Picea abies andAbies alba forests of the austrian alps: Numerical classification and ordination

A TWINSPAN classification of a representative set of 3026 relevés of spruce and fir forests from the Eastern Alps (Austria) is presented. Ecological features of relevé clusters and species groups are described by means of Ellenberg indicator values, site factors and stand characteristics. The most important floristic discontinuity in the data set separates acidophilous communities on mostly silicate substrates from basiphilous communities on mostly carbonate substrates. Further divisions reflect a combined gradient of temperature, nutrient regime and shading. This is supported by the correlation of average Ellenberg values of sample plots with DCA axes. A qualitative comparison between TWINSPAN clusters and a syntaxonomic system widely used in the Austrian Alps is drawn. The two ordersPiceetalia excelsae andAthyrio-Piceetalia largely coincide with the clusters of the first level of divisions. Alliances are partly reproduced by TWINSPAN. Clusters on the fifth and fourth level of division mostly correspond to associations. However, a considerable portion of the lower level clusters is of a transitional type. Out of thirteen fir and spruce associations described for the Austrian Alps, five associations are not reproduced by TWINSPAN, i.e.Bazzanio-Piceetum, Veronico-Piceetum, Adenostylo alliariae-Abietetum, Asplenio-Piceetum andCarici-Piceetum. Three associations are split on the second level of division, i.e.Larici-Piceetum, Luzulo nemorosae-Piceetum andCalamagrostio variae-Piceetum.     

Vegetation and environmental factors during primary succession on glacier forelands: Some outlines from the Italian Alps

ABSTRACT

Relationships between plant communities and the physical environment during primary succession on recently deglaciated glacier forelands were studied in 3 areas of the Italian Alps. The aim of the research was to relate traditional phytosociological data with environmental variables. Twenty-eight phytosociological relevés were performed, each associated with twenty-six environmental variables; quantitative parameters of richness and diversity were also calculated. Species/relevés, environmental variables/relevés and species/environmental variables matrices were analyzed by cluster analysis, PCA and Spearman correlation coefficient. Three main stages of succession were identified by floristic composition and confirmed by environmental parameter evaluation. A complex of environmental variables seems to be closely correlated with terrain age and richness/diversity parameters, even though diversity decreases in late successional stages. The phytosociological significance of species is in accordance with their position in the context of succession.     

Phytosociological data give biased estimates of species richness

Abstract. Large phytosociological data sets of three types of grassland and three types of forest vegetation from the Czech Republic were analysed with a focus on plot size used in phytosociological sampling and on the species‐area relationship. The data sets included 12975 relevés, sampled by different authors in different parts of the country between 1922 and 1999. It was shown that in the grassland data sets, the relevés sampled before the 1960s tended to have a larger plot size than the relevés made later on. No temporal variation in plot sizes used was detected in forest relevés. Species‐area curves fitted to the data showed unnatural shapes, with levelling‐off or even decrease in plot sizes higher than average. This distortion is explained by the subjective, preferential method of field sampling used in phytosociology. When making relevés in species‐poor vegetation, researchers probably tend to use larger plots in order to include more species. The reason for this may be that a higher number of species gives a higher probability of including presumed diagnostic species, so that the relevé can be more easily classified in the Braun‐Blanquet classification system. This attitude of phytosociologists has at least two consequences: (1) in phytosociological data bases species‐poor vegetation types are underrepresented or relevés are artificially biased towards higher species richness; (2) the suitability of phytosociological data for species richness estimation is severely limited.     

A phytosociological classification of Swiss mire vegetation

The mapping and monitoring of Swiss mires has so far relied on a classification system based on expert judgement, which was not supported by a quantitative vegetation analysis and which did not include all wetland vegetation types described in the country. Based on a spatially representative sample of 17,608 relevés from 112 Swiss mires, we address the following questions: (1) How abundant are wetland vegetation types (phytosociological alliances) in Swiss mires? (2) How are they distributed across the country––is there a regional pattern? (3) How clearly are they separated from each other? (4) How clear and reliable is their ecological interpretation? Using published wetland vegetation relevés and lists of diagnostic species for phytosociological units (associations and alliances) established by experts, we developed a numerical method for assigning relevés to units through the calculation of similarity indices. We applied this method to our sample of 17,608 relevés and estimated the total area covered by each vegetation type in Switzerland. We found that vegetation types not included in previous mapping were either rare in Switzerland (partly due to mire drainage) or poorly distinguished from other vegetation units. In an ordination, the Swiss mire vegetation formed a triangular gradient system with the Sphagnion medii, the Caricion davallianae and the Phragmition australis as extreme types. Phytosociological alliances were clearly separated in a subset of 2,265 relevés, which had a strong similarity to one particular association, but poorly separated across all relevés, of which many could not be unequivocally assigned to one association. However, ecological gradients were reflected equally well by the vegetation types in either case. Overall, phytosociological alliances distinguished until now proved suitable schemes to describe and interpret vegetation gradients. Nevertheless, we see the urgent need to establish a data base of Swiss wetland relevés for a more reliable definition of some vegetation units.     

Spatial patterns of historical growth changes in Norway spruce across western European mountains and the key effect of climate warming

Key message

Productivity changes in Norway spruce show important regional and local spatial variations, highlighting their context dependence at different spatial scales. These variations suggest the enhancing role of climate warming, and interplay with local water and nutrient limitations.

Abstract

While forest growth changes have been observed in many places worldwide, their spatial variation and environmental origin remain poorly documented. Analysis of these historical changes in contrasted regional contexts, and their mapping over continuous environmental gradients, may help uncover their environmental causes. The approach was tested on Norway spruce (Picea abies) in three western European mountain contexts (Massif Central, Alps and Jura), using National Forest Inventory (NFI) data. We explored the environmental factors influencing stand basal area increment (BAI) in each context. We then estimated and compared mean regional changes in BAI and related these to the regional environmental limitations evidenced. Within each region, we further mapped local BAI trends using a geographically weighted regression (GWR) approach. In each region, local estimates of BAI changes were finally correlated to environmental indicators. We found an increase in BAI in the three regions over 1980–2005, greater in the Massif Central (+71 %) than in the Alps (+19 %) and the Jura Mountains (+21 %). Inter-regional differences in BAI changes suggested the release of a thermal constraint—found more important in the Massif Central—by the strong temperature increase over the period, and a limitation by water availability in the Jura and the Alps Mountains. Spatial patterns of BAI change revealed significant local variations in the Massif Central and the Alps. From the correlation analysis, these were again found consistent with the hypothesis of an enhancing effect of climate warming in these mountain ranges. They were also related to local soil nutritional status in the two regions, and negatively related to nitrogen deposition level in the Massif Central. As a main outcome, a strong context and spatial scale dependence of productivity changes is emphasized. In addition, the enhancing effect of climate warming on productivity is suggested, with local modulation by climatic and nutritional conditions.     

Glacial refugia and postglacial expansion of the alpine–prealpine plant species Polygala chamaebuxus

The shrubby milkwort (Polygala chamaebuxus L.) is widely distributed in the Alps, but occurs also in the lower mountain ranges of Central Europe such as the Franconian Jura or the Bohemian uplands. Populations in these regions may either originate from glacial survival or from postglacial recolonization. In this study, we analyzed 30 populations of P. chamaebuxus from the whole distribution range using AFLP (Amplified Fragment Length Polymorphism) analysis to identify glacial refugia and to illuminate the origin of P. chamaebuxus in the lower mountain ranges of Central Europe. Genetic variation and the number of rare fragments within populations were highest in populations from the central part of the distribution range, especially in the Southern Alps (from the Tessin Alps and the Prealps of Lugano to the Triglav Massiv) and in the middle part of the northern Alps. These regions may have served, in accordance with previous studies, as long‐term refugia for the glacial survival of the species. The geographic pattern of genetic variation, as revealed by analysis of molecular variance, Bayesian cluster analysis and a PopGraph genetic network was, however, only weak. Instead of postglacial recolonization from only few long‐term refugia, which would have resulted in deeper genetic splits within the data set, broad waves of postglacial expansion from several short‐term isolated populations in the center to the actual periphery of the distribution range seem to be the scenario explaining the observed pattern of genetic variation most likely. The populations from the lower mountain ranges in Central Europe were more closely related to the populations from the southwestern and northern than from the nearby eastern Alps. Although glacial survival in the Bohemian uplands cannot fully be excluded, P. chamaebuxus seems to have immigrated postglacially from the southwestern or central‐northern parts of the Alps into these regions during the expansion of the pine forests in the early Holocene.     

A simple method for estimating homotoneity of sets of phytosociological relevés

The homogeneity of sets of phytosociological relevés or syntaxa—the homotoneity—is tested by means of a homotoneity-coefficient. This coefficient consists of the basic homotoneity-coefficient expressing the proportion of highly constant species (61–100%) in the species composition of the average relevé and the correcting factor which is proportional to the difference of species number between the richest and poorest relevé. Successive analysis of heterotoneity of sets of relevés for detecting sources of heterotoneity is proposed.     

Epiphytic lichen vegetation on Larix in the Italian Alps

Abstract

A synthesis of the epiphytic lichen vegetation on larch in the Italian Alps is presented. Vegetation types are described on the basis of a numerical classification of 291 relevés. They can be compared to a few well-known phytosociological unions found in other parts of the Alps. Some communities are restricted to subalpine areas with continental climate. Despite the homogeneity of the substrate, the flora is rich, with 98 species, several of which are rare, and hence of interest from a conservational point of view.     

Differentiation among Austrian populations of Norway spruce [Picea abies (L.) Karst.] assayed by mitochondrial DNA markers

Thirty-seven populations of Norway spruce [Picea abies (L.) Karst.] across the Austrian Alps and Bohemian Massif were sampled to elucidate the geographical pattern of genetic differentiation. Three polymorphic mitochondrial DNA (mtDNA) loci were surveyed. Two or three alleles were detected at each locus, resulting in seven multilocus mtDNA haplotypes (A–F). Western populations proved to be monomorphic, whereas eastern and central Austrian populations were slightly to highly polymorphic, respectively. As revealed by spatial analysis of molecular variance and Monmonier’s analysis, the two main haplotypes A and B are not randomly distributed. Haplotype A was restricted to central and eastern Austria, whereas haplotype B occurred in all Austrian populations but was the only haplotype identified in western populations. This pattern may be explained by different glacial refugia located in the Dinaric Alps and the Carpathian mountains.     

Differentiation and speciation in mountain streams: a case study in the caddisfly Rhyacophila aquitanica (Trichoptera)

Few studies have analysed the biogeography of mountain aquatic organisms, although this habitat provides stable conditions in which many species survived Pleistocene climatic oscillations, usually in the geographical vicinity of their present distribution ranges. The mountain caddisfly Rhyacophila aquitanica was selected as a model organism for this habitat type. Morphological measurements of genitalia and external characters of male individuals were obtained from almost the entire range of distribution of the species. Morphometric results were analysed by cluster analysis and multivariate statistics. Important differences were discovered among three population groups of R. aquitanica inhabiting different European mountain ranges: (i) mountain ranges north‐west of the Alps (Massif Central, Vosges, Schwarzwald and Fribourg); (ii) the southern Alps (Lombardia and Carinthia); and (iii) the western part of the southern Carpathians. This divergence suggests a long‐term isolation among these groups, which presumably took place long before the last Pleistocene glaciation, with no secondary contact among these populations. The differentiation centres of the southern Alps and Carpathian groups may have been mostly homotopic to their actual ranges, whereas the western group must have been distributed in the areas west or north‐west of the Alps with secondary expansions and disjunctions.     

Computer assisted floristic similarity analysis

A program for polythetic numerical evaluation of phytosociological material is described. Using Sörensen's coefficient of floristic similarity it computes the homogeneity of subjectively chosen sets of relevés, the affinity of each constituent relevé to the set as well as the similarity between any pair of sets. It is also able to plot a dendrogram of the hierarchic system obtained by agglomerating individual relevés into complex groupings. The program is suitable for processing phytosociological data arranged in any conventional table.     

Mountain forest growth response to climate change in the Northern Limestone Alps

Key message

Growth response to climate differs between species and elevation. Fir is the most drought-tolerant species. The mountain forests are robust to the climatic changes until now.

Abstract

Alpine mountain forests provide a wide range of ecological and socio-economic services. Climate change is predicted to challenge these forests, but there are still considerable uncertainties how these ecosystems will be affected. Here, we present a multispecies tree-ring network of 500 trees from the Berchtesgaden Alps (Northern Limestone Alps, Southeast Germany) in order to assess the performance of native mountain forest species under climate change conditions. The dataset comprises 180 spruce, 90 fir, 110 larch and 120 beech trees from different elevations and slope exposures. We analyse the species with respect to: (1) the general growth/climate response; (2) the growth reaction (GR) during the hot summer in 2003 and (3) the growth change (GC) resulting from increasing temperatures since the 1990s. Spruce is identified as the most drought-sensitive species at the lower elevations. Fir shows a high drought tolerance and is well suited with regard to climate change. Larch shows no clear pattern, and beech remains unaffected at lower elevations. The unprecedented temperature increase of the last decades did not induce any distinct GC. The mountain forests of the Berchtesgaden Alps appear to be robust within the climatic changes until now.     

Some notes on estimation of the basic homoteneity-coefficient of sets of phytosociological relevés

The formula for calculation of the basic homotoneity-coefficient of sets of phytosociological relevés does not give adequate values for sets of 4 or 3 relevés. Therefore an empirical correction procedure is proposed for the calculation of this coefficient for such small sets of relevés. For sets of 2 relevés the calculation of the basic homotoneity-coefficient gives values equal toSørensen’s coefficient of floristic similarity. A simplification of calculation of the basic homotoneity-coefficient for sets of more than 20 relevés is proposed.     

Relict occurrences of boreal brown-moss quaking rich fens in the Carpathians and adjacent territories

Quaking rich fens dominated by boreal semi-aquatic brown-mosses such as Scorpidium scorpioides and Calliergon trifarium are extremely rare in the Carpathians. These fens harbour endangered species persisting at few localities in the region. However, their phytosociological classification has not been sufficiently solved yet, because they lack Sphagnum species as well as calcicole species characteristic for the Caricion davallianae alliance. A recent pan-European synthesis on fen vegetation suggests that these fens belong to the Stygio-Caricion limosae alliance (boreal rich fen vegetation). The isolated occurrence of this alliance southward of the boreal zone and outside the Alps is rather exceptional and might represent a relict from an early post-glacial period. In this study, we compared phytosociological data for the Stygio-Caricion limosae alliance between Northern Europe and the Carpathians plus adjacent regions (the Bohemian Massif, the Dinaric Alps) using NMDS and cluster analysis. We found that the species composition of brown-moss quaking rich fens in Central and Southeastern Europe corresponds well with that in Northern Europe, confirming their assignment to Stygio-Caricion limosae. We further reconstructed the potential past distribution of the alliance in Czech Republic and Slovakia using available floristic and macrofossil data. Macrofossil data suggest that this vegetation type had been much more common in Central Europe and that today it persists only in ancient fens, showing the long-term stability of environmental conditions. The main causes of its present-day rarity are Middle-Holocene woodland phases in fens and recent water table decreases caused by anthropogenic deterioration of the water regime in the landscape.     

Plant communities and habitat differentiation in the Mediterranean coniferous woodlands of Mt. Parnon (Greece)

Coniferous forests are a significant feature in the natural vegetation of the Mediterranean mountains, but most stands are rather degraded and the present distribution is just a fraction of its potential natural area. The Parnonas range (Peloponnese, Greece) ranks among the most extensive areas with a well-preserved mountain coniferous forest in the Mediterranean. The present paper aims at describing the conifer-dominated vegetation of this mountain and the ecological roles of the prevailing speciesAbies cephalonica, Pinus nigra, andJuniperus drupacea. For comparison, a survey is provided of theAbies cephalonica forests in the total distribution area, using all published relevés. Species composition and abundance, together with structural and abiotic parameters were recorded in 118 relevés distributed throughout Mt. Parnon. The phytosociological classification reveals 8 interpretable vegetation types well separated by groups of diagnostic species and presented in a synoptic table. The ecology and distribution of the units are outlined, and they are assigned to 4 associations within theAbietion cephalonicae (Helictotricho convoluti-Abietetum cephalonicae, Junipero drupaceae-Abietetum cephalonicae, Lilio chalcedonicae-Abietetum cephalonicae, Pyrolo chloranthae — Pinetum nigrae). The syntaxonomy and nomenclature of these associations, two of which are described as new, are discussed. The most important gradients in the data matrix found by detrended correspondence analysis (DCA) are related to altitude/climate and rockiness/soil. The communities are well segregated in the ordination space, and the fairly distinct clusters of the three conifer species are discussed. These conifers may be arranged along a gradient of decreasing drought tolerance fromJuniperus throughAbies toPinus. The latter is predominant and most vital essentially on schistose soils, or else reflects previous disturbance by, e.g., wildfires.Juniperus drupacea is a subordinate low tree or shrub in Tripolitza limestoneAbies forest. The vitality ofAbies cephalonica as expressed by tree height and structure is best in theLilio-Abietetum, between 1300 and 1600 m. The community variation within the total area of theAbietion cephalonicae reflects a principal differentiation in xerophytic and mesophytic stands. This pattern is encountered in various mountains and suggests that water supply is the crucial factor governing the floristic variation in the mountain coniferous forests.     

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

The Sieversio montanae-Nardetum strictae 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 relevés 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 pratensis, 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.     

Habitat requirements of Cardaminopsis petraea — Rare and relict species of the Czech Republic

Vegetation with Cardaminopsis petraea was sampled by phytosociological relevés at almost all the known Czech sites, and characterized by indicator values, influence of potential direct solar irradiation, and bedrock type with the aid of multivariate methods. It has been declared that C. petraea prefers open natural or seminatural drier to mesophilous habitats without some preference to the bedrock. It occupies habitats with various slopes, but it prefers more extreme slants and northerly exposed slopes. The classification and ordination methods showed clear distinction between Moravian and Bohemian habitats. Apart from other types, in Bohemia this species occurs uniquely in acidophilous rock communities, whereas in Moravia it grows uniquely within calcareous grasslands. Relictness of the species was discussed in relation to the specific site conditions (light supply, disturbance regime, and geographical isolation) and to the occurrence of other rare taxa and habitats. Occurrence on some man-made habitats was also observed.     

Genetic variation and reproduction strategy of Gentiana pannonica in different habitats

Gentiana pannonicascop. (Gentianaceae) is a long-lived perennial mountain species. It is a typical east−Alpine element. The centre of its distribution is situated in the eastern Alps, where the species occurs in alpine and subalpine areas. Besides the Alps, G. pannonica occurs also in the Bohemian Forest, Czech Republic. The reproduction strategy and genetic variation of this species endangered in the Czech Republic were studied. Genetic variation within and among six populations of G. pannonica in the Bohemian Forest and two populations in the Alps was studied using random amplified polymorphic DNAs (RAPD) markers. The variation found between populations from primary habitats and populations situated in the secondary mountain meadows accounted for 5% of the total genetic variation, whereas no significant genetic diversity was found between two different regions (Bohemian Forest, the Alps). Most of the variation is distributed among individuals within populations in the primary habitats, specifically 77% in the Bohemian Forest and 79% in the Alps. The pattern of among-population variation was substantially different between primary and secondary habitats. While among-population variation in primary habitats was low in both of the regions (21–23%), the proportion of RAPD variation among populations in secondary habitats was distinctly higher (65%). Within-population genetic variation was higher in primary habitats than in the secondary ones. This may be a consequence of processes such as genetic drift or bottleneck, and founder effects caused by changes in their distribution during the Holocene period. The differences of vegetation composition, seedlings recruitment and seed set were investigated in the field. Vegetation composition differed between primary and secondary habitats of G. pannonica and seedlings recruitment was recorded only in primary habitats. The reproduction output of the species under study is probably influenced by the availability of pollinators.     

Neophyte Invasion in Moldavia (Eastern Romania) in Different Habitat Types

The actual state of neophyte invasion in Moldavia (Eastern Romania) is described in this paper on the basis of 11,055 phytosociological relevés. We analyzed the i) proportion of relevés with neophytes, ii) mean proportion of neophytes per relevés, and iii) mean coverage of neophytes per relevé for 36 EUNIS habitat types to identify general plant invasion patterns. The level of invasion differed considerably between habitats. The invasion of neophytes especially affected habitats strongly determined or influenced by man, such as anthropogenic woodlands, ruderal habitats, arable lands or trampled areas. Most natural habitats are either slightly invaded or entirely free of neophytes. Only riverine willow stands and wet tall-herb stands are relatively highly invaded. However, the absence of neophytes in some natural habitats less represented in the phytosociological dataset could be of artifactual nature. No significant relationship between the number of neophytes and non-neophytes was found in the analysis across different habitats. When the analyses were made within-habitats, both negative and positive relationships were found, which confirm that the relationship between alien and native species richness depends on the habitats. A total number of 105 neophyte species were recorded in the phytosociological relevés used in this study. Among these, 13 species that are currently considered invasive in Moldavia occur in at least 10 types of habitats.     

Determination of diagnostic species with statistical fidelity measures

Abstract. Statistical measures of fidelity, i.e. the concentration of species occurrences in vegetation units, are reviewed and compared. The focus is on measures suitable for categorical data which are based on observed species frequencies within a vegetation unit compared with the frequencies expected under random distribution. Particular attention is paid to Bruelheide's u value. It is shown that its original form, based on binomial distribution, is an asymmetric measure of fidelity of a species to a vegetation unit which tends to assign comparatively high fidelity values to rare species. Here, a hypergeometric form of u is introduced which is a symmetric measure of the joint fidelity of species to a vegetation unit and vice versa. It is also shown that another form of the binomial u value may be defined which measures the asymmetric fidelity of a vegetation unit to a species. These u values are compared with phi coefficient, chi‐square, G statistic and Fisher's exact test. Contrary to the other measures, phi coefficient is independent of the number of relevés in the data set, and like the hypergeometric form of u and the chi‐square it is little affected by the relative size of the vegetation unit. It is therefore particularly useful when comparing species fidelity values among differently sized data sets and vegetation units. However, unlike the other measures it does not measure any statistical significance and may produce unreliable results for small vegetation units and small data sets. The above measures, all based on the comparison of observed/expected frequencies, are compared with the categorical form of the Dufrêne‐Legendre Indicator Value Index, an index strongly underweighting the fidelity of rare species. These fidelity measures are applied to a data set of 15 989 relevés of Czech herbaceous vegetation. In a small subset of this data set which simulates a phytosociological table, we demonstrate that traditional table analysis fails to determine diagnostic species of general validity in different habitats and large areas. On the other hand, we show that fidelity calculations used in conjunction with large data sets can replace expert knowledge in the determination of generally valid diagnostic species. Averaging positive fidelity values for all species within a vegetation unit is a useful approach to measure quality of delimination of the vegetation unit. We propose a new way of ordering species in synoptic species‐by‐relevé tables, using fidelity calculations.