On the ecology of Thelypteris limbosperma — a synecological investigation of T. limbosperma-dominated stands in W Norway

The investigation is based on phytosociological and ecological analyses of T. limbosperma-dominated stands from different parts of W Norway. The distribution and floristical composition of the stands are related to different environmental parameters, such as climate, soil and topography. The data are analysed by numerical methods, of which two-way indicator species analysis (TWINSPAN), canonical correspondence analysis (CCA), and simple discriminant functions (DISCRIM) are the most useful. The broad-scale distribution of T. limbosperma stands is determined by climatic factors. Within its distribution area soil factors and human influence govern the floristic composition and development of different types of such stands. Based on the TWINSPAN classification 9 different stand types are distinguished.
The first major division of the TWINSPAN classification is mainly correlated with soil factors. Most of the stands growing on the richest soils are interpreted as representing regeneration stages after intensive human influence. Divisions at lower levels are well correlated with altitude and temperature conditions.
The stands are all characterized by low pH and base saturation. Some types are concentrated to certain geographical regions, indicating that both altitudinal and west-east-gradients are important in influencing the floristic composition of the different types. The investigations indicate that T. limbosperma has a rather narrow realised phytosociological niche. It is mainly concentrated to stands that are floristically most related to the poorest part of Vaccinio-Piceion communities. Occasionally it is also dominant in Alno-Padion and Lactucion alpinae communities.     

A synecological investigation of Matteuccia struthiopteris — dominanted stands in Western Norway

This investigation is based on phytosociological and ecological analyses of stands dominated by M. struthiopteris within its main distributional area in Western Norway. The distribution and floristical composition of such stands are related to different environmental parameters such as regional climate, soil and topography. The data are analysed by numerical methods, of which two-way indicator species analysis (TWINSPAN), canonical correspondence analysis (CCA), and simple discriminant functions (DISCRIM) are the most useful. Few stands occur in areas where the mean July temperature is below 12.0 °C, and where the mean January temperature is higher than 0 °C. The soil is rich in metal cations, with a base saturation mostly higher than 50% in the B-layer. Based on a TWINSPAN classification of the 230 investigated stands, 9 different groups (types) are distinguished. Many stands have a tree layer of Alnus incana, while stands with a canopy of Betula pubescens, Fraxinus excelsior and Ulmus glabra are considered to be marginal sites for M. struthiopteris. The types described are interpreted as belonging to the Pruno padi-Alnetum incanae, Alno incanae-Fraxinetum excelsioris and Alno incanae-Ulmetum glabrae associations within the Alno-Ulmion alliance. The floristic composition of the quadrats is highly correlated with the altitude/temperature gradient. The other important factor is interpreted to be soil moisture. Soil richness is poorly correlated with the floristic composition of the different stands. Several stands represent a regeneration stage after cessation in cultivation, while others appear to have been very little influenced by human activity.     

Plant communities in broad-leaved deciduous forests in Hordaland county, Western Norway

The vegetation of broad-leaved deciduous forests in Hordaland, Western Norway is described with regards to species composition and structure. The investigation is based on phytosociological analyses of native forest stands dominated by Corylus avellana, Fraxinus excelsior, Ulmus glabra , or Tilia cordata. Two-way-indicator analysis (TWINSPAN) and correspondence analysis (CA) are used to distinguish different vegetation types and assess possible gradients in the vegetation data. The vegetation types are discussed in relation to equivalent forest communities and to syntaxonomical units. Two forest types are described at the first hierarchical TWINSPAN level: 1) A hygrophilous and slightly thermophilous Fraxinus excelsior-Cirriphyllum piliferum forest characterised by an open tree canopy, a dense field layer of tall herbs and ferns, and a high cover of bryophytes. 2) A thermophilous and less hygrophilous Corylus avellana-Brachypodium sylvaticum forest characterised by a dense tree layer, a more open field layer with larger elements of small herbs, and a somewhat lower bryophyte cover. The CA analysis clearly separates the samples from the first TWINSPAN division along the first ordination axis. Five forest types have been described at the second hierarchical level, mainly associated with differences in mesotrophic and eutrophic species, but there is clearly a gradient structure in the species composition between these plant communities. In relation to syntaxonomy, the first TWINSPAN division supports the separation of the west Norwegian broad-leaved deciduous forests into a hygrophilous plant community, the Eurhynchio-Fraxinetum (Blom 1982) Øvstedal 1985 and a drier and more thermophilous community, the Primulo-Ulmetum (Blom 1982) Øvstedal 1985.     

Size and reproduction of Thelypteris limbosperma and Athyrium distentifolium along environmental gradients in Western Norway

The performance of the ferns Thelypteris limbospgrma and Athyrium distentifolium , as expressed by number of son on pinnules and frond size, has been related to environmental gradients. Data have been collected from homogenous fern-dominated stands from different geographical areas in Western Norway. Number of sori produced, which was assumed to indicate fern fertility, was highly variable. There were significant correlations between frond size and fertility, but only 35–46 % of the variation in fertility could be explained by differences in frond size. The highest number of sori was found on plants of medium size, and for A. distentifolium there was a statistically significant reduction in sori number on fronds taller than 125 cm. For the linear relationships between fertility and size, it is suggested that the small X-intercept and slope defines the high competitive ability of A. distentifolium in sites with a long-lasting snow cover. The ferns may dominate in large stands even though their fertility is low or zero, which indicates that they under certain ecological conditions mainly reproduce vegetatively. Both ferns show decreased fertility both toward their altitudinal and lowland distribution limits, but their fertility optima were different. The performances of the two fern species showed highly different responses to geographical and climatical variables. The fertility of T. limbosperma could be predicted by mean July- and January temperatures, humidity and canopy cover, while the fertility of A. distentifolium could not be predicted by any of the site environmental variables investigated.     

Nestedness,SLOSS and conservation networks of boreal herb‐rich forests

Question: Herb‐rich patches are biodiversity hotspots for vascular plants in boreal forests. We ask: Do species occurrences on herb‐rich patches show a non‐random, nested structure?; Does patch size relate to richness of edaphically demanding and red‐listed species?; Does a set of small patches support more edaphically demanding and red‐listed species than a few large patches of the equal area? Location: Eastern Finland (63°04′N, 29°52′E), boreal vegetation zone. Data: Vegetation mapping of 90 herb‐rich sites, varying from 0.05 to 6.93 ha in size and belonging to six different, predetermined forest site types. Results: Using the RANDNEST procedure, only one site type showed a significantly nested pattern, and patch area was not related to “nestedness” in any of the site types. The number of edaphically demanding and red‐listed plant species was positively correlated with a patch size in three forest site types. In all site types, a set of small patches had more edaphically demanding and red‐listed species than did a few large patches of the equal total area. Conclusions: For conservation, it is essential to protect representative sets of different herb‐rich forest site types because flora varies between the site types. Within herb‐rich forest site types, several small areas may support representative species composition. However, successful conservation requires thorough species inventories, because of the high level of heterogeneity between the herb‐rich patches.     

Quantitative vegetation-environment relationships in west norwegian tall-fern vegetation

The aims of this paper are to detect floristic variation within different types of tall-fern dominated vegetation and to interpret these patterns in terms of environmental variables. Numerical approaches have been applied to a large and varied vegetational data-set with associated environmental data from stands dominated by Athyrium distentifolium, Thelypteris limbosperma , and Matteuccia struthiopteris in different parts of western Norway. The numerical procedures of two-way indicator species analysis, simple discriminant functions, and canonical correspondence analysis have been used, and the strengths and weaknesses of these as tools in discerning vegetational-environmental relationships are discussed. For each of the 96 quadrats investigated, 17 environmental variables were measured. The investigation shows that some of the observed differences in vegetational composition can be explained in terms of relatively simple soil and climatic variables measured for each quadrat. The ferns appear to be ecologically well separated. T. limbosperma-dominaled stands are mainly characterised by low soil fertility, high January temperature, and high humidity. A. distentifolium-dominated stands are associated with low winter temperatures, and M. struthiopteris-dominated stands have high soil fertility and high summer temperatures.     

Classification of some North Norwegian forest types

The vegetation in 19 selected forest stands in N Norway north of Saltfjellet mountains have been described and classified. Two different classification methods have been used: An ordinary phytosociological classification, and an automatic, unweighted classification. The two methods give very similar results. All the investigated forests belong in the class Vaccinio-Piceetea. Some differences from South Norway seem to exist, e.g. Empetrum hermaphroditum and Vaccinium uliginosum are in the present investigation regarded as regional characteristic species of the class. The associations Barbilophozio-Pinetum and Eu-Piceetum are treated in some detail. Barbilophozio-Pinetum is divided into three variants: a "continental" variant without Calluna vulgaris , a "Rhacomitrium" and a "Piceetum" variant with dominance of Calluna vulgaris. The two former variants are clearly to be classified as belonging in Cladonio-Vaccinietalia, while Vaccinio-Piceetalia species are abundant in the "Piceetum" variant. The syntaxonomic position of this variant is uncertain. The material does not support a subdivision of Eu-Piceetum into a Myrtilletosum and a Dryopteridetosum subassociation in N Norway.     

Taxonomy and ecology of <Emphasis Type="Italic">Castanea sativa</Emphasis> Mill. forests in Greece

Castanea sativa Mill. is an important species of the Balkan Peninsula with high ecological and economic value. This article contributes to a phytosociological synsystematic approach of the C. sativa plant communities in Greece that covers the southern part of the Balkan Peninsula and describes the ecological parameters involved in their distribution and taxonomy. Phytosociological data have been collected from 14 mountainous areas across continental Greece, which are considered representative of the main forest types of C. sativa in the area. The 14 mountains demonstrate a variety of environmental conditions. Five communities (associations) and seven sub-communities (sub-associations) have been identified and described using TWINSPAN, Corespondence Analysis and the Braun-Blanquet classification scheme. Castanea sativa (chestnut) forests of Greece can be distinguished as three broad groups, defined primarily by their geographical distribution. In northern and central Greece chestnut stands are characterized by their high floristic homogeneity, which is reflected in the identification of only one community per region. Chestnut forests in northern Greece, in particular, exhibit strong floristic similarities with those of the rest of the Balkans. In southern Greece, on the other hand, the southern distribution limit of chestnut in the Balkans, there is a greater floristic diversity between the sampled mountains which is reflected in the presence of three confirmed communities and two sub-associations. Several factors have been identified as important in determining the structure and floristic composition of chestnut forests in Greece. Whilst climate and the grazing regime are influential, the degree of silvicultural management appears to be the most important factor determining the floristic composition of chestnut forests and their long-term sustainability.     

Threats affecting the distribution of Lygeum spartum Loefl. ex L. and its associated species in the Western Mediterranean desert of Egypt: Perspectives of conservation

The current study intends to evaluate the many hazards affecting the spread of the significant fodder grass Lygeum spartum Loefl. ex. L. and its associated species in Egypt's Western Mediterranean Desert over the past 60 years (from 1958 to 2020); depending on the data of the previous literature and the current situation evaluation by recent field trips. It also aims to assess its ecosystem services. Sixty-nine stands were collected from previous studies and unpublished data, in addition to 30 stands (10 m²) were added including 5 field trips during the period from spring 2019 to spring 2020. Measurements were taken to create a classification of this vegetation and to determine the main threats that affect its distribution composition. There are 142 species associated with the distribution of the studied plant in 6 habitats. While only 107 species were recorded in the recent stands in 7 habitats. Vegetation groups (G1- G8) obtained from TWINSPAN classification indicating that G1 (wadis) is the most diverse group with a species richness 34.7 species per stand, while G6 (dune) has the highest species turnover (4.57) during the period of 1958 to 2014. But in recent stands; G5 (saline and non-saline depression) is the most diverse group (30.875), while G2 (dunes) has the highest species turnover (4.819). Medicinal plants are the most represented (79.86%), followed by grazing plants (72.9%). The most common service is sand stabilization (92% of total services), followed by windbreak (natural) (40%). Threatened species represent 92.4%; habitat loss is the most affected threat (95 species), followed by over-collecting and over-cutting (82 species). Twenty out of 99 stands (20.2% of the total stands) have been destroyed due to human activity. Continuous monitoring of remaining patches of Egypt’s northwestern Mediterranean coast was required to save its natural flora from extinction.     

Bryophyte community composition and habitat specificity in the natural forests of Terceira,Azores

In order to characterize the main areas of natural forest on Terceira Island (Azores), some of the most interesting European forests due to their rich and diverse flora of endemic as well as relict species, six forest stands were studied and their flora inventoried using 226 randomized quadrats, which revealed one anthocerote, 64 liverworts, 41 mosses and 16 macrolichen taxa. The alpha-diversity of the samples is particularly species-rich, some quadrats (30 × 30 cm) including more than 25 bryophyte species. A quantitative analysis of the vegetationenvironment relationships consistently showed that the distributions of the native forest bryophytes and lichens of Terceira are governed by a complex set of factors related to water availability, the status of the substrata and the influences of the vascular plant community. Considering the generally high values of the Sørensen indices and the low number of specialist species found with Lloyd’s index, the differences are more in terms of dominance of species than in terms of species composition. In fact, substratum type was clearly important in the DCA and TWINSPAN analysis using species cover abundance values, largely explaining the distribution of bryophyte species between Juniperus brevifolia (Seub.) Antoine and Laurus azorica (Seub.) Franco bark. The eight putative plant community type groupings achieved with these multivariate methods were able to elucidate some major bryophyte – substratum relationships, that had not previously been considered and they offer a framework for future research.     

Habitat distribution and conservation of land bird populations in northern Norway

About 300 km line transect censuses of breeding land birds were made in northern Norway in 1974–76. The data (1020 pairs in 26 samples) were grouped into seven major habitats: stony ground and oceanic heath (C1), continental dry heath (C2). "'2Calcarius heath", dry peatland, wet peatland (C5), meagre and average birch forest, and rich birch forest (C7). Bird density ranged from 22 pairs km⁻² (C2) to 364 pairs km⁻² (C7), but most habitats had 67–96 pairs km⁻². The bird communities are described.
Defining the Optimal habitat for a species as the maximum-density habitat, the 43 species observed were found to comprise three major groups: tundra species (optimal habitat C1), peatland species (C5) and forest species (C7). Species whose central range includes northern Norway had a broader habitat amplitude than those with a southern or a very northern distribution. A new index was devised for the evaluation of the conservational importance of a habitat or a species. From the North European standpoint. C1 is certainly the most important habitat in northern Norway, as C1 supports many species which are rare in other parts of North Europe. However, population trends should also be taken into account when conservational preferences are judged.     

What is the upper size limit for cosmopolitan distribution in free‐living microorganisms?

A comparison of testate amoebae assemblages from the Arctic and Antarctic (areas of similar habitat a maximum distance apart) is used to try and answer the question ‘What is the upper size limit for cosmopolitan distribution in free‐living microbes?’ Species restricted to either the Arctic or Antarctic exhibited sizes up to 230 μm while the largest cosmopolitan species was 135 μm in size. Comparison of the testate assemblages using a multivariate classificatory technique (TWINSPAN ) also suggested more restricted distribution for the larger species. There was a negative relationship between species size and number of sites at which it was recorded (r_s=?0.261, P < 0.05), with all the more widespread species having a size of below 100 μm. It is suggested that for testate amoebae cosmopolitan distributions become common below 100–150 μm. This suggests that most species of testate (indeed most free‐living microbes) have low species richness because of lack of opportunities for allopatric speciation as most are below 100 μm and so geographical isolation is unlikely. It is suggested that if this is correct, only the largest free‐living microbes (> 150 μm) are likely to be of conservation concern because of their smaller ranges. However, I point out that currently different studies are giving very different answers to the question, how ubiquitous and species rich are free‐living microbes? The subject requires further work to try and reconcile these different results.     

Distributional patterns of vascular plants in Fennoscandia: a numerical approach

The distribution of 725 species and 4 subspecies of vascular plants in 522 grid squares of 50×50 km within Denmark, Finland, Norway and Sweden was analysed by detrended correspondence analysis (DCA) and two-way indicator species analysis (TWINSPAN). Two phytogeographical gradients were recognized: a major gradient running both latitudinally and altitudinally, from southern and lowland areas to northern and mountain districts, and from southerly distributed species to alpine and northerly distributed species, and a secondary gradient running from eastern districts and easterly distributed species to western districts and westerly distributed species. The results are in agreement with previous recognized phytogeographical patterns within the considered area. Possible ecological and historical causes for these patterns are briefly discussed.     

Upward shift in elevational plant species ranges in Sikkilsdalen,central Norway

Phytosociological studies can be an important tool to detect temporal vegetation changes in response to global climate change. In this study, we present the results of a resurvey of a plot‐based phytosociological study from Sikkilsdalen, central Norway, originally executed between 1922 and 1932. By using a detailed phytosociological study we are able to investigate several aspects of elevational shifts in species ranges. Here we tested for upward and downward shifts in observed upper and lower distribution limits of species, as well as changes in species optima along an elevational gradient, and related the observed range shifts to species traits that could explain the observed trends. More species shifted upwards than downwards, independently of whether we were investigating shifts in species’ upper or lower distribution ranges or in species optima. However, shifts in species upper range margins changed independently of their lower range margins. Linking different species traits to the magnitude of shifts we found that species with a higher preference for prolonged snow cover shifted upwards more in their upper elevational limits and in their optima than species that prefer a shorter snow cover, whereas no species traits were correlated with the magnitude of changes in lower limits. The observed change in species ranges concord both with studies on other mountains in the region and with studies from other alpine areas. Furthermore, our study indicates that different factors are influencing species ranges at the upper and lower range limits. Increased precipitation rates and increased temperatures are considered the most important factors for the observed changes, probably mainly through altering the pattern in snow cover dynamics in the area.     

Towards unification of national vegetation classifications: A comparison of two methods for analysis of large data sets

Abstract. In European phytosociology, national classifications of corresponding vegetation types show considerable differences even between neighbouring countries. Therefore, the European Vegetation Survey project urgently needs numerical classification methods for large data sets that are able to produce compatible classifications using data sets from different countries. We tested the ability of two methods, TWINSPAN and COCKTAIL, to produce similar classifications of wet meadows (Calthion, incl. Filipendulenion) for Germany (7909 relevés) and the Czech Republic (1287 relevés) in this respect. In TWINSPAN, the indicator ordination option was used for classification of two national data sets, and the extracted assignment criteria (indicator species) were applied crosswise from one to the other national data set. Although the data sets presumably contained similar community types, TWINSPAN revealed almost no correspondence between the groups derived from the proper classification of the national data set and the groups defined by the assignment criteria taken from the other national data set. The reason is probably the difference in structure between the national data sets, which is a typical, but hardly avoidable, feature of any pair of phytosociological data sets. As a result, the first axis of the correspondence analysis, and consequently the first TWINSPAN division, are associated with different environmental gradients; the difference in the first division is transferred and multiplied further down the hierarchy. COCKTAIL is a method which produces relevé groups on the basis of statistically formed species groups. The user determines the starting points for the formation of species groups, and groups already found in one data set can be tested for existence in the other data set. The correspondence between the national classifications produced by COCKTAIL was fairly good. For some relevé groups, the lack of correspondence to groups in the other national data set could be explained by the absence of the corresponding vegetation types in one of the countries, rather than by methodological problems.     

Restoration of the Cirsio dissecti‐Molinietum in The Netherlands: Can we rely on soil seed banks?

Abstract. Vegetation and soil seed banks of a threatened Atlantic fen meadow community were studied using recent phytosociological records and seedling emergence from soil samples. Similarly managed but differently degraded stands that suffered different levels of species impoverishment were compared. The actual vegetation was related to a set of phytosociological references representing the subassociations of the community. DCA positions of reference relevés from the different subassociations were overlapping, suggesting that in all references many common species occur. Recent records were positioned in‐between the seed bank samples and the references. The soil seed banks of all stands were dominated by ordinary species. Most character species had at most sparse seed banks and no seedlings of locally extinct character species, mentioned in historic floristic records, were detected. In contrast species of pioneer and small‐sedge communities as well as those of heathlands were abundant in the seed banks. Based on the vertical distribution of seeds in the soil layers most fen meadow species were classified into transient or short‐term persistent seed bank types. We concluded that complete restoration of the Cirsio dissecti‐Molinietum without reintroduc‐tion is only likely in stands that were degraded only a few years ago. On the other hand, the presence of viable seeds of Nanocyperion and Parvocaricetea species is promising for the restoration of these communities even after decades. Recreation of pioneer habitats by sod cutting will preserve these species.     

Geospatial modelling approach for identifying disturbance regimes and biodiversity rich areas in North Western Himalayas, India

The present study is a comprehensive effort for making spatially explicit vegetation type information, one of the basic inputs for species and habitat conservation, readily available to the decision makers, resource managers and nature conservationists. The present study was carried out to understand the vegetation composition and structure in Doda area of Western Himalayas, India. During the study, vegetation types were mapped using on-screen image interpretation technique of multispectral high resolution satellite data. A total of ten types of vegetation were delineated from the satellite data. Phytosociological data was collected for the forest, pasture and scrub classes using nested-quadrat approach to characterize the vegetation. A total of ten phytosociological parameters were analyzed. Pinaceae, Rosaceae and Asteraceae were the dominant plant families with most of the identified plant species having a very high medicinal value. Other important component of the study involved landscape modelling, using the Spatial Landscape Analysis Model for identifying disturbance regimes and biodiversity rich landscapes in the area. The model results indicate that most of the area contains a very rich biodiversity repository with only a few areas showing signs of disturbance where terrain is either complex or where the anthropogenic pressures on forest resources are apparent. The forest and nature conservation managers could use the conservation measures suggested on the basis of these research findings for developing biodiversity conservation strategies in the region.     

Rhythmic growth fluctuations in forest trees of Central Europe and the Front Range in Colorado

Summary Growth ring sequences in conifers and broadleaved trees show abrupt growth reductions persisting for several years. The frequency and duration of such reductions in almost 10000 trees in forest stands of Central Europe, in particular Switzerland, and in the Front Range, Colorado, USA, is related to species, climate and weather, and geographical unit. The growth reductions in the populations investigated are concentrated in certain periods and, in all species examined, essentially related to periods of drought in the spring and summer months. During the years covered by the study (1850–1985), periods of growth reduction occur in Central Europe, the Front Range, Colorado, and also in the Hudson Valley, New York, at irregular intervals of 6–34 years, and on average every 13–14 years. The growth reduction periods normally do not show synchronism between the three areas investigated. Regionally valid growth periodicities can only be definitely determined through the examination of a very large amount of material comprising a number of species growing in different forest stands and on different sites within a major phytosociological unit. Findings from studies on trees in different climatic areas or on few samples scarcely allow reliable comparisons.     

Ecogeography of the grass genusHelictotrichon (Poaceae: Aveneae) in the Mediterranean and adjacent regions

The geographical distribution of the grass genusHelictotrichon in the Mediterranean Region is analysed and mapped for 33 taxa. Based on only a single life form (perennial herb), a variety of edaphically, climatically and altitudinally differently adapted species complexes has evolved in the area. Most of these complexes show west-east disjunctions and contain geographically, sometimes even edaphically vicarious taxa with complementary distribution. A transition from mesomorphic to xeromorphic habit occurred independently in different species groups and led to the establishment of the modern Mediterranean taxa which are in part highly polyploid derivatives of more mesophilic diploids. The significance of polyploidy, patterns of parapatric and sympatric distribution, biogeographical borders, and centres of species diversity are discussed in context with the history of the Mediterranean vegetation. New combinations are:Helictotrichon setaceum subsp.petzense, H. pratense subsp.lusitanicum, H. praetutianum.