Classification of the Sieversio montanae-Nardetum strictaein a cross-section of the Eastern Alps

Christian Luth • Erich Tasser • Georg Niedrist •

Josef Dalla Via • Ulrike Tappeiner

Received: 21 April 2009 / Accepted: 15 June 2010

� Springer Science+Business Media B.V. 2010

Abstract The Sieversio montanae-Nardetum stric-

tae is one of the most widespread plant communities

in (sub-) alpine regions of the Alps. Our study

examines the composition, ecology and distribution

of this plant community in the Eastern Alps and

addresses the issue of how the community is to be

classified in the phytosociological system of Nardus-

rich grasslands. Therefore, 357 vegetation releves

were taken from the literature and 115 from our own

inventories were recorded from 2005 to 2007 in

Western Austria (mostly Tyrol) and Northern Italy

(mostly South Tyrol). Additionally, indicator values

of Ellenberg and land-use information were used to

help better interpret the ecological site conditions of

the subgroups. The HCA revealed there the existence

of four groups of the Sieversio montanae-Nardetum

strictae, which were classified to subassociations: (1)

typicum, (2) vaccinietosum, (3) trifolietosum praten-

sis, and (4) seslerietosum albicantis. Besides the

specific plant composition, altitude specifies the first,

land-use intensity the second and third, and the pH of

the topsoil the fourth subassociation. For the Eastern

Alps, the plant community of the Sieversio montanae-

Nardetum strictae should now be reclassified in the

order of Nardetalia and the class of Calluno-Ulicetea.

Finally, this plant community can be further classified

by using the four above-mentioned subassociations.

Keywords Indicator values � Indicator species �Differential species � Subassociation �Land-use types � European Alps

Abbreviations

ANOVA Analysis of Variance

a.s.l. Above sea level

DA Discriminant analysis

EIV Ellenberg indicator values

HCA Hierarchical cluster analysis

ISA Indicator Species Analysis

Introduction

Nardus-rich grasslands result from lightly used pas-

tures or meadows (Peppler 1992). In central Europe,

because of preindustrial low land use (Peppler-

Lisbach and Petersen 2001), such grasslands were

most common and were to be found from forest-free

lowlands up to high alpine regions (Preising 1949).

However, since the second half of the twentieth

century, due to intensification of land use in favorable

C. Luth � U. Tappeiner (&)

Institute of Ecology, University of Innsbruck,

Sternwartestraße 15, 6020 Innsbruck, Austria

e-mail: Ulrike.Tappeiner@uibk.ac.at

E. Tasser � G. Niedrist � U. Tappeiner

Institut for Alpine Environment, European Academy of

Bolzano/Bozen, Drususallee 1, 39100 Bozen, Italy

J. Dalla Via

Research Centre for Agriculture and Forestry Laimburg,

Laimburg 6, 39051 Pfatten, Italy

123

Plant Ecol

DOI 10.1007/s11258-010-9807-9

agricultural areas as well as abandonment of areas that

are difficult to manage (Lavorel et al. 1998; Tappeiner

et al. 1998; Bakker and Berendse 1999; Tasser and

Tappeiner 2002; Niedrist et al. 2008), these grassland

communities have been decreasing (Peppler-Lisbach

and Petersen 2001). Nowadays, even though grassland

communities dominated by Nardus stricta can be

found nearly all over the world (Krajina 1933; Kissling

et al. 2005; Trivedi et al. 2008), they are relicts of

traditional land use (Peppler 1992; Peppler-Lisbach

and Petersen 2001) and are only extensively found in

high alpine regions where they constitute a large part of

the cultural landscape (Grabherr and Mucina 1993).

Sieversio montanae-Nardetum strictae dominates

alpine pastures (Oberdorfer 1978), and the species

Nardus stricta establishes a close sward population

since grazing animals do not eat this plant species

(Ellenberg 1996). In addition to pastures, this plant

community also develops in meadows, where it

exhibits a variety of structural differences including

higher growth, increased frequency of tall forbs and

the absence of higher dwarf-shrubs such as Rhodo-

dendron ferrugineum (Grabherr and Mucina 1993).

Such structural differences can be influenced by land-

use types. For example, in high alpine regions

meadows are mostly mown once a year, unfertilized

or fertilized with manure, and grazed in autumn after

the return of livestock from high alpine summer

pastures (Knapp and Knapp 1952; Mucina et al.

1993). Moreover, abandonment of such areas does

not inevitably lead to the disappearance of Nardus

stricta, because deer start to graze favorably in such

areas, thereby keeping them clear and allowing scrub

and/or forest communities to begin to slowly colonize

the land (Stussi 1970; Peppler 1992). However,

literature detailing the differences or similarities in

the species composition of the Sieversio montanae-

Nardetum strictae is currently not available for the

Eastern Alps.

In subalpine and alpine regions, the Sieversio

montanae-Nardetum strictae establishes mostly

between 1,800 and 2,200 m a.s.l. (Oberdorfer 1978),

but is also found in lower regions at about 1,600 m

a.s.l. (Peppler-Lisbach and Petersen 2001). According

to the pH of the topsoil, the community establishes

usually above acidophilous bedrock (Gigon 1971;

Marschall and Dietl 1974; Oberdorfer 1978; Peppler

1992; Ellenberg 1996; Peppler-Lisbach and Schroder

2004), but is sometimes also found above calcareous

mica schist or calcareous marl (Grabherr and Mucina

1993). As the Sieversio montanae-Nardetum strictae

offers a wide range of growing conditions (Grabherr

and Mucina 1993), a broad spectrum of species can be

found, although some of them are regionally restricted.

Phytosociological classification has so far paid insuf-

ficient attention to the influencing factors such as land-

use intensity, altitude, pH and slope (Grabherr and

Mucina 1993; Peppler-Lisbach and Petersen 2001) and

their combination among each other.

For alpine regions in Austria, grasslands dominated

by Nardus stricta are already classified into different

alliances and classes (Grabherr and Mucina 1993;

Mucina et al. 1993). In the montane regions, Nardus

grasslands belong to the order Nardetalia (class of

Calluno-Ulicetea) (Krahulec 1985; Krahulec 1988;

Mucina et al. 1993). On the other hand, subalpine to

alpine Nardus-rich grasslands refer to the association

of Sieversio montanae-Nardetum strictae (Ludi 1948),

which belongs to the monotypic alliance Nardion

strictae and the order Festucetalia spadiceae (class of

Caricetea curvulae) (Grabherr and Mucina 1993).

However, classification problems arise, because the

Sieversio montanae-Nardetum strictae contains char-

acter species from lowlands that are present in the

order Nardetalia (e.g. Carex pallescens, Hypericum

perforatum) and species that are restricted to the

alpine zone, as well as character species of the

Festucetalia spadiceae (e.g. Geum montanum, Hypo-

chaeris uniflora) (Grabherr and Mucina 1993; Mucina

et al. 1993). Despite these classification difficulties, in

their seminal article Peppler-Lisbach and Petersen

(2001) support the notion that the alliance of the

alpine Nardion strictae can be integrated into the order

Nardetalia as proposed by other authors (Oberdorfer

1959, 1978; Marschall and Dietl 1974; Krahulec

1983).

The Sieversio montanae-Nardetum strictae has

already been classified by several authors for spatially

limited regions (e.g. Ludi 1948; Braun-Blanquet

1949; Hartl 1963; Bischof 1981). Although Peppler

(1992) and Peppler-Lisbach and Petersen (2001)

divide the association into two subassociations with

two variants each, their data mainly refer to Germany,

and only marginally include the Alps. For the Alps,

Heiselmayer (1985) divided the community into three

subassociations; however, the author only refers to the

Radstadter Tauern (Kleinarltal, Salzburg). Thus, to

date, a clear classification of this community in the

Plant Ecol

123

Eastern Alps is still missing (Grabherr and Mucina

1993). Therefore, the major objective of this study was

to clearly and comprehensively classify the Sieversio

montanae-Nardetum strictae for the entire Eastern

Alps by (1) generating a detailed inventory of this plant

community, (2) characterizing subgroups, and (3)

examining distribution patterns of this community.

Methods

Research area

Most of the vegetation releves were taken from Tyrol

(Austria) and South Tyrol (Italy) with some coming

from Vorarlberg (Austria) and Trentino (Italy). The

research area is situated between 47�360–46�140 N and

10�080–12�420 E and covers an area of about

20,000 km2 (Fig. 1). Annual precipitation ranges from

800 to 2,200 mm with maximum rainfall from June to

July and mean annual temperatures between 0 and 8�C

(Fliri 1998). High ranges of precipitation and annual

temperature are caused by the fact that the vegetation

releves were taken from 1,200 to 2,650 m a.s.l. and

that both the continental and oceanic climatic condi-

tions affect the vegetation in the research area (Fliri

1998). In general, the Eastern Alps are made up of

calcareous sedimentary rocks in the northern and

southern regions and of silicate bedrock in the central

massif, sometimes even with superimposed calcareous

isles (Bogel and Schmidt 1976, Fig. 1). The pH values

of the topsoil (0–10 cm) range from 3.7 to 7.8

(Niedrist et al. 2008), whereas high pH values are

mainly found above calcareous bedrock and low pH

values above silicate bedrock (Scheffer et al. 2002).

Data collection

A total of 357 vegetation releves from 27 different

sites were taken from the literature (Appendix 1)

after the method of Braun-Blanquet (1964). Thereby

the plot size ranges from 12 to 25 m2. We

incorporated only data on vegetation releves having

exact information on land use, geographical coordi-

nates, and site factors. Unfortunately, the data set

from literature just partially covers the research

area. To cover the research area entirely, we

consulted local experts (farmers, park rangers and

district agrarian decision-makers). Altogether we

detected a further 52 sites, where the Sieversio

montanae-Nardetum strictae occurs. We then

recorded 115 vegetation releves and took also soil

samples and pH measurements. Fieldwork took

place from 2005 to 2007. Vegetation releves were

Fig. 1 Location of the research area in the Eastern Alps. Dark gray lines rivers and valleys; light-grey areas: regions with calcareous

bedrock; white areas regions with silicate bedrock

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123

recorded according to the method of Braun-Blanquet

(1964): the plot size was about 16 m2 (4 9 4 m); in

some rare cases it was expanded up to 20 m2, when

rock outcrops constituted a considerable portion of

the plot area. At least two releves were recorded for

each site. The site factors of altitude and slope were

measured and the managing farmers were inter-

viewed to obtain exact information on land use.

Thus, we compiled a comprehensive data set of 472

vegetation releves (Fig. 1). Although the community

is established mostly above silicate bedrock (Grabh-

err and Mucina 1993), we were able to incorporate

149 releves, found above calcareous or mica schist

bedrock. For statistical calculations, we transformed

the scale of Braun-Blanquet (1964) to percental

dominance values (according to Tasser and Tappe-

iner 2004): r = 0.1%, ? = 0.3%, 1 = 2.8%,

2m = 4.5%, 2a = 10%, 2b = 20.5%, 3 = 38%,

4 = 63%, and 5 = 88%.

After analyzing the literature and collecting inter-

views about land use, we divided the grasslands into

three main groups (Table 1): (1) meadows, which

were subdivided into (a) fertilized meadows, mown

once, seldom twice a year, but mainly grazed after

mowing and (b) unfertilized meadows, mown every

year or infrequently every second to third year; (2)

pastures, with animals that are left more or less

unattended on the same site throughout the year; and

(3) young abandoned areas, lying fallow for not more

than 30 years, which were previously mown. Note

that older-abandoned areas were excluded from our

study and that the year of the last management event

was based on the literature or interviews. Land-use

intensity (LUi) was classified according to Tappeiner

et al. 1998 (Tables 1, 2). For meadows, we summed

every human impact Ih (mowing, fertilization) and

divided it by the frequency of these interferences in

years a. The same procedure was applied for

abandoned areas: thereby we summed the years from

the last human impact until now and took the

reciprocal value of it:

LUi ¼X

Ih � a�1

The pasture utilization was indicated as ordinal data

type, ranging from 1 (low grazing intensity) to 3

(intensive grazing) (for details see Tasser et al. 1998).

Data analysis

We first compiled information on vegetation releves

from the literature (Appendix 1), unifying the plant

taxa by using the nomenclature of Fischer et al.

(2005), since the incorporated data from the literature

span a period of more than 40 years. Species and

subspecies were aggregated unless taxonomically

uniformly used by the authors. This resulted in a

data set of 437 species.

We then integrated data from our studies on

vegetation releves from literature and used Hierarchi-

cal (agglomerative) Cluster Analysis (HCA) to reveal

groups. Thereby, we used the Euclidean distance

measure and Ward’s linkage method and the group

membership at each step of cluster formation was

written to a file. We used the Euclidean distance

measure, because the stems of the dendrogram were

much longer for the first 15 groups than the relative

Euclidean distance measure and, therefore, more

‘‘natural’’ (Fig. 2). This leads to aggregations appro-

priate to achieve the goals of this study (McCune and

Grace 2002). Moreover, large differences are weighted

more heavily than several small differences with the

Euclidean distance measure, which results in greater

sensitivity to outliers (McCune and Grace 2002).

We then used Indicator Species Analysis (ISA), as

this method combines information of the concentra-

tion of species abundance in a particular group with

the constancy of occurrence of a species in a particular

Table 1 Main land-use types, where the Sieversio montanae-Nardetum strictae establishes in the Eastern Alps along with their

respective land use (ID), land use intensity (LUi) and number of vegetation releves

ID Name Land use LUi No.of releves

FM Fertilized meadows Fertilized, mown once a year and grazed in autumn 3 48

UM Unfertilized meadows Mown every year, seldom only every 2nd or 3rd year 0.33–1 274

PA Pastures With low to intensive grazing 1–3 102

AA Abandoned areas Abandoned for not more than 30 years 0.03–0.1 48

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123

group (McCune and Grace 2002). The ISA was also

used to select the optimal number of clusters. There-

fore, we calculated indicator values for 335 species

that occur in three or more vegetation releves and

averaged the resulting P values for each cluster step up

to 15 levels of clustering (Fig. 3a). Furthermore, we

Table 2 Main ecological and floristic characteristics of the

four groups revealed by Hierarchical Cluster Analysis (HCA)

of the Sieversio montanae-Nardetum strictae in the Eastern

Alps and the respective land-use intensity (LUi). Abbreviations

of land-use types are given in Table 1

Group 1 Group 2 Group 3 Group 4

Number of releves 104 250 40 78

Number of sites 29 66 7 8

Altitude (m a.s.l.) Mean ± S.D. 1984 ± 240a 1855 ± 232b 1828 ± 171b 1846 ± 145b

Slope (�) Mean ± S.D. 17.6 ± 8.6a 18.3 ± 9.7a 18.2 ± 7.3a 14.9 ± 6.2a

pH (0–10 cm) Mean ± S.D. 4.54 ± 0.20a 4.56 ± 0.35a 4.54 ± 0.05a 4.75 ± 0.53b

Land use types in % referring to the data in Table 1; (effective number per group)

FM 5.1 (2) 23.5 (22) 61.7 (13) 28.4 (11)

UM 21.5 (48) 30.0 (159) 20.7 (22) 20.3 (45)

PA 58.0 (48) 23.3 (47) 7.3 (3) 4.8 (4)

AA 15.4 (6) 23.2 (22) 10.3 (2) 46.5 (18)

LUi mean ± S.D. 0.87 ± 0.47a 0.98 ± 0.75a 1.51 ± 1.02b 0.85 ± 0.93a

Indicator values of Ellenberg (EIV)

L = light Mean ± S.D. 7.43 ± 0.36a 7.09 ± 0.50b 7.26 ± 0.39a,b 6.57 ± 0.72c

T = temperature Mean ± S.D. 1.20 ± 0.46a 1.57 ± 0.53b 2.42 ± 0.53c 1.73 ± 0.73b

K = continentality Mean ± S.D. 3.15 ± 0.29a 3.18 ± 0.42a 3.48 ± 0.20b 3.15 ± 0.38a

F = moisture mean ± S.D. 2.56 ± 0.73a 3.45 ± 0.78b 4.10 ± 0.55c 3.49 ± 0.57b

R = soil reaction Mean ± S.D. 3.13 ± 0.63a 3.45 ± 0.85b 4.44 ± 0.63c 4.23 ± 0.99c

N = nitrogen Mean ± S.D. 2.43 ± 0.44a 2.65 ± 0.53b 3.26 ± 0.37c 3.09 ± 0.76c

Number of species Mean ± S.D. 35.5 ± 12.5a 36.4 ± 12.8a 48.4 ± 8.3b 44.9 ± 9.9b

10 characteristic species (indicator value of the Indicator Species Analysis) x = indicator species with a relative abundance and a

constancy of C50%

x Nardus stricta (62) x Calluna vulgaris(29)

x Festuca rubra agg.(67)

x Alchemilla vulgariss.l. (52)

Geum montanum (24) Vacciniumgaultherioides (23)

x Trifolium pratense(64)

x Agrostis capillaris(47)

Scorzoneroideshelvetica (15)

Vaccinium myrtillus(22)

Hypochaeris uniflora(38)

x Galiumanisophyllon (47)

Pedicularis tuberosa(13)

Vaccinium vitis-idaea(18)

Phleum com-

mutatum (36)

x Trollius europaeus(46)

Phyteuma hemis-

phaericum (13)

Anthoxanthumalpinum (17)

Mutellinaadonidifolia (35)

x Geraniumsylvaticum (45)

Hieracium hoppeanum(12)

Antennaria dioica(14)

x Luzula campestris(34)

x Rhinanthusglacialis (40)

Festuca halleri agg.(10)

Cetraria islandica(11)

Anthoxanthumodoratum (34)

x Phyteumaorbiculare (39)

Polygala comosa (10) Cladonia rangiferina(9)

Arnica montana (30) Briza media (39)

Festuca ovina agg. (9) Erica carnea (8) x Ranunculusmontanus (29)

x Sesleria albicans(33)

Carex pallescens (7) Picea abies (8) Lotus corniculatus(29)

x Trifolium badium(31)

Letters indicate significant differences at P \ 0.01 of Bonferroni post-hoc tests; S.D. standard deviation

Plant Ecol

123

also tallied the number of species that were shown to

be significant indicators (P \ 0.05, i.e. differential

species), then plotted this number against the cluster

steps (Fig. 3b). In order to pick the most ecologically

meaningful point to prune the dendrogram from the

HCA, we took the cluster step with one of the lowest

average P values and compared the highest number of

species with P \ 0.05. Finally, we tested the classi-

fication of the HCA with a Discriminant Analysis

(DA) using the stepwise method.

Differential species for each group were also

identified by the ISA, as P \ 0.05 after 3,000

permutations of the Monte Carlo test. Furthermore

with the ISA the relative abundance, the constancy

and the indicator value for 335 species were calcu-

lated. Thereby the relative abundance is the propor-

tional abundance of a particular species in a particular

group in relation to the abundance of that species in

all groups; the constancy expresses the proportional

frequency of a species in each group. Both values are

expressed as a percentage. Finally, the indicator value

was calculated by multiplying the relative abundance

and the constancy for each species, also expressed as

a percentage (McCune and Grace 2002). Species with

a relative abundance and a constancy of C50% were

identified as indicator species.

To explain ecological interrelations of the groups,

we took site factors and the Ellenberg indicator value

(EIV, Ellenberg et al. 1992) of all (437) species into

account. The EIVs used were L = light, T = tem-

perature, K = continentality, F = moisture, R = soil

reaction and N = nitrogen. We weighted the EIVs

with the abundance values of the species occurring in

each group and compared the means with an Analysis

of Variance (ANOVA). Tests of significance were

calculated with post-hoc tests of Bonferroni at

P \ 0.01.

DA and ANOVA were done with SPSS 15.0.1

(SPSS Inc. 1989–2006), HCA and ISA with PC-ORD

5.01 (McCune and Mefford 1999). Maps were plotted

in ArcViev 3.3 (ESRI Inc. 1992–2002).

Results

Classification of vegetation data

Figure 2 presents a truncated dendrogram of the HCA

showing the first 15 groups. Chaining of 1.86% is

higher than the one calculated with relative Euclidean

distance (0.61%), but the loss of information was

much higher for the first 15 groups (about 35%

retained) than with the Euclidean distance measure

(about 50% retained).

An objective criterion for choosing the number of

groups is given by the following ISA. This calcula-

tion presents the lowest average P value of the Monte

Carlo tests in group five, but with a difference of only

0.003 from group four (Fig. 3a). As there are four

more differential species (142) in group four than in

group five (Fig. 3b), which means a difference of

1.2%, we have decided to prune the dendrogram into

four groups. The dendrogram in Fig. 2 shows a loss

of information at 73.8% (26.2% retained) for four

groups.

Discriminant analysis explained 97.5% of the 472

vegetation releves as correctly classified. We then

compared the abundance values of the species of the

12 misclassified releves with the mean abundance

values of the four groups of the HCA, and discovered

that the species composition fitted more precisely to

the predicted group found by the DA. Thus, the

classification of the misclassified releves was chan-

ged accordingly. Table 2 summarizes the main

Fig. 2 Dendrogram for the first 15 groups of the HCA using the Euclidean distance measure and the Wards method

Plant Ecol

123

ecological and floristic characteristics of the four

groups and presents ten differential species with the

highest indicator values for each group. The results of

the ISA of all 142 differential species are presented in

Appendix 2.

Characterization of the four groups

of the Sieversio montanae-Nardetum strictae

The four groups are ranked as subassociations

because of ecological differences and numerous

specific indicator species that were found for each

group. On the basis of ecological characteristics and

floristic composition (see Table 2, Appendix 2), the

four subassociations of the Sieversio montanae-

Nardetum strictae can be characterized as follows:

Subassociation typicum

The first group describes mainly pastures and is

located at higher altitudes: the difference given by the

ecological factor altitude is, therefore, highly signif-

icant for the other groups. Of the EIVs, the T, F, R,

and N values are significantly lower than in the other

groups (Table 2). Additionally, the group is charac-

terized by having a low number of species, but the

highest abundance rates of the community’s epony-

mous species Nardus stricta and Geum montanum.

Together with Carex pallescens, Hieracium hoppea-

num, Phyteuma hemisphaericum, and Scorzoneroides

helvetica, Nardus stricta and Geum montanum are

typical species found in the Sieversio montanae-

Nardetum strictae (Grabherr and Mucina 1993). This

is why we identify this plant community as Sieversio

montanae-Nardetum strictae typicum (Peppler-Lis-

bach and Petersen 2001).

Subassociation vaccinietosum

The second group, including most of the vegetation

releves, contains nearly all four land-use types.

However, compared to the others, this group has

the highest number of unfertilized meadows and

abandoned areas (Table 2, numbers in brackets). In

addition to the abandoned areas, most of the unfer-

tilized meadows are mown every second to third year,

which explains the high number of dwarf-shrubs and

lichens present. A highly significant separation of the

means of the EIVs was found for the R and N values,

which are lower than in the third and fourth group,

but higher than in the first one. Similar to the first

group, the number of species is low and significantly

different from the third and fourth groups. Because of

the differential species of the genus Vaccinium, this

group is identified as Sieversio montanae-Nardetum

strictae vaccinietosum (Hartl 1963).

Subassociation trifolietosum pratensis

In the third group, slightly fertilized meadows are

found mainly. Therefore, the land-use intensity differs

quite significantly from the other groups, and the N

value of the EIVs separates this group from the first and

second ones. Despite having a constancy of not more

than 15%, the species Erigeron uniflorus occurs

exclusively in the third group (Appendix 2). This

subassociation is called Sieversio montanae-Nardetum

strictae trifolietosum pratensis (Braun-Blanquet 1949)

Fig. 3 Results of Indicator Species Analysis (ISA) from step 2 to 15 during the clustering process. a Change in P value after 3,000

permutations of the Monte Carlo test averaged across 335 species, b Number of species with P \ 0.05

Plant Ecol

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as Trifolium pratense (mostly Trifolium pratense ssp.

nivale) is used as the eponymous species.

Subassociation seslerietosum albicantis

The fourth group specifies the community related to

calcareous bedrock. Consequently, highly significant

differences from the other groups are given for the

pH values. Of the EIVs, the L value and—together

with the third group—the R and the N values show

significant differences. Finally, all releves belonging

to this group occur only in areas where limestone

forms the bedrock (Fig. 1) and most of the sites

(more than 80%) are abandoned areas or unfertilized

meadows (Table 2, number in brackets). The group is

also characterized by the highest number of indicator

species (Appendix 2). Because of the indicator

species Sesleria albicans, this fourth group is iden-

tified as Sieversio montanae-Nardetum strictae sesle-

rietosum albicantis; an original diagnosis for this

subassociation is given in Appendix 3.

Discussion

The Sieversio montanae-Nardetum strictae is the

most common plant community in alpine pastures of

the Eastern Alps (Grabherr and Mucina 1993), but it

can also be found in meadows and recently aban-

doned areas. After HCA and ISA, four distinct

subassociations of the Sieversio montanae-Nardetum

strictae were identified and this phytosociological

classification was confirmed by DA. After comparing

the Euclidean distance measure with Relative Euclid-

ean and Bray–Curtis measures, we found out that the

classification in the first four groups was the same for

all the releves. We, therefore, took the Euclidean

distance measure since most of the information is

retained in the first four groups. We furthermore

followed Whittaker (1962) and used species which

preferably occur in a specific group of the Sieversio

montanae-Nardetum strictae to classify the subasso-

ciations. Therefore, we took the ISA as a tool to

contrast performance of species across the groups of

releves. This method is well suited to species data for

describing community types (McCune and Grace

2002).

Our analyses led us to the conclusion that this

plant community should be integrated into the order

of the Nardetalia (class of Calluno-Ulicetea) as

already done by Oberdorfer (1978), Peppler (1992),

and Peppler-Lisbach and Petersen (2001) for central

Europe. Our suggested classification is in contrast to

Grabherr and Mucina (1993), who assigned the

Sieversio montanae-Nardetum strictae community

of the Eastern Alps to the Festucetalia spadiceae

(class of Caricetea curvulae). However, we believe

that the reclassification of this plant community is

valid for the following three key reasons:

1. Character species of the class Calluno-Ulicetea

and the order Nardetalia (as given in Mucina et al.

1993) are found more frequently in our vegetation

releves than the character species of the class

Caricetea curvulae and the order of Festucetalia

spadiceae (as given in Grabherr and Mucina 1993).

Table 3 summarizes the character species provided in

the literature to date for the above-mentioned groups

along with occurrence of the 472 vegetation releves

included in our data set for the Eastern Alps. Worthy

of note is the fact that the eponymous species Carex

curvula was found in just 3% and Festuca paniculata

(i.e. Festuca spadicea) in 4%, whereas Calluna

vulgaris occurs in 40% and Nardus stricta in 100%

of the vegetation releves studied.

2. Even though the Nardetalia communities are

dominated by grasses, we found a high number of

dwarf-shrubs in our 472 vegetation releves, such as

Calluna vulgaris (40%), Erica carnea (7%), Junipe-

rus communis ssp. nana (11%), Salix herbacea (3%),

Salix retusa (2%), Thymus pulegioides (12%), Thy-

mus serpyllum (6%), Vaccinium gaultherioides

(34%), Vaccinium myrtillus (49%), and Vaccinium

vitis-idaea (36%). This evidence warrants a classifi-

cation into the class of dwarf-shrub heather (Calluno-

Ulicetea). Moreover, in the 472 vegetation releves

studied, only 121 (25.6%) contain none of the above-

mentioned dwarf-shrubs. This means that in the

majority, i.e. 74.4%, of our releves dwarf-shrubs are

present. Growth conditions for dwarf-shrubs become

less favorable toward higher elevations (Ellenberg

1996), which is confirmed by our study presented by

the subassociation typicum. Finally, also land-use

type can affect dwarf-shrubs growth (Bischof 1981;

Zoller et al. 1984; Tasser and Tappeiner 2002), e.g.

when meadows are fertilized. This effect can be seen

in the subassociation trifolietosum pratensis.

3. As this plant community was subject to anthro-

pogenic and/or anthropo-zoogenic influence, we,

Plant Ecol

123

Ta

ble

3C

har

acte

rsp

ecie

sb

ased

on

Mu

cin

aet

al.(1

99

3)

for

the

clas

sC

allu

no

-Uli

cete

aan

dth

eo

rder

Nar

det

alia

and

bas

edo

nG

rab

her

ran

dM

uci

na

(19

93

)fo

rth

ecl

ass

Car

icet

ea

curv

ula

ean

dth

eo

rder

Fes

tuce

tali

asp

adic

eae

and

the

occ

urr

ence

in%

of

47

2re

lev

esfr

om

the

Eas

tern

Alp

s

Cal

lun

o-U

lice

tea

Occ

urr

ence

(%)

Nar

det

alia

Occ

urr

ence

(%)

Car

icet

eacu

rvu

lae

Occ

urr

ence

(%)

Fes

tuce

tali

asp

adic

aeO

ccu

rren

ce

(%)

An

ten

na

ria

dio

ica

26

.06

Arn

ica

mo

nta

na

63

.35

Ag

rost

isru

pes

tris

3.8

1C

am

pa

nu

lab

arb

ata

56

.78

An

tho

xan

thu

mo

do

ratu

m5

1.9

1C

are

xp

all

esce

ns

6.9

9A

ven

ula

vers

ico

lor

44

.92

Cen

tau

rea

ner

vosa

2.9

7

Ca

llu

na

vulg

ari

s4

0.0

4G

ali

um

pu

mil

um

12

.50

Gen

tia

na

aca

uli

s5

4.4

5C

rep

isco

nyz

ifo

lia

32

.42

Ca

rex

pil

uli

fera

0.6

4G

ali

um

saxa

tile

0.6

4G

enti

an

ap

un

cta

ta1

.69

Eri

ger

on

alp

inu

s1

.91

Da

nth

on

iad

ecu

mb

ens

4.0

3G

enis

tati

nct

ori

a0

.00

Gen

tia

na

pu

rpu

rea

0.0

0F

estu

cap

an

icu

lata

3.6

0

Gen

ista

sag

itta

lis

0.0

0H

iera

ciu

mla

ctu

cell

a0

.85

Ph

yteu

ma

con

fusu

m0

.00

Geu

mm

on

tan

um

48

.52

Hie

raci

um

pil

ose

lla

50

.85

Hyp

eric

um

ma

cula

tum

16

.31

Ph

yteu

ma

hem

isp

ha

eric

um

9.7

5H

ypo

cho

eris

un

iflo

ra2

6.6

9

Lu

zula

cam

pes

tris

20

.34

Hyp

eric

um

per

fora

tum

0.0

0P

ote

nti

lla

au

rea

58

.26

Pa

rad

isea

lili

ast

rum

3.1

8

Lu

zula

mu

ltifl

ora

47

.25

Na

rdu

sst

rict

a1

00

.00

Pu

lsa

till

aa

lpin

ass

p.

alb

a0

.00

Ph

yteu

ma

bet

on

icif

oli

um

20

.13

Lyc

op

od

ium

cla

vatu

m0

.00

Th

esiu

mp

yren

aic

um

2.3

3S

corz

on

ero

ides

hel

veti

ca1

8.8

6P

lan

tag

ose

rpen

tin

a0

.00

Po

lyg

ala

serp

ylli

foli

a0

.00

Tri

foli

um

alp

inu

m1

2.7

1P

uls

ati

lla

alp

ina

ssp

.a

pii

foli

a2

3.9

4

Po

ten

till

aer

ecta

76

.91

Ra

nu

ncu

lus

vill

ars

ii0

.00

Ver

on

ica

fru

ticu

losa

0.2

1

Mea

np

erce

nta

ge

26

.50

20

.30

18

.59

16

.95

Plant Ecol

123

therefore, integrated the community into the group of

mainly anthropogenic vegetation (Mucina et al.

1993) and not into the group of natural forest-free

vegetation (Grabherr and Mucina 1993). Although

many species of the Sieversio montanae-Nardetum

strictae occur in natural forest-free areas, such as

channels of avalanches and troughs (Oberdorfer

1978; Grabherr and Mucina 1993), the current

species composition of the community refers pri-

marily to anthropogenic influences. For example, it

is found growing on recently abandoned areas that

now attract deer for grazing (Stussi 1970; Peppler

1992).

Our study shows that the alliance Nardion strictae

with the association Sieversio montanae-Nardetum

strictae should remain unchanged. This alliance was

also proposed by Oberdorfer (1978), Krahulec

(1983), Peppler (1992), and Peppler-Lisbach and

Petersen (2001). We further suggest that the character

species of the Nardion strictae as given in Grabherr

and Mucina (1993) should be maintained, because

they separate the continental and lower located

alliance Nardo-Agrostion tenuis of the Eastern Alps.

However, we do not extend this classification to the

species Gnaphalium sylvaticum and Veratrum album,

because they are specified as character species in both

alliances by Mucina et al. (1993) and Grabherr and

Mucina (1993).

HCA confirmed by the DA and ISA of our data set

classified the plant community into four distinct

groups. Note that we merged two of the five groups

determined by ISA because of strong similarities with

the group of calciphile Sieversio montanae-Nardetum

strictae and since there are no significant differences

in site factors for these two groups. Finally, the EVIs

for the species in these two subjects were also not

significantly different. Therefore, we feel justified to

simplify the dendrogram produced by the HCA to

only four groups.

Since we found numerous indicator species for

each group (Appendix 2), as well as highly significant

differences in their ecology (Table 2), we further

classified this plant community into the subassocia-

tion level. We did not classify the groups into

independent associations for the following two key

reasons: (1) The differential species as determined by

ISA and the resulting indicator species for each group

also occur in other groups, albeit at differing

percentages of occurrence; (2) there are no species

that occur exclusively in one group, except Erigeron

uniflorus in group three, indicating that the groups are

highly correlated with each other.

The Sieversio montanae-Nardetum strictae plant

community in the Eastern Alps has already been

classified into subassociations (Braun-Blanquet 1949;

Hartl 1963; Bischof 1981; Heiselmayer 1985, see

Appendix 1 for diploma and doctoral theses). How-

ever, many of these classifications included regional

peculiarities of some restricted species, thus giving an

inaccurate reflection of the subassociation composi-

tion of this plant community. Nevertheless, in 1992

Peppler provided a comprehensive study and classi-

fication of this plant community for the German

region that marginally included the Eastern Alps.

Therefore, we have extended this study to include a

classification of this plant community for the Eastern

Alps. Furthermore, the classification into subassoci-

ations is confirmed by the fact that three of the four

groups were classified into the subassociation level

by Braun-Blanquet (1949) for the subassociation

trifolietosum pratensis, by Hartl (1963) for the

subassociation vaccinietosum and by Peppler-Lis-

bach and Petersen (2001) for the subassociation

typicum. Our fourth group revealed the highest

number of differential and indicator species (Appen-

dix 2), so that a classification of this group at

subassociation level is justified.

The first group represents the typical subassocia-

tion of the Sieversio montanae-Nardetum strictae.

The typical species composition detailed in Grabherr

and Mucina (1993) corresponds to most of our

differential species, such as Carex pallescens, Geum

montanum, Hieracium hoppeanum, Nardus stricta,

Phyteuma hemisphaericum, and Scorzoneroides hel-

vetica. This first group is mainly found in pastures,

which is the most common land-use type for the

Sieversio montanae-Nardetum strictae (Oberdorfer

1978; Peppler-Lisbach and Petersen 2001). The

significant difference in elevation (Table 2) is

explained by the fact that most of the lightly used

pastures are located near and above the timber line,

where grazing animals are left more or less unat-

tended (Tasser et al. 2003). This group also produced

the highest L value of the EVIs, which confirms this

subassociation to be found above the tree line.

Since unfertilized meadows are increasingly aban-

doned (Macdonald et al. 2000; Tasser et al. 2001,

2007), we focused on this land-use type in regard to

Plant Ecol

123

the classification of the Sieversio montanae-Narde-

tum strictae plant community. Moreover, also the

high number of dwarf-shrubs present in the second

group is explained by land-use type: 50.6% of the

releves which belong to meadows mown every

second to third year or to abandoned areas occur in

this group—conditions that are ideal for dwarf-shrubs

growth. Finally, we propose that species such as

Calluna vulgaris, Vaccinium gaultherioides, Vacci-

nium myrtillus, and Vaccinium vitis-idaea should be

used for defining this group in future, as done by

Hartl (1963) too.

Easily accessible meadows of the Sieversio mont-

anae-Nardetum strictae are mostly fertilized with

manure, mown once a year and grazed in autumn

after the return of livestock from high alpine summer

pastures (Knapp and Knapp 1952; Mucina et al.

1993). Such intensification of land use includes

massive extension of forest roads (Krausmann et al.

2003; Mottet et al. 2006) that make agricultural areas

accessible and subsequently lead to an increase of

vehicles and manure in the alpine and subalpine belt

(Pavlu et al. 2005; Stocklin et al. 2007; Liira et al.

2008). This intensification of land use results in a

subassociation of the community having a high

biodiversity (Table 2), because species that grow

well in either meadows or pastures or both occur

more frequently. In accordance with Braun-Blanquet

(1949) and Peppler-Lisbach and Petersen (2001), we

maintain the name of the subassociation, named after

the most common species Trifolium pratense. A

similar type was described by Heiselmayer (1982) as

Trifolio pratensis-Nardetum and by Dietl (1995) as

Hypochoero-Nardetum.

Currently, there is limited information on the

Sieversio montanae-Nardetum strictae plant commu-

nity that establishes above calcareous bedrock

(Grabherr and Mucina 1993). We found the highest

percentage of this plant community type in aban-

doned areas, i.e. in areas with the lowest land-use

intensity (Table 2). The ISA revealed 56 differential

species and 11 indicator species with more than 50%

relative abundance and constancy (Appendix 2),

which is by far the highest among all the groups.

The differential species Sesleria albicans, Phyteuma

orbiculare, and Trifolium badium are calciphile

(Fischer et al. 2005). Additionally, all vegetation

releves belonging to this group were found above

calcareous bedrock (Fig. 1) having a pH value

significantly higher than for the other three groups.

Therefore, a classification into a separate association

would be legitimate. However, the HCA shows that

this group is split from the other groups at last

(Fig. 2). Moreover, there are no species that occur

exclusively in this group. Therefore, the group must

be considered as an equivalent subgroup and not as

an independent association. Finally, Peppler-Lisbach

and Petersen (2001) describe a calciphile Soldanella

alpina variant of the subassociation—trifolietosum

pratensis, which corresponds to our subassociation—

seslerietosum albicantis of the Sieversio montanae-

Nardetum strictae. Appendix 3 compares the con-

stancy of our original diagnosis of the subassociation

seslerietosum albicantis with the constancy table of

Peppler-Lisbach and Petersen (2001). For acceptance

of the subassociation, we additionally present 10

releves from five different sites (two releves for each

site), according to Article 7, Recommendation 7A of

the International Code of Phytosociological Nomen-

clature (i.e. ICPN, Weber et al. 2000) and added the

constancy values extracted from all 78 releves

belonging to this subassociation.

The Sieversio montanae-Nardetum strictae is most

species-rich in the subassociation trifolietosum prat-

ensis with the land-use type mowing, slightly fertil-

izing and grazing in autumn. This traditional hay

meadow management preserves sites with high

biodiversity (Garcia 1992; Myklestad and Sætersdal

2004; Maurer et al. 2006). Therefore, agri-environ-

mental measures, which are among the most impor-

tant instruments for the promotion of environmentally

adapted agricultural land use (Matzdorf et al. 2008),

should focus on traditional hay management.

Acknowledgments We thank Prof. Dr. Robert Crawford for

his useful language revision and Dr. Ruth Willmott (BioScript

International) for her editorial support. We thank the provincial

government of Tyrol for enabling analysis of the soil samples.

This study was funded by the European Union within the

scope of the Interreg IIIA-Italy/Austria-Project ‘‘DNA-

Characterization for certification and valorisation of mountain

hay’’, supported by the provincial governments of Tyrol and

South Tyrol.

Appendix

See Tables 4, 5, 6

Plant Ecol

123

Table 4 Sources for information on the vegetation releves from literature

Author Title Type of

publication

Year of

publication

Location No. of

releves

Brunner, B. Die Vegetation von Bergmahdern im

Landschaftsschutzgebiet Noßlachjoch-

Obernberg-Tribulaun

Diploma thesis,

University of

Innsbruck (AT)

1999 Obernberg,

North Tyrol

83

Dalla Torre, M. Die Vegetation der subalpinen und alpinen

Stufe in der Puez-Geisler Gruppe

Dissertation,

University of

Innsbruck (AT)

1982 St. Christina,

Groden, South

Tyrol

4

Dierschke, H. Grunland-Gesellschaften im oberen

Paznauner Tal

Phytocoenologia6: 287-303

1979 Galtur, Paznaun,

North Tyrol

13

Dirrhammer, H. Die Vegetation im oberen Lechtal Diploma thesis,

University of

Innsbruck (AT)

2008 Elbigenalp,

Lechtal, North

Tyrol

17

Duelli, M. Die Vegetation des Gaißbergtales. Ein

Versuch, das Datenmaterial mit Hilfe der

EDV-Anlage zu bearbeiten

Dissertation,

University of

Innsbruck (AT)

1977 Gaißbergtal,

Obergurgl,

North Tyrol

24

Egger, G. Die Burstlingsrasen vom Nationalpark Hohe

Tauern in Tirol

Personal unpubl.

releves

2006 Hohe Tauern,

East Tyrol

15

Ender, M. Vegetation von gemahten Bergwiesen und

deren Sukzession nach Auflassung der

Mahd

Diploma thesis,

University of

Innsbruck (AT)

1997 Tannberg,

Vorarlberg

35

Flecker, K. Die Vegetation von Schipisten und

angrenzenden Bergmahdern im Raum

Hochtannberg

Diploma thesis,

University of

Innsbruck (AT)

1996 Lech, Vorarlberg 10

Gufler, R. Analyse der Vegetations- und

Erosionsverteilung in Abhangigkeit von

Bewirtschaftungsanderungen am Beispiel

Kaserstattalm

Diploma thesis,

University of

Innsbruck (AT)

1999 Kaserstatt,

Stubaital,

North Tyrol

5

Keim, K. Die Vegetationsverhaltnisse des

Pflerschertales

Dissertation,

University of

Innsbruck (AT)

1967 Pflersch, South

Tyrol

26

Lechner, C. Die Vegetation im Bereich des

Dreilanderecks bei Nauders am

Reschenpass

Diploma thesis,

University of

Innsbruck,

(AT)

1995 Nauders, North

Tyrol

19

Lechner, G. Die Vegetaion der inneren Pfunderer Taler Dissertation,

University of

Innsbruck (AT)

1969 Pfunders, South

Tyrol

16

Mayer, R. Die Vegetation der Bergmahder im Valsertal

und ihre Dynamik

Diploma thesis,

University of

Innsbruck (AT)

2002 Vals, North

Tyrol

11

Mulser, J. Analyse der Vegetationsverteilung in

Abhangigkeit der

Bewirtschaftungsanderung auf den Waltner

Mahdern

Diploma thesis

University of

Innsbruck (AT)

1998 Walten,

Passeiertal,

South Tyrol

12

Nieder brunner,

F.

Vegetation der Sextener Dolomiten

(subalpine und alpine Stufe)

Dissertation,

University of

Innsbruck (AT)

1975 Sexten, South

Tyrol

11

Oberhammer M. Die Vegetation der alpinen Stufe in den

ostlichen Pragser Dolomiten

Dissertation,

University of

Innsbruck

1979 Prags,

Dolomites,

South Tyrol

1

Plant Ecol

123

Table 5 Results of the Indicator Species Analysis (ISA) for 142 significant (P \ 0.05) species

Species name Rel. abundance (%) Constancy (%) Indicator values Monte Carlo test of significance of

observed maximum indicator values

for species with 3,000 permutations

No. of releves No. of releves No. of releves MaxGr Value Mean s.d. P

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Sieversio montanae-Nardetum strictae typicum

Nardus stricta 62 12 18 8 100 100 100 100 62 12 18 8 1 61.6 28.1 1.67 0.0003

Geum montanum 41 21 28 10 59 47 73 29 24 10 20 3 1 24.2 16.5 2.57 0.0123

Scorzoneriodes helvetica 51 21 4 25 30 18 8 13 15 4 0 3 1 15.2 7.7 1.97 0.0067

Pedicularis tuberosa 51 33 16 1 25 16 20 1 13 5 3 0 1 12.7 6.8 1.85 0.0160

Phyteuma hemisphaericum 58 30 11 0 22 8 8 0 13 2 1 0 1 12.9 5.1 1.78 0.0047

Hieracium hoppeanum 53 8 21 18 23 6 20 4 12 0 4 1 1 12.2 5.2 1.73 0.0077

Festuca halleri agg. 58 22 12 8 17 7 5 5 10 2 1 0 1 9.8 4.9 1.86 0.0237

Polygala comosa 73 18 0 9 14 5 0 3 10 1 0 0 1 10.3 3.6 1.45 0.0037

Festuca ovina agg. 70 30 0 0 13 5 0 0 9 2 0 0 1 9.1 3.5 1.51 0.0087

Carex pallescens 51 17 25 7 13 6 8 3 7 1 2 0 1 6.7 3.8 1.44 0.0500

Centaurea nervosa 67 15 19 0 10 0 8 0 7 0 1 0 1 6.7 2.5 1.31 0.0153

Danthonia decumbens 72 25 4 0 9 3 5 0 6 1 0 0 1 6.4 2.8 1.34 0.0273

Gentiana lutea 92 0 0 8 6 0 0 6 6 0 0 1 1 5.5 2.5 1.32 0.0363

Gnaphalium sylvaticum 98 2 0 0 6 0 0 0 6 0 0 0 1 5.9 1.6 0.93 0.0040

Carex capillaris 90 3 0 7 6 1 0 3 5 0 0 0 1 5.4 2.1 1.15 0.0230

Ranunculus bulbosus 89 11 0 0 5 2 0 0 4 0 0 0 1 4.4 1.9 1.06 0.0290

Table 4 continued

Author Title Type of

publication

Year of

publication

Location No. of

releves

Putzer, J. Pflanzengesellschaften im Raum von Brixen

mit besonderer Berucksichtigung der

Trockenvegetation

Dissertation,

University of

Innsbruck (AT)

1967 Brixen, South

Tyrol

5

Smettan, H. Die Pflanzengesellschaften des

Kaisergebirges/Tirol.

Dissertation,

University of

Innsbruck (AT)

1981 Kaisergebirge,

North Tyrol

2

Steinmair, V. Die Vegetation von unterschiedlich genutzten

Almflachen auf der Platzwiese

Diploma thesis,

University of

Innsbruck (AT)

1999 Platzwiese,

Prags, South

Tyrol

20

Tasser, E. Vegetationsaufnahmen von Burstlingsrasen

des Monte Bondone

Personal unpubl.

releves

2005 Mt. Bondone,

Trentino, Italy

22

Thomaser, J. Die Vegetation des Peitlerkofels in Sudtirol. Veroff. Museum

Ferdinandeum

47: 67–119

1967 Gardertal 2

Wallossek, C. Vegetationskundlich-okologische

Untersuchungen in der alpinen Stufe am

SW-Rand der Dolomiten

Dissertationes

Botanicae 154

(IT)

1990 Lafatscher Joch,

Latemar, South

Tyrol

4

Plant Ecol

123

Table 5 continued

Species name Rel. abundance (%) Constancy (%) Indicator values Monte Carlo test of significance of

observed maximum indicator values

for species with 3,000 permutations

No. of releves No. of releves No. of releves MaxGr Value Mean s.d. P

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Sieversio montanae-Nardetum strictae vaccinietosum

Calluna vulgaris 30 58 8 4 41 50 28 12 12 29 2 0 2 29 13.9 2.33 0.0003

Vaccinium gaultherioides 20 53 18 9 32 43 25 13 6 23 5 1 2 22.6 12.4 2.32 0.0017

Vaccinium myrtillus 24 39 15 23 39 56 35 47 9 22 5 11 2 21.5 16.3 2.31 0.0340

Vaccinium vitis-idaea 32 50 8 16 35 42 20 26 11 18 2 4 2 18.3 12.5 2.16 0.0203

Anthoxanthum alpinum 42 50 2 6 38 35 3 5 16 17 0 0 2 17.2 10.3 2.09 0.0097

Antennaria dioica 35 47 16 2 35 30 25 3 12 14 4 0 2 13.9 9.9 2.05 0.0473

Cetraria islandica 23 64 0 13 14 16 0 4 3 11 0 1 2 10.5 5.8 1.77 0.0250

Cladonia rangferina 12 82 0 6 3 11 0 3 0 9 0 0 2 8.7 3.8 1.46 0.0130

Erica carnea 7 78 0 15 4 10 0 1 0 8 0 0 2 7.9 4.1 1.62 0.0313

Picea abies 2 98 0 0 1 9 0 0 0 8 0 0 2 8.4 3.4 1.53 0.0147

Silene nutans 1 99 0 0 1 8 0 0 0 8 0 0 2 7.8 3.2 1.44 0.0147

Cladonia arbuscula 12 88 0 0 2 6 0 0 0 6 0 0 2 5.5 2.7 1.22 0.0343

Hieracium lachenalii 28 72 0 0 2 9 0 0 1 6 0 0 2 6.5 3.2 1.38 0.0327

Sieversio montanae-Nardetum strictae trifolietosum pratensis

Festuca rubra agg. 8 7 67 18 74 76 100 91 6 5 67 16 3 67.1 24.1 2.30 0.0003

Trifolium pratense 10 11 68 11 53 55 95 55 5 6 64 6 3 64.4 18.8 2.59 0.0003

Plantago alpina 6 10 76 9 9 5 50 3 1 0 38 0 3 38 4.7 1.60 0.0003

Hypochaeris uniflora 5 7 84 4 27 25 45 22 1 2 38 1 3 37.7 12.6 3.30 0.0003

Potentilla aurea 17 15 44 24 71 48 85 61 12 7 37 15 3 37.5 18.8 2.55 0.0003

Phleum commutatum 5 13 75 7 11 9 48 5 1 1 36 0 3 35.5 5.8 1.80 0.0003

Leucantthemum vulgare 15 12 45 28 39 45 80 68 6 5 36 19 3 36.3 16.4 2.34 0.0003

Mutellina adonidifolia 9 7 83 0 29 15 43 3 3 1 35 0 3 35.5 8.5 2.37 0.0003

Luzula campestris 15 13 55 17 20 15 63 18 3 2 34 3 3 34.5 8.2 2.03 0.0003

Anthoxanthum odoratum 10 14 40 35 37 46 85 74 4 7 34 26 3 34.2 16.8 2.32 0.0003

Rhinanthus alectorolophus 1 9 88 1 8 13 38 4 0 1 33 0 3 32.9 6.4 2.00 0.0003

Traunsteinera globosa 2 4 84 9 4 5 38 5 0 0 32 0 3 31.5 4.2 1.60 0.0003

Hieracium pilosella 15 19 48 18 41 52 65 53 6 10 31 9 3 31.3 17 2.59 0.0013

Campanula barbata 19 25 37 18 48 58 80 52 9 15 30 10 3 29.6 18.4 2.50 0.0023

Arnica montana 22 28 38 12 62 65 78 53 14 18 30 7 3 29.8 20.2 2.52 0.0053

Primula elatior 14 3 77 6 8 2 38 4 1 0 29 0 3 28.9 3.8 1.41 0.0003

Ranunculus montanus 13 12 58 17 26 21 50 21 3 3 29 3 3 29.1 9.3 2.05 0.0003

Campanula scheuchzeri 14 20 31 35 59 68 93 78 8 13 29 27 3 29.1 21 2.06 0.0027

Lotus corniculatus s.l. 16 20 33 31 58 67 90 78 9 13 29 25 3 29.4 21 2.17 0.0043

Crepis pontana 0 4 93 3 0 3 30 3 0 0 28 0 3 27.9 3.2 1.46 0.0003

Avenula versicolor 20 23 38 19 39 44 73 42 8 10 28 8 3 27.7 15 2.28 0.0010

Gymnadenia conopsea 19 25 48 8 31 38 58 16 6 10 28 1 3 27.7 12.7 2.58 0.0007

Gentiana acaulis 28 22 37 13 61 52 75 44 17 11 28 6 3 28.1 17.4 2.16 0.0020

Luzula sylvatica 5 24 67 5 2 18 38 5 0 4 25 0 3 25 6.3 1.83 0.0003

Plant Ecol

123

Table 5 continued

Species name Rel. abundance (%) Constancy (%) Indicator values Monte Carlo test of significance of

observed maximum indicator values for

species with 3,000 permutations

No. of releves No. of releves No. of releves MaxGr Value Mean s.d. P

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Anemone narcissiflora 0 5 79 16 0 4 30 16 0 0 24 2 3 23.7 3.9 1.50 0.0003

Heracleum sphondylium 0 10 87 3 0 2 28 5 0 0 24 0 3 23.9 3.6 1.57 0.0003

Myosotis alpestris 4 8 60 28 6 11 40 36 0 1 24 10 3 23.9 7.1 1.99 0.0007

Crepis conyzifolia 11 13 55 21 22 26 40 62 2 3 22 13 3 21.9 12.3 2.58 0.0070

Polygala alpestris 16 12 43 28 29 18 48 43 5 2 21 12 3 20.6 9.8 1.97 0.0010

Crepis aurea 17 12 44 27 21 15 38 45 4 2 17 12 3 16.6 9.3 2.33 0.0130

Pseudorchis albida 13 17 70 0 9 9 25 0 1 2 17 0 3 17.5 4.8 1.75 0.0003

Thesium alpinum 11 24 36 29 23 34 48 32 3 8 17 10 3 16.9 11.9 2.33 0.0397

Euphrasia officinalis s.l. 15 26 37 22 23 35 48 18 4 9 17 4 3 17.4 11 2.07 0.0117

Crocus albiflorus 23 17 41 19 33 25 40 39 7 4 16 7 3 16.5 10.9 2.05 0.0200

Erigeron uniflorus 0 0 100 0 0 0 15 0 0 0 15 0 3 15 1.5 0.89 0.0003

Euphrasia minima 25 28 42 4 16 12 35 1 4 3 15 0 3 14.7 6 1.74 0.0017

Phyteuma ovatum 1 6 93 0 1 2 15 0 0 0 14 0 3 14 2.1 1.15 0.0003

Ajuga reptans 1 3 84 13 1 0 15 4 0 0 13 0 3 12.5 2.1 1.10 0.0003

Gentianella campestris 17 21 56 6 13 11 23 4 2 2 13 0 3 12.6 5.5 1.86 0.0080

Pimpinella major 11 18 36 35 10 11 33 23 1 2 12 8 3 11.7 6.5 1.84 0.0190

Botrychium lunaria 20 16 36 28 14 16 33 25 3 3 12 7 3 11.6 7.7 1.95 0.0467

Trisetum flavescens 3 5 71 21 2 3 15 8 0 0 11 2 3 10.7 3.3 1.53 0.0033

Carduus defloratus 5 11 58 27 3 5 18 8 0 1 10 2 3 10.1 3.9 1.53 0.0053

Crepis pyrenaica 2 9 50 38 1 4 18 16 0 0 9 6 3 8.8 3.5 1.40 0.0090

Poa trivialis 0 4 69 26 2 1 13 4 0 0 9 1 3 8.7 2.1 1.09 0.0007

Phleum hirsutum 7 17 42 33 4 7 23 14 0 1 9 5 3 9.5 4.5 1.51 0.0140

Silene latifolia 5 19 46 30 7 6 20 8 0 1 9 2 3 9.2 4.2 1.61 0.0157

Aster alpinus 0 18 82 0 0 3 10 0 0 1 8 0 3 8.2 2.2 1.19 0.0027

Gnaphalium norvegicum 0 13 87 0 0 1 8 0 0 0 7 0 3 6.5 1.7 0.98 0.0043

Scabiosa canescens 0 1 99 0 0 0 8 0 0 0 7 0 3 7.4 1.4 0.84 0.0017

Poa annua 35 7 59 0 3 2 10 0 1 0 6 0 3 5.9 2.5 1.28 0.0243

Cyanus montanus 0 1 99 0 0 0 5 0 0 0 5 0 3 4.9 1.2 0.77 0.0070

Gentianella anisodonta 30 0 70 0 4 0 8 0 1 0 5 0 3 5.2 1.8 0.98 0.0113

Thesium pyrenaicum 10 24 67 0 2 2 8 0 0 1 5 0 3 5 2.1 1.16 0.0313

Dactylorhiza majalis 10 4 59 26 1 0 8 3 0 0 4 1 3 4.5 1.6 0.91 0.0147

Epipactis palustris 26 0 71 4 1 0 5 1 0 0 4 0 3 3.5 1.4 0.79 0.0317

Hieracium alpinum 16 6 78 0 1 0 5 0 0 0 4 0 3 3.9 1.3 0.80 0.0250

Sieversio montanae-Nardetum strictae seslerietosum albicantis

Alchemilla vulgaris agg. 16 8 17 60 42 38 83 87 7 3 14 52 4 52 16.9 2.44 0.0003

Agrostis capillaris 9 10 15 65 37 38 45 71 3 4 7 47 4 46.8 15 2.41 0.0003

Galium anisophyllon 17 13 18 52 42 28 40 91 7 4 7 47 4 46.9 14 2.14 0.0003

Trollius europaeus 7 13 24 56 23 39 52 83 2 5 13 46 4 46.2 14.9 2.31 0.0003

Geranium sylvaticum 3 1 18 78 12 5 33 57 0 0 6 45 4 44.7 7.8 2.08 0.0003

Plant Ecol

123

Table 5 continued

Species name Rel. abundance (%) Constancy (%) Indicator values Monte Carlo test of significance of

observed maximum indicator values

for species with 3,000 permutations

No. of releves No. of releves No. of releves MaxGr Value Mean s.d. P

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Rhinanthus glacialis 10 7 17 66 26 31 55 61 3 2 10 40 4 40 13.4 2.54 0.0003

Briza media 17 21 17 44 48 56 50 88 8 12 9 39 4 38.8 18.4 2.21 0.0003

Phyteuma orbiculare 11 7 27 55 23 18 48 71 3 1 13 39 4 39.1 10.9 2.08 0.0003

Rumex alpestris 1 4 20 75 4 4 25 45 0 0 5 34 4 34 6 1.90 0.0003

Leontodon hispidus 18 13 25 44 49 48 52 77 9 6 13 34 4 34 17.4 2.32 0.0003

Sesleria albicans 7 10 18 64 8 8 15 51 1 1 3 33 4 32.6 7.3 2.26 0.0003

Trifolium badium 7 9 28 56 24 20 55 56 2 2 15 31 4 31.2 10.9 2.15 0.0003

Potentilla erecta 24 21 23 32 75 72 90 88 18 15 21 29 4 28.6 22.7 1.96 0.0090

Silene vulgaris 2 12 33 53 12 23 45 56 0 3 15 29 4 29.5 10.3 2.09 0.0003

Plantago lanceolata 13 8 20 60 6 13 20 47 1 1 4 28 4 27.8 7.5 1.97 0.0003

Ranunculus nemorosus 13 7 34 46 19 19 25 61 2 1 9 28 4 27.9 10.3 2.34 0.0003

Achillea millefolium 10 27 19 44 25 35 38 61 2 9 7 27 4 26.8 12.9 2.25 0.0003

Phleum rhaeticum 10 8 33 49 13 11 15 48 1 1 5 24 4 23.8 7.3 1.85 0.0003

Deschampsia cespitosa 20 11 21 48 22 15 25 47 4 2 5 23 4 22.6 8.9 2.01 0.0010

Avenula pubescens 6 5 31 59 15 6 20 38 1 0 6 22 4 22.1 6.5 1.87 0.0003

Luzula luzuloides 9 37 5 49 13 23 13 45 1 9 1 22 4 22.2 9.5 2.26 0.0017

Anthyllis vulneraria ssp.alpicola

8 9 38 45 17 18 45 45 1 2 17 20 4 20.4 9.5 2.08 0.0010

Bartsia alpina 21 13 20 46 23 18 28 44 5 2 6 20 4 20.5 9 1.87 0.0007

Laserpitium latifolium 2 7 3 88 4 11 10 22 0 1 0 20 4 19.5 5.8 2.02 0.0003

Chaerophyllum villarsii 5 7 29 58 13 17 52 34 1 1 15 20 4 19.7 9 2.22 0.0017

Biscutella laevigata 16 17 13 53 13 12 15 35 2 2 2 19 4 18.7 7 1.93 0.0007

Centaurea pseudophrygia 1 6 33 59 5 15 28 32 0 1 9 19 4 19.3 7.5 2.11 0.0010

Trifolium medium 0 6 4 90 0 10 3 21 0 1 0 19 4 18.7 5.1 1.84 0.0007

Cerastium fontanum agg. 15 13 29 43 23 20 38 45 4 3 11 19 4 19.4 9.7 2.01 0.0030

Knautia maxima 1 7 29 63 6 8 20 29 0 1 6 18 4 17.9 5.9 1.84 0.0003

Plantago media 17 20 3 61 14 9 10 30 2 2 0 18 4 18.1 6.9 2.22 0.0027

Veronica chamaedrys 1 7 48 44 2 9 15 40 0 1 7 18 4 17.6 6.1 1.86 0.0010

Astrantia major 0 12 2 86 0 3 5 19 0 0 0 17 4 16.8 3.2 1.36 0.0003

Knautia longifolia 4 1 6 90 5 3 8 19 0 0 0 17 4 17.4 4 1.65 0.0003

Poa alpina 22 14 25 38 22 16 33 44 5 2 8 17 4 16.9 9 2.05 0.0067

Soldanella alpina 20 10 37 32 32 24 45 55 7 3 17 17 4 17.3 11.5 2.15 0.0203

Thymus pulegioides 22 18 4 57 4 11 5 30 1 2 0 17 4 16.9 5.6 1.68 0.0003

Trifolium montanum 22 23 13 42 25 27 30 42 6 6 4 17 4 17.4 10.9 2.09 0.0127

Dactylis glomerata 1 23 10 67 2 13 15 23 0 3 1 16 4 15.7 5.8 1.72 0.0010

Scabiosa lucida 15 11 6 69 14 14 28 22 2 2 2 15 4 15.2 7.7 2.29 0.0133

Carex montana 15 19 0 66 16 11 0 21 2 2 0 14 4 13.7 5.6 1.69 0.0027

Hippocrepis comosa 20 12 9 59 10 7 5 25 2 1 0 14 4 14.5 4.9 1.61 0.0017

Astragalus glyciphyllos 0 2 0 98 0 1 0 13 0 0 0 13 4 12.8 2.3 1.20 0.0003

Plant Ecol

123

Table 6 Constancy table of Peppler-Lisbach and Petersen

(2001) and constancy values of our original diagnosis and 10

releves after the method of Braun-Blanquet (1964) of the

subassociation seslerietosum albicantis with the factors alti-

tude, pH, slope, and land-use intensity (LUi). The fifth releve

(OV in bold) constitutes the nomenclatural type

Source

Subassociation

Peppler-Lisbach and Petersen (2001) Own

sesl.

10 Releves of the subassociation seslerietosum albicantis

typicum trifolietosum pratensis

Site LV LV RP RP OV OV SD SD HT HT

Number of releves 50 32 17 67 16 54 11 78 1 1 1 1 1 1 1 1 1 1

Mean altitude (*10; m a.s.l.) 161 194 158 153 192 154 196 185 178 187 164 162 176 180 198 198 192 175

Mean number of species 28 30 26 40 37 48 43 45 23 35 46 52 49 40 36 33 40 28

Mean pH (0–10 cm) 4.8 6.6 6.4 4.5 5.8 4.7 5.1 5 5.6 6.1 4.7

Mean slope (�) 15 5 10 24 18 12 10 15 20 10 10

LUi 0.5 0.5 0.1 0.1 0.5 3 0.3 0.3 1 1

Sieversio-Nardetum

Plantago alpina III II III III IV IV IV ?

Campanula barbata I IV I III V III V III 1

Gebtiana punctata III III III II III I II

Gentiana acaulis I III II IV II V III 1

Phleum alpinumagg. II III I II II II III III 1 1 2m 1 ?

Carex sempervirens I IV ? ? III III IV III 4 4 2a 4 3

Euphrasia minima I III II III II IV r

Persicaria vivipara r II ? II III II IV II 1 1 ? ?

Table 5 continued

Species name Rel. abundance (%) Constancy (%) Indicator values Monte Carlo test of significance of

observed maximum indicator values

for species with 3,000 permutations

No. of releves No. of releves No. of releves MaxGr Value Mean s.d. P

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Gr1

100

Gr2

255

Gr3

40

Gr4

77

Hypericum maculatum 12 14 32 42 14 11 30 31 2 1 10 13 4 13.2 7 1.91 0.0107

Avenula pratensis 8 4 0 88 3 3 0 13 0 0 0 11 4 11.5 3.1 1.43 0.0010

Carex ornithopoda 30 14 4 52 7 5 8 22 2 1 0 11 4 11.5 4.6 1.65 0.0040

Helianthemum ovatum 18 25 0 57 7 9 0 19 1 2 0 11 4 11.1 4.8 1.57 0.0040

Cirsium heterophyllum 0 6 28 66 0 4 10 14 0 0 3 9 4 9.4 3.6 1.51 0.0053

Linum catharticum 16 7 29 48 5 2 13 14 1 0 4 7 4 6.8 3.3 1.37 0.0233

Rhytidiadelphus squarrosus 7 0 37 56 2 0 5 12 0 0 2 7 4 6.5 2.5 1.27 0.0133

Taraxacum officinale 11 2 7 80 5 1 3 9 1 0 0 7 4 7.3 2.6 1.28 0.0090

Carex ferruginea 24 6 0 69 3 1 0 9 1 0 0 6 4 6.3 2.2 1.16 0.0097

Buphthalmum salicifolium 0 22 0 78 0 2 0 6 0 0 0 5 4 5.1 2 1.13 0.0240

Carex panicea 5 0 12 83 1 0 3 6 0 0 0 5 4 5.4 1.7 1.06 0.0127

Pedicularis rostratocapitata 0 22 0 78 0 1 0 5 0 0 0 4 4 4 1.7 0.98 0.0337

Lathyrus laevigatus 0 41 0 59 0 0 0 5 0 0 0 3 4 3 1.5 0.91 0.0487

Relative abundance (%) = Percent of average abundance of a given species in a given group of vegetation releves over the average

abundance of that species in all releves; Constancy (%) = Percent of releves in a given group where a given species is present;

Indicator values = Percent of perfect indication based on combining the values for relative abundance and constancy

Plant Ecol

123

Table 6 continued

Source

Subassociation

Peppler-Lisbach and Petersen (2001) Own

sesl.

10 Releves of the subassociation seslerietosum albicantis

typicum trifolietosum pratensis

Crocus alpiflorus ? II ? II ? III 1 1 1

Plantago atrata r ? II III

Galium anisophyllon ? r I I II III V 1 1 1 1 2m ? ? 1

Phyteume betonicifolium ? II ? II II I II r

Hieracium aurathiacum ? II I II ? ? ?

Gentiana purpurea I ? II ?

Crepisconyzifolia r II r II ? IV ?

Gentiana punctata r r ? ? I

Sieversio-Nardetum trifolietosum pratensis

Trifolium pratense r ? IV II IV III III 2m ?

Poa alpina I ? ? III II IV III II 1 1

Lotus corniculatus agg. r III II IV III IV 1 1 1 2m 2a ?

Crepis aurea ? ? ? III II IV III II 1 ?

Leucanthemum vulgare agg. r r III III III IV III ? 1 2m ? 1

Carlina acaulis s.l. ? ? III I III II V ? 1 1 1 ? 1 1 1

Prunella vulgaris III ? III I

Hieracium angustifolium et shaeroceph. r I II I II ? 1

Trifolium repens r II ? II ?

Trichophorum cespitosum-variant

Trichophorum cespitosum ? IV ?

Molinia caerulea r I IV r ? ? ? 2m

Carex nigra ? r III ? I r

Soldanella alpina-variant

Selagnella selaginoides r ? I I IV IV I 2m

Soldanella alpina r r I II IV IV III 1 2m 1 1 1

Bartsia alpina r III IV II 1 2m 1

Ranunculus montanus ? r ? I ? III III I 2m 1

Aster bellidiastrum r r II II I 1 1

Tofieldia calyculata ? r II r

Sieversio-Nardetum seslerietosum albicantis

Sesleria albicans IV 3 2a 2m 1 2m 1 1 1

Trifolioum badium III 2m ? ?

Silene vulgaris III ? 1 1 1 ? ?

Anthyllis vulneraria III ? ? 2m ? ?

Luzula luzuloides III 1 1 ?

Trifolium montanum II 1 ?

Carex pallescens-form

Carex pallescens V I V V I V I ?

Carex pilulifera II IV II II

Veratrum album II II III I III I ? r

Hieracium pilosella II I I III III I III 1 1 1 1

Helictotrichon versicolor-form

Avenochloa versicolor r V r V V III 1

Agrostis rupestris I III ? III I IV

Hypochaeris uniflora r III IV II II ? 1

Nardetalia

Nardus stricta V V V V V V IV V r r 2a 2m 2b 1 2b 2a ? 1

Plant Ecol

123

Table 6 continued

Source

Subassociation

Peppler-Lisbach and Petersen (2001) Own

sesl.

10 Releves of the subassociation seslerietosum albicantis

typicum trifolietosum pratensis

Arnica montana III IV IV IV V III V III ? 1 1 ?

Luzula multiflora III III III IV II III IV III 1 ?

Hieracium lactucella I ? I III I II I

Veronica officinalis I I II I ?

Antennaria dioica ? I I II II II ?

Danthonia decumbens r I ? II

Luzula campestris I II I I I I II 2m 1 ?

Gentianella campestris r ? ? II ?

Calluno-Ulicetea

Vaccinium myrtillus V IV V IV IV IV V III 2m 1 ?

Avenella flexuosa III V III II V II IV III ? ?

Callun a vulgaris II III IV III I II II I

Pleurozium schreberi I III II I ? I ?

Vaccinium vitis-idaea II II II II II I II II ? 1 1 ?

Remaining species

Alchemilla vulgaris s.l. ? r III III IV III IV 1 2a 1 1 1 ? ?

Anthoxanthum odoratum agg. V V V V V V V IV 2m 1 2m 2m 2a 4

Festuca rubra agg. V V V V V V V V 2a 2m 1 2a 2b 2a ? ?

Potentilla erecta V III V V III V III V 1 2m 2m 2m 2m 1 2m 2m ? ?

Agrostis capillaris IV II III V III IV IV IV 2a 2a 1 3 1 ?

Solidago virgaurea s.l. III IV III III IV II V I ?

Deschampsia cespitosa II I II III III IV III II ? 1

Vaccinium gaultherioides I III III I IV II IV I 2m 1

Leontodon hispidus s.l. I I III II IV III IV III 2m ? 20 1 2a

Ranunculus nemorosus r II I II II IV 1 1 3

Phyteuma orbiculare r ? ? I III IV 1 1 1 1 1 ? 1 1 ?

Trollius europaeus r ? I I I III V 1 1 1 ? 2a

Willemetia stipitata I II II ? II ?

Briza media r r I II II I V 1 1 1 1 2m 1 2m ? 2a

Carex panicea r ? r II I

Thymus praecox ssp. polytrichus r r I II II II II 1 2m ?

Hypericum maculatum I ? II ? I ? II 1 ?

Hieracium murorum r ? ? II ? I ? ? ?

Cerastium holosteoides r I I I I II III 1 1

Carexferruginea r r I I II I

Carex flacca ? r ? I II I 1

Rhinanthus glacialis r r II ? IV 2a 2m ? 2a

Geranium sylvaticum r ? I I II III 2a ? ? 1

Achillea millefolium r II ? ? IV 1 1 ? ?

Ranunculus acris II r I ? ? ?

Coeloglossum viride ? ? r II I I

Geum montanum II 2a

Polygala alpestris ? ? II III 1 1 1 1 ?

Knautia maxima r r II II ? ?

Plantago lanceolata III 1 2a ?

Rumex alpestris III 1 ?

Helianthemum nummularium s.l. I 2m 1 ? ?

Plant Ecol

123

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Source

Subassociation

Peppler-Lisbach and Petersen (2001) Own

sesl.

10 Releves of the subassociation seslerietosum albicantis

typicum trifolietosum pratensis

Trifolium medium I 2m 2m 3 3

Carex montana I 2m 2m 1 2a

Plantago media II 1 ? 1 ?

Dactylis glomerata II 1 1 2a ?

Pimpinella major II ? ? 1 ?

Biscutella laevigata II 1 1 1

Chaerophyllum villarsii II 2b 3 ?

Veronica chamaedrys II 1 ? ?

Thymus pulegioides II 1 2a 1

Centaurea pseudophrygia II ? 1 2a

Carex ornitopodoides II 1 1 ?

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Hippocrepis comosa II 2m

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Avenula pratensis I 3 2a

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Anthyllis vulneraria ssp. polyphylla I ? ? ?

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