Comparative analysis of the floristic richness of alpine communities in the Caucasus and the Central Alps

Abstract. Species richness of selected alpine plant communities in the Teberda Biosphere Reserve (Northwestern Caucasus) and the Davos area (Central Alps) was compared in series of plots from 0.0025 to 100 m². Communities developing under similar ecological conditions and with similar syntaxonomic positions were compared in order to estimate the role of recent environment versus regional historical factors in determining plant community structure and diversity. The floristic richness of the Caucasian and Alpic fens was very similar. The Grasslands and Meadows were quite similar as to floristic richness for plots > 25 cm × 25 cm, but the Caucasian communities had fewer species in smaller plots. The Lichen heath at Teberda was richer than the Caricetum curvulae cetrarietosum for all plot sizes, except the two smallest ones. On the other hand, the plots of the Salix Snowbed community were richer in species than the Caucasian snowbeds for all plot sizes. The Rhododendron Shrubland plots were very similar as to floristic richness in larger plots (4 — 100 m²). Generally, most Alpic communities near Davos were richer in species at small plot sizes than the corresponding communities from Teberda. Caucasian communities were floristically less similar to each other — and thus more discrete — than the Alpic ones. The possible role of different factors controlling floristic richness of the communities is discussed. Our results suggest that recent ecological conditions have a big influence on local floristic diversity and may lead to high similarities between ecologically similar communities from different regions. In addition, the general floristic richness of a regions as well as island effects should be taken into account.     

Nitrogen dynamics in alpine ecosystems of the northern Caucasus

Net N mineralization, nitrification, microbial biomass N and ¹⁵N natural abundance were studied in a toposequence of representative soils and plant communities in the alpine zone of the northern Caucasus. The toposequence was represented by (1) low-productive alpine lichen heath (ALH) of wind-exposed ridge and upper slope; (2) more productive Festuca varia grassland (FG) of middle slope; (3) most productive Geranium gymnocaulon/Hedusarum caucasicummeadow (GHM) of lower slope; (4) low-productive snowbed community (SBC) of the slope bottom. N availability, net N mineralization and nitrification were higher in soils of alpine grassland and meadow of the middle part of the toposequence compared with soils of lichen heath and snowbed community of extreme habitats in the alpine zone. There was no correlation between intensities of N transformation processes and favorable (low soil acidity, low C/N ratio, long vegetation period, relatively high temperature, absence of hydromorphic features) and unfavorable (opposite) factors, indicating that the intensity of N mineralization and nitrification in the alpine soils is controlled by a complex combination of these factors. Potential net N mineralization and nitrification in alpine soils determined in the short-term laboratory incubation were considerably higher than those determined in the long-term field incubation. The differences of potential nitrification between soils of various plant communities did not correspond to the field determined pattern indicating the importance of on-site climatic conditions for control of nitrification in high mountains. The result of comparison of N transformation potentials in incubated and native soils indicated that nitrification potential was significantly increased after long-term soil incubation. It means that net nitrification determined in the field was probably overestimated, especially in the meadow soils. A soil translocation experiment indicated that low temperature was an important factor limiting net N mineralization and nitrification in alpine soils: net N mineralization and especially nitrification increased when alpine soils were translocated into the subalpine zone and mean annual temperature increased by about 3°C. Additional N input increased N availability (NH₄ ⁺-N) and potential nitrification in soils of the lower part of the toposequense (GHM and SBC), and potential net N mineralization in two soils of extreme habitats (ALH and SBC). A positive correlation was found between soil ¹⁵N and net N mineralization and nitrification; the relative ¹⁵N enrichment was characteristic of grassland and meadow ecosystems. ¹⁵N of total soil N pool increased during the field mineralization experiment; there was a positive tendency between the change in ¹⁵N and net N mineralization and nitrification, however the relationship was not significant. Foliar ¹⁵N of dominant plant species varied widely within community, however, a tendency of higher foliar ¹⁵N for species growing on the soils with higher net N mineralization, nitrification and ¹⁵N was observed.     

Morphological analysis of herbaceous communities under different grazing regimes

Abstract. A methodology for the morphological analysis of herbaceous communities is presented, together with an example of its application in montane grasslands in the province of Córdoba (Argentina) subject to grazing and burning. The method, based on multivariate ordination and classification techniques, enabled the detection of morphological changes at three levels in response to disturbance: (a) characterization of the spatial structure of the vegetation; (b) identification of morphological plant groups; and (c) quantification of morphological shifts among different individuals of a single species. The architecture of the vegetation changed toward a progressive miniaturization of photosynthetic structures and concentration of biomass close to the ground, as disturbance intensity increased. Six morphological plant groups (modes of response) showing different behaviour in relation to competition for light and pressure from large herbivores were identified. Some species, highly preferred by ungulate grazers, showed high morphological variability among morphs growing in different grazing situations, whereas some others were morphologically uniform.     

Population strategies in severe environments: alpine plants in the northwestern Caucasus

Abstract. This study attempts to show a differentiation into strategy types among plant species of four alpine communities in the northwestern Caucasus, Russia. Four groups of population characteristics (traits) were examined: above-ground biomass, seed size, seed yield and occurrence of a persistent seed bank. In addition, values for relative growth rate and a morphology index were estimated. Agglomerative Cluster Analysis and Principal Component Analysis were applied to a data set with 42 species from these communities and six species characteristics. In the Geranium gymnocaulon meadow, the most productive community, large gaps occur created by wild boars and bears. Here, the response of 13 species to disturbance was also analysed. Strategy types were approached both according to Grime: ‘competitive’, ‘ruderal’, ‘stress-tolerating’ and according to Ramensky, Rabotnov and Romanovsky: ‘violent’ ≈ dominant (V), ‘explerent’ ≈ explorative (E) and ‘patient’ ≈ endurant (P). Differentiation into strategy types was more obvious in productive alpine grasslands than in lichen heaths and snowbed communities. The grassland dominants Festuca varia, Geranium gymnocaulon and Hedysarum caucasicum showed a more or less ‘violent’ strategy. Species approaching an Estrategy, e.g. Matricaria caucasica, Sibbaldia procumbens and Gnaphalium supinum, were also found in the grasslands but prevailed in the snowbed communities. Species with a ‘patient’ strategy prevailed in the lichen heaths. We conclude that plant strategy types are well-differentiated, this is also seen in plant communities occurring under severe high-mountain conditions. We also suggest that the approaches by Grime and Ramensky/Rabotnov/Romanovsky are not so similar as has been assumed earlier and that the R/R/R types are better differentiated and ecologically better interpretable. We emphasize the gradient character of strategy ‘types’ and would characterize plant species by the degree of adaptation to conditions: (1) from small seed size and large persistent seed bank to large seed size and absence of a seed bank; (2) from high to low biomass and seed production, with relative growth rate as a less important partially correlated trait.     

Response of Norwegian alpine communities to nitrogen

The effect of nitrogen pollution on species cover was tested in the Dovre mountains, south-central Norway, through a three-year fertilisation experiment in a low-alpine community and a two-year one in a middle-alpine community, both dominated by lichens. The doses used were 0.7, 3.5 and 7 g N/m², the highest doses corresponding to 1.75 and 3.5 times the annual deposition of nitrogen in southwestern Norway. The results were tested using the SAS/STAT/MIXED procedure. Using the Bonferroni correction, the procedure did not confirm a change in the cover of single species due to increased nitrogen supply, but there seemed to be a general tendency for an increase in cover in the low-alpine Cetrarietum nivalis community. When less rigorous tests were applied, Arctostaphylos uva-ursi, Vaccinium vitis-idaea, Polytrichum piliferum, Cetraria nivalis and Cladonia mitis in Cetrarietum nivalis, and Cladonia coccifera in Phyllodoco-Juncetum trifidi seemed to indicate that nitrogen had some effect on these communities. The lack of a clear response in single species may be due to the application of low doses over a short time-span in an environment characterised by large interannual variations in climate, the deficiency of other important nutrients, dryness, the low growth potential of the species involved, and, to some extent, the community structure.     

Dynamics of development of juvenile plants on the meadows of the alpine belt of northwest Caucasus

We investigated juvenile plants on the Hedysaro caucasicae-Gerametum gymnocauli meadows (Rabotnova and Onipchenko, 2002) of the alpine meadows of the northwest Caucasus. During the course of 5 years, we considered all juvenile plants on 16 plots of 0.5 m². We recorded 4485 sprouts and monitored their destiny. The dominant species were Hedysarum caucasicum Bieb. and Geranium gymnocaulon DC., and most species of Asteraceae had juvenile plants, but without seeds in soil bank of seeds. The grass species rarely produced sprouts and had seeds in the soil bank. The majority of sprouts died after the first winter. Sprouts of the dominant species were in their juvenile stage for a long time, and the sprouts of other species quickly shifted to the vegetative stage.     

Modelling species richness and diversity in grassland communities of the Central Caucasus

In this study I used small squares (4 cm×4 cm) as a sampling technique within plots (128 cm×128 cm) of different elevation, aspect and slope angle in grassland communities (20 plots examined). Then I used a rectangular hyperbole equation (the Michaelis-Menten model) to describe species richness and the Inverse of Simpson Concentration (ISC) as functions of sample size. I checked robustness and precision of the model both by interpolation and extrapolation. Interpolation was similarly good in both cases, while extrapolation produced reliable predictions of ISC but underestimated species richness. Dominance analysis indicated that the underestimation of richness depends on the proportion of bottom species, and that the predicted values of richness roughly coincide with the numbers of dominant species found in plots. Therefore, the model may be used to assess number of dominant species when precision is less important than saving time during a survey. However, the rectangular hyperbole equation appears to be precise and robust in the prediction of ISC, at least in grassland communities. This property may also be employed for extrapolation of diversity indices with a limited sampling effort.     

Insect-associated bacterial communities in an alpine stream

Hydrobiologia - The roles of macroinvertebrate and microbial communities in stream ecosystems are recognized to be important to energy flow and nutrient cycling. While the linkages of these major...     

Methane emissions by alpine plant communities in the Qinghai-Tibet Plateau

For the first time to our knowledge, we report here methane emissions by plant communities in alpine ecosystems in the Qinghai-Tibet Plateau. This has been achieved through long-term field observations from June 2003 to July 2006 using a closed chamber technique. Strong methane emission at the rate of 26.2+/-1.2 and 7.8+/-1.1microg CH4 m-2h-1 was observed for a grass community in a Kobresia humilis meadow and a Potentilla fruticosa meadow, respectively. A shrub community in the Potentilla meadow consumed atmospheric methane at the rate of 5.8+/-1.3microg CH4 m-2h-1 on a regional basis; plants from alpine meadows contribute at least 0.13Tg CH4 yr-1 in the Tibetan Plateau. This finding has important implications with regard to the regional methane budget and species-level difference should be considered when assessing methane emissions by plants.     

Life-form composition of alpine plant communities at the Eastern Qinghai-Tibetan plateau

Alpine plants of the eastern Qinghai-Tibetan plateau (Sichuan, China) are developed under long-lasting grazing by wild and domestic yaks. Among morphological features of plants, life forms may reflect their adaptation to grazing. We studied life-form composition of four typical communities within the alpine belt (3930–3960 m a.s.l.) subjected to grazing of various intensity: alpine fen (heavily grazed), alpine shrub meadow (heavily grazed), Spiraea alpina thicket (grazed), and Rhododendron thicket (practically not grazed). The following morphological traits were studied: (1) life form according to Raunkiaer, (2) life form according to Serebryakov, (3) canopy structure, and (4) rate of lateral spreading. We derived life-form spectra based on (1) the number of species per life form and (2) the cumulative abundance of species which have the same life form. One-way ANOVA and nonparametric ANOVA were run to test for significance of differences between spectra. The studied communities differed significantly by the proportion of different life forms. The main life forms are caudex and short rhizome hemicryptophytes, nonclonal species, or species with a low rate of lateral spreading. Therophytes made up 10–11% of the communities except in Rhododendron thickets, where such were absent. These life forms can indicate grazing in the study area.     

Contrasting altitudinal variation of alpine plant communities along the Swedish mountains

Changes in abiotic factors along altitudinal and latitudinal gradients cause powerful environmental gradients. The topography of alpine areas generates environmental gradients over short distances, and alpine areas are expected to experience greater temperature increase compared to the global average. In this study, we investigate alpha, beta, and gamma diversity, as well as community structure, of vascular plant communities along altitudinal gradients at three latitudes in the Swedish mountains. Species richness and evenness decreased with altitude, but the patterns within the altitudinal gradient varied between sites, including a sudden decrease at high altitude, a monotonic decrease, and a unimodal pattern. However, we did not observe a decline in beta diversity with altitude at all sites, and plant communities at all sites were spatially nested according to some other factors than altitude, such as the availability of water or microtopographic position. Moreover, the observed diversity patterns did not follow the latitudinal gradient. We observed a spatial modularity according to altitude, which was consistent across sites. Our results suggest strong influences of site‐specific factors on plant community composition and that such factors partly may override effects from altitudinal and latitudinal environmental variation. Spatial variation of the observed vascular plant communities appears to have been caused by a combination of processes at multiple spatial scales.     

Morphological and molecular systematics of Rocky Mountain alpine Laccaria

The alpine zone is comprised of habitats at elevations above treeline, and macromycetes play important ecological roles as decomposers and mycorrhizal symbionts here as elsewhere. Laccaria is an important group of ectomycorrhizal basidiomycetes widely used in experimental and applied research. A systematic study of alpine Laccaria species using morphological, cultural and molecular (ribosomal DNA internal transcribed spacer) data revealed five taxa in the Rocky Mountain alpine zone: L. laccata var. pallidifolia, L. nobilis (the first published report for arctic-alpine habitats), L. pumila, L. montana and L. pseudomontana (a newly described taxon similar to L. montana with more ellipsoidal, finely echinulate basidiospores). All occur in the southern Rocky Mountains of Colorado; however, only L. pumila and L. montana were found on the Beartooth Plateau in the northern Rocky Mountains of Montana and Wyoming. All are associated with dwarf and shrub Salix species, with L. laccata var. pallidifolia also associated with Dryas octopetala and Betula glandulosa. Maximum-parsimony phylogenetic analysis of rDNA-ITS sequences for 27 Laccaria accessions supports the morphological species delineations.     

Factors affecting bird communities in fragments ofsecondary pine forests in the north-western Mediterranean basin

We assessed the influence of size, extent of isolation and vegetation structure of secondary forest fragments on the richness and species composition of breeding bird communities in a sample of pine forest fragments surrounded by an agricultural matrix in the north-western Mediterranean basin. Fragment size was the main predictor of bird’s occurrence, since it accounted for 70 % of the model variation. Isolation was also a valuable predictor of species occurrence, especially for forest specialists. Finally, subarboreal vegetation such as holm oak and a well-developed tree layer of large pines favoured forest species occurrence. Therefore, in spite of the long history of human impact, forest birds in Mediterranean mosaics are sensitive to both habitat loss and isolation of remnant patches in a similar manner to the patterns found in other temperate fragmented landscapes where human impact is more recent.     

Microbial communities and activities in alpine and subalpine soils

Soil samples were collected along two slopes (south and north) at subalpine (1500–1900 m, under closed vegetation, up to the forest line) and alpine altitudes (2300–2530, under scattered vegetation, above the forest line) in the Grossglockner mountain area (Austrian central Alps). Soils were analyzed for a number of properties, including physical and chemical soil properties, microbial activity and microbial communities that were investigated using culture-dependent (viable heterotrophic bacteria) and culture-independent methods (phospholipid fatty acid analysis, FISH). Alpine soils were characterized by significantly ( P <0.01) colder climate conditions, i.e. lower mean annual air and soil temperatures, more frost and ice days and higher precipitation, compared with subalpine soils. Microbial activity (soil dehydrogenase activity) decreased with altitude; however, dehydrogenase activity was better adapted to cold in alpine soils compared with subalpine soils, as shown by the lower apparent optimum temperature for activity (30 vs. 37 °C) and the significantly ( P <0.01–0.001) higher relative activity in the low-temperature range. With increasing altitude, i.e. in alpine soils, a significant ( P <0.05–0.01) increase in the relative amount of culturable psychrophilic heterotrophic bacteria, in the relative amount of the fungal population and in the relative amount of Gram-negative bacteria was found, which indicates shifts in microbial community composition with altitude.