Simulation of multidimensional community patterns: towards a comprehensive model

Ecological gradients in the field layer of southern boreal forests in South Finland were studied in relation to the dominant tree species and the age of forest stands. The data are from a systematic sample of 529 plots from an area of 150 × 200 km, collected in the Third National Forest Inventory in 1951–53. Detrended correspondence analysis (DCA) was applied to log-transformed species cover values. It revealed three main gradients: fertility, moisture, and the effect of cattle grazing in forests (still extensive in the early 1950's). The fertility gradient dominated the first axis and the two latter sources of variation confounded with it in a complex manner in the first two axes of DCA. The second DCA axis was associated with canopy effects on understory pattern, with Pinus and Picea having opposite and Betula intermediate effects. These results were compared with an ordination model of Cajander's forest site types, based on DCA of independent, ideal data of 107 indicator species. The fertility gradient recovered by the model was almost identical to that obtained from the field data. The gradient was also stable from intermediate-age (40–69 yrold) to older forests. The forest site types showed rather large overlaps with main neighbouring types in composition of ground vegetation or nutrient status of the humus. Competitively efficient feather-mosses, which are dependent on nutrients released from the tree crowns, are considered important regulators of the understory vegetation. Accordingly, alternative approaches to the forest site type classification to be used in boreal forests treated by modern intensive forestry should give more weight to the effect of the canopy trees.     

Succession in sub‐boreal forests of West‐Central British Columbia

Abstract. Structural and compositional changes were analysed over the course of 400+ yr of post‐fire succession in the sub‐boreal forests of west‐central British Columbia. Using a chronosequence of 57 stands ranging from 11 to 438 yr in age, we examined changes in forest structure and composition with complementary PCA and DCA ordination techniques. To determine stand ages and timing of tree recruitment, approximately 1800 trees were aged. Most early successional forests were dominated by Pinus contorta, which established rapidly following fire. Abies lasiocarpa and Picea glauca × engel‐mannii were also able to establish quickly, but continued to establish throughout the sere. Few Pinus contorta survived beyond 200 yr, resulting in major changes in forest structure. In some stands P. contorta never established, which led to considerable variation among stands less than 200 yr old. The oldest forests converged on dominance by Abies lasiocarpa. Vascular plant diversity decreased during succession whereas canopy structure became more complex as gap dynamics developed. Although these sub‐boreal forests contain few tree species, successional changes were pronounced, with structure changing more than composition across the chronosequence.     

Implications of floristic and environmental variation for carbon cycle dynamics in boreal forest ecosystems of central Canada

Abstract. Species composition, detritus, and soil data from 97 boreal forest stands along a transect in central Canada were analysed using Correspondence Analysis to determine the dominant environmental/site variables that differentiate these forest stands. Picea mariana stands were densely clustered together on the understorey DCA plot, suggesting a consistent understorey species composition (feather mosses and Ericaceae), whereas Populus tremuloides stands had the most diverse understorey species composition (ca. 30 species, mostly shrubs and herbs). Pinus banksiana stands had several characteristic species of reindeer lichens (Cladina spp.), but saplings and Pinus seedlings were rare. Although climatic variables showed large variation along the transect, the CCA results indicated that site conditions are more important in determining species composition and differentiating the stand types. Forest floor characteristics (litter and humus layer, woody debris, and drainage) appear to be among the most important site variables. Stands of Picea had significantly higher average carbon (C) densities in the combined litter and humus layer (43530 kg‐C.ha^‐1) than either Populus (25 500 kg‐C.ha^‐1) or Pinus (19 400 kg‐C.ha^‐1). The thick surface organic layer in lowland Picea stands plays an important role in regulating soil temperature and moisture, and organic‐matter decomposition, which in turn affect the ecosystem C‐dynamics. During forest succession after a stand‐replacing disturbance (e.g. fires), tree biomass and surface organic layer thickness increase in all stand types as forests recover; however, woody biomass detritus first decreases and then increases after ca. 80 yr. Soil C densities show slight decrease with ages in Populus stands, but increase in other stand types. These results indicate the complex C‐transfer processes among different components (tree biomass, detritus, forest floor, and soil) of boreal ecosystems at various stages of succession.     

Changes in the understory of Canadian southern boreal forest after fire

Abstract. We investigated changes in the composition and abundance of understory species after fire in the southern boreal forest around Lake Duparquet, Québec. Ten plots of 100 m² were sampled in each of eight sites varying in post-fire age from 26 to 230 yr, with 20 1-m² quadrats in each of these 80 plots. Variation in the understory was described by DCA ordination and interpreted as a regeneration succession series. Thickness of the organic layers, stand age and canopy composition were all correlated with vegetational change. This change was not constant throughout succession; some old sites showed an increase in the diversity and abundance of certain pioneer species. This was partly related to openings in the canopy resulting from a major outbreak of spruce budworm, which affected sites dominated by Abies balsamea. The ordinations were performed on both the 100-m² plots and the 1-m² quadrats. Heterogeneity within sites was larger at the 1-m² scale and there was a great deal of overlap in the position of the quadrats in ordination space. At the smaller scale of analysis, stand age and thickness of the organic layers were not correlated with the changes observed in the understory.     

Classification of vegetational diversity in managed boreal forests in eastern Finland

Diversity of vegetation in managed forests is studied. A classification based on forest stand structure, the abundances of vegetation species and variations in these abundances is developed and diversity indices are calculated for the classes to describe the diversity of the vegetation within the classes. The classes were formed using detrended correspondence analysis (DCA), global nonmetric multidimensional scaling (GNMDS) and TWINSPAN classification. Discriminant analysis was used to determine the environmental variables differentiating between the classes, and Duncan's multiple range test was used to examine the ability of the diversity measures to distinguish the classes. Beta diversity was estimated with Økland's method based on DCA ordination of the sample plots. The results point to fertility and the successional stage of the stand as the main factors affecting species diversity, in addition to which soil type, the number of tree species, crown cover, basal area and certain variables describing the management of the stand were relevant to the classification. The most distinct diversity indices were the reciprocal of Simpson, Pielou's J' and species richness, while the highest alpha diversity was found in young stands with a low crown cover on herb-rich or mesic forest sites. Beta diversity was quite high, its highest values of all for the whole data being recorded along the fertility gradient.     

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.     

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     

Understorey vegetation change in a Picea mariana chronosequence

Eighteen black spruce (Picea mariana) stands, representing postfire ages of 26 to 120 yr, were surveyed for understorey vegetation and site/microsite characteristics at two spatial scales. This enabled comparison of within- versus among-stand compositional variation.Detrended correspondence analysis (DCA) ordination among the 18 stands revealed a complex age/moisture gradient. DCA ordination among 1 800 quadrats within the stands indicated a similar gradient with much compositional overlap. Quadrats were grouped, using two-way indicator species analysis (TWINSPAN), into 9 classes each representing a phase in understorey vegetation composition. These phases shifted in abundance from young to old stands with a high degree of concordance among replicates in the same age class. Understorey succession is strongly linked to the stages in tree growth, mortality and thinning coupled with the accumulation of site moisture.Abbreviations DCA Detrended Corrospondence Analysis     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

An Analysis of Vegetation Restoration on Opencast Oil Shale Mines in Estonia

We compared four types of 30‐year‐old forest stands growing on spoil of opencast oil shale mines in Estonia. The stand types were: (1) natural stands formed by spontaneous succession, and plantations of (2) Pinus sylvestris (Scots pine), (3) Betula pendula (silver birch), and (4) Alnus glutinosa (European black alder). In all stands we measured properties of the tree layer (species richness, stand density, and volume of growing stock), understory (density and species richness of shrubs and tree saplings), and ground vegetation (aboveground biomass, species richness, and species diversity). The tree layer was most diverse though sparse in the natural stands. Understory species richness per 100‐m² plot was highest in the natural stand, but total stand richness was equal in the natural and alder stands, which were higher than the birch and pine stands. The understory sapling density was lower than 50 saplings/100 m² in the plantations, while it varied between 50 and 180 saplings/100 m² in the natural stands. Growing stock volume was the least in natural stands and greatest in birch stands. The aboveground biomass of ground vegetation was highest in alder stands and lowest in the pine stands. We can conclude that spontaneous succession promotes establishment of diverse vegetation. In plantations the establishment of diverse ground vegetation depends on planted tree species.     

Understorey vegetation in boreal Picea mariana and Populus tremuloides stands in British Columbia

Abstract. We compared the species composition and species density of vascular plants in the understorey vegetation of boreal forest between Picea mariana (Black spruce) and Populus tremuloides (Trembling aspen) stands in British Columbia, Canada, and related differences in species composition and species density between the two forest types to dominant canopy tree species as well as a wide variety of environmental factors. We analysed 231 stands, distributed in three different climatic regions representing drier, wetter, and milder variations of montane boreal climate. Of these stands 118 were dominated by P. mariana and 113 by P. tremuloides. P. tremuloides stands had higher species density than P. mariana stands in all climatic regions, but species density of each dominance type varied among climatic regions. The floristic composition of the understorey vegetation was markedly different for P. mariana and P. tremuloides dominated stands. A detailed study on the effect of canopy dominance and local environmental factors on the understorey vegetation of the boreal forest was conducted using 88 stands from one of the three climatic regions. Using a combination of ordination and variation partitioning by constrained ordination we demonstrated a small but unique effect of canopy dominance type on the understorey vegetation, while a larger amount of compositional variation was shared with other factors. Our results accord with a scenario in which differences in primary environmental factors and humus form properties, the latter accentuated by the canopy dominants themselves, are the most important causes of higher species density in P. tremuloides stands than in P. mariana stands, as well as differences in species composition among the two canopy dominance types. Processes and time scales involved in the small but significant direct and indirect effects of the canopy dominant on understo‐ rey species composition are discussed.     

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。     

神农架川金丝猴栖息地森林群落的数量分类与排序

在神农架川金丝猴生境典型地段设立样方58块,根据样方资料对神农架川金丝猴栖息地的森林群落用组平均法分类和DCA排序.用组平均法将58块样地分为9个群系,依据《中国植被》的分类原则和系统将研究区植物群落划归为7个植被型.样地的DCA排序较好地揭示了该区森林群落的分布格局与环境梯度的关系;DCA第二轴明显地反映出森林群落的海拔梯度变化,沿DCA第二轴从右到左,海拔逐渐升高;第一轴表现了各植物群落或植物种所在环境的坡度、坡向,即水分和光照因素,沿第一轴从下到上,坡度渐缓、坡向渐向阳.其中海拔梯度是环境因子中对森林群落起决定性作用的因子.研究表明,巴山冷杉+糙皮桦-大齿槭+尾萼蔷薇-高原露珠草+星果草群系发育较好,高大树木占有较大的比例,是神农架川金丝猴最适宜栖息地.7个植被型物种丰富度指数在群落梯度上呈规律性波动.其中针叶林和针叶-阔叶混交林中,物种丰富度指数在群落梯度上的总体趋势表现为灌木层＞草木层＞乔木层;在常绿阔叶林和常绿-落叶阔叶混交林中,该趋势为灌木层＞乔木层＞草本层;在落叶阔叶林中,其丰富度指数的趋势为灌木层＞草本层和乔木层.不同植被类型川金丝猴食源植物种类在群落梯度上的变化趋势与物种丰富度指数相同,但地衣类植物作为川金丝猴冬季的重要食物只在针叶林和针叶-阔叶混交林中生长.本研究为制定栖息地保护计划,更好地保护神农架川金丝猴提供了科学理论依据.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Interactions with successional stage and nutrient status determines the life‐form‐specific effects of increased soil temperature on boreal forest floor vegetation

The boreal forest is one of the largest terrestrial biomes and plays a key role for the global carbon balance and climate. The forest floor vegetation has a strong influence on the carbon and nitrogen cycles of the forests and is sensitive to changes in temperature conditions and nutrient availability. Additionally, the effects of climate warming on forest floor vegetation have been suggested to be moderated by the tree layer. Data on the effects of soil warming on forest floor vegetation from the boreal forest are, however, very scarce. We studied the effects on the forest floor vegetation in a long‐term (18 years) soil warming and fertilization experiment in a Norway spruce stand in northern Sweden. During the first 9 years, warming favored early successional species such as grasses and forbs at the expense of dwarf shrubs and bryophytes in unfertilized stands, while the effects were smaller after fertilization. Hence, warming led to significant changes in species composition and an increase in species richness in the open canopy nutrient limited forest. After another 9 years of warming and increasing tree canopy closure, most of the initial effects had ceased, indicating an interaction between forest succession and warming. The only remaining effect of warming was on the abundance of bryophytes, which contrary to the initial phase was strongly favored by warming. We propose that the suggested moderating effects of the tree layer are specific to plant life‐form and conclude that the successional phase of the forest may have a considerable impact on the effects of climate change on forest floor vegetation and its feedback effects on the carbon and nitrogen cycles, and thus on the climate.     

Vegetation Structure,Composition, and Species Richness Across a 56‐year Chronosequence of Dry Tropical Forest on Providencia Island,Colombia1

We compared vegetation structure and species richness across a 56‐yr chronosequence of six replicated age classes of dry tropical forest on the island of Providencia, Colombia, in the Southwest Caribbean. Stand age classes were determined using sequential, orthorectified panchromatic aerial photos acquired between 1944 and 1996 and Landsat 7 ETM + satellite imagery from 2000. Along the chronosequence we established 59 plots of 2 × 50 m (0.01 ha) to document changes in species richness, basal area, tree height, stem density, and sprouting ability. All woody trees and shrubs >2.5 cm diameter at breast height (DBH) were censused and measured. Although woody species density reached a peak in stands from 32 to 56 yr old, rarefaction analysis showed that species richness increased linearly with stand age and was highest in stands 56 yr old or greater. Nonparametric, abundance‐based estimators of species richness also showed positive and linear associations with age. Basal area and mean tree height were positively associated with age since abandonment, while sprouting ability showed a negative relationship. Our results indicate rapid recovery of woody species richness and structural characteristics along this tropical dry forest chronosequence.     

Variation in terricolous bryophyte and macrolichen vegetation along primary gradients in Canadian boreal forests

Abstract. Compositional variation among 172 boreal forests stands, based upon the understorey terricolous bryophyte and macrolichen vegetation, is compared with that based upon vascular vegetation. Detrended correspondence analysis (DCA) of the two datasets yielded stand ordinations each indicating a single dominant upland-bottomland gradient. This was most clear among 138 stands in the cryptogam dataset. Canonical correspondence analysis (CCA) of the 138 stand subset yielded a pronounced first axis gradient from dry, nutrient poor pine dominated forest sites to moist, nutrient rich bottomland sites dominated by balsam poplar woodland. Individual species response curves, by the method of log-linear least squares regression, yielded three different respective patterns among the most abundant tree, understorey vascular and cryptogamic plant species. Whereas curves for the tree species were ‘Gaussian’ in shape and displaced at somewhat regular intervals along the gradient, the abundant understorey vascular plant species curves clustered in the mesic, mixed wood region two thirds of the way along. Curves for the abundant cryptogams were mostly linear in shape and absent from the mixed wood zone, showing concentrations at one end of the gradient or the other. Two interpretations of the understorey pattern are considered. One proposes competitive exclusion of cryptogams from the mesic mixed wood region by the vascular understorey plants. Another proposes that the cryptogams are adapted to the sharply contrasting ecosystems encountered at one end of the gradient or another but that none can effectively cope with the intermediate, mixed wood ecosystems. Evidence from the response curves favours this second interpretation.     

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.     

Use of Plant Strategy Ordination,DCA and ANOVA to elucidate relationships among habitats of Salix planifolia and Salix monticola

Abstract. Detrended Correspondence Analysis (DCA) was used on a data set of 112 species from 14 stands dominated by Salix planifolia and/or S. monticola. Environmental variables were determined which might be important to explain differences in stand vegetation composition, and willow stature (height), in willow carrs in the Rocky Mountain National Park. Correlation of environmental variables with DCA stand scores indicate that peat depth and soil redox potential are highly related to the primary DCA axis. Soils in stands of S. planifolia are more reduced in the early summer and have greater peat depths, than soils in stands of S. monticola. The second DCA axis is related to a water chemistry gradient of hydrogen ion, sulphate, and ammonium concentration. Four environmental variables selected by correlation analysis were regressed on stand scores from the first two DCA axes. Regression coefficients of August soil redox potential and soil water pH were significantly non-zero on the primary DCA axis. In addition to overall vegetation relationships elucidated by DCA, the relationships between willow stature and environmental variables were tested by simultaneous ANOVAs. ANOVA results and DCA ordinations indicate that S. planifolia and S. monticola plants achieve the greatest stature in the least reduced soils, and in shallow peat. Competitive, stress tolerant, and ruderal plant strategy theory applied to the vegetation data produced an arrangement of stands highly correlated to the DCA Axis 1 stand arrangement. These correlations indicate that the primary axis of vegetation structure can be extracted either by species composition, or species morphology/physiology relationships. Concurrent use of DCA and plant strategy theory is suggested as a tool for the prediction of community composition.