Plant communities of mountain grasslands in a broad cross-section of the Eastern Alps

Flora 206 (2011) 433–443

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Plant communities of mountain grasslands in a broad cross-section of the Eastern Alps Christian Lüth a , Erich Tasser b , Georg Niedrist b , Josef Dalla Via c , Ulrike Tappeiner a,∗ a b c

Institute of Ecology, University of Innsbruck, Sternwartestr. 15, 6020 Innsbruck, Austria European Academy of Bolzano/Bozen, Drususallee 1, 39100 Bozen, Italy Research Centre for Agriculture and Forestry Laimburg, Laimburg 6, 39051 Pfatten/Auer, Italy

a r t i c l e

i n f o

Article history: Received 2 March 2010 Accepted 27 July 2010 Keywords: Land-use types Land-use intensity Site factors Meadows Pastures Abandoned areas

a b s t r a c t Apart from forests, the landscape of the Alps is dominated by grasslands, where they account for up to 40% of the agricultural area. This study focuses on the main man-made grassland plant communities of the Eastern Alps, shows their current spatial distribution and examines how strongly the influence of land use and site factors determines the communities. Discriminant analysis was used to harmonize the phytosociological classification of 1502 vegetation relevés from the literature and 375 own recorded inventories from Western Austria and Northern Italy. Land-use intensity, altitude, slope and pH were also recorded, in order to assess the impact of the factors to plant communities, as calculated in nonmetric multidimensional scaling. We identified 39 plant communities and generated a table with the main ecological and floristic parameters as well as a map showing their present spatial distribution. Contrary to the literature, the pasture communities Crepido-Festucetum commutatae, Deschampsio cespitosaePoetum alpinae and Rumicetum alpini occur also in fertilized meadows. On the other hand we found meadow communities occurring in pastures, such as the Angelico-Cirsietum oleracei, the PastinacoArrhenatheretum, the Ranunculo repentis-Alopecuretum pratensis and the Trisetetum flavescentis. The most species-rich communities – the Caricetum ferruginei and the Seslerio-Caricetum sempervirentis – occur in unfertilized meadows above calcareous bedrock. Further species-rich communities – the Campanulo scheuchzeri-Festucetum noricae, the Gentianello anisodontae-Festucetum variae, the Pulsatillo alpinae-Festucetum noricae, the Trifolio thallii-Festucetum nigricantis and the Hypochoerido unifloraeFestucetum paniculatae – are endangered: they are regionally restricted and depend on the absence of fertilizer and on mowing once annually or every second or third year. Therefore agri-environmental measures should focus on unfertilized mountain meadows, in order to conserve these rare grassland communities. © 2010 Elsevier GmbH. All rights reserved.

Introduction Grasslands represent one of the most diverse man-made landscape formations in the Alps (Ellenberg, 1996; Kremer, 1991; Maurer et al., 2006). The percentage of meadows and pastures in the subalpine-alpine belt lies between 5% and 40% of the agriculturally used area, depending on the region (Tasser et al., 2009). Generally, due to the shorter growing season, the unfavourable climate and topographic conditions, farmers have tended very early to a marginal use of these areas (Niedrist et al., 2008). Alpine grasslands are mainly used as summer pastures (Ellenberg, 1996), but a small proportion are meadows, usually

Abbreviations: DA, discriminant analysis; NMS, nonmetric multidimensional scaling; a.s.l., above sea level. ∗ Corresponding author. Tel.: +43 512 507 5923; fax: +43 512 507 2975. E-mail address: [email protected] (U. Tappeiner). 0367-2530/$ – see front matter © 2010 Elsevier GmbH. All rights reserved. doi:10.1016/j.flora.2010.11.007

unfertilized and mown annually or every second to third year. Generally only the easily accessible meadows are fertilized and allow two cuts of hay in one year. Depending on land use, typical plant communities have become established over the past centuries and millennia (Dullinger et al., 2003; Liira et al., 2008; Marcos et al., 2003; Oomes, 1992; Tasser et al., 2003; von Arx et al., 2002). On the alpine belt meadows typically support a large percentage of herbs as well as dwarf-shrubs (Ellenberg, 1996; Ellmauer, 1996; Marini et al., 2007) and show a pronounced species-richness (Grabherr and Mucina, 1993; Mucina et al., 1993). Besides land-use management, small changes in site factors (altitude, slope angle and soil pH) increase ecosystem diversity (Kampmann et al., 2007; Marini et al., 2007). This is demonstrated impressively over large areas in alpine pastures, where animals are left more or less unattended (Tasser et al., 2003). Grazing animals prefer the gentle slopes of the pastures with their dry and undisturbed soils. Depending on topsoil pH, communities develop with Nardus stricta on acid soil and with Sesleria albicans on neutral to

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alkaline soils (Grabherr and Mucina, 1993; Mucina et al., 1993; Myklestad and Sætersdal, 2004). Plane places are used as animal resting places, where the increase in local nutrient load leads to the development of typical trodden swards. On steep pastures, the free range style of grazing in the Alps greatly reduces the frequency of use by grazing cattle and thus the level of nutrient deposition. The resulting vegetation covers exhibit a dwarf-shrub invasion or typical high alpine gramineous-rich communities. In the Alps land use combined with site factors results in a large variety of plant communities (Pykälä, 2000) and creates a landscape with high diversity (Tappeiner et al., 2008). Hence, this region is a ‘hot spot’ of biodiversity in Europe (World Wildlife Fund, 2004). Unfortunately, traditional land-use practices have become less important over the last few decades in the Alps (Streifeneder et al., 2007; Tasser et al., 2003). About 20% – in some areas even much more – of the agricultural land has been abandoned (Macdonald et al., 2000; Tappeiner et al., 2008; Tasser et al., 2007). The crucial factor here is that steep areas and small plots create higher production costs, thus mountain agriculture is hardly competitive on national and international markets. Consequently marginal productive areas in the Alps have been increasingly abandoned since 1950. However, the severity of this decline varies significantly, depending on the region (Tasser et al., 2007, 2008): in the region ‘Südtiroler Unterland/Überetsch’ – one of the most productive regions of the Alps – only 6% of formerly used agricultural areas are currently abandoned, while in the region ‘Innsbruck-Land’ they amount to 37%, and in the region ‘Carnia’ even to 67%. On the other hand, an intensification of land use can also be observed (Krausmann et al., 2003; Mottet et al., 2006), due to the massive extension of forest roads and farm tracks, which leads to a higher accessibility, mechanisation and manuring of agricultural areas in the alpine and subalpine altitudinal belt (Liira et al., 2008; Pavlu˚ et al., 2005; Stöcklin et al., 2007). All these changes lead to a reduction of semi-natural grasslands (Bakker and Berendse, 1999; Lavorel et al., 1998; Tappeiner et al., 1998), and an old and invaluable cultural landscape is gradually disappearing. Several studies exist on grasslands in the Eastern Alps (Bischof, 1981; Kohler et al., 2004; Maurer et al., 2006; Tasser and Tappeiner, 2002; Vonlanthen et al., 2006; Zimmermann and Kienast, 1999) as well as numerous diplomas and Ph.D. theses (see Appendix) containing field relevés of grasslands. However, they often refer only to small regions and are not related to one another. Unfortunately, their descriptions of plant communities do not focus on form and intensity of land use, even though plant communities are predominantly the result of differences in land-use management (Burnside et al., 2007; Studer, 2001). It is therefore about time that an inventory which characterizes the remaining grassland plant communities is created, in order to have a database for prospective decisions. With a huge and unique data set of vegetation relevés in the Eastern Alps, the present study aims at (1) providing a general overview of the man-made grassland plant communities and their distribution with a focus on montane to alpine regions, (2) clarifying how land use and site factors affect their establishment and (3) find out the most species-rich communities and how to maintain them.

Materials and methods Research area To obtain a cross-section of meadows through the Eastern Alps, vegetation relevés were taken from Tyrol (Austria) and South Tyrol (Italy) together with a number of further relevés from western Vorarlberg, eastern Salzburg (both Austria) and northern Trentino (Italy). The location of the region lies between 47◦ 36 –46◦ 02 N

and 10◦ 08 –12◦ 45 E (Fig. 1). Average annual precipitation ranges from 700 mm to 2000 mm, with maximum rainfall observed from June to July (Fliri, 1998). Mean annual temperature ranges from 0 ◦ C to 9 ◦ C. Strong climatic distinctions are caused by the fact that relevés were taken from 650 m to 2680 m a.s.l. and that two climatic regions – Continental and Atlantic climate – affect the vegetation in the research area (Fliri, 1998). The bedrock of the research area is comprised of calcareous sedimentary rocks in the northern and the southern regions and of primary rocks in the central massive, sometimes with superimposed calcareous isles (Bögel and Schmidt, 1976). The pH of the topsoil (0–10 cm), which ranges from 3.7 to 7.8 (Niedrist et al., 2008), is either affected by the bedrock (Scheffer et al., 2002) or else has been modified as a result of fertilizer applications (Seeber and Seeber, 2005). Data collection We first collected 1507 vegetation relevés from the literature (see Appendix), which had been recorded following the method of Braun-Blanquet (1964). This collection comprised meadows, pastures and abandoned areas from 60 different sites. Thereby the plot size ranges from 12 m2 to 25 m2 . All relevés included detailed information on land use, geographical coordinates and the site factors for altitude and slope angle. For regions where no literature data were found, we consulted local experts (farmers, park rangers and agrarian decision-makers of the particular districts) about the location of remaining mountain hay meadows and pastures. From this information we then recorded 265 vegetation relevés from 84 sites. Furthermore, at 30 sites we recorded 110 vegetation relevés of intensively used meadows from valley regions (between 650 and 1200 m a.s.l.), in order to compare them with lightly used ones. Fieldwork was carried out between 2005 and 2007. Vegetation relevés were recorded according to the method of Braun-Blanquet (1964): the minimum area ranged from 6 m2 (in intensively fertilized meadows from the valley) up to 20 m2 (in mountain meadows without fertilization), depending on the heterogeneity of the grassland area; most of the relevés covered an area of 12 m2 (i.e. 4 m × 3 m). At least two relevés were recorded for each study site, the variables altitude, slope, and pH (CaCl2 ) of the topsoil (0–10 cm) were measured and the managing farmers were interviewed to obtain exact information on land use. For statistical calculations we transformed the indications of BraunBlanquet (1964) to dominance values in percent (according to Tasser and Tappeiner, 2004): r = 0.1%, + = 0.3%, 1 = 2.8%, 2m = 4.5%, 2a = 10%, 2b = 20.5%, 3 = 38%, 4 = 63%, and 5 = 88%. Land use was divided into three main groups (Table 1): (1) meadows, which were subdivided into (a) unfertilized mountain meadows (UM) – mown every year or infrequently every second to third year, (b) fertilized mountain meadows (MM) – mown once a year, but grazed by animals before and/or after mowing and (c) fodder meadows (FM) – mown two to five times a year, mostly of valley regions; (2) pastures, which were subdivided into (a) lightly used pastures (LP) with unattended grazing animals during the vegetation period and (b) intensively used pastures (IP) in fenced sites or near stables with high stocking of grazing animals; (3) young abandoned areas (AA), which were formerly mown and having lain fallow for not more than 30 years; older abandoned areas were excluded, and the year of the last mowing was based either on literature or interviews. In alpine grasslands a smooth transition from fertilized meadows (mostly with cattle dung manuring) to meadows without fertilization, abandoned areas and pastures of different grazing intensities is observed. Therefore, land-use intensity (LUi) was classified according to Tappeiner et al. (1998): For meadows we summed every human impact Ih (mowing, fertilization) and divided it by the frequency of these interferences in years a. The same pro-

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Fig. 1. Location of the research area Tyrol (Western Austria) and South Tyrol (Northern Italy) and the sites of the vegetation relevés. Dark grey lines: rivers and valleys, light grey areas: limestone or calcareous mica schist regions, white areas: silicate regions.

cedure was applied for abandoned areas: thereby we summed the years from the last human impact until now and took the reciprocal value of it.



set and homogenized it; thereby the process of reviewing consisted of the following steps:

Data analysis

1. The species names had to be standardized, as the incorporated data from literature spanned a period of more than 50 years (e.g. Thimm, 1950, in Appendix), using the nomenclature of Fischer et al. (2005). Taxa of species and subspecies mostly had to be contracted to aggregates, if they had not been precisely classified by each author. This resulted in a data set of 769 species. 2. Syntaxonomic names of plant communities also had to be standardized, based on Grabherr and Mucina (1993) and Mucina et al. (1993); three newer association names belong to Ellenberg (1996), Wallossek (1999) and Grabner and Heiselmayer (2002).

The vegetation relevés from the literature were already classified by the corresponding authors (Appendix). As single authors classify communities in a slightly different way (due to subjectivity of estimating the abundance values), we reviewed the whole data

We then grouped our own recorded vegetation relevés with a hierarchical cluster analysis and integrated them into the data set from literature by comparing their species composition. Afterwards we used a discriminant analysis (DA) to obtain an automated clas-

LUi =

Ih × a−1

The pasture utilization was indicated as ordinal data type, ranging from 1 (low grazing intensity) to 4 (intensive grazing) (for details see Tasser et al., 1998). Thereby scaling is similar to the metric data of the land-use intensity for meadows and abandoned areas (i.e. high grazing intensity has a similar impact as intensively used meadows with 4 impacts per year).

Table 1 Main land-use types of grasslands in the Eastern Alps, their respective abbreviation codes (LU), number of relevés, land-use intensity quotient (LUi), site factors (altitude, slope angle and pH of the topsoil) and number of species. LU

Name and land-use type

No. of relevés

LUi

Altitude (m a.s.l) mean ± s.d.

Slope angle (◦ ) mean±s.d.

pH (0–10 cm) mean ± s.d.

No. of species mean ± s.d.

UM

Unfertilized mountain meadows, mown every year, seldom every 2nd or 3rd year Fertilized mountain meadows, mown once a year and mostly grazed in autumn Fodder meadows, mown two to five times a year Lightly used pastures, with unattended grazing animals Intensively used pastures, with high stocking of grazing animals Abandoned areas, abandoned for not more than 30 years

560

0.33–1

1797 ± 223

18.8 ± 11.3

5.07 ± 0.67

39.8 ± 13.0

260

2–3

1737 ± 260

13.9 ± 10.2

5.37 ± 0.49

31.9 ± 11.3

364

4–7

1246 ± 323

14.1 ± 11.8

5.83 ± 0.41

24.7 ± 6.9

335

1–2

2049 ± 303

20.5 ± 14.4

5.16 ± 0.86

31.3 ± 12.9

86

3–4

1752 ± 386

11.9 ± 11.0

5.52 ± 0.63

21.9 ± 9.8

194

0.03–0.1

1866 ± 243

22.8 ± 11.2

4.87 ± 0.67

41.7 ± 12.2

MM

FM

LP

IP

AA

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Table 2 Classification result of the DA of 1882 relevés shown in %; predicted group: 39 plant communities (see Table 3), independent variables: abundance values of 544 species. Overall 88.3% of the relevés are correctly classified. Identification numbers of the communities (ID) given in Table 3.

Predicted classification of plant communities of the Discriminant Analysis

O riginal classificaon of plant com m unies

ID-No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39

1 100 -

2 100 -

3 100 9.1 0.5

4 90 0.4 -

5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 1.4 1.4 - - - - - - - - - - - - - - - - - - - - - 7 - 82 - - - - - - - 9.1 - - - - - - - - 9.1 - - - - - - - - - - - - - - - - - 89 - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2.7 - 5.4 - 2.7 - - 100 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 100 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 80 - - - - - - - - - - - - 20 - - - - - - - - - - - - - - - - - - - - - 100 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 82 - - - - - - - - - - - - - - - - - - - - - - - - - 9.1 - 9.1 - - - - - - - 73 - 18 - - - 9.1 - - - - - - - - - - - - - - - - - - - - - - - - - - - - 85 5.4 2.2 - - 1.1 - - - - - - - - - - 1.1 - - - - - - - 5.4 - - - - - - - - - 1.9 87 - - - 1.9 - - - - - - - - - - - - - - - - - - 7.7 - 1.9 - - - - - - - - - - 93 - - - - - - - - - - - - - - - - - - - - - 7.4 - - - - - - - - - - - - 89 - - - - - - - - - - - - - - - - - - - - 11 - - - - - - - - - - 17 - - 83 - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2.9 - - - - - 79 - - - - - - - - - - - - - - - - - 2.9 15 - - - - - - - - - - - - - - - 98 - - - - - - - - - - - - - - - - - 2.5 - - - - - - - - - - - - - - - - 92 - - - - - - - - - - 4 - - - - - - - 4 - - - - - - - - - - - - - - - - 87 - - - - - - - - - - - - - - - 13 - - - - - - - - - - - - - - - - - - 75 - - - - - - - - - - - - - - 25 - - - - - - - - - - - - 4.8 - - - - - - 86 - - - - - - - - - - - - - 4.8 4.8 - - - - - - - - - - - - - - - - - - - 40 - - - - - - - - - - - - 60 - - - - - - - - - - - - - - - - - - - - - 79 - - 3.4 - - - - - - - - - - 17 - - - - - - - - - - - - - - - - - - - - - 100 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2.6 - 82 - - - 5.3 - - - - - - - 11 - - - - - - 1.5 - - - - - - - - - - - - - - - - 79 - - - - - - - - - - 20 - - - - - - - - - - - - - - - - - - - - - - - - 100 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 88 - - - - - - 13 - - - - - - - - - - - - - - - - - - - - - - - - - - - 86 14 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 73 - - - - - - 18 - - - - - - - - - - - - - - - - - - - - - - - - - - - - 100 - - - - - - - - - - - - - 9.1 - - - - - - - - - - - - - - - - - - - - 91 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 83 - 11 - 5.6 - - - - - - - - 0.7 - - - - - - 1.4 - - - - - - - - 0.7 - - - - - - 91 5.8 - - - - 0.4 0.4 - - - 0.4 0.2 0.2 0.6 - 2.8 - - - - - - - - - - - - - 0.2 - - - 0.2 94 - 0.2 - - - - - - - - - - - - 6.7 - - - - - - - - - - - - - - - - - - - 13 80 - 0.2 - 0.2 - - - - - 0.5 - - - 1.2 - 0.7 - - - - 0.7 0.2 1 - - - 0.5 1.4 - - - 0.2 1.2 - 91

sification of all (1882) relevés, based on species composition. To reduce the noise from rare species, we included only species occurring more than four times in the data set or with more than 60% abundance in a single community, giving an overall species pool of 544 species. The DA explained 88.3% of all vegetation relevés (i.e. 1662 of 1882 relevés) as correctly classified (Table 2), only 11.7% were misclassified. In the next step, the misclassified relevés calculated by the DA were reviewed thoroughly by comparing the species composition of these relevés with the typical species composition from the literature. As a consequence, we changed the classification of 83 relevés, according to the predicted syntaxonomic community of the DA. The reason of changing these relevés was the high abundance values of species belonging to the predicted community of the DA. Moreover, we could not detect in these relevés any character species given in literature, which confirmed the former stated plant community. The classification of other 54 relevés was left unchanged, because the character species and differential species – even if their abundance values were low – confirmed the original phytosociological classification. Finally, we eliminated 83

relevés, because they just had short species lists, so that a clear classification to a defined plant community was not possible. Since the number of relevés varies greatly between individual plant communities (Table 3), we used the mean abundance of species belonging to a community for a new data set. This data set was used in further nonmetric multidimensional scaling (NMS) to highlight the correlation between plant communities and the factors land-use intensity, altitude, slope and pH of the topsoil. As the pH is rarely indicated in literature, we calculated the missing pH values by using the R-values (=soil reaction) of Ellenberg et al. (1992) adapted for Austria by Karrer (Englisch et al., 1991; Karrer and Kilian, 1990; Karrer, 1992; Pichler and Karrer, 1991). According to Warmelink et al. (2005) we calculated a mean Rvalue for each relevé on the basis of the species presented in the relevé. A calibration curve was given for the measured pH-values of the 375 own recorded relevés, from which the missing pHvalues were afterwards predicted by linear regression. Thereby we achieved a highly significant correlation between R-value and soil pH (R2 = 0.575, p < 0.001), which is even higher than the one stated in Diekmann (2003).

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Table 3 Grassland plant communities of the Eastern Alps with main ecological and floristic characteristics and their respective identification numbers (ID); Abbreviation codes of land-use types (LU) given in Table 1; bold letters describe the main land-use type of each community. Id-No. Community name 1 2 3 4

5 6 7 8 9 10 11 12 13

14 15 16

Alchemillo-Poetum supinaeb Alnetum viridisa Angelico-Cirsietum oleraceib Campanulo scheuchzeri-Festucetum noricaea Caricetum curvulaea Caricetum davallianaea Caricetum ferrugineaea Caricetum sempervirentisa Carici curvulae-Nardetuma Carlino acaulis-Brometumb Chaerophyllo-Ranunculetum aconitifoliib Crepido-Cynosuretumb Crepido-Festucetum commutataeb Deschampsio cespitosaePoetum alpinaeb Elyno-Caricetum rosaea Empetro-Vaccinietum gaultherioidisa

17 18

Festucetum picturataea Festuco-Agrostietumc

19

Gentianello anisodontae-Festucetum variaee Geranio lividi-Trisetetumb Gymnadenio-Nardetumb Hygrocaricetum curvulaea Hypochoerido uniflorae-Festucetum paniculataea Loiseloirio-Caricetum curvulaea Lolietum multifloraeb Onobrychido viciifoliae-Brometumb Pastinaco-Arrhenateretumb Poo-Trisetetumb Potentillo erectae-Brachypodietum pinnatib Pulsatillo alpinae-Festucetum noricaed Ranunculo bulbosi-Arrhenatheretumb Ranunculo-repentisAlopecuretum pratensisb Rhododendretum ferrugineia

20 21 22 23

24 25 26 27 28 29

30 31 32

33

a

34

Rumicetum alpini

35

Selino-Molinetum caeruleaeb

36

Seslerio-Caricetum sempervirentisa Sieversio-Nardetum strictaea

37 38 39

a b c d e

Trifolio thalii-Festucetum nigricantisa Trisetetum flavescentisb

Grabherr and Mucina (1993). Mucina et al. (1993). Ellenberg (1996). Grabner and Heiselmayer (2002). Wallossek (1999).

Slope angle (◦ ) mean±s.d.

Land use types (LU)

Data set

No. of relevés

No. of sites

Altitude (m a.s.l) mean ± s.d.

IP UM; LP AA; LP AA; UM

Lit. & own Literature Literature Lit. & own

5 7 6 71

3 1 1 13

1428 1760 1100 2035

± ± ± ±

296 23 0 153

3.4 27.6 2.2 27.2

± ± ± ±

3.8 4.2 3.9 14.5

5.79 4.45 5.73 5.69

± ± ± ±

0.26 0.16 0.12 0.46

19.8 30.0 20.3 44.2

± ± ± ±

5.9 3.6 7.3 10.1

UM AA; UM AA; UM; LP UM UM; MM LP; IP UM; MM; LP; IP AA; UM; MM; LP; IP LP

Literature Lit. & own Literature Lit. & own Literature Literature Lit. & own

11 37 9 13 5 7 11

2 13 1 4 2 1 6

2349 1827 1903 1953 2064 1699 1577

± ± ± ± ± ± ±

87 209 33 162 340 63 250

4.5 11.9 17.1 26.5 12.6 20.6 10.0

± ± ± ± ± ± ±

4.2 8.8 5.7 13.1 8.1 14.9 5.5

3.63 5.64 5.54 4.43 4.22 5.81 5.42

± ± ± ± ± ± ±

0.28 0.44 0.14 0.63 0.50 0.19 0.50

16.2 28.9 46.8 30.8 36.8 33.4 22.0

± ± ± ± ± ± ±

7.4 8.4 3.1 6.8 11.8 5.7 7.0

Lit. & own

11

2

1301 ± 193

2.9 ± 4.6

6.35 ± 0.55

28.9 ± 7.4

Lit. & own

93

19

1916 ± 181

14.6 ± 11.9

5.69 ± 0.49

27.6 ± 12.2

AA; UM; LP LP AA; UM; MM; FM; LP AA; UM; LP UM; MM; FM AA; UM

Lit. & own Literature Lit. & own

52 27 18

22 1 7

1950 ± 170 2483 ± 24 1826 ± 130

10.5 ± 12.3 23.1 ± 13.6 15.0 ± 8.9

5.05 ± 0.57 5.63 ± 0.54 4.46 ± 0.31

23.5 ± 7.2 18.0 ± 8.7 42.3 ± 12.8

Literature Lit. & own

6 34

3 17

2162 ± 107 1724 ± 250

15.0 ± 10.9 20.0 ± 15.9

4.75 ± 0.46 4.91 ± 0.41

31.0 ± 8.2 32.3 ± 13.1

Literature

40

2

2080 ± 103

22.6 ± 7.1

4.40 ± 0.31

39.5 ± 9.4

AA; LP AA; UM LP FM

Literature Literature Literature Lit. & own

25 15 8 21

3 1 2 6

1426 1425 2440 2106

AA; UM

Literature

5

1

2460 ± 0

7.6 ± 5.3

4.11 ± 0.25

17.6 ± 11.6

MM; FM; IP Lit. & own MM; FM Own

29 3

7 1

900 ± 207 1240 ± 10

7.9 ± 10.6 9.3 ± 5.1

6.20 ± 0.24 6.60 ± 0.20

15.8 ± 3.4 32.0 ± 2.6

UM; LP Lit. & own UM Lit. & own MM; FM; IP Own

38 65 10

15 10 4

1026 ± 209 954 ± 232 1372 ± 157

15.5 ± 15.4 4.2 ± 2.1 18.4 ± 17.6

6.17 ± 0.39 5.66 ± 0.41 5.90 ± 0.36

20.7 ± 5.3 26.3 ± 5.1 35.4 ± 10.1

FM; IP

Literature

8

1

1832 ± 31

23.8 ± 5.2

5.68 ± 0.27

44.5 ± 6.6

LP

Literature

7

1

1100 ± 0

26.1 ± 10.9

5.81 ± 0.28

25.7 ± 6.8

MM; IP

Lit. & own

11

5

1048 ± 135

3.4 ± 4.8

5.92 ± 0.21

17.0 ± 4.5

AA; UM; MM AA; UM; MM; LP AA; UM; MM; LP AA; UM; LP

Literature

4

2

1918 ± 223

17.5 ± 15.5

4.64 ± 0.44

28.2 ± 12.3

Literature

11

3

2047 ± 197

8.4 ± 6.6

5.43 ± 0.86

12.5 ± 3.5

Literature

18

2

1486 ± 195

7.1 ± 3.0

5.61 ± 0.50

23.1 ± 12.7

Lit. & own

139

15

1817 ± 252

28.4 ± 10.1

6.06 ± 0.56

44.8 ± 12.9

UM; MM; FM; LP; IP IP

Lit. & own

468

74

1876 ± 219

17.6 ± 8.9

4.86 ± 0.62

38.7 ± 12.8

Literature

30

8

2016 ± 182

35.7 ± 17.3

5.56 ± 0.44

38.1 ± 12.5

UM; LP

Lit. & own

421

58

1559 ± 289

17.55 ± 11.6

5.72 ± 0.50

29.9 ± 10.1

± ± ± ±

147 52 170 133

22.7 9.7 6.6 18.9

± ± ± ±

7.8 6.5 6.9 12.1

pH (0–10 cm) mean ± s.d.

6.12 5.23 3.78 5.03

± ± ± ±

0.41 0.40 0.27 0.39

No. of species mean ± s.d.

41.2 33.6 14.4 43.7

± ± ± ±

9.2 4.7 7.7 10.9

438

C. Lüth et al. / Flora 206 (2011) 433–443

Fig. 2. NMS output graph of 39 communities and correlation between the factors land-use intensity (LUi), altitude, pH and slope: arrows indicate direction and strength of correlation of variables with R2 = 0.2. Identification numbers of the communities (ID) given in Table 3. Abbreviations of main land-use types given in Table 1.

The NMS calculation was performed with the slow and thorough autopilot mode (for parameter settings and options see McCune and Mefford, 1999) by using the Relative Sørensen distance measure; the cutoff R2 value for site factors was set at 0.2. For the diagram (Fig. 2) the stress in relation to dimensionality is stated, which visualizes the departure from monotonicity in the plot of distance – the lower the stress, the better the fit to monotonicity (McCune and Grace, 2002). Calculations were done with SPSS 15.0.1 (SPSS Inc., 2006) and PC-ORD 5.01 (McCune and Mefford, 1999). Map drawings were plotted in ArcView 3.3 (ESRI Inc., 2002). Results Phytosociological communities On the basis of floristic composition we distinguished 39 communities, listed in Table 3 with a summary of their main ecological and floristic characteristics. We could detect that 23 communities occur in unfertilized meadows: ten of them were established mainly in unfertilized meadows and three (Nos. 7, 10, 30) were restricted to this landuse type. Furthermore we found that some communities, stated in literature as pasture communities, were not just restricted to grazed sites: The Rumicetum alpini (No. 34) occurs also on fertilized meadows mown once a year and grazed in autumn. This is similar to the Crepido-Festucetum commutatae (No. 13), which additionally establishes also in unfertilized meadows. The Deschampsio cespitosae-Poetum alpinae (No. 14) occurs in every type of land use except in fodder meadows from valley regions. On the other hand, we found meadow communities which occur in pastures: The Angelico-Cirsietum oleracei (No. 3) occurs in pastures and the Ranunculo repentis-Alopecuretum pratensis (No. 32) in fertilized meadows as well as in intensively used pastures. The Pastinaco-

Arrhenatheretum (No. 27), which is typical for fertilized meadows mown 2–3 times a year, does also establish in meadows mown once a year with pasturing in autumn and in intensively used pastures. Finally the Trisetetum flavescentis (No. 39) occurs in every land-use type except in abandoned areas. Relating to the mean number of species, biodiversity is mainly affected by land use, but is also affected to a lesser extent by altitude and bedrock. Consequently, the most species-rich communities are found in mainly unfertilized mountain meadows, mown once a year or less frequently every second to third year as well as in rather young abandoned areas (Nos. 4, 7, 23, 30, 36). As a rule, these communities occur on limestone or calcareous mica schist bedrock (Fig. 3), which explains a pH of the topsoil of >5. On the other hand, we found lowest biodiversity in fertilized meadows, intensively used pastures and high alpine lightly used pastures above silicate bedrock (pH < 3.8), where just a few species (e.g. Carex curvula) are able to create a close vegetation cover (Nos. 5, 22, 25, 32, 34). The result of the NMS classification recommends a twodimensional solution (Fig. 2) with a final stress of 15.5% for all 39 plant communities. Land-use intensity and pH have an inverse relationship to altitude and slope, whereby among all factors slope has the lowest influence on plant communities of all site factors. Referring to site factors, the 39 plant communities can be characterized as follows (Fig. 2). Highly fertilized grasslands with either frequent mowing per year (mostly three to five times) or intensively used pastures are concentrated in the valleys (Nos. 1, 3, 25, 32, 34) or near flat plateaus. Due to fertilization the soil offers high pH values. Furthermore, plant communities mown twice a year (Nos. 27, 28, 31) are clearly located at higher altitudes than intensively used ones, but at lower altitudes than lightly used grasslands. Fertilized meadows, mown once a year and grazed in autumn, when cattle return from the high alpine pastures occur in a mountain range between 1300 m and 2000 m a.s.l. (e.g. Nos. 11, 20, 39). Unfertilized meadows were found in the northern limestone Alps at the lowest altitude of about 1200 m a.s.l. (No. 26, Fig. 3). Their species composition is very similar to fertilized grasslands from the valleys, but is distinguished by the absence of species that occur with the use of fertilizer. In the central massive the unfertilized meadows spread up to 2000 m a.s.l. Consequently, communities growing on calcareous bedrock (Nos. 6, 7, 10, 21, 29, 30) stand out from those on siliceous or mica schist bedrock (Nos. 4, 18, 37). Locations at high altitudes in the Eastern Alps, where grazing is possible, are characterized by pastures with the dominant taxon Carex curvula s.l. (Nos. 5, 9, 15, 22, 24). Abandoned areas are located on the steepest sites, with a mean slope of 22.5◦ (Table 1). They can be divided into hydrophilous abandoned areas on calcareous bedrock (No. 35), herb-rich ones (Nos. 8, 23, 36, 38) and abandoned areas dominated by shrubs (No. 2), which are similar to pastures dominated by shrubs (No. 33). Distribution patterns of plant communities The large data set of 1799 vegetation relevés provides detailed information about distribution patterns of current grassland communities over a broad cross-section of the Eastern Alps (Fig. 3). Besides fertilized grassland communities from the valleys, the most frequent plant communities of mountain grasslands are the usually fertilized Trisetetum flavescentis (No. 39, 421 relevés) and the usually unfertilized Sieversio-Nardetum strictae (No. 37, 468 relevés). Some communities are locally restricted (Fig. 3), based on the fact that some species only grow on specific bedrocks and the distribution area of some species is peripherally situated in the research area: The Crepido-Cynosuretum (No. 12) occurs only in northern limestone pastures. The Onobrychido viciifoliae-Brometum (No.

C. Lüth et al. / Flora 206 (2011) 433–443

439

Fig. 3. Distribution pattern of plant communities. Dark grey lines: rivers and valleys, light grey areas: limestone or calcareous mica schist regions, white areas: silicate regions. Identification numbers of the communities (ID) given in Table 3.

26) is located in the northern limestone alps, in montane altitude zones (at about 1200 m a.s.l.), similar to the Potentillo erectaeBrachypodietum pinnati (No. 29), which is found in steep slopes of Northern Tyrol in unfertilized, annually mown meadows. The Seslerio-Caricetum sempervirentis (No. 36) is found in limestone or calcareous mica schist regions over the entire Eastern Alps. The Campanulo scheuchzeri-Festucetum noricae (No. 4), however, is specific for the southern limestone and mica schist areas. The Festucetum picturatae (No. 17), the Gentianello anisodontaeFestucetum variae (No. 19) which occurs in the far south, the Hypochoerido uniflorae-Festucetum paniculatae (No. 23), the Trifolio thalii-Festucetum nigricantis (No. 38), as well as the Pulsatillo alpinae-Festucetum noricae (No. 30) are endemic in South Tyrol or East Tyrol, because the centers of distribution of the eponymous taxa Festuca picturata, Festuca varia, Festuca paniculata, Festuca nigricans and Festuca norica are in southern and/or south-eastern regions of the Eastern Alps.

Discussion In the Alps the geomorphology forces man to many different types of agricultural land use, resulting in a high number of grassland plant communities (Maurer et al., 2006). The communities reflect the anthropogenic influence and topographic, edaphic and climatic factors (Marini et al., 2007) and they show the dependency on those factors, which have a discriminative influence on them.

Classification The identified 39 grassland communities were clearly classified by the DA from their plant composition. Thereby mismatches in the diagonal matrix (Table 2) are explained by the absence of character species or incomplete species lists in single relevés. Most of these mismatches belong to the Sieversio-Nardetum strictae. The Sieversio-Nardetum strictae is one of the dominant plant communities in alpine pastures and meadows, with a wide range of growing conditions (Grabherr and Mucina, 1993). Therefore, a broad spectrum of species can be found, for which species with regional restrictions are responsible. This is similar to the mismatched relevés of the Trisetetum flavescentis. The community is the second most common grassland community in the Eastern Alps and displaces the Sieversio-Nardetum strictae in middle mountain range levels (Mucina et al., 1993). The community becomes established when grasslands are fertilized with manure once a year and mown once or twice a year, but grazed in autumn after the return of livestock from high alpine summer pastures (Knapp and Knapp, 1952; Mucina et al., 1993); in certain cases the community also occurs in lightly to intensively used pastures. The link between fertilized and unfertilized mountain meadows is represented by the Festuco-Agrostietum (Ellenberg, 1996), which was also found in nearly all land-use types (Table 3). Furthermore, we found that the communities Crepido-Festucetum commutatae, Rumicetum alpini and Deschampsio cespitosae-Poetum alpinae, which are associations of pasture alliances (Grabherr and Mucina,

440

C. Lüth et al. / Flora 206 (2011) 433–443

1993; Mucina et al., 1993), are not just restricted to pastures, but occur also in (predominantly fertilized) meadows. On the contrary we observed the Angelico-Cirsietum oleracei, usually described as a meadow community with upright forbs (Mucina et al., 1993), but here observed as a hydrophilous pasture community. We also identified the communities Pulsatillo alpinae-Festucetum noricae, Carlino acaulis-Brometum and Caricetum ferruginae belonging exclusively to unfertilized meadows. The same land-use type is necessary for the Caricetum davallianae, Caricetum sempervirentis, Gymnadenio-Nardetum and Hypochoerido uniflorae-Festucetum paniculatae. When the mowing is abandoned these communities can still be found in young abandoned areas, but they disappear in older ones. Impact of environmental factors Key factor in forming specific grassland plant communities in meadows and pastures is human influence. The higher the human impact, the fewer species and community types occur. In favourable areas (flat areas in lowlands) low diversity is a result of mowing several (3–5) times a year (Marini et al., 2007; Oberdorfer and Müller, 1993), which allows only those species to grow that have either very short life cycles or can clonally reproduce (Bahn et al., 1994). This is similar to highly stocked and fertilized pastures that result in grassland communities with a few species that resist trampling (Ellenberg, 1996). A strong dependency on land-use intensity is also revealed in fertilized meadows mown twice a year. Since accessibility is one of the major factors in fertilizing grasslands (Macdonald et al., 2000; Tasser and Tappeiner, 2002), poorly accessible unfertilized meadows form another group of meadow communities. Due to the shorter growth period, decrease of summer temperatures and increase of rainfall with increasing altitude (Ellenberg, 1996; Körner, 2007), the altitude site factor plays a decisive role in forming specific communities. Especially in unfertilized meadows the high number of 23 different plant communities is explained by the extensive amplitude of elevation, which ranges from about 1250 to 2500 m a.s.l. Here we found the Onobrychido-viciifoliae-Brometum and the Potentillo erectaeBrachypodietum pinnati from valley regions depending on low land-use intensity and the absence of fertilization (Mucina et al., 1993). In the highest located meadows the communities Campanulo scheuchzeri-Festucetum noricae and Sieversio-Nardetum strictae establish. In lightly stocked pastures and abandoned areas the species composition is similar to unfertilized meadows, as long as low anthropogenic influence and similar site factors affect the areas. The grassland communities at highest altitudes can be separated clearly by means of species, in that only few specific taxa (e.g. Carex curvula) generate a close vegetation cover (Choler and Michalet, 2002), and if at all, they are used as pastures. Slope is found to be relevant in meadows and abandoned areas. Meadow communities on steep slopes present a species composition similar to xerophilous grasslands, due to drainage (Seeber and Seeber, 2005). However, on steep slopes land use is no longer profitable and grasslands are nowadays often abandoned or infrequently managed (Giupponi et al., 2006; Tasser et al., 2007), which explains why abandoned areas were found on steepest sites. Near and below the timberline abandonment leads to an increase of shrubs which displace herbs, represented by the Alnetum viridis, the Empetro-Vaccinietum gaultherioidis or the Rhododendretum ferruginei. The pH value of the topsoil is affected by either fertilizing or by bedrock. High pH values result from the absence of humic acids (Marcos et al., 2003), due to intensive fertilizing (Seeber and Seeber, 2005). Apart from fertilizing, higher pH-values (about 5.5–7.2) can

be found in pastures with intensive grazing or in meadows and abandoned areas above calcareous bedrock. On the other hand, low pH-values (about 3.5–5.5) occur in unfertilized grasslands above silicate substrate.

Plant diversity We found grassland communities which refer to specific taxa (e.g. of the genus Festuca sp.), but with local restriction (Pils, 1980) or belonging to a specific bedrock (Fischer et al., 2005), represented by the communities Campanulo scheuchzeri-Festucetum noricae, Gentianello anisodontae-Festucetum variae, Pulsatillo alpinae-Festucetum noricae, Trifolio thalii-Festucetum nigricantis and Hypochoerido uniflorae-Festucetum paniculatae. These rarely occurring communities are characterized by high biodiversity and low land-use intensity (Table 3). The most species-rich meadows are characterized by soil of high pH (Myklestad and Sætersdal, 2004) and do also apply to unfertilized mountain meadows above calcareous bedrock in the Eastern Alps, represented by the Caricetum ferruginei, the Campanulo scheuchzeri-Festucetum noricae and the Seslerio-Caricetum sempervirentis (Table 3). According to the mean number of species we found that speciesrich communities must not be related in any case to altitude and pH of the topsoil. However, land-use intensity and slope significantly affect the number of species and correlate reciprocally to each other: species-poor communities occur in flat areas and with high anthropogenic influence, whereas on steep sites land-use intensity is low and communities provide high vascular plant biodiversity, which is confirmed by Niedrist et al. (2008). Traditional hay meadow management preserves sites with high biodiversity (Garcia, 1992; Maurer et al., 2006; Myklestad and Sætersdal, 2004), which has been confirmed by our studies. The highest number of species and the highest ecosystem diversity are found in unfertilized alpine meadows and young abandoned areas (Tables 1 and 3). Since agri-environmental measures are among the most important instruments for the promotion of environmentally adapted agricultural land use (Matzdorf et al., 2008), a register of the still remaining grassland communities is essential for such measures in the Eastern Alps. Moreover, unfertilized mountain meadows are nowadays increasingly abandoned (Macdonald et al., 2000; Tasser et al., 2001, 2007), nevertheless they play an important role in the culture of the Alps mountain regions (Ender and Grabner, 1997). Therefore, currently unfertilized meadows must be retained in order to ensure that rare communities – being endangered – will continue to be found today in the Eastern Alps and also in the future.

Acknowledgements We thank Prof. Dr. Robert Crawford, Dr. Avril Arthur-Goettig (BioScript International) and Daniela Dellantonio for language revision. We also thank the provincial government of Tyrol for analyzing the soil samples. This work was funded by the European Union within the scope of the Interreg IIIA-Project “DNACharacterisation for certification and valorisation of mountain hay”, supported by the provincial governments of Tyrol and South Tyrol.

Appendix. Sources for information on the vegetation relevés from literature.

C. Lüth et al. / Flora 206 (2011) 433–443

441

Author

Title

Type of publication

Year of publication

Location

No. of relevés

Brunner, B.

Die Vegetation von Bergmähdern im Landschaftsschutzgebiet Nößlachjoch-ObernbergTribulaune Die Vegetation der subalpinen und alpinen Stufe in der Puez-Geisler Gruppe Grünland-Gesellschaften im oberen Paznauner Tal Die Vegetation im oberen Lechtal Die Vegetation des Gaißbergtales. Ein Versuch, das Datenmaterial mit Hilfe der EDV-Anlage zu bearbeiten Die Wirtschaftswiesen des oberen Vinschgaues (Südseite) und ihre Bewirtschaftung Die Vegetation vom Nationalpark Hohe Tauern in Tirol Vegetation von gemähten Bergwiesen und deren Sukzession nach Auflassung der Mahd Die Vegetation von Schipisten und angrenzenden Bergmähdern im Raum Hochtannberg Die Lärchenwiesen im Nationalpark Trudner Horn – Pflanzensoziologische Untersuchungen in verschieden bewirtschafteten Wiesen und deren Vergleich mit aufgelassenen Flächen Die alpine Vegetation des hinteren Defreggentales Diversity of mountain meadows in the inner alpine valley Virgental/Eastern Tyrol Analyse der Vegetationsund Erosionsverteilung in Abhängigkeit von Bewirtschaftungsänderungen am Beispiel Kaserstattalm Die Vegetation und ihre Gliederung in den Leoganger Steinbergen Wirtschaftswiesen der Nordhänge und Tallagen im oberen Vinschgau aus vegetationskundlicher und futterbaulicher Sicht Die Vegetationsverhältnisse des Pflerschertales Auswirkungen des Pistenschilaufes auf die Pflanzengesellschaften der Komperdellalm (Tirol) Die Vegetation im Bereich des Dreiländerecks bei Nauders am Reschenpass Die Vegetation der inneren Pfunderer Täler Vegetation der Wirtschaftswiesen von Trafoi am Stilfser Joch

Diploma thesis, University of Innsbruck (AT)

1999

Obernberg, North Tyrol

118

Dissertation, University of Innsbruck (AT)

1982

St. Christina, Gröden, South Tyrol

11

Phytocoenologia 6: 287–303 Diploma thesis, University of Innsbruck (AT) Dissertation, University of Innsbruck (AT)

1979

Galtür, Paznaun, North Tyrol Elbigenalp, Lechtal, North Tyrol Gaißbergtal, Obergurgl, North Tyrol

54

Dalla Torre, M.

Dierschke, H. Dirrhammer, H. Duelli, M.

Ebner, S.

Egger, G.

Ender, M.

Flecker, K.

Florian, C.

Gander, M. Grabner, S., Heiselmayer, P

Gufler, R.

Gumpelmayer F.

Hellriegl, S.

Keim, K. Kirchmeir, H.

Lechner, C.

Lechner, G. Lüth, Ch.

2008 1977

18 27

Diploma thesis University of Innsbruck (AT)

1996

Vinschgau, South Tyrol

73

Personal unpubl. relevés

2006

Hohe Tauern, East Tyrol

86

Diploma thesis, University of Innsbruck (AT)

1997

Tannberg, Vorarlberg

63

Diploma thesis, University of Innsbruck (AT)

1996

Lech, Vorarlberg

29

Diploma thesis University of Innsbruck (AT)

1995

Bozner Unterland, South Tyrol

35

Diploma thesis University of Innsbruck (AT) Razprave IV. Razreda Sazu 43/3: 167–184

1984

Defreggental, East Tyrol Virgental, East Tyrol

13

2002

8

Diploma thesis, University of Innsbruck (AT)

1999

Kasertatt, Stubaital, North Tyrol

28

Dissertation, University of Innsbruck (AT)

1967

Leoganger Steinberge, Salzburg

10

Diploma thesis University of Innsbruck (AT)

1996

Vinschgau, South Tyrol

77

Dissertation, University of Innsbruck (AT) Diploma thesis University of Vienna (AT)

1967

Pflersch, South Tyrol

26

1996

Oberes Gericht, North Tyrol

19

Diploma thesis, University of Innsbruck, (AT)

1995

Nauders, Noth Tyrol

31

Dissertation, University of Innsbruck (AT) Personal unpubl. relevés

1969

Pfunders, South Tyrol

26

2002

Trafoital, South Tyrol

2

442

C. Lüth et al. / Flora 206 (2011) 433–443

Appendix A (Continued ) Author

Title

Type of publication

Year of publication

Location

No. of relevés

Mayer, Ch.

Landschaftsentwicklung in der Gemeinde St. Leonard in Passeier (Südtirol, Italien) unter besonderer Berücksichtigung der floristischen Biodiversität und der Bodendurchwurzelung Die Vegetation der Bergmähder im Valsertal und ihre Dynamik Die Vegetation des Grödner Tales/Südtirol Analyse der Vegetationsverteilung in Abhängigkeit der Bewirtschaftungsänderung auf den Waltner Mähdern Vegetation der Sextener Dolomiten (subalpine und alpine Stufe) Vegetationskundliche Untersuchung, Kartierung und Bewertung der Kulturlandschaft Kitzbühel-Aurach Die Vegetation der alpinen Stufe in den östlichen Pragser Dolomiten Pflanzengesellschaften im Raum von Brixen mit besonderer Berücksichtigung der Trockenvegetation Die Vegetation der alpinen Stufe der Texelgruppe

Diploma thesis University of Innsbruck (AT)

2004

Passeiertal, South Tyrol

13

Diploma thesis, University of Innsbruck (AT)

2002

Valsertal, Noth Tyrol

118

Giessener geogr. Schriften 47: 287 Diploma thesis University of Innsbruck (AT)

1980

Grödnertal, South Tyrol

7

1998

Walten, Passeiertal, South Tyrol

40

Dissertation, University of Innsbruck (AT)

1975

Sexten, South Tyrol

11

Diploma thesis University of Innsbruck (AT)

1997

Leukental, North Tyrol

50

Dissertation, University of Innsbruck(AT)

1979

Prags, Dolomites, South Tyrol

14

Dissertation, University of Innsbruck (AT)

1967

Brixen, South Tyrol

8

Diploma thesis University of Innsbruck (AT)

1982

18

Dissertation, University of Innsbruck (AT) Diploma thesis, University of Innsbruck (AT)

1981

Vinschgau, Texelgruppe, South Tyrol Kaisergebirge, North Tyrol Plätzwiese, Prags, South Tyrol

Mayer, R.

Meurer, M. Mulser, J.

Nieder-brunner, F.

Noichl, M.

Oberhammer M.

Putzer, J.

Raffl, E.

Smettan, H. Steinmair, V.

Tasser, E. Tasser, E.

Thimm, I.

Thomaser, J. Unterhofer, Ch.

Unterlug-gauer, P. Vorhauser, K.

Wallossek, C.

Winkler, J.

Die Pflanzengesellschaften des Kaisergebirges/Tirol Die Vegetation von unterschiedlich genutzten Almflächen auf der Plätzwiese Die Vegetation der Talwiesen des Stubaitales Vegetationsaufnahmen von Bürstlingsrasen des Monte Bondone Die Vegetation der alpinen und subalpinen Stufe des Sonnwendgebirges Die Vegetation des Peitlerkofels in Südtirol. Welche Landschaftsskala eignet sich zur Klärung der Fließgewässerqualität? Die Vegetation in Vent und Rofen (Ötztal, Tirol) Vegetationskundliche Untersuchungen im Bereich der Eggentaler Alm (Südtirol) Vegetationskundlichökologische Untersuchungen in der alpinen Stufe am SW-Rand der Dolomiten Populationsbiologische Untersuchungen an zwei eng verwandten Sippen in alpinen Rasen

1999

6 92

Personal unpubl. relevés

2004

Stubaital, North Tyrol

26

Personal unpubl. relevés

2005

Mt. Bondone, Trentino, Italy

22

Dissertation, University of Innsbruck (AT)

1950

Rofan, North Tyrol

20

Veröff Museum Ferdinandeum 47: 67–119 Diploma thesis University of Innsbruck (AT)

1967

Gardertal, South Tyrol

4

2006

Pfunderertal, South Tyrol

55

2003

Ötztal, North Tyrol

65

1998

Eggenberg, South Tyrol

116

Dissertationes Botanicae 154 (IT)

1990

Lafatscher Joch, Latemar, South Tyrol

27

Diploma thesis University of Innsbruck (AT)

1992

Ahrntal, Hasental, South Tyrol

41

Diploma thesis University of Innsbruck (AT) Diploma thesis University of Innsbruck (AT)

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