Long-term nutritional trends of conifer stands in Europe: results from the RECOGNITION project

Since 1999–2002 the European Commission has funded and the European Forest Institute has coordinated the interdisciplinary RECOGNITION project aiming at elucidating the causes of the growth acceleration which has been observed in some forest tree species in several parts of Europe. Within this project, it was our task to identify and quantify long-term changes in the nutritional status of representative forest stands that potentially could explain this growth increase, using available long-term series of foliar analyses. An inquiry among 25 forest research institutions in Europe resulted in 28 Scots pine (Pinus sylvestris L.) and 21 Norway spruce (Picea abies [L.] Karst.) stands for this historical development investigation (HDI). The stands generally are control plots of fertilization experiments and are located mainly in Central Europe and in Scandinavia. The monitoring periods vary from 15–40 years. The foliar data were given to us by our partner organisations, subjected to rigorous tests for plausibility and comprehensively evaluated using mainly single linear regression approaches. Most Scots pine stands under study in Central Europe, which grow predominantly on naturally poor or devastated soils in regions with relatively high atmospheric N deposition, suffered from N deficiency at the start of the monitoring period, but exhibited a considerable improvement in N nutrition over the past decades. The increase in N levels was usually associated with an increase in the ratios of N and P, K, Ca, and Mg, although critical values of these ratios are far from being reached. For the majority of the investigated Scots pine stands in Central Europe, growth acceleration due to a better N supply is highly probable. In contrast, N seems to be far less important as a potential driving factor for long-term growth changes in Scots pine in Scandinavia. Norway spruce stands examined in Central Europe were, in general, initially well supplied with N and characterised by a decreasing trend in foliar N levels although they received considerable N deposition from the atmosphere. This decrease in concentrations of N and other macronutrients is believed to be mainly a dilution effect as indicated by a simultaneous increase in needle weights. Spruce in Finland also turned out to be adequately provided with N at most sample sites, and no general trend of improving N nutrition was detected.     

Impacts of changing climate on the productivity of Norway spruce dominant stands with a mixture of Scots pine and birch in relation to water availability in southern and northern Finland

A process-based ecosystem model was used to assess the impacts of changing climate on net photosynthesis and total stem wood growth in relation to water availability in two unmanaged Norway spruce (Picea abies) dominant stands with a mixture of Scots pine (Pinus sylvestris) and birch (Betula sp.). The mixed stands were grown over a 100-year rotation (2000-99) in southern and northern Finland with initial species shares of 50, 25 and 25% for Norway spruce, Scots pine and birch, respectively. In addition, pure Norway spruce, Scots pine and birch stands were used as a comparison to identify whether species' response is different in mixed and pure stands. Soil type and moisture conditions (moderate drought) were expected to be the same at the beginning of the simulations irrespective of site location. Regardless of tree species, both annual net canopy photosynthesis (P(nc)) and total stem wood growth (V(s)) were, on average, lower on the southern site under the changing climate compared with the current climate (difference increasing toward the end of the rotation); the opposite was the case for the northern site. Regarding the stand water budget, evapotranspiration (E(T)) was higher under the changing climate regardless of site location. Transpiration and evaporation from the canopy affected water depletion the most. Norway spruce and birch accounted for most of the water depletion in mixed stands on both sites regardless of climatic condition. The annual soil water deficit (W(d)) was higher on the southern site under the changing climate. On the northern site, the situation was the opposite. According to our results, the growth of pure Norway spruce stands in southern Finland could be even lower than the growth of Norway spruce in mixed stands under the changing climate. The opposite was found for pure Scots pine and birch stands due to lower water depletion. This indicates that in the future the management should be properly adapted to climate change in order to sustain the productivity of mixed stands dominated by Norway spruce.     

Growth response of coniferous stands to whole‐tree harvesting in early thinnings

A series of 15 field experiments was established to quantify the growth response of first‐thinning stands of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst) to whole‐tree harvesting and to estimate the need for nutrient compensation. The experiments were undertaken in Finland, Norway and Sweden and represent a wide range of site conditions. The site index (H ₁₀₀) of Scots pine stands varied from 19 to 29 m, and that of Norway spruce stands from 28 to 36 m. Total amounts of biomass and nutrients removed were calculated based on data obtained from felled sample trees. During the first 5‐yr period the growth response to the removal of logging residues varied considerably in both pine and spruce stands. Regression analyses did not reveal any functions that explained the variation in results satisfactorily. In cases where whole‐tree harvesting influenced tree growth negatively, this effect was counteracted by compensatory fertilization. It was concluded that to determine the response of remaining trees to harvesting intensity reliably, the post‐harvest period analysed must be longer than 5 yrs.     

Fine root morphological adaptations in Scots pine, Norway spruce and silver birch along a latitudinal gradient in boreal forests

Variability in short root morphology of the three main tree species of Europe's boreal forest (Norway spruce (Picea abies L. Karst.), Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth)) was investigated in four stands along a latitudinal gradient from northern Finland to southern Estonia. Silver birch and Scots pine were present in three stands and Norway spruce was present in all stands. For three fertile Norway spruce stands, fine root biomass and number of root tips per stand area or unit basal area were assessed from north to south. Principal component analysis indicated that short root morphology was significantly affected by tree species and site, which together explained 34.7% of the total variability. The range of variation in mean specific root area (SRA) was 51-74, 60-70 and 84-124 m(2) kg(-1) for Norway spruce, Scots pine and silver birch, respectively, and the corresponding ranges for specific root length were 37-47, 40-48 and 87-97 m g(-1). The range of variation in root tissue density of Norway spruce, Scots pine and silver birch was 113-182, 127-158 and 81-156 kg m(-3), respectively. Sensitivity of short root morphology to site conditions decreased in the order: Norway spruce > silver birch > Scots pine. Short root SRA increased with site fertility in all species. In Norway spruce, fine root biomass and number of root tips per m(2) decreased from north to south. The differences in morphological parameters among sites were significant but smaller than the site differences in fine root biomass and number of root tips.     

Dynamics and potentials of carbon sequestration in managed stands and wood products in Finland under changing climatic conditions

Carbon (C) sequestration was studied in managed boreal forest stands and in wood products under current and changing climate in Finland. The C flows were simulated with a gap-type forest model interfaced with a wood product model. Sites in the simulations represented medium fertile southern and northern Finland sites, and stands were pure Scots pine and Norway spruce stands or mixtures of silver and pubescent birch.

Changing climate increased C sequestration clearly in northern Finland, but in southern Finland sequestration even decreased. Temperature is currently the major factor limiting tree growth in northern Finland. In southern Finland, the total average C balance over the 150 year period increased slightly in Scots pine stands and wood products, from 0.78 Mg C ha⁻¹ per year to 0.84 Mg C ha⁻¹ per year, while in birch stands and wood products the increase was larger, from 0.64 Mg C ha⁻¹ per year to 0.92 Mg C ha⁻¹ per year. In Norway spruce stands and wood products, the total average balance decreased substantially, from 0.96 Mg C ha⁻¹ per year to 0.32 Mg C ha⁻¹ per year. In northern Finland, the total average C balance of the 150 year period increased under changing climate, regardless of tree species: in Scots pine stands and wood products from 1.10 Mg C ha⁻¹ per year to 1.42 Mg C ha⁻¹ per year, in Norway spruce stands and wood products from 0.69 Mg C ha⁻¹ per year to 0.99 Mg C ha⁻¹ per year, and in birch stands and wood products from 0.43 Mg C ha⁻¹ per year to 0.60 Mg C ha⁻¹ per year.

C sequestration in unmanaged stands was larger than in managed systems, regardless of climate. However, wood products should be included in C sequestration assessments since 12–55% of the total 45–214 Mg C ha⁻¹ after 150 years' simulation was in products, depending on tree species, climate and location. The largest C flow from managed system back into the atmosphere was from litter, 36–47% of the total flow, from vegetation 22–32%, from soil organic matter 25–30%. Emissions from the production process and burning of discarded products were 1–6% of the total flow, and emissions from landfills less than 1%.     

Irradiance in young stands of picea abies (L.) karst. and pinus sylvestris L. and the possibilities to prevent suckers of broad‐leaved trees

Incoming shortwave global radiation (Q _g) and photosynthetically active radiation (PAR; Q _pa as a fraction of full daylight, relative irradiance (%Q), were measured at the same time in young stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.). Measurements were made on three levels above ground: 20 cm above ground and 50 and 15 % of stand height. Stands of three heights (75, 150 and 300 cm) were studied during two months. The stands were created by arranging young trees cut from natural stands, in nine quadratic spacings: 0.1, 0.2, 0.3, 0.4, 0.5, 0.7, 1.0, 1.4 and 2.0 m. The leaf area index (L) was estimated. Differences in %Q‐values for Q _pa and Q_g in the same species and at the same stand height and level of light measurements above ground were significant only in 150 and 300 cm stands. In 75 cm high stands of Scots pine, the %Q was <60% at spacings <0.2 m and in 75 cm high stands of Norway spruce, the %Q was <60% at spacings <0.3 m. Only at 0.1x0.1–0.3x0.3 m, %Q was <20% in Norway spruce stands. In 150 cm stands %Q<20% was measured at spacings <0.7 m in Norway spruce and <0.5 in Scots pine. In 300 cm stands of Norway spruce it was measured up to 1.2 and in Scots pine <1.1 m. Light extinction coefficients, K and K_g for spruce and pine stands were 0.17–0.40 and 0.16–0.31 respectively. Some practical implications of the study are presented. Lack of light (%Q<10%) as a single factor of seriously suppressed growth and development of broad‐leaved plants and suckers by competition in young stands of spruce and pine only occur in dense stands 0.3x0.3 m‐1.1xl.l m (8000–100000 stems/hectare). Competition by light on a regenerated area generally occurs in the level of 50% of tree height (150–300 cm) and higher due to the rapid growth of broad‐leaved trees (sprouts) compared with planted conifers but the light intensity at these levels %Q>10%.     

Risk of Snow Damage in Unmanaged and Managed Stands of Scots Pine,Norway Spruce and Birch

The aim of this study was to assess the risk of snow damage to trees in unmanaged and managed stands of Scots pine (Pinus sylvestris L.), Norway spruce [Picea abies (L.) Karst.] and birch (Betula spp.) over a rotation. The risk assessment was based on the prediction of critical snow loads in interaction with the windspeed at which trees can be expected to break or be uprooted, and on the frequency of long-term extremes of precipitation and of suitable temperature conditions for the accumulation of snow on the tree crowns. The Scots pine stands were found to be more susceptible to snow damage than the others, and an unmanaged stand of Scots pine to be more susceptible to break and uproot than a managed one. Correspondingly, an unmanaged stand of Norway spruce was more susceptible to stem breakage than a managed one, but less susceptible to uprooting. Neither unmanaged nor managed birch stands were likely to suffer any kind of snow damage. The susceptibility of unmanaged stands is caused by low tapering of the trees. Based on the frequency of long-term extremes in precipitation at the temperatures needed for snow accumulation on tree crowns, critical snow loads of 10-19, 20-29 and 30-39 kg m^-2 occurred 19.3, 3.3 and 1.3 times in a decade in southern Finland. Critical snow loads of 10-19, 20-29, 30-39 and 60-69 kg m^-2 occurred in northern Finland 17.0, 6.3, 1.7 and 0.3 times in a decade.     

Functions for Biomass Estimation of Young Pinus sylvestris,Picea abies and Betula spp. from Stands in Northern Sweden with High Stand Densities

New silvicultural regimes with high within-stand competition require new functions for estimation of standing stock and growth of biomass components, since the allometry of trees is changed by light competition. This paper presents functions for estimation of the aboveground biomass dry weights for stem wood, stem bark, branches and leaves of young (diameter at breast height <10 cm) Scots pine (Pinus sylvestris L.), Norway spruce [Picea abies (L.) Karst.] and birch (Betula pendula Roth. and Betula pubescens Ehrh.) trees growing in dense mixed stands. The functions were derived from a sample consisting of 84 Scots pine, 43 Norway spruce and 66 birch trees from six stands in northern Sweden with high stand densities (>10000 st ha^-1). The logarithmically transformed power function displayed a good ability to stabilize the variance of dry weights and showed a good fit to the material (0.37< R ² <0.99). A comparison with the most commonly used biomass functions in Sweden today showed that they overestimated the weight of stem wood and branches, while the weight of foliage was underestimated. The nature of these discrepancies suggested that the precision of biomass estimations might also be improved for young trees at wider spacing.     

Morphological and Ecological Variation of Gremmeniella abietina var. abietina in Pinus sylvestris,Pinus contorta and Picea abies Sapling Stands in Northern Finland and the Kola Peninsula

The morphological and ecological variation of two types of Gremmeniella abietina var. abietina causing scleroderris canker on conifers was investigated in Pinus spp. and Picea sp. sapling stands in northern Finland and the Kola Peninsula. Small - tree type (STT or B type) of G. abietina was detected alone in 13 Scots pine, three lodgepole pine and two Norway spruce sapling stands out of 26 stands investigated, both STT and large - tree type (LTT or A type) were observed in six Scots pine stands, and LTT was detected alone in two Scots pine stands. For the first time, G. abietina was found to injure Norway spruce saplings in a respective plantation in northern Fennoscandia. STT isolates produced statistically significantly more conidia in vitro than LTT isolates. Morphological variation in conidia septation revealed that STT produced conidia with more than five septa more frequently than did LTT. There was a greater range in variation in septation in STT than in LTT, with overlapping between the types. Isolates of both types were equally associated with cankers, coloured wood, pycnidia or apothecia in the infected saplings.     

From planting to 26 years of age – actual growth and estimated volume scenarios for spruces and pines

The objective of this study was to compare the survival and volume of conifer stands at 26 years of age with their status at planting. Survival, growth and damage were studied in eight clear felled stands regenerated in 1972. Five of the areas were planted with Norway spruce (Picea abies (L.) Karst.) and three with Scots pine (Pinus sylvestris L.). The plantings were examined in 1972 and 1974. In 1974, the number of living undamaged planted seedlings was low (10–15%). However, the number of undamaged seedlings was supplemented by naturally regenerated conifer and birch seedlings. The total number of undamaged seedling in 1974 was equivalent to 20–30% of the number of seedlings planted. In 1998, the main species in three stands had changed from Norway spruce to Scots pine, and in one stand from Norway spruce to birches. Actual volume in 1998 for the stands was compared to stand volume generate according to five scenarios based on recommended and actual seedling number in 1972 and 1974. The actual volume was 64% of that expected if the recommended number of trees had been planted. Naturally regenerated Scots pine and Norway spruce increased stand density in 1998. The actual volume was 37% higher than the average volume in the surrounding county. On average, 36% of the trees were damaged. More than 50% of the total damage was caused by moose (Alces alces L.). For Scots pine, moose or other browsing animals damaged 30% of the trees. The results of this study indicate that the 1998 volume was higher than expected, considering the low number of undamaged seedlings in 1974. This was mainly due to the large amount of naturally regenerated plants. In addition, the results indicate that the volume could have been higher if the initial conditions had been better. Despite the low number of undamaged seedlings in 1974, seven of the eight studied stands produced a higher volume than the average stand for the region. In practise, high numbers of seedlings should be planted on scarified areas. In most cases there will be a supply of naturally regenerated seedlings.     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

The spreading of the S type of Heterobasidion annosum from Norway spruce stumps to the subsequent tree stand

The occurrence of Heterobasidion annosum in stumps and growing trees was investigated on 15 forest sites in southern Finland where the previous tree stand had been Norway spruce (Picea abies) infected by H. annosum, and the present stand was either Scots pine (Pinus sylvestris), lodgepole pine (Pinus contorta), Siberian larch (Larix siberica), silver birch (Betula pendula) or Norway spruce 8–53 years old. Out of 712 spruce stumps investigated of the previous tree stand, 26.3% were infected by the S group and 0.3% by the P group of H. annosum. The fungus was alive and the fruit bodies were active even in stumps cut 46 years ago. In the subsequent stand, the proportion of trees with root rot increased in spruce stands and decreased in stands of other tree species. On average, one S type genet spreading from an old spruce stump had infected 3.0 trees in the following spruce stand, 0.5 trees in lodgepole pine, 0.3 trees in Siberian larch, 0.05 trees in Scots pine and 0.03 trees in silver birch stand. Although silver birch generally was highly resistant to the S type of H. annosum, infected trees were found on one site that was planted with birch of a very northern provenance.     

Stem Form and Taper Changes After Thinning and Nitrogen Fertilization in Picea abies and Pinus sylvestris Stands

Stem form and taper changes after thinning and thinning combined with N fertilization were studied in 23 Norway spruce [Picea abies (L.) Karst.] and 46 Scots pine (Pinus sylvestris L.) stands in Sweden observed for 5-28 yrs. Average taper was calculated for the stem section 1.3-6.0 m above ground level and as the ratio between the diameter at breast height and total tree height for periods of 5-10 yrs. At the start of the experiment, before the first thinning, the dominant height was 12-15 m. Once 65% of the basal area had been removed in a single thinning from below, the remaining trees showed a strong increase in taper compared with trees in unthinned stands. Stems in stands treated with heavily recurrent thinnings from below also increased in taper, although the increases were not always statistically significant. Trees in thinned, N-fertilized Scots pine stands in middle and northern Sweden developed a more pronounced taper compared with stems in equally thinned, unfertilized stands.     

Carbon and nitrogen dynamics in topsoils along forest conversion sequences in the Ore Mountains and the Saxonian lowland, Germany

Carbon and nitrogen stocks and their medium-term and readily decomposable fractions in topsoils were compared in relation to soil microbial biomass and activity along sequences from coniferous to deciduous stands. The study was carried out in the Ore Mountains and the Saxonian lowland, representing two typical natural regions in Saxony, Germany. In accordance with current forest conversion practices, the investigation sites represent different stands: mature conifer stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) (type A); Norway Scots spruce and pine with advanced plantings of European beech (Fagus sylvatica L.) or European beech/Common oak (Quercus petreae Liebl.) (type B); and mature deciduous stands of European beech and European beech/Common oak (type C). The investigated forest sites can be grouped into three silvicultural situations according to the development from coniferous stands to advanced plantings and finally mature deciduous forests (chronosequence A–B–C). The organic layer (L, F and H horizons) and uppermost mineral soil (0–10 cm) were analysed for potential C mineralisation, microbial biomass, concentrations of total C and N (TOC and TN) and for medium-term and readily decomposable C and N fractions, obtained by hot- and cold-water extraction respectively. The results showed an increase in organic layer thickness and mass as well as TOC and TN stocks along the forest sequences in the lowland. Yet, underplanted sites with two storeys revealed higher organic layer mass as well as TOC and TN stocks as compared to coniferous and deciduous stands. Stocks of hot- and cold-water-extractable C and N in relation to microbial biomass and its activity revealed a high turnover activity in deeper organic horizons of deciduous forests compared to coniferous stands. The stand-specific differentiation is discussed in relation to microbial biomass, litter quantity and quality and forest structure, but also with respect to the site-specific climatic factors and water budget as well as liming and fly-ash impacts. Results indicate higher dynamics in deciduous stands in the lowland especially during the initial turnover phase. The elevated microbial activity in deeper organic horizons of deciduous litter-influenced sites in spring is discussed as a specific indicator for long-term C sequestration potential as besides C mineralisation organic compounds are humified and thus, can be stored in the organic layer or in deeper soil horizons. Due to liming activities, stand-specific effects on organic matter turnover dynamics have evened out today in the Ore mountain region, but will presumably occur again once base saturation decreases. Here, the stand-specific effect on microbial biomass can currently be seen again as C_mic in the L horizon increased from spruce to beech. Our study sites in the lowland revealed no significant fly-ash impact. Differences between sites were evaluated by calculating the discriminance function. TOC and TN as well as medium-term degradable C and N were defined in this study as indicators for turnover dynamics along forest conversion sites.     

The chemical composition of needle and leaf litter from Scots pine, Norway spruce and white birch in Scandinavian forests

Needle litter from 14 stands of Scots pine (Pinus silvestris,L.), 13 stands of Norway spruce (Picea abies (L.) Karst.) andleaf litter from three stands of white birch (Betula pubescensEhrh.) were analysed for chemical composition. The concentrationsof the elements N, P, K, Ca, Mg and Mn as well as solid organiccomponents (lignin, cellulose and hemicelluloses) and solubleswere determined. When the average chemical compositions werecompared the Scots pine needle litter was clearly the most nutrient-poorlitter type. Of the solid organic-chemical components the ligninfraction dominated in the spruce and birch litter whereas thecellulose dominated in the pine needle litter. When Norway spruce and Scots pine were growing in adjacent standson soils with the same bedrock origin the spruce litter hadsignificantly higher concentrations of nutrients (N, P, K, Ca,Mg, Mn) than the pine needle litter. At sites where Norway spruceand white birch were growing in adjacent stands, the birch leaflitter had generally higher concentrations of nutrients. However, significant or nearly significant differences were onlyobtained for Mg (P = 0.002), K (P = 0.056) and N (P = 0.087),probably due to the few replicates of stands compared. Concerningorganic chemical components, the spruce needle litter had significantlyhigher concentrations of lignin and mannan than all the otherlitters and lower levels of ethanol-soluble substances, celluloseand galactan than the pine needle litter. Further, it had lowerconcentrations of water solubles, rhamnan and xylan than thebirch litter. No relationships were established between the nutrient statusof the conifer litters and the site index H100 (the dominantheight of the trees at a reference age of 100 years) of thestands. Concentrations of solid carbohydrates in the litterswere, however, positively correlated with site index (P <0.001). Further, the concentration of nitrogen in the pine needlelitter was negatively correlated with the latitude of the sites(P < 0.01). The influence of litter chemistry on the decompositionof litter and nutrient cycling of forests is discussed.     

Infection experiments with Gremmeniella abietina on seedlings of Norway spruce and Scots pine

Conidia of Gremmeniella abietina infected and caused disease symptoms in annual shoots of both Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings. In Norway spruce shoots the infection remained largely latent, with only a few seedlings showing symptoms. Mycelial growth inside the shoots was faster in Scots pine than in Norway spruce and was favoured by low temperature in both hosts. The shoots of Norway spruce seedlings had higher endophyte populations than those of Scots pine, and the populations were decreased by low temperatures. Reductions in the normal epiphytic or endophytic flora by acid mist treatments seemed to favour the development of G. abietina.     

Effect of uninucleate Rhizoctonia on Scots pine and Norway spruce seedlings

One‐year‐old container‐grown seedlings were planted in spring on clear cut areas: the Norway spruce (Picea abies) on a moist upland site (Myrtillus‐type) and Scots pine (Pinus sylvestris) on a dryish upland site (Vaccinium‐type). While still in the nursery, half of the seedlings of each species had been inoculated during the previous summer, with a uninucleate Rhizoctonia sp., a root dieback fungus. At outplanting all the seedlings appeared healthy and had a normal apical bud, although the height of the inoculated seedlings was less than that of the uninoculated control seedlings. At the end of the first growing season after planting, the mortality of inoculated Scots pine and Norway spruce seedlings was 25 and 69%, respectively. After two growing seasons the mortality of inoculated seedlings had increased to 38% for Scots pine and 93% for Norway spruce. The mortality of control seedlings after two growing seasons in the forest was 2% for Scots pine and 13% for Norway spruce. After outplanting the annual growth of inoculated seedlings was poor compared with the growth of control seedlings. These results show that, although Rhizoctonia‐affected seedlings are alive and green in the nursery, the disease subsequently affects both their survival and growth in the forest.     

Survival and Early Development of Lodgepole Pine

The effect of site fertility, spacing and mode of regeneration on the survival and stand development of lodgepole pine, Scots pine and Norway spruce was studied in a series of experiments comprising 22 study areas in Finland. After 13-14 yrs, lodgepole pine had a mean survival of 68% in planted and 61% in seeded plots, while Norway spruce had the highest (92%) and Scots pine the second highest survival (82%). The survival of planted lodgepole pine was better the wider the spacing. Best survival was achieved on subdry and dry sites, both with planting and with direct seeding. Dominant height was not affected by spacing, but both basal area and volume at the age of 13-14 yrs were significantly higher the denser the spacing. The average difference in the value of the estimated site index H ⁵⁰ was slightly under 3 m for the superiority of lodgepole compared with Scots pine. Seeding resulted in site indices almost as high as those of lodgepole planting.     

Height growth of planted conifer seedlings in relation to solar radiation and position in Scots pine shelterwood

Seedlings of different provenances of Scots pine (Pinus sylvestris L.), lodgepole pine (Pinus contorta Dougl., var. latifolia Engelm.) and Norway spruce (Picea abies (L.) Karst.) were planted in three Scots pine shelterwoods (125, 65 and 43 stems ha⁻¹) and a clear-cut, all in northern Sweden. The sites were mounded and planting took place during 2 consecutive years (1988 and 1989). The solar radiation experienced by the individual seedlings was determined using a simulation model. Height development of the seedlings was examined during their first 6 years after planting. During the final 3 years of the study, height growth of Norway spruce was relatively poor, both in the shelterwoods and the clear-cut area. Height growth of lodgepole pine was significantly greater than that of Scots pine, both in the shelterwoods and the clear-cut. In contrast to Norway spruce, Scots pine and lodgepole pine displayed significantly greater height growth in the clear-cut than in the shelterwoods. For all three species in the shelterwoods, regression analyses showed that height growth was more strongly correlated with the distance to the nearest tree than with the amount of radiation reaching the ground, i.e. growth was reduced in the vicinity of shelter trees. Therefore, we conclude that the significant reduction in height growth of seedlings of Scots pine and lodgepole pine in Scots pine shelterwoods was partially caused by factors associated with the distance to the nearest shelter tree. Because the substrate was a nitrogen-poor sandy soil, we suggest that root competition for mineral nutrients, especially nitrogen, accounts for the reduction in height growth.