High-altitude vegetation of the Western Carpathians — a syntaxonomical review

This paper presents a survey of high-altitude plant communities which occur in the Western Carpathians with an enumeration of the characteristic, transgressive and differential species of the individual alliances, orders and classes. It describes the tall-herb vegetation (Mulgedio-Aconitetea) and communities of the subalpine shrubs (Betulo carpaticae-Alnetea viridis), the montane and alpine calcareous swards (Elyno-Seslerietea), the wind-exposed cryophilous swards on ridge edges with low snow cover (Carici rupestris-Kobresietea bellardii), the chionophilous communities of snow beds and snow fields (Salicetea herbaceae), the arctic-boreal dwarf-shrub heathlands (Loiseleurio-Vaccinietea), the alpine acidophilous grasslands (Caricetea curvulae) and the high-mountain mat-grass swards of the alliance Nardion strictae (Nardetea strictae).     

Response of the moss Racomitrium lanuginosum to changes in sheep grazing and snow‐lie due to a snow‐fence

Question: What are the responses of Racomitrium lanuginosum moss to altered snow‐lie and sheep use? Location: ACarexbigelowii‐Racomitrium lanuginosum heath on a Scottish montane plateau affected since 1986 by a fenced ski corridor. Methods: Permanent quadrats were set up along transects 45 m long perpendicular to the snow‐fence. Cover was assessed over a 12‐year period from 1990. Pellet‐group clearance counts provided data on sheep usage between 1990 and 1996. Snow‐lie was mapped in the springs of 1991–1996. Results: The snow‐fence created a gradient in sheep use and altered the duration of snow‐lie. At the start of monitoring Racomitrium cover was lower immediately adjacent to the fence, and after 12 years its cover was significantly reduced within 10 m of the fence. Further away from the fence Racomitrium cover was relatively stable. The loss of Racomitrium was correlated both with increased snow‐lie and heavier sheep usage. Grass cover increased near the fence and was related to sheep use. Dicranum fuscescens responded differently to Racomitrium, increasing significantly near the fence. Conclusions: We found that changes in snow‐lie and grazing pressure quickly brought about vegetation change in this montane ecosystem. Racomitrium was the most sensitive species to the changes in grazing and snow‐lie caused by the fence, having the biggest initial changes. Loss of Racomitrium permitted increases of species more resistant to grazing including Dicranum fuscescens and grasses.     

Occurrence and distribution of arbuscular mycorrhizal fungal species in three types of grassland community of the Tibetan Plateau

Arbuscular mycorrhizal (AM) fungal spore communities and distribution patterns were surveyed in montane scrub grassland, alpine steppe, and alpine meadow sites at altitudes ranging from 3,500 to 5,200 m a.s.l. on the Tibetan Plateau. Thirty-two representative soil samples were collected from the root zone of the dominant and common plant species in late May 2004. Twenty-three AM fungal species representing six genera (Acaulospora, Entrophospora, Glomus, Pacispora, Paraglomus, and Scutellospora) were detected and species richness varied from 5.3 ± 0.8 to 10.5 ± 2.5 per site. Some AM fungal species were restricted to one vegetation type and Glomus mosseae, Glomus intraradices, and Scutellospora calospora were detected in all three vegetation types. Glomus species were found to be the most frequent and abundant in all three vegetation types. Acaulospora occurred mostly in the alpine steppe and alpine meadow. Scutellospora occurred mostly in montane scrub grassland. At the species level, Glomus mosseae was dominant in the montane scrub, Acaulospora laevis and Pacispora scintillans were dominant in the alpine steppe, and Acaulospora laevis, Pacispora scintillans, and Glomus claroideum dominated the alpine meadow. It was evident from the distribution pattern of AM fungi in the different vegetation types that the abundance and diversity of AM fungal species were lowest in the montane scrub grassland than the other two plant communities. Climatic conditions, especially temperatures, and intensity of land use may be the most important factors influencing the AM fungal community.     

Plant communities and species distribution of low alpine Betula nana heaths in Northernmost Sweden

Summary Low alpine Betula nana heaths were studied with regard to plant communities, species distribution, and environmental conditions. It is clear from cluster analyses and PCA-ordinations that the distribution of winter snow is the most important factor for the composition of plant communities and the distribution of most plant populations. Under deep snow conditions soil instability is also important as a differentiating factor.The numerical treatment of data gives a phytosociological grouping in eoncordance with earlier opinions based on stand tabulations.The cluster analysis group the species into chionophobous, chionophilous, moderate snow bed species, and an additional group of wide snow depth amplitude. Only a few species remain unclassified. The ecological species groups are readily identified also in PCA-ordinations. They are independent of differences in species numbers and combinations.Nomenelature follows Lid (1963), Nyholm (1954–1969), Arnell (1956), and Dahl & Krog (1972).I am grateful to Dr. Eddy van der Maarel, University of Nijmegen, for critical comments on the manuseript, to Dr. Ewa Kvillner, University of Lund, and to my colleagues in Göteborg for generous help and encouragement throughout the study, which forms part of a Ph. D. thesis in the University of Göteborg. The work has been supported by grants from The Royal Swedish Academy of Sciences and from the Faculty of Natural Sciences, University of Göteborg.     

Vegetation of rock crevices of the montane and colline zones in the Pamir-Alai and Tian Shan Mts in Tajikistan (Middle Asia)

The paper presents the results of phytosociological research conducted on the vegetation of rock habitats in the montane and colline zones of the Pamir-Alai Mountains (Tajikistan). During field surveys done in 2010–2013, 122 phytosociological relevés were sampled. Plant species were recorded according to the Braun-Blanquet cover-abundance scale. The vegetation studies were carried out generally in the western Pamir-Alai and south-western Tian Shan Mts, especially in the Zeravshan, Hissar, Hazratishoh, Darvaz, Rushan, Turkestan, Kuramin and Vanch ranges. A synopsis of the rock communities of the montane and colline zones in Tajikistan is proposed. The collected material presents most of the variability among the plant communities of rock crevices and fissures on calcareous as well as acidic substrates in the montane and colline zones in Tajikistan. As a result of field research and numerical analyses, eight associations have been distinguished: Scutellarietum hissaricae, Scutellarietumschugnanicae, Scutellarietum zaprjagaevii, Scutellarietum baldshuanicae, Tylospermetumlignosae, Dionysietum involucratae A. Nowak, M. Nobis, S. Nowak & A. Nobis, 2014, Nanorrhinetum ramosissimi, Campanuletum albertii and the community of Scutellaria adenostegia within new alliance – Caricion koshevnikovii. The main factors determining the species composition of classified associations seem to be the geography of the distribution range, elevation above sea level, rock type and exposition.     

兰坪铅锌矿区植被恢复初期土壤种子库与地上植被关系的研究

尾矿废弃地是一种极端的生态系统,其植被恢复的研究将丰富传统的生态学理论。该研究通过野外植被调查与室内萌发实验相结合的方法,探讨了兰坪铅锌矿区植被恢复初期不同群落类型地上植被、土壤种子库及其相互关系。结果显示:（1）与对照群落（云南松林、高山栎灌丛）相比,尾矿区恢复期各群落（早熟禾人工草地、魁蒿群落、马桑灌丛）地上植被及土壤种子库的物种数、物种多样性均较低。（2）植被恢复时间较短的2个群落（魁蒿群落、人工草地）土壤种子库较地上植被物种多样性高。（3）尾矿区恢复期各群落地上植被及土壤种子库的优势种均主要由风播、种子繁殖的植物组成,菊科、禾本科占较大比例,这些植物在尾矿区植被恢复初期起重要作用。（4）尾矿区恢复期各群落土壤种子库与地上植被的物种相似性较高,各群落之间地上植被及土壤种子库的相似性则较低。研究表明,尾矿区恢复初期土壤种子库与地上植被紧密联系,群落改造方式、恢复时间对土壤种子库具有重要影响。     

The lichens of high ground in the English Lake District

Abstract: A general account of the lichens of high-level acid habitats in the Lake District is provided. This is followed by detailed studies of base-rich sites in the Helvellyn range, acid outcrops in Langdale, and three upland tarns, one of which is of outstanding importance. The lichen vegetation is richer than in Snowdonia or the Northern Pennines, but its diversity is considerably less than that of certain individual mountains in the Scottish Highlands. The montane element is interpreted as representing the last remnants of communities that have declined gradually through the Postglacial Period. Many species are extremely rare. Current restraints on the alpine lichen flora include the climate, the predominance of wet, acid rock and heavy sheep grazing. Rhizocarpon simillimum is reported as new to the British lichen flora.     

Numerical phytosociology of the subalpine belt of the Kazbegi region,Caucasus, USSR

A vegetational transect across a glacial valley was investigated. Structure and species composition, as well as influence of disturbance by biotic factors (grazing of sheep and cattle) and abiotic factors (topography, wind, water) are discussed.According to species composition and community structure, 4 main vegetation types were distinguished by comparing field data from phytosociological relevés with results of numerical treatments (3 methods of numerical classification and DCA ordination): 1) Chamaesciadium acaule-Polygonum viviparum, 2) Festuca cf. woronowii-Carex tristis, 3) Veratrum lobelianum-Taraxacum stevenii and 4) Rhododendron caucasicum-Vaccinium myrtillus communities.Species diversity (evenness, and number of species) was low in the ungrazed dwarf-shrub alpine communities, which were dominated by Rhododendron caucasicum, in comparison with the grazed meadow. The highest values of these characteristics were found in the transitions among the types.     

Annual variation in the distribution of summer snowdrifts in the Kosciuszko alpine area,Australia, and its effect on the composition and structure of alpine vegetation

Abstract Australian alpine ecosystems are expected to diminish in extent as global warming intensifies. Alpine vegetation patterns are influenced by the duration of snow cover including the presence of snowdrifts in summer, but there is little quantitative information on landscape‐scale relationships between vegetation patterns and the frequency of occurrence of persistent summer snowdrifts in the Australian alps. We mapped annual changes in summer snowdrifts in the Kosciuszko alpine region, Australia, from Landsat TM images and modelled the frequency of occurrence of persistent summer snowdrifts from long‐term records (1954–2003) of winter snow depth. We then compared vegetation composition and structure among four classes that differed in the frequency of occurrence of persistent summer snowdrifts. We found a curvilinear relationship between annual winter snow depth and the area occupied by persistent snowdrifts in the following summer (r² = 0.9756). Only 21 ha (0.42% of study area) was predicted to have supported summer snowdrifts in 80% of the past 50 years, while 440 ha supported persistent summer snow in 10% of years. Mean cover and species richness of vascular plants declined significantly, and species composition varied significantly, as the frequency of summer snow persistence increased. Cushion plants and rushes were most abundant where summer snowdrifts occurred most frequently, and shrubs, grasses and sedges were most abundant in areas that did not support snowdrifts in summer. The results demonstrate strong regional relationships between vegetation composition and structure and the frequency of occurrence of persistent summer snowdrifts. Reductions in winter snow depth due to global warming are expected to lead to substantial reductions in the extent of persistent summer snowdrifts. As a consequence, shrubs, grasses and sedges are predicted to expand at the expense of cushion plants and rushes, reducing landscape vegetation diversity. Fortunately, few vascular plant species (e.g. Ranunculus niphophilus) appear to be totally restricted to areas where summer snow occurs most frequently. The results from this study highlight potential indicator species that could be monitored to assess the effects of global warming on Australian alpine environments.     

Climate and land use change impacts on Mediterranean high-mountain vegetation in the Apennines since the 1950s

Background: High-mountain ecosystems are centres of plant diversity that are particularly sensitive to land-use and climate change.

Aims: We investigated the ecological trends associated with land use and climate change since the 1950s in different vegetation types in high-mountain habitats in the central Apennines.

Methods: We analysed temporal changes in: Pinus mugo scrub, calcareous subalpine grasslands and alpine scree vegetation, comparing historical and recent vegetation records from vegetation plots from two periods (1955–1980 and 1990–2014) for their ecological indicator values (Landolt temperature and nutrient indicators) and structural traits (growth forms) over time using generalised linear models (GLMs).

Results: We observed significant temporal differences in the ecology and structure of the analysed habitats. In the Pinus mugo scrub we detected a reduction of subalpine and herbaceous species and in calcareous alpine screes we observed an increment of the lower montane, montane and subalpine species and of dwarf shrubs. Conversely, subalpine grasslands were stable over time.

Conclusions: Ecological changes that have occurred in the Central Apennines, following changes in type and intensity of land use and recent warming are consistent with those observed in other European mountains, for which climate and land-use changes are claimed as the main driving forces.     

The consequences of multiple resource shifts on the productivity and composition of alpine tundra communities: inferences from a long-term snow and nutrient manipulation experiment

Background: Gradients in the amounts and duration of snowpack and resulting soil moisture gradients have been associated with different plant communities across alpine landscapes.

Aims: The extent to which snow additions could alter plant community structure, both alone and in combination with nitrogen (N) and phosphorus (P) additions, provided an empirical assessment of the strength of these variables on structuring the plant communities of the alpine tundra at Niwot Ridge, Colorado Front Range.

Methods: A long-term snow fence was used to study vegetation changes in responses to snowpack, both alone and in conjunction with nutrient amendments, in plots established in dry and moist meadow communities in the alpine belt. Species richness, diversity, evenness and dissimilarity were evaluated after 20 years of treatments.

Results: Snow additions, alone, reduced species richness and altered species composition in dry meadow plots, but not in moist meadow; more plant species were found in the snow-impacted areas than in nearby controls. Changes in plant community structure to N and N + P additions were influenced by snow additions. Above-ground plant productivity in plots not naturally affected by snow accumulation was not increased, and the positive responses of plant species to nutrient additions were reduced by snow addition. Plant species showed individualistic responses to changes in snow and nutrients, and indirect evidence suggested that competitive interactions mediated responses. A Permanova analysis demonstrated that community dissimilarity was affected by snow, N, and P additions, but with these responses differing by community type for snow and N. Snow influenced community patterns generated by N, and finally, the communities impacted by N + P were significantly different than those affected by the individual nutrients.

Conclusions: These results show that changes in snow cover over a 20-year interval produce measureable changes in community composition that concurrently influence and are influenced by soil nutrient availability. Dry meadow communities exhibit more sensitivity to increases in snow cover whereas moist meadow communities appear more sensitive to N enrichment. This study shows that the dynamics of multiple limiting resources influence both the productivity and composition of alpine plant communities, with, species, life form, and functional traits mediating these responses.     

Rumex aquaticus L. at Loch Lomondside

Summary

A new alliance of spring vegetation dominated by the leafy liverwort Anthelia julacea is described from the montane regions of Britain. It is distinguished floristically and ecologically from similar alpine late snow bed communities in Scotland and Scandinavia. The classificatory position of the stands of this vegetation type is discussed with reference to continental and British systems and it is concluded that the erection of a new alliance is justified on floristic grounds.     

Athalamia hyalina (Sommerf.) S.Hatt. in Scotland,new to the British Isles

Abstract

The liverwort Athalamia hyalina (Sommerf.) S.Hatt. and the family Cleveaceae are reported as new to the British Isles from the eastern Scottish Highlands. The Scottish plants are described and illustrated and the significance of the discovery is assessed. In Scotland, Athalamia hyalina occurs on thin soil on eroding limestone ledges in a montane habitat with a continental climate. It is in active growth in the winter months, producing spores in spring. Its range is restricted but it occurs within a protected area and is not threatened at present. The species is widespread in the northern hemisphere.     

The lichen vegetation associated with areas of late snow-lie in the Scottish Highlands

Abstract:The lichen vegetation associated with areas of late snow-lie in the Scottish Highlands is described and three lichen-dominated communities recognized; from the tops of large boulders, from small pebbles, and from damp soil. The first of these is exclusive to this habitat throughout the Highlands, whereas the other two are restricted to areas of late snow-lie in the Eastern Highlands but are of more widespread occurrence further west. The community from damp soil has affinities with the Solorinion croceae Klement, which occupies a similar ecological niche in Central Europe, but the two other communities are apparently undescribed. The importance of the eu-oceanic climate of the Western Highlands in determining the distribution of these communities is emphasized. The new combination Micarea cinerea f. tenuispora (D. Hawksw. & Poelt) Fryday is made for the anamorph of M. cinerea (Schaer.) Hedl.     

Is vegetation inside Carex sempervirens tussocks highly specific or an image of the surrounding vegetation?

Questions : How do species diversity, frequency and composition in tussocks differ from those in similar sized plots outside tussocks? Does the extent of the differences depend on community types or environmental conditions? Location : A sub‐alpine grassland in the Swiss National Park. Methods : In each of the two communities (short grass and tall graminoid) differing in species composition, grazing intensity and soil nutrient availability, relevés were made in 40 pairs of small circular plots, with one plot located inside a randomly selected Carex sempervirens tussock and the other outside. Results : We found 92 vascular species, of which 46 had a frequency higher than 5%. Species richness (S), pooled cover, Shannon's diversity (H) and cumulative species number (CS) were higher outside than inside the C. sempervirens tussocks, but evenness (J) was lower. S, H and CS differed more in the tall graminoid community than in the short grass community. However, dissimilarity between the paired relevés inside and outside tussocks did not differ between the two communities. Of the 46 most frequent species, 12 were statistically more and only one less frequent outside than inside the tussocks. Vegetation inside and outside tussocks could be clearly distinguished in the ordination space. Conclusion : Vegetation inside C. sempervirens tussocks is different from that in the surrounding area and represents an impoverished but homogenized version of the surrounding vegetation. Although tussocks of C. sempervirens were systematically avoided by grazers, there is little evidence that tussocks facilitate the species growing inside them.     

川西高山林线三种灌木凋落叶分解中的无脊椎动物多样性

以无脊椎动物为主体的土壤动物是影响凋落物分解的重要生物因素,对维持陆地生态系统物质循环和能量流动具有重要作用。高山林线交错带是高山植被垂直带谱中重要的过渡区域,拥有比相邻生态系统更高的生境复杂性和物种多样性。林线上温度波动和冻融循环频率显著高于针叶林,为了了解林线交错带上环境差异对凋落物分解过程中的土壤动物群落结构和多样性的影响,采用凋落物分解袋的方法,于高山生态系统的两个主要时期,即雪被末期和生长季末期,研究了林线主要代表性灌木——高山柳(Salix cupularis)、高山杜鹃(Rhododendron lapponicum)和红毛花楸(Sorbus rufopilosa)凋落叶分解的土壤动物多样性特征。结果表明:凋落物中的无脊椎动物群落多样性及个体、类群密度随物种、海拔梯度和季节而变化,且季节差异对无脊椎动物多样性的影响比物种和海拔梯度更显著。3个因子的交互作用不仅影响土壤动物群落多样性和均匀度,而且影响群落个体密度和类群密度。雪被末期,凋落物中的无脊椎动物多样性指数H、均匀度指数J及丰富度指数D以针叶林最高,优势度指数C以林线最高;生长季节末期的无脊椎动物类群密度和个体密度显著高于雪被末期。总体上,凋落物中的无脊椎动物群落丰富度以生长季末期最高,林线较针叶林丰富。这意味着,未来气候变暖情景下,灌丛密度增加,凋落物输入量增大,可能导致无脊椎动物多样性增加。     

Soil seed banks of woodland, heathland, grassland, mire and montane communities, Cairngorm Mountains, Scotland

The size and species composition of soil seed banks were assessed at 111 altitudinally diverse sites in the Cairngorm Mountains. Mean densities of germinable seeds varied from 83 000 m^–2 in Scots pine (Pinus sylvestris L.) woodland at 230–490 m to 200 m^–2 in moss (Racomitrium lanuginosum (Hedw.) Brid.) heath at 1000–1120 m. Seed banks were dominated by Calluna vulgaris (L.) Hull, not only wherever it was prominent in the vegetation, but also at some sites with less than 5% cover of parent plants in the ground vegetation. Many species conspicuous in the vegetation were under-represented in or absent from the seed bank and surface vegetation generally was more species rich than was the underlying seed bank, especially in high montane communities. Multiple regression was used to examine the relationship between the density of buried Calluna seeds and the abundance of parent plants in the vegetation, site altitude and the organic matter content of the soil. The model fitted to woodland communities accounted for 95% of the variation in seed density. The heathland model was less predictive but still explained 52% of the variation in seed bank size. In mire communities there was no relationship, collective or individual, between buried seed density and the measured environmental variables, possibly due to variations in the duration and frequency of waterlogging at these sites. The potential role of seed banks for initiating the recolonisation of disturbed ground is discussed. Densities of buried seeds at most Calluna-dominant sites were probably sufficient to generate successful recolonisation but the prospects for recovery were poor at other sites, particularly in graminaceous communities at 800 m or higher.     

Phytosociological and palynological studies of alpine steppe communities on the northern Tibetan Plateau,Qinghai Province,China

The dry cold climate in the highland of the eastern Kunlun Mountains in Qinghai Province, north‐western part of China supports the establishment of alpine grassland, consisting of Poaceae and Cyperaceae species and dwarf shrub communities dominated by Potentilla and Krascheninnikovia shrubs. The phytosociological vegetation studies carried out around Lake Heihai (36°00′N/93°26′E, 4446 m a.s.l.) reveals a complex pattern of high mountainous vegetation, including three major vegetation communities. A Polygonum sibiricum community occupies wet and slightly saline sites close to the Heihai lake shoreline and Kobresia robusta and Poa pachyantha communities are characteristic for the drier slopes. These communities differ in ecological conditions, exposure and several characteristic species that form local subunits. Main ecological factors influencing plant growth are moisture coming from glacial melt water and the composition and texture of the surface sediments. The atmospheric circulation (i.e. monsoonal wind system) and the different amount of insolation of the southern and northern slopes are of minor importance for the establishment of the different communities. The reflection of the vegetation composition in the modern pollen rain is generally poor, since the pollen spectrum is highly influenced by the local appearance of taxa. Though the vegetation unit of a alpine steppe can be detected. (© 2013 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

21st century climate change threatens mountain flora unequally across Europe

Continental‐scale assessments of 21st century global impacts of climate change on biodiversity have forecasted range contractions for many species. These coarse resolution studies are, however, of limited relevance for projecting risks to biodiversity in mountain systems, where pronounced microclimatic variation could allow species to persist locally, and are ill‐suited for assessment of species‐specific threat in particular regions. Here, we assess the impacts of climate change on 2632 plant species across all major European mountain ranges, using high‐resolution (ca. 100 m) species samples and data expressing four future climate scenarios. Projected habitat loss is greater for species distributed at higher elevations; depending on the climate scenario, we find 36–55% of alpine species, 31–51% of subalpine species and 19–46% of montane species lose more than 80% of their suitable habitat by 2070–2100. While our high‐resolution analyses consistently indicate marked levels of threat to cold‐adapted mountain florae across Europe, they also reveal unequal distribution of this threat across the various mountain ranges. Impacts on florae from regions projected to undergo increased warming accompanied by decreased precipitation, such as the Pyrenees and the Eastern Austrian Alps, will likely be greater than on florae in regions where the increase in temperature is less pronounced and rainfall increases concomitantly, such as in the Norwegian Scandes and the Scottish Highlands. This suggests that change in precipitation, not only warming, plays an important role in determining the potential impacts of climate change on vegetation.