Corresponding author: Martin Magnes ( martin.magnes@uni-graz.at ) Academic editor: Arkadiusz Nowak
© 2021 Martin Magnes, Wolfgang Willner, Monika Janišová, Helmut Mayrhofer, Elías Afif Khouri, Christian Berg, Anna Kuzemko, Philipp Kirschner, Riccardo Guarino, Harald Rötzer, Elena Belonovskaya, Asun Berastegi, Idoia Biurrun, Itziar García-Mijangos, Ermin Masic, Jürgen Dengler, Iwona Dembicz.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Magnes M, Willner W, Janišová M, Mayrhofer H, Afif Khouri E, Berg C, Kuzemko A, Kirschner P, Guarino R, Rötzer H, Belonovskaya E, Berastegi A, Biurrun I, García-Mijangos I, Masic E, Dengler J, Dembicz I (2021) Xeric grasslands of the inner-alpine dry valleys of Austria – new insights into syntaxonomy, diversity and ecology. Vegetation Classification and Survey 2: 133-157. https://doi.org/10.3897/VCS/2021/68594
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Aims: We studied the syntaxonomic position, biodiversity, ecological features, nature conservation value and current status of dry grasslands investigated by Josias Braun-Blanquet more than 60 years ago. Study area: Inner-alpine valleys of Austria. Methods: We sampled 67 plots of 10 m2, following the standardized EDGG methodology. We subjected our plots to an unsupervised classification with the modified TWINSPAN algorithm and interpreted the branches of the dendrogram syntaxonomically. Biodiversity, structural and ecological characteristics of the resulting vegetation units at association and order level were compared by ANOVAs. Results: All the examined grasslands belong to the class Festuco-Brometea. From ten distinguished clusters, we could assign four clusters to validly published associations, while the remaining six clusters were named tentatively. We classified them into three orders: Stipo-Festucetalia pallentis (Armerio elongatae-Potentilletum arenariae, Phleo phleoidis-Pulsatilletum nigricantis, Medicago minima-Melica ciliata community, Koelerio pyramidatae-Teucrietum montani), Festucetalia valesiacae (Sempervivum tectorum-Festuca valesiaca community); Brachypodietalia pinnati (Astragalo onobrychidis-Brometum erecti, Agrostis capillaris-Avenula adsurgens community, Anthericum ramosum-Brachypodium pinnatum community, Ranunculus bulbosus-Festuca rubra community, Carduus defloratus-Brachypodium pinnatum community). Conclusions: The ten distinguished dry grassland communities of the Austrian inner-alpine valleys differ in their ecological affinities as well as their vascular plant, bryophyte and lichen diversity. We point out their high nature conservation importance, as each of them presents a unique habitat of high value.
Taxonomic reference: Names of vascular plants, bryophytes and lichens follow
Syntaxonomic reference: Names of orders and classes follow
Abbreviations: ANOVA = analysis of variance; DCA: detrended correspondence analyses; EDGG: Eurasian Dry Grassland Group; EIV: ecological indicator value; FL: Fließ; GR: Griffen; GU: Gulsen; KA: Kaunerberg; LA: Laudegg castle in Ladis; MA: Marin; NM: Neumarkt in der Steiermark; OM: Obermauern; PÖ: Pöls; PU: Puxer Loch; TWINSPAN = Two-way indicator species analysis; ZS: Zinizachspitze.
Austria, biodiversity, Brachypodietalia pinnati, Festucetalia valesiacae, Festuco-Brometea, inner-alpine dry valley, semi-natural grassland, soil parameter, resurvey, Stipo-Festucetalia pallentis, syntaxonomy, vegetation classification
The European Alps (“Alps”) form a nearly 1,200 km long arc from Nice in the southwest to Vienna in the northeast, with the widest sector in the centre of the Eastern Alps, stretching approximately 240 km from north to south (
The inner valleys of the Alps have been inhabited by humans since the Bronze Age (
Our study is part of a long-term project of the Eurasian Dry Grassland Group (EDGG) that will cover the inner-alpine dry grasslands in the whole Alps (
We mainly sampled in three regions of Austria: the Upper Inn valley in North Tyrol, the Virgen valley in East Tyrol and the central Mur valley in Styria. Additionally, one site in the Carinthian Basin was visited (Figure
Location of the study sites: blue: Upper Inn valley (FL: Fließ, LA: Laudegg castle in Ladis, KA: Kaunerberg); green: Virgen valley (OM: Obermauern, ZS: Zinizachspitze, MA: Marin), red: Mur valley and Carinthian Basin (PU: Puxer Loch, NM: Neumarkt in der Steiermark, PÖ: Pöls, GR: Griffen, GU: Gulsen).
Climate diagrams from weather stations adjacent to the study sites (from west to east): The Upper Inn valley (Landeck), East Tyrol (Lienz, the closest ZAMG climate station to the Virgen valley), and the Mur valley (Zeltweg), based on data period 1971–2000 (
Sampling localities: Fließ (FL), Kaunerberg (KA), Laudegg castle in Ladis (LA) (Figure
The sampling was performed in the Nature Park Kaunergrat, southwest of Landeck, in one of the driest regions of the Austrian Alps. Our study area is close to the Engadin in Switzerland and the most continental area of the Eastern Alps, the Vinschgau in South Tyrol (Italy).
Geologically, the Kaunerberg (KA) and the study sites below the castle Laudegg (LA) belong to the easternmost part of the Engadin window. Kauns-Kaunerberg is located above Bündner schist, a metamorphic marine sediment from the Cretaceous period that is rich in calcium carbonate (
The studied dry grassland sites bear witness of an old cultural landscape. Traditionally, these sites were communal pastures (“Allmende”) for small livestock (mainly sheep, to a lesser amount goats) in spring and autumn, before and after summering on alpine pastures at higher elevations. This kind of management is documented back to the medieval ages, but likely is much older. Until the 1940s and early 1950s, several hundred animals were herded in the slopes below Fließ. However, this practice was abandoned alongside the economic upturn of the post-war era, leading to a reforestation of the sites. This negative trend lasted until 2002, when, following the long-lasting pressure of regional conservationists, the sites at Fließ were finally put under protection and integrated into the Natura 2000 network. The sites at Kauns-Kaunerberg followed shortly after and are protected under Tyrolean law since 2006. Since then, both sites have been managed by the Kaunergrat Nature Park. Management measures include the initial re-establishment of abandoned areas by removal of bushes and trees, and subsequent, targeted grazing by cattle, goats and sheep. Since their implementation, these activities have been regularly evaluated via monitoring studies on vegetation and selected insect taxa (butterflies, ants). The area is the best example of inner-alpine dry valleys in Austria concerning size and extent of dry and semi-dry grassland vegetation.
Sampling localities: Obermauern (OM), Marin (MA), Zinizachspitze (ZS) (Figure
The Virgen valley runs in a west-east direction south of the main ridge of the Alps. It is quite remote from modern traffic routes and accessible only from the east, following the Isel river upstream. Due to its position south of the giant mountain massif Großvenediger (3,662 m), the climate is rather dry with an average annual precipitation of about 900 mm. The nearest climate station (Lienz, 30 km to the southeast) shows sub-Mediterranean characteristics with a relatively low precipitation in late summer (Figure
The valley has been populated since prehistoric times. Triticum aestivum was cultivated until the 1980s and up to an elevation of approximately 1,400 m. Cattle breeding is another important land use. Traditionally, the animals were brought across mountain passes to markets on the northern side of the main ridge (
The montane and subalpine zones reach from the valley bottom up to more than 2,000 m. On the northern slopes, Picea abies forests are dominant. On the sunny southern slopes, the forests were replaced by grasslands centuries ago. The forest remnants are usually dominated by Larix decidua with low cover and are still used as reserve pastures for dry years. The often very steep slopes were not only grazed by cattle, sheep and goats, but also used as mountain hay meadows as farmers could not produce enough hay in the lower parts of the valley. Hay was used as forage in winter in addition to harvested leaves and straw. The mountain meadows, sometimes even above the timberline, were traditionally mown only every second year. They are situated in the neighbourhood of the mountain pastures, which are mainly used for young cattle (
The bedrock of the sampling localities is mainly calcareous slate, partly covered with moraine material, and only on the southern border of OM there is also some dark phyllite together with the calcareous slate (
Sampling localities: Puxer Loch (PU), Neumarkt in der Steiermark (NM), Pöls (PÖ), Griffen (GR), Gulsen (GU) (Figure
The studied part of the Mur valley is situated along the upper reaches of the river, approximately 80 to 120 km east of its source.
Although all sites are situated in the Austrian Central Alps, the geological bedrock is very diverse (
In total, we sampled 67 plots of 10 m2 (15 nested-plot series with two 10-m2 subplots each and additional 37 normal plots, Suppl. materials 1, 2) with the EDGG methodology (
Soil of the uppermost 10 cm was collected in each plot by mixing five random samples. The following soil parameters were measured (all measured environmental variables as header data are provided in Suppl. material 1): skeleton content (mass fraction of particles > 2 mm), percentages of sand, clay and silt, texture class estimated with Robinson pipette method after removing organic matter with 6% H2O2, field capacity (%), drainage rate (cm/h), available water (%), saturation (%), permanent wilting point (%), pH (in a suspension of 10 g dry soil in 25 ml aqua dest.), electrical conductivity (EC) (in a suspension of 10 g dry soil in 50 ml aqua dest., dS/m), organic matter (as loss at ignition at 430°C, %), organic carbon (%), N content estimated with the Kjeldahl method (%), C/N, available P (estimated with the Mehlich 3 method (PM3), mg/kg).
Other recorded environmental and structural parameters were (for methodological details, see
The complete data are stored in and available from the GrassPlot database (
The compositional data, along with the header data, were entered into TURBOVEG (
We tried cluster numbers up to 15 and finally selected the solution with 11 clusters as the basis for our classification, as it corresponded most closely to what we consider the association level. One cluster containing a single relevé was joined with another cluster, so finally ten relevé groups were distinguished. Moreover, three clusters were subdivided into regional variants. These variants partly corresponded to TWINSPAN clusters of finer resolution, but the assignment of relevés to regional variants was based on their geographical distribution.
Fidelity of species to relevé groups was calculated using the phi coefficient of association (
We tried to assign our relevé groups to phytosociological associations, alliances and orders described in the literature by comparing the species composition. If such an assignment was ambiguous, we used informal community names. We refrained from formally describing new syntaxa, as this will be the task of a future, more comprehensive revision of all dry grasslands of the Eastern Alps. To facilitate the syntaxonomic interpretation, we calculated for each relevé the total percentage cover of the diagnostic species of orders according to
To visualize the relation of the species composition of the relevé groups (and relevés) to measured or calculated factors we performed a DCA using Canoco 5 (
The number and cover of red list species were calculated in JUICE based on
The first TWINSPAN division resulted in two groups roughly corresponding to the orders Stipo-Festucetalia pallentis (clusters 1–7) and Brachypodietalia pinnati (clusters 8–11) of the class Festuco-Brometea (Figure
Therefore, we classified our relevés into ten vegetation units at association level, which in turn were grouped in the three orders (A) Stipo-Festucetalia pallentis, (B) Festucetalia valesiacae and (C) Brachypodietalia pinnati (Table
Dendrogram of the 11-cluster resolution from modified TWINSPAN analysis. Below the cluster numbers, the corresponding association/community is indicated (A.1–A.4: Stipo-Festucetalia pallentis, B.1: Festucetalia valesiacae, C.1–C.5: Brachypodietalia pinnati). For the meaning of the community codes, see text.
Stacked total percentage covers of the diagnostic species of the orders Stipo-Festucetalia pallentis (A), Festucetalia valesiacae (B) and Brachypodietalia pinnati (C) in each relevé. The numbers below the diagram are the TWINSPAN clusters. The sequence of the relevés is the same as in Figure
Abridged synoptic table of the associations, based on the 10-m² plots. For each syntaxon, the 10 species with the highest phi-values are shown plus all species with 10% or higher constancy across all plots. The number of presented vs. total species in each group is given in brackets. Values are percentage constancies. Species are sorted by decreasing fidelity within the respective syntaxon and by decreasing overall constancy in case of accompanying species. The complete synoptic and relevé table is provided in Suppl. material 2. B: bryophyte, L: lichen, V: vascular plant.
Syntaxon | All | A | B | C | A.1 | A.2 | A.3 | A.4 | B.1 | C.1 | C.2 | C.3 | C.4 | C.5 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Number of plots | 67 | 14 | 15 | 38 | 4 | 3 | 2 | 5 | 15 | 8 | 5 | 7 | 5 | 13 | |
Diagnostic species order A (3) | |||||||||||||||
Hypnum cupressiforme | B | 16 | 43 | 7 | 11 | 50 | . | 50 | 60 | 7 | . | 20 | 14 | 20 | 8 |
Geranium columbinum | V | 7 | 21 | . | 5 | . | 33 | 50 | 20 | . | 13 | 20 | . | . | . |
Echium vulgare | V | 4 | 14 | . | 3 | . | 33 | 50 | . | . | . | . | 14 | . | . |
Diagnostic species association A.1 (11 of 15) | |||||||||||||||
Alyssum montanum | V | 6 | 29 | . | . | 100 | . | . | . | . | . | . | . | . | . |
Asplenium cuneifolium | V | 6 | 29 | . | . | 100 | . | . | . | . | . | . | . | . | . |
Dorycnium germanicum | V | 6 | 29 | . | . | 100 | . | . | . | . | . | . | . | . | . |
Koeleria pyramidata var. pubiculmis | V | 6 | 29 | . | . | 100 | . | . | . | . | . | . | . | . | . |
Potentilla incana | V | 6 | 29 | . | . | 100 | . | . | . | . | . | . | . | . | . |
Festuca pallens | V | 7 | 29 | . | 3 | 100 | . | . | . | . | . | . | 14 | . | . |
Lepraria lobificans | L | 4 | 21 | . | . | 75 | . | . | . | . | . | . | . | . | . |
Erysimum sylvestre | V | 15 | 50 | 13 | 3 | 100 | . | . | 60 | 13 | 13 | . | . | . | . |
Jovibarba globifera subsp. hirta | V | 6 | 21 | . | 3 | 75 | . | . | . | . | . | 20 | . | . | . |
Notholaena marantae | V | 3 | 14 | . | . | 50 | . | . | . | . | . | . | . | . | . |
Weissia brachycarpa | B | 12 | 21 | 7 | 11 | 75 | . | . | . | 7 | 25 | . | . | . | 15 |
Diagnostic species association A.2 (10 of 13) | |||||||||||||||
Alliaria petiolata | V | 3 | 14 | . | . | . | 67 | . | . | . | . | . | . | . | . |
Myosotis ramosissima | V | 3 | 14 | . | . | . | 67 | . | . | . | . | . | . | . | . |
Origanum vulgare | V | 10 | 21 | . | 11 | . | 100 | . | . | . | . | . | 57 | . | . |
Fallopia convolvulus | V | 10 | 21 | . | 11 | . | 100 | . | . | . | . | 60 | 14 | . | . |
Stachys recta | V | 12 | 29 | . | 11 | . | 100 | 50 | . | . | 13 | . | 43 | . | . |
Scabiosa ochroleuca | V | 6 | 14 | . | 5 | . | 67 | . | . | . | . | . | 29 | . | . |
Teucrium chamaedrys | V | 19 | 21 | 13 | 21 | . | 100 | . | . | 13 | 38 | 20 | 57 | . | . |
Aster amellus | V | 12 | 14 | 27 | 5 | . | 67 | . | . | 27 | 25 | . | . | . | . |
Bromus inermis | V | 1 | 7 | . | . | . | 33 | . | . | . | . | . | . | . | . |
Galeopsis pubescens | V | 1 | 7 | . | . | . | 33 | . | . | . | . | . | . | . | . |
Diagnostic species association A.3 (10 of 15) | |||||||||||||||
Medicago minima | V | 10 | 14 | 20 | 5 | . | . | 100 | . | 20 | 25 | . | . | . | . |
Agrimonia eupatoria | V | 12 | 14 | . | 16 | . | . | 100 | . | . | 25 | . | 57 | . | . |
Amblystegium serpens | B | 1 | 7 | . | . | . | . | 50 | . | . | . | . | . | . | . |
Clinopodium nepeta agg. | V | 1 | 7 | . | . | . | . | 50 | . | . | . | . | . | . | . |
Erigeron acris | V | 1 | 7 | . | . | . | . | 50 | . | . | . | . | . | . | . |
Lapsana communis | V | 1 | 7 | . | . | . | . | 50 | . | . | . | . | . | . | . |
Silene latifolia | V | 1 | 7 | . | . | . | . | 50 | . | . | . | . | . | . | . |
Torilis arvensis | V | 1 | 7 | . | . | . | . | 50 | . | . | . | . | . | . | . |
Melica ciliata | V | 18 | 21 | 40 | 8 | . | 33 | 100 | . | 40 | 38 | . | . | . | . |
Artemisia absinthium | V | 3 | 7 | 7 | . | . | . | 50 | . | 7 | . | . | . | . | . |
Galium lucidum | V | 24 | 36 | 33 | 16 | . | 33 | 100 | 40 | 33 | 50 | . | 14 | . | 8 |
Diagnostic species association A.4 (12 of 19) | |||||||||||||||
Poa badensis agg. | V | 9 | 36 | 7 | . | . | . | . | 100 | 7 | . | . | . | . | . |
- Poa molinieri | 4 | 14 | 7 | . | . | . | . | 40 | 7 | . | . | . | . | . | |
Toninia sedifolia | L | 6 | 29 | . | . | . | . | . | 80 | . | . | . | . | . | . |
Tortella inclinata | B | 6 | 29 | . | . | . | . | . | 80 | . | . | . | . | . | . |
Ditrichum flexicaule | B | 7 | 29 | . | 3 | . | . | . | 80 | . | . | . | . | 20 | . |
Peltigera rufescens | L | 7 | 29 | . | 3 | . | . | . | 80 | . | . | . | . | 20 | . |
Saxifraga paniculata | V | 4 | 21 | . | . | . | . | . | 60 | . | . | . | . | . | . |
Psora decipiens | L | 6 | 21 | 7 | . | . | . | . | 60 | 7 | . | . | . | . | . |
Cladonia symphycarpa | L | 15 | 29 | 40 | . | . | . | . | 80 | 40 | . | . | . | . | . |
Physconia muscigena | L | 7 | 21 | 13 | . | . | . | . | 60 | 13 | . | . | . | . | . |
Cladonia macrophyllodes | L | 3 | 14 | . | . | . | . | . | 40 | . | . | . | . | . | . |
Endocarpon pusillum | B | 10 | 21 | 27 | . | . | . | . | 60 | 27 | . | . | . | . | . |
Tortella tortuosa | B | 22 | 29 | 40 | 13 | . | . | . | 80 | 40 | . | . | . | 20 | 31 |
Diagnostic species order B (21 of 35) | |||||||||||||||
Festuca valesiaca | V | 33 | 14 | 93 | 16 | . | . | 50 | 20 | 93 | 38 | . | . | 20 | 15 |
Sempervivum tectorum | V | 16 | . | 67 | 3 | . | . | . | . | 67 | 13 | . | . | . | . |
Veronica spicata | V | 19 | . | 67 | 8 | . | . | . | . | 67 | 25 | . | . | 20 | . |
Artemisia campestris | V | 39 | 50 | 100 | 11 | . | 67 | . | 100 | 100 | 50 | . | . | . | . |
Koeleria macrantha | V | 22 | 14 | 73 | 5 | 25 | 33 | . | . | 73 | 25 | . | . | . | . |
Dianthus sylvestris | V | 33 | 36 | 87 | 11 | . | . | . | 100 | 87 | 25 | . | . | 40 | . |
Galium verum | V | 31 | 7 | 73 | 24 | . | . | 50 | . | 73 | 75 | . | . | 60 | . |
Bromus erectus | V | 39 | 7 | 80 | 34 | . | . | 50 | . | 80 | 100 | . | 14 | 40 | 15 |
Parmelia somloensis | L | 13 | . | 47 | 5 | . | . | . | . | 47 | . | 40 | . | . | . |
Phleum phleoides | V | 31 | 14 | 73 | 21 | . | . | 50 | 20 | 73 | 63 | 40 | . | 20 | . |
Tortula ruralis | B | 19 | 14 | 60 | 5 | . | . | . | 40 | 60 | 13 | 20 | . | . | . |
Potentilla pusilla | V | 48 | 43 | 93 | 32 | . | . | 50 | 100 | 93 | 63 | . | . | 60 | 31 |
Petrorhagia saxifraga | V | 24 | 21 | 67 | 8 | . | . | . | 60 | 67 | 38 | . | . | . | . |
Sempervivum arachnoideum | V | 19 | 21 | 60 | 3 | . | . | . | 60 | 60 | . | . | . | 20 | . |
Vincetoxicum hirundinaria | V | 45 | 14 | 80 | 42 | . | 67 | . | . | 80 | 100 | 20 | 86 | 20 | . |
Sedum album | V | 28 | 43 | 73 | 5 | . | 33 | . | 100 | 73 | 25 | . | . | . | . |
Festuca guestfalica | V | 15 | . | 40 | 11 | . | . | . | . | 40 | 38 | . | . | 20 | . |
Sedum sexangulare | V | 21 | 21 | 53 | 8 | . | . | . | 60 | 53 | 13 | . | . | 40 | . |
Hedwigia ciliata | L | 16 | 7 | 40 | 11 | . | . | 50 | . | 40 | . | 60 | . | 20 | . |
Cladonia pyxidata | L | 18 | 29 | 47 | 3 | . | . | . | 80 | 47 | . | . | . | 20 | . |
Scabiosa columbaria agg. | V | 28 | 36 | 53 | 16 | . | . | 50 | 80 | 53 | 25 | 20 | 14 | . | 15 |
- Scabiosa columbaria | 13 | 29 | 13 | 8 | . | . | . | 80 | 13 | . | 20 | 14 | . | 8 | |
- Scabiosa lucida | 1 | . | . | 3 | . | . | . | . | . | . | . | . | . | 8 | |
- Scabiosa triandra | 13 | 7 | 40 | 5 | . | . | 50 | . | 40 | 25 | . | . | . | . | |
Diagnostic species order C (22 of 28) | |||||||||||||||
Leontodon hispidus | V | 30 | . | . | 53 | . | . | . | . | . | 13 | . | 29 | 80 | 100 |
Achillea millefolium agg. | V | 42 | 14 | . | 68 | 25 | . | 50 | . | . | . | 60 | 100 | 100 | 85 |
Lotus corniculatus | V | 42 | . | 20 | 66 | . | . | . | . | 20 | 63 | . | 43 | 80 | 100 |
Dactylis glomerata | V | 33 | . | 13 | 53 | . | . | . | . | 13 | 50 | 20 | 86 | 40 | 54 |
Veronica chamaedrys | V | 18 | . | . | 32 | . | . | . | . | . | . | 60 | 43 | 80 | 15 |
Pimpinella saxifraga | V | 30 | 14 | . | 47 | . | . | 50 | 20 | . | . | 20 | 71 | 60 | 69 |
- Pimpinella saxifraga subsp. nigra | 6 | . | . | 11 | . | . | . | . | . | . | . | 43 | . | 8 | |
Anthoxanthum odoratum agg. | V | 16 | . | . | 29 | . | . | . | . | . | . | 20 | 14 | 60 | 46 |
Campanula glomerata | V | 15 | . | . | 26 | . | . | . | . | . | 13 | . | 43 | . | 46 |
Cerastium holosteoides | V | 12 | . | . | 21 | . | . | . | . | . | . | . | 14 | 40 | 38 |
Trifolium repens | V | 12 | . | . | 21 | . | . | . | . | . | 13 | 20 | 14 | 40 | 23 |
Carex ornithopoda | V | 10 | . | . | 18 | . | . | . | . | . | . | . | . | 40 | 38 |
Centaurea jacea | V | 10 | . | . | 18 | . | . | . | . | . | . | . | 14 | 20 | 38 |
Galium pumilum | V | 10 | . | . | 18 | . | . | . | . | . | . | 20 | 29 | 20 | 23 |
Larix decidua | V | 10 | . | . | 18 | . | . | . | . | . | . | . | . | 40 | 38 |
Brachypodium pinnatum agg. | V | 54 | 21 | 47 | 68 | . | 67 | 50 | . | 47 | 75 | . | 86 | 60 | 85 |
- Brachypodium pinnatum | 31 | 14 | . | 50 | . | 67 | . | . | . | . | . | 71 | 60 | 85 | |
- Brachypodium rupestre | 21 | 7 | 47 | 16 | . | . | 50 | . | 47 | 75 | . | . | . | . | |
Polygonatum odoratum | V | 15 | . | 7 | 24 | . | . | . | . | 7 | 13 | 40 | . | . | 46 |
Fragaria vesca | V | 15 | 7 | . | 24 | . | . | 50 | . | . | 13 | 20 | 57 | 60 | . |
Galium album | V | 25 | 21 | . | 37 | . | 67 | 50 | . | . | . | 100 | 71 | 40 | 15 |
Koeleria pyramidata var. pyramidata | V | 48 | 36 | 27 | 61 | . | . | . | 100 | 27 | 13 | 100 | 71 | 60 | 69 |
Seseli libanotis | V | 18 | 7 | 7 | 26 | . | . | 50 | . | 7 | 38 | . | 43 | . | 31 |
Brachythecium rutabulum | B | 12 | 7 | . | 18 | . | . | 50 | . | . | 13 | 20 | 14 | 20 | 23 |
Poa pratensis agg. | V | 34 | 21 | 20 | 45 | . | 33 | 100 | . | 20 | 75 | 40 | 57 | 40 | 23 |
Diagnostic species association C.1 (2) | |||||||||||||||
Fragaria viridis | V | 3 | . | . | 5 | . | . | . | . | . | 25 | . | . | . | . |
Astragalus onobrychis | V | 9 | . | 20 | 8 | . | . | . | . | 20 | 38 | . | . | . | . |
Diagnostic species association C.2 (12 of 16) | |||||||||||||||
Potentilla argentea | V | 10 | . | 13 | 13 | . | . | . | . | 13 | . | 100 | . | . | . |
Allium carinatum | V | 10 | . | 7 | 16 | . | . | . | . | 7 | . | 100 | 14 | . | . |
Genista tinctoria | V | 4 | . | . | 8 | . | . | . | . | . | . | 60 | . | . | . |
Jasione montana | V | 4 | . | . | 8 | . | . | . | . | . | . | 60 | . | . | . |
Populus tremula | V | 4 | . | . | 8 | . | . | . | . | . | . | 60 | . | . | . |
Potentilla rupestris | V | 4 | . | . | 8 | . | . | . | . | . | . | 60 | . | . | . |
Avenula adsurgens subsp. adsurgens | V | 15 | 7 | . | 24 | . | 33 | . | . | . | . | 100 | 43 | . | 8 |
Campanula rotundifolia | V | 10 | . | 7 | 16 | . | . | . | . | 7 | . | 80 | 14 | 20 | . |
Viola arvensis | V | 6 | . | . | 11 | . | . | . | . | . | . | 60 | 14 | . | . |
Ceratodon purpureus | B | 7 | . | 7 | 11 | . | . | . | . | 7 | 13 | 60 | . | . | . |
Silene nutans | V | 15 | . | . | 26 | . | . | . | . | . | . | 80 | 14 | 20 | 31 |
Cladonia fimbriata | B | 12 | 7 | 13 | 13 | 25 | . | . | . | 13 | . | 60 | . | 20 | 8 |
Diagnostic species association C.3 (10 of 18) | |||||||||||||||
Anthericum ramosum | V | 7 | . | . | 13 | . | . | . | . | . | . | . | 71 | . | . |
Campanula trachelium | V | 7 | . | . | 13 | . | . | . | . | . | . | . | 71 | . | . |
Centaurea scabiosa | V | 7 | . | . | 13 | . | . | . | . | . | . | . | 71 | . | . |
Tragopogon orientalis | V | 7 | . | . | 13 | . | . | . | . | . | . | . | 71 | . | . |
Peucedanum oreoselinum | V | 13 | . | 20 | 16 | . | . | . | . | 20 | 13 | . | 71 | . | . |
Astragalus glycyphyllos | V | 9 | 7 | . | 13 | . | 33 | . | . | . | . | . | 71 | . | . |
Polygala comosa | V | 4 | . | . | 8 | . | . | . | . | . | . | . | 43 | . | . |
Clinopodium vulgare | V | 13 | 7 | . | 21 | . | . | 50 | . | . | 13 | 20 | 86 | . | . |
Viola hirta | V | 16 | 7 | . | 26 | . | . | 50 | . | . | 38 | . | 86 | . | 8 |
Vicia cracca agg. | V | 10 | . | . | 18 | . | . | . | . | . | . | . | 57 | 20 | 15 |
Diagnostic species association C.4 (11 of 19) | |||||||||||||||
Ranunculus bulbosus | V | 9 | . | . | 16 | . | . | . | . | . | . | . | 14 | 100 | . |
Festuca rubra agg. | V | 12 | . | . | 21 | . | . | . | . | . | . | . | . | 100 | 23 |
- Festuca rubra | 6 | . | . | 11 | . | . | . | . | . | . | . | . | 60 | 8 | |
Knautia arvensis | V | 4 | . | . | 8 | . | . | . | . | . | . | . | . | 60 | . |
Lolium perenne | V | 4 | . | . | 8 | . | . | . | . | . | . | . | . | 60 | . |
Veronica officinalis | V | 4 | . | . | 8 | . | . | . | . | . | . | . | . | 60 | . |
Calliergonella cuspidata | B | 7 | . | . | 13 | . | . | . | . | . | . | . | . | 60 | 15 |
Ranunculus acris | V | 7 | . | . | 13 | . | . | . | . | . | . | . | . | 60 | 15 |
Thuidium philibertii | B | 7 | . | . | 13 | . | . | . | . | . | . | . | . | 60 | 15 |
Hieracium pilosella | V | 25 | . | 27 | 34 | . | . | . | . | 27 | . | 40 | . | 100 | 46 |
Avenula pratensis | V | 3 | . | . | 5 | . | . | . | . | . | . | . | . | 40 | . |
Plantago lanceolata | V | 25 | 7 | 20 | 34 | . | . | 50 | . | 20 | 38 | . | 43 | 100 | 15 |
Diagnostic species association C.5 (21 of 55) | |||||||||||||||
Persicaria vivipara | V | 13 | . | . | 24 | . | . | . | . | . | . | . | . | . | 69 |
Potentilla erecta | V | 24 | . | . | 42 | . | . | . | . | . | . | . | . | 60 | 100 |
Phyteuma orbiculare | V | 10 | . | . | 18 | . | . | . | . | . | . | . | . | . | 54 |
Plantago atrata | V | 10 | . | . | 18 | . | . | . | . | . | . | . | . | . | 54 |
Ranunculus nemorosus | V | 10 | . | . | 18 | . | . | . | . | . | . | . | . | . | 54 |
Thesium alpinum | V | 15 | . | . | 26 | . | . | . | . | . | . | . | . | 20 | 69 |
Centaurea pseudophrygia | V | 9 | . | . | 16 | . | . | . | . | . | . | . | . | . | 46 |
Gymnadenia conopsea | V | 9 | . | . | 16 | . | . | . | . | . | . | . | . | . | 46 |
Laserpitium latifolium | V | 9 | . | . | 16 | . | . | . | . | . | . | . | . | . | 46 |
Molinia caerulea | V | 9 | . | . | 16 | . | . | . | . | . | . | . | . | . | 46 |
Pimpinella major | V | 13 | . | . | 24 | . | . | . | . | . | . | . | . | 20 | 62 |
Polygala chamaebuxus | V | 13 | . | . | 24 | . | . | . | . | . | . | . | . | 20 | 62 |
Carlina acaulis | V | 27 | . | 7 | 45 | . | . | . | . | 7 | . | . | 29 | 60 | 92 |
Euphrasia officinalis | V | 16 | . | . | 29 | . | . | . | . | . | . | . | . | 40 | 69 |
Galium anisophyllon | V | 16 | . | . | 29 | . | . | . | . | . | . | . | . | 40 | 69 |
Carduus defloratus | V | 19 | 7 | . | 32 | 25 | . | . | . | . | 13 | . | . | 20 | 77 |
Trifolium montanum | V | 33 | . | 13 | 53 | . | . | . | . | 13 | 13 | . | 43 | 60 | 100 |
Campanula scheuchzeri | V | 12 | . | . | 21 | . | . | . | . | . | . | . | . | 20 | 54 |
Sesleria caerulea | V | 12 | . | . | 21 | . | . | . | . | . | . | . | . | 20 | 54 |
Anthyllis vulneraria | V | 19 | 14 | . | 29 | . | . | . | 40 | . | . | . | 14 | 20 | 69 |
Hippocrepis comosa | V | 12 | . | . | 21 | . | . | . | . | . | 13 | . | . | 20 | 46 |
Diagnostic species for more than one order (3) | |||||||||||||||
Carex caryophyllea | V | 52 | 7 | 60 | 66 | . | . | . | 20 | 60 | 63 | 100 | 29 | 80 | 69 |
Allium lusitanicum | V | 42 | 71 | 73 | 18 | 50 | 100 | . | 100 | 73 | 13 | . | 14 | . | 38 |
Thymus praecox | V | 33 | 50 | 67 | 13 | 100 | . | 50 | 40 | 67 | 50 | . | 14 | . | . |
Diagnostic species for more than one association (18 of 25) | |||||||||||||||
Thymus pulegioides | V | 37 | 29 | 20 | 47 | . | . | . | 80 | 20 | 25 | . | . | 80 | 92 |
Briza media | V | 33 | . | 13 | 53 | . | . | . | . | 13 | . | . | 71 | 60 | 92 |
Salvia pratensis | V | 31 | 7 | 60 | 29 | . | . | 50 | . | 60 | 63 | . | 86 | . | . |
Plantago media | V | 25 | . | 7 | 42 | . | . | . | . | 7 | 38 | . | . | 60 | 77 |
Teucrium montanum | V | 24 | 29 | 33 | 18 | . | . | . | 80 | 33 | 88 | . | . | . | . |
Agrostis capillaris | V | 22 | . | . | 39 | . | . | . | . | . | . | 100 | . | 80 | 46 |
Trifolium pratense | V | 22 | 7 | . | 37 | . | . | 50 | . | . | . | . | 14 | 100 | 62 |
Leucanthemum vulgare agg. | V | 21 | . | . | 37 | . | . | . | . | . | . | . | . | 80 | 77 |
Linum catharticum | V | 21 | . | . | 37 | . | . | . | . | . | . | . | . | 80 | 77 |
Verbascum chaixii subsp. austriacum | V | 21 | 36 | . | 24 | 50 | 100 | . | . | . | . | 80 | 57 | . | 8 |
Arrhenatherum elatius | V | 18 | . | 7 | 29 | . | . | . | . | 7 | . | 60 | 71 | 20 | 15 |
Taraxacum sect. Ruderalia | V | 16 | . | . | 29 | . | . | . | . | . | . | . | 71 | 60 | 23 |
Primula veris | V | 15 | . | . | 26 | . | . | . | . | . | . | . | 57 | . | 46 |
Hylotelephium maximum | V | 13 | 29 | . | 13 | 75 | 33 | . | . | . | . | 100 | . | . | . |
Genista sagittalis | V | 12 | . | . | 21 | . | . | . | . | . | . | 80 | 57 | . | . |
Homalotrichon (Avenula) pubescens | V | 12 | . | . | 21 | . | . | . | . | . | . | 60 | 43 | . | 15 |
Prunella vulgaris | V | 12 | . | . | 21 | . | . | . | . | . | . | . | . | 60 | 38 |
Trifolium arvense | V | 12 | 7 | 27 | 8 | . | . | 50 | . | 27 | . | 60 | . | . | . |
Other species (22 of 209) | |||||||||||||||
Abietinella abietina | B | 57 | 43 | 53 | 63 | . | 33 | . | 100 | 53 | 63 | 100 | . | 80 | 77 |
Euphorbia cyparissias | V | 57 | 36 | 60 | 63 | 50 | . | 50 | 40 | 60 | 63 | 100 | 71 | 60 | 46 |
Carex humilis | V | 55 | 57 | 73 | 47 | 75 | 100 | 100 | . | 73 | 88 | 20 | 29 | . | 62 |
Helianthemum nummularium subsp. obscurum | V | 52 | 36 | 47 | 61 | . | 33 | . | 80 | 47 | 63 | 20 | 71 | 60 | 69 |
Festuca rupicola | V | 48 | 50 | 27 | 55 | . | 100 | . | 80 | 27 | 38 | 100 | 86 | 60 | 31 |
Rhytidium rugosum | B | 46 | 43 | 53 | 45 | 50 | . | . | 80 | 53 | 25 | 80 | . | 40 | 69 |
Medicago falcata | V | 28 | 21 | 33 | 29 | . | 33 | 50 | 20 | 33 | 88 | . | 57 | . | . |
Bryum argenteum | B | 18 | 21 | 27 | 13 | . | . | . | 60 | 27 | 13 | 60 | . | 20 | . |
Hypericum perforatum | V | 16 | 14 | . | 24 | 25 | 33 | . | . | . | . | 40 | 71 | 40 | . |
Sanguisorba minor | V | 16 | 7 | 13 | 21 | . | 33 | . | . | 13 | 38 | . | 43 | 40 | . |
Cuscuta epithymum | V | 15 | 14 | 7 | 18 | . | 67 | . | . | 7 | 38 | 20 | 43 | . | . |
Arenaria serpyllifolia agg. | V | 13 | 29 | 20 | 5 | 50 | 33 | . | 20 | 20 | . | . | 14 | 20 | . |
Asperula cynanchica | V | 13 | 14 | 7 | 16 | 50 | . | . | . | 7 | 38 | . | 29 | 20 | . |
Medicago lupulina | V | 13 | 14 | . | 18 | . | 33 | . | 20 | . | 13 | . | 29 | 20 | 23 |
Rosa canina agg. | V | 12 | 14 | 7 | 13 | . | 33 | 50 | . | 7 | 25 | 20 | 29 | . | . |
Securigera varia | V | 12 | 14 | 13 | 11 | . | 33 | 50 | . | 13 | 50 | . | . | . | . |
Cladonia furcata | L | 10 | . | 13 | 13 | . | . | . | . | 13 | . | 20 | . | . | 31 |
Clinopodium alpinum | V | 10 | 7 | . | 16 | . | . | . | 20 | . | . | . | . | 40 | 31 |
Didymodon rigidulus | B | 10 | 14 | 27 | 3 | . | . | . | 40 | 27 | . | . | . | 20 | . |
Leucodon sciuroides | B | 10 | 14 | 27 | 3 | . | . | 50 | 20 | 27 | 13 | . | . | . | . |
Prunus spinosa | V | 10 | 7 | . | 16 | . | . | 50 | . | . | 25 | 40 | 29 | . | . |
Sedum acre | V | 10 | 21 | 20 | 3 | 25 | . | . | 40 | 20 | 13 | . | . | . | . |
The first axis of the DCA graph (Figure
DCA of the 10-m2 plots sampled during the 11th EDGG Field Workshop (gradient length/eigenvalue/cumulative explained variation of Axis 1: 6.12/0.646/8.56, Axis 2: 3.70/0.4086/13,97). A.1–C.5: clusters according to the order/association code in Table
Maximum species richness for all taxa increased from 9 in 0.0001 m² to 103 in 100 m² (Table
While we found the highest mean species richness of vascular plants in the mountain meadows of C.5 (58.5 species in 10 m²), the record (87 species on 10 m2) occurred in a plot of C.4 with shallow, acidic soil, grazed by cattle, sheep and goats in Fließ (Suppl. material 1). The highest cryptogam species richness was in a stand of A.4, a historically and recently grazed hill on calcareous slate in the Virgen valley in East Tyrol (Suppl. material 1). Concerning red list species, the non-managed steep grasslands on serpentinite (cluster A.1) had the highest values (Suppl. material 1), as many of the species are edaphic specialists with a restricted distribution.
Scale-dependent species richness of the studied nested-plot series. We provide maximum values across all 15 nested-plot series and means ± standard deviation for each of the three distinguished orders. Number of samples is given in brackets (first number for grain sizes up to 10 m², second number for 100 m²).
Area [m²] | All | Order | ||
---|---|---|---|---|
Max. | A (n = 7/3) | B (n = 9/4) | C (n = 14/6) | |
All taxa | ||||
0.0001 | 9 | 2.0 ± 2.0 | 2.1 ± 1.1 | 4.0 ± 1.9 |
0.001 | 11 | 3.0 ± 2.2 | 3.2 ± 1.6 | 6.6 ± 2.6 |
0.01 | 19 | 4.9 ± 2.4 | 5.6 ± 2.6 | 11.2 ± 3.9 |
0.1 | 36 | 11.6 ± 3.0 | 12.7 ± 3.2 | 22.3 ± 8.7 |
1 | 54 | 19.6 ± 7.4 | 25.3 ± 6.2 | 33.6 ± 11.5 |
10 | 71 | 29.9 ± 7.7 | 36.1 ± 5.4 | 51.2 ± 13.4 |
100 | 103 | 46.0 ± 13.9 | 60.8 ± 12.4 | 84.5 ± 14.3 |
Vascular plants | ||||
0.0001 | 8 | 1.3 ± 1.1 | 1.2 ± 0.8 | 3.8 ± 1.7 |
0.001 | 11 | 1.6 ± 1.5 | 2.1 ± 1.1 | 6.2 ± 2.3 |
0.01 | 18 | 2.4 ± 2.4 | 3.9 ± 2.0 | 10.6 ± 3.7 |
0.1 | 33 | 7.0 ± 1.9 | 9.2 ± 2.9 | 21.6 ± 8.2 |
1 | 49 | 14.7 ± 7.2 | 18.6 ± 3.8 | 31.3 ± 11.4 |
10 | 66 | 20.0 ± 4.6 | 26.9 ± 4.0 | 46.6 ± 14.8 |
100 | 94 | 30.0 ± 3.6 | 43.0 ± 8.6 | 75.7 ± 17.9 |
Bryophytes | ||||
0.0001 | 3 | 0.6 ± 1.1 | 0.8 ± 1.1 | 0.2 ± 0.6 |
0.001 | 3 | 1.1 ± 1.3 | 0.9 ± 1.3 | 0.3 ± 0.6 |
0.01 | 5 | 1.9 ± 2.0 | 1.3 ± 1.9 | 0.5 ± 0.7 |
0.1 | 7 | 3.7 ± 2.5 | 2.3 ± 2.3 | 1.1 ± 1.0 |
1 | 8 | 4.0 ± 2.7 | 3.8 ± 1.9 | 2.1 ± 1.5 |
10 | 14 | 6.6 ± 3.9 | 5.6 ± 2.1 | 3.5 ± 1.9 |
100 | 17 | 10.3 ± 5.9 | 9.0 ± 2.2 | 6.2 ± 1.7 |
Lichens | ||||
0.0001 | 1 | 0.1 ± 0.4 | 0.1 ± 0.3 | 0.0 ± 0.0 |
0.001 | 2 | 0.3 ± 0.8 | 0.2 ± 0.4 | 0.1 ± 0.3 |
0.01 | 2 | 0.6 ± 1.0 | 0.3 ± 0.5 | 0.1 ± 0.3 |
0.1 | 3 | 0.9 ± 1.2 | 1.1 ± 1.3 | 0.1 ± 0.3 |
1 | 9 | 1.9 ± 2.7 | 3.0 ± 2.4 | 0.1 ± 0.4 |
10 | 9 | 3.3 ± 3.7 | 3.7 ± 2.1 | 0.6 ± 1.2 |
100 | 17 | 5.7 ± 4.7 | 8.8 ± 5.6 | 2.7 ± 5.6 |
Biodiversity, structural and ecological characteristics of all 10 m2 plots across the three orders. The p-values and significance levels refer to ANOVAs.
Parameter | All | Order A | Order B | Order C | p-value | Signifi-cance | ||
---|---|---|---|---|---|---|---|---|
(n = 67) | (n = 14) | (n = 15) | (n = 38) | |||||
Mean | Min. | Max. | Mean ± SD | Mean ± SD | Mean ± SD | |||
Species richness | ||||||||
Species richness (all taxa) | 40.4 | 18 | 87 | 28.8 ± 6.9 | 34.4 ± 5.6 | 47.0 ± 15.8 | <0.001 | *** |
Species richness (vascular plants) | 34.2 | 16 | 71 | 20.9 ± 4.2 | 26.1 ± 3.7 | 42.4 ± 14.6 | <0.001 | *** |
Species richness (bryophytes + lichens) | 6.1 | 0 | 23 | 7.9 ± 7.9 | 8.3 ± 3.4 | 4.6 ± 4.1 | 0.021 | * |
Species richness (red-listed species) | 2.5 | 0 | 9 | 2.9 ± 2.5 | 3.1 ± 1.5 | 2.1 ± 1.6 | 0.109 | |
Vegetation structure | ||||||||
Cover vegetation [%] | 78 | 30 | 100 | 64 ± 15 | 58 ± 18 | 90 ± 15 | <0.001 | *** |
Cover herb layer [%] | 72 | 20 | 100 | 58 ± 22 | 47 ± 16 | 87 ± 18 | <0.001 | *** |
Cover bryophyte + lichen layer [%] | 13 | 0 | 60 | 14 ± 20 | 18 ± 20 | 11 ± 15 | 0.423 | |
Cover litter [%] | 29 | 0 | 95 | 22 ± 23 | 21 ± 16 | 35 ± 28 | 0.091 | |
Herb layer height mean [cm] | 14.1 | 1.4 | 62.6 | 11.5 ± 9.5 | 8.4 ± 5.6 | 17.3 ± 12.5 | 0.018 | * |
Herb layer height SD [cm] | 6.8 | 0.8 | 33.4 | 8.3 ± 9.0 | 5.6 ± 6.0 | 6.7 ± 6.4 | 0.564 | |
Ecological indicator values (not weighted by cover) | ||||||||
EIV light | 7.5 | 6.8 | 8.1 | 7.7 ± 0.3 | 7.8 ± 0.1 | 7.3 ± 0.2 | <0.001 | *** |
EIV temperature | 5.4 | 3.2 | 6.2 | 5.8 ± 0.3 | 5.9 ± 0.1 | 5.1 ± 0.9 | <0.001 | *** |
EIV continentality | 4.2 | 3.5 | 5.0 | 4.4 ± 0.3 | 4.5 ± 0.2 | 4.1 ± 0.3 | <0.001 | *** |
EIV moisture | 3.4 | 2.6 | 4.9 | 3.0 ± 0.3 | 2.8 ± 0.1 | 3.9 ± 0.6 | <0.001 | *** |
EIV reaction | 6.9 | 5.4 | 8.1 | 7.3 ± 0.4 | 6.8 ± 0.4 | 6.9 ± 0.7 | 0.031 | * |
EIV nutrients | 2.7 | 1.6 | 4.6 | 2.3 ± 0.7 | 2.2 ± 0.2 | 3.1 ± 0.6 | <0.001 | *** |
Topography | ||||||||
Elevation [m] | 1180 | 549 | 1945 | 1078 ± 274 | 1114 ± 120 | 1243 ± 390 | 0.191 | |
Inclination [°] | 36 | 8 | 59 | 40 ± 5 | 38 ± 7 | 33 ± 10 | 0.017 | * |
Southing | 0.76 | -0.97 | 1.00 | 0.79 ± 0.48 | 0.86 ± 0.31 | 0.71 ± 0.46 | 0.518 | |
Heat index | 0.36 | -0.98 | 1.19 | 0.36 ± 0.48 | 0.42 ± 0.38 | 0.34 ± 0.38 | 0.820 | |
Maximum microrelief [cm] | 15 | 2 | 105 | 20 ± 12 | 23 ± 25 | 11 ± 7 | 0.011 | * |
Soil parameters [general] | ||||||||
Cover rocks and stones [%] | 18 | 0 | 85 | 37 ± 25 | 34 ± 28 | 5 ± 11 | <0.001 | *** |
Cover gravel [%] | 7 | 0 | 50 | 16 ± 14 | 8 ± 11 | 3 ± 10 | 0.002 | ** |
Cover fine soil [%] | 73 | 1 | 100 | 41 ± 34 | 62 ± 29 | 89 ± 23 | <0.001 | *** |
Soil depth mean [cm] | 17 | 1 | 56 | 11 ± 6 | 9 ± 5 | 22 ± 15 | <0.001 | *** |
Soil depth CV | 58 | 12 | 225 | 73 ± 41 | 89 ± 66 | 40 ± 21 | <0.001 | *** |
Soil parameters [physical] | ||||||||
Skeleton content [%] | 31 | 0 | 77 | 44 ± 18 | 26 ± 8 | 28 ± 12 | <0.001 | *** |
Sand content [%] | 70 | 48 | 81 | 70 ± 7 | 74 ± 3 | 68 ± 9 | 0.036 | * |
Silt content [%] | 14 | 6 | 31 | 14 ± 5 | 12 ± 3 | 15 ± 6 | 0.170 | |
Clay content [%] | 16 | 11 | 29 | 16 ± 4 | 14 ± 1 | 17 ± 4 | 0.060 | |
Water at saturation [%] | 42.7 | 4.7 | 48.3 | 43.3 ± 1.5 | 42.4 ± 0.6 | 42.6 ± 6.6 | 0.863 | |
Water at field capacity [%] | 20.5 | 17.9 | 27.7 | 20.5 ± 1.9 | 19.3 ± 0.7 | 21.1 ± 2.5 | 0.028 | * |
Water at permanent wilting point [%] | 11.3 | 9.4 | 16.7 | 11.3 ± 1.5 | 10.5 ± 0.5 | 11.6 ± 1.8 | 0.063 | |
Plant-available water [%] | 9.2 | 8.1 | 12.2 | 9.2 ± 0.7 | 8.8 ± 0.4 | 9.4 ± 1.0 | 0.080 | |
Drainage rate [cm/h] | 1.3 | 0.3 | 2.2 | 1.3 ± 0.5 | 1.6 ± 0.3 | 1.2 ± 0.5 | 0.053 | |
Soil parameters [chemical] | ||||||||
pH [in aqua dest.] | 6.87 | 4.34 | 8.34 | 7.37 ± 0.56 | 6.87 ± 0.55 | 6.67 ± 1.02 | 0.040 | * |
Electrical conductivity [µS/cm] | 148 | 25 | 511 | 214 ± 138 | 90 ± 53 | 147 ± 110 | 0.011 | * |
Organic matter [%] | 14.2 | 6.0 | 32.3 | 16.4 ± 7.5 | 11.8 ± 5.1 | 14.4 ± 7.2 | 0.209 | |
Organic carbon [%] | 8.2 | 3.5 | 18.8 | 9.5 ± 4.3 | 6.8 ± 3.0 | 8.3 ± 4.2 | 0.212 | |
N content [%] | 1.4 | 0.1 | 4.6 | 2.0 ± 0.9 | 1.0 ± 0.6 | 1.4 ± 0.8 | 0.004 | ** |
C/N ratio | 7.9 | 2.8 | 61.8 | 4.8 ± 1.7 | 9.0 ± 6.8 | 8.6 ± 10.6 | 0.326 | |
P available [ppm] | 18.5 | 8.8 | 47.3 | 23.1 ± 12.6 | 17.5 ± 5.0 | 17.2 ± 8.9 | 0.120 |
Biodiversity, structural and ecological characteristics across the 10 association-level communities. The p-values and significance levels refer to ANOVAs.
Parameter | A.1 | A.2 | A.3 | A.4 | B.1 | C.1 | C.2 | C.3 | C.4 | C.5 | p-value | |
---|---|---|---|---|---|---|---|---|---|---|---|---|
(n = 4) | (n = 3) | (n = 2) | (n = 5) | (n = 15) | (n = 8) | (n = 5) | (n = 7) | (n = 5) | (n = 13) | |||
Species richness | ||||||||||||
Species richness (all taxa) | 22.3 | 25.0 | 28.5 | 36.4 | 34.4 | 29.3 | 38.4 | 43.7 | 58.4 | 58.5 | <0.001 | *** |
Species richness (vascular plants) | 17.5 | 24.7 | 24.5 | 19.8 | 26.1 | 26.6 | 30.6 | 42.4 | 48.8 | 54.1 | <0.001 | *** |
Species richness (bryophytes + lichens) | 4.8 | 0.3 | 4.0 | 16.6 | 8.3 | 2.6 | 7.8 | 1.3 | 9.6 | 4.5 | <0.001 | *** |
Species richness (red-listed species) | 6.0 | 2.7 | 2.0 | 1.0 | 3.1 | 3.1 | 3.2 | 2.1 | 1.2 | 1.2 | <0.001 | *** |
Vegetation structure | ||||||||||||
Cover vegetation [%] | 63 | 82 | 70 | 53 | 58 | 76 | 86 | 98 | 88 | 98 | <0.001 | *** |
Cover herb layer [%] | 62 | 82 | 70 | 37 | 47 | 73 | 83 | 98 | 73 | 96 | <0.001 | *** |
Cover bryophyte + lichen layer [%] | 4 | 0 | 2 | 37 | 18 | 5 | 5 | 1 | 29 | 15 | 0.001 | *** |
Cover litter [%] | 8 | 43 | 55 | 8 | 21 | 48 | 40 | 56 | 19 | 20 | <0.001 | *** |
Herb layer height mean [cm] | 13.9 | 16.7 | 10.3 | 7.0 | 8.4 | 10.4 | 22.8 | 32.6 | 11.9 | 13.4 | <0.001 | *** |
Herb layer height SD [cm] | 12.0 | 7.2 | 6.1 | 7.0 | 5.6 | 3.3 | 17.4 | 9.5 | 4.0 | 4.1 | 0.008 | ** |
Ecological indicator values | ||||||||||||
EIV light | 7.7 | 7.5 | 7.5 | 8.0 | 7.8 | 7.5 | 7.3 | 7.1 | 7.1 | 7.3 | <0.001 | *** |
EIV temperature | 6.0 | 6.0 | 6.0 | 5.5 | 5.9 | 5.8 | 5.7 | 5.6 | 5.2 | 4.0 | <0.001 | *** |
EIV continentality | 4.6 | 4.6 | 4.2 | 4.2 | 4.5 | 4.5 | 4.0 | 4.2 | 3.6 | 3.9 | <0.001 | *** |
EIV moisture | 2.9 | 3.2 | 3.4 | 2.8 | 2.8 | 3.1 | 3.6 | 3.8 | 4.2 | 4.3 | <0.001 | *** |
EIV reaction | 7.3 | 7.6 | 7.7 | 7.1 | 6.8 | 7.6 | 5.8 | 7.4 | 6.3 | 6.8 | <0.001 | *** |
EIV nutrients | 2.0 | 3.1 | 3.3 | 1.8 | 2.2 | 2.6 | 2.9 | 3.7 | 3.4 | 3.0 | <0.001 | *** |
Topography | ||||||||||||
Elevation [m] | 810 | 899 | 1055 | 1410 | 1114 | 1112 | 905 | 854 | 1212 | 1675 | <0.001 | *** |
Inclination [°] | 45 | 41 | 35 | 38 | 38 | 36 | 22 | 38 | 30 | 33 | 0.010 | ** |
Southing | 0.49 | 0.94 | 0.98 | 0.87 | 0.86 | 0.95 | 0.89 | 0.92 | 0.16 | 0.62 | 0.031 | * |
Heat index | 0.32 | 0.40 | 0.40 | 0.35 | 0.42 | 0.45 | 0.20 | 0.71 | 0.16 | 0.21 | 0.322 | |
Maximum microrelief [cm] | 30 | 11 | 26 | 15 | 23 | 11 | 15 | 8 | 17 | 7 | 0.083 | |
Soil parameters [general] | ||||||||||||
Cover rocks and stones [%] | 41 | 22 | 8 | 54 | 34 | 9 | 11 | 1 | 9 | 0 | <0.001 | *** |
Cover gravel [%] | 18 | 10 | 25 | 15 | 8 | 3 | 1 | 7 | 8 | 0 | 0.047 | * |
Cover fine soil [%] | 24 | 68 | 68 | 29 | 62 | 77 | 88 | 92 | 83 | 99 | <0.001 | *** |
Soil depth mean [cm] | 9 | 18 | 13 | 6 | 9 | 14 | 8 | 33 | 11 | 31 | <0.001 | *** |
Soil depth CV | 79 | 46 | 95 | 77 | 89 | 37 | 54 | 29 | 45 | 40 | 0.019 | * |
Soil parameters [physical] | ||||||||||||
Skeleton content [%] | 53 | 52 | 37 | 36 | 26 | 26 | 30 | 26 | 22 | 31 | 0.002 | ** |
Sand content [%] | 67 | 70 | 74 | 71 | 74 | 75 | 62 | 67 | 61 | 70 | 0.004 | ** |
Silt content [%] | 14 | 15 | 10 | 15 | 12 | 10 | 22 | 16 | 20 | 14 | 0.002 | ** |
Clay content [%] | 18 | 16 | 16 | 14 | 14 | 15 | 16 | 18 | 20 | 16 | 0.182 | |
Water at saturation [%] | 44.3 | 43.3 | 43.2 | 42.6 | 42.4 | 42.8 | 44.0 | 44.0 | 45.0 | 39.9 | 0.739 | |
Water at field capacity [%] | 21.6 | 20.4 | 20.1 | 19.8 | 19.3 | 19.7 | 21.7 | 21.5 | 23.0 | 20.7 | 0.037 | * |
Water at permanent wilting point [%] | 12.3 | 11.2 | 11.4 | 10.5 | 10.5 | 11.1 | 11.3 | 12.0 | 12.8 | 11.5 | 0.178 | |
Plant-available water [%] | 9.4 | 9.3 | 8.7 | 9.3 | 8.8 | 8.6 | 10.4 | 9.5 | 10.2 | 9.2 | 0.002 | ** |
Drainage rate [cm/h] | 1.0 | 1.3 | 1.3 | 1.6 | 1.6 | 1.4 | 1.3 | 1.1 | 1.0 | 1.3 | 0.192 | |
Soil parameters [chemical] | ||||||||||||
pH [in aqua dest.] | 7.12 | 7.27 | 7.35 | 7.64 | 6.87 | 7.22 | 4.73 | 7.15 | 6.28 | 7.00 | <0.001 | *** |
Electrical conductivity [µS/cm] | 275 | 263 | 112 | 175 | 90 | 84 | 77 | 203 | 193 | 167 | 0.010 | ** |
Organic matter [%] | 19.9 | 25.7 | 11.3 | 10.0 | 11.8 | 12.7 | 25.8 | 13.6 | 15.0 | 11.0 | <0.001 | *** |
Organic carbon [%] | 11.5 | 14.8 | 6.5 | 5.8 | 6.8 | 7.3 | 14.9 | 7.9 | 8.6 | 6.3 | <0.001 | *** |
N content [%] | 1.7 | 3.3 | 1.6 | 1.7 | 1.0 | 0.9 | 2.1 | 1.4 | 2.1 | 1.2 | <0.001 | *** |
C/N ratio | 7.0 | 4.4 | 4.1 | 3.6 | 9.0 | 14.1 | 8.2 | 5.7 | 4.6 | 8.4 | 0.589 | |
P available [ppm] | 17.1 | 38.5 | 29.8 | 16.0 | 17.5 | 14.6 | 28.2 | 21.6 | 15.5 | 12.7 | <0.001 | *** |
According to all ecological indicator values except soil reaction, orders A and B were indistinguishable, but different from the meso-xeric order C, whose species composition stands for less light, lower temperature, lower continentality, more moisture and more nutrients (Table
At the association level, there were also strong differences for most of the measured variables (Table
Comparison of six mean ecological indicator values among the plots of the three orders A (Stipo-Festucetalia pallentis), B (Festucetalia valesiacae) and C (Brachypodietalia pinnati). Letters represent homogeneous groups (at α = 0.05) according to Tukey’s post-hoc test following a significant ANOVA.
In the following paragraphs, the ten accepted vegetation units at association level are described, and their syntaxonomic position is discussed. Four units could be unambiguously assigned to described associations, while the six remaining ones are labelled with informal community names.
(relevés 1–4 in Suppl. material 2, Table
Photos of associations and communities I. a. A.1 Armerio elongatae-Potentilletum arenariae, site GU; b. A.2 Phleo-Pulsatilletum nigricantis, PÖ; c. A.2 Phleo-Pulsatilletum nigricantis, PÖ; d. A.2 Phleo-Pulsatilletum nigricantis, PU; e. A.3 Medicago minima-Melica ciliata community, LA; f. A.4 Koelerio pyramidatae-Teucrietum montani, OM. Photos: J. Dengler (a, b, e, f) and M. Magnes (c, d).
Ecology and distribution: Our relevés were recorded on very steep slopes with predominantly southern aspect and elevation between 785 and 830 m. The rugged terrain had a high cover of rocks and gravel on the surface and very shallow soils rich in skeleton. The high soil pH is typical for serpentinite (in this case it corresponds quite well with mean EIV-R of 7.2). N and P are rather low, but organic matter is high, even if there is a strikingly low litter cover (Table
Syntaxonomy: Our relevés are from the locus classicus of this association, which was described by
(relevés 5–7 in Suppl. material 2, Table
Characterisation: Rocky grasslands on steep, south-facing slopes at elevations between 861 and 935 m. The vegetation is almost closed with bryophyte and lichen layers nearly absent (Table
Ecology and distribution: The sampled stands were not managed. They have deep soils, the second highest organic matter content among all clusters and also relatively high litter cover, P and N (Table
Syntaxonomy: This unit perfectly matches the association Phleo-Pulsatilletum nigricantis described by
(relevés 8–9 in Suppl. material 2, Table
Characterisation: Cluster A.3 only includes two relevés from the Upper Inn valley, dominated by Melica ciliata. The vegetation cover is 70% while the cryptogam layer is poorly developed, covering only below 2%. Along with Melica ciliata, some other species have a high cover, such as Artemisia absinthium, Brachypodium pinnatum, Bromus erectus, Galium lucidum and Thymus praecox. Less abundant are Agrimonia eupatoria, Carex humilis and Medicago minima.
Ecology and distribution: The shallow soils on the steep south-facing upper slopes show a high skeleton content, high pH and P, but low N and organic matter, although the litter cover is very high (Table
Syntaxonomy: This unit does not match any described association from the Eastern Alps. The closest syntaxon from the literature is the Astragalo-Brometum stipetosum capillatae of
(relevés 10–14 in Suppl. material 2, Table
Characterisation: Rocky grasslands on extremely shallow soils on calcareous slate (Table
Ecology and distribution: Most of the surface (30–90%) is covered by rocks or gravel. Litter cover and organic content are low, like N and P content, with a high pH (Table
Syntaxomy: This unit corresponds to the association Koelerio pyramidatae-Teucrietum montani, first proposed by
(relevés 15–29 in Suppl. material 2, Table
Photos of associations and communities II. a. B.1 Sempervivum tectorum-Festuca valesiaca community, site FL; b. C.1 Astragalo onobrychidis-Brometum erecti, KA; c. C.2 Agrostis capillaris-Avenula adsurgens community, NM; d. C.3 Anthericum ramosum-Brachypodium pinnatum community, PÖ; e. C.4 Ranunculus bulbosus-Festuca rubra community, MA; f. C.5 Carduus defloratus-Brachypodium pinnatum community, ZS. Photos: J. Dengler.
Ecology and distribution: Shallow soils with rather low litter cover and organic content, and low N and P (Table
Syntaxonomy: This cluster is transitional between the orders Stipo-Festucetalia pallentis and Festucetalia valesiacae. In five relevés, the diagnostic species of the first order are more abundant, in seven relevés those of the second order, while in three relevés both species groups have more or less the same total cover (Figure
(relevés 30–37 in Suppl. material 2, Figure
Characterisation: Grasslands in the Upper Inn valley dominated by Bromus erectus and/or Brachypodium rupestre, Carex humilis and Festuca rupicola. This unit is similar to the previous one, but it has a much lower cover of stones and deeper soils. Festuca valesiaca, Sempervivum tectorum and other diagnostic species of unit B.1 are rare or absent.
Ecology and distribution: Our plots have open and shallow soils, which are poor in skeleton, organic matter, N and P, with a rather high litter cover (Table
Syntaxonomy: This cluster can be assigned to the Astragalo-Brometum, described from the Lower Engadine (Switzerland) by
(relevés 38–42 in Suppl. material 2, Table
Characterisation: This is a semi-dry grassland with dominant Agrostis capillaris and Koeleria pyramidata and the presence of several acidophytes like Avenella flexuosa, Potentilla argentea or Trifolium arvense. Remarkable is the occurrence of some very rare lichens like Rinodina cana and R. vezdae (= R. moziana, new for Styria!).
Ecology and distribution: This unit occurs on siliceous soils (significantly lowest pH of all clusters, Table
Syntaxonomy: This community has some similarity with the associations Chamaecytiso hirsuti-Koelerietum and Genisto sagittalis-Festucetum rupicolae, both (invalidly) described from southern Styria – near to the Slovenian border – by
(relevés 43–49 in Suppl. material 2, Table
Characterisation: These semi-dry grasslands on calcareous soils are dominated by Brachypodium pinnatum and Festuca rupicola. Clinopodium vulgare, Salvia pratensis, Vincetoxicum hirundinaria, Viola hirta, and together with Molinio-Arrhenatheretea taxa such as Achillea millefolium agg., Arrhenatherum elatius, Dactylis glomerata, Taraxacum sect. Ruderalia and Tragopogon orientalis are also common.
Ecology and distribution: This cluster mainly includes the semi-dry grasslands on base rich, very deep and fine soils near Pöls (PÖ) that had been abandoned for years before grassland management was started again by clearing shrubs only a few years ago. Because of the occurrence of Stipa styriaca, the site was declared as a Natura 2000 protected area. With the reintroduction of grazing by sheep in 2020, there is a good chance that Stipa will be able to spread again from the rocky areas (see unit A.2) to the parts with deep soils. In this community, there is nearly no rock and gravel cover and also skeleton soil content, P and N are low (Table
Syntaxonomy: Three geographical variants can be distinguished in this cluster: The relevé from Griffen (variant a) is a strongly disturbed, ruderalized grassland of doubtful phytosociological position. The relevé from Kaunerberg (variant c) might represent a more mesic variant of the Astragalo-Brometum (see unit C.1). The remaining relevés (variant b) could be assigned to the Scabioso ochroleucae-Brachypodietum
(relevés 50–54 in Suppl. material 2, Table
Characterisation: This unit lacks dominant species, but some Molinio-Arrhenatheretea species show high constancy, like Achillea millefolium agg., Festuca rubra agg., Plantago lanceolata and Trifolium pratense. Species number is very high, especially in variant (b) from FL, as well as cryptogam species number and cover.
Ecology and distribution: The soils are shallow and pH is remarkably low (with the exception of variant c) and both litter cover and organic soil matter are low (Table
Syntaxonomy: This is a rather heterogeneous cluster consisting of three geographical variants. Variant (a) includes a single relevé from Neumarkt in der Steiermark (NM), which is dominated by Arrhenatherum elatius and might belong to the alliance Arrhenatherion (class Molinio-Arrhenatheretea). Variant (b) contains two relevés from the Upper Inn Valley (FL), which have a high portion of acidophilous species (e.g. Calluna vulgaris, Danthonia decumbens, Viola canina). These relevés have some similarity with the Agrostio-Dianthetum deltoidis, which was described by
(relevés 55–67 in Suppl. material 2, Table
Characterisation: Semi-dry grasslands in the montane and subalpine belt on steep southern slopes dominated by Brachypodium pinnatum agg. and Laserpitium latifolium and with (sub)alpine species such as Astragalus penduliflorus, Campanula scheuchzeri, Carex sempervirens, Festuca norica, Plantago atrata, Rhinanthus glacialis and Soldanella alpina.
Ecology and distribution: Grasslands on deep soils with low organic matter and low P and N (Table
Syntaxonomy: This community is characterised by the presence of several subalpine and alpine species and therefore is transitional towards the class Elyno-Seslerietea. In this respect, it is similar to the Carlino-Caricetum sempervirentis Lutz et Paul 1947 of the northern Alps. According to the alliance concept of
While the assignment of the TWINSPAN clusters to phytosociological orders was relatively straightforward, the identification of the alliances remains doubtful for the most part.
The rocky grasslands of the Eastern Alps were included in three alliances by
Grass steppes of the Festucetalia valesiacae are represented by a single community in our data set (Sempervivum tectorum-Festuca valesiaca community), which probably can be assigned to the Festucion valesiacae. It must be noted, however, that the current concept of the order Festucetalia valesiacae is much narrower than that of
Among the semi-dry grasslands of the Brachypodietalia pinnati, the communities of Styria (Agrostis capillaris-Avenula adsurgens community, Anthericum ramosum-Brachypodium pinnatum community) clearly belong to the Cirsio-Brachypodion, which includes the meadow steppes of eastern Central Europe (
Perhaps surprisingly, only four clusters could be unambiguously assigned to described associations. The main reason is that a comprehensive syntaxonomic revision of the dry and semi-dry grasslands of the inner Eastern Alps is still lacking. Instead of adopting names of associations that might or might not correspond to our units, we preferred to use informal community names, thus making the insufficient knowledge clearly visible. The relationship between these communities and described associations such as the Achnathero-Stipetum capillatae and Teucrio-Caricetum humilis (
Beside the lack of comprehensive studies another reason for the lacking match with associations from the literature might be the unstable management of most of the sites during the last thirty years. Even at sites that have recently become protected under the Natura 2000 regime (like the grasslands of FL and KA in the Upper Inn valley or PÖ in the Mur valley), the current management is very different from the historical, much more intensive use, and it will take quite some time for the vegetation to get in balance with the new type of management.
Our results do not support a closer relationship of the dry and semi-dry grasslands of the Austrian inner-alpine dry valleys with the Pannonian steppes (see
Class: Festuco-Brometea Br.-Bl. et Tx. ex Klika et Hadač 1944
A. Order: Stipo pulcherrimae-Festucetalia pallentis Pop 1968
Alliance: ??? [Avenulo adsurgentis-Festucion pallentis Mucina in Mucina et Kolbek 1993 nom. inval. (Art. 3f), “Diantho lumnitzeri-Seslerion albicantis” sensu Mucina et Kolbek 1993 p.p.]
Nomenclatural remark: The name Diantho-Seslerion (
A.1 Armerio elongatae-Potentilletum arenariae Br.-Bl. 1961
A.2 Phleo phleoidis-Pulsatilletum nigricantis Br.-Bl. 1961
A.3 Medicago minima-Melica ciliata community
A.4 Koelerio pyramidatae-Teucrietum montani Franz in Mucina et al. 1993
B. Order: Festucetalia valesiacae Br.-Bl. et Tx. ex Br.-Bl. 1950 nom. cons. propos. [= Festucetalia Soó 1940 ≡ Festucetalia valesiacae (Soó 1940) 1947 nom. illeg.]
Nomenclatural remark: For the nomenclatural problems surrounding the name Festucetalia valesiacae and a formal proposal for a nomen conservandum, see
Alliance: Festucion valesiacae Klika 1931 nom. cons. propos. (= Festucion sulcatae Soó 1930; incl. Stipo-Festucion xerophilae Br.-Bl. et Richard 1950)
Nomenclatural remark:
B.1 Sempervivum tectorum-Festuca valesiaca community
C. Order: Brachypodietalia pinnati Korneck 1974 nom. cons. propos. (= Brometalia erecti Koch 1926)
Nomenclatural remark:
Alliance: Cirsio-Brachypodion pinnati Hadač et Klika in Klika et Hadač 1944
C.1 Astragalo onobrychidis-Brometum erecti Br.-Bl. 1950
C.2 Agrostis capillaris-Avenula adsurgens community
C.3 Anthericum ramosum-Brachypodium pinnatum community
C.4 Ranunculus bulbosus-Festuca rubra community
C.5 Carduus defloratus-Brachypodium pinnatum community
For vascular plants, our findings that meso-xeric stands were substantially richer in species than either rocky or xeric grasslands across all grain sizes, is consistent with results from various other regional studies (
For bryophytes and lichens, the situation was reversed in comparison to mean values from nested-plot sampling throughout the Palaearctic (GrassPlot Diversity Explorer v.2.10; https://edgg.org/databases/GrasslandDiversityExplorer;
Based on the similar floristic composition of their vegetation, traditional vegetation ecology has anticipated close ties between the inner-alpine dry valleys and the Eastern European steppes (
Our study provides the first phytosociological overview of dry and semi-dry grasslands of the Austrian inner-alpine valleys using new plot data since the seminal work of
The plot data presented in this paper were recorded on the occasion of the 11th EDGG Field Workshop 2018 in Austria. They are included in the Suppl. material of this paper and are stored in and available from the GrassPlot database (https://edgg.org/databases/GrassPlot;
M.M. organized the 11th EDGG Field Workshop and guided it together with H.M. and P.K. As former and current EDGG Field Workshop Coordinators, J.D. and I.D. ensured consistent application of the EDGG methodology. M.M identified critical vascular plant species and led the writing of the manuscript. H.M. identified the lichens and provided information about rare or stenoecious lichens. E.A.K. analysed the soil samples, C.B. identified the bryophytes and added ecological aspects. W.W. prepared the syntaxonomic table and framework, wrote most of the syntaxonomic discussions and revised several other parts of the manuscript, M.J. contributed the descriptions of the Stipo-Festucetalia pallentis, J.D. prepared the vegetation tables. P.K. wrote the description of the Upper Inn valley and contributed to the discussion of the nature conservation value. R.G. supported the description of the Brachypodietalia pinnati and revised the Introduction. H.R. wrote the description of the Virgen valley. I.D. prepared the map and conducted the inferential statistical analyses, while J.D. wrote the Results and Discussion on biodiversity aspects. All authors revised and approved the whole text.
We are grateful to the IAVS for financial support for some of the participants. We would like to give special thanks to Dr. Ernst Partl, director of the Naturpark Kaunergrat for authorizing sampling in the protected areas and for helping with the organization of the field workshop.