Current issue: 53(1)

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Silva Fennica 1926-1997
Acta Forestalia Fennica

Articles containing the keyword 'stand density'.

Category: Research article

article id 1141, category Research article
Isabel Miranda, Jorge Gominho, Helena Pereira. (2015). Heartwood, sapwood and bark variation in coppiced Eucalyptus globulus trees in 2nd rotation and comparison with the single-stem 1st rotation. Silva Fennica vol. 49 no. 1 article id 1141.
Highlights: Coppiced E. globulus trees in the 2nd rotation have similar heartwood and sapwood development as single-stem trees in the 1st rotation; The initial tree planting density did not influence heartwood development of coppiced E. globulus trees; Heartwood diameter and height can be modelled with tree diameter and height respectively; Sapwood width is approximately constant within and between coppice and single-stem E. globulus trees.
Coppiced Eucalyptus globulus trees with 18 years in a 2nd rotation were analysed in relation to heartwood, sapwood and bark content taking into account the effect of the initial planting density by using a spacing trial. A total of 25 stumps, with a variable number of stems per stump from 1 to 3, were analysed. Comparison was made to the previous 1st rotation single stem trees, also harvested at 18 years. In the 2nd rotation, the stump density did not significantly affect stem height and diameter, in opposition to the 1st rotation where spacing significantly impacted on tree dimensions. The effect of the initial planting density is somewhat lost in the coppiced stand in relation with i.e. the number of stems per stump. Heartwood was present in all the coppiced trees up to 49.9% of the total tree height and heartwood volume amounted to 38.9–51.7% of the total tree volume. Within the tree, heartwood content decreased from the base upwards, representing, on average, 54.1% at the base and decreasing to 5.1% at 15.3 m. The sapwood width remained relatively constant with an average radial width of approximately 2 cm. The average stem bark content of coppiced trees was 17.4% of the total stem volume. The comparison of heartwood and sapwood development in the coppiced trees did not show significant differences to the 1st rotation trees, nor did the initial spacing. Heartwood diameter could be modelled using the tree diameter both for 1st and 2nd rotation trees.
  • Miranda, Centro de Estudos Florestais, Instituto Superior de Agronomia, Universidade Lisboa, 1349-017 Lisboa, Portugal ORCID ID:E-mail:
  • Gominho, Centro de Estudos Florestais, Instituto Superior de Agronomia, Universidade Lisboa, 1349-017 Lisboa, Portugal ORCID ID:E-mail: (email)
  • Pereira, Centro de Estudos Florestais, Instituto Superior de Agronomia, Universidade Lisboa, 1349-017 Lisboa, Portugal ORCID ID:E-mail:
article id 1002, category Research article
Pentti Niemistö. (2013). Effect of growing density on biomass and stem volume growth of downy birch stands on peatland in Western and Northern Finland. Silva Fennica vol. 47 no. 4 article id 1002.
Highlights: The thinning response in young downy birch stands was low and the mortality of merchantable stems was tolerable even in dense unthinned thickets during the rotation of 50 years; The production of above-ground woody biomass and stem volume or even the production of pulpwood increased with stand density, with these values being greatest for unthinned or very lightly thinned plots.
The objective of the study was to ascertain the relationship of thinning intensity of downy birch (Betula pubescens Ehrh.) stands with height, crown, and diameter development as well as pulpwood, stem volume, and biomass increment using long-term (20−30 years) field experiments. Diameter growth of birches increased with thinning intensity during the first 15 years from thinning in all development phases, though after that it did so only for the youngest stands. The thinning response was low. Thinning intensity had no influence on increase in height. In terms of stem volume with bark, the mortality in unthinned stands during the study period was 30–45 m3 ha-1. The mean stem number in unthinned birch thickets fell from 25 000 ha-1 at a dominant height of 7 m to 3000 ha-1 at 18 m. The stem volume increment over the first 15 years was highest (5–6 m3 ha-1 a-1) on the very lightly thinned or unthinned plots, but later there was no significant difference between initial thinning intensities. The maximum above-ground leafless biomass (over 100 Mg ha-1) was achieved on very lightly thinned plots. Also, the total production (including thinning removal) of biomass or stem volume or even the production of pulpwood increased with stand density, with these values being greatest for very lightly thinned or unthinned plots. During 50-year rotation, the highest leafless above-ground biomass production was 2.5 Mg ha-1 a-1 as a mean value from the experiments. The highest mean annual production of pulpwood (d > 6.5 cm) was 3.2 m3 ha-1 a-1, and, in practice, no saw timber or veneer timber was produced, because of the small size and low quality of the stems. A thinning in downy birch stands increased slightly the size of stems to be removed in future cuttings, but with exception for very light thinning it decreased the production of biomass and merchantable wood.
  • Niemistö, Finnish Forest Research Institute, Parkano Unit, FI-39700 Parkano, Finland ORCID ID:E-mail: (email)
article id 938, category Research article
Lars Karlsson, Tommy Mörling, Urban Bergsten. (2013). Influence of silvicultural regimes on the volume and proportion of juvenile and mature wood in boreal Scots pine. Silva Fennica vol. 47 no. 4 article id 938.
Highlights: Initial stand densities have a large impact on the proportion of mature wood within trees throughout most of their life cycle; The difference between regimes in volume of long fibres in crop trees could be substantial; Long-term silvicultural strategies implemented at juvenile stand ages can be important tools in order to produce wood raw material suited for specific end-uses.
Trees from 48 to 78 years old, exposed to three different long-term silvicultural regimes, were examined for transition ages between juvenile (JW), transition (TW) and mature wood (MW), total wood volume and proportions of the same wood types, as defined by fibre length. Twenty one sample trees were collected at sites with similar growing conditions within the same geographical area. Stem discs and fibre samples from breast height (BRH), 20% of tree height, green crown height and 70% of tree height were analysed. Trees growing in an environment with few neighboring trees (Sparse regime) started to produce MW, on average, five years later at BRH and six to nine years later at 20% of total tree height than trees in stands with high stem numbers (Dense regime) and trees growing in stands where the stem number had been heavily reduced after an initial high stand density (Dense/Sparse regime). For all regimes, the greatest mean fibre length was found below the green crown and high initial stem numbers were found to positively influence fibre length. The proportion of MW in the whole stem was 34% at Sparse regime sites, 57–69% at Dense/Sparse sites and 63–64% in sites where there was a Dense regime. The proportion of MW was significantly lower for trees from the Sparse regime on each stem section compared. In conclusion, the results suggest that the initial condition a tree faces affects the stem wood properties and quality output at the end of the rotation period.
  • Karlsson, Department of Forest Biomaterials and Technology, Swedish University of Agricultural Sciences, SE-901 83 Umeå, Sweden ORCID ID:E-mail: (email)
  • Mörling, Department of Forest Ecology and Management, Swedish University of Agricultural Sciences, SE-901 83 Umeå, Sweden ORCID ID:E-mail:
  • Bergsten, Department of Forest Biomaterials and Technology, Swedish University of Agricultural Sciences, SE-901 83 Umeå, Sweden ORCID ID:E-mail:
article id 404, category Research article
William L. Mason, Colin Edwards, Sophie E. Hale. (2004). Survival and early seedling growth of conifers with different shade tolerance in a Sitka spruce spacing trial and relationship to understorey light climate. Silva Fennica vol. 38 no. 4 article id 404.
Alternative silvicultural systems to clearfelling are being adopted in Great Britain as a means of increasing the species and structural diversity of conifer plantation forests. One area where knowledge is lacking is the critical level of below-canopy light for survival and growth of young seedlings. This was investigated by planting seedlings of European larch Larix decidua (Mill.), Scots pine Pinus sylvestris L., Sitka spruce Picea sitchensis (Bong.(Carr.)), Douglas fir Pseudotsuga menziesii (Mirb.(Franco.)), and western hemlock Tsuga heterophylla (Raf. (Sarg.)) in a Sitka spruce plantation thinned to 3 different spacings. The incident light intensity beneath the canopy ranged from about 2 to over 60 per cent of full light. Planting in an adjoining open area provided an indication of growth under full light. Growth and survival of these seedlings were followed for 4 growing seasons. The highest seedling survival was found under the widest spacing and declined with closer spacing and lower light intensity. Only Douglas fir and western hemlock seedlings survived at the closest spacing, and in low percentages. The tallest seedlings of each species were found in the open grown conditions but survival was variable due to increased weed competition. Species-specific growth responses showed little difference under high light conditions but performance at low light was generally consistent with shade tolerance rankings in the literature except that Sitka spruce shade tolerance was slightly lower than expected. Minimum light requirements for these species increased from 10 to 30 per cent of full light with decreasing shade tolerance. Other studies of incident light in Sitka spruce plantations indicated that target basal areas in the range 25–30 m2 ha–1 are required if these light conditions are to be met, which suggests an irregular shelterwood system with frequent interventions should be favoured.
  • Mason, Forest Research, Northern Research Station, Roslin, Midlothian, Scotland, UK, EH25 9SY ORCID ID:E-mail: (email)
  • Edwards, Forest Research, Northern Research Station, Roslin, Midlothian, Scotland, UK, EH25 9SY ORCID ID:E-mail:
  • Hale, Forest Research, Northern Research Station, Roslin, Midlothian, Scotland, UK, EH25 9SY ORCID ID:E-mail:
article id 511, category Research article
Luis Diego Pérez Cordero, Markku Kanninen. (2003). Heartwood, sapwood and bark content, and wood dry density of young and mature teak (Tectona grandis) trees grown in Costa Rica. Silva Fennica vol. 37 no. 1 article id 511.
The aim of this study was to evaluate the heartwood, sapwood and bark content, and wood dry density in young and mature teak (Tectona grandis) trees. For this, 17 plantations were selected from 11 sites representing different climatic conditions and plantation densities (156 to 1600 trees ha–1, and line planting). From these plantations, a total of 87 trees with ages between 5 and 47 years were felled for stem analysis. The highest heartwood proportion of stem volume (over bark) was 61% and the lowest 0.4%. The sapwood proportion ranged between 24 and 72%, while bark represented from 14 to 37% of the total volume. Heartwood proportion was significantly different (P < 0.05) among climatic zones: ‘wet’ sites producing less heartwood than ‘dry’ sites. Stem diameter (under bark) and heartwood diameter at different stem heights differed among sample trees, even when plotted in relative values to avoid dependency with stem size. Dry density was statistically different between 8-year-old trees or younger and 47-year-old trees, and between line planting trees and 13-year-old trees or younger, but did not differ statistically between line planting trees and mature trees. No significant differences were found between climatic zones or between different stand densities. Dry density values for T. grandis plantations in Costa Rica are similar to those reported elsewhere.
  • Pérez Cordero, Centro Agronómico Tropical de Investigación y Enseñanza (CATIE), Turrialba, Costa Rica ORCID ID:E-mail: (email)
  • Kanninen, Center for International Forestry Research (CIFOR), Bogor, Indonesia ORCID ID:E-mail:
article id 663, category Research article
Lennart Moberg. (1999). Variation in knot size of Pinus sylvestris in two initial spacing trials. Silva Fennica vol. 33 no. 2 article id 663.
The objective of this study was to investigate the variation in internal knot size of Scots pine (Pinus sylvestris L.) stems sampled from mature permanent plots, and for which the silvicultural history was known. It was based on a sample of mature trees removed from two different spacing trials representing a moderate and high site index. Knot size was measured with non-destructive methods using a CT-scanner and digital image analysis. Initial spacing varied between 0.75 and 3 m on the high site-index trial and between 1.5 and 2.5 m on the moderate site-index trial. Wider initial spacing on the high site index resulted in larger knots near the base of the stem. However, due to successive thinnings which gradually equalised stand density among plots, the difference between most plots was less further up in the stems. The effect of silvicultural regime was much more limited on the lower site index. Within-stand differentiation resulted in a variation of tree diameter (DBH); larger trees had significantly larger knots. Furthermore, knots were larger towards south than towards north in both trials. These results illustrated that, by using non-destructive measurements on trees sampled from permanent research plots, it was possible to simultaneously study the variation of internal knot size at stand (such as site and silviculture effects), within-stand (such as relative tree size) and within-tree levels (such as height and azimuth). However, lack of replication prevented valid statistical inference as to stand-level effects.
  • Moberg, Swedish University of Agricultural Sciences, Department of Forest Management and Products, P.O. Box 7060, S-750 07 Uppsala, Sweden ORCID ID:E-mail: (email)

Category: Article

article id 7164, category Article
Veijo Heiskanen. (1965). Puiden paksuuden ja nuoruuden kehityksen sekä oksaisuuden ja sahapuulaadun välisistä suhteista männiköissä. Acta Forestalia Fennica vol. 80 no. 2 article id 7164.
English title: Relation between the development of the early age and thickness of trees and their branchiness in Scots pine stands in Finland.

The objective of the study was to establish the influence of the founding density of a stand and the intensity of intermediate cutting on the quality of pine saw logs stems, primarily on their branchiness. Measurements were carried out in 68 Myrtillus-type and 32 Vaccinium-type Scots pine (Pinus sylvestris L.) stands. The quality of 1,982 sample trees was assessed.

According to the results, the branchless part of the stem is longest in the older age classes of trees. In all age classes the percentage of the branchless part is highest in medium sized stems. The relative height of the crown limit is greatest in small diameter classes and continues as the thickness of the tree increases. The crown is longer in the thicker tree. The grade of the butt log is on average highest in medium sized stems. Knottiness of a log made it unsuitable for a saw log only among the thickest stems. The relative share of the u/s grade decreased as the thickness of the trees increased.

From the point of view of early development of the trees it was concluded that in all age classes the branchless part is the shorter the faster the tree has grown in diameter when it was young. Also, branches of the butt log are the bigger the faster the tree has developed when it was young. The grade of the butt log improves as the thickness of the annual rings diminishes.

To produce good quality sawn timber, the pine stands should be established dense, and the first thinnings should be delayed as much as possible. The best time for the thinning would be when the diameter of the dominant trees at stump height is 12–15 cm and when all the branches have died on the length of the butt log. After the first thinning, comparatively intense intermediate thinning may be applied.

The PDF includes a summary in English.

  • Heiskanen, ORCID ID:E-mail:
article id 5608, category Article
Harri Mäkinen. (1997). Possibilities of competition indices to describe competitive differences between Scots pine families. Silva Fennica vol. 31 no. 1 article id 5608.

Possibilities of distance-independent and -dependent competition indices to describe the competition stress of an individual tree was studied in Southern Finland. Five half-sib open-pollinated families and one check lot of Scots pine (Pinus sylvestris L.) was used as study material in order to analyse competitive interactions of crown form and stand density variation. Almost all competition indices correlated strongly with radial increment. Thus distance-independent indices were adequate to describe competition in young row plantations, where distance effects between trees were implicitly eliminated. Correlations between indices and height increment were not significant. Along with the increase in competition, the width and length of the crown and the diameter increment of the stem of some narrow-crowned families decreased slowly compared to wide-crowned families.

  • Mäkinen, ORCID ID:E-mail:
article id 5490, category Article
Timo Pukkala, Jari Karsikko, Taneli Kolström. (1992). A spatial model for the diameter of thickest branch of Scots pine. Silva Fennica vol. 26 no. 4 article id 5490.

The model predicts the base diameter of the thickest living branch of a tree growing in a planted or naturally regenerated even-aged stand. A mixed model type was used in which the residual variation was divided into within-stand and between-stand components. The study material consisted of 779 trees measured in 12 plots located in 20 to 35 years old Scots pine (Pinus sylvestris L.) stands (breast height age 10 to 20 years). Branch diameter was closely connected to the breast height diameter of the stem. In a stand of a certain age, competition by close neighbours slightly decreased branch diameter in a given diameter class. According to the model, the greatest difference is between trees subjected to very little competition and those subjected to normal competition. The model was used in simulated stands with varying age, density, and tree arrangement. The simulations showed that trees with rapid diameter growth at young age had thicker branches at a given breast height diameter than trees with slower diameter growth. However, a very slow growth rate did not produce trees with branches thinner than those possessing a medium growth rate.
The PDF includes an abstract in Finnish.

  • Pukkala, ORCID ID:E-mail:
  • Karsikko, ORCID ID:E-mail:
  • Kolström, ORCID ID:E-mail:
article id 5302, category Article
Seppo Kellomäki, Heikki Hänninen, Taneli Kolström, Ahti Kotisaari, Timo Pukkala. (1987). A tentative model for describing the effects of some regenerative process on the properties of natural seedling stands. Silva Fennica vol. 21 no. 1 article id 5302.

The effect of the size of seed crop, dispersal of seeds and the early development of seedlings on the density and spatial distribution of young Scots pine (Pinus sylvestris L.) stands are evaluated on the basis of theoretical models. The models include (i) number and spatial distribution of parent trees on the regeneration area, (ii) size of annual seed crop, (iii) seed dispersal from a particular parent tree, (iv) germination of the seeds (germination percentage), (v) death of ageing seedlings after the establishment process, and (vi) height growth of the seedlings.

As expected, stand density and spatial distribution varied within a large range in relation to the density of the parent trees and the distance from them. The simulations also showed that natural seedling stands can be expected to be heterogenous due to the geometry of seed dispersal, emphasizing the frequency of young and small trees. The properties of the seedling stands were, however, greatly dependent on the density of the parent trees and the length of the regeneration period.

The PDF includes an abstract in Finnish.

  • Kellomäki, ORCID ID:E-mail:
  • Hänninen, ORCID ID:E-mail:
  • Kolström, ORCID ID:E-mail:
  • Kotisaari, ORCID ID:E-mail:
  • Pukkala, ORCID ID:E-mail:
article id 5207, category Article
Seppo Kellomäki. (1984). Havaintoja puuston kasvatustiheyden vaikutuksesta mäntyjen oksikkuuteen. Silva Fennica vol. 18 no. 2 article id 5207.
English title: Observations on the influence of stand density on branchiness of young Scots pines.

The study based on young Scots pine (Pinus sylvestris L.) of varying density showed that number of living branches per whorl and total number of living branches per tree were negatively correlated with stand density. On the contrary, the number of dead branches increased with increasing stand density. The diameter of living and dead branches decreased with increasing stand density. Consequently, the branchiness, i.e. the share of the branch cross-sectional area from the surface area of the stem, decreased in dense stands compared with the thin stands. At the densest stands the branchiness, however, levelled of indicating a greater decrease of the radial growth at stems than at branches. The 2/3 power law described relatively well the relationship between stand density and mean squared branch diameter of living branches.

The PDF includes a summary in English.

  • Kellomäki, ORCID ID:E-mail:
article id 5200, category Article
Seppo Kellomäki, Timo Nevalainen. (1983). Näkökohtia puuston tiheyden ja puiden koon välisestä suhteesta. Silva Fennica vol. 17 no. 4 article id 5200.
English title: Relationship between stand density on tree size.

Two Japanese models regarding the within-stand competition have been reviewed on the basis of relevant literature. Competition-density and 3/2 th power models seem to be applicable also into tree stands. The latter model has been applied into the material obtained from literature. Computations showed consistancy with the results obtained elsewhere in the world. It is concluded that also in Finnish conditions the 3/2 th power law may have great potentials in describing the effects of stand density on tree size.

The PDF includes a summary in English.

  • Kellomäki, ORCID ID:E-mail:
  • Nevalainen, ORCID ID:E-mail:
article id 5111, category Article
Seppo Kellomäki. (1981). Effect of the within-stand light conditions on the share of stem, branch and needle growth in a twenty-year-old Scots pine stand. Silva Fennica vol. 15 no. 2 article id 5111.

The share of stem, branch and needle growth was dependent on the within-stand light regime in a young Scots pine (Pinus sylvestris L.) stand. The share of needle growth increased at the expense of stem and branch growth in poor light conditions. In good light condition the share of branch wood increased substantially. The share of stem wood growth was greatest in moderate shading, emphasizing the role of an adequate stand density for growing high-quality timber. The basic density of the stem wood was considerably greater in suppressed trees than in dominating trees. The differences were related to the illumination of the crown system.

The PDF includes a summary in Finnish.

  • Kellomäki, ORCID ID:E-mail:
article id 5079, category Article
Seppo Kellomäki, Pertti Hari, Pirkko Ilonen, Markku Kanninen. (1980). Eco-physiological studies on young Scots pine stands. II. Silva Fennica vol. 14 no. 3 article id 5079.

The technique of double normalizing, i.e. normalizing the relative needle biomass and the length of the living crown system, is applied to the modelling of the distribution of needle biomass in the canopy of young Scots pine (Pinus sylvestris L.) stands. The study based on the parameters of β-function shows that at the individual-tree level, the variance in needle distribution was not closely associated with any tree characteristics. A shift in the point of maximum needle biomass upwards unsuppressed trees was, however, evident. This was associated with an increase in the height of the trees. At the stand level, the stand mean height and stand density had an equal and a rather high potential for explaining the variance in the needle distribution. The normalized crowns are utilized in models for determining light extinction in the crown. A special technique for determining the amount of photosynthates available for growth in a particular tree is presented.

The PDF includes a summary in Finnish.

  • Kellomäki, ORCID ID:E-mail:
  • Hari, ORCID ID:E-mail:
  • Ilonen, ORCID ID:E-mail:
  • Kanninen, ORCID ID:E-mail:
article id 4964, category Article
Seppo Kellomäki, Pertti Hari, Eero Väisänen. (1977). Annual production of some forest mosses as a function of light available for photosynthesis. Silva Fennica vol. 11 no. 1 article id 4964.

The aim of the present paper was to study the annual production of Pleurozium schreberi (Brid.) Mitt., Hylocomnium splendens (Hedw.) B.S.G and Dicranum polysetum Sw. as a function of light available for photosynthesis. The productivity of the above moss species is studied using the harvested quadrats method in Norway spruce (Picea abies) stands of the Myrtillus site type representing different stand density classes (basal area from 0 to 34 m2/ha) in Southern Finland.

The annual production of each species in different stands was correlated with the amount of light available for photosynthesis i.e. with the photosynthetic production. Functions for the dependence of productivity on light conditions were produced for each species. The individual functions and their ecological significance is discussed. The adaptation of each species to low light intensity is evident since no meaningful addition to production takes place when the photosynthetic light ratio reaches values greater than 0.3–0.4. In other words, the level of photosynthesis which is 30–40% of that possible in the open, provides sufficient supply of carbohydrates or the basic functions of the moss species studied. Pleurozium schreberi and Dicranum polysetum seem to have greater light requirements than Hylocomnium splendens.

The PDF includes a summary in Finnish.

  • Kellomäki, ORCID ID:E-mail:
  • Hari, ORCID ID:E-mail:
  • Väisänen, ORCID ID:E-mail:
article id 4784, category Article
Matti Leikola, Pentti Pylkkö. (1969). Verhopuuston tiheyden vaikutus metsikön minimilämpötiloihin hallaöinä. Silva Fennica vol. 3 no. 1 article id 4784.
English title: Influence of stand density on the minimum temperatures during frost nights.

The objective of this investigation was to study the influence of stand density of white birch (Betula pubescens Ehrl.) on the minimum temperatures in the stand during the growing season, and the actual minimum temperatures of the leading shoot of Norway spruce (Picea abies (L.) H. Karst.) seedlings growing in the open. The 40-year-old uniform white birch stand was situated in 142 m above the sea level in Southern Finland. The stand was treated with thinnings of three different densities in 1961.

Air temperature was recorded in four sample plots at heights of 0.1 m, 0.5 m, 1.0 m, 2 m and 4 m. In the stand of moderate density, temperatures were measured at heights of 6.0 m, and in the stand of full density at 6.0 m, 8.0 m and 10.0 m.

The temperature differences between stands of various densities proved to be rather small. Especially the thinnest stand differed very little from the open area. The soil surface has in all cases been warm compared with the higher air layers indicating meadow-fog-type by Geier (1965). On cloudy or windy weather all the temperature profiles in the various stands resembled each other. The difference between the air temperature and temperature of the spruce shoot was greatest at midnight and decreased steadily thereafter.

The problem in using shelter stands for spruce regeneration areas is that optimum shelter stand density is difficult to define. Already a thin shelter stand causes drawbacks to the young seedlings, but in order to be effective enough against early frosts, the shelter stand should be comparatively dense.

The PDF includes a summary in English.

  • Leikola, ORCID ID:E-mail:
  • Pylkkö, ORCID ID:E-mail:
article id 7677, category Article
Risto Heikkilä, Timo Mikkonen. (1992). Effects of density of young Scots pine (Pinus sylvestris) stand on moose (Alces alces) browsing. Acta Forestalia Fennica no. 231 article id 7677.

The study was carried out at Padasjoki, Southern Finland, where moose (Alces alces L.) density on the winter range had been over 1.5 individuals/km2. Moose browsing intensity, expressed in terms of number of twigs eaten and biomass used, increased with stand density (biomass). Total biomass consumption (dry weight) per sample plot and per sapling. The number of bites increased, but the percentage biomass removed did not differ when stand density increased. A relatively large bite size was observed on the plots of low stand density. The quantity of food, which on average was of relatively low quality, was obviously important due to the benefit gained through reducing the search time.

The nutritive value of the browse, expressed in terms of chemical compounds indicating low food digestibility, was lower in the dense than in the sparse Scots pine stand. However, the amount of crude protein and arginine were relatively high in the dense stand. We concluded that shading affected the nutritional status of saplings on high density plots.

Although the biomass removed by moose per sapling was high for low density plots, the remaining biomass was larger than that on the high-density plots owing to the relatively large twig biomass of saplings. The number of saplings per hectare without main stem breakage increased significantly as stand density increased.

The PDF includes a summary in Finnish.

  • Heikkilä, ORCID ID:E-mail:
  • Mikkonen, ORCID ID:E-mail:

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