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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.
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.
We studied the dry grasslands of shallow, skeletal soils (Sedo-Scleranthenea, Koelerio-Corynephoretea) in northern Europe, based on a combination of new relevés from southern Oland (Sweden, n = 182) and Saaremaa (Estonia, n = 73) as well as a comprehensive evaluation of literature data, of which 65 suitable relevés were directly included in our analyses. Apart from a few vague indications of acidophytic Sedo-Scleranthenea communities (order Sedo-Scleranthetalia), all data refer to basiphytic communities (Alysso-Sedetalia); our analyses are thus focussed on the latter. The Nordic Alysso-Sedetalia communities proved to be quite different from their temperate counterparts and thus are included in a separate alliance, Tortello tortuosae-Sedion albi, which forms the northern counterpart to the central European Alysso-Sedion. Within the northern alliance, we distinguish two suballiances. The more widespread central suballiance Tortello tortuosae-Sedenion albi inhabits different types of base-rich substrata in both natural and anthropogenic sites, and is comprised of the Cladonio symphicarpiae-Sedetum albi and the Ditricho flexicaulis-Sedetum acris. The second suballiance Tortello rigentis-Helianthemenion oelandici is restricted to the alvar sites (= treeless limestone plateaus) in Oland, Gotland, Västergötland and Estonia. It is characterised by several endemic taxa and a large number of cryptogams typical of alvar. It is comprised of four associations, Crepido pumilae-Allietum alvarensis, Fulgensio bracteatae-Poetum alpinae, Helianthemo oelandici-Galietum oelandici and Gypsophilo fastigiatae-Globularietum vulgaris. All six Nordic associations are described in detail with respect to their floristic composition, ecology, distribution and lower-ranked units, and each is represented by a vegetation table. The floristic differences within the Nordic communities are worked out in a synoptic table. Whereas several vegetation scientists have pointed out that vegetation types occurring at the limits of their distribution ranges in northern Europe are generally difficult to classify, our application of the Braun-Blanquet approach, which is based on a priori separated structural types and the general application of the central syntaxon concept, has enabled us to characterise and adequately define all Nordic communities. The Tortello-Sedion associations are two to three times as species-rich as those of the Alysso-Sedion and are among the most diverse small-scale plant communities ever described. We discuss the reasons for this exceptionally high plant diversity and the peculiar species mixture in the Tortello-Sedion and compare the relationship between Alysso-Sedion and Tortello-Sedion to the situation of other Nordic syntaxa of predominantly temperate vegetation types. Our results further underline the uniqueness of Baltic alvars and their paramount importance for conservation at the European level.
The Transylvanian Plateau in Romania is well known to host large areas of a variety of dry grassland types, still traditionally managed by low-intensity mowing or grazing. While this natural heritage is now under threat from changes in agricultural practices, the diversity of Transylvanian dry grasslands is still little understood. There is a lack of both field data sampled with standardised methods and a syntaxonomic treatment with modern statistical methods and supra-regional perspective. Therefore, the European Dry Grassland Group (EDGG) carried out its first international Research Expedition in Transylvania 2009 to study syntaxonomy, vegetation-environment relationships, and biodiversity patterns of these communities. In various locations across Transylvania, we sampled 10-m² vegetation plots (n = 82) and nested-plot series from 0.0001 m² to 100 m² (n = 20), including all vascular plant, bryophyte, and lichen species, as well as structural and soil data. The vegetation classification was carried out with modified TWINSPAN, followed by determination of diagnostic species with phi values and a small-scale re-assignment of relevés with the aim of crispness maximisation. Both TWINSPAN and ordination revealed three major groups of syntaxa, which were matched to three orders from the class of basiphilous dry grasslands, Festuco-Brometea, represented by one alliance each: rocky dry grasslands (Stipo pulcherrimae-Festucetalia pallentis: Seslerion rigidae); xeric grasslands on deep soils (Festucetalia valesiacae: Stipion lessingianae) and meso-xeric grasslands on deep soils (Brachypodietalia pinnati: Cirsio-Brachypodion pinnati). We accepted nine association-level units plus two that potentially merit association status but were only represented by one relevé each. Most of the units could be identified with one or several previously described associations. To support nomenclatural stability, we provide a nomenclatural revision and designate nomenclatural types where previously there were none. Further, we used DCA ordination and analysis of variance to determine the main environmental drivers of floristic differentiation and to determine ecological and structural differences between the vegetation types. The strongest differentiation occurred along the aridity gradient with the dense, particularly diverse stands on more or less level sites on the one hand (Brachypodietalia pinnati) and the more open, less diverse stands on steep south-facing slopes on the other end of the gradient (Stipo pulcherrimae- Festucetalia pallentis, Festucetalia valesiacae). The two xeric orders were then separated along the second DCA axis, with the Stipo pulcherrimae-Festucetalia pallentis inhabiting the stone-rich sites at higher altitudes while the Festucetalia valesiacae occur on soft, deep substrata at lower altitudes. The analysed dry grassland communities have extraordinarily high -diversity at all spatial scales for all plants and for vascular plants, but are relatively poor in bryophytes and lichens. Some formerly mown stands of the Festuco sulcatae-Brachypodietum pinnati (Brachypodietalia pinnati) are even richer in vascular plant species than any other recorded vegetation type worldwide on the spatial scales of 0.1 m² (43) and 10 m² (98); the respective relevés are documented here for the first time. Also, the b-diversity of the grasslands was unexpectedly high, with a mean z-value of 0.275. Despite its limited extent, the methodological thoroughness of this study allows us to shed new light on the syntaxonomy of dry grasslands in Romania and to raise the awareness that Transylvania still hosts High Nature Value grasslands that are bio - diversity hotspots at a global scale but at the same time are highly endangered through changes in agricultural practices.
In many regions of Central Europe, semi-natural grasslands have experienced severe vegetation changes, e.g. compositional change and overall species loss, because of land use changes, atmospheric nitrogen input and also climate change. Here we analysed the vegetation change in a dry grassland complex (Gabower Hänge) in the Biosphere Reserve Schorfheide-Chorin (NE Brandenburg, Germany), one of the driest regions of the country. We resampled four 10 m² plots of each of four typical alliances (Festucion valesiacae, Koelerion glaucae, Armerion elongatae, Arrhenatherion elatioris) about 20 years after their original sampling with a recovery accuracy of approx. 10 m. The cover of vascular plants, bryophytes and lichens was recorded in both samplings. The overall compositional change was analysed with a detrended correpondence analysis (DCA). To interpret this change, we calculated unweighted mean Ellenberg indicator values for old and new plots. Furthermore we tested differences in constancy of individual species between old and new plots as well as differences in species richness, cover of herb and cryptogam layer, ecological indicator values and unweighted proportion of species groups (vascular plants, bryophytes, lichens), floristic status (native or not), life forms, CSR-strategy types and Red List species. The results of the ordination indicated no significant vegetation change, but revealed tendencies towards more nutrient-rich conditions. Ellenberg indicator values for nutrients and soil reaction were significantly correlated with the axes of the ordination. There were 28 species exclu-sively found in the new plots and 45 species of the old plots missing. While no species decreased signif-icantly, there were seven species that increased significantly. Mean species richness was significantly increased in the new plots. There were no significant differences in mean Ellenberg indicator values. Proportions of vascular plants, neophytes, hemicryptophytes and CS-strategists decreased. We conclude that overall vegetation changes are small, indicating that the dry grassland complex at the Gabower Hänge is still in a good state and of high conservation value. This relative stability over time compared to the situation in many other dry grasslands throughout temperate Europe is likely attributable to low nitrogen deposition and the dryness of the local climate. However, the detected tendency towards more nutrient-rich conditions should be taken into account in future management.
Trockenrasen (v. a. Klassen Koelerio-Corynephoretea und Festuco-Brometea) gehören zu den auf kleinen Flächen artenreichsten Pflanzengesellschaften überhaupt und stellen ideale Modellsysteme für die Analyse von Phytodiversitätsmustern und deren Ursachen dar. Ich gebe einen Überblick der in verschiedenen Trockenrasentypen Europas auftretenden Durchschnitts- und Maximalwerte der Artenzahlen von Gefäßpflanzen, Moosen und Flechten auf unterschiedlichen Flächengrößen zwischen 1 mm2 und 100 m2. Gesellschaften der Festuco-Brometea sind generell artenreicher als jene der Koelerio-Corynephoretea. Die bislang höchsten publizierten Artendichten stammen jedoch aus dem Gypsophilo fastigiatae-Globularietum vulgaris, einer basiphilen Felsgrusflur der schwedischen Insel Öland (Ordnung Alysso alyssoidis-Sedetalia, Koelerio-Corynephoretea). Sie betragen auf 4 m2 durchschnittlich 53,6 und maximal 80 Arten. Die geringsten Artendichten unter den Trockenrasen weisen das Caricetum arenariae und das Corniculario-Corynephoretum (beide Ordnung Corynephoretalia canescentis, Koelerio-Corynephoretea) mit durchschnittlich weniger als 10 Arten auf 4 m2 auf. Die Artenzahl-Areal-Beziehung von Trockenrasen lässt sich über den ganzen betrachteten Dimensionsbereich exzellent durch eine Potenzfunktion S = c • A z beschreiben. Dabei unterscheiden sich die z-Werte verschiedener Trockenrasentypen nur geringfügig und betragen im Mittel 0,21. Das führt dazu, dass die Reihung verschiedener Trockenrasengesellschaften hinsichtlich ihrer Artenzahl auf unterschiedlichen Skalenebenen nahezu unverändert bleibt, und eröffnet die Möglichkeit, Artenzahlen auf andere Flächengrößen zu extrapolieren.
Es folgt eine Erörterung der wichtigsten Faktoren, welche die Verteilung der Phytodiversität in Trockenrasen beeinflussen. Die größte Bedeutung hat hier die Bodenreaktion. Im Allgemeinen wurden stark steigende Artendichten mit zunehmendem pH-Wert, teilweise auch ein erneuter leichter Abfall oberhalb des Neutralpunktes gefunden. Eine Abnahme der Artendichte mit zunehmender Höhenlage im Gebirge konnte zumindest in zwei Fällen auch für Probeflächen gleicher Größe nachgewiesen werden. Abschließend diskutiere ich die Gründe für den überdurchschnittlichen Artenreichtum von Trockenrasen, weise auf Forschungsdefizite hin und unterbreite Empfehlungen für die Konzeption und Durchführung künftiger Studien der Biodiversitätsforschung wie auch der pflanzensoziologischen Datenerhebung allgemein. Als vordringliche Qualitätskriterien erachte ich die Arbeit mit standardisierten Probeflächengrößen und die sorgfältige Berücksichtigung von Moosen und Flechten.