Habitat factors for land snails in European beech forests with a special focus on coarse woody debris

Analyses of land snails and habitat factors in acid beech forests were conducted in southern Germany (northern Bavaria). The objectives were to study the effects of habitat characteristics on snail density and species richness. Habitat structures were determined for 37 plots in one big forest. We found a significant relationship between the number of snail species and individuals and the following set of habitat factors coverage of herbaceous layer, growing stock, mean diameter at breast height of the three largest trees (DBHmax), stand age, total dead wood volume per ha, and advanced decomposed dead wood volume per ha. We use maximally selected rank statistics to estimate cutpoints separating stands with low densities, from stands with high snail densities. Here, we define cutpoints for a significant higher snail density at a stand age of 187 years, 57 m³/ha dead wood, 40 m³/ha advanced decomposed dead wood, 63 cm DBHmax and more than 1% herbaceous layer. For species richness, cutpoints are estimated at 338 m³/ha stand volume, 170 years stand age, 50 m³/ha total dead wood amount, 15 m³/ha advanced decomposed dead wood and 56 cm DBHmax. The microhabitat analysis shows a higher pH value and a higher Calcium content at the bottom of large snags and under large lying dead wood pieces in comparison to litter, upper mineral soil and at the bottom of vital living trees. Snail species and individual density are significantly linked to these patterns of chemical parameters. The identified cutpoints are a good base for ecological management decisions in forest management.     

Structure of a Central-European mountain spruce old-growth forest with respect to historical development

This study examines the structural characteristics of the tree layer, dead wood, canopy openings, and regeneration patterns of a spruce old-growth forest in the Bohemian Forest, Czech Republic. An old-growth stand with minor human influence and a stand that was presumably logged about 200 years ago were analyzed and compared, as some forest managers considered the presumable human impact as a reason for salvage logging. Even though the stands differed in tree density, height and DBH structure, it was not possible to conclude whether it was due to management history or the environmental differences. The volume of dead wood also differed between the stands. There was about 142 and 83 m³ ha⁻¹ of dead wood in the old-growth stand and presumably logged stand, respectively. The amount of dead wood found in the old-growth stand was comparable with values reported from spruce old-growth stands across Central Europe. In both stands, many canopy trees were arranged in linear patterns, which was a result of spruce regeneration on nurse logs. This suggests that the origin and development of the stands were characterized by natural processes and during the past 200 years typical old-growth structural characteristics have already evolved.     

Diversity and spatio-temporal dynamics of dead wood in a temperate near-natural beech forest (<Emphasis Type="Italic">Fagus sylvatica</Emphasis>)

The diversity, spatial patterns and temporal dynamics of dead wood were examined within the near-natural beech forests (Fagus sylvatica) of Serrahn (North-eastern Germany). Data were collected in an 8 ha sample plot and in two permanent plots (0.36 and 0.25 ha) that had been established at the end of the 1960s. The mean volume of dead wood was 94 m³ ha⁻¹, amounting to 14% of the total volume of all trees. The dead wood displayed a large variation in dead wood type, tree size and decay class. Standing dead wood accounted for about one-third of the total dead wood volume. The densities of standing dead trees were about 10% of the densities of the living trees over a wide range of diameters. The overall spatial distribution of dead trees exhibited a random pattern. Among the different dead wood types, standing entire dead trees and uprooted trees deviated from this pattern by displaying a significantly aggregated pattern. In the permanent plots a high mortality of overstorey trees was observed (1.3% year⁻¹) and the average amount of dead wood increased greatly from 2.9 to 111.6 m³ ha⁻¹ over the 35-year observation period. The near-natural beech forests of Serrahn have experienced a long period of low human interference. Nevertheless, our results suggest that the structure and dynamics of dead wood are strongly affected by the last major disturbance events that took place at the end of the Middle Ages. Information about the forest history is therefore a basic requirement when interpreting the results obtained in near-natural forests.     

Stand characteristics of mixed oriental beech (<Emphasis Type="Italic">Fagus orientalis</Emphasis> Lipsky) stands in the stem exclusion phase,northern Iran

The structure of forest stands changes through developmental phases. This study is carried out in the unmanaged, oriental beech (Fagus orientalis Lipsky) stands in the north of Iran. The aim of this research was to quantify structural characteristics of stands in the stem exclusion phase using common structural indices, which include mingling, tree–tree distance, stem diameter, and tree height differentiation. According to our measurements from three stands, naturally regenerated stands tend to be mixed in species composition have slightly heterogeneous diameter distributions and uniform tree height. The average distance between trees was 3.3 m. Stocking volume of the stands had an average of 540 m³ ha^?1 and 412 stem ha^?1. Dead wood volume was 24 m³ ha^?1, and as a standing volume, the most frequent species in dead wood pool was oriental beech (F. orientalis) (48 %). The common form of dead trees was snag (41 %). The mean value of mingling and tree-to-tree interval indices revealed that beech was mixed intensively with hornbeam and appears to be a more successful competitor for space and light compared with hornbeam; moreover, we found relatively high evidence of inter-species competition in this phase. A better understanding of stand characteristics in the stem exclusion phase as a critical part of the natural dynamics of forest ecosystems could facilitate predictions about the future changes within the stand.     

Spatiotemporal Availability of Dead Wood in Protected Old-growth Forests: A Case Study from Boreal Forests in Eastern Finland

Spatiotemporal patterns of standing and fallen dead trees were examined in two protected Scots pine ( Pinus sylvestris L.)-Norway spruce ( Picea abies (L.) Karst.) forests in eastern Finland (Pahkavaara and Pönttövaara). In Pahkavaara the volume of standing dead trees was 10 m 3 ha -1 and the volume of fallen logs was 66 m 3 ha -1 , and in Pönttövaara the values were 48 m 3 ha -1 and 107 m 3 ha -1 , respectively. The areas differed with respect to the tree species composition, whereas the proportion of different decay stages was similar. Decay stage and dendrochronological analyses revealed the continuity of dead wood formation. The spatial pattern of standing dead trees was usually towards clustered. The volumes of fallen logs showed a spatial autocorrelation up to distances of 10-20 m. The results also suggest that the current amount and diversity of dead wood is rather high, but that forest succession is likely to lead to a less diverse state in the future.     

Stand structure and successional trends in virgin boreal forest reserves in Sweden

Fire history and stand structure was examined in twelve virgin forest stands situated within forest reserves in northern Sweden. The selected stands represented fire refuges as well as different successional stages after fire. Six of the stands were dominated by Norway spruce (Picea abies L. Karst.), three were dominated by Scots pine (Pinus sylvestris L.), and three were dominated by hairy birch (Betula pubescens Ehrh.) or aspen (Populus tremula L.). In 3 of the southernmost stands, the average fire interval was 34 to 65 years during the late 1600s to late 1800s, but since 1888 no fires had occurred in any of the stands. The absence of fire disturbance since 1888 is probably caused by the fire suppression in the overall landscape. The standing volume of living trees ranged between 87 and 511 m³ ha⁻¹ while the volume of dead trees, including both snags and logs, ranged between 27 and 201 m³ ha⁻¹. The volume of dead trees constituted ca. 30% of the total stem volume. In the conifer dominated stands, there was a statistically significant relationship between total stem volume, including both living and dead trees, and site productivity. A comparison between the amount of dead and living trees indicated substantial changes in tree species composition in several stands. It is suggested that data on the amount of dead trees, especially logs, and its distribution over decay classes could be used to examine the continuity of certain tree species. All stands had a multi-sized tree diameter distribution, which in most cases was similar to a reversed J-shaped distribution. In general spruce was numerous in the seedling cohort and in small diameter classes, indicating that its proportion in the stands was stable, or was increasing at the expense of pioneer tree species such as pine, aspen and silver birch (Betula pendula Roth.). The most numerous species in the seedling cohort, rowan (Sorbus aucuparia L.), was almost totally missing in the tree layer, indicating a high browsing pressure preventing rowan seedlings from growing into trees. The general increase of spruce and the sparse regeneration of pioneer species, in the stands previously affected by fire, are discussed in relation to natural disturbance regimes, biological diversity and nature conservation policies. It is proposed that reintroduction of fire disturbance is a necessity for future management plans of forest reserves. Other management practices to increase species diversity within forest reserves are also discussed.     

Short- and long-term effects of thinning and prescribed fire on carbon stocks in ponderosa pine stands in northern Arizona

Euro-American logging practices, intensive grazing, and fire suppression have increased the amount of carbon that is stored in ponderosa pine (Pinus ponderosa Dougl. Ex Laws) forests in the southwestern United States. Current stand conditions leave these forests prone to high-intensity wildfire, which releases a pulse of carbon emissions and shifts carbon storage from live trees to standing dead trees and woody debris. Thinning and prescribed burning are commonly used to reduce the risk of intense wildfire, but also reduce on-site carbon stocks and release carbon to the atmosphere. This study quantified the impact of thinning on the carbon budgets of five ponderosa pine stands in northern Arizona, including the fossil fuels consumed during logging operations. We used the pre- and post-treatment data on carbon stocks and the Fire and Fuels Extension to the Forest Vegetation Simulator (FEE-FVS) to simulate the long-term effects of intense wildfire, thinning, and repeated prescribed burning on stand carbon storage.The mean total pre-treatment carbon stock, including above-ground live and dead trees, below-ground live and dead trees, and surface fuels across five sites was 74.58 Mg C ha⁻¹ and the post-treatment mean was 50.65 Mg C ha⁻¹ in the first post-treatment year. The mean total carbon release from slash burning, fossil fuels, and logs removed was 21.92 Mg C ha⁻¹. FEE-FVS simulations showed that thinning increased the mean canopy base height, decreased the mean crown bulk density, and increased the mean crowning index, and thus reduced the risk of high-intensity wildfire at all sites. Untreated stands that incurred wildfire once within the next 100 years or once within the next 50 years had greater mean net carbon storage after 100 years compared to treated stands that experienced prescribed fire every 10 years or every 20 years. Treated stands released greater amounts of carbon overall due to repeated prescribed fires, slash burning, and 100% of harvested logs being counted as carbon emissions because they were used for short-lived products. However, after 100 years treated stands stored more carbon in live trees and less carbon in dead trees and surface fuels than untreated stands burned by intense wildfire. The long-term net carbon storage of treated stands was similar or greater than untreated wildfire-burned stands only when a distinction was made between carbon stored in live and dead trees, carbon in logs was stored in long-lived products, and energy in logging slash substituted for fossil fuels.     

Growth and productivity of black spruce in even- and uneven-aged stands at the limit of the closed boreal forest

The increasing commercial interest and advancing exploitation of new remote territories of the boreal forest require deeper knowledge of the productivity of these ecosystems. Canadian boreal forests are commonly assumed to be evenly aged, but recent studies show that frequent small-scale disturbances can lead to uneven-aged class distributions. However, how age distribution affects tree growth and stand productivity at high latitudes remains an unanswered question. Dynamics of tree growth in even- and uneven-aged stands at the limit of the closed black spruce (Picea mariana) forest in Quebec (Canada) were assessed on 18 plots with ages ranging from 77 to 340 years. Height, diameter and age of all trees were measured. Stem analysis was performed on the 10 dominant trees of each plot by measuring tree-ring widths on discs collected each meter from the stem, and the growth dynamics in height, diameter and volume were estimated according to tree age. Although growth followed a sigmoid pattern with similar shapes and asymptotes in even- and uneven-aged stands, trees in the latter showed curves more flattened and with increases delayed in time. Growth rates in even-aged plots were at least twice those of uneven-aged plots. The vigorous growth rates occurred earlier in trees of even-aged plots with a culmination of the mean annual increment in height, diameter and volume estimated at 40–80 years, 90–110 years earlier than in uneven-aged plots. Stand volume ranged between 30 and 238 m³ ha⁻¹ with 75% of stands showing values lower than 120 m³ ha⁻¹ and higher volumes occurring at greater dominant heights and stand densities. Results demonstrated the different growth dynamics of black spruce in single- and multi-cohort stands and suggested the need for information on the stand structure when estimating the effective or potential growth performance for forest management of this species.     

Availability of standing trees for large cavity-nesting birds in the eastern boreal forest of Québec,Canada

Large cavity-nesting birds depend on large-diameter trees for suitable nest sites. The increased spatial extent of commercial timber harvesting is modifying forest structure across the land base and may thus compromise the availability of large trees at the landscape scale. In this study, our objectives were to (1) characterize the availability of large living and dead trees in old-growth stands dominated by different tree species and surficial deposits that encompass the range of natural cover types of eastern Québec's boreal forest; (2) analyze the distribution of trees among decay-classes; and (3) compare the availability of large trees in unharvested, remnant, and harvested stands for the entire range of decay-classes. A total of 116 line transects were distributed across unharvested forests, remnant linear forests, and cutblocks in cutover areas. Unharvested forest stands (black spruce [Picea mariana], balsam fir [Abies balsamea]–black spruce, balsam fir–white spruce [Picea glauca] and balsam fir) reflected a gradient of balsam fir dominance. The remnant forests selected were isolated for 5–15 years. Analyses were performed at two diameter cut-off values. Trees with DBH ≥20 cm were considered for availability of total trees whereas trees with DBH ≥30 cm were considered for availability of large trees. Forest stands comprised high proportions of standing dead trees (33% of all stems, 8% were large dead stems). Availability of total and large standing trees increased with the dominance of balsam fir in stands. Forest stands located on thick surficial deposits showed higher densities of large dead trees for every stand type suggesting a higher productivity on those sites. Availability of stems according to decay-classes showed a dome-shaped distribution with higher densities of snags in intermediate decay stages. However, for large stems, black spruce stands showed a significantly lower availability that was consistent across all decay-classes. In linear remnant forests, pure balsam fir stands were absent. Remnant stands thus showed a much lower availability in large trees when compared with unharvested balsam fir stands. Clearcuts had the lowest densities of dead trees across sampled stands. Current even-aged management practices clearly affect availability and recruitment of large trees, therefore forest-dwelling wildlife relying on these structures for breeding is likely to be affected by large-scale harvesting in coniferous boreal forests.     

Dead trees and protected polypores in unmanaged north-temperate forest stands of Lithuania

The availability of coarse woody debris (CWD) and distribution of dead trees into categories of mortality (dead standing, broken and uprooted) were investigated in north-temperate forests of central Europe (Lithuania). The studied area comprised 188.7 ha and included 18 different stands 40–130 years of age with a variety of tree species (spruce (Picea abies (L.) Karst.), pine (Pinus sylvestris L.), alder (Alnus glutinosa (L.) Gaertn.), birch (Betula pendula Roth and B. pubescens Ehrh.), aspen (Populus tremula L.), oak (Quercus robur L.), forest types (caricus-sphagnum, vaccinium-myrtillus, oxalis, myrtillus-oxalis, caricus-calamagrostis) and edaphic conditions (peaty, sandy, loamy soils of different moisture). The stands were excluded from wood harvesting for at least 30 years. A total of 11 365 dead trees (over 10 cm in DBH) or 6160.7 m³ of dead wood was found (60.2 trees/ha and 32.6 m³/ha). The volume of CWD per hectare was larger in older stands (r_S=0.78, P<0.01). Tree mortality during the last 2 years consisted of 482 trees and 381 m³, or 1.28 trees/ha×year and 1.01 m³/ha×year. In 25–33% of cases it was wind-related. Uprooted and broken trees were of larger DBH than dead standing. The distribution into the categories of mortality was strongly dependent on tree species (chi-square test, d.f.=10,P=0). Dead standing dominated in CWD of pine and alder. Broken trees comprised almost a half in CWD of aspen, and about one-third in birch, alder and oak. Uprooting most often occurred in spruce, aspen and birch. Edaphic conditions and stand age had a pronounced impact on distribution into mortality categories for spruce (chi-square test, d.f.=20, P<0.00001) and pine (d.f.=8, P≤0.0003). On peat soil, only a minority of trees of both pine and spruce was uprooted, and standing dead prevailed. In CWD of spruce and pine, the proportions of both dead standing and broken decreased and that of uprooted trees increased on mineral soils of higher moisture and bulk density in older stands. By contrast, uprooting in birch and alder occurred less often on more wet sites, where the proportions of standing snags were higher. A total of 41 species of wood-decomposing polypores were found in the study area. Among those, 10 (24%) were of conservation value.     

Tree mortality after low-intensity prescribed fires in managed Pinus sylvestris stands in southern Finland

Abstract

Tree mortality, its causes, and the input of dead charred wood were studied in 11 managed 30–45-year-old Scots pine (Pinus sylvestris L.) stands 1 year after experimental low-intensity prescribed burnings in southern Finland. First, the relationship between fire-induced tree damage and several external variables, e.g. stand density, within-stand wind speed, open-air wind speed, the Finnish Forest Fire Index (FFI) and flame height, was studied. Secondly, the study examined which damage and morphological characteristics best predicted tree mortality. Tree mortality was very variable in the experimental plots, ranging from 0% to 48% on the basis of stem number and from 0% to 41% in terms of wood volume. The input of dead and charred wood decreased with stand age, being 19.4 m³ ha^?1 in 30–35-year-old stands, but only 1.7 m³ ha^?1 in 45-year-old stands. The input of dead wood was on average 10 m³ ha^?1, representing less than 5% of the mean volume before the prescribed fire. The external variables that best explained fire-induced damage were within-stand wind speed, flame height and FFI. Tree mortality was best predicted by charred stem ratio with bark thickness, and by charred stem ratio with tree diameter. The results indicate that prescribed burning that is conducted downwind increases tree mortality and changes subsequent stand structure with increasing within-stand wind speed.     

Deadwood in New Zealand's indigenous forests

We present the results of a systematic, unbiased national survey of deadwood volume and biomass in New Zealand's remaining indigenous forests based on an 8-km grid of 894 permanent plots. New Zealand's old growth evergreen temperate forests are largely comprised of long-lived, slow-growing tree species typically growing in cool, humid conditions; collectively these conditions are thought to promote accumulation of high deadwood stocks. We estimated deadwood biomass and volume in New Zealand's forests and compared these stocks with published values from other broadleaved evergreen temperate forests. Mean deadwood biomass in New Zealand was 54 Mg ha⁻¹ but ranged across plots from 0 to 550 Mg ha⁻¹. Mean deadwood volume was 158 m³ ha⁻¹ and ranged across plots from 0 to 1890 m³ ha⁻¹. Fallen logs accounted for 63% of total deadwood volume and 65% of total deadwood biomass, with standing dead trees being the remainder. Each piece of deadwood was classified into one of three broad decay classes and >40% of deadwood was fallen logs of the intermediate decay class. Deadwood biomass and volume varied 1.8- and 1.9-fold, respectively, among forest types and was greatest in broadleaved forests, dominated by Weinmannia racemosa (Cunoniaceae), Metrosideros umbellata (Myrtaceae) and Metrosideros robusta, and broadleaf-Nothofagus (Nothofagaceae) forests supporting the large tree species Nothofagus fusca. Deadwood biomass and volume were least in broadleaf-conifer admixtures. We used structural equation models to determine whether deadwood biomass could be predicted from climate and environment (vapor pressure deficit, elevation and slope), live tree biomass, forest composition (captured by two ordination axes), wood density of live trees, and tree size (a proxy for stand age). The model that best fit the data retained only vapor pressure deficit, live tree biomass and the first ordination axis as predictors of deadwood biomass. However, this model predicted just 2.4% of the variation in deadwood biomass, suggesting that additional factors not captured by this dataset, such as disturbance dynamics, may control deadwood abundance. Comparisons with other temperate and tropical forests did not support the hypothesis that New Zealand's cool temperate rainforests support higher than expected biomass or volume of deadwood.     

Stand structure of hemiboreal old-growth forests: Characteristic features,variation among site types,and a comparison with FSC-certified mature stands in Estonia

To improve the silvicultural targets for ecologically sustainable forestry, we quantified functionally important structural features for the first time in a representative set of old-growth forests in hemiboreal Europe. Altogether, 23 old-growth stands of four site-type groups were compared with mature commercial stands nearby in the Estonian state forests that hold the Forest Stewardship Council (FSC) certificate of sustainable forestry. These two treatments did not differ significantly in terms of tree-species diversity, volumes of woody debris of <20 cm diameter (including fine woody debris) and its decay-stage composition. However, mature stands had many more early-successional trees and lacked late-successional deciduous species; they also had a higher overall density and volume of live trees, due to abundant individuals of 10–39 cm diameter at breast height. Old-growth stands had at least twice as many live trees ≥40 cm, standing dead trees ≥30 cm and lying wood ≥20 cm in diameter, any freshly fallen debris, and regeneration. For lying wood ≥20 cm in diameter, the treatment effect depended on site type: both treatments of Vaccinium-type dry boreal forests were remarkably deadwood-poor (indicating historical management of the old-growth stands), while mature eutrophic stands of Aegopodium-type were most impoverished relative to old-growth levels. We conclude that many functional characteristics of old growth were present in the FSC-certified, mostly naturally regenerated, commercial stands. The main problem is the lack of very large trees, particularly of late-successional deciduous species, which should be addressed by their well-planned retention in cut areas and reconsideration of salvage logging strategies. A dense regeneration in old-growth stands also indicated the potential of selection cuttings. The study highlighted the need for region- and site-type specific numerical targets for sustainable forest management, which in the hemiboreal region should address the characteristic occurrence of late-successional deciduous trees on fertile soils and higher natural deadwood volumes than in typical boreal forests. For certification, the issues of structural impoverishment revealed both the inadequacy of some silvicultural practices and some indicators set by the national FSC-standard in Estonia.     

Accumulation of dead wood in abandoned beech (Fagus sylvatica L.) forests in northwestern Germany

Currently, there is much debate about what strategy is most suitable for increasing old-growth attributes in forests that have been managed intensively for wood production in the past. Passive restoration, i.e. cessation of forestry interventions, should be considered when the old-growth attributes desired can be restored within a feasible period of time.Our study focuses on standing and lying coarse dead wood (≥20 cm diameter) in beech-dominated forests in northwestern Germany. We analyzed monitoring data of 545 sample plots (sized 500-1000 m²) from 12 strict forest reserves (SFRs). The SFRs had been without forestry intervention for up to 28 years.Both, number of dead objects and volume of dead wood (m³ ha⁻¹) increased significantly with ongoing time since abandonment from forestry interventions. The mean amount doubled from 9 to 18 m³ ha⁻¹ within 10 years. The proportion of standing dead wood was about 40% of the total dead wood pool ≥20 cm diameter.With mixed linear modeling we showed that dead wood increased by a mean net rate of about 1 m³ ha⁻¹ a⁻¹. Therefore, after three decades critical values for restoring the dead wood pool could be reached. We hypothesized that the rate of dead wood input is mainly determined by disturbance driven tree mortality such as oak decline, bark beetle infestations and storms.A comparison with primeval forests or reserves abandoned more than 100 years ago showed that the SFRs studied are at the beginning of a long process of dead wood accumulation.Based on our results, the abandonment of forest activities in harvestable pure and mixed beech stands is an effective strategy for restoring the dead wood pool.     

Assemblages of wood-inhabiting fungi related to silvicultural management intensity in beech forests in southern Germany

Current silvicultural treatments in beech forests are aimed at achieving thick logs without discoloured hardwood. Therefore intensive thinning is applied already in younger stands with the objective of large-sized trunks at an age of 100 years. However, this approach bears the risk that dead wood structures and broken trees are completely removed from the forest. The impact of three different silvicultural management intensity levels on wood-inhabiting fungi over decades was investigated in a large beech forest (>10,000 ha) in southern Germany in 69 sampling plots: A Intensive Thinning and Logging with high-value trees, B Conservation-Oriented Logging with integration of special structures such as dead wood and broken trees and C Strict Forest Reserves with no logging for 30 years. The analysis of community showed marked differences in the fungus species composition of the three treatments, independent of stand age. The relative frequencies of species between treatments were statistically different. Indicator species for naturalness were more abundant at sites with low silvicultural management intensity. Fomes fomentarius, the most common fungus in virgin forests and strict forest reserves, is almost missing in forests with high-management intensity. The species richness seemed to be lower where intensive thinning was applied (P = 0.051). Species characteristic for coarse woody debris were associated to low management intensity, whereas species with a significant preference for stumps became more frequent with increasing management intensity. A total amount of dead wood higher than 60 m³/ha was found to enable significantly higher numbers of species indicators of naturalness (P = 0.013). In conclusion, when applying intensive silvicultural treatment, the role of dead wood needs to be actively considered in order to maintain the natural biocoenosis of beech forests.     

Natural development and regeneration of a Central European montane spruce forest

Montane Norway spruce forests of Central Europe have a very long tradition of use for timber production; however, recently there has been increasing concern for their role in maintaining biological diversity. This concern, coupled with recent severe windstorms that led to wide-spread bark beetle outbreaks, has brought the management of montane spruce forests to the forefront of public policy discussions in Central Europe. In order to shed light on the natural development and current structure of mature montane spruce forests, we established four 0.25 ha research plots in a semi-natural montane spruce forest in the Šumava Mountains (The Bohemian Forest), Czech Republic. We mapped all trees, extracted increment cores for age and growth-pattern analyses, and inventoried all current tree regeneration, including the substrates on which it was found. Stands were characterized by uni-modal tree diameter distributions and high basal areas (56.6 m² ha⁻¹ on average), indicating a natural transition from the stem exclusion phase towards the understory reinitiation phase. The stands showed largely single-cohort recruitment age structures, however, with recruitment spanning seven decades. Our analyses suggest that this cohort existed as advance regeneration prior to major disturbances in the late 1800s, which included post-bark beetle salvage logging. Spatial pattern analyses of living and dead stems combined, showed an increase in uniformity of living trees, pointing to the role of natural density-dependent mortality. However, past growth patterns and historical documentation suggest that low intensity canopy disturbances (wind and snow) also caused mortality and diversified canopy structure. Because the stands developed naturally over the past 120+ years and thus escaped thinning operations, high volumes of coarse woody debris (94 m³ ha⁻¹) and snag densities (546 stems ha⁻¹) have accrued. Advance spruce regeneration was quite abundant and existed primarily on deadwood substrates, even though these occupied only a small percent of stand area. Because of salvage logging in the late 1880s, these stands do not qualify, according to the traditional paradigm, as natural spruce forests. As a result, they are recently subject to active management practices including salvage logging that remove dead and dying trees. Given the importance of deadwood for forest regeneration and recovery from disturbance, as demonstrated in this study, we argue that dead wood removal may limit future natural regeneration in these stands. Thus, the purported benefits of removing dead and dying trees from semi-natural forests must be carefully weighed against the potential detrimental impacts on natural spruce forest regeneration and biodiversity.     

Functions for Biomass Estimation of Young Pinus sylvestris,Picea abies and Betula spp. from Stands in Northern Sweden with High Stand Densities

New silvicultural regimes with high within-stand competition require new functions for estimation of standing stock and growth of biomass components, since the allometry of trees is changed by light competition. This paper presents functions for estimation of the aboveground biomass dry weights for stem wood, stem bark, branches and leaves of young (diameter at breast height <10 cm) Scots pine (Pinus sylvestris L.), Norway spruce [Picea abies (L.) Karst.] and birch (Betula pendula Roth. and Betula pubescens Ehrh.) trees growing in dense mixed stands. The functions were derived from a sample consisting of 84 Scots pine, 43 Norway spruce and 66 birch trees from six stands in northern Sweden with high stand densities (>10000 st ha^-1). The logarithmically transformed power function displayed a good ability to stabilize the variance of dry weights and showed a good fit to the material (0.37< R ² <0.99). A comparison with the most commonly used biomass functions in Sweden today showed that they overestimated the weight of stem wood and branches, while the weight of foliage was underestimated. The nature of these discrepancies suggested that the precision of biomass estimations might also be improved for young trees at wider spacing.     

Site, plot, and individual tree yield reduction of interior Douglas-fir associated with non-lethal infection by Armillaria root disease in southern British Columbia

Root pathogens are one of the principle factors affecting forest productivity in many forests, but few estimates of impact are available. Non-lethal root infections associated with Armillaria root disease were studied to determine their effect on stem volume yield in seven planted Douglas-fir stands and a naturally regenerated stand in British Columbia's southern interior. Trees were removed from the soil and the infection date of a random selection of trees was determined. The volume reduction attributable to disease was determined as a comparison of diseased to disease-free trees over time since infection. Volume reductions per tree ranged from 0 to 30 dm³ (0-27%) depending on the tree age and disease duration. Yield reduction reached 27 m³/ha, averaging 15 m³/ha for the three oldest planted sites by age 30 (7-15%), but was lower at the naturally regenerated site. Yield reduction at the site level correlated best with the number of diseased trees and an unknown site factor. Sites with slow juvenile growth had the least yield reduction owing to their lower incidence of disease over time. Yield was less affected by the proportion of diseased primary roots per tree than by the cumulative time since infection. A few of the diseased trees maintained growth rate after infection similar to disease-free trees; interestingly, these trees were smaller than average to begin with. Overall, trees suffer accumulating growth reduction without recovery. Root diseases prevent full expression of site potential even without mortality. Minimizing disease impact in respect to other forest management goals is also discussed.     

The choice of definition has a large effect on reported quantities of dead wood in boreal forest

A survey was conducted to assess the impact of the choice of definition on reported quantities of dead wood in Swedish forests, which to more than 90% are located in the boreal zone. The data collection was made on a subsample of the permanent plots of the Swedish national forest inventory. The objects included were standing dead trees and snags down to 5-cm diameter at breast height, dead lying stems and branches down to a threshold diameter of 1 cm and stumps down to a threshold diameter of 5-cm at normal stump height. Standing trees, snags and stumps were inventoried on 10-m radius circular plots while the downed objects were inventoried using both circular plots and line intersect sampling; thin objects (diameter 1–5 cm) were assessed only through line intersect sampling. The results showed that the estimated volume of dead wood was as high as 25 m³ ha^?1 when all components were included. With the standard Swedish definition, the corresponding estimate was only 10.9 m³ ha^?1, or 43% of the total value. Since definitions of dead wood vary greatly between countries we conclude that great caution must be exercised when figures are compared in connection with international reporting. For example, adding stumps to the Swedish definition would increase the amounts of dead wood from 10.9 to 15.7 m³ ha^?1, i.e. with 44%.     

A silvicultural strategy for managing uneven-aged beech-dominated forests in Thuringia,Germany: a new approach to an old problem

Maintaining a permanent forest canopy cover and eventually harvesting wood in a final harvest according to predefined dimensions is often considered as prototype for future management of deciduous forests. An uneven-aged structure is considered by the public to resemble “natural” conditions, and by forest engineers it is considered as being more resilient to disturbances. In the Hainich-Dün region of Thuringia, Germany, beech-dominated selection forests covering about 10,000?ha have been managed for almost 1000 years, initially by irregular use, but as regular selection system since about 200 years. Managing these stands remains difficult, due to the lack of yield tables and a quantification of harvest of uneven-aged stands considering differences in site conditions and handling of over-sized trees. It is the objective of the present study to develop tables of target stand volumes, increments, and harvest for different diameter ranges of uneven-aged stands according to site conditions. The present study is based on repeated grid-based inventories of about 2150 plots, which were partly re-inventoried 3 times over the past 20 years. The recommended target wood volumes vary between 296 and 388 m³ ha^?1. Stand growth rates of different yield classes were estimated to range between 6.7 and 7.7 m³ ha^?1 yr^?1 which is 30% lower than for age class forest. Nevertheless, the economic returns are higher. Thus, selective cutting with single tree selection remains a viable silvicultural system, but it may change over time into small-scale shelter-woods for improving growth of regeneration.