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Articles containing the keyword 'composition'

Category : Article

article id 5600, category Article
Anton K. Chtchoukine. (1996). North European platyphyllous forests: biodiversity dynamics and climate changes in northwest European Russia. Silva Fennica vol. 30 no. 2–3 article id 5600. https://doi.org/10.14214/sf.a9245
Keywords: biodiversity; reforestation; climatic change; tree species composition; broadleaved forests; deciduous forests
Abstract | View details | Full text in PDF | Author Info

Distribution, biodiversity and reforestation dynamics of the platyphyllous forests in the Northwest European Russia were investigated. Data assembled from 21 landscape regions (250–350 km2 each) show special features of small-leaved lime (Tilia cordata Mill., Norway maple (Acer platanoides L.), mountain elm (Ulmus glabra Mill.) and English oak (Qurecus robur L.) reforestation during the last two decades. New tendencies were found for the taiga areas with natural Norway spruce (Picea abies (L.) H. Karst.) and Scots pine (Pinus sylvestris L.) vegetation. Natural platyphyllous reforestation in cut spruce areas poses as supposed a special question for forest management policy in the relationship to global climate changes. Feasible unsustainability of the common types of succession (Norway spruce - European birch (Betula pendula Roth); Norway spruce - European aspen (Populus tremula L.)) is discussed. Biodiversity of herbs, shrubs and tree species of platyphyllous forests is high and complex and is situated in 4–15 old-growth relics in each landscape region. Low-level genotype heterogeneity of nemoral flora species of such isolated populations is presumed. Special biodiversity conservation regulations are proposed.

  • Chtchoukine, E-mail: ac@mm.unknown (email)
article id 5593, category Article
Malle Mandre, Jaan Klõseiko, Vaike Reisner, Hardi Tullus. (1996). Assessment of CO2 fluxes and effects of possible climate changes on forests in Estonia. Silva Fennica vol. 30 no. 2–3 article id 5593. https://doi.org/10.14214/sf.a9238
Keywords: climate change; CO2; forestry; Estonia; carbon dioxide; forest composition; Forest Gap model
Abstract | View details | Full text in PDF | Author Info

The present study is the first attempt to carry out an inventory of greenhouse gas (GHG) fluxes in the forests of Estonia. The emission and uptake of CO2 as a result of forest management, forest conversion and abandonment of cultivated lands in Estonia was estimated. The removal of GHG by Estonian forests in 1990 exceeded the release about 3.3 times. Changes in the species composition and productivity of forest sites under various simulated climate change scenarios have been predicted by using the Forest Gap Model for the central and coastal areas of Estonia. The computational examples showed that the changes in forest community would be essential.

  • Mandre, E-mail: mm@mm.unknown (email)
  • Klõseiko, E-mail: jk@mm.unknown
  • Reisner, E-mail: vr@mm.unknown
  • Tullus, E-mail: ht@mm.unknown
article id 5574, category Article
Jouni Vettenranta. (1996). Effect of species composition on economic return in a mixed stand of Norway spruce and Scots pine. Silva Fennica vol. 30 no. 1 article id 5574. https://doi.org/10.14214/sf.a9219
Keywords: Pinus sylvestris; Picea abies; net present value; mixed forests; growth models; tree species composition; forest management practices; felling methods; valuation; soil expectation value; treatment programme
Abstract | View details | Full text in PDF | Author Info

The effect of species mixture was studied in a mixed stand of Norway spruce (Picea abies (L.) H. Karst.) and Scots pine (Pinus sylvestris L.) by simulating around 100 different treatment schedules during the rotation in a naturally regenerated even-aged stand located on a site of medium fertility in North Karelia, Finland. Both thinning from below and thinning from above were applied. Optimum rotations were determined by maximising the net present value calculated to infinity and different treatment schedules were compared with the net present value over one rotation as per rotation applied. In the optimum treatment programme, the proportion of pines was decreased by half of the basal area in the first thinning stage and by the end of the rotation to about one third. In thinning from above, the proportion of pines can be maintained at a slightly higher level. It is economically profitable to maintain the growing stock capital at approximately the level recommended by Forest Centre Tapio, a semi-governmental forestry authority. With non-optimum species composition, the loss in net present value over one rotation can be about 10 % in thinning from below and about 20 % in thinning from above.

  • Vettenranta, E-mail: jv@mm.unknown (email)
article id 5471, category Article
Hannu Fritze. (1992). Effects of environmental pollution on forest soil microflora - a review. Silva Fennica vol. 26 no. 1 article id 5471. https://doi.org/10.14214/sf.a15632
Keywords: soil respiration; forest soil; litter decomposition; acidification; lime; heavy metals; pollution; microflora; fungal hyphae
Abstract | View details | Full text in PDF | Author Info

The article is a literature review focusing on the reaction of soil respiration, litter decomposition and microflora of forest soils to various pollutants like acidic deposition, heavy metals and unusual high amounts of basic cations. There is a great deal of evidence indicating that environmental pollution affects soil microbial activity and community structure. Much of the data originates from experimental designs where high levels of pollutants were applied to the soil under field or laboratory conditions. Furthermore, many were short-term experiments designed to look for large effects. These experiments have an indicative value, but it has to be kept in mind that environmental pollution is a combination of many pollutants, mostly at low concentrations, acting over long periods of time. There is therefore consequently a demand for research performed in natural forest environments polluted with anthropogenic compounds. 

  • Fritze, E-mail: hf@mm.unknown (email)
article id 5239, category Article
Jussi Kuusipalo. (1985). On the use of tree stand parameters in estimating light conditions below the canopy. Silva Fennica vol. 19 no. 2 article id 5239. https://doi.org/10.14214/sf.a15418
Keywords: Pinus sylvestris; basal area; Picea abies; canopy; tree species composition; light conditions; light climate; number of stems; canopy coverage
Abstract | View details | Full text in PDF | Author Info

Especially in forest vegetation studies, the light climate below the canopy is of great interest. In extensive forest inventories, direct measurement of the light conditions is too time-consuming. Often only the standard tree stand parameters are available. The present study was undertaken with the aim to develop methods for estimation of the light climate on the basis of readily measurable tree stand characteristics. The study material includes 40 sample plots representing different kinds of more or less mature forest stands of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) H. Karst.).

In each forest stand, a set of hemipherical photographs was taken and standard tree stand measurements were performed. A regression approach was applied in order to elaborate linear models for predicting the canopy coverage. The total basal area of the stand explained 63% of variance in the canopy coverage computed from hemipherical photographs. A coefficient representing the relative proportion of Norway spruce in the stand increased the explanatory power into 75%. When either the stand density (stems/unit area) or dominant age of the stand was included into the model, increment of the explanatory power into 80% was achieved. By incorporating both of the preceding predictors, an explanatory power of 85% was reached.

The PDF includes a summary in Finnish.

  • Kuusipalo, E-mail: jk@mm.unknown (email)
article id 5110, category Article
Helena Lehtiö. (1981). Effect of air pollution on the volatile oil in needles of Scots pine (Pinus sylvestris L.). Silva Fennica vol. 15 no. 2 article id 5110. https://doi.org/10.14214/sf.a15051
Keywords: Pinus sylvestris; needles; Scots pine; needle damages; air pollution; monoterpene composition; volatile oil
Abstract | View details | Full text in PDF | Author Info

The amount of volatile oil and monoterpene composition in Scots pine (Pinus sylvestris L.) needles were studied in trees growing near two factories and in the city of Kuopio, Central Finland. Trees in these locations are exposed principally to sulphur dioxide and fluoride emissions.

The amount of volatile oil increased with increasing injury class in the trees growing near the fertilizer factory. The amounts of volatile oil in trees near the pulp mill differed in the various injury classes. More oil was found in younger needles. The greatest differences in monoterpene composition were in the amounts of camphene, α-pinene, myrcene and tricyclene.

The PDF includes a summary in Finnish.

  • Lehtiö, E-mail: hl@mm.unknown (email)
article id 4730, category Article
Erkki Lähde. (1966). Kokeita selluloosan hajaantumisnopeudesta erilaisissa metsiköissä. Silva Fennica no. 119 article id 4730. https://doi.org/10.14214/sf.a14288
English title: Experiments on the decomposition rate of cellulose in different stands.
Original keywords: kuusi; koivu; puulajit; hajotus; maatuminen; selluloosa; hajoaminen; hajotusnopeus
English keywords: Norway spruce; birch; Betula; Picea abies; tree species; decompostition; decomposition rate; cellulose
Abstract | View details | Full text in PDF | Author Info

The aim of this project was to investigate the cellulose decomposition rate in the soil on the ecological conditions created by different tree species, particularly birch (Betula sp.) and Norway spruce (Picea abies (L.) H. Karst.). Therefore, comparable sample plots were established in adjoining birch and spruce stands. Data on the stands, the vegetation, and the soil in the sample plots were collected. The experiment was carried out in the Ruotsinkylä Experimental Forest near Helsinki in Southern Finland.

Five pieces (3x5x0.15 cm) of cellulose (bleached sulphite pulp) were dried, weighed, and fastened in a row into a nylon bag. The bags were placed into the soil at a slant so that the upmost piece of cellulose was in the depth of 0–1.5 cm and the bottom one 6–7.5 cm. The weight losses of the pieces were measured after periods ranging from 6 to 12 months.

The results show that even within the same forest type, decomposition is much more rapid in birch stands than in spruce stands. In all the stands the decomposition rate decreased rapidly with increasing depth. The difference between birch and spruce stand, as well as the decrease with increasing depth, was probably mainly due to different thermal conditions.

The PDF includes a summary in English.

  • Lähde, E-mail: el@mm.unknown (email)

Category : Article

article id 7172, category Article
Ilmari Schalin. (1966). Studies on the microfungi in the forest floor of subarctic pine forests. Acta Forestalia Fennica vol. 81 no. 7 article id 7172. https://doi.org/10.14214/aff.7172
Keywords: forest soil; decomposition; microgungi; microbes; microfungal populations; organic layer; Mucor; Penicillium; Trichoderma
Abstract | View details | Full text in PDF | Author Info

Dilution plate method was used in studying the density and composition of the microfungal populations of the organic layer of Scots pine forests, and the soil-plate method in studying the part of these populations decomposing cellulose. The media used were rose bengal agar (Martin’s medium for fungi) and cellulose medium.

The microfungal density depended to a considerable extent on the moisture content and temperature of the organic layer. Only the combination of relatively high moisture content and temperature, but neither of these factors alone, influenced considerably the microfungal population density. The correlation of the populations to the changes in this combined factor was stronger than the correlation to the seasonal variations of spring, summer and autumn.

The microfungal population consisted of only a few species. Mucor, Mortierella and Penicillium were the most common genera isolated from the rose bengal agar. The first and the last of these comprised almost 90% of the total population. For the Mucor fungi, increases in the moisture content up to the maximum values found (75%) were favourable; the Penicillium fungi, on the contrary, were intolerant of high moisture content.

Among the cellulose decomposing microfungi grown on cellulose medium, Trichoderma sp. was the most common; also, it formed the most colonies, tolerated the lowest temperatures, and was most efficient. The others were of the genera Pullularia, Verticillium, Scopulariopsis and Penicillium. In addition, there were some unidentified Phycomycetes fungi. Only the two first-mentioned caused observable changes in cellulose.

  • Schalin, E-mail: is@mm.unknown (email)
article id 7171, category Article
Erkki Lähde. (1966). Vertical distribution of biological activity in peat of some virgin and drained swamp types. Acta Forestalia Fennica vol. 81 no. 6 article id 7171. https://doi.org/10.14214/aff.7171
Keywords: drained peatlands; peat; decomposition; peatlands; cellulose decomposition; gas exchange; biological activity
Abstract | View details | Full text in PDF | Author Info

The objective of this project was to determine the amount of gas exchange in peat samples collected from several swamps, using the Warburg method in the laboratory measurements. Special attention was directed on the influence of the lowering of the ground water level through drainage, on oxidation-reduction conditions in the samples from both forested and treeless peatlands, by measuring oxygen uptake and CO2 release. The biological activity in situ was determined by the cellulose decomposition rate in the sample plots. The six areas examined were both in drained peatlands and peatlands in natural condition.

The results show that in the sample plots in open swamps there was no consistent differences in the CO2 release rate in peat samples taken from different depths. However, in the sample plots on forested swamps rapid decrease is seen with increasing depth. The decreased biological activity of peat is caused by the oxidation-reduction conditions. The CO2 release rate may also be due to the respiration of tree roots, which are very shallow in peatlands.

The rate of in situ cellulose decomposition experiment and CO2 release indicated by the Warburg measurements appear to be correlated. The results indicate improved conditions for cellulose-decomposing microbes after draining. It is also possible that the biological activity of peat after draining increases to a considerable depth until the decrease of easily decomposable substances limit the activity in an old drainage area. The cellulose decomposition rate would still increase as the oxidation-reduction conditions improve.

  • Lähde, E-mail: el@mm.unknown (email)
article id 7154, category Article
Ilmari Schalin. (1964). Some factors affecting the quantitative determination of aerobic bacteria in forest humus. Acta Forestalia Fennica vol. 78 no. 1 article id 7154. https://doi.org/10.14214/aff.7154
Keywords: decomposition; soil; humus; methods; bacteria; quantitative analysis; dilution plate method
Abstract | View details | Full text in PDF | Author Info

Quantitative investigation of bacteria in the humus is needed when the intensity of their function in decomposition in the soil is studied. In this study bacterial density of humus was measured using dilution plate method, which was subjected to thorough investigation. The method was chosen, despite its complexity, because it is quite consistent, each stage can be studied separately and the reliability can be tested. The aim of the study was to determine the best way to take samples so that the sample will represent the bacterial population as closely as possible, and to optimize homogenization and dilution of the sample and the assays.

On the basis of the results of the investigation, a procedure was developed for quantitative determination of aerobic bacteria in the humus by the dilution plate method. The paper recommends that subsamples are collected systematically from at least 25 different points. The moisture and temperature of the samples should remain similar to the natural environment until preparation of the dilution. The sample was homogenized with the Bühler homogenizer, which is constructed so that a certain degree of asepsis can be maintained and the speed of the apparatus can be regulated. The content of the mineral nutrients of the sample must be determined when choosing the way of homogenization to obtain the highest number of colonies per plate. The sample was diluted with either 0.1–0.01% peptone solution or 0.5% soil extract. The most advantageous degree of dilution was obtained by testing with the aid of Fisher’s dispersion index the probability of the Poisson distribution in the results.

  • Schalin, E-mail: is@mm.unknown (email)
article id 7111, category Article
Juhani Sarasto. (1960). Turpeen maatuneisuuden määrittämisestä : v. Postin maatumisasteen ja Pjavtshenkon maatumisprosentin vertailu. Acta Forestalia Fennica vol. 71 no. 2 article id 7111. https://doi.org/10.14214/aff.7111
English title: Determining decomposition decree of peat with von Post Humification Scale and Pjavtshenko's method.
Original keywords: suot; menetelmät; turve; turpeen maatumisaste
English keywords: peat; peatlands; methods; decomposition of peat; decomposition decree
Abstract | View details | Full text in PDF | Author Info

Decomposition of the peat using von Post Humification Scale (1–10), developed by Lennart von Post, can be determined based on characteristics of peat, such as fibre integrity, colour and viscosity of exudate, and presence of colloidal particles, of handful of peat squeezed in the hand. It is easy to use and has proved useful in the practical work. The method developed by Pjavthenko is mostly based on specific weight off a dried sample in percentage, and requires analysis in the laboratory. The aim of this study was to compare the results of these two methods by measuring 156 peat samples representing different stages of decomposition.

The methods are based on different principles, which is reflected in small differences of the results. The maximum scores of the methods are clearly in different level. The maximum grade of 9–10 in von Post scale correspond decomposition percentage 51 in the scale of Pjavtsheko. However, the decomposition values in von Post scale are placed evenly on the scale of Pjavtshenko. This suggests that the von Post Scale is consistent and accurately developed. According to the study, the Pjavtshenko method is a good method to validate results of von Post Humification Scale, and can be used when decomposition of peat samples is determined in laboratory.

The PDF includes a summary in German.

  • Sarasto, E-mail: js@mm.unknown (email)
article id 7451, category Article
Helge Gyllenberg, Pauli Hanioja, Unto Vartiovaara. (1954). Havaintoja eräiden viljelemättömien maatyyppien mikrobiston koostumuksesta. Acta Forestalia Fennica vol. 62 no. 2 article id 7451. https://doi.org/10.14214/aff.7451
English title: Observations on the composition of the microbial population in some virgin soils.
Original keywords: metsätyyppi; metsämaa; bakteerit; mikrobit; sienet; mykobakteerit; hajottajat; typensitojabakteeri
English keywords: forest soil; decomposition; fungi; forest type; bacteria; microbes; nitrogen fixing bacteria
Abstract | View details | Full text in PDF | Author Info

The purpose of this investigation was to obtain a preliminary picture of the composition of the microbial population in some virgin soils on forest land in Finland. Four different forest types were studied, Oxalis-Myrtillus type birch (Betula sp.) stand, Oxalis-Myrtillus type Norway spruce (Picea abies (L.) Karst.) stand, Vaccinium type Scots pine (Pinus sylvestris L.) stand, and drained pine bog. In addition, a flood meadow was selected as a comparison.

The methods used captured only part of the fungi growing in the soil. Rapidly growing types, especially Mucor and Penicillium species, were mainly isolated. In addition, fungi showing activity of decomposition, such as Fusarium, Monosporium and Spicaria, as well as an ascomycete of the genius Ascobolus, were isolated. Autochthonous bacteria were most abundant in the soils of Oxalis-Myrtillus type forests and in the flood meadow. In the birch stand 90% of the autochthonous bacterial flora were gram-negative bacteria, in the Oxallis-Myrtillus spruce and Vaccinium type pine stand 60% were gram-negative, while the share was only 25% in the pine bog. Nitrogen-fixing bacteria of the type Clostridium pasteurianum were found in all soils. Actinomycetes were found in all sites. The numbers of protozoa were highest in the soils of Oxalis-Myrtillus type forests.

There were no big differences between the forest soils and the flood meadow. Some groups of micro-organisms seem to be absent from the forest soils, which is probably due to the more favourable pH in the meadow. The occurrence of myxobacteria is interesting since no earlier data exist of this organism in Finnish soils.

The PDF includes a summary in English.

  • Gyllenberg, E-mail: hg@mm.unknown (email)
  • Hanioja, E-mail: ph@mm.unknown
  • Vartiovaara, E-mail: uv@mm.unknown
article id 7295, category Article
O. J. Lukkala. (1934). Lounais-Suomen metsien puulajihistoriasta. Acta Forestalia Fennica vol. 40 no. 18 article id 7295. https://doi.org/10.14214/aff.7295
English title: The history of tree species in southwest Finland.
Original keywords: suo; puulajisuhteet; siitepölyanalyysi
English keywords: peatland; tree species composition; pollen analysis
Abstract | View details | Full text in PDF | Author Info

Pollen analysis has given information on development of the tree species composition after the ice age, but this kind of studies have not been published in Finland. In this study, pollen analysis was performed in five peatlands in the southwest Finland. According to the analysis, the forests of the area have had similar tree species composition for many thousands of years. Scots pine (Pinus sylvestris L.) has been the dominant species as long as there has been Norway spruce (Picea abies (L.) H. Karst.) in the area. Norway spruce seems to have arrived about 4,500 years ago. It increased slowly in the beginning, and after reaching a maximum has been slowly decreasing. Before spruce arrived, Betula sp. was more abundant, and seemed to be the dominant tree species in some places. Traces of fire in the peat layers indicate that forest fires have been common before people arrived in the area, and may have beneficial to birch. Like Betula sp., also Alnus sp. were more common before spruce arrived. Also pollen of other broadleaved species, for example, Tilia sp., Ulmus sp. and Corylus could be found. However, Quercus pollen was not found. The paludification of the peatlands had begun at different times which indicates that there has not been a common cause for the development of the peatlands in the area.

The PDF includes a summary in German.

  • Lukkala, E-mail: ol@mm.unknown (email)
article id 7280, category Article
Gust. Komppa. (1934). Knowledge on the needle oil of certain exotic coniferous trees growing in Finland. Acta Forestalia Fennica vol. 40 no. 3 article id 7280. https://doi.org/10.14214/aff.7280
Keywords: needle oil; coniferous-needle oil; Pseudotsuga Douglasii cecian Schwerin; Pinus Murrayana Engelm; chemical composition
Abstract | View details | Full text in PDF | Author Info

The aim of the study was to find out firstly whether there are differences between the etheric oils of botanically alike but in different climatic conditions growing trees, and secondly whether the oils from alike trees are also alike. This article shortly describes the chemical composition of  Pseudotsuga Douglasii cecian Schwerin and Pinus Murrayana Engelm. The needle samples are from Mustiala Arboretum.

The first mentioned contains α-pinene, camphene, bornyl acetate, probably also limonene but no geraniol. Pinus Murraya’s oil contains pinene, camphene, borneol and cadinene.

The PDF contains a summary in Finnish.  

  • Komppa, E-mail: gk@mm.unknown (email)
article id 7601, category Article
Erkki Lähde. (1969). Biological activity in some natural and drained peat soils with special reference to oxidation-reduction conditions. Acta Forestalia Fennica no. 94 article id 7601. https://doi.org/10.14214/aff.7601
Keywords: drained peatlands; peat; decomposition; ground water table; oxidizing conditions; reducing conditions
Abstract | View details | Full text in PDF | Author Info

The aim of the present study was to collect information on biological activity in the topmost 30 cm peat layer in certain natural and drained peatlands of different fertility, covered by different stands.

The results showed that if the ground water table in peatland sites is located in the immediate vicinity of the ground surface (about 5-10 cm in depth), conditions are reducing, and often even anaerobic, up to the ground surface. By means of drainage the aerobic limit can be dropped to a greater depth. This will occur because of the aerobic limit closely follows the fluctuation of the ground water table.

Although, by means of drainage, the aerobic limit can be lowered to more than 50 cm in depth, rains are followed by a rise of a ground water table and the aerobic limit; hereby a change from oxidizing to reducing conditions takes place. Only by keeping the ground water table and the aerobic limit constantly at the depth of more than 50 cm is it possible to obtain oxidizing conditions in the topmost 20-30 cm peat layer. The anaerobic conditions prevent the tree roots penetrating deeper in the peat.

In reducing conditions cellulose decomposition as well as carbon dioxide release from peat samples is slower than in oxidizing conditions. The rate of cellulose decomposition, however, is essentially dependent on the nitrogen content and the acidity of the peat.

  • Lähde, E-mail: el@mm.unknown (email)

Category : Special section

article id 290, category Special section
Mikko Peltoniemi, Esther Thürig, Stephen Ogle, Taru Palosuo, Marion Schrumpf, Thomas Wutzler, Klaus Butterbach-Bahl, Oleg Chertov, Alexander Komarov, Aleksey Mikhailov, Annemieke Gärdenäs, Charles Perry, Jari Liski, Pete Smith, Raisa Mäkipää. (2007). Models in country scale carbon accounting of forest soils. Silva Fennica vol. 41 no. 3 article id 290. https://doi.org/10.14214/sf.290
Keywords: National Forest Inventory; soil carbon; greenhouse gas inventory; decomposition; IPCC; regional and national modeling; soil model
Abstract | View details | Full text in PDF | Author Info
Countries need to assess changes in the carbon stocks of forest soils as a part of national greenhouse gas (GHG) inventories under the United Nations Framework Convention on Climate Change (UNFCCC) and the Kyoto Protocol (KP). Since measuring these changes is expensive, it is likely that many countries will use alternative methods to prepare these estimates. We reviewed seven well-known soil carbon models from the point of view of preparing country-scale soil C change estimates. We first introduced the models and explained how they incorporated the most important input variables. Second, we evaluated their applicability at regional scale considering commonly available data sources. Third, we compiled references to data that exist for evaluation of model performance in forest soils. A range of process-based soil carbon models differing in input data requirements exist, allowing some flexibility to forest soil C accounting. Simple models may be the only reasonable option to estimate soil C changes if available resources are limited. More complex models may be used as integral parts of sophisticated inventories assimilating several data sources. Currently, measurement data for model evaluation are common for agricultural soils, but less data have been collected in forest soils. Definitions of model and measured soil pools often differ, ancillary model inputs require scaling of data, and soil C measurements are uncertain. These issues complicate the preparation of model estimates and their evaluation with empirical data, at large scale. Assessment of uncertainties that accounts for the effect of model choice is important part of inventories estimating large-scale soil C changes. Joint development of models and large-scale soil measurement campaigns could reduce the inconsistencies between models and empirical data, and eventually also the uncertainties of model predictions.
  • Peltoniemi, Finnish Forest Research Institute, Vantaa Research Unit, P.O. Box 18, FI-01301 Vantaa, Finland E-mail: mikko.peltoniemi@metla.fi (email)
  • Thürig, Swiss Federal Institute for Forest, Snow and Landscape Research (WSL), Birmensdorf, Switzerland; European Forest Institute, Joensuu, Finland E-mail: et@nn.ch
  • Ogle, Natural Resources Ecology Laboratory, Colorado State University, Fort Collins, USA E-mail: so@nn.us
  • Palosuo, European Forest Institute, Joensuu, Finland E-mail: tp@nn.fi
  • Schrumpf, Max-Planck-Institute for Biogeochemistry, Jena, Germany E-mail: ms@nn.de
  • Wutzler, Max-Planck-Institute for Biogeochemistry, Jena, Germany E-mail: tw@nn.de
  • Butterbach-Bahl, Institute for Meteorology and Climate Research, Forschungszentrum Karlsruhe GmbH, Garmisch-Partenkirchen, Germany E-mail: kbb@nn.de
  • Chertov, St. Petersburg State University, St. Petersburg-Peterhof, Russia E-mail: oc@nn.ru
  • Komarov, Institute of Physicochemical and Biological Problems in Soil Science of Russian Academy of Sciences, Pushchino, Russia E-mail: ak@nn.ru
  • Mikhailov, Institute of Physicochemical and Biological Problems in Soil Science of Russian Academy of Sciences, Pushchino, Russia E-mail: am@nn.ru
  • Gärdenäs, Dept. of Soil Sciences, Swedish University of Agricultural Sciences, Uppsala, Sweden E-mail: ag@nn.se
  • Perry, USDA Forest Service, Northern Research Station, St. Paul, MN USA E-mail: cp@nn.us
  • Liski, Finnish Environment Institute, Helsinki, Finland E-mail: jl@nn.fi
  • Smith, School of Biological Sciences, University of Aberdeen, Aberdeen, UK E-mail: ps@nn.uk
  • Mäkipää, Finnish Forest Research Institute, Vantaa Research Unit, P.O. Box 18, FI-01301 Vantaa, Finland E-mail: raisa.makipaa@metla.fi
article id 288, category Special section
Björn Berg, Per Gundersen, Cecilia Akselsson, Maj-Britt Johansson, Åke Nilsson, Lars Vesterdal. (2007). Carbon sequestration rates in Swedish forest soils – a comparison of three approaches. Silva Fennica vol. 41 no. 3 article id 288. https://doi.org/10.14214/sf.288
Keywords: carbon sequestration; stable humus; forest floor C; litter decomposition; limit value
Abstract | View details | Full text in PDF | Author Info

Carbon sequestration rates in forest soil can be estimated using the concept of calculable stable remains in decomposing litter. In a case study of Swedish forest land we estimated C-sequestration rates for the two dominant tree species in the forest floor on top of the mineral soil. Carbon sequestration rates were upscaled to the forested land of Sweden with 23 x 106 ha with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (Karst.) L.). Two different theoretical approaches, based on limit-value for litter decomposition and N-balance for vegetation and SOM gave rates of the same magnitude. For the upscaling, using these methods, 17 000 grids of 5 x 5 km were used.

The ‘limit-value approach’ gave a sequestration of 4.8   106 tons of C, annually sequestered in the forest floor, with an average of 180 kg C ha–1 yr–1 and a range from 40 to 410 kg C ha–1 yr–1. The ‘N-balance approach’ gave an average value of c. 96 kg ha–1 yr–1 and a range from –60 to 360 kg ha–1 yr–1. A method based on direct measurements of changes in humus depth over 40 years, combined with C analyses gave an average rate that was not very different from the calculated rates, viz. c. 180 kg ha–1 yr–1 and a range from –20 to 730 kg ha–1 yr–1. These values agree with forest floor C sequestration rate based on e.g. sampling of chronsequences but differ from CO2 balance measurements.

The three approaches showed different patterns over the country and regions with high and low carbon sequestration rates that were not always directly related to climate.

  • Berg, Dept. of Forest Ecology, University of Helsinki, Finland (present address: Dipartimento Biologia Strutturale e Funzionale, Complesso Universitario, Monte S. Angelo, Napoli, Italy E-mail: bjorn.berg@helsinki.fi (email)
  • Gundersen, Forest & Landscape Denmark, University of Copenhagen, Denmark E-mail: pg@nn.dk
  • Akselsson, Swedish Environmental Research Institute, IVL, Gothenburg, Sweden E-mail: ca@nn.se
  • Johansson, Department of Forest Soils, SLU, Uppsala, Sweden E-mail: mbj@nn.se
  • Nilsson, Department of Forest Soils, SLU, Uppsala, Sweden E-mail: an@nn.se
  • Vesterdal, Forest & Landscape Denmark, University of Copenhagen, Denmark E-mail: lv@nn.dk

Category : Research article

article id 10084, category Research article
Mihails Čugunovs, Eeva-Stiina Tuittila, Jari Kouki. (2020). Proximity to charred logs in burned forests likely affects decomposition processes in the soil. Silva Fennica vol. 54 no. 1 article id 10084. https://doi.org/10.14214/sf.10084
Keywords: coarse woody debris; forest fire; prescribed burning; cellulose decomposition; tea-bag method
Highlights: Standardised organic substrate decomposition was tentatively observed to be faster adjacent to non-charred downed logs than away from the logs or adjacent to charred logs; A spatial linkage was observed between non-charred logs and decomposition in the soil in burned boreal forests; Proximity to a charred log may provide a micro-environment where decomposition rates differ from the surrounding forest soil.
Abstract | Full text in HTML | Full text in PDF | Author Info

We studied the spatial decomposition rates of standardised organic substrates in soils (burned boreal pine-dominated sub-xeric forests in eastern Finland), with respect to charred and non-charred coarse woody debris (CWD). Decomposition rates of rooibos plant litter inside teabags (C:N = 42.870 ± 1.841) and pressed-sheet Nordic hardwood pulp (consisting of mainly alpha-cellulose) were measured at 0.2 m distance from 20 charred (LC0.2) and 40 non-charred logs (LNC0.2). We also measured decomposition at 60 plots located 3–10 m away from downed logs (L3,10). The rooibos decomposition rate constant ‘k’ was 8.4% greater at the LNC0.2 logs than at the L3,10 or LC0.2 logs. Cellulose decomposed more completely in 1 micron mesh bags at LNC0.2 (44% of buried bags had leftover material) than at LC0.2 (76%) or L3,10 (70%). Decomposition of cellulose material was rapid but varied greatly between sampling plots. Our results indicate that decomposition of the standardised organic matter was more rapid close to CWD pieces than further away. However, only the plots located near non-charred logs (LNC0.2) exhibited high decomposition rates, with no corresponding increase observed at the charred logs (LC0.2). This suggests a possible noteworthy indirect effect of forest burning on soil organic matter (SOM) decomposition rates close to charred CWD after forest fires. We urge for more studies on this tentative observation as it may affect the estimates on how fires affect carbon cycling in forests.

  • Čugunovs, University of Eastern Finland, School of Forest Sciences, Yliopistokatu 7, P.O. Box 111, FI-80101 Joensuu, Finland E-mail: mihails.cugunovs@gmail.com (email)
  • Tuittila, University of Eastern Finland, School of Forest Sciences, Yliopistokatu 7, P.O. Box 111, FI-80101 Joensuu, Finland ORCID https://orcid.org/0000-0001-8861-3167 E-mail: eeva-stiina.tuittila@uef.fi
  • Kouki, University of Eastern Finland, School of Forest Sciences, Yliopistokatu 7, P.O. Box 111, FI-80101 Joensuu, Finland ORCID https://orcid.org/0000-0003-2624-8592 E-mail: jari.kouki@uef.fi
article id 1403, category Research article
Kristina Mjöfors, Monika Strömgren, Hans-Örjan Nohrstedt, Annemieke Ingrid Gärdenäs. (2015). Impact of site-preparation on soil-surface CO2 fluxes and litter decomposition in a clear-cut in Sweden. Silva Fennica vol. 49 no. 5 article id 1403. https://doi.org/10.14214/sf.1403
Keywords: boreal forest; soil disturbance; soil respiration; carbon stock; needle decomposition; coarse root decomposition
Highlights: Disturbances of the soil did not lead to higher CO2 emissions from the soil; Heavy mixing of the soil lead to lower CO2 emissions from the soil; Buried needles and coarse roots decomposed faster than those on the surface; Abundance of δ15N decreased in needles and roots after site preparation.
Abstract | Full text in HTML | Full text in PDF | Author Info

Boreal forest soil contains significant amounts of organic carbon. Soil disturbance, caused for example by site preparation or stump extraction, may increase decomposition and thus lead to higher CO2 emissions, contributing to global warming. The aim of this study was to quantify responses of soil-surface CO2 fluxes (Rs) and litter (needle and root) decomposition rates following various kinds of soil disturbance commonly caused by mechanical site preparation and stump harvest. For this purpose four treatments were applied in a clear-cut site in central Sweden: i) removal of the humus layer and top 2 cm of mineral soil, ii) placement of a humus layer and 2 cm of mineral soil upside down on top of undisturbed soil, forming a double humus layer buried under mineral soil, iii) heavy mixing of the humus layer and mineral soil, and iv) no disturbance (control). Rs measurements were acquired with a portable respiration system during two growing seasons. To assess the treatments’ effects on litter decomposition rates, needles or coarse roots (Ø = 6 mm) were incubated in litterbags at positions they would be located after the treatments (buried, or on top of the soil). The results indicate that site preparation-simulating treatments have no effect or may significantly reduce, rather than increase, CO2 emissions during the following two years. They also show that buried litter decomposes more rapidly than litter on the surface, but in other respects the treatments have little effect on litter decomposition rates.

  • Mjöfors, Swedish University of Agricultural Sciences (SLU), Department of Soil and Environment, P.O. Box 7014, 150 07 Uppsala, Sweden E-mail: kristina.mjofors@slu.se (email)
  • Strömgren, Swedish University of Agricultural Sciences (SLU), Department of Soil and Environment, P.O. Box 7014, 150 07 Uppsala, Sweden E-mail: Monika.stromgren@slu.se
  • Nohrstedt, Swedish University of Agricultural Sciences (SLU), Department of Soil and Environment, P.O. Box 7014, 150 07 Uppsala, Sweden E-mail: Hans-orjan.nohrstedt@slu.se
  • Gärdenäs, Swedish University of Agricultural Sciences (SLU), Department of Soil and Environment, P.O. Box 7014, 150 07 Uppsala, Sweden E-mail: Annemieke.gardenas@slu.se
article id 1337, category Research article
Leszek Bujoczek, Małgorzata Bujoczek, Jan Banaś, Stanisław Zięba. (2015). Spruce regeneration on woody microsites in a subalpine forest in the western Carpathians. Silva Fennica vol. 49 no. 3 article id 1337. https://doi.org/10.14214/sf.1337
Keywords: Picea abies; coarse woody debris; stumps; decomposition; regression model; fallen deadwood
Highlights: The occurrence probability of Picea abies seedlings on fallen deadwood was found to increase with diameter and decay stage of deadwood and with the volume of living trees, and to decrease with the density of living trees, sapling density, and land slope. It was also higher on stumps with greater diameter and in plots with higher sapling density, but decreased with increasing stump height.
Abstract | Full text in HTML | Full text in PDF | Author Info

The density of Picea abies [L.] Karst. regeneration on different microsites, the quantity and quality of woody microsites, and seedling occurrence probability on stumps and fallen deadwood were studied in a subalpine forest that has been under protection for approximately 30–40 years (Gorce Mountains in the western Carpathians). Thirty percent of seedlings and 29% of saplings grew on stumps and fallen deadwood, while the remaining regeneration occurred on soil surface and mounds created by uprooted trees. The occurrence probability of Picea seedlings on fallen deadwood increased with deadwood diameter and decay stage and with the volume of living trees, and decreased with increased density of living trees, sapling density, and land slope. Furthermore, seedlings were more likely to grow on stumps with a greater diameter and in plots with higher sapling density, but less likely to grow on higher stumps. Stumps and fallen deadwood covered about 4% of the forest floor, but the material that is most important for promoting regeneration (strongly decomposed logs and those of a diameter exceeding 30 cm) took up only about 22 m2 ha-1. We have concluded that in a subalpine forest that has been protected for 30–40 years regeneration processes take place mostly on soil surface and stumps. The role of fallen deadwood increases over time as a greater number of suitable logs (in terms of size and decay stage) become available.

  • Bujoczek, University of Agriculture in Krakow, E-mail: lbujoczek@gmail.com (email)
  • Bujoczek, University of Agriculture in Krakow, E-mail: bujoczek.m@gmail.com
  • Banaś, University of Agriculture in Krakow, E-mail: rlbanas@cyf-kr.edu.pl
  • Zięba, University of Agriculture in Krakow, E-mail: rlzieba@cyf-kr.edu.pl
article id 1279, category Research article
Andreas Kreutz, Tuomas Aakala, Russell Grenfell, Timo Kuuluvainen. (2015). Spatial tree community structure in three stands across a forest succession gradient in northern boreal Fennoscandia. Silva Fennica vol. 49 no. 2 article id 1279. https://doi.org/10.14214/sf.1279
Keywords: Picea abies; age structure; spatial structure; natural forest; Betula spp.; point pattern analysis; northern boreal; community composition
Highlights: We studied the tree community spatial structure in three 1.2-ha plots representing naturally developed northern boreal forests of varying ages; Spatial structure showed little differences between the mid-successional, late-successional and old-growth stands; The occurrence of Picea abies relative to Betula spp. indicated a mosaic-like spatial assembly; Mosaics are likely maintained by species-specific replacement, not reciprocal replacement as thought earlier.
Abstract | Full text in HTML | Full text in PDF | Author Info
Development of species composition during succession is well studied in natural boreal forests, but empirical assessments of how within-stand spatial structure develops in late-successional stages are few. Here, we quantified spatial patterns in three unmanaged stands consisting of Picea abies (L.) Karst. and Betula pubescens Ehrh. and Betula pendula Roth (hereafter Betula spp.) in northern boreal Fennoscandia. We conducted a comprehensive analysis of small-scale spatial point patterns in three fully mapped 1.2-ha sample plots, representing different forest developmental stages: mid-successional, late-successional and old-growth forest. We used several variants of Ripley’s K-function to analyze the spatial point patterns along the successional gradient. Univariate analyses showed that mature trees of both species were either randomly distributed or clumped. P. abies saplings were clumped, and Betula spp. saplings occurred in a random or clumped manner. In the bivariate analyses, saplings were more likely to be found in the surroundings of mature trees of the same species, but occurred independent of the individuals of other tree species. Mature trees showed interspecific repulsion. Only modest differences occurred in the univariate patterns between the three successional stages, but in the bivariate analyses the most evident patterns, i.e. intraspecific attraction and interspecific repulsion, were stronger in the older successional stages. Overall, the studied stands appear structured as species-specific mosaics. These mosaics, along with mixed species composition, seem to be maintained by species self-replacement, which contrasts with findings from earlier studies.
  • Kreutz, Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: andreas.kreutz@wald-rpl.de
  • Aakala, Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 University of Helsinki, Finland ORCID http://orcid.org/0000-0003-0160-6410 E-mail: tuomas.aakala@helsinki.fi (email)
  • Grenfell, Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: russell.grenfell@gmail.com
  • Kuuluvainen, Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: timo.kuuluvainen@helsinki.fi
article id 1211, category Research article
Xiao Chen, Deborah Page-Dumroese, Ruiheng Lv, Weiwei Wang, Guolei Li, Yong Liu. (2014). Interaction of initial litter quality and thinning intensity on litter decomposition rate, nitrogen accumulation and release in a pine plantation. Silva Fennica vol. 48 no. 4 article id 1211. https://doi.org/10.14214/sf.1211
Keywords: litter quality; litter decomposition; nitrogen cycling; thinning intensity; Pinus tabulaeformis
Highlights: Litter quality and thinning showed an interaction on one year litter decomposition rates, N accumulation, and net N release; N accumulated until the underlying critical acid-unhydrolyzable residue to nitrogen ratio (approximately 57–69) was met; Increased N concentration in litter and thinning intensity induced rapid litter decomposition and N cycling in coniferous plantation with a slow decomposition rate.
Abstract | Full text in HTML | Full text in PDF | Author Info
Thinning alters litter quality and microclimate under forests. Both of these two changes after thinning induce alterations of litter decomposition rates and nutrient cycling. However, a possible interaction between these two changes remains unclear. We placed two types of litter (LN, low N concentration litter; HN, high N concentration litter) in a Chinese pine (Pinus tabulaeformis Carrière) plantation under four thinning treatments to test the impacts of litter quality, thinning or their combination on decomposition rate and N cycling. In our study, N was accumulated to approach an underlying critical acid-unhydrolyzable residue to nitrogen ratio (approximately 57–69) in litter. Moreover, an interaction between litter quality and thinning on decomposition rates, N accumulation and net release did exist. On one hand, one year decomposition rate of LN was elevated after thinning while that of HN remained the same or even lower (under light thinning); N accumulation of LN declined with light thinning and was restored with the increase of thinning intensity whereas that of HN did not decline with thinning and increased under heavy thinning; Net N release from LN was only found in light and heavy thinning while that from HN was found in all treatments, moreover net N release from LN and HN were both elevated under heavy thinning. On the other hand, HN decomposed faster, accumulated less and released more N than LN did under all treatments. Generally, high N concentration in litter and high-intensity thinning can lead to rapid litter decomposition and N cycling in coniferous plantations.
  • Chen, Key Laboratory for Silviculture and Conservation of Ministry of Education, Beijing Forestry University, Beijing, 100083, China E-mail: chenxiao_0123@126.com
  • Page-Dumroese, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, 1221 South Main Street, Moscow, ID 83843, USA E-mail: ddumroese@fs.fed.us
  • Lv, College of Plant Science and Technology, Tarim University, Alar Xinjiang, 843300, China E-mail: lvrh514723@126.com
  • Wang, Key Laboratory for Silviculture and Conservation of Ministry of Education, Beijing Forestry University, Beijing, 100083, China E-mail: fuyuerdejia@126.com
  • Li, Key Laboratory for Silviculture and Conservation of Ministry of Education, Beijing Forestry University, Beijing, 100083, China E-mail: glli226@163.com
  • Liu, Key Laboratory for Silviculture and Conservation of Ministry of Education, Beijing Forestry University, Beijing, 100083, China E-mail: lyong@bjfu.edu.cn (email)
article id 98, category Research article
Meeri Pearson, Markku Saarinen, Kari Minkkinen, Niko Silvan, Jukka Laine. (2011). Mounding and scalping prior to reforestation of hydrologically sensitive deep-peated sites: factors behind Scots pine regeneration success. Silva Fennica vol. 45 no. 4 article id 98. https://doi.org/10.14214/sf.98
Keywords: soil preparation; peatland forest regeneration; mound; scalp; Scots pine; OM decomposition; excess moisture
Abstract | View details | Full text in PDF | Author Info
Watering up typically ensues after clearcutting forestry-drained peatland forests. Thus, the effectiveness of maintenance drainage and soil preparation procedures becomes paramount for establishing a new generation of commercial forest. Mounding is the primary method of soil preparation applied in regeneration sites lying on deep peat. As raised planting spots, mounds are resistant to waterlogging and assumed to be beneficial for organic matter (OM) decomposition via, e.g., increased soil aeration and temperature, which would also enhance seedling growth. In recent years, however, less intensive and cheaper alternatives like scalping have been sought with some reported cases of success. Our case study investigated the survival and growth of Scots pine outplants in mounds, scalps, and unprepared microsites along a moisture gradient. After three growing seasons, mounding accelerated neither seedling growth nor OM decomposition relative to the unprepared treatment. Survival in mounds was nonetheless superior overall. Scalps behaved as water collecting depressions leading to a catastrophic regeneration result. Based on our findings, water table level (WTL) overrides other growth-controlling factors in excess moisture conditions. To combat watering up coupled with greater than normal rainfall, we recommend reforestation strategies which provide elevated, prepared planting spots (i.e., mounds) or utilize unprepared, higher microforms.
  • Pearson, Finnish Forest Research Institute, Western Finland Regional Unit, Kaironiementie 15, FI-39700 Parkano, Finland E-mail: meeri.pearson@metla.fi (email)
  • Saarinen, Finnish Forest Research Institute, Western Finland Regional Unit, Kaironiementie 15, FI-39700 Parkano, Finland E-mail: ms@nn.fi
  • Minkkinen, University of Helsinki, Department of Forest Sciences, Helsinki, Finland E-mail: km@nn.fi
  • Silvan, Finnish Forest Research Institute, Western Finland Regional Unit, Kaironiementie 15, FI-39700 Parkano, Finland E-mail: ns@nn.fi
  • Laine, Finnish Forest Research Institute, Western Finland Regional Unit, Kaironiementie 15, FI-39700 Parkano, Finland E-mail: jl@nn.fi
article id 130, category Research article
Yongqing Yang, Yinan Yao, Xuejiang Zhang. (2010). Comparison of growth and physiological responses to severe drought between two altitudinal Hippophae rhamnoides populations. Silva Fennica vol. 44 no. 4 article id 130. https://doi.org/10.14214/sf.130
Keywords: sea buckthorn (Hippophae rhamnoides L.); physiological response; abscisic acid (ABA); carbon isotope composition
Abstract | View details | Full text in PDF | Author Info
Growth and physiological differences in response to drought were compared between two sea buckthorn (Hippophae rhamnoides L.) populations inhabited in the southeast of the Qinghai-Tibetan Plateau of China. The experimental design included two water regimes (100% and 25% of field capacity) and two populations from the low and high altitude zone. Our experiments were conducted in a naturally lit greenhouse under semi-controlled environmental conditions for a whole growing season in a dry valley (1800 m above the sea level). We found that drought tolerance is highly related to the plant antioxidant capacity and water use efficiency as well as leaf nutrient status in H. rhamnoides. The highland population (HP) experienced a greater inhibition in plant growth and leaf enlargement, lower leaf nitrogen and phosphorus content, lower root nodule biomass and root mass/foliage area ratio, and higher leaf water content loss paralleling with higher enhancement of abscisic acid level in response to drought, as compared with lowland population (LP). Additionally, reduction of leaf lignin content in HP further reduced its drought tolerance. On the contrary, LP showed effective adaptation strategies such as improvement of water economy and maintaining high ascorbic acid content. Therefore, we conclude that LP was more tolerant to drought than HP, and could be selected for reforestation in the dry valleys of upper Minjiang River regions in China.
  • Yang, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi, 830011, China & College of Life Sciences, Chongqing Normal University, Chongqing, 400047, China E-mail: yy@nn.cn
  • Yao, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi, 830011, China E-mail: yaoya@ms.xjb.ac.cn (email)
  • Zhang, Institute for Plant Protection and Soil sciences, Hubei Academy of Agricultural Sciences, Wuhan, 430064, China E-mail: xz@nn.cn
article id 151, category Research article
Janne Miettinen, Pekka Helle, Ari Nikula, Pekka Niemelä. (2010). Capercaillie (Tetrao urogallus) habitat characteristics in north-boreal Finland. Silva Fennica vol. 44 no. 2 article id 151. https://doi.org/10.14214/sf.151
Keywords: forest management; forestry; grouse; thinnings; landscape composition
Abstract | View details | Full text in PDF | Author Info
This study aimed to identify tools for taking capercaillie habitats into consideration in forest management. This would provide new alternatives for ecologically more sustainable forest management. Capercaillie summer and winter locations, from wildlife monitoring counts (1998–2004) in northern Finland, and reference, non-capercaillie locations were combined with forest planning data, and the area proportions of different landscape classes in an 800-m radius circle surrounding capercaillie and reference locations were compared. Thinning stands (in summer and winter) and spruce mires (in summer) were more abundant in capercaillie habitats than in reference landscapes, whereas e.g. seedling stands, mature stands and waste land areas were less abundant. The relative habitat use was highest in mean tree diameter (DBH) classes from 10.5 to 14.5 cm in summer habitats of adult capercaillie in heath forests, whereas in peatland forests, in brood habitats and in winter habitats it peaked in diameter classes 14.5 to 18.5 cm. The tree layer density was positively associated with the relative habitat use. A trend of lower habitat use was detected in the largest diameters (17–40 cm) in comparison to middle-sized diameters (10–16 cm) in heath forests, but not in peatland forests. Relatively young managed forests (age 30–40 years or more) can form suitable capercaillie habitats in north-boreal forests. However, this suitability is not necessarily permanent. Understorey management, longer rotations and multicohort forest management are suitable tools for capercaillie habitat management, because they can increase the available cover close to the ground, canopy cover, overall forest cover at the landscape scale and bilberry cover.
  • Miettinen, Finnish Game and Fisheries Research Institute, Finland E-mail: janne.miettinen@rktl.fi (email)
  • Helle, Finnish Game and Fisheries Research Institute, Finland E-mail: ph@nn.fi
  • Nikula, Finnish Forest Research Institute, Finland E-mail: an@nn.fi
  • Niemelä, University of Turku, Dept of Biology E-mail: pn@nn.fi
article id 171, category Research article
Takuo Nagaike. (2009). Snag abundance and species composition in a managed forest landscape in central Japan composed of Larix kaempferi plantations and secondary broadleaf forests. Silva Fennica vol. 43 no. 5 article id 171. https://doi.org/10.14214/sf.171
Keywords: snags; long-rotation plantation; plantation age; species composition
Abstract | View details | Full text in PDF | Author Info
Larix kaempferi is the main plantation species in the low-snow, cool-temperate zone of Japan. I studied L. kaempferi plantations of various stand ages in central Japan to examine and compare the effect of stand age on the abundance, size, and species composition of snags (standing dead trees) compared to those in secondary broadleaf forests. Plantations that were older than the standard rotation age had more and larger snags than young plantations, and the species diversity of snags was positively correlated with stand age. Because the density of living planted L. kaempferi showed little correlation with snag variables, whereas that of naturally regenerated tree species was positively correlated with snag variables, the density dependence of snag occurrence was stronger in naturally regenerated trees than in planted L. kaempferi. Snag species that were positively correlated with stand age were the main species in secondary broadleaf forests in this area. Basal area, density, and number of species of snags in standard-rotation plantations were significantly lower than in long-rotation plantations and secondary broadleaf forests. Long-rotation plantations are useful for retaining snags compared to standard-rotation plantations.
  • Nagaike, Yamanashi Forest Research Institute, Masuho, Yamanashi 400-0502, Japan E-mail: nagaike-zty@pref.yamanashi.lg.jp (email)
article id 182, category Research article
Janne Miettinen, Pekka Helle, Ari Nikula, Pekka Niemelä. (2009). Changes in landscape-scale habitat selection of capercaillie (Tetrao urogallus) in managed north-boreal forest. Silva Fennica vol. 43 no. 4 article id 182. https://doi.org/10.14214/sf.182
Keywords: forestry; landscape composition; capercaillie; wildlife triangle
Abstract | View details | Full text in PDF | Author Info
We studied changes in habitat selection of capercaillie in northern Finnish boreal forests at two spatial scales during two time periods, 1989–1992 and 2000–2003. We studied capercaillie densities and their changes between the study periods in relation to the landscape class proportions within 3-km buffer zones around the wildlife triangle center points. Furthermore, we compared the landscape class proportions in 800-meter buffer zones around capercaillie wildlife triangle count observations and around the counted wildlife triangle transects using t-tests and compositional analysis. At the local population scale (3 km) the change in adult density between the study periods was associated positively with the proportion of young thinning stands in 2003 and reversely with the mature stand (1992 and 2003) and clear-cut (1992) proportions. Capercaillie juvenile and pooled densities during 2000–2003 were positively associated with the advanced thinning stand proportion in 2003. At the capercaillie home range scale (800 m) habitats were rich in mature stands during 1989–1992 in relation to available habitats, but not during 2000–2003 when young thinning stands were more abundant in relation to available habitats. Relatively young managed forests can be suitable for capercaillie, but mature managed forests as capercaillie habitats may have deteriorated between the study periods. Spatial planning may help to form suitable areas that are large enough for the species, but the highest potential may lay in the forest stand scale, where increased cover on the ground could promote the habitat quality.
  • Miettinen, Kankurinhaka 14, FI-90450 Kempele, Finland E-mail: janne.miettinen@rktl.fi (email)
  • Helle, Finnish Game and Fisheries Research Institute, Tutkijantie 2 E, FI-90570 Oulu, Finland E-mail: ph@nn.fi
  • Nikula, Finnish Forest Research Institute, Rovaniemi Research Unit, Eteläranta 55, FI-99600 Rovaniemi, Finland E-mail: an@nn.fi
  • Niemelä, University of Turku, Department of Biology, FI-20014 University of Turku, Finland E-mail: pn@nn.fi
article id 255, category Research article
Claude Vidal, Adrian Lanz, Erkki Tomppo, Klemens Schadauer, Thomas Gschwantner, Lucio di Cosmo, Nicolas Robert. (2008). Establishing forest inventory reference definitions for forest and growing stock: a study towards common reporting. Silva Fennica vol. 42 no. 2 article id 255. https://doi.org/10.14214/sf.255
Keywords: national forest inventories; reference definitions; growing stock; harmonisation; analytical decomposition
Abstract | View details | Full text in PDF | Author Info
International agreements such as the Kyoto protocol and Convention on Biological Diversity (1992), as well as, criteria and indicator processes require reports on the status of nations’ forests. Any comparison of the current status and trends of forest resources among nations presumes that the nations’ applied definitions and concepts produce comparable estimates of the status of forests. In spite of this, the FAO has already collected global information for 60 years and made noticeable efforts in creating common definitions, but forest related data are still collected using diverse definitions, even regarding basic concepts such as forest and forest area. A simple consequence is that the cross-countries estimates are not comparable. The reasons behind the differences in the definitions are diverse histories, and sometimes different use of forests. In an ideal case, national forest inventories should fulfil both national and international needs. In addition to the FAO’s Forest Resources Assessment process, other efforts are made to assess the status of forests in European countries, e.g. European Forest Information and Communication System (EFICS). EFICS produced reports about forest inventories but does not suggest any common definition or method to convert estimates from one definition to another one. This article presents principles and methods to create commonly acceptable and adoptable definitions for forest inventories. The principles and methods are demonstrated using two examples: the reference definitions of forest and growing stock. The article is based on the work of COST Action E43 (http://www.metla.fi/eu/cost/e43/).
  • Vidal, Inventaire Forestier National, Château des Barres, Nogent-sur-Vernisson, France E-mail: claude.vidal@ifn.fr (email)
  • Lanz, WSL/FNP, Abteilung Landschaftsinventuren, Birmensdorf, Switzerland E-mail: al@nn.ch
  • Tomppo, Finnish Forest Research Institute, Vantaa Research Unit, Vantaa, Finland E-mail: et@nn.fi
  • Schadauer, Bundesamt und Forschungszentrum für Wald, Wien, Austria E-mail: ks@nn.at
  • Gschwantner, Bundesamt und Forschungszentrum für Wald, Wien, Austria E-mail: tg@nn.at
  • di Cosmo, ISAFA, Villazzano, Italy E-mail: ldc@nn.it
  • Robert, Inventaire Forestier National, Ch‰teau des Barres, Nogent-sur-Vernisson, France E-mail: nr@nn.fr
article id 294, category Research article
Ülle Püttsepp, Krista Lõhmus, Andres Koppel. (2007). Decomposition of fine roots and α-cellulose in a short rotation willow (Salix spp.) plantation on abandoned agricultural land. Silva Fennica vol. 41 no. 2 article id 294. https://doi.org/10.14214/sf.294
Keywords: decomposition; α-cellulose; fine roots; acid detergent lignin; short rotation forest; Salix spp.
Abstract | View details | Full text in PDF | Author Info
Decomposition of fine roots (<1 mm in diameter) of the clones of Salix viminalis, S. dasyclados and α-cellulose sheets (50 x 10 x 1 mm) was studied in a 6-years old Salix spp. plantation established on abandoned agricultural land in Estonia. The substrates were incubated in litterbags (mesh size 0.14 mm) in 5–10 cm topsoil, in non-fertilised plots for one year. Changes in the ash-free weight of the fine roots were best described by negative exponential models (S. viminalis R2 = 0.98, S. dasyclados R2 = 0.96), and by a linear model for α-cellulose (R2 = 0.63). The sheets of α-cellulose decomposed roughly twice as rapidly as the fine roots (S. viminalis k = 0.325, S. dasyclados k = 0.165). The remaining (of the initial) ash-free weights of the fine roots were 73.3 ± 0.8% (mean ± SE) and 85.8 ± 2.2% respectively, and of the α-cellulose 35.9 ± 8.5%, in the end of the one year of decomposition. The amount of acid detergent (AD) lignin in the fine-roots of S. viminalis increased significantly and did not change in S. dasyclados, suggesting higher activity of microbial decomposers in the first substrate. Of the studied quality parameters, the AD lignin was the major factor determining the different rate of decomposition of the fine roots of S. viminalis and S. dasyclados. Nitrogen was recycled in the fine root sub-system in both Salix species. This knowledge can be applied in the management of Salix plantations, aimed at bioenergy production.
  • Püttsepp, Department of Ecology, Swedish University of Agricultural Sciences, P.O. Box 7072, SE-75007 Uppsala, Sweden; Estonian University of Life Sciences, Kreuzwaldi 64, Tartu 51014, Estonia E-mail: ulle.puttsepp@ekol.slu.se (email)
  • Lõhmus, Institute of Geography, University of Tartu, Vanemuise 46, Tartu 51014, Estonia E-mail: kl@nn.ee
  • Koppel, Estonian University of Life Sciences, Kreuzwaldi 64, Tartu 51014, Estonia E-mail: ak@nn.ee
article id 305, category Research article
Anna-Maria Veijalainen, Marja-Liisa Juntunen, Arja Lilja, Helvi Heinonen-Tanski, Leo Tervo. (2007). Forest nursery waste composting in windrows with or without horse manure or urea – the composting process and nutrient leaching. Silva Fennica vol. 41 no. 1 article id 305. https://doi.org/10.14214/sf.305
Keywords: microbial hygiene; lignocellulose; nutrients; organic matter decomposition; temperature; tree seedling waste; waste management
Abstract | View details | Full text in PDF | Author Info
In order to find the best management practices for forest nursery waste composting, organic waste was composted without or with horse manure or urea in six windrows for two years. The windrows were built in four consecutive years during 1999–2002. In 1999, no extra-nutrients were added to the windrow (N99). In 2000, urea fertilizer was used as a nitrogen source (U00). Despite this, the process did not function properly. In 2001, two windrows were built, one (H01) with and the other (N01) without horse manure. Horse manure slightly accelerated the heating process. Consequently, two windrows with more horse manure were built in 2002. One was aerated passively (H02) as earlier windrows, and the other was aerated forcedly (HA02). Horse manure and forced aeration were needed to keep the temperature above 55°C for long enough to ensure microbial hygiene of the material. The degradation of cellulose was greater during the curing stage. Nutrient leaching was low, although the additives increased leaching in conjunction with the inefficient process. The results showed that forest nursery waste alone is ineffective at raising the temperature of the compost, and degrades slowly due to its low nutrient and easily available carbon content. The best management practice for forest nursery waste composting is to use horse manure and aeration to ensure the heating process. Environmental contamination can be avoided by collecting the leachates. Further research is needed to evaluate the usability of the compost.
  • Veijalainen, Finnish Forest Research Institute, FI-77600 Suonenjoki, Finland E-mail: amv@nn.fi (email)
  • Juntunen, Finnish Forest Research Institute, FI-77600 Suonenjoki, Finland E-mail: mlj@nn.fi
  • Lilja, Finnish Forest Research Institute, PO Box 18, FI-01301 Vantaa, Finland E-mail: al@nn.fi
  • Heinonen-Tanski, Univ. of Kuopio, Dept. of Environm. Sc., PO Box 1627, FI-70211 Kuopio, Finland E-mail: hht@nn.fi
  • Tervo, Finnish Forest Research Institute, FI-77600 Suonenjoki, Finland E-mail: lt@nn.fi
article id 348, category Research article
Chunyang Li, Xuejiang Zhang, Xingliang Liu, Olavi Luukkanen, Frank Berninger. (2006). Leaf morphological and physiological responses of Quercus aquifolioides along an altitudinal gradient. Silva Fennica vol. 40 no. 1 article id 348. https://doi.org/10.14214/sf.348
Keywords: carbon isotope composition; leaf nitrogen content; specific leaf area; stomata
Abstract | View details | Full text in PDF | Author Info
Quercus aquifolioides Rehder & E.H. Wilson, an evergreen alpine and subalpine shrub species, occupies a wide range of habitats on the eastern slopes of the Himalaya in China. In this study, we measured leaf morphology, nitrogen content and carbon isotope composition (as an indicator of water use efficiency) of Q. aquifolioides along an altitudinal gradient. We found that these leaf morphological and physiological responses to altitudinal gradients were non-linear with increasing altitude. Specific leaf area, stomatal length and index increased with increasing altitude below 2800 m, but decreased with increasing altitude above 2800 m. In contrast, leaf nitrogen content per unit area and carbon isotope composition showed opposite change patterns. Specific leaf area seemed to be the most important parameter that determined the carbon isotope composition along the altitudinal gradient. Our results suggest that near 2800 m in altitude could be the optimum zone for growth and development of Q. aquifolioides, and highlight the importance of the influence of altitude in research on plant physiological ecology.
  • Li, Chengdu Institute of Biology, Chinese Academy of Sciences, P.O. Box 416, Chengdu 610041, P. R. China E-mail: licy@cib.ac.cn (email)
  • Zhang, Chengdu Institute of Biology, Chinese Academy of Sciences, P.O. Box 416, Chengdu 610041, P. R. China; Graduate School of the Chinese Academy of Sciences, Beijing 100039, P.R. China E-mail: xz@nn.cn
  • Liu, Sichuan Academy of Forestry, Chengdu 610081, P. R. China E-mail: xl@nn.cn
  • Luukkanen, Viikki Tropical Resources Institute, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: ol@nn.fi
  • Berninger, Département des sciences biologiques, Cp 8888 succ centre ville, Université du Québec à Montréal, Montréal (QC) H3C 3P8, Canada E-mail: fb@nn.ca
article id 502, category Research article
Janne Levula, Hannu Ilvesniemi, Carl Johan Westman. (2003). Relation between soil properties and tree species composition in a Scots pine–Norway spruce stand in southern Finland. Silva Fennica vol. 37 no. 2 article id 502. https://doi.org/10.14214/sf.502
Keywords: Norway spruce; Scots pine; tree species composition; soil texture; soil-water retention; spatial variability
Abstract | View details | Full text in PDF | Author Info
It is commonly known in Finland that Scots pine (Pinus sylvestris L.) is a tree of dry soils and Norway spruce (Picea abies (L) Karst.) is a tree of fresh soils. However, the concepts of dry and fresh soils still lack a precise definition. Consequently, the discussion on which soil/site is a pine or spruce habitat has continued over several decades. Moreover, in forest regeneration, the practice of tree species selection between the pine and the spruce has varied. We investigated the relationship between soil properties and pine–spruce species composition in a mature, naturally regenerated stand in southern Finland. We applied spatial analysis to divide the stand area up into 3–7 classes based on selected soil properties and then investigated the variations in species composition among those classes. The pine–spruce basal area ratio (BA of pines / BA of spruces) increased along with increasing mean particle size and proportion of coarse sand and gravel particle size fraction (0.6–20 mm) of mineral soil, and was lowest in classes, with the highest proportions of fine texture fractions. The results suggest that in southern Finland on sorted soils, pine is more competitive in regeneration and growth than spruce when mean particle size is above 0.44 mm or percentage of coarse sand and gravel is higher than 50%.
  • Levula, University of Helsinki, Department of Forest Ecology, FIN-00014 University of Helsinki, Finland E-mail: janne.levula@helsinki.fi (email)
  • Ilvesniemi, University of Helsinki, Department of Forest Ecology, FIN-00014 University of Helsinki, Finland E-mail: hi@nn.fi
  • Westman, University of Helsinki, Department of Forest Ecology, FIN-00014 University of Helsinki, Finland E-mail: cjw@nn.fi
article id 567, category Research article
Sonia Légaré, Yves Bergeron, David Paré. (2002). Influence of forest composition on understory cover in boreal mixedwood forests of western Quebec. Silva Fennica vol. 36 no. 1 article id 567. https://doi.org/10.14214/sf.567
Keywords: biomass; understory; canopy; composition; nutrient availability
Abstract | View details | Full text in PDF | Author Info
Forest overstory composition influences both light and nutrient availability in the mixed boreal forest. The influence of stand composition on understory cover and biomass was investigated on two soil types (clay and till deposits). Four forest composition types were considered in this study: aspen (Populus tremuloides Michx.), paper birch (Betula papyrifera Marsh.), jack pine (Pinus banksiana Lamb.) and a mixture of balsam-fir (Abies balsamea (L.) Mill.) and white spruce (Picea glauca (Moench) Voss). The cover of all understory species was recorded while the biomass of two important and ubiquitous species was measured: mountain maple (Acer spicatum Lam.) of the shrub layer and large-leaved aster (Aster macrophyllus L.) of the herb layer. Soil analyses were conducted to evaluate the influence of overstory composition on understory biomass through its influences on soil characteristics. Analyses of variance showed a significant effect of forest canopy type on mountain maple biomass, understory cover and shrub cover but not on herb cover and large-leaved aster biomass. Path analysis was performed to explore the relationships between canopy type, nutrient availability and understory biomass. Contrary to what was expected, the variation in plant biomass associated with forest composition was weakly related to soil nutrient availability and more strongly related to stand structural attributes.
  • Légaré, Université du Québec en Abitibi-Témiscamingue, Groupe de recherche en écologie forestière interuniversitaire, 445, boulevard de l'Université, Rouyn-Noranda, QC, Canada J9X 5E4 E-mail: sonia.legare@uqat.uquebec.ca (email)
  • Bergeron, NSERC-UQAT-UQAM, Industrial Chair in sustainable forest management, CP 8888, succ. Centre-Ville, Montréal, QC, Canada H3C 3P8 E-mail: yb@nn.ca
  • Paré, Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, P.O. Box 3800, Sainte-Foy, QC, Canada G1V 4C7 E-mail: dp@nn.ca
article id 565, category Research article
Sybille Haeussler, Lorne Bedford, Alain Leduc, Yves Bergeron, J. Marty Kranabetter. (2002). Silvicultural disturbance severity and plant communities of the southern Canadian boreal forest. Silva Fennica vol. 36 no. 1 article id 565. https://doi.org/10.14214/sf.565
Keywords: site preparation; species composition; species diversity; ecosystem integrity; degradation; biodiversity indicators; life history strategies
Abstract | View details | Full text in PDF | Author Info
Boreal forest ecosystems are adapted to periodic disturbance, but there is widespread concern that conventional forest practises degrade plant communities. We examined vegetation diversity and composition after clearcut logging, mechanical and chemical site preparation in eight 5- to 12-yr old studies located in southern boreal forests of British Columbia and Quebec, Canada to find useful indicators for monitoring ecosystem integrity and to provide recommendations for the development and testing of new silvicultural approaches. Community-wide and species-specific responses were measured across gradients of disturbance severity and the results were explained in terms of the intermediate disturbance hypothesis and a simple regeneration model based on plant life history strategies. Species richness was 30 to 35% higher 5 to 8 years after clearcut logging than in old forest. Total and vascular species diversity generally peaked on moderately severe site treatments, while non-vascular diversity declined with increasing disturbance severity. On more-or-less mesic sites, there was little evidence of diversity loss within the range of conventional silvicultural disturbances; however, there were important changes in plant community composition. Removing soil organic layers caused a shift from residual and resprouting understory species to ruderal species regenerating from seeds and spores. Severe treatments dramatically increased non-native species invasion. Two important challenges for the proposed natural dynamics-based silviculture will be 1) to find ways of maintaining populations of sensitive non-vascular species and forest mycoheterotrophs, and 2) to create regeneration niches for disturbance-dependent indigenous plants without accelerating non-native species invasion.
  • Haeussler, C2 Site 81 RR#2 Monckton Rd., Smithers, B.C., Canada V0J 2N0 E-mail: skeena@bulkley.net (email)
  • Bedford, B.C. Ministry of Forests, P.O. Box 9513 Stn. Prov. Govt., Victoria, B.C., Canada, V8W 9C2 E-mail: lb@nn.ca
  • Leduc, Groupe de recherche en écologie forestière interuniversitaire, Université du Québec à Montréal, CP 8888, Succursale A, Montréal, Québec, Canada, H3C 3P8 E-mail: al@nn.ca
  • Bergeron, Groupe de recherche en écologie forestière interuniversitaire, Université du Québec à Montréal, CP 8888, Succursale A, Montréal, Québec, Canada, H3C 3P8 E-mail: yb@nn.ca
  • Kranabetter, B.C. Ministry of Forests, Bag 5000, Smithers, B.C., Canada, V0J 2N0 E-mail: jmk@nn.ca
article id 599, category Research article
Chun-Jiang Liu, Carl J. Westman, Hannu Ilvesniemi. (2001). Matter and nutrient dynamics of pine (Pinus tabulaeformis) and oak (Quercus variabilis) litter in North China. Silva Fennica vol. 35 no. 1 article id 599. https://doi.org/10.14214/sf.599
Keywords: Pinus; Quercus; decomposition; nutrients; fall; forest floor; litter; North China
Abstract | View details | Full text in PDF | Author Info
In the mountainous area of North China, a distinguishing feature of climate is the serious drought of spring and the humidity and high temperature of summer by which the litter production and decomposition of forest litter were strongly characterized. We investigated the dynamic and nutrient characteristics of litter in a 30-year-old mixed stand of Chinese pine (Pinus tabulaeformis Carr.) and deciduous orient oak (Quercus variabilis Bl.) and two comparable pure stands. Oak litterfall peaked in November and pine litterfall in December. The oak stand had the largest annual litterfall (347 g m–2) and the forest floor mass (950 g m–2), the mixed stand the second (236 g m–2 and 634 g m–2), and the pine stand the least (217 g m–2 and 615 g m–2). The nutrient return through litterfall and the storage in forest floor followed corresponding order between three stands. The weight loss of pine and oak foliage litter in first year was 25% and 20%. For senesced pine and oak leaves, the translocation rates of N, P and K were 56–83%. Nutrient concentrations were higher in oak leaf litter than pine needle litter, and the concentration of N and Ca appeared to rise while K concentration decreased in both decomposing litter.
  • Liu, Department of Forest Ecology, P.O. Box 27, FIN-00014 University of Helsinki, Finland E-mail: cliu@silvia.helsinki.fi (email)
  • Westman, Department of Forest Ecology, P.O. Box 27, FIN-00014 University of Helsinki, Finland E-mail: cjw@nn.fi
  • Ilvesniemi, Department of Forest Ecology, P.O. Box 27, FIN-00014 University of Helsinki, Finland E-mail: hi@nn.fi

Category : Research note

article id 91, category Research note
Raisa Mäkipää, Tapio Linkosalo. (2011). A non-destructive field method for measuring wood density of decaying logs. Silva Fennica vol. 45 no. 5 article id 91. https://doi.org/10.14214/sf.91
Keywords: coarse woody debris; carbon stock; decaying wood; wood decomposition; penetrometer; pilodyn
Abstract | View details | Full text in PDF | Author Info
Decaying dead wood density measurements are a useful indicator for multiple purposes, such as for estimating the amount of carbon in dead wood and making predictions of potential diversity of dead wood inhabiting fungi and insects. Currently, qualitative decay phases are used as wood density estimates in many applications, since measuring the density is laborious. A quantitative measure of density would, however, be preferred over the qualitative one. Penetrometers, which are commonly used for measuring the density of standing trees, might also be applicable to dead wood density measurements. We tested the device for making quick, quantitative measurements of decaying logs. The penetrometer measures the depth into which a pre-loaded spring forces a pin in the wood. We tested pins of 5 and 10 mm diameter together with an original 2.5 mm pin and compared the results with gravimetric density measurements of the sample logs. Our results suggest that the standard pin works for less decayed wood, but for more decomposed wood, the thicker 5 mm pin gave more reliable estimates when the penetration measures were converted to densities with a linear regression function (R2 = 0.62, F = 82.9, p = 0.000). The range of wood densities successfully measured with the 5 mm pin was from 180 to 510 kg m–3. With the 10 mm pin, the measuring resolution of denser wood was compromised, while the improvement at the other end of density scale was not large. As a conclusion, the penetrometer seems to be a promising tool for quick density testing of decaying logs in field, but it needs to be modified to use a thicker measuring pin than the standard 2.5 mm pin.
  • Mäkipää, The Finnish Forest Research Institute, Vantaa, Finland E-mail: raisa.makipaa@metla.fi (email)
  • Linkosalo, The Finnish Forest Research Institute, Vantaa, Finland E-mail: ts@nn.fi

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