Vegetation of Sphagnum bogs in highly disturbed landscapes: relative influence of abiotic and anthropogenic factors

Question: Has the vegetation of Sphagnum bogs been affected by more than 200 years of human activities? Location: Bas‐Saint‐Laurent region, southeastern Québec, Canada. Methods: Data (species assemblages, abiotic and spatio‐historical variables) were collected in 16 bogs ranging from 2 to 189 ha, and incorporated in a geographical information system. Major gradients in vegetation composition were identified using DCA. CCA was used to relate vegetation gradients to abiotic and spatio‐historical variables. Results: A clear segregation of species assemblages was observed, from open and undisturbed bogs to forested and highly disturbed sites. Among abiotic factors, tree basal area, water table level and peat thickness had a significant influence on plant species composition. Among spatio‐historical factors, disturbance level, area loss and fire were the most influential factors. Variance partitioning between these groups of factors suggests that spatio‐historical factors had a major influence on peatlands, representing 22% of the variation observed in the plant species assemblages while abiotic factors represent only 17% of the variation. Conclusions: The results highlight the influence of agricultural and other anthropogenic activities on plant assemblages and suggest that even wetlands apparently resistant to disturbances, such as peatlands, can be severely affected by anthropogenic factors. Plant species assemblages of ombrotrophic peatlands of the Bas‐Saint‐Laurent region were, and still are, largely influenced by human activities.     

Influence of historical,geographical and environmental variables on understorey composition and richness in Danish forests

Question: What is the relative influence of forest continuity, environmental differences and geographical context on vegetation and species richness in ancient and recent forests? Location: Himmerland and Hornsherred in Denmark. Methods: Lists of forest species from deciduous forests were subjected to CCA with variation partitioning to quantify the relative amount of variation in species composition attributable to historical, present geographical and environmental variables. GLM was used to estimate the importance of the variables to species richness. Results: The importance of temporal forest continuity in one region was negligible but was considerable in the other. The variation in species composition explained by geographical, environmental and historical variables showed little overlap in both regions, particularly at the fine scale. Conclusions: This paper does not support the idea that differences in the flora between ancient and recent forests is mainly caused by environmental differences. Furthermore, species richness seemed unaffected by isolation and forest connectivity.     

Spider (Aranei) assemblages in bog-forest ecotone of the northern taiga subzone (Eastern Fennoscandia)

Soil-dwelling and ground spiders were studied in oligotrophic and mesotrophic bog-forest ecotones of the northern taiga (Eastern Fennoscandia) in 2005–2007. The number of spider species was greater in the forest as compared to the bog. The spider assemblages of the bogs and swampy forests was characterized by the prevalence of Lycosidae both in abundance and in species richness, whereas members of the family Linyphiidae prevailed in the forest. The species diversity of soil-dwelling spiders was lower in oligotrophic bogs than in mesotrophic ones. In all the bog-forest ecotones studied, the spider assemblages showed no edge effect (an increase in diversity and abundance), indicating a high degree of interpenetration of the spider complexes from adjacent ecosystems. Stenotopic spider species were not found in the ecotones studied either. The major ecological factors responsible for the specific features of spider assemblages in the biotopes studied are the temperature regime and soil humidity.     

Correlations among vegetation strata and environmental variables in subalpine spruce-fir forests,southeastern British Columbia

A set of forest vegetation-environment data consisting of 110 plots from subalpine spruce-fir forests in southeastern British Columbia was analyzed with multivariate methods. The aims were to describe and compare relationships among plots using species composition data from four strata of the forest (trees, shrubs, herbs, bryophytes), and to compare patterns in the separate strata with patterns in the environmental data. Weak but statistically significant correlations were detected among PCA axes from all four strata, with a tendency for correlations to be stronger between axes of adjacent strata. Relationships between stratal patterns and environmental variation were examined by correlating PCA axes with environmental variables separately, and jointly with canonical correlation analysis. Linear composites of the environmental data identified by the first canonical axis were similar for shrub, herb, and bryophyte strata implying similar responses to environmental variation in understorey strata; however, < 16% of the total variance was explained. No clear relationships emerged between tree stratum and environmental variation. Probable determinants of pattern in the tree stratum include historical events such as fire and winter snowpack, as well as possible interactions between species. Collectively, these events bear little resemblance to existing environmental conditions in the forest.Nomenclature used by BCMF follows Crum, Steere & Anderson (1973), Hale & Culberson (1970), and Stottler & Crandall-Stottler (1977) for non-vascular plants, and Taylor & MacBryde (1977) for vascular plants.The British Columbia Ministry of Forests provided the data on which this paper is based, as well as a grant to pay for the computer time used. We are grateful to G. Utzig for initially suggesting that a project of this nature be done, and to S. Phelps for helping with the retrieval of computer data files from tape. This paper has greatly benefited from discussions with P. Comeau and R. K. Scagel, and the comments of two anonymous reviewers.     

Local and regional trends in the ground vegetation of beech forests

We sampled moss and vascular forest vegetation in five ancient beech forests from northwest France, embracing in each a wide array of environmental conditions. Indirect (PCA) and direct (RDA) gradient analysis were used to discern local and regional ecological factors which explain the observed variation in species composition. Our results point to a global factor encompassing a large array of soil and light conditions, unravelled when local particularities of studied forests are singled out. The humus form, numerically expressed by the Humus Index, explains a large part of the observed variation in ground vegetation. Our study confirmed opposite trends in vascular and moss species richness according to humus condition. Ecological factors to which vascular and moss forest species respond at the regional level can be estimated directly in the field by visually inspecting humus forms and vegetation strata despite of the confounding influence of local factors.     

Assembly rules for New England ant assemblages

Community assembly rules specify patterns of species co-occurrence and morphology dictated by interspecific competition. We collected data on the occurrence of ground-foraging ant species in 22 ombrotrophic bogs and adjacent forest plots of New England to test two general assembly rules: reduced co-occurrence of species among communities, and even spacing of body sizes of species within communities. We used null models to generate random communities unstructured by competition and evaluated patterns at regional and local spatial scales. At the regional scale, species co-occurrence in forests, but not bogs, was less than expected by chance, whereas, at the local scale, co-occurrence in both habitats was not different from random. At the regional scale, spacing of body size distributions was random (in bogs) or aggregated (in forests). At the local scale, body size patterns were weakly segregated in bogs, but random or weakly aggregated in forests. In bogs, size ratio constancy was accompanied by greater generic diversity than expected. Although assembly rules were originally developed for vertebrate communities, they successfully explained some patterns in New England ant assemblages. However, the patterns were contingent on spatial scale, and were distinctly different for bog and forest communities, despite their close proximity and the presence of many shared species in both assemblages. The harsh physical conditions of bogs may act as a habitat filter that alters community assembly rules.     

Travelling to a former sea floor: colonization of forests by understorey plant species on land recently reclaimed from the sea

Questions: What are important forest characteristics determining colonization of forest patches by forest understorey species? Location: Planted forests on land recently reclaimed from the sea, the Netherlands. Methods: We related the distribution of forest specialist species in the understorey of 55 forests in Dutch IJsselmeer polders to the following forest characteristics: age, area, connectivity, distance to mainland (as a proxy for distance to seed source) and path density. We used species of the Fraxino‐Ulmetum association for the Netherlands as reference for species that could potentially occur in the study area. Results: Area and age of the surveyed forests explained a large part of the variation in overall species composition and species number of forest plant species. The importance of connectivity and distance to the mainland of forest habitats became apparent only at a more detailed level of dispersal groups and individual species. The importance of forest parameters differed between dispersal groups and also between individual species. After 60 years, 75% of the potential pool of wind‐dispersed species has reached the polders, whereas this was only 50% for species lacking specific adaptations to long‐distance dispersal. However, the average percentage of successful colonizing species present per forest was substantially lower, ranging from 15 to 37%. Conclusions: The data strongly suggest that the colonization process in polder forests is still in its initial phase, during which easily dispersed species dominate the vegetation. Colonization success of common species that lack adaptations to long‐distance dispersal is affected by spatial configuration of the forests, and most rare species that could potentially occur in these forests are still absent. Implications for conservation of rare species in fragmented landscapes are discussed.     

Development of ombrotrophic raised bogs in North-east Germany 17?years after the adoption of a protective program

The present study reports on the development of ombrotrophic raised bogs in NE Germany after the adoption of a protective program with a focus on the hydrological and floristic changes in two simultaneously restored cut-over bogs as a reference for all other bogs. At the moment, four fifths of the bogs area is dominated by woody vegetation forms and <5?% is characterized by typical raised bog vegetation. The trend of increasing areas with woody vegetation forms and decreasing areas with vegetation typical for ombrotrophic raised bogs could not be prevented despite increasing the portion of revitalized areas. The reason for the negative developments is founded in the different types of raised bogs in NE Germany, which differ in regard to the historical lagg area and the original grown up raised peat body. This as well as the kind and intensity of anthropogenic use have significant implications for the possibilities of revitalization and protection. Except for those bogs where no peat remnants remain, the revitalization of cut-over raised bogs in NE Germany can succeed if an almost closed hydrological protection zone (HPZ which is equal to the historical lagg area) is established so that the gradient of entire lateral water flow between the higher lying peat remnant and the lower lying surrounding area is reduced. The attainment of a lateral water flow gradient comparable to the original state and of a water-saturated peat body succeeds in turn in the precondition for growing Sphagnum species, which readjust the system??s self regulation. The comparison of two bogs with similar preconditions concerning location, climate, and anthropogenic transformation has shown that an incompletely closed HPZ is one important reason for the decreasing portion of intact raised bogs with their typical vegetation. The influence of climate change on current hydrological and floristic changes in NE German ombrotrophic raised bogs has less influence on the success of revitalization than the historic grown type of bog and the intensity of anthropogenic use.     

Types and Succession of Forest Bogs in Hingganling and Changbaishan

Climate and geomorphy arc the main factors of the formation of forest bogs in Hingganling and changbaishan. The bogs are both multiple in types and widespread in distribution. There exist not only the type of lower bog, but also that of middle bog and raised bog. They are chiefly distributed in the alluvial flat or valley and in the watershed. The succession process of forest bog is generalized into two kinds: (1) The formation of forest bogs as a result of the natural succession of forest vegetation under the influence of natural conditions. (2) The formation of forest bogs as a result of the bogginess in the vicinity of forestland.     

The role of landscape configuration in plant composition of floodplain forests across different physiographic areas

Questions: What is the relative importance of landscape variables compared to habitat quality variables in determining species composition in floodplain forests across different physiographic areas? How do species composition and species traits relate to effects of particular landscape variables? Do lowland and mountain areas differ in effects of landscape variables on species composition? Location: Southern Czech Republic. Methods: A total of 240 vegetation relevés of floodplain forests with measured site conditions were recorded across six physiographic areas. I tested how physiographic area, habitat quality variables and landscape variables such as current land‐cover categories, forest continuity, forest size and urbanization influenced plant species composition. I also compared how mountain and lowland areas differ in terms of the relative importance of these variables. To determine how landscape configuration affects the distribution of species traits, relationships of traits and species affinity with landscape variables were tested. Results: Among landscape variables, forest continuity, landscape forest cover and distance to nearest settlement altered the vegetation. These variables also influenced the distributions of species traits, i.e. life forms, life strategies, affinity to forest, dispersal modes, seed characteristics, flooding tolerance and Ellenberg indicator values for nitrogen, light, moisture and soil reaction. Nevertheless, physiographic area and habitat quality variables explained more variation in species composition. Landscape variables were more important in lowland areas. Forest continuity affected species composition only in lowlands. Conclusions: Although habitat quality and physiographic area explained more vegetation variability, landscape configuration was also a key factor influencing species composition and distribution of species traits. However, the results are dependent on forest geographical location, with lowland forests being more influenced by landscape variables compared to mountain forests.     

Overstorey Control of Understorey Species Composition in a Near-natural Temperate Broadleaved Forest in Denmark

Little is known about the importance of the forest overstorey relative to other factors in controlling the spatial variability in understorey species composition in near-natural temperate broadleaved forests. We addressed this question for the 19 ha ancient forest Suserup Skov (55°22′ N, 11°34′ E) in Denmark, one of the few old-growth temperate broadleaved forest remnants in north-western Europe, by inventorying understorey species composition and environmental conditions in 163 100 m² plots. We use unconstrained and constrained ordinations, variation partitioning, and Indicator Species Analysis to provide a quantitative assessment of the importance of the forest overstorey in controlling understorey species composition. Comparison of the gradients extracted by unconstrained and constrained ordinations showed that the main gradients in understorey species composition in our old-growth temperate broadleaved forest remnant are not caused by variability in the forest overstorey, but are related to topography and soil, edge effects, and unknown broad-scale factors. Nevertheless, overstorey-related variables uniquely accounted for 15% of the total explained variation in understorey species composition, with the pure overstorey-related (R_po), topography and soil (R_pt), edge and anthropogenic disturbance effects (R_pa), and spatial (R_ps) variation fractions being of equal magnitude. The forward variable selection showed that among the overstorey-related variables understorey light availability and to a lesser extent vertical forest structure were most important for understorey species composition. No unique influence of overstorey tree species identity could be documented. There were many indicator species for high understorey light levels and canopy gap centres, but none for medium or low light or closed canopy. Hence, no understorey species behaved as obligate shade plants. Our study shows that, the forest overstorey has a weak control of understorey species composition in near-natural broadleaved forest, in contrast to results from natural and managed forests comprising both conifer and broadleaved species. Nevertheless, >20% of the understorey species found were indicators of high light conditions or canopy openings. Hence, variability in canopy structure and understorey light availability is important for maintaining understorey species diversity.     

Forest snail diversity and its environmental predictors along a sharp climatic gradient in southern Siberia

Diversity patterns of forest snail assemblages have been studied mainly in Europe. Siberian snail faunas have different evolutionary history and colonization dynamics than European faunas, but studies of forest snail diversity are almost missing from Siberia. Therefore, we collected snails at 173 forest sites in the Russian Altai and adjacent areas, encompassing broad variation in climate and forest types. We found 51 species, with a maximum of 15 and an average of seven species per site. The main gradient in species composition was related to soil pH, a variable that also positively correlates with snail abundances. The second gradient was associated with climate characteristics of winter. We observed significant differences in both species richness and composition among six forest types defined based on vegetation classification. Hemiboreal continental forests were the poorest of these types but hosted several species characteristic of European full-glacial stages of the Late Pleistocene. A high snow cover in Temperate coniferous and mixed forests, protecting the soil from freezing, allowed the frost-sensitive large-bodied (>10 mm) species to inhabit this forest type. In contrast to most of the European snail assemblages studied so far we found that the factors responsible for the variation in species richness differed from those driving species composition. This may be attributed to the sharp climatic gradient and the presence of the cold-adapted species typical of the Pleistocene cold stages. We suggest that southern Siberian forests hosting these species can serve as modern analogues of full-glacial forests in periglacial Central and Eastern Europe.     

The Auchenorrhyncha fauna of peat bogs in the Austrian part of the Bohemian Forest (Insecta,Hemiptera)

The first overview on the Auchenorrhyncha fauna of peat bogs of the Austrian Bohemian Forest is presented. Seven oligotrophic peat bog sites were studied in 2011 by suction sampler (“G-Vac”) and 93 Auchenorrhyncha species (with 7465 adult specimens) were recorded. Eleven species (about 18 % of the individuals) are tyrphobiontic or tyrphophilous. The relative species abundance plot is not very steep; the six most abundant species represent 50 % of the individuals. The most common species is Conomelus anceps (17 % of the individuals). Compared to the whole Austrian Auchenorrhyncha fauna, the fauna of peat bogs comprises distinctly more univoltine species and more species hibernating in nymphal stage. Densities of adult Auchenorrhyncha in peat bogs are low in spring (about 10–60 individuals per m²) and high in July, with up to 180 (±50) individuals per m². Disturbed peat bogs have higher species numbers and higher Auchenorrhyncha densities in total, but lower numbers and densities in peat bog specialists.     

The Regeneration of a Highly Disturbed Ecosystem: A Mined Peatland in Southern Québec

We studied the natural regeneration of an ombrotrophic peatland (Cacouna bog) located in southern Québec that was disturbed by peat mining and other anthropogenic activities over a 200-year period. Using an extensive collection of historical documents, as well as dendrochronological data, we reconstructed the history of the peatland. We also sampled vegetation and environmental variables, and integrated the data in a geographic information system. More than 60% of the total area of the bog was mined between 1942 and 1975, and 98 km of ditches were dug to drain the site. The peatland lost 34% of its initial peat volume between 1946 and 1998. Although the bog was severely disturbed, the spontaneous revegetation of the site by vascular plants was successful (90%–100% cover). However, only 10% of the total mined area has been recolonized by Sphagnum species, mainly because drainage ditches are still operational and contribute to drying out the bog. Water table level, peat deposit thickness, and pH are abiotic factors strongly influencing the vegetation composition in the bog. Spatial and historical factors are also important components in this study since they explain, either alone or in interaction with abiotic factors, 44% of the variation of the species data. The intensity of mining activities and the pattern of abandonment of mined sectors strongly influenced abiotic factors, which in turn affected the revegetation process. Even if the Sphagnum cover of the bog is low, the rapid “recovery” of the vegetation cover in the peatland indicates that after the reestablishment of an appropriate hydrological regime, a highly disturbed peatland has a considerable potential for regeneration. Received 24 April 2001; accepted 30 October 2001.     

Diversity of plant assemblages in isolated depressional wetlands from Central-Western Europe

A closed depression is defined as a geomorphologic element where a sediment depository is encircled by hillslopes. Despite the fact that closed depressions are often the only stagnant water points in many European landscapes, few ecological researches on their plant assemblages have been done. The main goal of this study was to give first results of the environmental factors responsible for the vegetation composition, richness and rarity in the closed depressions of the Lorraine biogeographical district (Belgium, France and grand-duché de Luxembourg). We surveyed for plant presence 85 forest and 77 grassland closed depressions. For each site, wetland area, local environmental factors and regional connectivities registered. For each species, the Ellenberg values were compiled. To investigate the main source of variation in species composition and in species richness (including richness in rare species), Non-metric Multidimensional Scaling analyses and Generalized Linear Models were respectively used. Species pools in forest (forest and preforest species) and in grassland (bog plants, pioneers, helophytes) were quite different. In both landscapes, a gradient from plants typical of basic high-productive soils to plants typical of acid low-productive soils reflects a shared successional gradient. The accumulation of organic matter allowed the establishment of Sphagnum spp., which slowly acidified the soil and thus acted as ecosystem engineers for the arrival of bog plants. Moreover, the species composition was additionally driven by the plant light tolerance in forests and by the plant water requirements in grasslands. Mechanisms of species accumulation (increase in species richness) were different in forests and in grasslands: respectively related to the plant light tolerance and to the wetland area. At the regional level, the averaged soil productivity was negatively related to the richness in rare species. Indeed, perennial highly-competitive plants such as Glyceria spp., Iris pseudacorus or Urtica dioica impeded the establishment of smaller and rarer species. At the habitat level, isolated closed depressions (due to either low connectivity or low grazing pressure) have more habitat rare species, giving evidences of dispersal limitation in plant assemblages of closed depressions.     

Patterns in specific leaf area and the structure of a temperate heath community

Relationships between the distribution and specific leaf area (SLA: leaf area per unit dry mass) of six heath (Ericaceae) species were investigated along an environmental gradient between peat bogs and conifer forest in British Columbia, Canada. I asked whether patterns in SLA could help to identify the processes shaping plant distributional patterns. Specifically, I assessed whether (i) species’ distributions across the environmental gradient are correlated with SLA (ii) relationships between plant distributional patterns and SLA are similar among bogs with different shrub species (iii) intraspecific patterns in SLA parallel interspecific relationships between distributions and SLA, and (iv) intraspecific patterns are environmentally determined. Results showed that distributional patterns were often correlated with SLA; species with lower SLA were more abundant towards the centre of bogs, while species with higher SLA were more abundant in forest. Intraspecific patterns in SLA paralleled distributional patterns across the gradient; individuals located towards the centre of bogs had lower SLA than those growing in forest. A transplantation experiment showed that plants typically altered their SLA according to local environmental conditions. However, one bog showed no relationship between species’ distributions and SLA. This bog lacked the two species with lowest SLA, which typically occurred at the centre of other bogs. In their absence, species with higher SLA that typically occurred in forest increased in abundance towards the centre of the bog, where they obtained lower values of SLA. Therefore, while distributional patterns were often closely associated with SLA, plasticity in SLA was associated with increased breadth of species’ distributions across the gradient. Overall results indicate SLA may serve as a useful proxy for a range of life history traits to help elucidate the processes structuring plant communities.     

新疆天山森林公园土壤螨类群落多样性与环境因子的相关性

为探讨西北干旱区森林土壤螨类群落和环境因子的相互关系,于2014年对新疆天山森林公园七种不同生境进行土壤螨类群落调查与环境因子测定,并采用除趋势对应分析法(DCA)和冗余分析法(RDA)对土壤螨类群落结构和多样性特征及其与环境因子之间的关系进行相关分析。结果表明,共捕获土壤螨类成体标本24399只,隶属4目56科108属(包括9个中国新记录属),其中小甲螨属Oribatella为优势类群。方差分析表明,在7种不同生境之间土壤螨类群落多样性指标均存在显著差异(P<0.05),Shannon-Wiener多样性指数(H)依次为针叶林>苗圃林>阔叶林>灌木林>针阔混交林>草甸草原>林中草地。RDA分析结果表明,第一主轴和第二主轴分别解释了土壤螨类主要群落总变量的34.8%和27.3%,所有环境因子共解释了土壤螨类群落物种组成变异的82.1%。蒙特卡罗置换检验显示,十种环境因子与全部排序轴(F=7.355,P=0.002)均存在极显著的相关性。研究表明,海拔、土壤含水量和有机质含量对螨类群落结构和多样性的影响显著。     

Moorland pools as refugia for endangered species characteristic of raised bog gradients

In intact raised bog landscapes transitions from ombrotrophic into minerotrophic conditions occur. These gradients are lost from many bogs due to peat harvesting and drainage, and are difficult to restore. To determine which endangered species are characteristic of pristine raised bog gradients and their current status in degraded bogs, plants and macroinvertebrates were surveyed in Estonian intact raised bogs and Dutch degraded bog remnants. Dutch national distribution data were used to determine whether communities with these species occurred outside bog habitats. Water chemistry data were used to describe associated environmental conditions. Intact bog gradients were the preferred habitat for six plant species and fifteen macroinvertebrate species, all of which are endangered. In degraded bogs these species were scarce or not recorded. In intact bogs these species lived at sites where runoff from the bog massif came into contact with regional ground water resulting in a gradient in pH, alkalinity, Ca, Fe and ionic ratio. Analysis of Dutch national distribution data revealed aggregations of these endangered species in moorland pools. These pools contained water chemistry gradients similar to those found in pristine bogs, which occurred at sites were groundwater seepage and stream water came in contact. In the past, stream water has been used to increase pH and trophic status of moorland pools facilitating fisheries. Today, this practice offers a conservation strategy for the protection of endangered species for which no short-term alternatives are available.     

海南岛霸王岭热带天然林景观中木本植物功能型分布的影响因素

为了探讨在热带天然林景观中不同因素对木本植物(限于乔木和灌木)功能型分布的影响,在对海南岛霸王岭的热带天然林进行样方调查的基础上,运用数量分类方法对热带天然林的木本植物进行了功能型划分,并运用冗余度分析(Redundancy analysis, RDA)分别探讨了功能型出现与否、功能型物种丰富度和功能型木本植物多度3个矩阵与环境、空间和干扰因素之间的关系。运用偏冗余度分析对影响功能型分布变化的环境、干扰、空间及其交互作用等因素进行了定量分解。结果表明:利用木材密度和潜在最大高度两个指标划分出的9个功能型,能较好的区分功能型间对生境的适应响应差异;因子分解表明,纯环境、纯人为干扰和混合的环境-人为干扰是影响3个功能型矩阵分布变化的主要因素,纯空间和与空间位置相耦合的因素相对较低;在诸多因子中,干扰类型、地形因子、土壤类型、砂砾含量、土层厚度以及经纬度坐标是影响功能型分布变化的主导因子;RDA排序分析表明,一般来讲,硬木功能型多在生境条件好和远离人为干扰的立地高发生。软木功能型则更多地出现于生境条件恶劣和人为干扰频繁的立地。而中等硬度功能型则生态幅度较宽。但除了软木灌木功能型外,其它功能型物种丰富度和木本植物多度多在土层深厚和生境条件较好的立地较高。     

The shrubland vegetation in western Shewa,Ethiopia and its possible recovery

Shrubland vegetation and environmental data in western Shewa, Ethiopia have been analysed. Vegetation data include cover-abundance values of vascular plant species; environmental data comprise physical and chemical properties of the soil, altitude, slope, grazing and browsing pressure. The vegetation data were subj ected to hierarchical and non-hierarchical classification and ordination with correspondence analysis. The classification resulted in seven different vegetation types, ranging from grassland with scattered shrubs to degenerated forest. Ordination of the data and biplot analysis showed that the vegetation is influenced by anthropogenic factors and altitudinal variation. Sand content is related to a low level of anthropogenic influence whereas silt content is related to a high level. This is explained by historical events rather than by the present situation. Total nitrogen, organic carbon, altitude and slope are positively correlated and these variables are negatively related to anthropogenic influences. The shrubland vegetation may have expanded from lower altitudes and drier sites as forests gradually disappeared. The recovery of an economically more rewarding vegetation type may be achieved through pro viding alternative sources of fuel and construction and through prohibiting cultivation and grazing in the shrublands on the hillsides. Regeneration can be accelerated by actively introducing seedlings of tree species that do not need a heavy canopy cover for establishment and growth.