Differences in soil properties in adjacent stands of Scots pine, Norway spruce and silver birch in SW Sweden

Soil properties were compared in adjacent 50-year-old Norway spruce, Scots pine and silver birch stands growing on similar soils in south-west Sweden. The effects of tree species were most apparent in the humus layer and decreased with soil depth. At 20-30 cm depth in the mineral soil, species differences in soil properties were small and mostly not significant. Soil C, N, K, Ca, Mg, and Na content, pH, base saturation and fine root biomass all significantly differed between humus layers of different species. Since the climate, parent material, land use history and soil type were similar, the differences can be ascribed to tree species. Spruce stands had the largest amounts of carbon stored down to 30 cm depth in mineral soil (7.3 kg C m⁻²), whereas birch stands, with the lowest production, smallest amount of litterfall and lowest C:N ratio in litter and humus, had the smallest carbon pool (4.1 kg C m⁻²), with pine intermediate (4.9 kg C m⁻²). Similarly, soil nitrogen pools amounted to 349, 269, and 240 g N m⁻² for spruce, pine, and birch stands, respectively. The humus layer in birch stands was thin and mixed with mineral soil, and soil pH was highest in the birch stands. Spruce had the thickest humus layer with the lowest pH.     

Effects of liming on chemical properties of soil, needle nutrients and growth of Scots pine transplants

The effects of limestone (2.0 and 4.0 Mg ha⁻¹) on chemical properties of soil, nutrient concentrations of needles and growth of Scots pine (Pinus sylvestris L.) transplants were investigated on three reforestation areas on infertile acidic sites in southern Finland. The limestone was applied either on the soil surface (unploughed plots) or was mixed into the topsoil (ploughed plots). All the treatments were replicated four times. Surface broadcast of lime elevated the pH in the organic layer and in the 0-10 cm layer of mineral soil. The increase in the pH of the organic layer after 21 years was, on average, 0.7 and 1.1 pH units, with a dose of lime 2 and 4 Mg ha⁻¹, respectively. On the ploughed plots, the pH in the uppermost 0-10 cm soil layer was 0.4-0.5 units higher than on the corresponding unlimed plots. Both doses of lime significantly increased the amount of exchangeable Ca and the base saturation (BS) in the topsoil on the ploughed plots, and the amount of exchangeable Ca and Mg, as well as the base saturation (BS) in the organic layer + the 0-10 cm layer of mineral soil on the unploughed plots. Regardless of the techniques used for application of lime, after 21 growing seasons the Ca and Mg concentrations in needles were significantly higher on the limed plots than on the controls. In needles, the Ca/Mn ratio was the best indicator for measuring the response to liming. Only on the unploughed plots did liming increase stand volume and dominant diameter of pines. Intensive disc ploughing produced significantly more stems and increased both stand volume and the dominant height of pines compared to unploughed plots.     

Modelling stand structure in young Scots pine dominated stands

The purpose of this study was to construct models for predicting the structure of young Scots pine (Pinus sylvestris L.) stands. The two-parameter Weibull function characterized the height distribution of the stands. In young stands height was preferred to dbh as a random variable because of its continuous feature. Tree diameters were predicted using a multiplicative model, fitted as a linearized mixed-effect model. The modelling data consisted of repeatedly measured Scots pine dominated juvenile stands, carried out on a sub-sample of the 7th National Forest Inventory. The data covered a dominant height range from 0.2 up to 17 m. Two independent data sets were used to validate the models. The Weibull function was fitted using the maximum likelihood method. Four methods for predicting the distributions were compared: (1) parameter prediction models (PPM) consisting of seemingly unrelated regression equations, (2) a generalized linear model (GLM) which was a one-stage distribution and model fitting procedure, (3) a hybrid method including PPM for the shape parameter together with moment-based parameter recovery for the scale parameter, and (4) inclusion of moment-based parameter recovery for the scale parameter in the estimated GLM. Goodness-of-fit were tested in terms of Kolmogorov–Smirnov and error index statistics. Parameter recovery showed no improvement when used with PPM, but it improved GLM and gave the overall best performance for this new method. The constructed diameter–height model showed quite flexible and unbiased behaviour. Models are recommended as practical tools for Finnish forest management planning purposes.     

Allometric equations for estimating the foliage biomass of Scots pine

The research described in this paper was performed in the Niepolomice Forest (Southern Poland) in 2001 as part of the Forest Environmental Monitoring and Management System (FOREMMS; 5FP IST) project. The material for the present study consisted of the measurement results of the biomass of Scots pine shoots with needles and needles alone carried out on 113 felled sample trees. The purpose of this study was to construct empirical equations for estimating the foliage biomass of Scots pine from easy to measure parameters. To achieve this aim, the dependence of the foliage biomass of Scots pine on stem diameter, height, age, crown length, basal area increment of the trees was analyzed. Using the biometric characteristics such as: tree diameter at breast height (dbh), basal area increment, age, height, and crown length empirical equations for estimating the foliage biomass of Scots pine reasonably precisely have been established. The created empirical equation gives accurate foliage biomass estimates. The explained variability varies between 65 and 85%, it depends on the number of variables applied in the equation. The equations presented in this paper were created with a view to their possible use in ecological studies where biomass quantity may be used, for example, in modeling carbon circulation in the forest ecosystem. From the point of view of forestry practice, these equations may help to assess biomass production in Scots pine stands.     

Modelling annual individual-tree growth and mortality of Scots pine with data obtained at irregular measurement intervals and containing missing observations

An individual-tree growth model was developed with data from 54 permanent plots of Scots pine (Pinus sylvestris L.) located in Galicia (northwestern Spain). The study involved two model fitting approaches, one considering constant growth and mortality rates in the period between two consecutive inventories, and another considering variable growth and mortality rates in the same period. The individual-tree growth model was based on annual basal area growth, height growth and survival probability. The model included variables from groups pertaining to tree size, competition and age. Weighted regression was used as a tool for dealing with missing height observations in model fitting. Evaluation of the model via simulation of growth and mortality in the period between inventories showed that the variable growth rate approach provided slightly better results than the constant growth rate approach. The final model was consistent with expected diameter growth, height growth, dominant height growth, stand basal area growth and reduction in number of stems per hectare.     

The effects of species mixture on the growth and yield of mid-rotation mixed stands of Scots pine and silver birch

The effect of tree species mixture on stand volume yield and on tree-species-specific diameter and height growth rates were analysed in managed mixed stands of Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Ehrn.).Data were obtained from 14 repeatedly measured stands located in Southern Finland on mineral soil sites with varying admixture of Scots pine and silver birch. Statistical analysis was carried out for studying the effect of species mixture on the development of stand characteristics. For the analysis, the plots were categorised into three groups (plot types) according to the species dominance. In order to analyse species-specific growth rates, individual-tree mixed linear growth models for tree diameter and height growth were developed for both tree species.The results clearly show that the yield of the managed mid-rotation, mixed stands was greater for stands dominated by Scots pine than for stands dominated by birch, and the stand volume increment decreased with an increasing proportion of silver birch. Analysis of diameter and height growth by tree species revealed that the main reason for this pattern is the negative impact of birch competition on the growth of pine trees. The increase in diameter of pine was clearly hampered if the proportion of birch was high. An abundance of birch also slightly decreased the growth in height of Scots pine, although the effect was less than on diameter growth. Species mixture did not affect the diameter growth of birch but did have a significant effect on height development. Height growth of birch was considerably greater in pine-dominated stands than in birch-dominated stands. In pine-dominated mixed stands, the height growth of birch was quite close to that of dominant pine trees, and birches can endure in competition with pines for light.The results apply for even-aged and single-storey managed stands, where stocking density and structure are controlled with pre-commercial and commercial thinnings. The results are not applicable to unmanaged mixed stands undergoing self-thinning. This study provides new information on mixed stands from a silvicultural perspective, which can be applied in decisions involving the management of mixed stands.     

The effect of late season damage on the apical bud development of Scots pine seedlings

The effect of Lygus rugulipennis Popp. feeding and artificial damage on the apical bud of Scots pine (Pinus sylvestris L.) late in the growing season was studied on two-year-old seedlings. Lygus feeding in late July and August caused inhibition of bud formation in the subsequent year's shoots, which led to loss of apical dominance and formation of interfascicular buds. Mechanical damage caused by piercing the apical bud with a needle in July, August and October produced scars and malformations on subsequent year's shoots and buds, but did not inhibit bud formation. Damage to the apical meristem could not be detected visually on the dormant bud before shoot elongation.     

Modelling mortality of Scots pine (Pinus sylvestris L.) plantations in the northwest of Spain

Mortality is an important element of growth and yield models, especially if only low intensity silvicultural treatments are carried out. The objective of the present study was to develop a model for predicting tree number decline in planted even-aged stands of Scots pine (Pinus sylvestris L.) in Galicia (northwestern Spain). The model was constructed using data from two inventories of a trial network involving 68 permanent plots located in unthinned stands, or stands thinned lightly from below. Two alternatives were tested. In one alternative, a two-step modelling strategy was applied. First, a binary response function predicting the survival probability of all the trees in the stand was constructed, and an equation for reduction in tree number was developed, using only data where death had occurred over the period analyzed. Three different approaches were then used to compare the application of the above-mentioned functions together. In the other alternative, a mortality function for directly predicting the reduction in tree number was fitted, including all plots (with and without occurrence of mortality). Both alternatives provided similar results, showed logical behavior, and performed satisfactorily in evaluation tests. However, in choosing the best strategy for inclusion in a stand-level simulator, the use of the second alternative is recommended because it possesses the path invariance property required in a mortality model.     

帽儿山地区不同种源樟子松树轮对气候因子的响应

[目的]为了进一步了解不同种源的樟子松在帽儿山地区的生长差异,研究该地区气候因子对樟子松径向生长的影响,本文建立了来自于阿尔山、高峰、红花尔基、卡伦山、图强罕达盖和金山等7个种源樟子松树轮年表。[方法]通过年轮-气候响应分析、相关分析、主成分分析,探讨了不同种源樟子松径向生长与气候因子的响应差异。[结果]表明:7个种源地樟子松树轮年表整体变化趋势基本相似,但局部也有不同之处,除图强种源外,其余6个种源地樟子松在生长初期年轮宽度呈增加趋势。阿尔山、高峰、红花尔基和卡伦山种源地樟子松径向生长与当年2月平均温度显著正相关,罕达盖种源地樟子松径向生长与前一年4月平均温度显著正相关;高峰、图强、卡伦山种源地樟子松径向生长与当年6月降水显著正相关,阿尔山、高峰、金山、卡伦山、红花尓基、图强种源樟子松的径向生长主要与前一年8月降水显著负相关。[结论]多数种源的樟子松在初期径增长较快,与当年2月份的温度和6月份的降水量有正相关的响应性,而与前一年8月份的降水量有负相关的响应性。     

Reduction of frost heaving of Norway spruce and Scots pine seedlings by planting in mounds or in humus

This study was carried out in northern Sweden to determine the effects of frost heaving on the establishment of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings in relation to planting methods. For this purpose, one year old containerised seedlings were planted on two sites and on two dates: during the spring (early planting) and during the fall (late planting). In each case, two planting depths (normal and deep planting) and four planting sites (in mineral soil in the depressions, in the scalp/trench area, on the top of the mound and in the untreated humus layer) were used. On each site, 50 seedlings were planted for each treatment. Frost heaving was observed and measured during two years. The amount of heaving was highest in the hole and almost insignificant on the top of the mound and in the humus layer. Planting depth influenced the degree of heaving only for Scots pine planted in the hole and was not related to the planting time.     

Peroxidase activity, isoenzymes and histological localisation in sapwood and heartwood of Scots pine (Pinus sylvestris L.)

Peroxidase activity and isoenzymes of fresh wood samples of the third shoot of 12-year old trees and from the sapwood, transition zone and heartwood of c. 60-year old stems of Scots pine (Pinus sylvestris L.) were investigated. Wood samples were ground at −30°C, extracted, and the extracts concentrated c. 20-fold for peroxidase activity assays (guaiacol method) and for IEF-PAGE. At least 11 major isoenzymes could be found in the gels. Even the heartwood contained some peroxidase isoenzymes. Isoenzyme patterns of the juvenile wood did not change with the season. However, juvenile wood showed the highest peroxidase activity at the end of the growing season. Peroxidase activity decreased from the outer sapwood towards the heartwood. Thin sections of different wood zones stained for peroxidase revealed activity in ray parenchyma and resin canal epithelial cells. Intensive staining was localised in the bordered pits of vertical and ray tracheids, and in the end walls of ray parenchyma cells.     

Climate and historical stand dynamics in the tropical pine forests of northern Thailand

Forest recruitment is the outcome of local- and regional-scale factors such as disturbances and climate. The relative importance of local- and regional-scale factors will determine the spatial scale at which temporal pulses of recruitment occur. In seasonal tropical forests, where the annual dry-season is a critical bottleneck to seedling survival, multi-year periods of relatively cool, wet dry seasons may be required for successful tree recruitment. Consequently, when such conditions are present, region-wide synchronisation of recruitment may occur. To examine the case for regional synchronisation of forest dynamics in the seasonal tropical pine forests of northern Thailand, we investigated forest age structures at three spatial scales: stand, site and region. We compared forest age structures with instrumental climatic records beginning in 1902. We found significant statistical evidence of synchronous recruitment at the stand- and site-scales, but not at the regional-scale. While correlations between recruitment and climate were not statistically significant, recruitment success was often linked to favourable climatic conditions. For example, recruitment at all sites was associated with multi-year periods of cool-wet dry seasons. The lack of significant correlations between recruitment and climate appears to reflect complex interactions among local disturbance history, regional climate variability and pine recruitment.     

Increased mortality can promote evolutionary adaptation of forest trees to climate change

Forecasts of rapid climate change raise the question how quickly species can evolutionarily adapt to future climates. The adaptability of forest trees to environmental changes is generally promoted by high levels of genetic diversity and gene flow, but it can also be slowed down by long generation times and low mortality of established trees. Here, we investigate the adaptation of Scots pine (Pinus sylvestris) and Silver birch (Betula pendula) to climate change induced prolongation of the thermal growing season. We use quantitative genetic individual-based simulations to disentangle the relative roles of mortality, dispersal ability and maturation age for the speed of adaptation. The simulations predict that after 100 years of climate change, the genotypic growth period length of both species will lag more than 50% behind the climatically determined optimum. This lag is reduced by increased mortality of established trees, whereas earlier maturation and higher dispersal ability had comparatively minor effects. The evolutionary lag behind environmental change shown in our simulations stresses the importance of accounting for evolutionary processes in forecasts of the future dynamics and productivity of forests. Sensitivity of the adaptation speed to mortality suggests that species experiencing high mortality rates as well as populations subject to regular disturbances such as storms or fires might be the quickest to adapt to a warming climate.     

Species substitution for carbon storage: Sessile oak versus Corsican pine in France as a case study

Species choice is potentially an important management decision for increasing carbon stocks in forest ecosystems. The substitution of a slow-growing hardwood species (Quercus petraea) by a fast-growing conifer plantation (Pinus nigra subsp. laricio) was studied in central France. Simulations of carbon stocks in tree biomass were conducted using stand growth models Fagacées for sessile oak and PNL for Corsican pine. The changes in soil carbon were assessed using the Century model and data from two European soil monitoring networks: 16 km × 16 km grid and RENECOFOR. Carbon in wood products was assessed with life cycle analysis and lifespan of final products. However, only carbon stocks and their variation were accounted for: effects of energy-consuming materials or fossil fuel substitution are excluded from the analysis. To compare the growth of these two types of forest stands, an important part of the study was to assess the productivity of both species at the same site, using National Forest Inventory data.     

Does selecting for improved growth affect wood quality of Pinus pinaster in Portugal?

The selection criteria for the first generations in the Portuguese Pinus pinaster improvement program have been the growth rate and form traits. In this work we study the consequences of this selection on wood quality traits. This study assesses genetic and phenotypic correlation between growth, wood density components, lignin content and mechanical traits (radial modulus of elasticity and radial modulus of rupture) of 46 half sib families from a progeny trail located in Leiria, Portugal, originated from seed collected in a clonal seed orchard. A total of 552 seventeen-years-old trees (about half of full rotation age) were sampled at 2 m height. Height measured at 12 years old presented a higher genetic control (h² = 0.34) relatively to DBH, measured at 12 and 17 years old respectively (h² = 0.17 and h² = 0.15). The results of this study also showed that DBH growth is more dependent on latewood components than earlywood components and that higher growth in Mediterranean regions can be due to an increase of the period of latewood formation. Further, we can conclude genetic selection based on growth will not result in a decrease of wood density, will not affect the occurrence of spiral grain, and is possible to obtain an increase in the radial modulus of elasticity. The present study also showed that it is possible to select for increased growth with lower lignin proportion. Results also suggest that selection for growth at 12 years will probably not affect negatively the wood properties at 17 years.     

The role of germination microsite in the establishment of sugar pine and Jeffrey pine seedlings

Mature shrubs can provide microhabitats that are beneficial to tree seedling growth and development. Sugar pine trees (Pinus lambertiana) grow in a narrow zone on the eastern slope of the Carson Range in extreme western Nevada, whereas Jeffrey pine (Pinus jeffreyi) is the dominant tree species in the region, an area extensively disturbed by wild fire. This study compares seedling establishment of sugar pine and Jeffrey pine relative to mature shrubs. In the fall of 2002 (cohort 1) and 2003 (cohort 2), 13,600 seeds of both species were planted in wire mesh enclosures, at three sites, under a variety of microhabitat treatments: under shade and in the open, under two species of shrub cover, and with and without plant litter. Seedlings were monitored for survival through two growing seasons. Even though more sugar pine seedlings emerged, more Jeffrey pine seedlings survived, and Jeffrey pine was the more drought tolerant species, better suited for the xeric climate found in the Carson Range. Litter slightly hindered seedling emergence but had no effect on survival and there was no significant species × litter interaction. Supplemental water facilitated survival in all treatments with highest survival in shade treatments. Sugar pine seedlings showed a significant increase in survival over Jeffrey pine seedlings with the addition of water, particularly in open treatments and more of both species survived under manzanita shrubs with water. The highest seedling mortality occurred when shrub canopy was removed, and seedlings experienced the effect of full sun and competition for soil water. For either species, microhabitat is a significant factor in determining success or failure in rehabilitation efforts after disturbance.     

Modeling jack pine branch characteristics in Eastern Canada

A total of 83 trees were sampled in three regions of Eastern Canada in order to model branch characteristics (number of branches per annual shoot, branch insertion angle and diameter) using linear mixed-effects models. Differences in branch characteristics according to branch type (pseudo-whorl at annual shoot apical end (PWA) versus pseudo-whorl between shoot apical ends (PWB)) were also studied. The number of both PWA and PWB branches are proportional to annual shoot length, whereas the number of PWB branches also decreases with tree age. Insertion angle was mainly driven by annual shoot number from apex (branch age). The diameter models showed the most complexity with branch vertical position and tree size (DBH and total height) among the statistically significant variables. Region and plot random effects were minimal compared with tree and annual shoot levels. Tree-level random effects were significant for every model and might be a symptom of genetic control over the number of branches and, to a small extent, branch diameter. Interaction between insertion angle and diameter is relatively strong because all the models using them as independent variables (except for the model of insertion angle for PWB branches) showed better fit statistics. These results lead us to believe that tree-, annual shoot- and branch-level variables should be further explored in order to better understand branch dynamics.     

Irrigation regime affects water and aeration conditions in peat growth medium and the growth of containerized Scots pine seedings

Containerized Scots pine seedlings growing in light Sphagnum peat growth medium in a greenhouse were subjected to three irrigation treatments. The seedlings were irrigated by saturating the growth medium with a nutrient solution (i) daily or when (ii) –5 or (iii) –10 kPa medium matric potential was achieved. In general, the nutrient concentrations in the foliage of the seedlings were rather similar for all irrigation treatments. In the wet (daily) irrigation treatment, however, the foliar concentration of N was lower and the foliar concentrations of P, Ca, S, Mn and Na were higher than in the other two treatments. In the wet irrigation treatment, height growth was lower than that in the other treatments. In this treatment the shoot to root ratio and several other morphological characteristics were also reduced, while the number of root lenticels and seedling mortality were increased compared to the other treatments. Rapid growth was obtained by irrigating at –5 to –10 kPa matric potential. In the wet irrigation treatment, the median matric potential and air-filled porosity of the peat medium were –0.4 kPa and 11%, respectively, which indicated waterlogging and very suboptimal aeration to the seedlings. The respective values for the irrigations at –5 and –10 kPa were –2.7 to –3.3 kPa and 39 to 43%, which were considered to cover the range from optimum to slightly suboptimum for average water availability and aeration.     

Predicting sapstain and degrade in fallen trees following storm damage in a Pinus radiata forest

Storm damage in production forests constitutes a major source of economic loss world wide, yet the retrieval of salvageable timber remains problematic. In particular, an inability to anticipate when sapstain and degrade will appear hampers the planning of log recovery operations. A study was conducted to monitor the deterioration of fallen trees following two winter storms causing wind and snow damage in a Pinus radiata plantation forest in the upper South Island of New Zealand. Percentage sapstain, incidence of basidiomycete decay fungi, and frequency of bark beetle infestation increased, while percentage sapwood moisture content decreased, over a period of 1 year. These changes proceeded more rapidly in fallen trees that were severed at stump height, to simulate breakage, than in those that were left partially rooted. There was little beetle activity at the time of the storms, but Arhopalus ferus (Coleoptera: Cerambycidae), and Hylastes ater, Hylurgus ligniperda and Pachycotes peregrinus (Coleoptera: Curculionidae: Scolytinae), were collected in flight traps during the following spring and summer. The predominant fungal species associated with sapstain was Diplodia pinea, while Ophiostoma piceae and Grosmannia huntii were isolated near the end of the period. The main decay fungi obtained were Phlebiopsis gigantea, Stereum sanguinolentum, and Schizophyllum commune. A generalized linear mixed model was constructed to predict the development of sapstain in fallen trees for conditions prevailing during the study after a storm at the same time of year. According to the model, a 10 m long butt log of 22 cm mid length diameter will have minimal stain (<10% of the cross sectional area affected) when cut from severed stems up to 4 months after the storm; if taken from still-rooted trees this period will extend to 1 year. However, because of large between-tree variation, economically productive log recovery will also depend on the proportion of trees that lie below an acceptable sapstain threshold. Further research is needed to determine regional and seasonal influences on the development of sapstain in fallen trees.