Vertical distribution of the ectomycorrhizal community in the top soil of Norway spruce stands

The vertical distribution of the ectomycorrhizal (ECM) community was studied in four old high-mountain Norway spruce (Picea abies [L.] Karst.) stands in northern Italy. The aim was to verify if the variability in the community structure could be explained by characteristics of the organic and mineral soil horizons. The community structure was evaluated in terms of both fungal species and their ability to explore soil (exploration types). From the 128 humus profiles sampled over the two study periods, 31 ECM species were recorded. The study demonstrated that the number of both non-vital tips and vital non-mycorrhized tips decreases with soil depth, from organic to mineral horizons, while the number of ectomycorrhizal tips mainly increases with soil depth. A preference was found of some ECM species and exploration types for specific organic or mineral soil layers and their features, especially moisture and available nitrogen. These results can help in understanding how the functional role of the single consortia and the ecological features determining this “adaptive diversity” in ectomycorrhizal communities could be of major importance to assess the resilience in forest soil ecosystems.     

Norway spruce emblings as cutting donors for tree breeding and production

In the Nordic countries, Norway spruce (Picea abies) is a major species in tree breeding. In order to facilitate breeding work and availability of highly bred forest regeneration material, the time required for breeding and implementation of results should be shortened. This could be done by accelerating production of clonal material for field testing, and possibly for planting stock, by combining production of rooted cuttings with somatic embryogenesis (SE). This would allow efficient production of numerous plants of the same genotype, with equal age and propagation history between genotypes. In the present work, we studied the rooting potential of cuttings from Norway spruce emblings. Altogether 36 clones from 12 families representing elite breeding materials and ornamental forms were examined under different rooting conditions (container type and rooting media) in 2015 and 2016. Our results show that Norway spruce emblings are good donors for cuttings. Best combination (peat–vermiculite mixture and Plantek 81f containers) resulted in 91% rooting, variation among the tested clones of elite breeding materials being 55–100% per treatment. The rooting variation between families is acceptable for breeding purposes. High rooting (87–96%) of ornamental forms indicates propagation potential with the combination of SE and rooted cuttings.     

Modeling individual tree height growth of Norway spruce and Scots pine from national forest inventory data in Norway

We developed individual tree height growth models for Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) in Norway based on national forest inventory data. Potential height growth is based on existing dominant height growth models and reduced due to competition by functions developed in this study. Three spatially explicit and two spatially non-explicit competition indices were tested. Distance effects and diameter ratio effects were estimated from the data simultaneously with parameters of the potential modifier functions. Large height measurement errors in the national forest inventory data caused large residual variation of the models. However, the effects of competition on height growth were significant and plausible. The potential modifier functions show that height growth of dominant trees is largely unaffected by competition. Only at higher levels of competition, height growth is reduced as a consequence of competition. However, Scots pine also reduced height growth at very low levels of competition. Distance effects in the spatially explicit competition indices indicated that the closest neighbors are most important for height growth. However, for Scots pine also competitors at larger distance affected height growth. The five competition indices tested in this study explained similar proportions of the variation in relative height growth. Given that unbiased predictions can only be expected for the same plot size, we recommend a spatially explicit index, which describes the distance function with a negative exponential, for use in growth simulators.     

Snow disturbances in secondary Norway spruce forests in Central Europe: Regression modeling and its implications for forest management

Snow is an important ecological factor limiting vegetation distribution, growth, and regeneration, and the importance of snow in the latitudes of Central Europe is expected to increase in the future. We assessed snow damage to secondary spruce stands (not of native provenance therefore not adapted to local conditions) in a mountainous region of 14500 ha in Central Europe (Moravian-Silesian Beskids, Czech Republic). We used neural networks-based regression modeling to study the relationship between stand and environmental parameters and four types of snow damage (top tree, crown, and stem breakage, and uprooting) that occurred during heavy snowfalls in winter 2005/2006 and 2009. Almost 40000 trees were sampled in 345 plots after each of these two events.The results suggest that parameters that can be controlled by forest management (mainly stand density and taper) were not closely associated with spruce forest resistance to snow damage. Investigated snow damage types were primarily related to the developmental stage of the stand, as indicated by stand volume, age, height, and diameter. Damage in 2009, which was caused by shorter-lasting and lower snow loads than the damage in 2005/2006, was also associated with elevation and snow depth. The response of snow damage to all stand development-related variables was clearly unimodal.We infer that forest management can reduce snow damage to secondary spruce forests in Central Europe only to a limited extent, especially under heavy snow loads. This conclusion is supported by the heavy snows that have frequently fallen on forests in Central Europe in the past and the projected increase in winter precipitation in mid- and northern latitudes; thereby increasing snow damage to forest in the future. Therefore, managers of such spruce forests should not specifically consider forest resistance to snow damage but should apply general practices that maintain forest health and productivity.     

Decomposing cones of Norway spruce (Picea abies (L.) H. Karst.): dry weight loss,chemical changes,and vertical transport in a Norwegian raw humus soil profile

Naturally shed cones of Norway spruce (Picea abies) were placed on the forest floor, partly on moss-covered ground, and partly on vegetation-free needle mats. Numbered sticks allowed individual recovery after 1, 3, 5, 7, 9, 11, and 13 years. During the first nine years, cones decomposed fastest in the moss site. After 13 years, dry weight loss was about 60% in both habitats, but cones still retained their shape. In the moss site, cones were then situated in the middle of the organic layer, with their underside about 3?cm above the bleached layer. Cones in the needle mat profile had sunk less. One cone with rather intact shape was found below a nylon strip laid out 28 years earlier. Compared to needle litter from Norway spruce and Scots pine, spruce cones had low concentrations of Ca, Mn, and Fe. The concentration of these elements, as well as N, Al, and S, was greatly increased during decomposition. Ca, Mn, Fe, and Al were immobilized for a long time, as their total amount per cone increased with time. Cones decomposed slower than needles, so their role in humus composition and carbon storage may be greater than indicated by their fraction of fresh litter.     

Using drill resistance to quantify the density in coarse woody debris of Norway spruce

To evaluate the mass of coarse woody debris (CWD), it is necessary to quantify its density. Drill resistance measurements are introduced as a approach to estimate the density of CWD in different stages of decay. Dead logs of Norway spruce [Picea abies (L.) Karst.] from a Central European mountainous site were used as a test system to compare the new method with conventional predictors of wood density such as fast quantitative field estimates (e.g., knife probe) and classification of decay classes based on a set of qualitative traits and quantitative estimates. The model containing only drill resistance as a predictor explained 65% of the variation in wood density and was markedly better than models containing one or more of several conventional predictors. However, we show that the relationship between drill resistance and gravimetric wood density relationship is sensitive to the decay status. Therefore, the best model combines drill resistance and decay class (adj. R2 = 0.732). An additional experiment showed that drill resistance is also sensitive to the moisture state (fresh vs. oven-dry) of the sample. The major potential of the method lies in its non-destructive nature which allows repeated sampling in long-term ecosystem studies or in protected areas where destructive sampling is prohibited. The limitations of the method are discussed and recommendations for applications are given.     

Effects of repeated urea doses on soil chemistry and nutrient pools in a Norway spruce stand

Pools of macro-nutrients in soil and vegetation were studied in an old fertilization experiment with a large previous input of N. Different doses of N, in the form of urea, had been added four times during a 20-year period. In total, between 480 and 2400 kg N ha⁻¹ had been given. The experiment was established in a relatively productive Norway spruce stand and the expectation was that the large N input would cause an accelerated leaching of N, especially nitrate, accompanied by soil acidification and losses of several nutrients. The aim was to test for possible residual effects. Thirteen years after the last N addition, samples from the aboveground part of trees, field layer, S-layer, humus layer and mineral soil (0–10 cm) were analyzed for concentrations of most major nutrients. Nutrient pools were calculated. In the humus layer, the concentration of N increased and the C/N-ratio decreased with increasing N dose. The calculated recovery of added N in soil including ground vegetation was complete for the lowest N dose, while it was 25–50% for higher doses. The amount of N retained was unaffected by the N dose. The amount of extractable P in the upper part of the mineral soil was negatively correlated with N dose, as was also the concentration of total P in the S-layer. Neither soil pH, nor concentrations or amounts of Ca, Mg and K were affected by the previous fertilization. The calculated total soil-plant pool was only influenced by N dose in the case of P, which was 20% lower at the highest N dose compared with unfertilized conditions. Despite the large extra N input, the nutritional changes in plants and soil of the actual study site seemed surprisingly small.     

Durability of thermally modified sapwood and heartwood of Scots pine and Norway spruce in the modified double layer test

In the present study, durability of untreated and thermally modified sapwood and heartwood of Scots pine and Norway spruce was examined using a modified double layer test. Base layer samples were partly on contact with ground where exposure conditions were harder than that in a double layer test above the ground. The base layer on ground contact gave results already after one year of exposure in Finnish climate, but the top layer of a double layer test element simulated more the situation of decking exposure.

Significant differences in durability and moisture content (MC) between the wood materials were detected after six years of exposure in the field. Thermally modified pine heartwood performed very well in all layers of the test element and only minor signs of decay were found in some of the base samples. Both sapwood and heartwood of thermally modified spruce were suffering only slight amounts of decay while thermally modified pine sapwood was slightly or moderately decayed. Untreated sapwood samples of pine and spruce were severely decayed or reached failure rating after six years in the field. Untreated heartwood samples performed clearly better. The highest MCs were measured from untreated and thermally modified pine samples. Thermal modification increased significantly the durability and decreased the MC values of all wood materials.     

Crown condition as a function of soil,site and tree characteristics

One of the aims of this work is to describe how the target variable “tree vitality” in terms of needle loss is affected by other explanatory variables. To describe such a relationship in a realistic way, we use generalized additive mixed models (GAMMs) which allow to take spatial correlation of the data into account and in addition allow the inclusion of explanatory variables as predictors with the possibility of having non-linear effects. The GAMMs are fitted in a Bayesian framework using Markov chain Monte Carlo techniques. Data are available for two years 1988 and 1994. We select a set of best explanatory variables from a large set of variables including tree-specific variables, such as species, age, nutrients in the needles and site-specific variables such as altitude, relief type, soil depth and content of different nutrients in the top soil. In the two models for 1988 and 1994, different sets of explanatory variables were selected as best predictors. In both models, the effects of explanatory variables allowed a plausible interpretation. For example, the site-specific variables such as relief and soil depth were significant predictors, since these factors determine how well water and nutrient supply is balanced at a specific site. The selected sets of explanatory variables differed between 1988 and 1994, giving an indication of a possible change in the main causes of forest deterioration between 1988 and 1994. From the set of nutrient variables measured in the soil and in the needles, in 1988 altitude a.s.l. and magnesium supply were among the explanatory variables, in 1994 a combination of Al in the soil and the N/K-ratio (in the needles) was selected in the model. In 1988 altitude a.s.l. was among the most important predictors in the model. This is in contrast to 1994 where altitude was not selected. This may have to do with the fact that in the early phase of forest health monitoring (1988) one of the main causes of forest deterioration was magnesium deficiency. Later on this may have changed to a combination of soil acidification and nitrogen eutrophication. Thus by using an adequate model such as the GAMM, sets of explanatory variables for needle loss may be identified. By fitting two GAMMs, with different sets of “best” predictors, at two time points 1988 and 1994, we can detect changes in these sets of “best” predictors over time. This allows us to use the monitoring data with the tree vitality indicator crown condition/needle loss as a tool for forest health management, which may involve decisions about concrete counter measures like e.g. forest liming.     

Applicability of non-destructive substitutes for leaf area in different stands of Norway spruce (Picea abies L. Karst.) focusing on traditional forest crown measures

Since individual tree leaf area is an important measure for productivity as well as for site occupancy, it is of high interest in many studies about forest growth. The exact determination of leaf area is nearly impossible. Thus, a common way to get information about leaf area is to use substitutes. These substitutes are often variables which are collected in a destructive way which is not feasible for long term studies. Therefore, this study aimed at testing the applicability of using substitutes for leaf area which could be collected in a non-destructive way, namely crown surface area and crown projection area. In 8 stands of Norway spruce (Picea abies L. Karst.), divided into three age classes and two thinning treatments, a total of 156 trees were felled in order to test the relationship between leaf area and crown surface area and crown projection area, respectively. Individual tree leaf area of the felled sample trees was estimated by 3P-branch sampling with an accuracy of ±10%. Crown projection area and crown surface area were compared with other, more commonly used, but destructive predictors of leaf area, namely sapwood area at different heights on the bole. Our investigations confirmed findings of several studies that sapwood area is the most precise measure for leaf area because of the high correlation between sapwood area and the leaf area. But behind sapwood area at crown base and sapwood area at three tenth of the tree height the predictive ability of crown surface area was ranked third and even better than that of sapwood area at breast height (R² = 0.656 compared with 0.600). Within the stands leaf area is proportional to crown surface area. Using the pooled data of all stands a mixed model approach showed that additionally to crown surface area dominant height and diameter at breast height (dbh) improved the leaf area estimates. Thus, taking dominant height and dbh into account, crown surface area can be recommended for estimating the leaf area of individual trees. The resulting model was in line with many other findings on the leaf area and leaf mass relationships with crown size. From the additional influence of dominant height and dbh in the leaf area model we conclude that the used crown model could be improved by estimating the position of the maximum crown width and the crown width at the base of the crown depending on these two variables.     

Organic matter decomposition in an acidic forest soil in Denmark as measured by the cotton strip assay

Decomposition of organic matter in forest soils controls mineralization of nutrients. The decomposition is to a large extent controlled by climatic and soil conditions. Four different soil treatments of water and nutrients were applied to a Norway spruce (Picea abies) stand in Denmark, and the relative changes in decomposition rate were quantified by the cotton strip assay. Additions of water and nutrients to the soil increased the decomposition of the cotton strips significantly, especially in the deeper soil layers. Reduced water addition during the summer, thereby creating ‘summer drought’, caused no changes in the decomposition compared to an untreated control situation, but the spatial variability in the decomposition within the treatment plot was strongly increased. The decomposition of the cotton strips decreased linearly with the soil depth (1st order).     

Compositional changes of forest-floor vegetation in young stands of Norway spruce as an effect of repeated fertilisation

Forestry practices that aim to increase biomass production may mitigate climate change through increased carbon sequestration and the potential of substituting fossil fuels with renewable biofuels. Fertilising young stands of Norway spruce in Sweden have shown to increase tree growth by more than 200%. Fertilisation, however, also has other effects on forest ecosystems. Here, we studied the response of the species composition of forest-floor vegetation to three different frequencies of fertilisation in young stands of Norway spruce. Fertiliser was applied every year, every second year or every third year. The total amount of N ranged from 425 kg ha⁻¹ to 625 kg ha⁻¹, in combination with P, K, Ca, Mg, S, Mn, Zi, B and Cu. The largest effects of the fertilisation were found among bryophytes and lichens, which lost substantial cover. Unexpectedly, Deschampsia flexuosa, commonly known to be favoured by fertilisation, was negatively affected. Species that increased in frequency were Oxalis acetosella, Brachythecium sp. and Plagiothecium sp. Decreased availability of light, as an indirect effect of fertilisation through increased tree canopy cover, was found to be the most important factor behind the change in species composition of vascular plants. The total cover of bryophytes, however, did not show any significant response to the changes in canopy cover, indicating that the effects seen in this group may be a result of more direct effects of the fertiliser. Few significant differences were found between the two most intensive fertilisation frequencies, although fertilisation every third year was often distinguished from both the control and the other fertilised treatments. Even though the effects at the stand level were substantial, the effects on biodiversity and function of ecosystems on a landscape or regional level need further investigation.     

Effect of removal of logging residue on nutrient leaching and nutrient pools in the soil after clearcutting in a Norway spruce stand

Following clearcutting applying the conventional stem-only harvesting method in a Norway spruce (Picea abies (L.) Karst.) stand and different levels of removal of logging residue, the nutrient fluxes from the heaps of logging residue and from the O horizon were monitored over four growing seasons and the soil nutrient pools were determined. Three levels of removal of logging residue were carried out using (i) conventional stem-only harvesting (no residues removed); (ii) residues removed; and (iii) removal of branches (foliage left on site). The heaps of logging residue were a minor source of inorganic N entering the soil in the water percolating through the heaps, but they were a significant source of organic N, P, Ca, Mg, and especially K. Nutrient fluxes from the O horizon were in general greater under the heaps of logging residue as compared to soils without overlying logging residue. The leaching of inorganic N from the O horizon under the heaps of logging residue resulted in a net loss of these compounds, while the O horizon without overlying logging residue gained N. The removal of logging residue significantly decreased the extractable K pools in the soil while it or conversely, the presence of residue heaps had no significant effect on the pools of organic matter and the pools of N, P, Ca, and Mg in the O horizon and in the 0–10 cm soil layer. The results show that the short-term effects of logging residue on nutrient dynamics in the soil can be complex and difficult to interpret in terms of site productivity as there are changes in the nutrient fluxes, which imply the opposite effects on site productivity. However, the results do indicate that, in the short-term, the removal of logging residue does not impair pools of N in the soil nor site productivity on sites where the availability of N limits productivity.     

Reforestation with native tree species using site preparation techniques for the restoration of woodlands degraded by air pollution in the Erzgebirge,Germany

Reforestation of woodlands with native species in the Erzgebirge, where large-scale deforestation has been caused by severe air pollution, was investigated. In an experiment, three tree species (Norway spruce, rowanberry, and birch) were studied with regard to fencing (no protection versus protection against game browsing) and site preparation techniques with eight levels: a control and seven amelioration techniques (soil cultivation, weed control, liming, and their combinations). Four criteria, survival, growth, production, and vitality, were used in assessing the success of the plantings. Repeated-measures analyses were performed to examine the development of the young plantations over an observation period of 7 years, and to determine whether this development was dependent on the experimental factors. Spruce showed high survival rates, reasonable growth and production, and good individual vitality. With the exception of survival, birch responded similarly to spruce. The low survival rate of birch resulted from the initial small size of the seedlings. By providing appropriate seedling material, both species could be used for reforestation. Seedlings of rowanberry were unsuccessful due to severe mouse damage, confirming the necessity of rodent control when planting cleared areas with this species is undertaken. Game browsing was not significant. None of the amelioration techniques had a major effect on the studied criteria of the three tree species. Slightly positive effects occurred only when spruce was limed and when birch received soil cultivation. Weed control by scything showed undesirable results, particularly for birch. For reasons of financial thriftiness, none of the amelioration techniques could be justified for the study area.     

Effect of soil chemical properties on growth,foliation and nutrition of Norway spruce stand affected by yellowing in the Bohemian Forest Mts., Czech Republic

In 1998, a fertilizer experiment aiming to investigate the effects of slow-release N, P, K and Mg fertilizer (SILVAMIX Mg NPK^®) on a 60-year-old spruce stand with symptoms of yellowing was established. In this paper, trees were selected to investigate the relation between annual diameter increment, yellowing, foliation, needle and soil chemical properties: ten from the fertilized treatment (F), ten green trees from the control (CG) and ten yellow trees from the control (CY). CG and CY trees were growing in close proximity at a distance of only several meters apart under the same soil conditions. In treatment F, increased annual diameter increment, improved foliation, needle Mg concentration, plant-available Mg and P concentrations in the soil and absence of yellow trees were recorded 7 years after a single application of the fertilizer. During the last 15 years, annual growth increment and foliation of CY trees have continuously decreased while relatively stable values were recorded for CG trees and increased for F trees. In 2006, CG and CY trees differed significantly in Mg concentration in needles, foliation, yellowing and annual diameter increment. Although differences in soil chemical properties between CG and CY treatments were not significant, lower concentrations of plant-available Mg²⁺ and higher concentrations of H⁺ and Al³⁺ were found in soils under CY trees. There was a negative correlation between soil concentration of Mg and yellowing, but this correlation was relatively weak, indicating that there is no simple relation between soil and needle concentrations of Mg. In the investigated locality, the “new type” of yellow tree decline has been a long-term gradual process.     

Variation in the chemical composition of green leaves and leaf litters from three deciduous tree species growing on different soil types

The chemical composition of green leaves and leaf litters of sweet chestnut (Castanea sativa), oak (Quercus robur) and beech (Fagus sylvatica) were determined for 26 sites grouped into high fertility (HF) and low fertility (LF) soils according to base saturation and N-mineralization potentials. Measurements were made of total carbon, acid detergent fibre (ADF), Klason lignin, holo-cellulose, sugar constituents of hemicellulose and phenylpropanoid derivatives of lignin, and nutrient concentrations (N, Ca, P, Mg, K and Mn). Leaf and litter constituents varied within and between species according to soil groups, but beech showed contrasting responses to oak and chestnut. Beech leaves had lower ADF, lignin and cellulose on HF soils than LF soils, whereas oak and chestnut leaves had higher ADF, lignin and cellulose on HF than the LF soils. Conversely, the same constituents in beech leaf litter were higher on HF soils than LF soils, but lower in oak and chestnut leaf litter on HF soils than LF soils. The phenylpropanoid derivatives of lignin and sugar constituents of hemicellulose also showed similar variations in relation to soil groups with contrasting patterns for in leaves and litters. Re-absorption of N from leaves before litter fall was negatively correlated with soil N mineralization potential for beech (highest on LF soils) but showed an unexpected, positive relationship for oak and chestnut (highest on HF soils). These intra-specific differences of leaf and litter chemistry in relation to soil fertility status are unprecedented and largely unexplained. The observed patterns reflect phenotypic responses to soil type that result in continuum of litter quality, within and between tree species, that have been shown in related studies to significantly influence litter decomposition rates.     

Concept and feasibility study for the integrated evaluation of environmental monitoring data in forests

In the 1970s unexpected forest damages, called “new type of forest damage” or “forest decline”, were observed in Germany and other European countries. The Federal Republic of Germany and the German Federal States implemented a forest monitoring system in the early 1980s, in order to monitor and assess the forest condition. Due to the growing public awareness of possible adverse effects of air pollution on forests, in 1985 the ICP Forests was launched under the convention on long-range transboundary air pollution (CLRTAP) of the United Nations Economic Commission for Europe (UN-ECE). The German experience in forest monitoring was a base for the implementation of the European monitoring system. In 2001 the interdisciplinary case study “concept and feasibility study for the integrated evaluation of environmental monitoring data in forests”, funded by the German Federal Ministry of Education and Research, concentrated on in-depths evaluations of the German data of forest monitoring. The objectives of the study were: (a) a reliable assessment of the vitality and functioning of forest ecosystems, (b) the identification and quantification of factors influencing forest vitality, and (c) the clarification of cause-effect-relationships leading to leaf/needle loss. For these purposes additional data from external sources were acquired: climate and deposition, for selected level I plots tree growth data, as well as data on groundwater quality. The results show that in particular time series analysis (crown condition, tree growth, and tree ring analysis), in combination with climate and deposition are valuable and informative, as well as integrated evaluation of soil, tree nutrition and crown condition data. Methods to combine information from the extensive and the intensive monitoring, and to transfer process information to the large scale should be elaborated in future.