Corresponding author: Maximilian Lübben ( m.luebben@posteo.de ) Academic editor: Jozef Šibík
© 2021 Maximilian Lübben, Brigitta Erschbamer.
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:
Lübben M, Erschbamer B (2021) Long term changes of the inner-alpine steppe vegetation: the dry grassland communities of the Vinschgau (South Tyrol, Italy) 40-50 years after the first vegetation mapping. Vegetation Classification and Survey 2: 117-131. https://doi.org/10.3897/VCS/2021/65217
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Aims: The Vinschgau is the driest inner-alpine valley in the Eastern Alps and harbours a unique steppe vegetation. We studied these dry grassland communities and aimed to answer the following questions: Which plant communities can be found currently? Do the syntaxa described by Braun-Blanquet in the 1960s still prevail in the area? Has there been any change in species composition over the last 40–50 years? Study area: Along an approximately 40 km transect, the south-facing slopes of the Vinschgau valley (South Tyrol, Italy) from Mals to Plaus were investigated. Methods: For the classification, 92 relevés were sampled in 2019 and compared with 76 relevés from the 1960s and ´70s by means of vegetation tables and ordinations (Detrended Correspondence Analysis). Results: Based on our investigation, the majority of dry grassland communities can be classified as Festuco-Caricetum supinae. Three subassociations were defined by the dominant species Stipa capillata, Bothriochloa ischaemum and Stipa pennata agg. The comparison of new and old relevés shows an increase in species from the class Sedo-Scleranthetea (e.g. Trifolium arvense, Erodium cicutarium) and the association Artemisieto-Agropyretum. In addition, ruderal elements (e.g. Erigeron annuus, Convolvulus arvensis) have also migrated into dry grasslands. A shift in the dominance over time can be recognized as well. In particular, Festuca rupicola and to some extent also Stipa capillata, have increased in abundance and frequency. Conclusions: We suggest to include the investigated closed dry grasslands in the alliance Festucion valesiacae. The rank of the character species at association, alliance and order level should be re-analysed. In order to obtain a better syntaxonomic overview of western and eastern alpine dry grassland communities in relation to Eastern European dry grasslands, a comprehensive study is absolutely necessary. Furthermore, long-term vegetation dynamics and vegetation change need to be studied in more detailed future studies.
Taxonomic reference:
Syntaxonomic references:
Abbreviations: agg. = aggregate; cf. = confer (means ‘compare’); DCA = Detrended Correspondence Analysis; s. lat. = sensu lato; s. str. = sensu stricto
biodiversity, Festuco-Brometea, Festucetalia valesiacae, inner-alpine steppes, syntaxonomy, vegetation change
The Eurasian steppe belt is the largest steppe region and stretches from the Amur in the east to the Hungarian basin in the west (
Distribution of the inner-alpine dry valleys (with friendly permission by Angelika Schwabe-Kratochwil, according to
In the past, three local scientists were concerned with vegetation mapping and ecophysiological investigations (
Due to the essential impacts on vegetation, such as climate change (
In the present study we aimed to repeat the relevés performed in the 1960s and 1970s by the three local authors
First, we were interested to check if the character species of the syntaxa described by
Over the approximately 40 km long transect from Mals to Plaus (Figure
In the 1960s and ´70s mostly all of the lower slopes in Vinschgau were used as pastures (
The investigated study area in the Vinschgau (South Tyrol, Italy) at the south-facing slopes from Mals and Laatsch to Plaus, spanning a length of approximately 40 km (Source: Office for Geology and Building Materials Testing of the Autonomous Province of Bolzano and ISPRA (big map); Eurostat (https://ec.europa.eu/eurostat/web/gisco/geodata/reference-data/administrative-units-statistical-units) EuroGeographics for the administrative boundaries (small map)).
Climate diagram from Schlanders (1981–2010) based on data from the 3PCLIM-project (Source: www.3pclim.eu). The red line shows the monthly mean temperature and the blue line the precipitation. Overall, there is an average temperature of 9.5°C and an annual precipitation rate of approximately 530 mm.
Together with Dr. Strimmer, Prof. Dr. Florineth and Dr. Köllemann, 76 relevés were selected from their studies, relocated in spring 2019, and new relevés were sampled in June 2019. Since it became apparent during the field inspections that there are currently only a few dry grassland occurrences in Dr. Köllemann’s study area and that these were hardly accessible, only the area between Mals and Staben was investigated. Due to the lack of GPS information, a congruent resurvey was not possible. The old relevés could not be spatially assigned exactly to one plot but to larger areas or slopes, thus, a “one-to-one” comparison of old and new plots was not possible. The comparison, therefore, was more focused on the vegetation type so that all 92 new relevés from June 2019 were compared with the 76 old ones in order to investigate the general changes in the species spectrum. Our relevés were sampled using the same cover scale as the three initial investigators (i.e.
Mosses and lichens were not recorded. In the first mapping, plots of 100 m² were used for the Vinschgau dry grasslands (according to
The raw table with the relevés from 2019 was sorted iteratively using the frequency of the species as a phytosociological characteristic. Relevés with similar floristic composition form a group which is characterized by character and differential species (
In addition to the vegetation tables, a Detrended Correspondence Analyses (DCA) was performed in R (
The nomenclature of the plant species follows
Due to a few floristic peculiarities in the Vinschgau, some taxa should be considered closer: Festuca valesiaca (2n = 2x = 14) and Festuca rupicola (2n = 6x = 42) belong to the Festuca valesiaca aggregate. F. valesiaca s. str. is, in addition to the microscopic sclerenchyma features, characterized vegetatively by hair-thin and darker blue-green, frosted leaves. The leaf of F. rupicola usually has a larger leaf cross-section (typically 0.6–0.7 mm), which can be practiced relatively quickly visually and haptically. Furthermore, it is often characterized by a comparatively warmer shade of green (although a blue-green colour, as is mandatory for F. valesiaca, is common). In the Vinschgau also higher-ploidy forms occur which can differ significantly from these two types in their vegetative characteristics. In addition to the number of chromosomes these characteristics primarily concern height, leaf width, leaf cross section and spikelet dimensions. In dry grasslands of lower and middle locations, the two octoploid species F. bauzanina (s. str.) and Festuca bauzanina subsp. rhaetica occur as well (Thomas Wilhalm, pers. comm.;
From the Stipa pennata complex four elements occur in South Tyrol: Stipa pennata s. str. (quite common), Stipa eriocaulis (by far the most common species, with subspecies subsp. eriocaulis and subsp. austriaca), S. epilosa (very rare). The taxonomic value of these clades is the subject of current research (Thomas Wilhalm, pers. comm.;
The delimitation between the (sub)species within the genus Thymus is not clear in every case. In our investigation this particularly concerns e.g. the alliance character species Thymus serpyllum subsp. carniolicum (= T. pulegioides subsp. carniolicus) (
According to
Hieracium pilosella s. str. and Hieracium velutinum are included in the Hieracium pilosella agg. (
Syntaxonomy and classification were essentially based on
Based on the character species Astragalus exscapus, Carex liparocarpos, Festuca rupicola, F. valesiaca, Oxytropis xerophila, Petrorhagia saxifraga, Potentilla pusilla, Pulsatilla montana, Silene otites and Stipa capillata dry grassland communities recorded in 2019 (Table
The Festuco-Poetum xerophilae could be documented in only five relevés from the northwest of the study area, near Laatsch (Figure
The Festuco-Caricetum supinae (87 relevés) occurred on the south-west to south-facing slopes from Tartsch near Mals approximately to Staben-Plaus (Figure
Synoptic table of the dry grassland communities in the Vinschgau (South Tyrol, Italy) with all relevés from 2019. Values are percentage frequencies. Only companion species with frequency > 15% are stated. I = Festuco-Poetum xerophilae (col. 1); II = Festuco-Caricetum supinae; II.1 = subassociation stipetosum capillatae (cols. 2-4); II.2 = subassociation bothriochloetosum ischaemi (cols. 5, 6); II.3 = subassociation stipetosum pennatae (cols. 7, 8). The name giving species Stipa capillata, Bothriochloa ischaemum and Stipa pennata agg. are indicated in bold. Variants (D1–D7, cols 2–8) were identified based on the similarity of the species composition: Veronica verna-variant (cols. 2, 5), typical variant (cols. 3, 6, 8), species-poor variant (col. 4), Melica ciliata-variant (col. 7). Abbreviations: AC = association character species, agg. = aggregate, cf. = confer (means ‘compare’), juv. = juvenile, KC = class character species, OC = order character species, s. lat. = sensu lato, sp. = species, ssp. = subspecies, VC = alliance character species.
Vegetation type | I | II.1 | II.2 | II.3 | ||||
---|---|---|---|---|---|---|---|---|
Column number | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 |
Number of relevés | 5 | 12 | 25 | 8 | 13 | 8 | 9 | 12 |
AC1: Festuco-Poetum xerophilae | ||||||||
Achillea nobilis | 100 | – | – | – | – | – | – | – |
Thesium linophyllon | 80 | 33 | 12 | – | 31 | – | – | 8 |
D1 | ||||||||
Bromus erectus | 60 | 25 | 16 | – | 8 | 13 | 11 | 8 |
Pimpinella saxifraga | 80 | – | 4 | – | 8 | – | – | – |
Prunus spinosa | 60 | 8 | – | – | – | – | – | – |
Potentilla argentea | 40 | 8 | – | – | 8 | – | – | – |
Anthoxanthum odoratum | 40 | – | – | – | – | – | – | – |
AC2: Festuco-Caricetum supinae | ||||||||
Astragalus onobrychis | 20 | 92 | 84 | 13 | 100 | 88 | 100 | 58 |
Carex supina | – | 67 | 80 | 25 | 100 | 75 | 78 | 33 |
Achillea tomentosa | – | 67 | 28 | 50 | 77 | 38 | 11 | 25 |
D2 | ||||||||
Artemisia absinthium | – | 75 | 8 | – | 38 | – | – | – |
Buglossoides incrassata | 20 | 58 | 4 | – | 31 | – | – | – |
Erodium cicutarium | – | 50 | 4 | 13 | 38 | – | – | – |
Convolvulus arvensis | – | 50 | 4 | – | 8 | 13 | – | – |
D3 | ||||||||
Veronica verna agg. | 20 | 92 | 44 | 63 | 77 | 38 | 11 | 8 |
Trifolium arvense | 100 | 75 | 12 | 88 | 85 | 13 | 11 | 25 |
Trifolium campestre | – | 50 | – | 13 | 77 | 13 | 11 | – |
Plantago lanceolata | – | 58 | 8 | – | 46 | 13 | – | – |
Turritis glabra | 40 | 50 | 12 | – | 69 | – | – | 8 |
D4 | ||||||||
Silene nutans | – | – | 16 | – | – | 13 | – | – |
Plantago media | – | 8 | 16 | – | – | 13 | – | – |
Carduus nutans | 20 | – | 20 | – | 8 | – | – | – |
Achillea cf. collina | – | – | 8 | – | – | 13 | – | – |
Trifolium repens | – | – | 8 | – | – | – | – | – |
D5 | ||||||||
Erigeron annuus | – | – | – | 75 | – | 13 | – | 8 |
Chondrilla juncea | – | 25 | 16 | 50 | – | – | 11 | 8 |
Quercus pubescens juv. | – | – | – | 38 | – | – | – | 8 |
Prunus mahaleb | – | – | 8 | 38 | 8 | 13 | – | – |
Filago arvensis | 20 | 17 | 12 | 50 | 8 | – | – | 8 |
D6 | ||||||||
Melica ciliata | 40 | 42 | 8 | 13 | 15 | 13 | 100 | – |
Allium sphaerocephalon | 100 | 25 | 8 | 25 | 23 | 13 | 67 | 8 |
D7 | ||||||||
Scorzonera austriaca | – | – | 8 | – | – | – | – | 25 |
Ephedra helvetica | – | – | – | – | – | – | – | 17 |
Telephium imperati | – | – | – | – | – | – | – | 17 |
Seseli pallasii | – | – | – | 38 | – | 13 | – | 17 |
Kengia serotina | – | – | – | 13 | – | – | – | 25 |
VC: Stipo-Poion xerophilae | ||||||||
Centaurea stoebe | 80 | 67 | 52 | 100 | 85 | 50 | 44 | 42 |
Thymus pulegioides agg. | – | 17 | – | – | 31 | – | 11 | – |
Verbascum chaixii agg. | – | – | 4 | – | – | – | – | 17 |
Poa molinerii | – | – | – | – | – | 13 | – | – |
OC: Festucetalia valesiacae | ||||||||
Potentilla pusilla | 100 | 83 | 96 | 75 | 100 | 100 | 100 | 83 |
Festuca valesiaca | 80 | 83 | 84 | 88 | 92 | 88 | 78 | 83 |
Festuca rupicola | 80 | 58 | 88 | 100 | 69 | 100 | 89 | 83 |
Stipa capillata | 80 | 100 | 100 | 100 | 62 | 88 | 44 | 25 |
Petrorhagia saxifraga | 80 | 75 | 36 | 88 | 77 | 75 | 100 | 75 |
Silene otites | 100 | 58 | 56 | 75 | 85 | 63 | 56 | 42 |
Carex liparocarpos | 20 | 33 | 32 | 63 | 8 | 13 | 22 | 33 |
Pulsatilla montana | 40 | 8 | 8 | – | 23 | – | – | 8 |
Astragalus exscapus | – | 8 | 16 | – | 15 | 13 | – | – |
Oxytropis xerophila | – | – | 4 | – | – | 25 | – | – |
KC: Festuco-Brometea | ||||||||
Veronica spicata | 100 | 17 | 16 | 25 | 69 | 38 | – | 17 |
Galium lucidum | 100 | 42 | 44 | 25 | 15 | – | 44 | 58 |
Stipa pennata agg. | 100 | 33 | 16 | 50 | 8 | 25 | 100 | 100 |
Artemisia campestris | 80 | 100 | 92 | 88 | 100 | 88 | 100 | 100 |
Phleum phleoides | 100 | 92 | 88 | 88 | 85 | 75 | 78 | 75 |
Koeleria macrantha | 80 | 83 | 84 | 75 | 100 | 50 | 100 | 67 |
Thymus praecox agg. | 100 | 67 | 96 | 63 | 77 | 100 | 67 | 67 |
Verbascum lychnitis | 60 | 67 | 76 | 63 | 54 | 63 | 44 | 25 |
Alyssum alyssoides | 60 | 83 | 56 | 25 | 85 | 50 | 89 | 50 |
Arenaria serpyllifolia | 60 | 83 | 52 | 50 | 92 | 63 | 67 | – |
Bothriochloa ischaemum | 20 | 75 | 40 | 88 | 100 | 88 | 78 | 50 |
Stachys recta subsp. recta | 60 | 50 | 40 | 13 | 8 | 38 | 78 | 50 |
Carex humilis | 60 | 42 | 16 | 13 | 15 | 50 | 22 | 33 |
Lotus corniculatus | 20 | 42 | 32 | – | 31 | – | – | 8 |
Astragalus glycyphyllos | – | 25 | 4 | – | – | – | – | – |
Fumana procumbens | – | 8 | 44 | 50 | 23 | 75 | 22 | 50 |
Helianthemum nummularium subsp. obscurum | 20 | – | 24 | 75 | 8 | 13 | 33 | 58 |
Medicago minima | 40 | 42 | 32 | 38 | 23 | 25 | 11 | 17 |
Clinopodium acinos | 20 | 33 | 4 | – | 62 | 13 | 33 | 17 |
Companion species | ||||||||
Sempervivum arachnoideum | 100 | 50 | 56 | 75 | 85 | 88 | 67 | 92 |
Hieracium pilosella agg. | 100 | 50 | 68 | 25 | 54 | 100 | 44 | 50 |
Teucrium chamaedrys | 100 | 42 | 56 | 75 | 23 | 13 | 44 | 67 |
Erysimum rhaeticum | 60 | 83 | 40 | 13 | 77 | 63 | 44 | 25 |
Sempervivum tectorum | 40 | 25 | 32 | 63 | 46 | 38 | 78 | 92 |
Dianthus sylvestris | 80 | 42 | 36 | 25 | 46 | 75 | 44 | 67 |
Plantago strictissima | 80 | 58 | 44 | – | 46 | 63 | 22 | – |
Sedum montanum s. lat. | 100 | 58 | 12 | 25 | 31 | 25 | 78 | 33 |
Scabiosa columbaria s. lat. | – | 25 | 36 | 50 | 38 | 38 | 56 | 17 |
Teucrium montanum | 20 | 25 | 40 | – | 23 | 50 | 44 | 42 |
Berberis vulgaris | 60 | 50 | 40 | – | 15 | 50 | 11 | 33 |
Chenopodium album | – | 33 | 24 | 38 | 54 | 13 | 11 | 8 |
Medicago falcata | 40 | 58 | 28 | 25 | 8 | – | – | 17 |
Juniperus communis | – | 25 | 28 | – | 23 | 13 | 33 | 25 |
Sedum sexangulare | 20 | 17 | 12 | – | 38 | 25 | – | 17 |
Lactuca perennis | 40 | 17 | 12 | – | 8 | 13 | 22 | 25 |
Euphorbia cyparissias | – | 8 | 8 | 38 | 15 | 13 | 22 | 8 |
Tragopogon dubius | 20 | 33 | 8 | – | 23 | – | 22 | – |
Carex caryophyllea | 20 | 17 | 16 | 25 | 15 | – | – | – |
Saponaria ocymoides | – | 8 | 12 | – | 15 | – | 33 | 17 |
Veronica prostrata | – | 25 | 4 | – | 31 | – | – | – |
Arabidopsis thaliana | 60 | 8 | 4 | – | 15 | 13 | – | – |
Anthericum liliago | – | – | 4 | 13 | 15 | – | 22 | 17 |
Viola cf. kitaibeliana | – | 33 | 4 | – | 15 | – | – | – |
Rosa cf. micrantha | 20 | 8 | 8 | – | – | 13 | 11 | – |
Rosa sp. | 20 | 8 | 4 | – | 15 | – | 11 | – |
Allium lusitanicum | – | – | 12 | – | – | 13 | – | 17 |
Asplenium septentrionale | 40 | 8 | – | 25 | – | 13 | – | – |
Fraxinus ornus juv. | – | – | – | 13 | 8 | – | 22 | 17 |
Phelipanche bohemica | 20 | 17 | – | – | – | – | 22 | – |
Securigera varia | 20 | – | 4 | – | 15 | – | 11 | – |
Bromus japonicus | – | – | – | 25 | 15 | – | – | – |
Robinia pseudacacia juv. | – | – | – | – | – | – | 11 | 25 |
Poa angustifolia | 20 | 8 | 4 | – | – | – | – | 8 |
Astragalus vesicarius subsp. pastellianus | – | – | – | – | – | – | 11 | 25 |
Descurainia sophia | – | 25 | – | – | 8 | – | – | – |
Echium vulgare | – | 17 | – | – | 8 | – | – | – |
Torilis arvensis | – | – | – | 25 | 8 | – | – | – |
Hypericum maculatum | – | – | – | – | 23 | – | – | – |
Calina acaulis | – | 17 | 4 | – | – | – | – | – |
Verbascum nigrum | – | 17 | – | – | – | – | – | – |
Silene vulgaris | 20 | – | – | – | – | – | 11 | – |
Linaria angustissima | 20 | – | – | – | – | – | – | 8 |
Trifolium alpestre | 20 | – | – | – | – | 13 | – | – |
Cuscuta epithymum | 40 | – | – | – | – | – | – | – |
Vicia tetrasperma | – | – | – | 25 | – | – | – | – |
Geum montanum | 20 | – | – | – | – | – | – | – |
Idealised scheme of the ecological occurrence of the three subassociations of the Festuco-Caricetum supinae (cf.
The subassociation with Stipa capillata was found on low mountain terraces and lower slopes with deeper soils. The community was grass-rich and very often contained tall herbs (Figure
This subassociation dominated on the heavily grazed areas of the terraces and the lowest slopes, especially near Mals. The more open and very low-growing vegetation was dominated by Bothriochloa ischaemum, Festuca valesiaca, Potentilla pusilla and Thymus praecox agg. (Figure
The subassociation stipetosum pennatae characterized the rockier slopes, which were at great risk of erosion, and on rocky outcrops so that the canopy layer showed more gaps (Figure
The comparison of new and old relevés showed a clear vegetation change. Over the last 40–50 years, a large group of species newly immigrated (D1 in Table
The quantitative analysis of the relevés (DCA ordination) confirmed the discrimination of old and new relevés (Figure
Synoptic table of the dry grassland communities in the Vinschgau (South Tyrol, Italy) from 2019 in comparison to the 1960s/´70s. Values are percentage frequencies. The columns 1 to 9 show the different relevé groups based on the similarity of the species composition (2019, cols. 1-5; 1960s/70s, cols. 6-9). Different species groups (D1 – D5) were identified which separate the new and old relevés. The floristical shift over time is illustrated by these groups. In D5 species are stated which do not show a clear change over time in their frequencies. Only species with frequency > 15% are stated in this group. Abbreviations: agg. = aggregate, cf. = confer (means “compare”), juv. = juvenile, s. lat. = sensu lato, sp. = species.
Sampling period | 2019 | 1960/70 | |||||||
---|---|---|---|---|---|---|---|---|---|
Column number | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 |
Number of relevés | 17 | 16 | 19 | 17 | 23 | 15 | 24 | 26 | 11 |
D1 | |||||||||
Camelina microcarpa | 6 | – | – | – | – | – | – | – | – |
Lolium perenne | 6 | – | – | – | – | – | – | – | – |
Medicago sativa | 6 | – | – | – | – | – | – | – | – |
Geum montanum | 6 | – | – | – | – | – | – | – | – |
Reseda luteola | 6 | – | – | – | – | – | – | – | – |
Silene vulgaris | 6 | 6 | – | – | – | – | – | – | – |
Descurainia sophia | 18 | 6 | – | – | – | – | – | – | – |
Astragalus glycyphyllos | 18 | 6 | – | – | – | – | – | – | – |
Bromus japonicus | 12 | 13 | – | – | – | – | – | – | – |
Linaria angustissima | 6 | – | – | 6 | – | – | – | – | – |
Papaver dubium | 6 | 6 | – | 6 | – | – | – | – | – |
Torilis arvensis | 6 | 6 | – | 6 | – | – | – | – | – |
Viola cf. kitaibeliana | 29 | 6 | – | – | 4 | – | – | – | – |
Plantago lanceolata | 65 | 19 | 11 | – | – | – | – | – | – |
Erodium cicutarium | 59 | 13 | 5 | – | – | – | – | – | – |
Trifolium campestre | 59 | 44 | – | 6 | 4 | – | – | – | – |
Chenopodium album | 47 | 50 | 21 | – | 13 | – | – | – | – |
Arabidopsis thaliana | 18 | 6 | 5 | 6 | 9 | – | – | – | – |
Anthericum liliago | 6 | 13 | 5 | 18 | 4 | – | – | – | – |
Rosa cf. micrantha | 6 | – | 11 | 12 | 4 | – | – | – | – |
Plantago media | 6 | – | 21 | – | 4 | – | – | – | – |
Taraxacum laevigatum | 6 | – | 16 | – | 4 | – | – | – | – |
Phelipanche bohemica | 12 | – | – | 6 | 9 | – | – | – | – |
Poa angustifolia | 12 | – | – | 6 | 4 | – | – | – | – |
Asplenium septentrionale | 6 | – | – | 18 | 9 | – | – | – | – |
Prunus spinosa | 12 | – | – | – | 9 | – | – | – | – |
Lactuca serriola | 6 | – | 5 | 12 | – | – | – | – | – |
Erigeron annuus | – | 25 | 5 | 18 | – | – | – | – | – |
Quercus pubescens | – | 13 | – | 6 | 4 | – | – | – | – |
Sanguisorba minor | – | 13 | – | – | 4 | – | – | – | – |
Senecio inaequidens | – | 13 | – | 6 | – | – | – | – | – |
Vicia tetrasperma | – | 13 | – | – | – | – | – | – | – |
Orobanche artemisiae-campestris | – | 6 | – | – | – | – | – | – | – |
Ononis spinosa | – | 6 | – | – | – | – | – | – | – |
Salvia pratensis | – | 6 | 5 | – | – | – | – | – | – |
Veronica fruticans | – | – | 5 | 6 | 4 | – | – | – | – |
Silene nutans | – | – | 26 | – | – | – | – | – | – |
Achillea cf. collina | – | – | 16 | – | – | – | – | – | – |
Trifolium repens | – | – | 11 | – | – | – | – | – | – |
Myosotis stricta | – | – | 5 | – | – | – | – | – | – |
Carlina vulgaris | – | – | 5 | – | – | – | – | – | – |
Anchusa arvensis | – | – | 5 | – | – | – | – | – | – |
Cynoglossum officinale | – | – | 5 | – | – | – | – | – | – |
Lonicera xylosteum | – | – | 5 | – | – | – | – | – | – |
Cerastium semidecandrum | – | – | 5 | – | – | – | – | – | – |
Kengia serotina | – | – | – | 24 | – | – | – | – | – |
Viscaria vulgaris | – | – | – | 18 | – | – | – | – | – |
Quercus petraea | – | – | – | 6 | – | – | – | – | – |
Ulmus minor | – | – | – | 6 | – | – | – | – | – |
Carduus defloratus | – | – | – | 6 | – | – | – | – | – |
Trifolium alpestre | – | – | – | 6 | 4 | – | – | – | – |
Cuscuta epithymum | – | – | – | – | 9 | – | – | – | – |
Bromus tectorum | – | – | – | – | 9 | – | – | – | – |
Anthoxanthum odoratum | – | – | – | – | 9 | – | – | – | – |
Telephium imperati | – | – | – | – | 9 | – | – | – | – |
Aster alpinus | – | – | – | – | 4 | – | – | – | – |
Ononis natrix | – | – | – | – | 4 | – | – | – | – |
D2 | |||||||||
Artemisia absinthium | 71 | – | – | 6 | 13 | – | 4 | – | 9 |
Buglossoides incrassata | 76 | – | – | – | – | – | – | – | 9 |
Turritis glabra | 59 | 38 | 11 | – | 13 | – | 8 | – | – |
Chondrilla juncea | 18 | 19 | 5 | 35 | – | – | 4 | – | 9 |
Trifolium arvense | 82 | 88 | – | 41 | 22 | 7 | 38 | 12 | 9 |
Sedum sexangulare | 29 | 6 | 21 | – | 22 | – | – | 15 | 9 |
Veronica verna agg. | 76 | 100 | 32 | 18 | 22 | – | 4 | – | – |
Arenaria serpyllifolia | 82 | 75 | 74 | 24 | 39 | 7 | – | – | – |
Festuca rupicola | 71 | 69 | 89 | 88 | 91 | 13 | – | 27 | 36 |
D3 | |||||||||
Saponaria ocymoides | 18 | 6 | 5 | 12 | 17 | 47 | 33 | 4 | 9 |
Rosa sp. | 6 | 13 | – | – | 13 | 60 | 38 | 4 | 9 |
Oxytropis pilosa | – | – | – | – | – | 13 | 4 | – | – |
Onosma helvetica subsp. tridentata | – | – | – | – | – | 13 | 13 | – | – |
Asparagus officinalis | – | – | – | – | – | 7 | 13 | – | – |
Orobanche gracilis | – | – | – | – | – | 7 | – | – | 9 |
Veronica teucrium | – | – | – | – | – | 7 | – | – | – |
Phelipanche arenaria | – | – | – | – | – | – | 8 | – | – |
Cirsium sp. | – | – | – | – | – | – | 4 | – | – |
Clinopodium alpinum | – | – | – | – | – | – | 4 | – | – |
Medicago lupulina | – | – | – | – | – | – | 4 | – | – |
D4 | |||||||||
Stipa pennata agg. | 24 | 38 | – | 82 | 74 | 73 | 8 | 35 | 27 |
Alyssum alyssoides | 82 | 69 | 42 | 29 | 87 | 47 | 29 | 15 | 36 |
Achillea tomentosa | 76 | 63 | 21 | 29 | 17 | 53 | 54 | 19 | 55 |
Berberis vulgaris | 35 | 6 | 53 | 6 | 52 | 93 | 79 | 4 | 9 |
Juniperus communis | 24 | 6 | 32 | – | 39 | 80 | 92 | 12 | 9 |
Hieracium pilosella agg. | 65 | 25 | 79 | 29 | 87 | 73 | 92 | 81 | 9 |
Dianthus sylvestris | 47 | 31 | 37 | 41 | 74 | 80 | 83 | 81 | 36 |
Plantago strictissima | 65 | 13 | 53 | – | 52 | 67 | 88 | 92 | 64 |
Teucrium montanum | 29 | 13 | 42 | 24 | 48 | 80 | 88 | 62 | – |
Thymus praecox agg. | 71 | 75 | 100 | 59 | 91 | 87 | 75 | 88 | 82 |
Stipa capillata | 82 | 81 | 100 | 59 | 65 | 80 | 83 | 88 | 82 |
Bothriochloa ischaemum | 71 | 94 | 42 | 71 | 57 | 93 | 92 | 81 | 64 |
Centaurea stoebe | 88 | 75 | 47 | 47 | 57 | 87 | 92 | 88 | 73 |
D5 | |||||||||
Artemisia campestris | 94 | 100 | 95 | 88 | 96 | 100 | 96 | 85 | 100 |
Potentilla pusilla | 88 | 88 | 95 | 94 | 96 | 93 | 83 | 96 | 100 |
Festuca valesiaca | 82 | 88 | 95 | 71 | 87 | 73 | 100 | 96 | 100 |
Astragalus onobrychis | 82 | 69 | 89 | 59 | 78 | 100 | 79 | 85 | 82 |
Phleum phleoides | 94 | 94 | 79 | 82 | 74 | 73 | 79 | 69 | 64 |
Sempervivum arachnoideum | 71 | 69 | 58 | 71 | 87 | 80 | 96 | 65 | 36 |
Koeleria macrantha | 88 | 88 | 68 | 100 | 70 | 80 | 75 | 46 | 36 |
Petrorhagia saxifraga | 71 | 88 | 47 | 76 | 65 | 100 | 83 | 62 | 45 |
Silene otites | 71 | 75 | 42 | 47 | 78 | 93 | 88 | 50 | 27 |
Carex supina | 71 | 69 | 74 | 47 | 65 | 33 | 50 | 50 | 64 |
Erysimum rhaeticum | 88 | 50 | 32 | 24 | 57 | 80 | 67 | 23 | 18 |
Carex humilis | 35 | 13 | 26 | 24 | 35 | 67 | 58 | 85 | 36 |
Verbascum lychnitis | 59 | 63 | 89 | 35 | 48 | 53 | 25 | 8 | 27 |
Sempervivum tectorum | 35 | 31 | 16 | 88 | 70 | 93 | 33 | 12 | 18 |
Teucrium chamaedrys | 41 | 38 | 47 | 53 | 65 | 73 | 50 | 4 | 18 |
Galium lucidum | 29 | 31 | 26 | 24 | 74 | 67 | 58 | 15 | 27 |
Scabiosa columbaria s. lat. | 47 | 25 | 26 | 41 | 30 | 53 | 75 | 38 | – |
Fumana procumbens | 18 | 38 | 37 | 53 | 35 | 73 | 54 | 27 | – |
Stachys recta subsp. recta | 29 | 25 | 47 | 47 | 48 | 73 | 38 | 4 | 9 |
Veronica spicata | 41 | 38 | 11 | 18 | 39 | 20 | 29 | 46 | 45 |
Sedum montanum s. lat. | 53 | 13 | 11 | 41 | 61 | 60 | 33 | 12 | – |
Helianthemum nummularium subsp. obscurum | 6 | 25 | 11 | 65 | 30 | 73 | 50 | 15 | 9 |
Medicago falcata | 53 | 13 | 32 | 6 | 13 | 27 | 46 | 31 | 36 |
Allium sphaerocephalon | 24 | 25 | – | 29 | 43 | 40 | 33 | 8 | 18 |
Lotus corniculatus | 53 | 19 | 21 | 6 | 9 | 13 | 50 | 27 | – |
Carex liparocarpos | 35 | 38 | 26 | 24 | 22 | – | – | 31 | 36 |
Clinopodium acinos | 41 | 31 | 5 | 18 | 17 | 40 | 33 | 4 | 18 |
Melica ciliata | 35 | 19 | – | 24 | 39 | 40 | 17 | 8 | 18 |
Medicago minima | 29 | 31 | 37 | 29 | 17 | – | 13 | – | 9 |
Tragopogon dubius | 29 | 19 | – | 18 | 4 | 27 | 25 | 8 | 18 |
Bromus erectus | 29 | 13 | 11 | 6 | 17 | 33 | 17 | – | 9 |
Thesium linophyllon | 41 | 13 | 11 | – | 22 | 7 | 17 | 4 | 9 |
Carex caryophyllea | 29 | 19 | 11 | 6 | – | – | 4 | 31 | 18 |
Astragalus exscapus | 6 | 13 | 16 | – | 9 | – | 4 | 38 | 9 |
Euphorbia cyparissias | 12 | 13 | 11 | 12 | 17 | 13 | 17 | – | – |
Filago arvensis | 12 | 19 | 11 | 18 | 9 | – | 13 | 8 | 9 |
Carduus nutans | 12 | – | 26 | – | – | 7 | 33 | 4 | 9 |
Lactuca perennis | 6 | 13 | 5 | 24 | 26 | 13 | 4 | – | – |
Verbascum nigrum | 6 | 6 | – | – | – | – | – | 42 | 27 |
Galium verum | 6 | 6 | 5 | – | – | 27 | 25 | 8 | 9 |
Thymus pulegioides agg. | 24 | 13 | – | – | 4 | 13 | 13 | 12 | – |
Allium lusitanicum | – | 6 | 11 | 6 | 9 | 27 | 13 | 4 | – |
Pulsatilla montana | 12 | 13 | 11 | – | 13 | – | – | 15 | – |
Pimpinella saxifraga | 12 | – | 5 | – | 13 | – | 21 | 8 | – |
Seseli pallasii | – | 6 | – | 24 | 4 | 20 | 13 | – | 9 |
Calina acaulis | 12 | – | 5 | – | – | – | 8 | 19 | 18 |
Oxytropis xerophila | – | – | 11 | – | 4 | – | 13 | 19 | 9 |
Astragalus vesicarius subsp. pastellianus | – | – | – | 6 | 13 | 13 | 21 | 4 | – |
Convolvulus arvensis | 35 | – | – | 6 | 9 | 7 | 4 | – | 9 |
Prunus mahaleb | – | 19 | 11 | – | 9 | – | 17 | – | – |
Securigera varia | 6 | 6 | 5 | – | 9 | 7 | 17 | – | – |
Veronica prostrata | 24 | 13 | 5 | – | 4 | – | – | 4 | 9 |
Potentilla argentea | 6 | – | – | – | 13 | 7 | 4 | 4 | 18 |
Scorzonera austriaca | – | 6 | – | 12 | 9 | 27 | – | – | – |
Fraxinus ornus juv. | 6 | – | – | 18 | 9 | – | – | – | 9 |
Robinia pseudacacia | – | – | – | 18 | 4 | 7 | 4 | – | 9 |
Lappula squarrosa | 6 | – | 16 | 6 | – | – | 4 | – | – |
Achillea nobilis | 6 | – | – | – | 17 | – | – | – | 9 |
Echium vulgare | 18 | – | – | – | – | – | – | – | 9 |
Following
On the association level, we were able to identify two associations (Festuco-Poetum xerophilae and Festuco-Caricetum supinae). According to
The Festuco-Caricetum supinae was well represented (87 relevés). The association is well justified by the character species. This holds also for the subassociations, based on the dominance of Bothriochloa ischaemum, Stipa capillata and Stipa pennata agg. Nevertheless, as our investigation points out, the three subassociations were floristically closely related and showed transitions. A mosaic distribution of different dry grassland ”fragments” and fluent transitions of dry grassland communities in Vinschgau were already mentioned by
On the whole, a more precise determination of some species on the subspecies level could perhaps lead to a more exact delimitation of the associations, subassociations and variants. A comprehensive phytosociological study of the entire inner-alpine steppe vegetation is definitely needed to gain a better syntaxonomical overview and classification in the context of western and eastern dry grassland communities (cf.
Over the last 40–50 years, considerable changes in species composition were recognized. The abundance and constancy of Stipa capillata and – to a weaker extent – of Stipa pennata agg. increased, while that of Bothriochloa ischaemum slightly decreased. The most impressive increase was shown by Festuca rupicola. These changes of the character species have to be interpreted with caution. Especially for Festuca rupicola, determination problems in the past cannot be excluded. According to our own observation, F. rupicola inhabits more mesophilic and deeper soils, while F. valesiaca grows dominantly on shallower and drier soils. The different requirements of the two species were already mentioned by
Besides these uncertain changes, ruderalisation trends, mentioned already by
We also recognized that species from the Sedo-Scleranthetea class (e.g. Erodium cicutarium, Trifolium campestre; D1, Table
According to our investigation the three subassociations of the Festuco-Caricetum supinae generally inhabit different parts in the Vinschgau. The subassociation stipetosum pennatae occurs on rockier and usually steeper areas, while the bothriochloetosum ischaemi stocks on heavily grazed pastures and the stipetosum capillatae grows mainly on deeper soils (Figure
It is known that, in addition to ecological factors, changes in management, i.e. over- or undergrazing respectively abandonment of use, strongly influence species composition and community changes in steppe vegetation (
Our investigation shows that current dry grassland communities in the Vinschgau can be identified mainly as the Festuco-Caricetum supinae (order Festucetalia valesiacae). We suggest to include the investigated dry grasslands to the alliance Festucion valesiacae. Although the presented classification is sufficiently justified, the delimitation of the associations and subassociations still needs further discussion. Likewise, the rank of the character species at association, alliance and order level should be re-analysed. A comprehensive study is definitely needed not only to gain a better syntaxonomical overview of western and eastern alpine dry grassland communities but also to evaluate their relation to Eastern European dry grasslands. Especially, relevés from different years and over the whole season of one year are necessary for a more precise classification of the inner-alpine steppe vegetation, to be able to estimate the fluctuations in abundance between years. In addition, a more precise identification of some (sub)species could lead to a more exact delimitation on association-, subassociation- and variant level. The current classification and delimitation of dry grassland communities of the class Festuco-Brometea is neither uniform nor free of contradictions (cf.
M.L. performed the fieldwork, did the statistical analysis, and wrote the first draft of the manuscript. B.E. supervised the work and improved the drafts of the manuscript.
We would like to thank Dr. Thomas Wilhalm (Nature Museum, Bolzano) for his floristic support, Dr. Strimmer, Prof. Dr. Florineth and Dr. Köllemann for their support and information about their study in the Vinschgau, the Department of Botany (University of Innsbruck) for the financial support and the editors as well as the anonymous reviewers for their helpful comments.