Silva Fennica vol. 30 | 1996

This study investigates the relationship between Finnish sulphate pulp export prices and international pulp inventories using the Johansen cointegration method. Long-run equilibrium is found to exist between pulp price and NORSCAN inventory for the study period, 1980-94. Granger causality is found to exist from inventory to price but not vice versa. A simple short-run forecasting model for the Finnish pulp export price is formed. In preliminary analysis, the explanatory power of model is found to be acceptable but only under stable market conditions.

• Toppinen, E-mail: at@mm.unknown
• Hänninen, E-mail: rh@mm.unknown
• Laaksonen, E-mail: sl@mm.unknown

This paper is an assessment of what in Finland are referred to as local forest management associations (LFMAs); the local units of a non-profit, forestry-promotion institution. First, the concept of organisational effectiveness is explored and an attempt is made to define it with respect to the LFMAs. The study then seeks to identify the environmental constraints, organisational characteristics and managerial practices differentiating the most effective and least effective associations. Discriminant analysis revealed four determinants of effectiveness: agrarian prosperity in the given area, activeness in marketing services to forest owners, the board of governors' role, and goal setting practices. The results thus indicate that the comparison of managerial policies and practices among LFMAs can provide useful information for improving their effectiveness.

• Viitala, E-mail: ev@mm.unknown

Growth data were collected from 40 European aspen (Populus tremula L.) stands growing on eight localities in Sweden. The stands ranged in latitude from 56 to 66°N. The mean age of the stands was 32 years (range, 12–63), the mean stand density 1978 stems ha^-1 (range, 300–6,000), and the mean diameter at breast height (on bark) 17 cm (range, 8–34).

Site index curves were constructed for total age. Curves for H₄₀ (dominant height at 40 years total age) were made for total Sweden. Curves fitted for H₄₀ total age have another shape than curves presented by other Nordic studies. The curves from the present study have slower growth for young aspens than curves from Norwegian and Finnish conditions. For 50–70-year-old aspen stands, curves from the present study indicate taller heights than from Nordic studies.

Classified soil types from the stands were grouped into three groups: sandy till (17), light clay (15) and medium clay till (4). As there was only one stand growing in the fine sand group and one stand in the heavy clay till group and two stands in the silty till group, these stands were not presented with growth curves. There were no statistically significant differences in site index between the three soil type groups. Some recommendations for management of aspen stand are given. Damages caused by moose, fungi and other injuries are discussed as a problem for height yield production and a good timber quality.

• Johansson, E-mail: tj@mm.unknown

The colonisation of a burned clear-cut by ants in southern Finland was monitored using pitfall traps, artificial nest sites, and direct nest sampling from the ground and stumps. Clearcutting and fire seemed to have destroyed wood-ant colonies (Formica rufa group), and also other mature-forest species suffered from fire. Myrmica ruginodis Nylander was able to survive only in less severely burned moist sites, whereas it benefitted from the enhanced light conditions in a non-burned clear-cut. The fire resulted in an essentially ant-free terrain into which pioneering species immigrated. The mortality of nest-founding queens appeared to be high. The results supported the hypothesis that the pioneering species tend to be those that are capable of independent colony founding, followed by species founding nests through temporary nest parasitism. The succession of the burned clear-cut differed from that of the non-burned one, suggesting that habitat selection in immigration and priority effects, i.e. competition, introduce deterministic components in the successional pathways of boreal ant communities.

• Punttila, E-mail: pp@mm.unknown
• Haila, E-mail: yh@mm.unknown

The prefire fungal flora (polypores and corticoid fungi) of 284 dead trees, mainly fallen trunks of Norway spruce (Picea abies (L.) H. Karst.), was studied in 1991 in an old, spruce-dominated mesic forest in Southern Finland. Species diversity of the prefire fungal flora was very high, including a high proportion of locally rare species and four threatened polypore species in Finland.

In 1992 part of the study area (7.3 ha) was clear-cut and a 1.7 ha forest stand in the centre of study area was left standing with a tree volume of 150 m³/ha, and later on (June 1st) in the same year the whole area was burned. Burning was very efficient and all trees in the forest stand were dead one year after the fire. Also, the ground layer burned almost completely.

In 1993 the fungal flora of the 284 sample trees was studied again. Most of the trees had burned strongly and the fungal species diversity and the evenness in community structure had decreased considerably as compared with the prefire community. Species turnover was also great, especially in corticoid fungi. Greatest losses in the species numbers occurred in moderately and strongly decayed trees, in coniferous trees and in very strongly burned trees. Fungal flora of non-decayed and slightly decayed trees, deciduous trees and slightly burned trees seemed to have survived the fire quite well, and in these groups the species numbers had increased slightly as compared with the prefire community.

Fungal species suffering from fire (anthracophobe species) were mainly growing in moderately and strongly decayed trees before the fire, whereas species favoured by fire (anthracophile species) were growing in less decayed trees. No fruitbodies of threatened polypores or other "old-forest species" of polypores were found again after fire. Some very common and effective wood-rotting fungi (e.g. Fomitopsis pinicola, Fomes fomentarius, Antrodia serialis) survived the fire quite well (anthracoxene species). Species favoured by fire were mainly ruderal species which can utilize new, competition-free resources created by fire, and species that have their optima in dry and open places also outside forest-fire areas. Some rarities, e.g. Phanerochaete raduloides and Physisporinus rivulosus, were favoured by fire.

• Penttilä, E-mail: rp@mm.unknown
• Kotiranta, E-mail: hk@mm.unknown

Seasonal fluctuations in free polyamines, spermidine, spermine, putrescine and potassium concentrations were studied for two years (1992–1993) in three needle years of Scots pine (Pinus sylvestris L.) grown on a drained mire in western Finland. Seven different fertilizer treatments involving five different sources of potassium were used.

Putrescine concentrations were high in winter and in May but low in summer. High peaks in putrescine in March and May could be found in unrefertilized or rock phosphate treatments. Spermidine and spermine concentrations were high in March and May. In December spermine concentrations were low. Biotite increased the needle potassium concentrations less than the other potassium fertilizers but the putrescine concentrations or the putrescine/spermidine ratio to about the same level. This suggests that biotite, although very slowly soluble, can reasonably satisfy potassium nutrition of young pine trees.

The potassium concentrations of needles in all the fertilization treatments were higher in winter than in summer. The response of putrescine to the potassium concentration was strongly negative in all the needle years and sampling times. In March, May and December the response of putrescine to potassium was fairly similar in both years but not in June and August. The results suggest that the potassium concentrations during the growing season cannot be used for estimating the potassium nutrition of trees, because the variation between the years may be substantial, whereas the needle putrescine concentration or putrescine/spermidine ratio indicates the suboptimum potassium status of Scots pine fairly well. Needle putrescine concentrations over 500 nmol g^-1FW quite regularly coincided with a unsatisfactory potassium nutrition and concentrations over 1,000 nmol g^-1FW were a reliable indication of potassium deficiency. Putrescine/spermidine ratios below 5 indicated a satisfactory potassium nutrition in all needle years throughout the year.

• Sarjala, E-mail: ts@mm.unknown
• Kaunisto, E-mail: sk@mm.unknown

Addressing the potential impact of climate change on boreal forest ecosystems will require a range of new conservation techniques. During the early 1990s, the scope of WWF's (the World Wide Fund for Nature) forest policy work has broadened from a focus on tropical moist forests to a more general consideration of all the world's forests. Climate change is only one of a series of threats currently facing boreal forests.

Planning conservation strategies that take account of global warming is not easy when there are many computer models of climate change, sometimes predicting very different ecological effects. Climate change could result in some particularly extreme problems for the boreal forest biome. A summary of the problems and opportunities in boreal forests is presented. WWF has also been drawing up strategies for conservation on a global, regional and national level. The organization has concluded that conservation strategies aimed at combatting climate change need not be in direct conflict with other conservation planning requirements. However, proposals have emerged for ways to address the impacts of climate change that would have detrimental impacts on existing conservation plans.

• Dudley, E-mail: nd@mm.unknown
• Jeanrenaud, E-mail: jj@mm.unknown
• Markham, E-mail: am@mm.unknown

Four governmental efforts are underway to reach consensus on indicators of sustainable forestry. Through the Helsinki process, European countries have developed and reached a pan-European, binding consensus, The Montreal process includes non-European Temperate and boreal forest countries, the International Tropical Timber Organization (lTTO) have developed guidelines for the sustainable management of natural tropical forests, and the countries around the Amazon basis have developed a joint initiative for creating guidelines of sustainable forest management of the Amazonian tropical rain forests. It is estimated that as many as 15–20 distinct processes are under way in the private sector by non-profit organizations and for-profit companies, some domestic and other international in scope. Perhaps the most wide-ranging definition work of non-governmental organizations is the undertake by the Forest Stewardship Council, FSC. The paper discusses the Helsinki and Montreal processes and the tasks for research.

• Parviainen, E-mail: jp@mm.unknown

A presentation based on the historical development of Russia is given in the form of an overview of the development of Russian forest resources, of the wood, non-wood, and biological aspects of the forest ecosystem. The list of non-wood forest resources includes resin, saps, oils, berries, wild nuts, mushrooms, hay harvesting, game animals, etc. The dynamics of the system are presented in the light of the data of the Forest State Account (FSA) of Russia for the period 1956–1993. The most significant changes in the dynamics of Russia's forest resources are related to concentrated, large-scale wood harvesting operations. The dynamics of non-wood resources follow the process of the economic recession in all parts of the forest sector of Russia, the said recession having begun in the mid-1980s. The forests of Russia are considered to be of immense social and cultural value and a globally significant factor contributing to the sustainable development of forest resources.

• Pisarenko, E-mail: ap@mm.unknown
• Strakhov, E-mail: vs@mm.unknown

Nearly every forest land in Finland has been burnt down by a wildfire at least once during the past 400–500 years. Slash and burn cultivation (1700–1920) was practised on 50–75 percent of Finland's forests, while prescribed burning (1920–1990) has been applied to 2–3 percent of the country's forests. Because of land-use changes and efficient fire prevention and control systems, the occurrence of wildfires in Finland has decreased considerably during the past few decades. Owing to the biodiversity and ecologically favourable influence of fire, the current tendency is to revive the use of controlled fire in forestry in Finland. Prescribed burning is used in forest regeneration and endeavours are being made to revert old conservation forests to the starting point of succession through forest fires.

• Parviainen, E-mail: jp@mm.unknown

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

Postfire recovery of species diversity (including a number of species, entropy of species relative coverage (Shannon index of species diversity) was studied in lichen and green moss site types of Scots pine (Pinus sylvestris L.) forests in the central part of the Kola Peninsula. The results obtained indicate the difference in the dynamics of characteristics of biodiversity of forest components during postfire recovery. The stabilization of separate components of forest community varies in time from 5–15 to 120–140 years after the fire. Characteristics of the dwarf shrub and herb stratum recovered and stabilized 5–15 years after fire, while the complete stabilization of characteristics of moss-lichen cover is observed in community with fire ages of 90–140 years. Species richness of tree stratum recovered 120–140 years after fire. Time of complete stabilization of species richness of the community was estimated 120–140 years after fire. The size of the area over which characteristics of the biodiversity were estimated effected the mean values and, in most cases, the character of variation of studied characteristics. Over an area of 1 x 1 m dynamics of characteristics of species diversity coincide in forests of the studied types. Regardless of forest type within the area of 100 m² species richness recovered 30 years after the fire (i.e. 3–5 times earlier than the establishment of the complete stabilization of the forest structure). That means that floristic composition of the forest remained unchanged from 30 to 210 years after the fire.

• Gorshkov, E-mail: vg@mm.unknown
• Bakkal, E-mail: ib@mm.unknown

The horizontal and vertical stand structure of living trees was examined in a managed and in a primeval Norway spruce-dominated forest in Southern Finland. Tree size distributions (DBHs, tree height) were compared using frequency histograms. The vertical distribution of tree heights was illustrated as tree height plots and quantified as the tree height diversity (THD) using the Shannon-Weaver formula. The horizontal spatial pattern of trees was described with stem maps and quantified with Ripley's K-function. The spatial autocorrelation of tree sizes was examined with semivariogram analysis. In the managed forest the DBH and height distributions of trees were bimodal, indicating a two-layered vertical structure with a single dominant tree layer and abundant regeneration in the understory. The primeval forest had a much higher total number of trees which were rather evenly distributed in different diameter and tree height classes. The K-function summaries for trees taller than 15 m indicated that the primeval stand was close to complete random pattern. The managed stand was regular at small distances (up to 4 m). The semivariograms of tree sizes (DBH tree height) showed that the managed forest had a clear spatial dependence in tree sizes up to inter-tree distances of about 12 meters. In contrast, the primeval spruce forest had a variance peak at very short inter-tree distances (< 1 m) and only weak spatial autocorrelation at short inter-tree distances (1–5 m). Excluding the understory trees (h < 15 m) from the analysis drastically changed the spatial structure of the forest as revealed by semivariograms. ln general, the structure of the primeval forest was both horizontally and vertically more variable and heterogeneous compared to the managed forest. The applicability of the used methods in describing fine-scale forest structure i discussed.

• Kuuluvainen, E-mail: tk@mm.unknown
• Leinonen, E-mail: kl@mm.unknown
• Nygren, E-mail: mn@mm.unknown
• Penttinen, E-mail: ap@mm.unknown

In a silviculture experiment in east-central Maine, USA, natural regeneration was sampled to measure the effects of: (1) a range of partial harvest intensities, and (2) repeated partial harvest at one intensity. Under the first objective, five treatments were compared with residual basal areas ranging from 15 to 24 m² ha^-1 for trees ≥1.3 cm diameter at breast height. For the second objective, regeneration was evaluated after four harvests at 5-year intervals. Prior to harvests, the overstory of all the treated stands was dominated by Tsuga canadensis (L.) Carr., Picea spp. A Dietr., and Abies balsamea (L.) Mill. Eleven species or species groups were identified among the regeneration: A. balsamea, T. canadensis, Picea spp., Thuja occidentalis L., Pinus spp. L., Betula papyrifera Marsh., Acer rubrum L., Betula populifolia Marsh., Populus spp. L., Fagus grandifolia Ehrh. and Prunus serotina Ehrh. Regeneration abundance was measured as counts of seedlings or sprouts taller than 15 cm but with diameters less than 1.3 cm at breast height (1.37 m). Regardless of harvest treatment, total regeneration was profuse, ranging from over 25,000 to nearly 80,000 trees ha^-1. Regeneration was dominated by conifers with a total angiosperm component of 10 to 52 percent approximately 5 years after harvest and 11 to 33 percent after 10 years. Consequently, in forests of similar species composition, tree regeneration following partial harvests should be sufficiently abundant with an array of species to meet a variety of future management objectives.

• Brissette, E-mail: jb@mm.unknown

As a follow-up on acid rain programmes many countries, e.g. Finland, the Netherlands, Sweden, launched national research programmes on Climate Change by the end of the eighties. Other countries centred new programmes on Global Change, such as Belgium, United Kingdom, Germany, Canada. Also, the European Community included the climate issue in the research programme 'Environment & Climate'. The conclusion of the Intergovernmental Panel on Climate Change (IPCC) shifted in the successive assessment reports from possible climate change to actual climate change. The paper describes the first and second phase of the Dutch Climate Change Research Programme, and discusses the future of the programme.

• Rompaey, E-mail: rr@mm.unknown

There is no doubt that tree survival, growth, and reproduction in North America's boreal forests would be directly influenced by the projected changes in climate if they occur. The indirect effects of climate change may be of even greater importance, however, because of their potential for altering the intensity, frequency, and perhaps even the very nature of the disturbance regimes which drive boreal forest dynamics. Insect defoliator populations are one of the dominating disturbance factors in North America's boreal forests and during outbreaks trees are often killed over vast forest areas. If the predicted shifts in climate occur, the damage patterns caused by insects may be considerably changed, particularly those of insects whose temporal and spatial distributions are singularly dependent on climatic factors. The ensuing uncertainties directly affect depletion forecasts, pest hazard rating procedures, and long-term planning for pest control requirements. Because the potential for wildfire often increases in stands after insect attack, uncertainties in future insect damage patterns also lead to uncertainties in fire regimes. In addition, because the rates of processes key to biogeochemical and nutrient recycling are influenced by insect damage, potential changes in damage patterns can indirectly affect ecosystem resilience and the sustainability of the multiple uses of the forest resource.

In this paper, a mechanistic perspective is developed based on available information describing how defoliating forest insects might respond to climate warming. Because of its prevalence and long history of study, the spruce budworm, Choristoneura fumiferana Clem. (Lepidoptera: Tortricidae), is used for illustrative purposes in developing this perspective. The scenarios that follow outline the potential importance of threshold behaviour, historical conditions, phenological relationships, infrequent but extreme weather, complex feedbacks, and natural selection. The urgency of such considerations is emphasized by reference to research suggesting that climate warming may already be influencing some insect lifecycles.

• Fleming, E-mail: rf@mm.unknown

The impact of carbon sequestration on the financial profitability of four tree plantation cases in Finland and the Philippines were examined. On the basis of stem wood growth; the accumulation of carbon in forest biomass, the formation and decomposition of litter, and the carbon flow in wood-based products were assessed for each reforestation case representing boreal (Finland) and moist tropical conditions (the Philippines). Using different unit values for carbon sequestration the profitability of reforestation was estimated for a fixed 100-year period on a per hectare basis. The financial profitability of reforestation increased notably when the sequestered carbon had high positive values. For example, when the value of carbon sequestration was set to be Twenty-five United States Dollar per megagram of carbon (25 USO/Mg C), the internal rate of return (IRR) of a reforestation investment with Norway spruce (Picea abies (L.) H. Karst.) in Finland increased from 3.2% to 4.1 %. Equally, the IRR of reforestation with mahogany (Swietenia macrophylla King) in the Philippines increased from 12.8% to 15.5%. The present value of carbon sequestration ranged from 39–48% and from 77–101% of the present value of the reforestation cost in Finland and the Philippines, respectively when a 25 USO/Mg C shadow price and a 5% discount rate were applied. Sequestration of one mg of carbon in reforestation in Finland and the Philippines was estimated to cost from 10.5–20.0 and from 4.0–13.6 USO, respectively.

• Niskanen, E-mail: an@mm.unknown
• Rantala, E-mail: tr@mm.unknown
• Saastamoinen, E-mail: os@mm.unknown

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
• Klõseiko, E-mail: jk@mm.unknown
• Reisner, E-mail: vr@mm.unknown
• Tullus, E-mail: ht@mm.unknown

A gap-model was used with forest inventory data in taking ground-true site, soil and tree characteristics into account in predicting the effects of climate change on forests. A total of 910 permanent sample plots established in the course of national forest inventory (NFI) in Finland and located on mineral soil sites in southern Finland were selected as the input data. The climatological input used in the simulations consisted of interpolated means of and deviations from long-term local temperature and precipitation records. The policy-oriented climate scenarios of SILMU (Finnish Research Programme on Climate Change) were used to describe the climate change. The temperature changes in the climate scenarios were increases of ca. +1.1 °C (low), +4.4 °C (medium) and +6.6 °C (high) compared to the current climate in 110 years. The simulation period was 110 years covering the time years 1990–2100.

Southern Finland, divided into fifteen forestry board districts, was used as the study region. Regional development of stand volume, cutting yield, and total wood production of forests under different climate scenarios were examined. The annual average growth in simulations under current climate was close to that observed in NFL Forests benefited from a modest temperature increase (Scenario 2), but under Scenario 1 the growing stock remained at a lower level than under the current climate in all parts of the study region. In wood production and cutting yield there were regional differences. In the southern part of the study regional wood production under Scenario 1 was ca. 10% lower than under the current climate, but in the eastern and western parts wood production was 5–15% higher under Scenario 1 than under the current climate. The relative values of total wood production and cutting yield indicated that the response of forests to climate change varied by geographical location and the magnitude of climate change. This may be a consequence of not just varying climatic (e.g. temperature and precipitation) and site conditions, but of varying responses by different kind of forests (e.g. forests differing in tree species composition and age).

• Talkkari, E-mail: at@mm.unknown

To project the changes in wood production of Norway spruce (Picea abies (L.) H. Karst.) and Scots pine (Pinus sylvestris L.) in Finland as a result of climate change, two separate studies were made. The first study, at the Faculty of Forestry, University of Joensuu, based its projections on mathematical models; the second one, at the Finnish Forest Research Institute, based projections on measurements of wood production in two series of aged provenance experiments. The results of the two studies were similar for both species: after a 4°C increase of the annual mean temperature a drastic increase in wood production in northern Finland, but little effect, or even some decrease in the southern part of the country. However, the assumptions used in the two studies differed. One important difference was that in the models the temperature is assumed to be increasing gradually over the years, whereas in the provenance experiments, climate changed immediately when the seedlings were transferred to the planting sites. Another problem with the provenance experiments is that when material is moved in a north-south direction in Finland, not only temperature but also photoperiod changes markedly. To compare these two studies, site factors (e.g. soil type, temperature, precipitation) and silvicultural factors (e.g. plant spacing, survival, time of thinning, thinning intensity) from the provenance experiments were included a variable in the mathematical models.

• Beuker, E-mail: eb@mm.unknown
• Kellomäki, E-mail: sk@mm.unknown
• Kolström, E-mail: mk@mm.unknown

The productivity of Scots pine (Pinus sylvestris L.) under changing climatic conditions in the southern part of Finland was studied by scenario analysis with a gap-type forest ecosystem model. Standard simulations with the model predicted an increased rate of growth and hence increased productivity as a result of climatic warming. The gap-type model was refined by introducing an overwintering sub-model describing the annual growth cycle, frost hardiness, and frost damage of the trees. Simulations with the refined gap-type model produced results conflicting with those of the standard simulation, i.e., drastically decreased productivity caused by mortality and growth-reducing damage due to premature dehardening in the changing climate. The overwintering sub-model was tested with frost hardiness data from Scots pine saplings growing at their natural site 1) under natural conditions and 2) under elevated temperature condition, both in open-top chambers. The model predicted the frost hardiness dynamics quite accurately for the natural conditions while underestimating the frost hardiness of the saplings for the elevated temperature conditions. These findings show that 1) the overwintering sub-model requires further development, and 2) the possible reduction of productivity caused by frost damage in a changing climate is less drastic than predicted in the scenario analysis. The results as a whole demonstrated the need to consider the overwintering of trees in scenario analysis carried out with ecosystem model for boreal conditions. More generally, the results revealed a problem that exists in scenario analysis with ecological models: the accuracy of a model in predicting the ecosystem functioning under present climatic condition does not guarantee the realism of the model, nor for this reason the accuracy for predicting the ecosystem functioning under changing climatic conditions. This finding calls for the continuous rigorous experimental testing of ecological models used for assessing the ecological implications of climatic change.

• Hänninen, E-mail: hh@mm.unknown
• Kellomäki, E-mail: sk@mm.unknown
• Leinonen, E-mail: il@mm.unknown
• Repo, E-mail: tr@mm.unknown

Willows (Salix phylicifolia) were grown for four months in organic rich soil at four nutrient levels (fertilization with a micronutrient-macronutrient mixture of 0, 100, 500 and 1,000 kg ha^-1 per month) and four CO₂ concentrations (300, 500, 700 and 1,000 ppm). Nitrogen and phosphorus concentration of the willows were reduced at CO₂ enhancement, the decrease being larger in the leave and roots than in the stems. Nitrogen content of the willows plus extractable nitrate-N in the soil coincided well with the doses of nitrogen supplied, but the corresponding sum of phosphorus in the plants and soil were smaller. The total nitrogen content of willows grown in unfertilized soil was nearly two times higher than the sum of the extractable nitrate-N in soil and N content of the cutting at the beginning of the experiment. The contents of nitrogen and phosphorus of the unfertilized willows were independent of CO₂ concentration, suggesting that CO₂ concentration did not affect through increased mineralization the availability of those nutrients to the willows.

• Silvola, E-mail: js@mm.unknown
• Ahlholm, E-mail: ua@mm.unknown

Investigations carried out in the Kola peninsula (northern taiga) and in the South-western part of Western Siberia (southern taiga and forest-steppe) revealed identical course of the postfire restoration process of forest litter thickness in Scots pine (Pinus sylvestris L.) forests. Despite the differences in mean annual temperature (2°C) and other climatic characteristics the recovery time for thickness of forest litter in both regions amounts to 90–100 years after fire in pine forests of lichen site type and 120–140 years – in green moss type; the thickness of forest litter therewith corresponds 3–4 cm and 7–8 cm respectively. That mean that within the natural borders of pine forests, communities of a specific type possess uniform characteristics of restoration. On the basis of empirical data, it appears that the predicted increase of mean annual temperature of earth surface by (2°C) will not bring changes into the character of postfire recovery of forest litter thickness. It was shown that during the period of the recovery, which spans about 90 years after fire in pine forests of lichen and green moss-lichen site types and 140 years in ones of green moss site types, the rate of increasing of carbon store in the forest litter averaged 0.6 t ha^-1 year^-1, 0.1 t ha^-1 year^-1 and 0.2 t ha^-1 year^-1, respectively.

• Gorshkov, E-mail: vg@mm.unknown
• Bakkal, E-mail: ib@mm.unknown
• Stavrova, E-mail: ns@mm.unknown

A system of zonality in Siberia has been formed under the control of continentality, which provides the heat and humidity regimes of the forest provinces. Three sectors of continentality and four to six boreal sub-zone form a framework for the systematization of the different features of land cover in Siberia. Their climatic ordination provides the fundamental basis for the principal potential forest types (composition, productivity) forecasting the current climate. These are useful in predicting the future transformations and succession under global change.

• Nazimova, E-mail: dn@mm.unknown
• Polikarpov, E-mail: np@mm.unknown

An equilibrium model driven by climatic parameters, the Siberian Vegetation Model, was used to estimate changes in the phytomass of Siberian vegetation under climate change scenarios (CO₂ doubling) from four general circulation models (GCM's) of the atmosphere. Ecosystems were classified using a three-dimensional climatic ordination of growing degree days (above a 5 °C threshold), Budyko's dryness index (based on radiation balance and annual precipitation), and Conrad's continentality index. Phytomass density was estimated using published data of Bazilevich covering all vegetation zones in Siberia. Under current climate, total phytomass of Siberia is estimated to be 74.1 ± 2.0 Pg (petagram = 1,015 g). Note that this estimate is based on the current forested percentage in each vegetation class compiled from forest inventory data.

Moderate warming associated with the GISS (Goddard Institute for Space Studies) and OSU (Oregon State Univ.) projections resulted in a 23–26 % increase in phytomass (to 91.3 ± 2.1 Pg and 93.6 ± 2.4 Pg, respectively), primarily due to an increase in the productive Southern Taiga and Sub-taiga classes. Greater warming associated with the GFDL (General Fluid Dynamics Laboratory) and UKMO (United Kingdom Meteorological Office) projections resulted in a small 3–7 % increase in phytomass (to 76.6 ± 1.3 Pg and 79.6 ± 1.2 Pg, respectively). A major component of predicted change using GFDL and UKMO is the introduction of a vast Temperate Forest-Steppe class covering nearly 40% of the area of Siberia, at the expense of Taiga; with current climate, this vegetation class is nearly non-existent in Siberia. In addition, Sub-boreal Forest-Steppe phytomass double with all GCM predictions. In all four climate change scenarios, the predicted phytomass stock of all colder, northern classes is reduced considerably (viz., Tundra, Fore Tundra, northern Taiga, and Middle Taiga). Phytomass in Sub-taiga increases greatly with all scenarios, from a doubling with GFDL to quadrupling with OSU and GISS. Overall, phytomass of the Taiga biome (Northern, Middle, Southern and Sub-taiga) increased 15% in the moderate OSU and GISS scenarios and decreased by a third in the warmer UKMO and GFDL projections. In addition, a sensitivity analysis found that the percentage of a vegetation class that is forested is a major factor determining phytomass distribution. From 25 to 50% more phytomass is predicted under climate change if the forested proportion corresponding to potential rather than current vegetation is assumed.

• Monserud, E-mail: rm@mm.unknown
• Denissenko, E-mail: od@mm.unknown
• Kolchugina, E-mail: tk@mm.unknown
• Tchebakova, E-mail: nt@mm.unknown

Concentration of the phytotoxic air pollutant, ozone (O₃) is continually increasing in the lower layer of the troposphere. The purpose of this study was to compare performance of pine sawflies on Scots pine (Pinus sylvestris L.) seedlings in ambient and future levels of ozone. Scots pine seedlings were grown in field fumigation system where the ozone doses in fumigated plots were 1.5–1.6 times the ambient level. Larvae of the European pine sawfly (Neodiprion sertifer Geoffroy and Gilpinia pallida Klug) were reared on the foliage of Scots pine. The levels of resin acids and monoterpenes in foliage were analysed. There were no significant effects of ozone fumigation on sawfly performance or levels of defence compounds in pine foliage. The results suggest that the elevated ozone concentrations do not strongly affect the needle quality of young Scots pine and the importance of these two diprionid sawfly species forest pests.

• Lyytikäinen, E-mail: pl@mm.unknown
• Holopainen, E-mail: jh@mm.unknown
• Kainulainen, E-mail: pk@mm.unknown
• Nerg, E-mail: an@mm.unknown
• Neuvonen, E-mail: sn@mm.unknown
• Virtanen, E-mail: tv@mm.unknown

The European Pine Sawfly (Neodiprion sertifer Geoffroy) is one of the most serious defoliators of Scots pine (Pinus sylvestris L.) in northern Europe. We studied the pattern in the regional occurrence of the outbreaks of N. sertifer in Finland in years 1961-90, and made predictions about the outbreak pattern to the year 2050 after predicted winter warming. We tested whether minimum winter temperatures and forest type and soil properties could explain the observed outbreak pattern. We analysed outbreak patterns at two different spatial levels: forest board- and municipal-level.

The proportion of coniferous forests on damage-susceptible soils (dry and infertile sites) explained a significant part of the variation in outbreak frequency at small spatial scale (municipalities) but not at large spatial scale (forest boards). At the forest board level, the incidence of minimum temperatures below -36 °C (= the critical value for egg mortality) explains 33% of the variation in the outbreak pattern, and at the municipal level the incidence of cold winters was also the most significant explaining variable in northern Finland. Egg mortality due to cold winters seems to be the most parsimonious factor explaining why there have been so few N. sertifer outbreaks in northern and north-eastern Finland. We predict that climate change (increased winter temperatures) may increase the frequency of outbreaks in eastern and northern Finland in the future.

• Virtanen, E-mail: tv@mm.unknown
• Neuvonen, E-mail: sn@mm.unknown
• Niemelä, E-mail: pn@mm.unknown
• Nikula, E-mail: an@mm.unknown
• Varama, E-mail: mv@mm.unknown

Two dynamic models predicting the development of frost hardiness of Finnish Scots pine (Pinus sylvestris L.) were tested with frost hardiness data obtained from trees growing in the natural conditions of Finland and from an experiment simulating the predicted climatic warming. The input variables were temperature in the first model, and temperature and night length in the second. The model parameters were fixed on the basis of previous independent studies. The results suggested that the model which included temperature and photoperiod as input variables was more accurate than the model using temperature as the only input variable to predict the development of frost hardiness in different environmental conditions. Further requirements for developing the frost hardiness models are discussed.

• Leinonen, E-mail: il@mm.unknown
• Hänninen, E-mail: hh@mm.unknown
• Repo, E-mail: tr@mm.unknown

How will global warming affect southern populations of boreal trees? In paper birch, Betula papyrifera (Betulaceae), alpine trees with an evolutionary history of relatively cool summers may be more sensitive to climate warming than valley populations. We evaluated this scenario by growing seedlings from different populations in four temperature treatments (mountain field site, valley field site, and two greenhouse rooms).

Populations from low elevations germinated earlier and had higher germination success than population from high elevations (16.8 vs. 22.0 d; 72% vs. 11%). At the valley site, seedlings from native populations grew faster than seedlings from higher elevations (mean ± SE = 0.25 ± 0.02 vs. 0.09 ± 0.04 mm · cm^-1 · d^-1) while at the mountain site, all seedlings grew at similar rates. Seedling grown in cooler environments had higher root : shoot ratios, perhaps to compensate for temperature limitations in nutrient uptake by roots. Leaf area varied among populations but was not affected by environmental differences across the field sites. Net photosynthetic rates at valley temperatures were higher for seedlings grown in the valley than for seedling grown in the mountains or the warm greenhouse (12.0 vs. 10.3 and 5.8 μmoles · m^-2 · s^-1), perhaps due to adaptive phenotypic adjustments. Climatic warming could rapidly produce important phenotypic changes in birch trees (e.g. decreased root : shoot ratio, reduced growth in alpine populations). On a longer time-scale, warming could also result in genetic changes as natural selection favours valley genotypes in alpine sites where they are presently rare.

• Ruel, E-mail: jr@mm.unknown
• Ayres, E-mail: ma@mm.unknown

A total of 1,800 3-year old seedlings of Norway spruce (Picea abies (L.) H. Karst.) from two Norwegian and one German provenance were treated with two different nitrogen levels during the 1992 growth season. The plants were kept during the following winter at two different temperature levels. In the spring of 1993, the nutrient application was resumed, and the plants were divided between three different treatments, 350 and 650 p.p.m. in open top chamber and a control plot outside the chambers. This treatment was repeated also during the following 1994 growth season.

The growth and primary production were studied by photosynthesis experiments and by non-destructive growth measurements. The result indicate that raised winter temperature may lead to increased needle loss and reduced growth the following season, particularly in northern provenances. Carbon dioxide significantly influenced growth in addition to nutrient level and winter temperature. High CO2 also seemed to cause increased photosynthesis at early season, and earlier budbreak and growth cessation than in control plants.

• Skre, E-mail: os@mm.unknown
• Nes, E-mail: kn@mm.unknown

The effects of realistically elevated O₃ and CO₂ concentrations on the needle ultrastructure and photosynthesis of ca. 20-year-old Scots pine (Pinus sylvestris L.) saplings were studied during one growth period in open-top field chambers situated on a natural pine heath at Mekrijärvi, in eastern Finland. The experiment included six different treatments: chamberless control, filtered air, ambient air and elevated O₃, CO₂ and O₃ + CO₂. Significant increases in the size of chloroplast and starch grains were recorded in the current-year needles of the saplings exposed to elevated CO₂ These responses were especially clear in the saplings exposed to elevated O₃ + CO₂ concentrations. These treatments also delayed the winter hardening process in cells. In the shoots treated with O₃, CO₂ and combined O₃ + CO₂ the P_max was decreased on average by 50% (ambient CO₂) and 40% (700 ppm CO₂). Photosynthetic efficiency was decreased by 60% in all the treated shoots measured under ambient condition and by 30% in the CO₂ and O₃ + CO₂ treated shoots under 700 ppm. The effect of all the treatments on photosynthesis was depressive which was probably related to evident accumulation of starch in the chloroplasts of the pines treated with CO₂ and combined O₃ + CO₂. But in O₃ treated pines, which did not accumulate starch in comparison to pines subjected to ambient air conditions, some injuries may be already present in the photosynthetic machinery.

• Palomäki, E-mail: vp@mm.unknown
• Holopainen, E-mail: th@mm.unknown
• Kellomäki, E-mail: sk@mm.unknown
• Laitinen, E-mail: kl@mm.unknown

BOREAS is a four-year, regional-scale experiment to study the forested continental interior of Canada. It aims at improving our understanding of the interaction between the earths' climate system and the boreal forests at short and intermediate time scales, in order to clarify their role in global change.

During the winter, spring and summer of 1994, five field campaigns were conducted. About 85 investigation teams including nearly 300 scientists participated, including forest ecologists and ecophysiologists, atmospheric physicists, boundary-layer meteorologists, hydrologists, biochemists, atmospheric chemists and remote sensing specialists.

The findings so far have been significant in terms of their implication for global change. The boreal ecosystem, occupying roughly 17 percent of the vegetated land surface and thus an important driver of global weather and climate, absorbs much more solar energy than is assumed by operational numerical weather prediction models. Albedo measurement show that this forest absorbs nearly 91% of the sun's incident energy. Additionally, while it is known that much of the boreal ecosystems consists of forested wetlands, lakes, bogs and fens, the measurements show that the atmosphere above was extremely dry; humidity and deep boundary layer convection (3,000 m) mimicked conditions found only over deserts. Physiological measurements of the trees show that this atmospheric desiccation was a result of the forests' strong biological control limiting surface evaporation. This tight control was linked to the low soil temperature and subsequently reduced rates of photosynthesis. BOREAS measurement also focused on net ecosystem carbon exchange. Data acquired during the late spring and summer, showed the boreal forests to be a net carbon sink. However, no measurements were taken in the early spring following thaw, and in the late fall, where the balance between photosynthesis and respiration is poorly understood. During 1996 additional data will be acquired to resolve the annual carbon budget and how it might depend on interannual climate differences.

• Hall, E-mail: fh@mm.unknown
• Sellers, E-mail: ps@mm.unknown
• Williams, E-mail: dw@mm.unknown

The eddy covariance technique is a novel micrometeorological method that enables the determination of the atmosphere-biosphere exchange rate of gases such as ozone and carbon dioxide on an ecosystem scale. This paper describes the technique and presents results from the first direct measurements of turbulent fluxes of O₃, CO₂ and H₂O above a forest in Finland. The measurements were performed during 15 July-5 August 1994 above a Scots pine (Pinus sylvestris L.) stand near the Mekrijärvi research station in Eastern Finland.

The expected diurnal cycles were observed in the atmospheric fluxes of O₃, CO₂ and H₂O. The data analysis includes interpretation of the O₃ flux in terms of the dry deposition velocity and evaluation the dependency of the net CO₂ flux on radiation. The eddy covariance method and the established measurement system has proved suitable for providing high-resolution data for studying ozone deposition to a forest as well as the net carbon balance and related physiological processes of an ecosystem.

• Aurela, E-mail: ma@mm.unknown
• Laurila, E-mail: tl@mm.unknown
• Tuovinen, E-mail: jt@mm.unknown

Investigation on the relationship between the size of the Megastigmus suspectus Borr. (Hymenoptera, Torymidae) and the size of Abies alba Mill. seeds from which those insects emerged was conducted. The examined insects (n = 46) originated from southern Poland, and were obtained from cones collected in 1993. M. suspectus thorax length was used as an index of adult insect size, because that was correlated with 8 other body measurements. In addition, the diameter of the emergence hole of this insect was measured. All data were log-transformed and analysed using regression analysis. There was no linear correlation between adult size and seed size, using either length of thorax or any of the other 8 body traits tested.

• Skrzypczynska, E-mail: ms@mm.unknown

In the winter of 1977/78, a White-backed Woodpecker (Dendrocopos leucotos Bechstein) was observed in the archipelago of southwestern Finland 200 km from its breeding areas. It foraged on insects living in mall dead alder and birches. The potential prey species were identified by rearing the insects from the trunks used by the White-backed Woodpecker. Altogether 628 adult insects emerged. In addition to the big larvae the potential food also included larvae of Sciaridae and Cecidomyidae (Diptera) living in dense clumps.

• Nuorteva, E-mail: mn@mm.unknown
• Saari, E-mail: ls@mm.unknown

This paper deals with the testing of dynamic stratification for estimating stand level forest characteristics (basal areas, mean diameter, mean height and mean age) for a 117 ha study areas in Finland. The results do not show possibilities to achieve more accurate estimates using only Landsat TM principal components as auxiliary data opposed to static stratification. It was found that in dynamic stratification non-measured observations should be assigned the mean characteristics of the measured observations that belong to the same cube (class) instead of stratification variable classes until a certain limit. If only one principal component is used the number of classes has, however, little influence. Low field values are overestimated and high values underestimated.

The only successful results were obtained using two variables of different origin – the qualitative development stage class and the quantitative 1st principal component. The lowest root mean square error in estimating basal area was 6.40 m²/ha, mean diameter 3.34 cm, mean height 2.65 m and mean age 14.06 years. This increase of stratification accuracy is mainly resulted by the use of development stage class as an auxiliary variable.

• Mozgeris, E-mail: gm@mm.unknown

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

The prescribed burning of a 7.3 ha clear-cut and a 1.7 ha partially cut forest (volume 150 m³/ha) was carried out in Evo (61 °12'N, 25°07'E) on 1 June 1992. The forest was a mesic Myrtillus site type forest dominated by Norway spruce (Picea abies (L.) H. Karst.). Practically all the trees and the above-ground parts of the understorey vegetation died in the fire, while the mor layer was thinned by an average of 1.5 cm.

A study was made on the change of germinated seedling population in time and their dependence on environmental factors. Seedlings of Norway spruce, Scots pine (Pinus sylvestris L.), silver birch (Betula pendula Roth), pubescent birch (B. pubescens Ehrh.) and rowan (Sorbus aucuparia L.) were inventoried in 1993 and in 1994 on permanent plots, four times per growing season. Autoregression models were used to compare regeneration of tree species in the burned forest with regeneration in the burnt clear-cut area, and to study the effect of distance from nearest seed source to regeneration.

The average number of seedlings germinating in 1993 was higher than in 1994, probably because of differences between these consecutive years in regard to the amount of seed rain and weather conditions. The number of Norway spruce and rowan seedling was higher inside the forest area than in the clear-cut area. The distance to the bordering forest and to the closest seed tree did not explain the result. It is suggested that the more stable microclimatic conditions under the shade of dead tree promote germination and seedling establishment in the forest area. As rowan is a bird-dispersed species, it is likely that dead trees help the dispersal of rowan seed by providing birds place to sit and defecate. The shade provided by dead trees may influence the further succession of the tree stand and vegetation composition and diversity.

• Vanha-Majamaa, E-mail: iv@mm.unknown
• Suominen, E-mail: rs@mm.unknown
• Tonteri, E-mail: tt@mm.unknown
• Tuittila, E-mail: et@mm.unknown

Fresh and herb-rich upland forest sites in the north-western part of the central boreal vegetation zone in Finland were studied with respect to vegetation structure and vegetation-environment relationships (soil, stand characteristics). Two fresh heath vegetation data sets, one from the northern boreal zone and the other from the central boreal zone, were compared with the data of this study using multivariate methods.

The variation in heath forest vegetation within the climatically uniform area was mainly determined by the fertility of the soil (primarily Ca and Mg) and the stage of stand development. N, P and K content of the humus layer varied little between the vegetation classes. Fertile site types occurred, in general, on coarse-textured soils than infertile site types, may be due to the fact that the sample plots were located in various bedrock and glacial till areas, i.e. to sampling effects.

The place of the vegetational units of the study area in the Finnish forest site type system is discussed. The vegetation of the area has features in common with the northern boreal zone as well as the southern part of the central boreal vegetation zone. The results lend some support to the occurrence of a northern Myrtillus type or at least that intermediate form of fresh and herb-rich mineral soil sites commonly occur in the studied area. It is argued that the older name Dryopteris-Myrtillus type is more suitable than Geranium-Oxalis-Myrtillus type for herb-rich heath sites in the study area.

• Lehtelä, E-mail: ml@mm.unknown
• Hotanen, E-mail: jh@mm.unknown
• Sepponen, E-mail: ps@mm.unknown

Germination of Norway spruce (Picea abies (L.) H. Karst.) and Scots pine (Pinus sylvestris L.) pollen decreased during exposure to open air conditions. Usually more than half of the pollen remained germinative after a few days outdoors, but following more than four days outdoors the germination became very low. This study supports the opinion that pollen in the atmosphere remains viable long enough to allow for long-distance gene flow by pollen migration, as an important factor in genetic management of conifers and in evolution, maintaining diversity and potential for adaptation.

• Lindgren, E-mail: kl@mm.unknown
• Lindgren, E-mail: dl@mm.unknown