The role of historical processes in determining tree species richness in the forests of Western Caucasus

To estimate the role of history in variation of tree species richness in the forests of the Western Caucasus we analyzed correlation between their local richness (S--the mean number of species per 300 m2) and size of actual species pool (N--the number of species per 1 ha). If compared communities are differently distant from the point of evolutionary equilibrium one should expect a significant variation in correlation between S and N (determined with the greater sensitivity of N than S in respect of historical factors). The lower value of N/S corresponds to less saturated level of historically determined species richness. A mean N/S ratio in Japana temperate broadleaved forests (Masaki et al., 1999) provided the basis for analysis. The present tree species richness of the forest communities in the 1 ha plots seem essentially determined by the historical processes. The mountain forest communities of Western Caucasus are characterized on the average with lower saturation level of the actual tree species pool in comparison with the Japan temperate broad-leaved forest communities. On the Western Caucasus the middle mountain beech and coniferous-broadleaved communities (400-1600 m a.s.l.) are characterized with the higher saturation level of the actual species pool in comparison with communities located lower and higher. These results confirm published historical reconstructions, according to which the middle mountain forest communities in the Western Caucasus are older than forests located higher or lower. Present low mountain forests of the southern (to Black Sea) and the northern macroslopes of the Western Caucasus are characterized with similar saturation level of the actual species pool. These data agree with the assumption of Dolukhanov (1980) that low mountain zone of the southern macroslope was not a refuge for tree species in Pleistocene.     

Relationship between Degree of Dominance and Species Richness in Grass Communities with Different Productivities

The aim of this study is to test the assumption that the relationship between the degree of dominance and local species richness may be different in grass communities with different productivities. Alpine, subalpine, and low-mountain grasslands, as well as subalpine mires, alpine communities of low-snow habitats and those with long-term snow cover, steppe communities, and the grass layer of low-mountain forest communities of the Western Caucasus and Ciscaucasia, are used as objects of research. The data on the phytomass of 419 plots with an area of 0.25 m² are studied. The results show that, the higher the mean productivity of communities is, the closer the relationship between the degree of dominance and species richness is, and the closest relationship is observed in meadow communities. Possible causes of these relationships are considered. It is reasonably suggested that this may be due to the features of the organization of plant communities with high and low productivity (in particular, high or low intensity of interspecific competition).     

Relationships between plant diversity,vegetation cover,and site conditions: implications for grassland conservation in the Greater Caucasus

Overgrazing, land use abandonment and increasing recreational activities have altered the vegetation of high-montane and subalpine grassland of the Caucasus. The failure of previous restoration efforts with unsuitable and exotic plant species indicates the need for information on the present vegetation and in which way it might change. Within the Greater Caucasus, we have described and quantified the mountain grassland which develops under characteristic overgrazed and eroded site conditions. Further, we have proposed potential native plant species for revegetation to restore and conserve valuable mountain grassland habitats. We used non-metric dimensional scaling ordination and cluster comparison of functional plant groups to describe a gradient of grassland vegetation cover. For our study region, we identified four major vegetation types with increasing occurrence of ruderal pasture weeds and tall herb vegetation on abandoned hay meadows within the subalpine zone. Within high-montane grassland a decline of plant diversity can be observed on sites of reduced vegetation cover. Due to a low potential of the grassland ecosystem to balance further vegetation cover damage, the long-term loss of diverse habitats can be expected. We conclude with management recommendations to prevent erosion and habitat loss of precious mountain grasslands.     

Snowbeds are more affected than other subalpine–alpine plant communities by climate change in the Swiss Alps

While the upward shift of plant species has been observed on many alpine and nival summits, the reaction of the subalpine and lower alpine plant communities to the current warming and lower snow precipitation has been little investigated so far. To this aim, 63 old, exhaustive plant inventories, distributed along a subalpine–alpine elevation gradient of the Swiss Alps and covering different plant community types (acidic and calcareous grasslands; windy ridges; snowbeds), were revisited after 25–50 years. Old and recent inventories were compared in terms of species diversity with Simpson diversity and Bray–Curtis dissimilarity indices, and in terms of community composition with principal component analysis. Changes in ecological conditions were inferred from the ecological indicator values. The alpha‐diversity increased in every plant community, likely because of the arrival of new species. As observed on mountain summits, the new species led to a homogenization of community compositions. The grasslands were quite stable in terms of species composition, whatever the bedrock type. Indeed, the newly arrived species were part of the typical species pool of the colonized community. In contrast, snowbed communities showed pronounced vegetation changes and a clear shift toward dryer conditions and shorter snow cover, evidenced by their colonization by species from surrounding grasslands. Longer growing seasons allow alpine grassland species, which are taller and hence more competitive, to colonize the snowbeds. This study showed that subalpine–alpine plant communities reacted differently to the ongoing climate changes. Lower snow/rain ratio and longer growing seasons seem to have a higher impact than warming, at least on plant communities dependent on long snow cover. Consequently, they are the most vulnerable to climate change and their persistence in the near future is seriously threatened. Subalpine and alpine grasslands are more stable, and, until now, they do not seem to be affected by a warmer climate.     

Population history of European mountain pines Pinus mugo and Pinus uncinata revealed by mitochondrial DNA markers

The dwarf mountain pine (Pinus mugo) and the Pyrenean pine (P. uncinata) constitute a pair of closely related coniferous taxa of poorly resolved evolutionary history and affinity, which inhabit numerous stands scattered over subalpine environments of European mountain ranges. The aim of the study was to investigate their phylogeography and mutual relationships, shedding new light on their taxonomy and the past of the alpine flora. Previous evolutionary reconstructions of the mountain pines relied mainly on bi‐parentally or paternally inherited markers that quickly homogenize between populations, showing rather shallow and recent differentiation of gene pools. Therefore, to contrast these pictures, we analyzed diversity and differentiation within a large set of new mitochondrial loci, inherited in maternal line and distributed by seeds at short geographical distances. Samples of the taxa were taken from 27 natural populations representing their range‐wide distributions—17 populations of P. mugo and 10 of P. uncinata. All markers appeared polymorphic, providing a total of 31 multilocus haplotypes. Two of the loci proved to be species‐diagnostic and nearly fixed between analyzed samples. Distribution of mitotypes indicate that allopatric populations of the taxa constitute separate mitochondrial haplogroups, and the two mountain pines have independent evolutionary history. However, introgression of P. mugo mitotypes by P. uncinata specimens revealed in the species contact zone in Western Alps shows that their speciation is not fully completed.     

山西五台山高山林线的植被景观

 过草本植物群落的分类和排序,结合对乔木和灌木分布的分析,确定了五台山高山林线的几条植被界线以及五台山森林上限附近植被的性质。结果表明：1）阳坡林线的海拔范围为2 605～2 790 m,阴坡林线的海拔范围为2 810～3 015 m;2）草本植物群落随海拔高度的变化比较明显,阴坡和阳坡从郁闭林到山顶均依次分布林下草本层、林缘草甸、亚高山灌丛草甸、高山草甸,草本植物的分布很好地体现了林线内部景观的差异性;3）海拔高度是高山林线附近草本植物群落空间分异的决定性因素。     

Virtual issue: Alpine and subalpine plant communities: importance of plant growth,reproduction and community assemblage processes for changing environments

Natural plant communities are exposed to environmental changes such as global warming and increased human activities. It is thought that alpine and subalpine ecosystems with cool climatic conditions are sensitive to environmental changes. This virtual issue introduces multidisciplinary research at alpine and subalpine plant communities. The articles include research on (1) species diversity, vegetation and biomass, (2) species assembly, (3) climate and growth of alpine plants, (4) reproduction of alpine plants, (5) differences of growth traits among coexisting species, (6) vegetation changes by human activities and overgrazing of deer, and (7) differentiation of growth traits among ecotypes in relation to climatic conditions. These thirteen articles provide valuable information for future research on the effects of environmental changes on alpine and subalpine plant communities.     

The Relationship between Local Species Richness and Species Pool: A Case Study from the High Mountains of the Greater Caucasus

The causes of linear relationships between local species richness and the size of the actual species pool in closed subalpine meadow communities and open plant communities of the alpine stony substrate (the Greater Caucasus Mountains) were analyzed using a computer simulation model. The results demonstrated that this relationship is insufficient evidence for the variation of local species richness among communities is wholly or partly determined by regional processes (the species-pool hypothesis). A relatively proportional ratio between these variables can also arise where local species richness and the size of the species pool both depend on local processes, or where local species richness is determined by local factors alone while the size of the species pool is determined by both local and regional factors.     

Facilitation and interference in subalpine meadows of the central Caucasus

Abstract. We studied the effects of neighbours on the biomass of seven randomly chosen species in species‐rich sub‐alpine meadows in the central Caucasus Mountains by comparing the performance of plants with neighbours removed experimentally to that of paired plants with their neighbours left intact. In most cases the removal of neighbours led to significant increases in vegetative and total above‐ground biomass implying the species were limited by competition. However, the neighbour removal led also to an increased leaf wilting for target plants, as well as to strong decline in reproductive effort for some species. We hypothesise that competition may be the prevailing type of interaction in species‐rich sub‐alpine meadow communities, but competitive effects on vegetative production may be balanced, if not outweighed, by facilitation, at least for some species. Such a balance may enhance species coexistence in communities.     

DOES INTERFERENCE CAUSE NICHE DIFFERENTIATION? EVIDENCE FROM SUBALPINE PLANT COMMUNITIES

The effects of interference on community structure of subalpine meadows were investigated. Adults greatly reduced seedling survival in the greenhouse, and natural seedling survival was low. In fell fields, nurse plant effects were common, while survival was confined to gaps in productive meadows. A greenhouse experiment demonstrated that community dominants were strong competitors able to suppress subordinates. Simulated grazing of Festuca, a dominant, reduced its yield relative to most subordinates. Interference intensity appears to be a function of productivity and proximity. The proportion of negative species associations increased as productivity increased. Morphological similarity between species is least in the most productive community, but greatest where productivity is only moderate. Interference may permit only relatively dissimilar species to coexist unless it is ameliorated by factors such as grazing or heterogeneity. Spacing patterns suggest that minimal contact between dominants and other species is a characteristic of communities with intense interference. The evidence points to this hypothesis: interference acts contemporaneously to limit niche width in productive communities, but evolutionary changes are unlikely where complete dominance is possible and specialization is not a viable option. In contrast, species in unproductive communities, where abiotic stress is likely to have been an evolutionary force, appear genetically more niche differentiated. Where contemporary interference is moderate, evolutionary effects are possible because species may use resources not preempted by the dominants.     

Changes in eco-morphological parameters of alpine plants' leaves as an effect of fertilization

Plants growing on rich soil usually have thin leaves with large specific leaf area. On the other hand, at intraspecific level; soil fertilization results in leaves size increasing which, in turn, can lead to reduction in specific leaf area. To what extent soil fertilization implies only leaves increasing in size and does not affect other eco-morphological characteristics is a question that is still open. To assess coherence between plants intraspecific reactions to changes in soil richness and general tendencies in changes of leaves parameters in communities with different productivity, an experiment has been conducted in alpine plant communities of the north-western Caucasus. Changes in leaf traits are studied in four types of alpine plant communities after long term application of mineral nutrients (NP and lime treatment). It is shown that in all species, except legume Hedysarum caucasicum, fertilization results in size leaf characteristics (leaf area, wet and dry mass) increase. Specific leaf area appears to decrease in plants inhabiting alpine heathlands and increase in plants inhabiting alpine snow beds and in dominant species of Geranium-Hedysarum meadows, Geranium gymnocaulon. After correction of specific leaf area that accounts for changes in leaf size, it becomes discernable that in most species the increase in leaf area per se results in specific leaf area reduction while changes in leaf structure under influence of fertilization leads to this trait increasing. Those species demonstrating the increase in specific leaf area as an effect of fertilization, also gain more in terms of biomass.     

Syntaxonomy of vegetation of Svjatoj Nos Peninsula,Lake Baikal 1. Non forest communities

Natural and semi-natural plant communities of Svjatoj Nos Peninsula on the East coast of Lake Baikal, Eastern Siberia, Russia, are classified and described using the methods of Braun-Blanquet phytosociology. A total of 48 associations and communities were recognized, comprising alpine tundra, subalpine forb vegetation, aquatic macrophyte vegetation, tall-herb and poor fens, mires, bogs, meadows, sand-dune and steppe vegetation. Twenty six syntaxa of ranks ranging from subassociation to class are described or validated for the first time. All communities are documented by phytosociological relevés.     

Experimental evidence for the immediate impact of fertilization and irrigation upon the plant and invertebrate communities of mountain grasslands

The response of montane and subalpine hay meadow plant and arthropod communities to the application of liquid manure and aerial irrigation – two novel, rapidly spreading management practices – remains poorly understood, which hampers the formulation of best practice management recommendations for both hay production and biodiversity preservation. In these nutrient-poor mountain grasslands, a moderate management regime could enhance overall conditions for biodiversity. This study experimentally assessed, at the site scale, among low-input montane and subalpine meadows, the short-term effects (1 year) of a moderate intensification (slurry fertilization: 26.7–53.3 kg N·ha⁻¹·year⁻¹; irrigation with sprinklers: 20 mm·week⁻¹; singly or combined together) on plant species richness, vegetation structure, hay production, and arthropod abundance and biomass in the inner European Alps (Valais, SW Switzerland). Results show that (1) montane and subalpine hay meadow ecological communities respond very rapidly to an intensification of management practices; (2) on a short-term basis, a moderate intensification of very low-input hay meadows has positive effects on plant species richness, vegetation structure, hay production, and arthropod abundance and biomass; (3) vegetation structure is likely to be the key factor limiting arthropod abundance and biomass. Our ongoing experiments will in the longer term identify which level of management intensity achieves an optimal balance between biodiversity and hay production.     

新疆博格达山地面生苔藓植被的数量分类与排序研究

 应用数量分类(TWINSPAN)和排序(DCA)方法,对新疆博格达山苔藓植被进行了多元分析。TWINSPAN结果表明,该地区苔藓植被可分为6个主要群落类型,即牛角藓-真藓(Cratoneuron-Bryum)群落、扭口藓-墙藓(Barbula-Tortula)群落、提灯藓-细罗藓(Mnium-Leskeella)群落、对叶藓-提灯藓(Distichium_Mnium)群落、墙藓-连轴藓(Tortula_Schistidium)群落、扭口藓-紫萼藓(Barbula_Grimmia)群落,它们分别分布于山地荒     

Floristic richness of three perhumid New Zealand alpine plant communities in comparison with other regions

Abstract Results are presented on vascular species richness in three representative alpine plant communities at 1040–1410 m on Mt Burns in the perhumid Fiordland region, a hotspot of alpine plant diversity, in south‐western South Island, New Zealand. Overall species richness was not dissimilar between the three communities in any of the eight plot sizes (mean values of 20.8–24.4 species in the largest plots of 100 m²), even though coefficients of floristic similarity were small (17.9; 23.5) between both low‐alpine communities (snow tussock‐shrubland and snow tussock grassland) and the high‐alpine cushion fellfield. Vascular species richness was generally similar to that in the few other oceanic New Zealand alpine communities for which data are available. The decline in richness from the low‐alpine to high‐alpine zones, revealed in more comprehensive records from two other regions with generally similar oceanic environments, was not recorded, indeed was reversed, on Mt Burns. Whether the recognized biodiversity hotspot of Fiordland has a generally richer high‐alpine flora than other regions in New Zealand needs further examination. The general pattern of alpine floristic richness in relation to elevation, in New Zealand, also prevails in most alpine regions abroad, usually under much more extreme continental environments. This pattern is usually ascribed to the associated decrease in temperature. Both the small size of the land mass and/or associated environmental conditions may be implicated but clarification awaits further data, preferably collected with standardized procedures.     

Integral assessment of the regrow capacity of alpine plants

A method for the comparative assessment of the regrow capacity (the ability of plants to the after-mowing recovery of above-ground biomass) of herbs in natural plant communities has been proposed. The method is based on the combined use of biomass parameters and the data on the number of shoots for the cases of the single and double removal of the biomass. The regrow capacity of 42 alpine species from four different plant communities has been assessed. The average weighted regrow capacity score decreases in the following series: Geranium-Hedysarum meadows > alpine lichen heaths ≥ alpine snow-bed communities ≥ Festuca varia meadows.     

Plant diversity changes and succession along resource availability and disturbance gradients in Kamchatka

Changes in plant species richness across environmental and temporal gradients have often been explained by the intermediate disturbance hypothesis and a unimodal diversity–productivity relationship. We tested these predictions using two sets of mountain plant communities assembled along postglacial successional and snow depth (disturbance and stress) gradients in maritime Kamchatka. In each community, we counted the number of species in plots of increasing sizes (0.0025–100 m²) and analyzed them using species–area curves fitted by the Arrhenius power function and the Gleason logarithmic function. A comparison of successional communities along a 270-year-old moraine chronosequence behind the receding Koryto Glacier—representing gradients of increasing productivity and resource competition—confirmed the unimodal species richness pattern. The plant diversity peaked in a 60–80-year-old Salix–Alnus stand where light availability was sufficient to sustain a rich understory combining pioneer and late successional herbs. The closed Alnus canopy on older moraines caused a pronounced decrease in species richness for all plot sizes (interactive stage 80–120 years since deglaciation). A slight increase in species richness in the oldest assortative stages (120–270 years), when Alnus stands are mature, was found only at the smaller spatial scales. This reflects (i) the consolidation of clonal understory dominants and (ii) the absence of other woody species such as Betula ermanii whose invasion would eliminate Alnus and increase diversity at larger spatial scales. A comparative study of major mountain plant communities distributed above the Koryto Glacier foreland did not confirm the highest species richness at intermediate levels of disturbance and stress. Contrary to our expectation, the species richness was highest in alpine tundra and snowbed communities, which are subjected to severe winter frost and a short summer season, while less disturbed communities of subalpine meadows, heaths, and Betula ermanii woods were less species-rich. We attribute this pattern to differences in habitat area and species pool size.     

新疆吉木萨尔县蝴蝶群落多样性

蝴蝶作为指示生物, 被广泛地应用于生物多样性监测及环境质量评估。探究新疆吉木萨尔县蝴蝶群落多样性, 可为当地蝴蝶多样性的保护及环境监测提供基础数据。本研究采用样线法在新疆吉木萨尔县选取山前荒漠、农田、山地草原、山地森林、亚高山草甸5种不同的生境类型, 对蝴蝶种类和群落多样性进行调查。共记录蝴蝶4,401号, 隶属于7科26属38种。其中蛱蝶科有9属12种, 为优势科; 粉蝶科的个体数最多, 占比55.01%; 绢蝶科、凤蝶科和弄蝶科的种类数和个体数最少, 均为单科种, 是该地区的稀有类群。对不同生境蝴蝶群落多样性和相似度分析比较的结果显示: 5种生境中多样性指数从高到低依次为亚高山草甸、山地森林、山地草原、农田及山前荒漠, 其中山地森林和亚高山草甸的相似性系数较高, 达到0.77, 山前荒漠和山地草原的相似性系数最低, 为0.37。蝴蝶物种数及多样性指数随海拔的增加呈上升趋势。蝴蝶群落随月份发生变化, 蝴蝶种类和数量在5月发生、7月达到峰值。蝶类个体数在3年内呈下降趋势。研究结果表明, 蝴蝶物种的组成和多样性与生境类型具有密切联系, 保护生态环境, 维持该地区植物群落的多样性、降低人为干扰程度是保护蝶类多样性的关键。     

Long‐term geobotanical observations of climate change impacts in the Scandes of West‐Central Sweden

In the context of projected future human‐caused climate warming, the present study reports and analyses the performance of subalpine/alpine plants, vegetation and phytogeographical patterns during the past century of about 1 °C temperature rise. Historical baseline data of altitudinal limits of woody and non‐woody plants in the southern Scandes of Sweden are compared with recent assessments of these limits at the same locations. The methodological approach also includes repeat photography, individual age determinations and analyses of permanent plots. At all levels, from trees to tiny herbs, and from high to low altitudes, the results converge to indicate a causal association between temperature rise and biotic evolution. The importance of snow cover phenology is particularly evident. Treeline advance since the early‐20th century varies between 75 and 130 m, depending on species and site. Tendencies and potentials for further upshift in a near future are evident from the appearance of young saplings of all tree species, growing 400–700 m atop of the treeline. Subalpine/alpine plant species have shifted upslope by average 200 m. In addition, present‐day repetitions of floristic inventories on two alpine mountain summits reveal increases of plant species richness by 58 and 67%, respectively, since the early‐1950s. Obviously, many plants adjust their altitudinal ranges to new climatic regimes much faster than generally assumed. Nevertheless, plants have migrated upslope with widely different rates. This produces non‐analogous alpine plant communities, i.e. peculiar mixtures of alpine and silvine species. The alpine region is shrinking (higher treeline), and the character of the remaining alpine vegetation landscape is changing. For example, extensive alpine grasslands are replacing snow bed plant communities.     

Microbotryum heliospermae, a new anther smut fungus parasitic on Heliosperma pusillum in the mountains of the European Alpine System

The members of the smut genus Microbotryum are pathogens of a wide range of host plant species from nine dicotyledonous families. Within the genus, the species sporulating in anthers of Caryophyllaceae form a monophyletic group that in recent years attracted much interest in various biological studies. The phylogenetic framework developed for species delimitation within Microbotryum revealed that high level host-specificity is a major feature of most caryophyllaceous anther smuts. However, the great number of anther smut specimens on diverse host plant species reported worldwide has still not been included in phylogenetic analyses due to the inaccessibility of recently collected specimens, and thus many species remain still undiscovered. In this study, anther smut specimens on Heliosperma pusillum originating from all main mountain ranges of the European Alpine System were examined using partial rDNA sequence and/or morphological analyses. The investigation revealed that all specimens are morphologically uniform and phylogenetically represent a monophyletic lineage, sister to Microbotryum lagerheimii complex on Atocion rupestre/Silene lacera/Silene vulgaris/Viscaria vulgaris. This lineage cannot be attributed to any of the previously described species, and therefore the smut in anthers of H. pusillum is described and illustrated here as a new species, Microbotryum heliospermae. The species is known from subalpine zone of the Alps, the Carpathians, the Dinaric Alps, and the Pyrenees, inhabiting host plants growing in open spring communities or semihumid mountain meadows.