Regeneration strategies of dominant boreal forest dwarf shrubs in response to selective removal of understorey layers

Abstract. The deciduous Vaccinium myrtillus and the evergreen Vaccinium vitis‐idaea were subjected to five removal treatments of understorey layers: control, removal of the moss layer, removal of the field layer, removal of both moss and field layers and removal of moss, field and humus layers. A second factor, sowing, was included to investigate sexual reproduction after disturbance. Density of new ramets and seedlings and growth of annual shoots were studied for the first two growing seasons, whereas cover was measured for five growing seasons after disturbance treatment. Initially, vegetative production of new ramets and species cover increased rapidly in all disturbed plots, except for the most severe treatment, in which production of new ramets was virtually absent throughout the study. Full recovery following removal of the field layer only or both field and moss layers was reached after four years for V. myrtillus. V. vitis‐idaea recovered more quickly, after one year (removal of field layer only) and four years (removal of field and moss layers). The relative growth of V. myrtillus and V. vitis‐idaea increased in the latter treatment in terms of production of annual shoots and length of annual shoots, respectively. Seedling density increased after sowing in the most severe treatment. The results underscore the importance of vegetative growth for recovery of these species at moderate‐level disturbances. The high rate of sexual reproduction in the most severe treatment implies that strong mechanical disturbance is needed to enhance the establishment of new genotypes in these species.     

Seedling establishment in relation to microhabitat variation in a windthrow gap in a boreal Pinus sylvestris forest

Abstract. The characteristics of microhabitats of established Pinus sylvestris and Betula seedlings were studied in a small windthrow gap in a mature P. sylvestris-dominated forest in the Petkeljärvi National Park in eastern Finland. Seedlings were strongly clustered in disturbed microhabitats, particularly uprooting pits and mounds, formed by tree falls. They covered 3% of the 0.3.ha study area consisting of the gap and some of the forest edge. Although Betula occurred only as scattered individuals in the dominant canopy layer of the forest, it accounted for 30% of the seedlings found in the study area. Betula regeneration was almost completely restricted to pits and mounds, where 91% of the seedlings were found. Uprooting spots were also the most important regeneration microhabitats for Pinus, where 60% of the seedlings grew, even though the seedlings were found in other substrates as well, particularly on sufficiently decomposed coarse wood. Undisturbed field- and bottom-layer vegetation had effectively hindered tree seedling establishment, which emphasises the role of soil disturbance for regeneration. While the establishment of seedlings was found to be clearly determined by the availability of favourable regeneration microhabitats, the early growth of seedlings was affected by a complex interaction of environmental variables, including the type of microhabitat, radiation environment and interferences caused by competing seedlings and adjacent trees. In the most important regeneration microhabitats, i.e. in uprooting pits and on mounds, the distributions of the local elevations of Pinus and Betula seedlings were different. Pinus seedlings occurred closer to ground level, i.e. on the fringes of pits and lower on mounds, while Betula seedlings grew deeper in pits and higher on mounds. The position of the Betula seedlings indicate that they may have a competitive advantage over Pinus seedlings in the dense seedling groups occurring in uprooting spots. We suggest that this initial difference in Pinus and Betula establishment may affect the subsequent within-gap tree species succession and can, in part, explain the general occurrence of Betula in conifer-dominated boreal forests.     

Digitized photographs in vegetation analysis ‐ a comparison of cover estimates

Abstract. The use of digital images in measuring plant species cover and species number in boreal forest vegetation was studied. Plant cover was estimated by manual delineation on photographs and subsequent digitized measurement of the areas. This value was regarded as the reference and compared with cover obtained by visual estimate, point frequency and automatic image analysis methods. The automatic image analysis was based on scanned photographs. Supervised image classification with ERDAS software was then used to distinguish between the covers of different plant species. When comparing the ability of the methods to detect species, the visual estimate method gave values similar to the reference. The study material was collected from three sites along a heavy‐metal pollution transect in western Finland. All four methods detected, in a similar manner, differences between the plant species abundances along the transect. Compared with the reference, the digital images underestimated the cover of the lichens Cladina spp. and Cetraria islandica, but gave similar estimates for the dwarf shrubs Empetrum nigrum and Vaccinium vitis‐idaea. The point frequency method overestimated the cover of all the species studied. The visual estimates of lichens were close to the reference, while the dwarf shrub covers were overestimated. The number of species detected using supervised image analysis and the point frequency method was lower than that with visual estimation. Visual estimation was faster, and the estimate closer to the reference cover values than the others. Digital images may be useful in detecting changes in some selected species in vegetation with a simple vertical structure but with taller, multilayered vegetation and a higher species number, the reliability of the cover estimates is lower.     

Sixty years of vegetation dynamics in a south boreal coniferous forest in southern Norway

Abstract. Vegetation data from permanent plots were collected in 1931, 1961 and 1991 in a south boreal forest 20 km north of Oslo in southern Norway. Major changes were found in the vegetation composition during those 60 years. The main changes were a reduction in the frequency of species and the frequency of joint occurrences of vascular species such as Andromeda polifolia, Calluna vulgaris, Cornus suecica, Eriophorum vaginatum, Maianthemum bifolium, Melampyrum pratense, Trientalis europaea, Vaccinium uliginosum and V. oxycoccus, and mosses, e.g. Dicranum fuscescens, Hylocomium splendens, Pleurozium schreberi, Ptilidium ciliare and Ptilium crista-castrensis. The observed changes were interpreted as being induced by internal processes e.g. notably a long-term change from paludified forest to mesic forest. In particular the growth of Picea abies seems to be a main driving force. The dominance of Picea abies and Vaccinium myrtillus appears to have rendered the conditions more unfavourable for other species. A doubling of the living stem biomass of P. abies during the last 67 yr shows that this old-growth forest has not yet reached a steady state. It was demonstrated that species such as Deschampsia flexuosa and Molinia caerulea did not increase in frequency in response to nitrogen deposition, as has occurred elsewhere in northern Europe. pH in the humus layer increased with 0.2 unit from 1961 to 1991. The results of this study indicate that protection from logging has initiated the reduction of species in the field layer and bottom layer. This study questions if monitoring of forest vegetation should be restricted to protected forests as is the practice in Scandinavia today. We recommend that also areas with some kind of selective cutting will be used for monitoring of forest vegetation.     

Effects of reindeer grazing on understorey vegetation in dry Pinus sylvestris forests

Abstract. Data on floristic composition and environmental variables were collected in floristically homogeneous oligotrophic pine (Pinus sylvestris) forests with heath-like under- storey vegetation in eastern Fennoscandia, and ordinated by non-linear multidimensional scaling (NMDS) in order to study the effect of lichen grazing by reindeer on the understorey vegetation. The study sites included areas with varying grazing pressure, as well as 50-yr old grazing exclosures. Sites rich in respectively bryophytes and lichens were placed at opposite ends of the ordination axes, and heavily grazed sites were placed in between them. Reindeer grazing increased the abundance of bryophytes, especially Dicranum spp. and Pleurozium schreberi. Grazing changed the vegetation to the extent that it resembled more mesotrophic sites, but this did not show any relationship with tree volume or other site productivity indicators. This was observed both in the ordination and, in a more compelling way, when exclosures with adjacent grazed areas were compared. No such signs were evident at ungrazed sites, where especially Cladina spp. spatially replace Cladonia spp. and tiny bryophytes like Barbilophozia spp., Polytrichum spp. and Pohlia nutans during succession. Cladina stellaris had almost disappeared from the most intensively grazed sites. The soil at ungrazed sites was characterized by high Al and Fe concentrations and bryophyte-rich sites by high Mn concentrations. Shannon's diversity index, depth of humus layer and proportion of bare ground also increased in sites getting richer in bryophytes.     

Large‐scale changes in abundance of terricolous bryophytes and macrolichens in Finland

Abstract. Major changes in forest floor vegetation were identified on the basis of three nationwide surveys conducted as part of national forest inventories in 1951–1953, 1985–1986 and 1995. These surveys provided objectively selected, statistically representative samples of all forested land in Finland. The 1951–1953 data consist of over 10000 sample plots, while the later surveys were conducted on ca. 3000 permanent plots. Changes in relative abundance of dominant species (i.e. in the proportions of species of the total cover of forest floor vegetation) were analysed across biogeographical provinces. Spatial correlation, systematic sampling, partial re‐measurement and multiple testing were taken into account in assessment of the statistical significance of the observed changes. The most notable changes in forest floor vegetation were a decrease in the relative abundance of Hylocomium splendens and an increase in Dicranum polysetum. In N Finland, where forests are grazed by semi‐domestic reindeer, we observed a decline in the abundance of Cladina lichens and an increase in Dicranum mosses. Peltigera aphthosa declined throughout the country. Polytrichum juniperinum, Pohlia nutans, and Brachythecium species, which occupy disturbed sites or grow on litter, increased in abundance. The relative abundance of Sphagnum species decreased, particularly in W Finland, where Pleurozium schreberi was favoured. A major decline in S. fuscum was also recorded in C and E Finland. Many of the changes detected in this study are apparently related to intensified forest management; but solely on the basis of this study, its effects cannot be distinguished from those of other large‐scale environmental changes.     

Nutrient limitation in boreal plant communities and species influenced by scything

Abstract. The nutrient status was studied in permanent plots of four plant communities, two rich-fen communities and two wooded grassland communities, all formerly used for haymaking. The concentrations of N, P and K in plant material of dominant and subdominant species (above- and below-ground) were measured in plots experimentally scythed annually or biennially for two decades, and in plots unscythed for four decades. Three of the communities had an N:P ratio of 14 or less, indicating N-limitation; the most fertile grassland community had particularly low values for the N:P ratio (6–12), as did a majority of the species, including all tall-herb species. A species-rich community of fen-margin vegetation in the lowest productive rich fen, had an N: P ratio of 17–19 in the above-ground biomass, which indicates P-limitation of nutrients. Molinia caerulea and Thalictrum alpinum were found to be the vascular plants with the highest N:P ratio, indicating P-limitation of nutrients. Calculations of N:K and K:P ratios indicated possible K-limitation in the rich-fen communities, especially for Thalictrum alpinum, the species with the highest N:K value. No expected change from N- to P-limited growth was found; in contrast, a reduction in the N:P ratio was found in the annually scythed plots of the rich fens, suggesting that reduced biomass production is mainly a result of disturbance by scything. As expected, a reduction in the concentration of K was detected in the scythed plots.