Inherited narrow crown form, harvest index and stem biomass production in Norway spruce, Picea abies

The effect of crown form on stem biomass production was investigated in an 18-19-year-old Norway spruce stand (Picea abies (L.) Karst.). The harvest index was 0.271 in pendula trees, which have a heritable narrow crown form, and 0.235 in normal-crowned trees and the dry weights of stem biomass were 2.57 kg and 3.37 kg, respectively. However, the production of stem biomass per crown projected area was more than twice as much in pendula trees as in normal-crowned trees. Results indicate that the crown form of pendula spruce is less plastic than that of normal-crowned spruce suggesting that such trees might maintain a high stemwood yield per unit ground area with increasing stand density. Because pendula spruce has a higher harvest index and a less flexible crown than normal-crowned spruce, it might be useful for crop tree ideotype breeding.     

Patterns in aboveground carbon allocation and tree architecture that favor stem growth in young Scots pine from high latitudes

The production and allocation of aboveground biomass and the characteristics of tree architecture were examined in eight-year-old Scots pine (Pinus sylvestris L.). Considerable among-tree variation existed in tree architecture, total aboveground dry mass production, and dry mass partitioning among tree parts. A linear relationship existed between needle and branch mass. Stem mass was directly proportional to tree height, which in turn was directly proportional to the allocation ratio between stem mass and total needle + branch mass production. The architectural characteristics that were related to a high proportional allocation to stem and high stemwood production were a large mean shoot volume, large mean number of branches per whorl, long needle retention and a high crown length/crown width ratio. Individual trees were found that combined high stemwood production with both high harvest index and high stemwood specific gravity.     

Crown architecture and stemwood production in Norway spruce (Picea abies (L.) Karst.)

The morphological data for Norway spruce (Picea abies (L.) Karst.) trees published by Burger (1953) were reanalyzed to examine the relationships between crown shape, needle and branch mass density of crown volume, needle efficiency in stemwood production, harvest index and stemwood production efficiency per unit of crown projected area. Production of stemwood per unit of crown projected area was higher in narrow-crowned trees than in broad-crowned trees because the narrow-crowned trees had (i) smaller horizontal space requirements for equal crown volumes, (ii) higher leaf area index due to both the geometrical crown shape and higher needle density per unit of crown volume, and (iii) higher harvest index indicating high allocation of dry matter production to stem.     

Within-crown structural variability of dwarfed mature Abies mariesii in snowy subalpine parkland in central Japan

We investigated mature dwarf Abies mariesii trees growing in conifer thicket–meadow parklands on a snowy subalpine plateau, where these dwarf trees are buried in the accumulated snow in winter. We focused on structural variation in the needles, shoots, and branchlets within different crown positions (leader crown vs lower crown) of the dwarf trees. In the leader crown, which appears above the snow surface earlier than the lower crown, current-year shoots and branchlets had greater total biomass, and foliage was more closely packed along the stem axis than in the lower crown, whereas current-year shoots in the leader crown had a lower needle mass ratio than in the lower crown. These results suggest that current-year shoots and branchlets in the leader crown have a specific structure that allows them to harvest more light, although construction and maintenance costs would be higher. In contrast, the structural characteristics of current-year shoots and branchlets in the lower crown efficiently concentrate incoming light by avoiding mutual shading within foliage, thus leading to increased biomass of photosynthetic needles within shoot and branchlet biomass. Such within-crown variability at various hierarchical levels from needles to branches in mature, but very dwarf, A. mariesii trees maintains the crown and allows survival within conifer clumps in areas of subalpine parklands that receive heavy snowfall.     

Needle and stem wood production in Scots pine (Pinus sylvestris) trees of different age,size and competitive status

We studied effects of tree age, size and competitive status on foliage and stem production of 43 Scots pine (Pinus sylvestris L.) trees in southern Finland. The tree attributes related to competition included foliage density, crown ratio and height/diameter ratio. Needle mass was considered to be the primary cause of growth through photosynthesis. Both stem growth and foliage growth were strongly correlated with foliage mass. Consequently, differences in growth allocation between needles and stem wood in trees of different age, size, or position were small. However, increasing relative height increased the sum of stem growth and foliage growth per unit foliage mass, indicating an effect of available light. Suppressed trees seemed to allocate more growth to stem wood than dominant trees, and their stem growth per unit foliage mass was larger. Similarly, trees in dense stands allocated more growth to stem wood than trees in sparse stands. The results conformed to the pipe model theory but seemed to contradict the priority principle of allocation.     

Generic biomass functions for Norway spruce in Central Europe--a meta-analysis approach toward prediction and uncertainty estimation

To facilitate future carbon and nutrient inventories, we used mixed-effect linear models to develop new generic biomass functions for Norway spruce (Picea abies (L.) Karst.) in Central Europe. We present both the functions and their respective variance-covariance matrices and illustrate their application for biomass prediction and uncertainty estimation for Norway spruce trees ranging widely in size, age, competitive status and site. We collected biomass data for 688 trees sampled in 102 stands by 19 authors. The total number of trees in the "base" model data sets containing the predictor variables diameter at breast height (D), height (H), age (A), site index (SI) and site elevation (HSL) varied according to compartment (roots: n = 114, stem: n = 235, dry branches: n = 207, live branches: n = 429 and needles: n = 551). "Core" data sets with about 40% fewer trees could be extracted containing the additional predictor variables crown length and social class. A set of 43 candidate models representing combinations of lnD, lnH, lnA, SI and HSL, including second-order polynomials and interactions, was established. The categorical variable "author" subsuming mainly methodological differences was included as a random effect in a mixed linear model. The Akaike Information Criterion was used for model selection. The best models for stem, root and branch biomass contained only combinations of D, H and A as predictors. More complex models that included site-related variables resulted for needle biomass. Adding crown length as a predictor for needles, branches and roots reduced both the bias and the confidence interval of predictions substantially. Applying the best models to a test data set of 17 stands ranging in age from 16 to 172 years produced realistic allocation patterns at the tree and stand levels. The 95% confidence intervals (% of mean prediction) were highest for crown compartments (approximately +/- 12%) and lowest for stem biomass (approximately +/- 5%), and within each compartment, they were highest for the youngest and oldest stands, respectively.     

Improvement in accuracy of aboveground biomass estimation in Eucalyptus nitens plantations: Effect of bole sampling intensity and explanatory variables

Two sets of aboveground biomass equations were fitted for stem only and stem plus crown predictive variables in Eucalyptus nitens plantations in Northern Spain. A sample of 40 trees was chosen after a complete study of variation in tree height and diameter in the region. The trees were felled and the biomass was divided into the following components: wood, bark, thick branches, thin branches, twigs, leaves and dead branches along the stem. Bole biomass was estimated by systematic subsampling of one 5 cm-thick disk every 0.5 m. Such intensive subsampling enabled determination of the effect of subsampling intensity on accuracy and bias of wood estimation, considering two ratio-type estimators: stem weight to dry matter, determined by the complete weighing (CW) method (i.e. of the fresh weight of the entire stem) and volume to dry matter, determined by the partial weighing (PW) method. The changes in moisture content and basic density along the stem explained the serious risk of dry mass or weight overestimation when a systematic subsample is considered. The average basic density was usually found at a relative height of 30-35% along the stem. The default choice of the bottom disk or log as the first section resulted in overestimations for the CW method and underestimations for the PW one. The biomass equations were fitted by seemingly unrelated regression, with corrections for heteroscedasticity carried out by weighted fitting. Diameter at breast height was the best explanatory variable, and the inclusion of height did not improve the accuracy, except for wood. The inclusion of crown variables improved the predictive ability for crown fractions, increasing the accuracy for estimating thick branches (by 10.8%), twigs (by 19.1%) and leaves (by 17.3%). The biomass of each fraction decreased in the following order: wood > bark > thick branches > dead branches along the stem > leaves > thin branches > twigs. The changes in these percentages with diameter class and the predictive ability of the fitted equations were also studied.     

Radiation-use efficiency and dry matter partitioning of a young olive (Olea europaea) orchard

Radiation-use efficiency (RUE) relates biomass production to the photosynthetically active radiation (PAR) intercepted by a plant or crop. We determined RUE and biomass partitioning coefficients of young olive (Olea europaea) trees for use in a general growth model. In 1995, 1-year-old olive trees var. 'Picual' were planted at a density of either 0.5 or 2.0 trees m(-2) near Córdoba, Spain, at a site providing favorable growth conditions. During the experiment (1995-1997), both PAR interception by the canopy and plant area index (PAI) were measured with radiation sensors. Regular harvests were performed to determine leaf area and biomass accumulation in roots, wood (stem, branches and trunk) and leaves. Leaf, wood and root biomass partitioning coefficients were calculated. The leaf area partitioning coefficients were also estimated. Dry matter production was linearly related to cumulative intercepted PAR. Seasonal RUE, calculated as the slope of the regression of aboveground biomass and cumulative intercepted PAR, was 1.35 g (MJ PAR)(-1). Radiation-use efficiency appeared to respond to environmental conditions, but was independent of planting density and PAI. The young olive trees allocated 0.26 of their total biomass to roots. Partitioning of aboveground dry matter was 0.60 to wood and 0.37 to leaves. As competition increased, dry matter partitioning to wood increased to 0.70.     

Genetic and environmental control of crown development in Picea sitchensis and its relation to stem wood production

Differences in stem wood production were found among four clones of Picea sitchensis (Bong.) Carr. at both an agricultural and a forest site. Clonal rankings were not consistent between the sites. Four variables were identified that may influence stem wood production, foliage amount, total aboveground production per unit foliage, proportional allocation of production to new foliage and to branch wood thickening. Comparison of clonal performance between sites showed that stem wood production could be influenced by differences in each of these variables. The four variables were themselves determined by components of crown structure, these were (i) production of new branches from the main stem, (ii) the numbers and lengths of branchlets supported on branches from the main stem, (iii) foliage production per unit branchlet length and foliage longevity, and (iv) branch wood thickening per unit branch length. These components varied both among clones and between sites. Branch production from the mainstem varied among clones, and for some clones it varied substantially between sites. In some clones branch production was positively correlated with mainstem height increment. There were differences among clones in the way that branchlet production varied between the agricultural and forest sites. In the absence of needle fall, foliage weight/branch length varied threefold among clones, but for each clone, varied little between sites. Branch wood weight/branch length was generally greater when foliage weight/branch length was large, but total branch wood increment/tree was strongly influenced by total branch length. The dynamics of crown development and its relation to stem wood production are discussed with reference to (i) the importance of needle longevity, (ii) the importance of tallness and narrowness of crowns and (iii) the importance of branch thickening, a process that competes with stem growth in the utilization of photosynthate. It is concluded that although each of these factors may be important, none singly accounts for the observed differences in stem wood production among genotypes and the way these change in response to the environment.     

Structural development of pinus sylvestris stands with varying initial density: A simulation model

The growth and structural development of Scots pine (Pinus sylvestris L.) trees growing at different spacing was simulated using a model based on the dry matter production per needle biomass unit and its allocation to needles, branches and stem. Special emphasis was given to the effect of stand density on the growth of the crown system and its implications on branchiness and timber quality. The simulations showed that the needle biomass culminates considerably earlier than the branch biomass with a time lag inversely related to the stand density. The lengths of living and dead crown were also inversely related to stand density. The resulting differences in branchiness were especially obvious in the early development of the tree stands. In the long run these differences tend to disappear, indicating equal external branchiness independently of the initial spacing for mature stands of Scots pine. The internal branchiness, however, was particularly sensitive to the initial spacing.     

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.     

黄连木苗期年生长节律及其生物量分配规律

为给黄连木幼苗生产管理提供参考,在定期观测的基础上,研究黄连木苗期年生长动态、生物量积累与分配规律以及植物生长指标与生物量组分的相关性。结果表明:在整个苗期生长过程中,黄连木的苗高、地径、冠幅生长均呈现"S"型曲线,用Logistic生长方程和生长曲线能对其动态生长节律进行很好地拟合。结合实际测量值将黄连木1年生幼苗的生长过程分为4个阶段:出苗期、生长初期、速生期和生长末期。各指标速生期的生长量占其总生长量50%以上;黄连木1年生苗木的地上与地下部分生物量比例为1.78∶1.00;苗木生物量各指标中,地径与苗高、茎干质量、叶干质量显著相关,苗高与茎干质量、叶干质量显著相关,茎干质量与叶干质量极显著相关。     

Carbon budget for Scots pine trees: effects of size, competition and site fertility on growth allocation and production

Time series of carbon fluxes in individual Scots pine (Pinus sylvestris L.) trees were constructed based on biomass measurements and information about component-specific turnover and respiration rates. Foliage, branch, stem sapwood, heartwood and bark components of aboveground biomass were measured in 117 trees sampled from 17 stands varying in age, density and site fertility. A subsample of 32 trees was measured for belowground biomass excluding fine roots. Biomass of fine roots was estimated from the results of an earlier study. Statistical models were constructed to predict dry mass (DW) of components from tree height and basal area, and time derivatives of these models were used to estimate biomass increments from height growth and basal area growth. Biomass growth (G) was estimated by adding estimated biomass turnover rates to increments, and gross photosynthetic production (P) was estimated by adding estimated component respiration rates to growth. The method, which predicts the time course of G, P and biomass increment in individual trees as functions of height growth and basal area growth, was applied to eight example trees representing different dominance positions and site fertilities. Estimated G and P of the example trees varied with competition, site fertility and tree height, reaching maximum values of 22 and 43 kg(DW) year(-1), respectively. The site types did not show marked differences in productivity of trees of the same height, although height growth was greater on the fertile site. The G:P ratio decreased with tree height from 65 to 45%. Growth allocation to needles and branches increased with increasing dominance, whereas growth allocation to the stem decreased. Growth allocation to branches decreased and growth allocation to coarse roots increased with increasing tree size. Trees at the poor site allocated 49% more to fine roots than trees at the fertile site. The belowground parts accounted for 25 to 55% of annual G, increasing with tree size and decreasing with site fertility. Annual G and P per unit needle mass varied over the ranges 1.9-2.4 and 3.5-4.0 kg(DW) kg(-1), respectively. The relationship between P and needle mass in the example trees was linear and relatively independent of competition, site fertility and age.     

Effect of wide spacing on increment and branch properties of young Norway spruce

The effects of stand density on increment and branch properties were studied in three spacing experiments of Norway spruce [Picea abies (L.) Karst.]. The stand densities ranged from 350 stems ha⁻¹, regarded as open-grown trees, up to 1,600 stems ha⁻¹, corresponding to the density recommended for forestry practice. Properties of all the branches were measured from the stem apex downwards. The study material included a total of 5,661 branches from 45 trees. Increasing stand density resulted in a decrease in radial increment as well as shorter and narrower crowns, but it had no effect on height increment. The average number of spike knots per tree was 0.87, 0.27, and 0.33 in densities of 350, 700 and 1,600 ha⁻¹, respectively. Additionally, in the widely spaced stands of 350 stems ha⁻¹, the fraction of trees having spike knots was high (over 50%). At a density of 1,600 ha⁻¹, the sample trees had somewhat less branches in a whorl compared with the more widely spaced plots. The most pronounced effect of stand density was the increase in branch diameter with decreasing stand density. At a density of 350 ha⁻¹, the maximum branch diameter of all the sample trees exceeded the diameter limit of quality class B in the European quality requirements for round wood. The results give some indication that trees subjected to severe competition would produce smaller branches per unit of crown projection area. However, the possibilities for reducing branch dimensions relative to stem and crown size through competition appear quite restricted.     

Biomass distribution in 40-year-old trees of Japanese black pine

Measurements were carried out to survey the quantity of above- and below-ground biomass and its distribution of five Japanese black pines (Pinus thunbergii Parl.) growing on a sandy soil. The roots, divided into diameter groups, were surveyed using two methods—soil coring and excavation. Average dry weight of total biomass of the trees was 176,185 g. Roots represented 13.2%, below-ground stump 6.5%, stem 70.4% and branches with needles 9.9% of total biomass. Roots made up about two thirds and stump one third of below-ground biomass. Total length of below-ground biomass (except roots with diameter < 0.1 cm) was 479.1 m/tree. Roots with diameter of 0.1–0.2 cm represented only 0.7% of below-ground biomass, however as much as 49.9% of their total length. Roots with diameter over 10.0 cm constituted as much as 21.6% of below-ground biomass, however were only 0.3% of its total length. Root systems had well developed tap roots to maximal depth of 231 cm. The results indicated that mass and length of roots with diameter 0.5–2.0 cm had a close correlation with branch mass. Mass and length of roots with diameter 2.0–10.0 cm closely correlated to stem mass. Stem mass, root mass and root length closely correlated to DBH. A rather low correlation was found between DBH and mass of branches and below-ground stump. DBH was a suitable variable for predicting total biomass.     

Allometric equations for aboveground biomass estimation by size class for Pinus brutia Ten. trees growing in North and South Aegean Islands, Greece

Empirical allometric equations relating biomass of aboveground components to dendrometric variables for Pinus brutia Ten. trees are derived in this paper. They are based on data collected from Lesvos (North Aegean Sea) and Crete (South Aegean Sea) Islands. Comparisons to published equations for the same species growing in northwestern and southeastern Turkey, for Pinus nigra A. growing in Turkey and Pinus halepensis Mill. found in Western Aegean (island of Evia), are also presented. The biomass of branches from destructively sampled trees (twelve in Crete and six in Lesvos) was divided into four size classes (0?C0.63 cm, 0.64?C2.5 cm, 2.51?C7.61 cm, and 7.62?C22.8 cm). Tree crown biomass was calculated as the sum of the biomass in the four classes plus the fraction of stem above crown base. Over bark stem biomass was estimated through bole volume conversion based on wood density. The results showed clearly that, for a given diameter, the Cretan trees had more crown biomass and a higher share of small branches than trees on Lesvos, probably due to differences in environment and stand structure. Comparisons to published diameter versus crown biomass equations reveal a lower crown biomass for Turkish sites of Calabrian pine and Aleppo pine on Evia Island, while only Turkish Black pine seems to be comparable to the Calabrian pine on Crete. The derived allometries can be used for landscape fire behavior modeling, for ecophysiological studies and for the Kyoto protocol requirements of carbon changes in Pinus brutia Ten. forests located in northern and southern Greek sites.     

Assessment of crown woody biomass in <Emphasis Type="Italic">Eucalyptus grandis</Emphasis> and <Emphasis Type="Italic">E. globulus</Emphasis> plantations

Young trees were harvested to explore non-destructive methodologies to estimate live branch dry weights in young fast-growing Eucalyptus species under different spacing and fertilizer treatments. Branch growth can vary with silvicultural management such as spacing, fertilizing and thinning, and over relatively short periods in response to environmental conditions. Many published regressions based on standard measurements of height and diameter are site, age and treatment specific. The aim of this study was to improve our capacity to predict woody crown dry weight, based on stem measurements, and to minimize (or eliminate) treatment effects on the resulting model. In young trees, branches are temporary support structures for foliage and are often discarded as the base of the green crown rises. As temporary structures they represent an investment of biomass and nutrient elements, and are subject to selection pressures to maximize the return on investment by the tree. Trees were harvested from existing plantation experiments located in south-eastern Queensland for E. grandis W. Hill ex Maiden (ranging from 0.28 to 15.85 m in height, to 5 years old) and south-western Australia for E. globulus Labill. (0.10–34.4 m in height, to 10.2 years) in order to examine the impact of spacing, nitrogen and phosphorus fertilization on early growth. Relationships to estimate crown woody biomass from non-destructive measurements were developed, and these relationships tended to have different slopes and intercepts for trees with predominantly juvenile foliage and those with intermediate or adult foliage. Dry weight of whole-crown live branch wood (W_branch) was related to heights and/or diameter at breast height (D_BH), but the regressions parameters were different, depending on treatment. The relationships became more generic (i.e. less dependent on treatment effects) between W_branch and stem sectional area at the height of the base of the green crown (SA_CB), consistent with the pipe model theory (R² > 0.91 for the two species for trees with intermediate/adult leaves). However, W_branch was more closely related again to the stem volume above the base of the green crown and treatment effects were not significant (V_Con,gc, R² > 0.93). Branches exit the stem below the green crown, and for E. grandis the best relationship was on stem volume above the lowest live branch (V_Con,llb, R² 0.94). Limited sampling from four other species with similar or contrasting crown characteristics indicated that the relationship could be applied quite generally. Individual E. grandis branch woody dry weight was closely related to the conical volume of the main (first order) branch (V_con,br, R² 0.98₎. The whole crown equivalent, branch woody dry weight plus stem dry weight above the lowest live branch, was also closely related to the stem volume within the woody crown (V_Con,llb, R² 0.97–0.99). While the slope of this relationship was still significantly different between trees with juvenile and intermediate/adult foliage, it had a similar form, suggesting that trees with juvenile foliage allocated a different proportion of their woody biomass within the crown to branches than older trees.     

Modelling the economically viable wood in the crown of European beech trees

Long-term forest development programs in Germany aim on an increase of close-to-nature broadleaf forest stands. This means that the economic importance of European beech is expected to increase. The economic potential of a tree basically consists of the stem as well as the economically viable wood volume in the crown. Due to the high morphological variability of European beech crowns, taper models are often not satisfactory for predicting the economically viable wood volume arising from crowns. Prediction models with a higher precision are recently still lacking. Aim of this study is thus the development of prediction model for the economically viable crown wood volume of European beech trees.We determined the distribution of the wood volume in the crown over the branch diameters using the multistage ‘randomized branch sampling’ method (RBS). The tree-specific wood volume distribution on the branch diameters were used to cluster all sampled trees into 3 groups. Additionally, we developed a method able to distinguish between economically viable and unviable crown branches. Basing on the RBS measurements as well as revenues and processing costs, we modeled the economically viable wood volume from the crown for each tree. To calculate the wood volume under bark, we parameterized a bark thickness function from disk samples of the trees.We showed that the European beech crowns could be clustered into 3 groups differing in their wood volume distribution. The economically viable wood volume in the crown significantly depended on this grouping parameter as well as diameter at breast height (DBH). By contrast, the total amount of wood in the crown only depended on DBH. The differing viable wood volumes in the crowns were thus explained by different wood distributions and not by differing total crown wood volume. To make the results applicable in practice forestry, the modelling results were used to develop a regression formula able to predict the economically viable wood volume in the crown depending on the DBH and the crown type. As the crown type can also be predicted via measurable tree covariates, the regression model of the viable wood volume in the crown can be used as a support tool for the management of European beech stands. Sensitivity analysis quantifies how harvest revenues and costs translate into different viable tree volume.     

云南省云杉立木生物量模型研建

以云南云杉为研究对象,对云杉地上生物量和地下生物量模型进行研建。建立云杉地上总生物量、树干、树冠、干材、干皮、树枝、树叶独立模型与材积相容模型,采用分级联合控制和度量误差模型方法,建立地上总生物量和所有分量相容的立木生物量模型,建立根茎比模型对云杉地下生物量进行估计。结果表明：建立的云杉地上总生物量、树干、干材生物量二元模型预估精度均达95％以上,干皮生物量模型预估精度达94％以上,树冠、树叶、树枝生物量的预估精度均在92％以上,地下生物量模型预估精度在88％以上;所建立的模型可以用于云杉生物量的估计。     

Relationships between crown projected area and components of above-ground biomass in Norway spruce trees in even-aged stands: Empirical results and their interpretation

Sample tree material was reanalyzed in order to study the relationships between horizontal crown projected area and components of above-ground biomass in Norway spruce (Picea abies (L.) Karst.) trees growing in even-aged stands. The needle mass of dominant trees increased linearly with the increase in crown projected area, but in co-dominant and dominanted trees the increase in needle mass levelled off toward larger crown projected areas. The branch mass of dominant and co-dominant trees accumulated faster than linearly with increasing crown projected area, whereas in dominated trees an approximately linear relationship existed between these two variables. The increase in needle and branch mass per unit increase in crown projected area was highest in dominant trees and decreased to co-dominant and dominated trees, i.e. with tree position in the canopy. The stem mass accumulated obviously faster than linearly and similarly in all tree classes with the increase in crown projected area. The narrow crown shape indicated a high density of all components of above-ground biomass per unit of crown projected area.