A stochastic model of tree architecture and biomass partitioning: application to Mongolian Scots pines

Background and Aims

Mongolian Scots pine (Pinus sylvestris var. mongolica) is one of the principal species used for windbreak and sand stabilization in arid and semi-arid areas in northern China. A model-assisted analysis of its canopy architectural development and functions is valuable for better understanding its behaviour and roles in fragile ecosystems. However, due to the intrinsic complexity and variability of trees, the parametric identification of such models is currently a major obstacle to their evaluation and their validation with respect to real data. The aim of this paper was to present the mathematical framework of a stochastic functional–structural model (GL2) and its parameterization for Mongolian Scots pines, taking into account inter-plant variability in terms of topological development and biomass partitioning.

Methods

In GL2, plant organogenesis is determined by the realization of random variables representing the behaviour of axillary or apical buds. The associated probabilities are calibrated for Mongolian Scots pines using experimental data including means and variances of the numbers of organs per plant in each order-based class. The functional part of the model relies on the principles of source–sink regulation and is parameterized by direct observations of living trees and the inversion method using measured data for organ mass and dimensions.

Key Results

The final calibration accuracy satisfies both organogenetic and morphogenetic processes. Our hypothesis for the number of organs following a binomial distribution is found to be consistent with the real data. Based on the calibrated parameters, stochastic simulations of the growth of Mongolian Scots pines in plantations are generated by the Monte Carlo method, allowing analysis of the inter-individual variability of the number of organs and biomass partitioning. Three-dimensional (3D) architectures of young Mongolian Scots pines were simulated for 4-, 6- and 8-year-old trees.

Conclusions

This work provides a new method for characterizing tree structures and biomass allocation that can be used to build a 3D virtual Mongolian Scots pine forest. The work paves the way for bridging the gap between a single-plant model and a stand model.     

A review of ozone responses in Scots pine (Pinus sylvestris)

The data on Scots pine responses to elevated ozone (O₃) mainly come from experimental studies with young seedlings and trees. Based on the 38 experiments reviewed here, Scots pine may be considered as an O₃-sensitive conifer species, with mature pines more sensitive than younger trees. This is due to their relatively small proportion of current (c) year needles with the highest photosynthetic capacity. Moreover, young seedlings and trees seem to acclimate to slightly elevated realistic O₃ exposures, and hence do not often exhibit growth and biomass reductions in spite of the visible and microscopic needle injuries and changes in needle chemistry. The O₃ sensitivity in Scots pine is thought to relate to impaired water status due to the malfunction of stomata and subsequent increase in transpiration. This may lead to reduced wood biomass in the long term, if Scots pines try to maximise the biomass of c needles and root biomass to maintain efficient water and nitrogen (N) supply to support the photosynthesis of c needles. Tree water status also contributes to the spring-time recovery of photosynthesis. We call especially for studies on atmosphere–needle surface interaction that would yield novel information on the impact of O₃ on epicuticular waxes and stomatal functioning, which both regulate O₃ flux and tree water status and hence also modify photosynthesis. The need for flux-based field studies is especially important in the light of future climatic change, since the risk presented by O₃ to Scots pine forests in Northern and Central Europe seems to be equal.     

Branch transpiration of pine and spruce scaled to tree and canopy using needle biomass distributions

Branch water exchange and total tree water uptake were measured in a mixed Norway spruce and Scots pine stand in central Sweden during the 1995 and 1996 growing seasons. Branch transpiration was scaled to canopy level on the basis of a branch conductance model, using vertical needle-area distributions obtained by destructive sampling. Comparison with total tree water uptake scaled to canopy level showed agreement within 10%, for periods when the canopy was not affected by climatically induced stress. Comparison of scaled fluxes on individual trees showed that measurements of transpiration at branch level provide information on the direct response of transpiration to variations in weather, and furthermore that the time-lag between transpiration and tree water uptake was as much as 3 h. The vertical needle-area distribution of Scots pine was similar to that found by other authors. Needle-area distribution on Norway spruce, which has not been described before, showed that it has its largest needle area at the top of the crown. Specific needle area varied considerably both within trees and between trees. For spruce, mean specific needle area (±SD) varied from 2.4±0.5 mm² mg^–1 at the top of the crown to 7.1±1.9 mm² mg^–1 at the base. Corresponding figures for Scots pine were 3.4±2.0 and 9.1±2.1. Received: 5 March 1999 / Accepted: 17 March 2000     

The role of below-ground competition during early stages of secondary succession: the case of 3-year-old Scots pine (Pinus sylvestris L.) seedlings in an abandoned grassland

In abandoned or extensively managed grasslands, the mechanisms involved in pioneer tree species success are not fully explained. Resource competition among plants and microclimate modifications have been emphasised as possible mechanisms to explain variation of survivorship and growth. In this study, we evaluated a number of mechanisms that may lead to successful survival and growth of seedlings of a pioneer tree species (Pinus sylvestris) in a grass-dominated grassland. Three-year-old Scots pines were planted in an extensively managed grassland of the French Massif Central and for 2 years were either maintained in bare soil or subjected to aerial and below-ground interactions induced by grass vegetation. Soil temperatures were slightly higher in bare soil than under the grass vegetation, but not to an extent explaining pine growth differences. The tall grass canopy reduced light transmission by 77% at ground level and by 20% in the upper part of Scots pine seedlings. Grass vegetation presence also significantly decreased soil volumetric water content (Hv) and soil nitrate in spring and in summer. In these conditions, the average tree height was reduced by 5% compared to trees grown in bare soil, and plant biomass was reduced by 85%. Scots pine intrinsic water-use efficiency (A/g), measured by leaf gas-exchange, increased when Hv decreased owing to a rapid decline of stomatal conductance (g). This result was also confirmed by δ ¹³C analyses of needles. A summer ¹⁵N labelling of seedlings and grass vegetation confirmed the higher NO₃ capture capacity of grass vegetation in comparison with Scots pine seedlings. Our results provide evidence that the seedlings' success was linked to tolerance of below-ground resource depletion (particularly water) induced by grass vegetation based on morphological and physiological plasticity as well as to resource conservation.     

Differentiation in adaptive traits between neighbouring bog and mineral soil populations of Scots pine Pinus sylvestris

I used a reciprocal sowing experiment in the field to reveal differentiation in adaptive traits between two neighbouring northern populations of Scots pine Pinus sylvestris. 1 compared a peat bog population with a mineral soil population. Seedling survival was monitored during seven growing seasons and the plants were then harvested to obtain data on pine traits associated with growth and resource allocation, i.e. height, needle length, total dry weight, relative growth, proportion root and proportion needles. Seeds from the peat bog populations had lower germination capacity and were smaller than those from the mineral soil population. Despite their smaller size, the seeds from the bog population were superior for establishment of pines on the bog. On the mineral soil, the traits were strikingly similar in the two pine populations. In contrast, the traits were more variable and differentiated on the bog. Here, the native bog pines grew faster and had a larger proportion root than those originating from the population on the adjacent mineral soil. It is suggested that the differentiation between peat bog populations and mineral soil populations might represent a major direction of differentiation in northern Scots pine populations.     

Nutritional disturbances of young Scots pines caused by pine bark bugs in a dry heath forest

This study deals with the effect of pine bark bugs (Aradus cinnamomeus Panzer) on the nutrition of young Scots pines (Pinus sylvestris L.). Soil and needle samples for analytical purposes were collected from a young pine stand growing on a dry barren mineral soil afflicted by pine bark bugs.The damage to vascular tissues caused by pine bark bugs disturbed the nutrition of the trees, especially in the top part of the crown. The foliar calcium, magnesium, manganese and sulphur concentrations were highly reduced. Scots pines suffered from a lack of calcium, magnesium, nitrogen and phosphorus. These deficiencies were secondary by nature, because no differences were observed between the nutrient concentrations of the underlying soil of the healthy and affected trees. The symptoms of trees damaged by pine bark bugs resembled most of all calcium deficiency symptoms.     

Nine Years of Irrigation Cause Vegetation and Fine Root Shifts in a Water-Limited Pine Forest

Scots pines (Pinus sylvestris L.) in the inner-Alpine dry valleys of Switzerland have suffered from increased mortality during the past decades, which has been caused by longer and more frequent dry periods. In addition, a proceeding replacement of Scots pines by pubescent oaks (Quercus pubescens Willd.) has been observed. In 2003, an irrigation experiment was performed to track changes by reducing drought pressure on the natural pine forest. After nine years of irrigation, we observed major adaptations in the vegetation and shifts in Scots pine fine root abundance and structure. Irrigation permitted new plant species to assemble and promote canopy closure with a subsequent loss of herb and moss coverage. Fine root dry weight increased under irrigation and fine roots had a tendency to elongate. Structural composition of fine roots remained unaffected by irrigation, expressing preserved proportions of cellulose, lignin and phenolic substances. A shift to a more negative δ¹³C signal in the fine root C indicates an increased photosynthetic activity in irrigated pine trees. Using radiocarbon (¹⁴C) measurement, a reduced mean age of the fine roots in irrigated plots was revealed. The reason for this is either an increase in newly produced fine roots, supported by the increase in fine root biomass, or a reduced lifespan of fine roots which corresponds to an enhanced turnover rate. Overall, the responses belowground to irrigation are less conspicuous than the more rapid adaptations aboveground. Lagged and conservative adaptations of tree roots with decadal lifespans are challenging to detect, hence demanding for long-term surveys. Investigations concerning fine root turnover rate and degradation processes under a changing climate are crucial for a complete understanding of C cycling.     

Partitioning of carbohydrates and biomass of needles in Scots pine canopy

The study was aimed at the quantitative evaluation of the temporal and spatial partitioning of non-structural carbohydrates and needle biomass in a canopy of Scots pine (Pinus sylvestris L.) growing in a Myrtillus site type forest stand (predominant in Estonia). The tree canopy was divided into ten equal layers and the material for the spatial partitioning of the investigated characteristics was sampled from all layers. Our findings revealed a significant variation in morphology and in the partitioning of carbohydrates in needles in different layers of the canopy. The study of the temporal dynamics of carbohydrates showed that starch content in needles started to increase in early spring before budbreak, which was accompanied by a decline in soluble carbohydrates. In October, the starch content of needles was low, but the concentration of soluble sugars started to increase attaining a maximum in winter. Regression analysis indicated that before budbreak, the partitioning of soluble sugars in different canopy layers was relatively weakly correlated with the height of the layer; however, a strong correlation was observed for starch. In autumn, when the growth of trees stopped and daily temperatures decreased, the allocation of soluble sugars was correlated with the height of the canopy layer.     

Temperature sensitivity of nitrogen productivity for Scots pine and Norway spruce

Environmental conditions control physiological processes in plants and thus their growth. The predicted global warming is expected to accelerate tree growth. However, the growth response is a complex function of several processes with both direct and indirect effects. To analyse this problem we have used needle nitrogen productivity, which is an aggregate parameter for production of new foliage. Data on needle dry matter, production, and nitrogen content in needles of Scots pine ( Pinus sylvestris) and Norway spruce ( Picea abies) from a wide range of climatic conditions were collected and needle nitrogen productivities, defined as dry matter production of needles per unit of nitrogen in the needle biomass, were calculated. Our results show that the nitrogen productivity for spruce is insensitive to temperature. However, for pine, temperature affects both the magnitude of nitrogen productivity at low needle biomass and the response to self-shading but the temperature response is small at the high end of needle biomass. For practical applications it may be sufficient to use a species-specific nitrogen productivity that is independent of temperature. Because temperature affects tree growth indirectly as well as through soil processes, the effects of temperature change on tree growth and ecosystem carbon storage should mainly be derived from effects on nitrogen availability through changes in nitrogen mineralization. In addition, this paper summarises data on dry matter, production and nitrogen content of needles of conifers along a temperature gradient.     

Biomass allocation and growth responses of Scots pine saplings to simulated herbivory depend on plant age and light availability

This study experimentally analyses the response to simulated herbivory of juvenile Scots pine of two different ages in contrasting abiotic scenarios, focusing on the potential dual role of browsing ungulates: negative, by removing aerial biomass, and positive, by stimulating compensation capacity and providing nutrients by depositing their excrement. Compensation against herbivory was investigated by experimentally clipping a set of Scots pine (Pinus sylvestris L. nevadensis) juveniles, grown under different levels of light and nutrient availability. The responses analysed were survival, trunk-diameter growth, leader-shoot growth, increment in number of meristems, RGR, biomass of needles, shoots, root and whole plant, and root-to-shoot ratio. Clipping consistently resulted in a worse survival and performance of pines with respect to unclipped ones. From the factors analysed, light availability was responsible mainly for the variations in plant performance, while the addition of nutrients was much less important. Age was also important, with older pines showing in general better performance after clipping. Overall, clipping invariably had a negative effect on Scots pine, since none of the combinations of abiotic factors used resulted in overcompensation. However, the intensity on this negative effect proved quite variable, from almost an exact compensation in clipped older pines under full sunlight availability to very poor performance and high death probability in younger pines in shade. Scots pine cannot overcompensate after clipping, but, depending of the environmental conditions, the negative result of clipping varies from severe undercompensation to almost exact compensation. Also, small differences in sapling age can promote significant differences in sapling response to clipping and light environment.     

Laccaria bicolor S238N improves Scots pine mineral nutrition by increasing root nutrient uptake from soil minerals but does not increase mineral weathering

The role of ectomycorrhizal fungi on mineral nutrient mobilization and uptake is crucial for tree nutrition and growth in temperate forest ecosystems. By using a “mineral weathering budget” approach, this study aims to quantify the effect of the symbiosis with the ectomycorrhizal model strain Laccaria bicolor S238N on mineral weathering and tree nutrition, carrying out a column experiment with a quartz/biotite substrate. Each column was planted with one Scots pine (Pinus sylvestris L.) non-mycorrhizal or mycorrhizal with L. bicolor, with exception of the abiotic control treatment. The columns were continuously supplied with a nutrient-poor solution. A mineral weathering budget was calculated for K and Mg. The pine shoot growth was significantly increased (73%) when plants were mycorrhizal with L. bicolor. Whatever their mycorrhizal status, pines increased mineral weathering by factors 1.5 to 2.1. No difference between non-mycorrhizal and mycorrhizal pine treatments was revealed, however, mycorrhizal pines assimilated significantly more K and Mg. This suggests that in our experimental conditions, L. bicolor S238N improved shoot growth and K and Mg assimilation in Scots pine mainly by increasing the uptake of dissolved nutrients, linked to a better exploration and exploitation of the soil by the mycorrhizal roots.     

辽宁章古台樟子松人工林的生物量与营养元素分布的初步研究

樟子松(Pinus sylvestris var.mongolica)人工林是章古台沙地生物生产量较高的林分,其生长主要受沙丘部位和造林类型所影响。樟子松林乔木层与凋落物中贮存了大量营养元素,土壤上层的有机质与N、P元素含量较高,下层含量较低,出现一定的降低趋势。樟子松林混以小青杨可提高生物产量,增加土壤腐殖质,且改善土壤腐殖酸组成成分。     

Pine invasion impacts on plant diversity in Patagonia: invader size and invaded habitat matter

Conifers, which are widely planted as fast growing tree crops, are invading forested and treeless environments across the globe, causing important changes in biodiversity. However, how small-scale impacts on plant diversity differ according to pine size and habitat context remains unclear. We assessed the effects of different stages of pine invasion on plant communities in forest and steppe sites located in southern Chile. In each site, we sampled plant diversity under and outside the canopy of Pinus contorta individuals, using paired plots. We assessed the relative impact of pine invasion on plant species richness and cover. In both sites, richness and cover beneath pine canopy decreased with increasing pine size (i.e. height and canopy area). A significant negative impact of pines on species richness and plant cover was detected for pines over 4 m in height. The impact of pines on plant richness and cover depended on pine size (i.e. canopy area) and habitat type. Larger pines had more negative impacts than smaller pines in both sites, with a greater impact for a given pine size in the Patagonian steppe compared to the A. araucaria forest. Species composition changed between under and outside canopy plots when pines were 4 m or taller. Pine presence reduced cover of most species. The impacts of pine invasions are becoming evident in forested and treeless ecosystems of southern Chile. Our results suggest that the magnitude of pine invasion impacts could be related to how adapted the invaded community is to tree cover, with the treeless environment more impacted by the invasion.     

The simulation of soil organic matter and nitrogen accumulation in Scots pine plantations on bare parent material using the combined forest model EFIMOD

The individual-based combined forest model EFIMOD including the soil-sub model SOMM has been used for the simulation of Scots pine stand growth and soil organic matter (SOM) accumulation on a humus-free bare mineral surface. The growth of Scots pine plantation, with an initial density of 10 000 trees ha⁻¹ and average tree biomass of 0.01 kg was simulated for 50 yr under Central European climatic conditions (i) with varying atmospheric nitrogen inputs and (ii) different rates of initial application of raw undecomposed organic material or compost, on humus-free parent material. The accumulation of typical raw humus was simulated in all cases. The accumulation was most intensive in the simulation of high atmospheric nitrogen input. The humus pool in the mineral topsoil was small but achieved its maximum value with compost application. SOM nitrogen accumulation was scant in all cases, except the compost applications with low atmospheric nitrogen input. No statistically significant differences of SOM and stand parameters were found between variants without organic matter and those with low input of organic manure. However, the maximum relative rate of SOM and nitrogen accumulation was found in the scenario without organic manure, under slowly growing unstable Scots pine plantation. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date.     

Connections between climatic variables and the growth and needle dynamics of Scots pine (Pinus sylvestris L.) in Estonia and Lapland

The relationships between climatic variables and Scots pine (Pinus sylvestris L.) growth and needle dynamics were studied in three stands in Estonia and in four stands located near the northern timberline in Lapland. The trees sampled in Estonia had low correlations with the analysed climatic variables (air temperature, precipitation and indices of atmospheric circulation). Moreover, the weak cross-correlation of the time-series of the Estonian sample trees indicated that Scots pine is affected mainly by local factors in that region. In Lapland, however, height increment and needle production correlated strongly among trees within a stand (mean r=0.45 and 0.46, respectively) and between stands (r=0.32 and 0.37). Radial increment also showed a high inter-correlation among the trees within a stand in Lapland (r=0.45). Both height increment and needle production were strongly influenced by the temperature regime of the previous summer in Lapland (mean r=0.64 and 0.64, respectively). Radial increment was correlated with the mean July temperature of the current year (mean r=0.29). The correlations between the indices of atmospheric circulation and tree attributes were weak, while the strongest correlation was between the Ponta Delgada NAO index (PD–NAO) and height increment and needle production in Lapland. Height increment, needle production and radial increment have increased since the 1990s in the trees growing in Lapland. This may indicate a positive effect of climate warming on tree growth in Lapland. In Estonia, where climatic conditions do not limit tree growth, the climate warming seems not to directly influence the growth and needle dynamics of Scots pine.     

Long term effect of liming and fertilization on ectomycorrhizal colonization and tree growth in old Scots pine (Pinus sylvestris L.) stands

In this study, we surveyed the long term effects of liming and fertilizing in old Scots pine stands on the ectomycorrhiza (ECM) colonization, tree growth and needle nutrient concentration 35 years later. Four mature stands of Scots pine on low productive mineral soil were limed in 1959 and 1964 with total doses of limestone ranging from 3 to 15 Mg ha^?1 and fertilized with nitrogen (N) in 1970. Thirty-five years after the first liming treatment, all stands were analysed for tree growth and needle nutrient concentrations and two of the stands were also analysed for ECM colonization. ECM colonization increased significantly with liming from 61.5% in the control plots to 88% in the plot with the highest limestone dose. ECM colonization increased with increasing pH in the humus layer from 62% colonization at pH?=?3.5 to 90% at pH?=?6.5 and decreased with increasing amount of extractable phosphorus (P) in the humus. Liming did not affect the frequencies of different ECM morphotypes or dead short root tips, the fine root biomass or necromass. ECM colonization was uncorrelated with needle nutrient concentrations or tree increment. Liming did not significantly affect tree growth. However, nutrient concentrations of current-year needles were affected by prior liming. Ca concentrations in current-year needles increased from approximately 15 mg g^?1 in control treatments to more than 30 mg g^?1 in limed plots, whereas concentrations of Mn, Al, Fe, and in two stands, B, decreased due to liming. In conclusion, liming with doses up to 15 Mg ha^?1 was detectable in stands 35 years after treatment. The liming significantly increased the ECM colonization of Scots pine fine roots, increased the needle nutrient concentration of Ca and decreased the needle concentrations of Mn, Al, and Fe.     

Quantitative characterization of clumping in Scots pine crowns

Background and Aims

Proper characterization of the clumped structure of forests is needed for calculation of the absorbed radiation and photosynthetic production by a canopy. This study examined the dependency of crown-level clumping on tree size and growth conditions in Scots pine (Pinus sylvestris), and determined the ability of statistical canopy radiation models to quantify the degree of self-shading within crowns as a result of the clumping effect.

Methods

Twelve 3-D Scots pine trees were generated using an application of the LIGNUM model, and the crown-level clumping as quantified by the crown silhouette to total needle area ratio (STAR_crown) was calculated. The results were compared with those produced by the stochastic approach of modelling tree crowns as geometric shapes filled with a random medium.

Key Results

Crown clumping was independent of tree height, needle area and growth conditions. The results supported the capability of the stochastic approach in characterizing clumping in crowns given that the outer shell of the tree crown is well represented.

Conclusions

Variation in the whole-stand clumping index is induced by differences in the spatial pattern of trees as a function of, for example, stand age rather than by changes in the degree of self-shading within individual crowns as they grow bigger.     

Fatal interactions between Scots pine and Sphagnum mosses in bog ecosystems

In this study, we explore how Sphagnum mosses and Scots pine, Pinus sylvestris , interact on different spatial and temporal scales in a boreal bog ecosystem. We were particularly interested in relationships between the occurrence of Sphagnum- dominated habitats and the occurrence of Scots pines of different age and size.
Juvenile and adult pines occurred in different habitats. While juveniles mainly occurred in Sphagnum- dominated habitats, predominantly with Sphagnum rubellum , adult pines were found in habitats dominated by lichens, or with a sparse vegetation cover. Examination of surface peat cores sampled close to adult pines revealed that almost all pines (97%) had established in a Sphagnum -dominated environment and that the habitat had changed since pine establishment. Scots pine is thus capable of changing and exterminating the Sphagnum -dominated environment preferred for germination and establishment. Pines impede Sphagnum growth and peat accumulation significantly once they have reached a stem diameter of approximately 20 mm. It takes from 30 to 90 yr for a pine to reach that size.
Our results show the importance of interactions between Scots pine and Sphagnum mosses in bog ecosystems. We conclude that interactions between trees and Sphagnum mosses are important driving forces behind the vegetation change that has characterised boreal bogs during the Holocene.     

Perch selection by singing chaffinches: a better view of surroundings and the risk of predation

Predation risk varies among perches, and the vulnerability ofsinging chaffinches (Fringilla coelebs) might differ dependingon where they perch in a tree. To find out how singing of thechaffinch is associated with antipredatory behavior, I studiedperch selection in mature pine forest and in pine saplings,two habitats differing in the amount of cover for protection from predators. My results show that male chaffinches preferto sing below the canopy of mature pines and in the uppermostparts of sapling pines. Although these are the canopy partsmost exposed to sparrowhawk (Accipiter nisus) attacks, stayingin open sites may be a beneficial strategy for singing chaffinchesbecause it makes it possible to improve antipredatory vigilance.This assumption was supported by the hawk experiments. Afterbeing exposed to the sparrowhawk model, all of the singingbirds selected the conspicuous perch below the canopy of maturepines. The males that perched higher in the canopy before theexperiment moved to the lower canopy, whereas those males thatsang below the canopy did not change their singing perch.     

Patterns of Forest Decline and Regeneration Across Scots Pine Populations

To predict future changes in forest ecosystems, it is crucial to understand the complex processes involved in decline of tree species populations and to evaluate the implications for potential vegetation shifts. Here, we study patterns of decline (canopy defoliation and mortality of adults) of four Scots pine populations at the southern edge of its distribution and characterized by different combinations of climate dryness and intensity of past management. General linear and structural equation modeling were used to assess how biotic, abiotic, and management components interacted to explain the spatial variability of Scots pine decline across and within populations. Regeneration patterns of Scots pine and co-occurring oak species were analyzed to assess potential vegetation shifts. Decline trends were related to climatic dryness at the regional scale, but, ultimately, within-population forest structure, local site conditions, and past human legacies could be the main underlying drivers of Scots pine decline. Overall, Scots pine regeneration was negatively related to decline both within and between populations, whereas oak species responded to decline idiosyncratically across populations. Taken together, our results suggest that (1) patterns of decline are the result of processes acting at the plot level that modulate forest responses to local environmental stress and (2) decline of adult Scots pine trees seems not to be compensated by self-recruitment so that the future dynamics of these forest ecosystems are uncertain.