Long-term trends in height growth of Picea obovata and Pinus sylvestris during the past 100 years in Komi Republic (north-western Russia)

Abstract

Adapting forest management practices to the changing environment of Komi Republic requires an understanding of the response of unmanaged natural forests to climatic changes. Komi Republic is a region of north-western Russia where large areas of natural boreal forest still exist. Apical growth of Scots pine (Pinus sylvestris L.) (108 trees, 529 discs) and Siberian spruce (Picea obovata Ledeb.) (88 trees, 423 discs) was analysed using stem analysis techniques. Mean apical growth curves were calculated for four zones of boreal forests in two subsequent 50 year periods starting from 1900. A statistically significant increase in height increment of 40% for Siberian spruce and 30% for Scots pine was identified from samples representing the Komi Republic. Within this region statistically significant height increment increases were found in the middle taiga zone for Siberian spruce of 240% and Scots pine of 140%, while northern taiga Siberian spruce increased by 164%. Taking into account that trees were samples in remote untouched pristine forests, the main causes of increased height increment are suggested to be climatic, which shows a large increase in temperature (0.43°C during the past 30 years) and a modest decrease in precipitation (2.2% over the past 30 years).     

Distribution and ecology of Armillaria species in Greece

Five Armillaria species were identified in a nationwide survey in Greece. Armillaria mellea was present in coniferous and broad-leaved forests in most of the areas examined, except the high altitudes (above 1100 m) of the mountains of north Greece. It was found to cause significant damage in fir forests as well as in fruit orchards and vineyards. Armillaria gallica was common in coniferous and broad-leaved forests in the high altitudes of central and northern Greece, predominating in the beech forests. The fungus was a weak parasite or a saprophyte of forest trees and was occasionally found on cultivated plants. Armillaria ostoyae was not found in southern and central parts of the country, but it has a wide distribution in the mountain forests of northern Greece and causes significant damage on fir, black pine, Scots pine and spruce. Armillaria cepistipes was recorded at high altitudes (1400–1800 m) on two mountains of northern Greece, mostly as a saprophyte in coniferous and broad-leaved forests. Armillaria tabescens was rare in the forests of Greece; it was found to cause disease in almond tree orchards.     

Modelling the distribution of wood properties along the stems of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as affected by silvicultural management

In this work, empirical ring-based models were developed to predict the distribution of early wood percentage, wood density and fibre length along the stems of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as affected by silvicultural management. The performance of the ring-based models was also compared for Scots pine and Norway spruce with corresponding disc-based (cross-sectional) models. Moreover, both models were integrated with example simulations by a process-based growth and yield model to analyze how management, such as thinning, affects the growth and wood properties of Scots pine trees over a rotation as an average for the tree stem, but also along the stem.The ring-based models built for annual early wood percentage (explained by ring width), air dry wood density (explained by early wood percentage and cambial age) and fibre length (explained by radial growth percentage and cambial age) predicted reasonably well the wood properties both at an intra-ring level, but also at a cross-sectional level. These predictions were also reasonably well in line with corresponding cross-sectional predictions by the disc-based models (which predicted the properties based on the number of annual rings and diameter at breast height and/or the cross-section being considered and temperature sum). The example simulations also demonstrated that both models predicted slightly lower wood density for dominant trees compared to dominated ones grown in thinned and unthinned Scots pine stands over a rotation. Unlike the disc-based model, the ring-based model predicted, on average, higher early wood percentage in dominant trees than in dominated ones. However, fibre length was not significantly affected when the averages of the whole stems were predicted, and this held true for both ring- and disc-based models.In summary, the incorporation of empirical ring-based wood property models into a process-based growth and yield model, offers a means to study in detail how environmental conditions, forest structure and management affect the quantity and properties of stem wood produced over a rotation. The disc-based wood property models used in this work are based on data with large geographical and genetic variation, and therefore may turn out to be more applicable for predicting future wood and fibre resources at a regional and national level. This kind of integrated use of wood property models with a process-based growth and yield model could help us to evaluate the forest resources under current and changing climate.     

Larval performances and life cycle completion of the Siberian moth, <Emphasis Type="Italic">Dendrolimus sibiricus</Emphasis> (Lepidoptera: Lasiocampidae), on potential host plants in Europe: a laboratory study on potted trees

The Siberian moth, Dendrolimus sibiricus, Tschtv. is the most harmful defoliator of coniferous forests in North Asia. The pest has already spread over the Urals and continues moving westwards. Recently, it has been recommended for quarantine in member countries by European and Mediterranean Plant Protection Organization (EPPO). The performances of the pest on coniferous species planted in Europe were assessed on a range of potted trees corresponding to the spectrum of economically important conifers in the EU: European larch Larix decidua, Norway spruce Picea abies, Scots pine Pinus sylvestris, European black pine Pinus nigra, and the North American species: Douglas fir Pseudotsuga menziesii and grand fir Abies grandis. Larvae showed a potential to survive and complete the development on all these host tree species. Favorable hosts were grand fir, European larch, and Douglas fir that allowed higher survival, better larval development, and as a result, yielded heavier pupae and adult moths with higher longevity. Black pine was a poor host but, however, could still support larval and pupal development. Norway spruce and Scots pine had an intermediate behavior. If accidentally introduced to Europe, the Siberian moth may become especially damaging in forest stands predominated by European larch and by the North American firs. Norway spruce and especially the two-needle pines will be less prone to intensive defoliation by this species. The fact that the pest may damage the range of economically important coniferous species should be taken into account in the pest risk assessment for Europe and also for North America where the Siberian moth occurrence is considered likely.     

Quantifying the effect of pine mistletoe on the growth of Scots pine

Mistletoe infection results in substantial growth losses in mistletoe‐infected forests. This study reports and evaluates the results of retrospective analyses of radial growth of Scots pine (Pinus sylvestris) in relation to the level of infection of pine mistletoe (Viscum album ssp. austriacum). A total of 43 Scots pine trees were destructively sampled from different sites. Of these trees, 14 were uninfected and 29 were infected. Infection classes were determined using six‐class dwarf mistletoe rating system (DMRS). All needle and mistletoe biomass were removed completely and weighed for each sampled tree. Subsamples from needles and all mistletoe biomass were taken to the laboratory for oven‐dried weight determinations. Five‐cm‐thick wood discs were cut from the stem at the breast height (1.3 m) to determine annual basal area increment for the last 25 years. In addition to DMRS, new infection classes were created using mistletoe‐to‐needle biomass (MB/NB) ratio. The results showed that the radial growth losses could be as much as 41% to 64% at different infection levels. The rate of growth loss in relation to DMRS and MB/NB ratio was similar, but with a larger variability in DMRS values. The results showed that both DMRS rating and MB/NB ratio seem to be important for quantifying growth loss on Scots pine trees infected with mistletoe. The results of this study can also be invaluable in modelling the effects of mistletoe on the growth of Scots pine trees.     

Low- to moderate-severity historical fires promoted high tree growth in a boreal Scots pine forest of Norway

Abstract

Fire is the most important ecological factor governing boreal forest stand dynamics. In low- to moderate-severity fires, the post-fire growth of the surviving trees varies according to fire frequency, intensity and site factors. Little is known about the growth responses of Scots pine (Pinus sylvestris L.) following fires in boreal forests. We quantified changes in tree growth in the years following 61 historical forest fires (between 1210 and 1866) in tree-ring series collected from fire-scarred Scots pine trees, snags and stumps in Trillemarka nature reserve in south-central Norway. Basal area increment 10 years pre-, 5 years post-, and 11–20 years post-fire were calculated for 439 fire scars in 225 wood samples. We found a slight temporary growth reduction 5 years post-fire followed by a marked growth increase 11–20 years post-fire. Beyond 20 years post-fire, the long-term tree growth declined steadily up to approximately 120 years. Our results indicate that recurring fires maintained high tree growth in remnant Scots pines, most probably due to a reduction in tree density and thus decreased competition.     

Forest Management and Regeneration Success in Protection Forests near the Timberline in Finnish Lapland

Sustainable forestry is practised in Finnish Lapland north of latitude 69°, i.e. farther north than anywhere else in the world. The Protection Forests Act was passed in 1922 to guarantee the stability of timberline forests in this climatically extreme, unique area. This review, based on earlier studies, covers forest and climate statistics, and legislation, regulations and recommendations of forest management, and the effects of management practices and climatic trends on the results of natural and artificial regeneration of conifer forests, as well as on shifts in the timberline. The review is mainly based on Finnish research, but references from the other Fennoscandian countries have also been used. Temperature plays the main role in forest regeneration success, and the effective temperature sum has been found to express well the location of the Scots pine (Pinus sylvestris L.) timberline. Natural regeneration of pine has been satisfactory, especially when combined with soil preparation. In many cases the pine forests regenerate in the form of advance growth without any silvicultural action. The success of artificial regeneration seems to be associated with favourable, warm periods lasting for at least 20 yrs. The effects of cutting, forest fires, reindeer herding, biotic damage and air pollution on forest regeneration and the conifer timberlines are discussed.     

Growth responses of three coexisting conifer species to climate across wide geographic and climate ranges in Yukon and British Columbia

This work aimed to compare radial growth–climate relationships among three coexisting coniferous tree species across a wide geographic and climate range from southern British Columbia (BC) to central Yukon, Canada. Tree-ring data were collected from 20 mature stands of white spruce (Picea glauca), lodgepole pine (Pinus contorta var. latifolia), and subalpine fir (Abies lasiocarpa). Linear relationships between annual growth variation and monthly and seasonal climate were quantified with correlation and regression analyses, and variation in climate–growth responses over a climatic gradient were quantified by regressing growth responses against local mean climatic conditions. Temperatures had more consistent and stronger correlations with growth for all three species than precipitation, but growth–climate responses varied among species and among sites. In particular, pine and fir populations showed different responses between BC and Yukon, whereas spruce showed a more consistent response across the study domain. Results indicate that (1) the response and sensitivity of trees to seasonal climate variables vary among species and sites and (2) winter temperatures prior to growth may have significant impacts on pine and fir growth at some sites. The capacity to adapt to climate change will likely vary among the study species and across climatic gradients, which will have implications for the future management of mixed-species forests in Yukon and BC.     

Damage by pine weevil Hylobius abietis to conifer seedlings after shelterwood removal

Abstract

Pine weevil (Hylobius abietis L.) damage to seedlings after overstorey removal was investigated in a survey study in six shelterwoods in the south–central part of Sweden. The shelterwoods predominantly consisted of Scots pine, except at one site where the shelter trees mainly consisted of Norway spruce. Before final cutting, 10 plots were laid out at each site and measurements of shelter trees and marked seedlings were taken. The seedlings were examined during the 2 years after final cutting. The study showed that removal of shelter trees increases the risk of severe damage by pine weevil and the variable that was most strongly correlated with the risk was the seedling root collar diameter. Both Scots pine and Norway spruce seedlings were severely damaged by pine weevil, and most of the feeding occurred during the first year after cutting. The amount of debarked area was significantly larger for Scots pine than for Norway spruce seedlings. Vitality (growth of the leading shoot before final cutting) of the seedlings also affected the probability of damage. Seedlings with high vitality were less damaged by pine weevil than seedlings with low vitality. For Scots pine the shelterwood density before final cutting was correlated to the intensity of pine weevil feeding after cutting. In conclusion, after the final cutting of a pine or spruce shelterwood, pine weevils will probably invade the area. To avoid serious damage, Norway spruce and Scots pine seedlings should have reached a diameter of at least 10–12 mm.     

The effects of nitrogen fertilization on the ultrastructure of mesophyll cells of conifer needles in northern Finland

Cytological injuries in needles of Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.] were studied at light and electron microscope level in northern Finland. Needle samples were taken from fertilized (mainly with nitrogen) and nonfertilized trees during winter 1980. The results are discussed in relation to the severe climate and growth response of the trees.     

The pine pathogen Diplodia sapinea is associated with the death of large Douglas fir trees

Climate change poses severe pressures to European conifer forests. Using non-native tree species, such as Douglas fir (Pseudotsuga menziesii), is one proposed strategy to circumvent adverse effects for forest management. However, novel forest health risks can impair the cultivation of non-native trees. In 2022, we observed large Douglas fir trees (approximately 40–50 years old, diameter at breast height (dbh) 21–41 cm) that had recently died in spring or summer 2022 in three forest stands in Eastern Austria. Intensive resin flow, blue-staining of the sapwood and the absence of bark- and wood-boring insects indicated a fungal infection. Isolations from blue-stained sapwood of the dead trees consistently yielded cultures of the opportunistic pathogen Diplodia sapinea. In a greenhouse wound inoculation experiment, seven D. sapinea isolates obtained from Douglas fir caused phloem necrosis, blue-staining of sapwood and mortality and thus displayed pathogenicity towards seedlings of both Ps. menziesii and its common host, Scots pine (Pinus sylvestris). Diplodia sapinea produced significantly longer areas of blue-stain as well as higher and faster mortality in Ps. menziesii compared to P. sylvestris. We conclude that D. sapinea substantially contributed to the death of seven of the 13 examined large Douglas fir trees. While this fungus has been described as a pathogen of young Douglas fir trees before, this is the first report that it can potentially kill large individuals of this conifer species under drought conditions. Thus, our results indicate that D. sapinea could represent a severe threat to the cultivation of Ps. menziesii in European forestry.     

Ectomycorrhizal communities associated with silver fir seedlings (Abies alba Mill.) differ largely in mature silver fir stands and in Scots pine forecrops

Context

The requirement for rebuilding forecrop stands besides replacement of meadow vegetation with forest plants and formation of soil humus is the presence of a compatible ectomycorrhizal (ECM) fungal community.

Aims

This study aims to assess ectomycorrhizal fungi diversity associated with silver fir (Abies alba Mill.) seedlings regenerating in silver fir stands and Scots pine forecrops.

Methods

One-year-old seedlings were sampled in six study sites: three mature fir forests and three pine forests. ECM fungi were identified by polymerase chain reaction amplification and sequencing of the internal transcribed spacer of rDNA.

Results

The mean mycorrhizal colonization exceeded 90 %. Thirty-six ectomycorrhizal taxa were identified in fir stands and 23 in pine forecrops; ten out of these species were common to both stands. The fungal communities were different between study sites (R?=?0.1721, p?=?0.0001). Tomentella stuposa was the only species present at all sites.

Conclusion

Silver fir seedlings in Scots pine forecrops supported smaller ECM fungal communities than communities identified in mature silver fir stands. Nevertheless, fungal colonization of seedling roots was similar in both cases. This suggests that pine stands afforested on formerly arable land bear enough ECM species to allow survival and growth of silver fir seedlings.     

Effects of silvicultural cleaning in mixed pine‐deciduous stands on moose damage to scots pine (Pinus sylvestris)

Silvicultural cleaning is commonly carried out in young stands to control competition between conifers and deciduous tree species. However, it has been questioned whether this practice affects moose (Alces alces L.) browsing on Scots pine (Pinus sylvestris L.) with respect to damage at the stand level. In this study, moose winter browsing was examined in relation to food availability in 19 Scots pine stands divided into silviculturally cleaned and non‐cleaned blocks. Cleaning had a negative effect on moose foraging in terms of reduced density, biomass availability, and height of the deciduous tree species. The total biomass consumed by moose was significantly lower on the cleaned blocks than on the non‐cleaned ones. There were no differences between the treatments in the amounts of pine consumed and in moose damage to pines. The density or biomass available of different deciduous tree species did not explain the variation in moose browsing on pine. In conclusion, timing of silvicultural cleaning with reference to moose browsing should be taken into account. If pines are not overtopped by deciduous trees, as in the present study, silvicultural cleaning is not necessary to prevent moose damage.     

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.     

Crystalline structure of heat-treated Scots pine [<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.] and Uludağ fir [<Emphasis Type="Italic">Abies nordmanniana</Emphasis> (Stev.) subsp. <Emphasis Type="Italic">bornmuelleriana</Emphasis> (Mattf.)] wood

The aim of this study was to determine changes in crystallinity and crystalline unit cell type of heat-treated Scots pine (Pinus sylvestris L.) and Uludağ fir (Abies nordmanniana stev. subsp. bornmuelleriana Mattf.) wood samples by means of FT-IR spectroscopic method. Heat treatment was applied on the test samples in an oven at three different temperatures (120, 150, and 180°C) and for two different periods of time (6 and 10 h) under atmospheric pressure. It was designated that crystallinity of both Scots pine and Uludağ fir wood samples increased during heat treatment depending on the duration. However, monoclinic structure in crystalline unit cells of Scots pine and Uludağ fir wood samples converted to triclinic structure when heat treated. It was estimated that monoclinic structure was dominant in the crystalline unit cell. It was established that the crystalline structure of Scots pine wood samples was more affected by heat treatment than that of Uludağ fir wood samples.     

Inter- and intra-annual variations in canopy fine litterfall and carbon and nitrogen inputs to the forest floor in two European coniferous forests

? Context

The amount and chemistry of litterfall have been known to strongly vary among the years with important implications for ecosystem nutrient cycles, but there are few quantitative data describing such variations.

? Aims

We studied the climatic implications on the variation in litterfall and its C and N input to soil in two distinct European coniferous forests.

? Methods

Year-to-year variations in canopy litterfall were measured in a Scots pine stand (Hyytiälä, Finland) over 13 years, and a Douglas fir stand (Speulderbos, The Netherlands) over 3 years.

? Results

Important inter-annual variations in litterfall were observed in Scots pine. Litterfall was mainly driven by leaf senescence; however, premature needle fall was observed in high wind speed and early frost events. The seasonal variation in litterfall was characterized by a maximum in September in Scots pine, and by a biphasic variation pattern in Douglas fir, in May and November. Lower seasonal variations and lower annual average in litterfall N content were observed in Scots pine.

? Conclusion

Significant inter- and intra-annual variations in litterfall and chemistry and between the sites were demonstrated; and it depended on year-to-year differences in climate and extreme weather events.     

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.     

The effects of pine mistletoe (Viscum album subsp. austriacum) on the growth of Scots pine and Crimean pine in Turkey

In this study, the effect of pine mistletoe (Viscum album subsp. austriacum) on basal area increment of Crimean pine and Scots pine was investigated. Dendrochronological data were collected from 223 (71 uninfected and 152 infected) Crimean pines and 195 (77 uninfected and 118 infected) Scots pines located in Kastamonu province of Turkey in 2014. Infected sample trees were classified as light, moderate or severe infection levels. Growth trends and basal area increment loses were compared between uninfected and infected trees for the periods of the last 10, 20 and 30 years. In addition, infection status of forest stands was investigated using temporary sample plots; 27 plots in Crimean pine stands and 26 plots in Scots pine. Results demonstrated that basal area increments were negatively affected by pine mistletoe for both species. Mean basal area increment losses of infected trees for the last decade were determined as 24% for Scots pine and 26% for Crimean pine. Basal area increment losses varied by infection levels (light, moderate and severe) as follows: 25%, 20% and 28% for Scots pines and 20%, 32% and 9% for Crimean pines. Scots pine stands were more severely infected by pine mistletoe than Crimean pine stands. There were negative correlations between number of infected trees and stand density for both species, while positive correlation was detected between the number of infected trees and mean diameter for Scots pine. The results of this study indicate that the pine mistletoe infection has negative effect on radial growth of Scots pine and Crimean pine trees. The results can be an important contribution to the forest management and protection activities in mistletoe-infected stands.     

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.     

The effect of canopy position on growth and mortality in mixed sapling communities during self-thinning

This research investigates how species in the sapling phase differ in growth and survival depending on light availability (as estimated by canopy position) by means of tree-ring analysis and modelling mortality. We harvested 120 live and 158 dead saplings in self-thinning communities consisting of Silver birch (Betula pendula Roth.), Scots pine (Pinus sylvestris L.), Japanese larch (Larix kaempferi Carr.) and Douglas fir (Pseudotsuga menziesii Mirb. Franco) in the Netherlands. Results are evaluated within the framework of a trade-off between high-light growth and low-growth survival. Radial growth, measured at ground level, generally declined over time. In addition, a decreasing light availability further reduced growth in all species except Douglas fir. Trees died when radial growth was reduced to about 0.5 mm year⁻¹. Mortality in all species except Scots pine was significantly related to recent growth, but mortality curves were not different. The light-demanding Silver birch and Japanese larch differed from the shade-tolerant Douglas fir in both high-light growth and low-growth mortality, in line with a growth-survival trade-off. The light-demanding Scots pine did not fit this pattern as it was unable to transfer high radial growth into height gain, leaving it in suppressed canopy positions. This indicates the importance of height growth in the growth-survival trade-off. Differences in mortality probabilities affect the potential for coexistence, however, in all species also fast-growing individuals died suggesting additional factors causing mortality during self-thinning, other than direct competition for light.