引入草木樨对半干旱黄土高原区早期植物群落演替和土壤养分的影响

植被演替早期的优势物种是影响演替进程的重要因素。以半干旱黄土高原地区退化耕地植被的自然恢复为对照,于植被恢复的早期引入两年生豆科植物草木樨(Melilotus officinalis L.),在种植密度为11.3kg.hm-2条件下,考察了6年内引入草木樨对退化耕地植被恢复过程中植物群落演替进程的影响。结果表明,与自然恢复植被相比,草木樨的引入不仅可以提高地表植被覆盖度和加速植物群落演替进程,促使一些群落演替后期物种如冷蒿(Artemisia frigida)、阿尔泰狗哇花(Heteropappus altaicus)和短花针茅(Stipa breviflora)等在植被恢复的第3年出现,同时有利于提高植被生产力和稳定性。同自然恢复相比,试验期间引入草木樨后的植被地上平均生物量提高67.90%。同时,植被恢复早期生物多样性与土壤全氮含量呈显著正相关,而草木樨的引入有利于提高土壤全氮含量;在植被恢复初期(2003～2005)引入草木樨的土壤全氮平均含量比自然恢复高7.32%;试验期间土壤速效磷含量与物种丰富度呈显著负相关。因此,在半干旱区黄土高原退化耕地植被恢复早期阶段,两年生豆科植物草木樨的引入和磷肥的施用对于加快植被恢复有着积极作用。     

乌兰布和沙漠东北缘三种豆科绿肥植物生物量和养分含量及其对土壤肥力的影响

土壤肥力是限制乌兰布和沙漠东北缘沿黄灌区建立高产绿洲的主要因子之一。在内蒙古自治区磴口县试验地种植3种绿肥植物,比较它们改良土壤的效果。结果表明,1～2年生紫花苜蓿的总生物量干重为最高,而沙打旺则因其抗风沙能力强,地上部分生长较其它两种植物为好,由于植株中水分含量较高,其生物量的鲜重在三者中是最高的。2年生绿肥植物总生物量干重和养分含量的测定结果则显示,全氮和全磷含量的变化趋势为沙打旺〉紫花苜蓿〉草木樨,而全钾含量则为紫花苜蓿〉草木樨〉沙打旺;从3种植物对土壤改良效果来看,改良作用为沙打旺〉紫花苜蓿〉草木樨。与两种常用的绿肥作物相比,沙打旺因其对风沙较强的抗性而具有较高的改良土壤能力,因此在同类型的沙漠绿洲开发初期可以作为改良土壤的首选植物。     

Secondary Succession and Natural Habitat Restoration in Abandoned Rice Fields of Central Korea

Floristic composition and soil characteristics (moisture, pH, nutrient contents) in abandoned upland rice paddies of different ages were analyzed to clarify the regenerative aspects of succession as a tool for habitat restoration. The study sites represented five seral stages: newly abandoned paddy fields; successional paddy fields abandoned for 3, 7, and 10 years; and a 50‐year‐old Alnus japonica forest. A vegetation sere was apparent in changes of dominant plant species in the order Alopecurus aequalis var. amurensis (annual grass), Aneilema keisak (annual forb), Juncus effusus var. decipiens (rush), Salix koriyanagi (willow), and Alnus japonica (alder) communities. These temporal stages resemble the spatial zonation of vegetation in local riparian floodplain ecosystems, indicating a hydrosere, with soil moisture decreasing over time. Age distributions and life forms of the dominant plant species support a “tolerance” model of secondary succession, in which the established species persist into later successional stages. Persistence of earlier colonizers led to a net cumulative increase in species richness and a more even distribution of species cover with increasing field age. Between 10 and 50 years, vegetation stabilizes as an alder community. Soil moisture content decreased steadily with paddy field age after an initial rise immediately after their abandonment, whereas pools of organic matter, N, P, K, Ca, and Mg, increased with field age. The pace and direction of recovery of native vegetation and natural soil properties in these abandoned rice paddies resembled classic old field succession, a form of secondary succession that often serves as a template for guiding restoration efforts. Active intervention, in particular dismantling artificial levees, could accelerate the recovery process, but natural habitat recovery generally appears sufficiently robust to achieve “passive” restoration of this rare community without intervention.     

Temporal changes in species diversity and composition in abandoned fields in a <Emphasis Type="Italic">trans</Emphasis>-Himalayan landscape,Nepal

Secondary succession is an increasing phenomenon due to global changes in agriculture policies and practices. The empirical findings are biased towards the temperate zone. Abandonment of agriculture fields is less frequent in the subtropical and tropical zones where agriculture areas are, in general, expanding. But there are exceptions; a rapid rate of abandonment of agricultural fields have taken place in the arid trans-Himalayan region, due to today’s globalization of economy. We analysed agriculture fields that were abandoned between 1950 and 2003 in a large u-valley in central Nepal (3400 m a.s.l.). The potential forest vegetation is dominated by Pinus wallichina and shrubs of junipers and cotoneaster species. We tested the intermediate richness hypothesis in relation to vegetation cover, soil development and whether old-field succession is convergent or divergent with species data from 242 1 m² plots in 5 age-classes. The main species compositional turnover expressed by Detrended Correspondence Analyses (DCA) correlated, as expected, with time after abandonment. Fields that were abandoned a long time ago are closer to forest at the periphery of the agricultural landscape. Moisture of the soil significantly increased with age of abandonment, but total vegetation cover and pH were negatively related to age. Beta diversity expressed in DCA SD-units showed an increasing trend with age of abandonment, supporting the divergence pattern in old-field succession. The reason why the succession is not converging may be due to browsing by domestic animals that prevent a closed canopy of pines and juniper to develop. There was a significant hump-shaped pattern in species richness along the temporal gradient, which agrees with the intermediate species-richness hypothesis. There was a rapid increase in species richness in plots close to the villages that were used for haymaking which increased the seed input significantly.     

Long-term Effects of <Emphasis Type="Italic">Lupinus lepidus</Emphasis> on Vegetation Dynamics at Mount St. Helens

The nitrogen-fixing legume Lupinus lepidus is the most abundant herb on new volcanic surfaces at Mount St. Helen. We compared vegetation structure in 30 Lupinus colonies in three age classes (old, mature, or young based on known years of their establishment) to adjacent sites that were sparsely populated by Lupinus. Our goals were to determine if the age of colonies affected either species composition or vegetation structure and if colonies altered vegetation structure of a site. Species richness increased with site age and colonies had significantly greater richness. Mature colonies had lower cover and frequency of other species, while old colonies had significantly higher cover of other species, particularly mosses. Frequencies in colonies were higher than in adjacent sites. Diversity measures were highest in old colonies and least in mature ones. ANOVA showed that density effects of Lupinus were stronger than age effects on richness, cover, frequency, dominance, and diversity. The floristic similarity between colonies and adjacent sites declined with colony age, and sites without Lupinus became heterogeneous with age. Detrended correspondence analysis demonstrated floristic change as colonies aged and that the differences between colonies and sparse sites increased with age. Regressions of Lupinus frequency with measures of structure showed weak second order relationships that suggested that low densities of Lupinus inhibited other species, while higher densities promoted abundance and diversity. Together these results suggest that Lupinus is a key element that alters floristic successional trajectories and accelerates succession. However, its effects are complicated by the wide ranges of Lupinus density, the sequence of invasion, and by herbivores.     

Management impacts on vegetation dynamics of hyper‐eutrophicated fields at Berlin,Germany

Abstract. This paper reports on vegetation dynamics on terrestrial, temperate grassland sites at the upper range of the productivity scale, i.e. on abandoned sewage fields (fields once used for waste water disposal) at Berlin‐Blankenfelde, Germany. I studied regeneration and the influence of different management practices (removal of top soil and mowing in late summer). Changes in species composition and cover were followed on permanent plots of 2m × 2m size through five years of vegetation development. At the outset of the experiment the abandoned fields were dominated by dense Urtica dioica /Elymus repens stands. Species richness was 7 species/ 4m², and it remained low on unmanaged plots during the time of observation (7.6 species/plot in year 5). Removal of 20 cm of top soil caused a severe decline of Urtica and a large increase in species richness (21 species in year 1 after disturbance). Mowing was slightly higher compared with unmown plots on both initially excavated and unexcavated plots. Total cover was always near 100 % (except immediately after top soil removal). Colonization of bare soil was very rapid and in late summer of the first year after disturbance cover already increased towards 100%. On all plots the vegetation was mostly dominated by perennial herbs and grasses. Winter season gaps are occupied by Galium aparine, a large‐seeded annual scrambling climber. Monocarpic perennials behaved as winter annuals in most cases. Woody species were inhibited by dense above‐ground biomass and litter cover. The paper questions whether succession on abandoned sewage fields may proceed towards a woodland stage and advises how vegetation of such hyper‐eutrophicated sites can be managed towards higher diversity.     

Efficient nitrogen economy: Key to the success ofPolygonum bistorta in an abandoned mountain meadow

The dominance ofPolygonum bistorta in abandoned mountain meadows was studied. Successional and seasonal changes in biomass, nitrogen concentration and content were studied at the community level. During the successionP. bistorta increases its biomass both in absolute and mainly in relative values compared to the surrounding species. Nitrogen stored in its rhizomes seems to be quickly translocated into above-ground parts at the beginning of the growing season. High amounts of nitrogen allow rapid growth of both vegetative and generative plant parts. The success ofP. bistorta is probably the result of efficient use of stored nitrogen, suppression of surrounding vegetation by above-ground cover of leaves and by decrease in the level of soil nitrogen, and the effect of a large amount of litter. Management consisting in the combination of mowing and fertilization is proposed.     

Dynamics of vesicular-arbuscular mycorrhizae during old field succession

Summary The species composition of vesicular-arbuscular mycorrhizal (VAM) fungal communities changed during secondary succession of abandoned fields based on a field to forest chronosequence. Twenty-five VAM fungal species were identified. Seven species were clearly early successional and five species were clearly late successional. The total number of VAM fungal species did not increase with successional time, but diversity as measured by the Shannon-Wiener index tended to increase, primarily because the community became more even as a single species, Glomus aggregatum, became less dominant in the older sites. Diversity of the VAM fungal community was positively correlated with soil C and N. The density of VAM fungi, as measured by infectivity and total spore count, first increased with time since abandonment and then decreased in the late successional forest sites. Within 12 abandoned fields, VAM fungal density increased with increasing soil pH, H₂O soluble soil C, and root biomass, but was inversely related to extractable soil P and percent cover of non-host plant species. The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.     

Early Plant Succession in Abandoned Pastures in Ecuador1

We compared early plant succession in four abandoned pastures of differing age since abandonment and a nearby secondary forest site in northwestern Ecuador. Two “Open” pastures had no tree canopy covering, and two “Guava” pastures had a well-developed canopy cover of Psidium guajava. No site had been seeded with pasture grasses. All pastures were compared in a chronological sequence; two were monitored for 18 months. Species richness was consistently higher in Guava sites than in Open sites and it continued to increase over time, whereas it remained static in Open sites. Species richness was highest in secondary forest. Recruitment of tree saplings in Guava sites was lower than in secondary forest; however, it was nearly absent in Open sites. The seed bank contained predominantly herbaceous species at all sites, and was highly dissimilar to aboveground vegetation. Dominance-diversity curves for Guava sites showed a more equitable distribution of species that increased over time. In contrast, dominance-diversity curves for Open sites were static and indicated dominance by a few aggressive species. Soil characteristics among sites were variable; however, a principal components analysis on soils isolated the older Open site from all others. The older Open site had the lowest species richness and was dominated by Baccharis trinervis, an aggressive shrub species. The site appears to be in a state of arrested succession and some form of restorative intervention may be necessary to initiate succession toward a forested condition. Succession in Guava sites appears headed toward secondary forest, whereas it does not in Open sites.     

Vegetation dynamics of Mediterranean shrublands in former cultural landscape at Grazalema Mountains,South Spain

Plant community dynamics in Mediterranean basin ecosystems are mainly driven by an alternation of episodes of human intervention and land abandonment. As a result, a mosaic of plant communities has evolved following different stages of degradation and regeneration. Some authors has relate secondary succession to abandoned culture lands and regeneration to natural systems with abandonment of livestock or forestry exploitation. In this paper, the dynamics of shrublands in mid-mountain areas in the South of Spain after disturbance and land abandonment has been studied. The plant cover and 13 environmental variables of 137 selected sites on the Grazalema mountains was analysed to determine the vegetation pattern in relation to environmental factors and the succession types, either regenerative or secondary succession. The results show that today the Grazalema mountains have a heterogeneous vegetation pattern. Besides physical factors such as altitude or soil , human disturbance has modulated current vegetation patterns and dynamics. Two main types of vegetation dynamics can be distinguished in the study area. In areas affected by cutting, regeneration results in rich and dense shrub land, with resprouters as dominant species. In areas affected by recurrent wildfires or agriculture, secondary succession became dominant, resulting in less diverse shrubland, due to the dominance of seeders and decrease in resprouter species richness and cover.     

黄土高原地区不同草地退耕模式水分利用效率的比较

研究了半干旱黄土高原区不同退耕模式下植被恢复的比较,对3种不同人工豆科牧草多年生紫花苜蓿alfalfa(Medicago sativa)、多年生沙打旺erect milkvetch(Astragalus adsurgens)、2年生草木樨sweetclover(Melilotus officinalis))和一种自然撂荒(fallow)进行了实地种植比较。通过3a研究发现:紫花苜蓿是耗水最严重的牧草,水分利用效率仅高于撂荒;沙打旺具有最高的地上总生物量和水分利用效率。紫花苜蓿和沙打旺地块中杂草生物量逐年降低,物种数量最低且没有增加。2年生草木樨地物种数和地上生物量逐年增高,草木樨对暴雨的入渗贮存能力最大,显著高于自然撂荒。草木樨结束生活史后第1年地上总生物量(和撂荒一样全为杂草)是撂荒地的两倍,且略高于紫花苜蓿的地上总生物量,同时物种数量也和物种数目最多的撂荒地没有显著差异(p<0.05)。草木樨显著降低了10～40cm土壤剖面的容重,草木樨结束生活史后残留根系有助于深层土壤水分恢复,水分状况恢复良好且优于紫花苜蓿和沙打旺。可见短期的人工干扰下两年生草木樨的种植有利于促进自然植被的恢复,优于自然撂荒和其他牧草种植的方式,容易推广且实际可行。     

Plant distribution patterns during early succession on an artificial protosoil

Abstract. Plant succession has been studied since 1977 in permanent plots on a calcareous protosoil in the northern outskirts of Göttingen (Central Germany). At the beginning the substrate was extremely poor in organic carbon and nitrogen. On 128 plots of 100 m², the temporal and spatial distribution of the most conspicuous plant species showed remarkable changes and correlations to soil site factors. While presenting data on Melilotus alba, Medicago lupulina, Agropyron repens, Arrhenatherum elatius, Dactylis glomerata, and Picris hieracioides the mechanisms of successional changes on nutrient-poor soils are discussed. It is demonstrated that a high cover of legumes favours the accumulation of soil nitrogen. Grassland species were more abundant in areas with low humus content whereas ruderal plant species preferred those parts of the area which contained more humus in the substrate. Life-form, colonization and population strategies play an important role in the described successional processes, in addition to resource competition. Finally, some methodological conclusions for permanent plot studies are presented.     

Assigning plant communities to a successional phase: Time trends in abandoned limestone quarries

We investigated spontaneous vegetation succession and the relationship between time and vegetation patterns in several abandoned quarries of the Botticino extraction basin (Lombardy, Italy) and then assigned plant assemblages to a predetermined theoretical successional phase using an original procedure. To recognise and validate the gradient due to time, an ordination approach of vegetation plots linked to constant variables and time since last mining Canonical Correspondence Analysis was conducted first. Then, to determine the durations of the vegetation succession phases and trends between the colonisers and late successional species, we used an original six-step procedure based primarily on the regression curve of the percent relative abundance of life forms (RALFs) over time. The results demonstrated that time is the primary factor that significantly affects life form turnover during succession. Vegetation establishment and development in the “pioneer phase” (0–6 years) were affected by abiotic filters, which determined the dominance of a few ruderal and annual/alien species, mostly therophytes. The successive phases were characterised by an increasing presence of perennial species (mostly phanerophytes) with a consequent influence of biotic filters. The RALF procedure may be applied to other environments to investigate the time trends of plant communities during successions.     

Influence of a non-native invasive tree on primary succession at Mt. Koma, Hokkaido, Japan

The slopes of Mt. Koma in Japan are undergoing primary succession as a result of a 1929 eruption. Understory vegetation below a non-native invasive tree species, Larix kaempferi, a native tree, Betula ermanii, and in the open were compared to determine if the non-native tree species was influencing species composition. Larix canopies are significantly larger than Betula canopies. Vegetation under Larix canopies had significantly greater richness and diversity than vegetation in the open, vegetation under Betula was intermediate but was significantly greater than the open in diversity. Vegetation cover was highest under Betula and significantly lower in the open. Larix canopy size was positively correlated with size and number of Salix reinii shrubs. Betula canopy size was positively correlated with size but not with number of Salix reinii shrubs. Species assemblages in the three sites are slightly different as shown by DCA. Due to the limited species pool on Mt. Koma the greatest possible extent of differences between the three microsites is not large. At this point Larix certainly appears to be accelerating succession for the non-tree species. If Larix persists on the slopes then succession would be permanently deflected towards a Larix forest. This would be a case of succession being deflected towards dominance by the introduced species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Seed germination and seedling growth of five desert plants and their relevance to vegetation restoration

Due to significant decreases in precipitation in northern China, knowledge of the response of seed germination and plant growth characteristics to key limiting factors is essential for vegetation restoration. We examined seed germination under different temperatures and water potentials, and we examined seedling growth under different amounts of water supply. Experiments were carried out in automatic temperature‐, humidity‐, and light‐controlled growth chambers. Under low water potentials, the final germination percentages of four herbaceous species were high, while seed germination of the shrub species Caragana microphylla was significantly inhibited. Under the different water supply amounts, seedlings of Agropyron cristatum allocated more biomass to the root and had a higher growth rate than those of Elymus dahuricus and C. microphylla. In light of these results and drier environmental conditions (annual mean precipitation is 366 mm, which falling mainly between June and August), potential selections for revegetation of different landscapes include the following: A. cristatum for shifting sand dunes, the establishment of the pioneer species Agriophyllum squarrosum, C. microphylla for semifixed sand dunes, E. dahuricus for fixed sand dunes, and Melilotus suaveolens and Medicago sativa for cultivation.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.     

Limits to convergence of vegetation during early primary succession

Questions: Primary succession, measured by changes in species composition, is slow, usually forcing a chronose‐quence approach. A unique data set is used to explore spatial and temporal changes in vegetation structure after a 1980 volcanic eruption. On the basis of data from a transect of 20 permanent plots with an altitudinal range of 250 m sampled through 2005, two questions are asked: Do changes along the transect recapitulate succession? Do plots converge to similar composition over time? Location: A ridge between 1218 and 1468 m on Mount St. Helens, Washington, USA. Methods: Repeat sampling of plots for species cover along a 1‐km transect. Floristic changes were characterized by techniques including DCA, clustering and similarity. Results: Species richness and cover increased with time at rates that decreased with increasing elevation. The establishment of Lupinus lepidus accelerated the rate of succession and may control its trajectory. Diversity (H) at first increased with richness, then declined as dominance hierarchies developed. Primary succession was characterized by overlapping phases of species assembly (richness), vegetation maturation (diversity peaks, cover expands) and inhibition (diversity declines). Each plot passed through several community classes, but by 2005, only four classes persisted. Succession trajectories (measured by DCA) became shorter with elevation. Similarity between groups of plots defined by their classification in 2005 did not increase with time. Similarity within plot groups converged slightly at the lower elevations. Despite similarities between temporal and spatial trends in composition, trajectories of higher plots do not recapitulate those of lower plots, apparently because Lupinus was not an early colonist. Any vegetation convergence has been limited to plots that are in close proximity.     

Root development in simple and complex tropical successional ecosystems

Fine and coarse root mass and fine root surface area were studied during 5 yr following the felling and burning of a tropical forest near Turrialba, Costa Rica. Five experimental ecosystems were established: 1) natural successional vegetation, 2) successional vegetation enriched by seed applications, 3) imitation of succession (built by substituting investigator-selected species for natural colonizers), 4) monocultures (two maize crops followed by cassava andCordia alliodora), and 5) a bare plot. Fine roots grew rapidly in all treatments during the first 15 wk, at which time there were 75 gm⁻² in the monoculture and 140 gm⁻² in the enriched and natural successions. Subsequent growth was slower, and fine-root mass decreased during the first dry season. After 5 yr coarse root mass to a depth of 85 cm was about 800, 1370, and 1530 gm⁻² in the succession, enriched succession and imitation of succession, respectively. At the final harvest, the 3.5 yr-oldC. alliodora plantation had 1000 g m⁻² of coarse-root biomass. Roots <1 mm in diameter were concentrated in the upper 5 cm of soil and accounted for most fine-root surface area. Total fine-root surface area was greatest in the enriched successional vegetation and usually lowest in the monoculture.     

半干旱黄土高原苜蓿草地撂荒过程土壤水分变化特征

土壤水分是黄土高原植被恢复和生态建设的主要限制因子,明确土壤水分随植被演替的变化规律是阐明黄土高原植被与水分相互作用机制的重要基础。以半干旱黄土高原小流域苜蓿草地撂荒过程为研究对象,通过对2016-2018年生长季苜蓿群落、苜蓿+赖草群落、赖草群落和长芒草群落四种草地群落0-1.8 m土壤水分进行动态监测以及0-5 m深度土壤水分测定,分析不同演替阶段苜蓿草地土壤水分的动态特征,探讨土壤水分对苜蓿草地撂荒过程的响应。结果表明：（1）在苜蓿草地撂荒演替过程中,土壤水分随群落恢复时间的延长呈先增加后降低的变化,降水的年际动态显著影响不同演替群落的土壤水分响应;（2）0-0.4 m土壤水分主要受降水影响,使得各草地群落在这一层次没有显著差异（P>0.05）,而1 m以下的土壤水分含量则主要受植被类型的影响,各草地群落之间存在显著差异（P<0.05）;（3）0-5 m深层土壤水分随群落的演替,1 m以下各土层土壤水分含量逐渐增加,表明撂荒过程中使土壤水分得到了一定程度的恢复。研究结果揭示了苜蓿草地撂荒过程土壤水分的变化规律,可为黄土高原生态恢复提供理论依据。     

Succession secondaire et perte de diversité végétale après réduction du broutage dans un paturage boisé des Alpes centrales suisses

Freléchoux F., Meisser M. and Gillet F. 2007. Secondary succession and loss in plant diversity following a grazing decrease in a wooded pasture of the central Swiss Alps. Bot. Helv. 117: 37 – 56. Reduced cattle grazing pressure in the Alps has caused the reforestation of many subalpine pastures during the last decades. To understand the dynamics of natural reforestation and to evaluate how this change affects plant species diversity, we described the vegetation of a wooded pasture in the central Swiss Alps (Sembrancher, Valais) using the integrated synusial method. Based on stratified vegetation relevés in 27 plots,we defined 11 community types at the synusial level (two tree-layer, five shrub-layer, and four herb-layer synusiae), and four community types at the phytocoenosis level (pasture, tall forbs and scrub, wooded pasture and forest). The spatial distribution of these four phytocoenoses suggests that they represent successional stages after abandonment, and that the pathway of vegetation succession depends on the aspect. We suppose that on northern oriented, cool and shady locations, abandoned pastures first develop towards tall-forb meadows and scrub with Alnus viridis, and then to a preforested stage with Picea abies and Larix decidua. In contrast, on western oriented, warm and sunny location, Larix decidua (mainly) and Picea abies directly colonize the abandoned pastures, but further succession finally leads to the same pre-forested stage as on northern slopes. Plant species richness was highest in open areas and decreased by 25% as tree cover increased from 6% to 65%. According to our successional model, plant species diversity is lost more rapidly on northern slopes (with species-poor green alder scrub) than on western slopes (with species-rich young larch forests), suggesting that northern slopes most urgently need an appropriate grazing management. Manuscrit accepté le 28 mars 2007