effects of the tree stand and successional stage of the stand have been underestimated. Most of the present-day forest stands in Finland represent young successional stages and are subjected to intensive management. This should result in an additional difficulty in the application of the forest site types in the field.The present study is based on three independent data sets representing forests on mineral soil in southern part of Finland. TWINSPAN classification, DCA ordination and canonical correspondence analysis (CCA) techniques were applied in successive stages of the data analysis. It was found that the definition of the intermediately fertile, mesic site types was clearly confused by the effects of the tree species and age of the stand. The analyses also revealed that the succession pathways on mesic forest sites are largely determined by the tree species composition. In stands dominated by Pinus sylvestris, the succession follows the competitive hierarchy model, whereas in stands dominated by Picea abies, severe shading of the tree canopy governs the development of understorey vegetation.Abbreviations CCA Canonical correspondence Analysis - DCA Detrended correspondence Analysis - TWINSPAN Two-way indicator species analysis     

The influence of soil cover organization on the floristic and structural heterogeneity of a Guianan rain forest

The impact of soil cover organization on the forest community has been studied in a 19-ha tract at Piste de St Elie station in French Guiana. 195 species each represented by at least 10 individuals were chosen from records of the position, diameter at breast height (dbh) and precise identification by botanical sampling of 12104 ligneous plants (dbh 10 cm).Spatial variations in the soil were mapped using the method proposed by Boulet et al. (1982). The soil mapping units correspond to the successive stages of evolution of a currently unbalanced ferralitic cover. These stages describe firstly the thinning by erosion of the microaggregated upper horizon and secondly the mineralogical changes under more or less extended hydromorphic conditions. The degree of evolution of ferralitic cover is also related to the hydrodynamic functioning and chemical properties of the soil. Geological substrate, topography and slope have also been taken into account.Analysis of the influence of environmental variables on plant cover has been performed using the Ecological Profiles method and Correspondence Analysis (CA) of the table of ecological profiles.The forest community seems to be dependent on the soil and the topographical features that govern it. There are significant, exclusive soil-species links for each soil functioning mapping unit. However, the highest proportion of significant positive links is connected with a thick microaggregated horizon (25%). Several species are of real value as indicators and more particularly enable differentiation between the forest stands of typical ferralitic soil and the ones of thinned out, transformed and hydromorphic soils. The CA of the species by environmental variables matrices reveals two significant factorial axes. The first can be associated with the drainage mainly related to the thinning of the soil and the second with the hydromorphic conditions related to the topography. The vegetation ordination of the stands ( 0.25 ha) delimited in the various soil domains clearly shows that changes in ferralitic cover and in particular the transition from soil with deep vertical drainage to soil with superficial lateral drainage is accompanied by substantial changes in the forest community.     

Insect community response to switchgrass intercropping and stand age of loblolly pine (Pinus taeda) plantations

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