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We performed a survey of grassland communities in the Ukrainian Carpathians with the aim of: (1) syntaxonomically classifying the meso- and subxerophilous grassland vegetation; (2) analysing the main gradients in their species composition; (3) estimating the effect of selected environmental factors on grassland species composition; (4) assessing the species richness of vascular plants and bryophytes in relation to the measured environmental variables. We collected 46 phytosociological relevés during the growing seasons of 2010 and 2011. Species composition and species richness were studied at two spatial scales (1 m² and 16 m²) in relation to soil parameters (soil depth, pH (KCl), content of P, K, Mg, N and C), management regime (mowing, grazing, ploughing in the past and burning), and other factors (altitude, litter cover, open soil, inclination, solar radiation and animal excrement). Seven grassland types were distinguished belonging to 3 classes and 4 alliances, namely the Nardetea strictae including the Violion caninae (mesic pastures at altitudes of 400–600 m mostly on moderate slopes) and the Nardo strictae-Agrostion tenuis (grasslands on moderate slopes at altitudes of 700–900 m usually managed by mowing and grazing the aftermath); the Molinio-Arrhenatheretea, including the Arrhenatherion elatioris (submontane grasslands originated mostly on former fields after their abandonment in the past) and the Cynosurion cristati (intensive pastures); and the Festuco-Brometea including the Cirsio-Brachypodion pinnati (abandoned grasslands dominated by Brachypodium pinnatum and Inula salicina). Detrended correspondence analysis indicated that the major compositional turnover was related to altitude and soil reaction. A canonical correspondence analysis confirmed that altitude had the strongest effect on species composition in the analysed dataset, followed by management treatments (former ploughing, grazing intensity). For vascular plant species richness, regression tree analysis identified grazing intensity as the most important predictor at the 1 m² scale. At the 16 m² scale, soil humus content was evaluated as the most important predictor of vascular plant species richness, followed by litter cover and grazing intensity. The number of bryophytes was not determined by the studied environmental factors at either of the two spatial scales. Although the number of analysed relevés in this study was limited, our results significantly contribute to the understanding of submontane grasslands in the Ukrainian Carpathians.
A long-term systematic survey of grassland communities was performed in the Biele Karpaty Mts. in Slovakia. The main aims of the research were i) syntaxonomical classification of meso- and subxerophilous grassland vegetation, ii) analysis of the main gradients in species composition, iii) evaluation of the effect of environmental factors on species composition of grasslands. The data set included 342 phytosociological relevés of grasslands recorded between 1991 and 1999. For the classification of relevés to associations, the expert system for identification of grassland vegetation of Slovakia was used. The main environmental gradients of species composition were analysed by detrended correspondence analysis (DCA). For the ecological interpretation of ordination axes Ellenberg indicator values were used. The relationship between species composition and environmental factors (geology, pedology, climate, topography, management) was analysed by canonical correspondence analysis (CCA). The expert system identified (according to association definitions) 220 phytosociological relevés (64% of the whole data set). Grassland communities were classified within seven associations belonging to four alliances and three classes: Festuco-Brometea: Bromion erecti and Cirsio-Brachypodion pinnati; Molinio- Arrhenatheretea: Arrhenatherion; Nardetea strictae: Violion caninae. The results of the DCA support our assumption that the main environmental gradient in species compositions of grasslands is related to moisture and soil reaction (content of CaCO3 in the soil). The results of the direct gradient analysis (CCA) show that all 23 environmental variables explained 16.15% of the variability of the species data. The most important factors affecting the data variation were precipitation and geological bedrock.
Syntaxonomic revision of the Pannonian grasslands of Austria : 1. Introduction and general overview
(2013)
The Pannonian part of Austria is a diverse landscape situated in the transition zone between the Alps, the Bohemian Massif and the Carpathian Basin. Although the grasslands of this region have been investigated in many botanical and vegetation studies, their phytosociological classification has remained confusing. With this paper, we start a series aiming at a developement of a revised, consistent system of the Austrian Pannonian grasslands. Here we present a general overview focusing on the higher syntaxonomic units. We define grasslands as all types of meadows, pastures, fens and primary steppes. We selected all available relevés of Pannonian grasslands from the Austrian Vegetation Database. Additional unpub-lished data were included from the Danube National Park and the Biosphere Reserve Vienna Woods. To account for the comparatively low number of relevés from the northern part of the Pannonian region of Austria (Weinviertel), we included also data from southern Moravia (Czech Republic). This set of 3384 relevés was classified using TWINSPAN. Relevés that were considered as misclassified at the alliance level according to the summarised cover of diagnostic species were manually re-arranged, and the data-set specific fidelity of species to alliances was calculated using the phi coefficient. The first TWINSPAN division largely corresponded to the traditional border between the classes Festuco-Brometea and Molinio-Arrhenatheretea. The conventional alliance concepts were generally well supported. As an exception, the distinction between Diantho-Seslerion and Bromo-Festucion pallentis was not reproduced at all. Therefore, we unite all rocky grasslands on calcareous soils in a single alliance Seslerio-Festucion pallentis. We also advocate the inclusion of all basiphilous semi-dry grasslands of the study area within a single alliance Cirsio-Brachypodion. Each of the corresponding TWINSPAN clusters showed a clear prevalence of Cirsio-Brachypodion species. Moreover, two separate alliances of semi-dry grasslands would have almost no regional character species.
As second part of a series focusing on the Pannonian grasslands of Austria, we present a detailed classification of the grasslands of the Vienna Woods (Wienerwald). This region, although geographically belonging to the Alps, has strong floristic affinities to the Pannonian Basin. The eastern slopes of the Vienna Woods are a hotspot of xero-thermophytic vegetation and exhibit the highest vascular plant species richness in Austria at a scale of 3' x 5'. We used the TWINSPAN classification of a large data set reported in the first part of this series as starting point. Relevés that were considered as misclassified at the level of alliances were manually re-arranged. From this table, an excerpt containing only the relevés of the Vienna Woods was used for the present study (1055 plots). We re-classified the relevés preliminary labelled as Arrhenatherion with another TWINSPAN run. Clusters for which no ecological difference could be detected were merged. The final delimitation of associations and subassociations was achieved by adjusting their diagnostic species so as to get units that were most informative in terms of environmental conditions. All re-arrangements were based on the summarised cover of diagnostic species within individual relevés. As a result, the grasslands of the Vienna Woods are classified within 22 associations (plus two subassociations) belonging to ten alliances and four classes. Within the Seslerio-Festucion pallentis, a new association Scorzonero austriacae-Caricetum humilis is described. The semi-dry grasslands of the study area previously classified as Onobrychido-Brometum are de-scribed as new association Filipendulo vulgaris-Brometum erecti and assigned to the Cirsio-Brachypodion. The Anthoxantho-Agrostietum tenuis is reported for Austria for the first time, and its syntaxonomy is discussed within a broader geographical context.
Semi-dry grasslands were once widely distributed communities, but today they represent some of the most vulnerable habitats in Central Europe. European and national legislation and non-governmental organizations have managed to protect some of the remaining fragments. However, despite their status as Natura 2000 habitats, they are often endangered due to improper management, fragmentation and edge effects from adjacent croplands. By using a sample of 44 semi-dry hay meadows in the south-eastern Alpine Foreland of Styria, we investigated how species-richness and trait composition of semi-dry grassland species respond to variation in patch size, connectivity, abiotic site factors and management regimes. We used linear regression models to identify the most important drivers for richness of typical semi-dry grassland species and thus conservation value. The number of typical semi-dry grassland species was highest in well-connected fragments, i.e. units that shared two or more borders with neighbouring species-rich grasslands. Furthermore, large semi-dry grasslands (> 8000 m²) had highest numbers of semi-dry grassland species and highest relevance for conservation; no difference was found among smaller fragment sizes. Unregular management was associated with increased presence of competitive species which replaced stress-tolerant specialists. Our study indicates that under eutrophication, small fragment size and isolation, only large semi-dry grasslands can sustain a high number of species with high conservation value. The conservation value of smaller semi-dry grassland fragments could be improved by buffer zones, adapted mowing treatments and periodical sheep grazing.
In this paper we aim to present data on current state of steppe-like grasslands dominated by feather grasses (Stipa spp.) in the Transylvanian Lowland (Romania), and to investigate whether management type affects species composition, biodiversity, or the occurrence of rare and endangered vascular plant species in these grasslands. Twelve grassland stands, originating from nine areas and belonging to two associations, Stipetum lessingianae and Stipetum pulcherrimae from the alliance Stipion lessingianae (Festucetalia valesiacae, Festuco-Brometea), were investigated by phytosociological sampling. Our data set contains 60 relevés originating from differently managed grasslands: grazed and abandoned stands as well as former grasslands afforested with Pinus nigra. Transylvanian stands of the Stipion lessingianae occur on steep, south-facing slopes. The species composition has an accentuated sub-continental character, including several steppe elements of Siberian and Pontic origin, many of them reaching the western edge of their distribution in this region. By comparing the species composition of the two associations, we identified 12 differential species for the Stipetum lessingianae and 19 for the Stipetum pulcherrimae, but these associations also contained many species in common, which underlines their close syntaxonomic relatedness. Grazing vs. abandonment had only a slight influence on species composition and vegetation structure, while afforestation with pines resulted in strong transformations: decreased cover of the herb layer, increased cover of litter, decreased representation of diagnostic (i.e. typical) species of the two associations but increased frequency of woody, ruderal, and mesophytic grassland species. As the Stipion lessingianae belongs to the priority habitats of the Habitats Directive, these alterations call for grassland restoration measures. Our study revealed a lack of high-quality data on the vegetation of steppe-like dry grasslands in southeast Europe, where they have their largest and best-developed stands on the continent. Thus, we emphasize the need for compiling such data and for developing a consistent supranational vegetation classification of Festucetalia valesiacae communities in this region.
Using four exclosures, the impact of mouflon grazing and weather on plant communities of the phytosociological alliances Koelerio-Phleion phleoidis and Hyperico perforati-Scleranthion perennis occurring on shallow soils within a forest landscape was studied in the Křivoklátsko Biosphere Reserve (Czech Republic) during seven years. In the years 2004–2010, the vascular plant species composition was recorded annually on a total of eight fenced and eight control plots, each 2 m × 1 m in size. Treatment, time, and weather data were used as explanatory variables in ordination and correlation analyses. Cover values of different life forms and indicator values varied significantly in dependence on the weather conditions of the preceding five months; however, correlations varied according to the vegetation type and were rather rarely detected. The effect of fencing appeared important in all study plots; however, the temporal trends were significant only in half of them. We did not find a significant interaction between treatment and time in the total dataset. A successional change was detected in one fenced plot only; in all other cases, the species composition fluctuated – a phenomenon that is not directly attributable to weather conditions. In the fenced plots, the herbaceous vegetation cover decreased, mainly due to litter accumulation and partly due to shrub encroachment. At least some parts of the valuable and species-rich habitats could be maintained under high game density, but some parts are endangered by eutrophication and game grazing. Nature conservation management should balance both mechanisms.
Dry grasslands of NW Bulgarian mountains : first insights into diversity, ecology and syntaxonomy
(2013)
We present the data of the 3rd research expedition of the European Dry Grasslands Group (EDGG), which was conducted in 2011 in two contrasting areas in NW Bulgarian mountains. The aim was to collect plot data for comparing Bulgarian dry grasslands with those of other parts of Europe in terms of syntaxonomy and biodiversity. We sampled 15 nested-plot series (0.0001–100 m²) and 68 normal plots (10 m²) covering the full variety of dry grassland types occurring in the Vratsa area (Balkan Mts.) and the Koprivshtitsa area (Sredna Gora Mt.). In the plots all vascular plants, terricolous non-vascular plants and a set of soil and other environmental parameters were determined. By applying modified TWIN-SPAN, we distinguished 10 floristically well characterised vegetation types at the association level. After comparison with the regional and European literature, we propose to place them within three classes and five orders: Festuco-Brometea with the orders Stipo pulcherrimae-Festucetalia pallentis (xerophilous dry grasslands of base-rich rocks; alliance Saturejion montanae), Brachypodietalia pinnati (meso-xeric, basiphilous grasslands; alliances Cirsio-Brachypodion pinnati and Chyrsopogono grylli-Danthonion calycinae), Calluno-Ulicetea with the order Nardetalia stricae (lowland to montane Nar-dus swards; alliance Violion caninae), and Koelerio-Corynephoretea with the orders Sedo-Scleranthetalia (open communities of skeleton-rich, acidic soils; alliance unclear) and Trifolio arvensis-Festucetalia ovinae (closed, meso-xeric, acidophilous grasslands; alliance Armerio rumelicae-Potentillion). The Violion caninae with the association Festuco rubrae-Genistelletum sagittalis is reported from Bulgaria for the first time, while the two occurring Koelerio-Corynephoretea communities are described as new associations (Cetrario aculeatae-Plantaginetum radicatae, Plantagini radicatae-Agrostietum capillaris). According to DCA the main floristic gradient was largely determined by soil conditions, differentiating the Festuco-Brometea communities on soils with high pH and high humus content from the Koelerio-Corynephoretea communities on acidic, humus-poor soils, while the Calluno-Ulicetea stands are the connecting link. At 10 m2 Festuco-Brometea and Calluno-Ulicetea stands were richer in species across all investigated taxa and in vascular plants than Koelerio-Corynephoretea stands; the latter were richest in lichen species, while bryophyte richness did not differ significantly among syntaxa. Among the Bulgarian classes, the species-area relationships tended to be steepest in the Festuco-Brometea (i.e. highest beta diversity), but both alpha and beta diversity clearly fell behind the Festuco-Brometea communities in the Transylvanian Plateau, Romania, located less than 500 km north of the study region. Overall, our study contributes to a more adequate placement of the Bulgarian dry grasslands in the European syntaxonomic system and provides valuable data for large-scale analyses of biodiversity patterns.
Classification of higher level vegetation units (orders and alliances) based on numerical methods often yields different results than traditional plant community classification concepts. We performed a numerical cluster analysis of phytosociological relevés from the class Festuco-Brometea in Slovakia with the aim of identifying areas of overlap between the two classification approaches. The research was carried out using a database of approximately 1500 phytosociological relevés sampled in the period between 1927 and 2004. The outputs of the numerical classification form six clusters. Diagnostic taxa of individual clusters were determined using species constancy and fidelity. The cluster analysis enabled us to differentiate the alliances Seslerio-Festucion pallentis, Diantho lumnitzeri-Seslerion albicantis, Festucion valesiacae, Cirsio-Brachypodion pinnati and Asplenio septentrionalis-Festucion pallentis (inch Festucenion pseudodalmaticae). However, it did not permit the differentiation of the alliances Koelerio-Phleion phleoidis and Bromion erecti. It also did not allow us to differentiate the orders Brometalia erecti and Festucetalia valesiacae. The reason for this may be the peripheral occurrence of plant communities of Brometalia erecti in Slovakia.
We present the data of the 2nd research expedition of the European Dry Grassland Group (EDGG), which was conducted in 2010 in Central Podolia, Ukraine. The aim was to collect plot data to compare Ukrainian dry grasslands with those of other parts of Europe in terms of syntaxonomy and biodiversity. We sampled 21 nested-plot series (0.0001–100 m2) and 184 normal plots (10 m2) covering the full variety of dry grassland types occurring in the study region. For all plots, we recorded species composi-tion of terrestrial vascular plants, bryophytes and lichens, while for the 226 10-m2 plots we estimated and measured percentage cover of all species, structural, topographic, soil and landuse parameters. The 10-m² plots were used for phytosociological classification based on iteratively refined TWINSPAN classification as well as for DCA ordination. Differences between the derived vegetation types with respect to environmental conditions and species richness were assessed with ANOVAs. We assigned our plots to nine association-level units but refrained from placing them into formal associations with two exceptions. In the study area, dry grasslands of the Festuco-Brometea were far more common than those of the Koelerio-Corynephoretea. Among the Festuco-Brometea, xeric Festucetalia valesiacae grasslands were more frequent and represented by the Festucion valesiacae (2 associations, including the Allio taurici-Dichanthietum ischaemi ass. nova) and the Stipion lessingianae (1) compared to the Brachypodietalia pinnati with the Agrostio vinealis-Avenulion schellianae (3). The Koelerio-Corynephoretea were represented by three associations, each from a different order and alliance: basiphilous outcrops (Alysso alyssoidis-Sedetalia: Alysso alyssoidis-Sedion?), acidophilous outcrops (Sedo-Scleranthetalia: Veronico dillenii-Sedion albi?) and mesoxeric sandy grasslands (Trifolio arvensis-Festucetalia ovinae: Agrostion vinealis). We discuss the issue of the mesoxeric order Galietalia veri placed within the Molinio-Arrhenatheretea by Ukrainian authors and conclude that the content of that order would probably be better placed in the mesoxeric orders of the Koelerio-Corynephoretea and Festuco-Brometea. Other syntaxonomic questions could not be solved with our geographically limited dataset and await a supraregional analysis, e.g. whether the Ukrainian outcrop communities should be assigned to the same alliances as known from Central Europe or rather represent new vicariant units. The analysis of the biodiversity patterns showed that at a grain size of 10 m2, Podolian Koelerio-Corynephoretea communities were overall richer than Festuco-Brometea communities (46.4 vs. 40.6 species). This difference was due to the Koelerio-Corynephoretea containing twice as many bryophytes and nine times more lichens, while vascular plant species richness did not differ significantly between classes. The orders within the classes showed no real differences in species richness. The richness patterns observed in Podolia were almost the opposite of those usually found in dry grasslands, where Brachypodietalia pinnati are richer than Festucetalia valesiacae, and these richer than stands of the Koelerio-Corynpehoretea – and we do not have a good explanation for these idiosyncrasies. In conclusion, Podolian dry grasslands behave quite unexpectedly regarding biodiversity, and their syntaxonomy is still poorly understood. These knowledge gaps can only be addressed with supranational analyses based on comprehensive datasets.