Effects of Forest Management on Field Layer Vegetation: A Comparison Between Finnish and Russian Karelia

The effects of forest management on the field layer vegetation were compared between Finnish and Russian Karelia in a total of 46 mature stands representing pine, spruce and deciduous forests. In Finland, the number of species was slightly higher in each forest type. Many grassland species, such as Galium boreale , Lathyrus pratensis , and Veronica officinalis , were more abundant in Finnish deciduous stands (14.4 species, mean cover 15.8%) than in Russian stands (9.6 species, mean cover 11.5%). Dwarf shrubs, such as Calluna vulgaris (mean cover: Fin 3.3%, Rus 7.7%) and Vaccinium vitis-idaea (Fin 25.0%, Rus 26.9%) in pine stands, and Vaccinium myrtillus (Fin 22.9%, Rus 31.5%) in spruce stands, were more abundant in Russia. Ferns and graminoids exhibited no differences between the two countries. The observed differences were likely to result from the lower tree stand density, more extensive fragmentation and fertilisation of forests in Finland, the higher frequency of forest fires and more intensive grazing in Russian forests, and from differences in natural conditions between Finnish and Russian Karelia.     

Evaluation of mature conifer plantations as secondary habitat for epigeic forest arthropods (Coleoptera: Carabidae; Araneae)

Spiders (Araneae) and ground beetles (Coleoptera, Carabidae) were studied in a woodland of the Northwest German lowland. An ancient oak–beech stand (170 years old) growing on mineral soil as well as a 110-year-old Scots pine (Pinus sylvestris) and a 55-year-old spruce (Picea spp.) forest growing on peat were investigated by pitfall trapping. A total of 155 species (39 carabids, 116 spiders) and 16,887 individuals (5269 carabids, 11,618 spiders) was recorded. Beetle diversity was high in the oak–beech stand and the spruce forest, but significantly lower in the pine forest. In both conifer plantations the activity density of carabids was considerably lower. Spider diversity was significantly higher in the spruce forest when compared to the beech and pine stand, respectively. Analyses of assemblage similarity distinguished clearly the fauna of all three stands. For each forest type, indicator species were detected. Although both conifer plantations were planted on former bogs, spider species typical of bogs were present only in the pine stand, not in the spruce stand. In both animal taxa, species typical of deciduous forests were more numerous and abundant in the oak–beech stand when compared to the conifer plantations. Although they were in direct contact, the conifer stands on peat only to a very low extent serve as secondary habitats for the epigeic fauna of the autochthonous deciduous woodland. During the 1990s, various agricultural programmes in Central Europe promoted such conifer plantations – in contrast, such afforestation measures on extensively used or fallow land of former bogs are not supported by the results of this study.     

Variation in post-wildfire regeneration of boreal mixedwood forests: underlying factors and implications for natural disturbance-based management

To test the direct regeneration hypothesis and support natural disturbance-based forest management we characterized the structure and composition of boreal mixedwood forests regenerating after large wildfires and examined the influence of pre-fire stand composition and post-fire competing vegetation. In stands which had been deciduous (Populus sp.)-dominated, conifer (white spruce)-dominated, or mixed pre-fire we measured regeneration stocking (presence in 10 m² plots), density and height 10–20 years post-burn in five wildfires in Alberta, Canada. Most plots regenerated to the deciduous or mixed stocking types; plots in the older fire and in stands that were pure conifer pre-fire had higher amounts of conifer regeneration. Surprisingly, regeneration in pre-fire ‘pure’ white spruce stands was most often to pine, although these had not been recorded in the pre-fire inventory. Pre-fire deciduous stands were the most resilient in that poplar species dominated their post-fire regeneration in terms of stocking, density and height. These stands also had the highest diversity of regenerating tree species and the most unstocked plots. High grass cover negatively affected regeneration density of both deciduous and conifer trees. Our results demonstrate the natural occurrence of regeneration gaps, pre- to post-fire changes in forest composition, and high variation in post-fire regeneration composition. These should be taken into consideration when developing goals for post-harvest regeneration mimicking natural disturbance.     

Tree regeneration and future stand development after bark beetle infestation and harvesting in Colorado lodgepole pine stands

In the southern Rocky Mountains, current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks and associated harvesting have set millions of hectares of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) forest onto new stand development trajectories. Information about immediate, post-disturbance tree regeneration will provide insight on dynamics of future stand composition and structure. We compared tree regeneration in eight paired harvested and untreated lodgepole pine stands in the Fraser Experimental Forest that experienced more than 70% overstory mortality due to beetles. New seedlings colonized both harvested and untreated stands in the first years after the beetle outbreak. In harvested areas the density of new seedlings, predominantly lodgepole pine and aspen, was four times higher than in untreated stands. Annual height growth of pine and fir advance regeneration (e.g., trees established prior to the onset of the outbreak) has doubled following overstory mortality in untreated stands. Growth simulations based on our regeneration data suggest that stand basal area and stem density will return to pre-beetle levels in untreated and harvested stands within 80-105 years. Furthermore, lodgepole pine will remain the dominant species in harvested stands over the next century, but subalpine fir will become the most abundant species in untreated areas. Owing to terrain, economic and administrative limitations, active management will treat a small fraction (<15%) of the forests killed by pine beetle. Our findings suggest that the long-term consequences of the outbreak will be most dramatic in untreated forests where the shift in tree species composition will influence timber and water production, wildfire behavior, wildlife habitat and other forest attributes.     

The response of ground vegetation to structural change during forest conversion in the southern Black Forest

The aim of this study is to investigate the effects of forest conversion on forest floor vegetation. ‘Ecological’ forest conversion, as defined by an interdisciplinary southern Black Forest project group, describes the transformation of even-aged spruce (Picea abies L. H. Karst.) stands to structured continuous cover forests consisting of spruce (Picea abies), silver fir (Abies alba Mill.) and beech (Fagus sylvatica L.). In order to analyse the conversion process, four conversion stages were defined in a conceptual forest development model. Four forest districts deemed to be representative of the southern Black Forest region were selected for the study. The ground vegetation was initially classified independently from the stand structures. Subsequently, the relationship between stand structures, as determined by development stage, and ground vegetation was analysed. It was revealed that forest conversion modified the ground vegetation. The main factors influencing ground vegetation were the influence of broadleaves, predominately beech (F. sylvatica), on the canopy composition and litter coverage on the one hand; and the canopy coverage of spruce, the proportion of needle litter and the associated light penetration on the other. The prevalence of moss and vascular plant species preferring acidic sites found in spruce forests decreased during the transition process, whereas, species requiring a moderate base supply increased in abundance. The continuous cover forest representing the final stage of conversion increasingly contained a mixture of ground vegetation species normally associated exclusively with either conifer or deciduous forest. Due to the fact that there is an associated ground vegetation specific to the different stages of forest conversion in stands dominated by Norway spruce or European beech and a mixture in the latest conversion stages, large-scale forest conversion should be avoided in favour of management promoting a diversity of silvicultural goals and treatments in neighbouring stands. Only a variety of treatments ensures the maintenance of floristic diversity in the long-term.     

Carbon and nitrogen dynamics in topsoils along forest conversion sequences in the Ore Mountains and the Saxonian lowland, Germany

Carbon and nitrogen stocks and their medium-term and readily decomposable fractions in topsoils were compared in relation to soil microbial biomass and activity along sequences from coniferous to deciduous stands. The study was carried out in the Ore Mountains and the Saxonian lowland, representing two typical natural regions in Saxony, Germany. In accordance with current forest conversion practices, the investigation sites represent different stands: mature conifer stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) (type A); Norway Scots spruce and pine with advanced plantings of European beech (Fagus sylvatica L.) or European beech/Common oak (Quercus petreae Liebl.) (type B); and mature deciduous stands of European beech and European beech/Common oak (type C). The investigated forest sites can be grouped into three silvicultural situations according to the development from coniferous stands to advanced plantings and finally mature deciduous forests (chronosequence A–B–C). The organic layer (L, F and H horizons) and uppermost mineral soil (0–10 cm) were analysed for potential C mineralisation, microbial biomass, concentrations of total C and N (TOC and TN) and for medium-term and readily decomposable C and N fractions, obtained by hot- and cold-water extraction respectively. The results showed an increase in organic layer thickness and mass as well as TOC and TN stocks along the forest sequences in the lowland. Yet, underplanted sites with two storeys revealed higher organic layer mass as well as TOC and TN stocks as compared to coniferous and deciduous stands. Stocks of hot- and cold-water-extractable C and N in relation to microbial biomass and its activity revealed a high turnover activity in deeper organic horizons of deciduous forests compared to coniferous stands. The stand-specific differentiation is discussed in relation to microbial biomass, litter quantity and quality and forest structure, but also with respect to the site-specific climatic factors and water budget as well as liming and fly-ash impacts. Results indicate higher dynamics in deciduous stands in the lowland especially during the initial turnover phase. The elevated microbial activity in deeper organic horizons of deciduous litter-influenced sites in spring is discussed as a specific indicator for long-term C sequestration potential as besides C mineralisation organic compounds are humified and thus, can be stored in the organic layer or in deeper soil horizons. Due to liming activities, stand-specific effects on organic matter turnover dynamics have evened out today in the Ore mountain region, but will presumably occur again once base saturation decreases. Here, the stand-specific effect on microbial biomass can currently be seen again as C_mic in the L horizon increased from spruce to beech. Our study sites in the lowland revealed no significant fly-ash impact. Differences between sites were evaluated by calculating the discriminance function. TOC and TN as well as medium-term degradable C and N were defined in this study as indicators for turnover dynamics along forest conversion sites.     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.     

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.     

Tree effects on the chemical topsoil features of oak, beech and pine forests

We aimed to study tree effects on the chemical properties of forest soils. We compared soil features of three types of forest ecosystems, each with four stands (replicates): beech forests (Fagus sylvatica), oak forests (dominated by Quercus pyrenaica) and pine plantations (Pinus sylvestris). Five samples from the top 10 cm of soil were taken per stand, from which pH, organic matter content (O.M.), total nitrogen (N) and available calcium (Ca²⁺), magnesium (Mg²⁺), potassium (K⁺) and sodium (Na⁺) were determined. Litter layer depth was measured at each soil sampling point. We also measured tree density and crown diameters at each stand. Our results indicated that soil samples from the four pine plantation stands were more similar while oak and beech stands were characterised by great variability in terms of soil properties and leaf litter depth. Although the identity of the dominant tree species significantly influenced several topsoil chemical properties (increase in pH and available cations in oak forests and higher organic matter and total nitrogen in beech and pine ecosystems), there were other important factors affecting soil features that may be taken under consideration. Differences between soil properties of the three types of forest ecosystems were mainly related to the characteristics of the litter layer and less related to the tree layer structure. Finally, the establishment of pine plantations in naturally deciduous tree areas made the topsoil features more homogeneous.     

Commercial thinning that maintained species diversity of a mixed black spruce–jack pine stand enhanced productivity

Commercial thinning enables forest managers to meet timber production objectives. Thinning reduces tree density to alleviate competition for resources and favour growth of selected tree species. However, in doing so, thinning can homogenize the composition of mixed-species forests and raise biodiversity issues. There is increasing evidence that species richness can lead to higher productivity through a complementarity effect. Hence, thinning that would maintain species diversity of mixed-species forests could enhance stand productivity and help forest managers to reconcile timber production objectives and biodiversity issues. The objective of this study was to compare post-thinning stand production, experimentally over 10 years, in mixed and monospecific stands of black spruce (Picea mariana [Mill.] B.S.P.) and jack pine (Pinus banksiana Lamb.). The post-thinning stand production curve of the mixed stand converged toward that of the unthinned mixed stand while the production curves of the thinned and unthinned monospecific stands remained parallel. The convergent productivity of the mixed stand could be explained by a positive interaction between effects of thinning and niche complementarity. We propose that thinning that maintains species diversity of mixed stands could help forest managers who are implementing ecosystem management to reconcile timber production objectives with biodiversity issues.     

Tree species composition affects productivity and carbon dynamics of different site types in boreal forests

The objective was to analyse how differences in the initial proportions of tree species and site fertility affect carbon sequestration in living biomass and soil. We used the individual-based simulation model EFIMOD, which is able to simulate spatially explicit competition between trees for light and nutrients. Simulations were carried out for three site types with distinct initial stocks of soil nutrients. For each site, the 100-years undisturbed dynamics of monocultures and mixtures of three tree species (Betula pendula Roth, Pinus sylvestris L. and Picea abies (L.) H. Karst.) was predicted. Changes in the proportions of competing tree species were dependent on the fertility of the site: on poor sites, pine was the most competent species, while on rich sites, spruce increased its proportion during stand succession. Net primary production (NPP) and soil respiration were the highest in stands of two coniferous species and in stands with a high initial proportion of pine. Mixed stands were more productive than monocultures; the highest overyielding was observed with mixtures of two coniferous species. Simulated NPP and carbon stocks in all pools increased from poor to rich sites. The highest carbon stocks in standing biomass were observed for mixtures of conifer species and three-species mixtures; the greatest accumulation of forest floor occurred in stands with high proportions of pine.     

Conversion of single-layered Scots pine monocultures into close-to-nature mixed hardwood forests: effects on parasitoid wasps as pest antagonists

In the north Saxon lowlands (near Torgau), effects of conversion of pure pine stands (Pinus sylvestris L.) into mixed stands by planting deciduous trees [Fagus sylvatica L. and Quercus petraea (Matt.) Liebl] were investigated on parasitoid wasps as pest antagonists. The effects of planting deciduous trees in pine stands were investigated using the strategy of space-for-time substitution including advanced plantations of beech and oak of different age classes. Wasps were captured at 4-week intervals by ground photoeclectors (GPE, n=6, 1 m²) and flight-interception traps (FIT, n=8), placed in the tree crown layer during the vegetation period (April–October) of 2000. A total of 32,479 parasitoid wasps belonging to 30 families were caught in the traps. Fifty-nine percent of individuals were representatives of families relevant as antagonists of forest insect pests: Ichneumonidae, Braconidae, Pteromalidae, Eulophidae, Mymaridae, Trichogrammatidae, and Scelionidae. Compared to the pure pine stand, individual numbers of most families of parasitoid wasps were statistically significantly higher in mixed stands with deciduous trees. The clearest promoting effects of deciduous trees on parasitoid wasps were manifested in the oldest age classes of advanced beech and oak plantations. Oak exerted stronger effects than beech. This applied especially to the tree crown layer, the stratum with the highest spatial correlation between important phytophagous pine pests and relevant parasitoids. The increased structural diversity of the stands and the broader spectrum of potential hosts are regarded as key factors for the promoting effects of advanced plantings of deciduous trees in pine stands on the community of parasitoid wasps.     

Stand specific occurrence of coarse woody debris in a managed boreal forest landscape in central Sweden

Forests long subjected to management for timber production contain only a fraction of the volumes of coarse woody debris (CWD) found in pristine forests. This is a threat to many organisms that depend on CWD. Forest management practices have been altered to achieve an increase in the amounts of CWD. Few studies have attempted to analyze the occurrence of CWD at the landscape level. We studied the occurrence of CWD in stands of different ages and management background in a boreal forest landscape in central Sweden. Volume of CWD in unmanaged stands (nature reserves and set-asides) was twice that in managed stands. The composition of CWD was influenced by stand age and management regime. Standing CWD was more common in unmanaged stands than in managed stands. Pine CWD was particularly prevalent in young forest stands (8–59 years of age). Bark-covered CWD was most common on deciduous and spruce wood and uncommon on pine. Bark area in young forest stands was almost 10 times lower than that in other managed stands. Using the age distribution of stands, we estimated the volume and bark area of CWD in the landscape. Recent clear-cuts harvested in accordance with new management guidelines contained more early decay CWD per ha than old managed stands. Young forests covered over half the landscape and had significantly lower volumes of spruce and deciduous CWD compared with other stands. The consequences of these results for biodiversity-oriented forest management are discussed.     

How to quantify forest management intensity in Central European forests

Existing approaches for the assessment of forest management intensity lack a widely accepted, purely quantitative measure for ranking a set of forest stands along a gradient of management intensity. We have developed a silvicultural management intensity indicator (SMI) which combines three main characteristics of a given stand: tree species, stand age and aboveground, living and dead wooden biomass. Data on these three factors are used as input to represent the risk of stand loss, which is a function of tree species and stand age, and stand density, which is a function of the silvicultural regime, stand age and tree species. Consequently, the indicator consists of a risk component (SMI_r) and a density component (SMI_d). We used SMI to rank traditional management of the main Central European tree species: Norway spruce (Picea abies [Karst.] L.), European beech (Fagus sylvatica L.), Scots pine (Pinus sylvestris L.), and oak (Quercus robur L. and Quercus petraea L.). By analysing SMI over their whole rotation period, we found the following ranking of management intensity: oak<beech<pine?spruce. Additionally, we quantified the SMI of actual research plots of the German Biodiversity exploratories, which represent unmanaged and managed forest stands including conifer forests cultivated outside their natural range. SMI not only successfully separate managed from unmanaged forests, but also reflected the variability of forest management and stand properties across the entire sample and within the different management groups. We suggest using SMI to quantify silvicultual management intensity of stands differing in species composition, age, silvicultural system (even-aged vs. uneven-aged), thinning grade and stages of stand conversion from one stand type into another. Using SMI may facilitate the assessment of the impact of forest management intensity on biodiversity in temperate forests.     

Habitat use of large ungulates in northeastern Germany in relation to forest management

Estimating large herbivore density has been a major area of research in recent decades. Previous studies monitoring ungulate density, however, focused mostly on determining animal abundance, and did not interpret animal distribution in relation to habitat parameters. We surveyed large ungulates in the Biodiversity Exploratory Schorfheide-Chorin using faecal pellet group counts. This allowed us to explore the link between relative ungulate abundance, habitat use, and browsing damage on trees in a region with several types of forest, including unharvested and age-class beech forests, as well as age-class pine forests. Our results demonstrate that roe deer and fallow deer relative abundance is negatively correlated with large tree cover, and positively correlated with the cover of small shrubs (Rubus spec., Vaccinium spec.), and winter food supply. Habitat use of roe deer and fallow deer, as estimated by counting faecal pellet groups, revealed a preference for mature pine forests, and avoidance of deciduous forests. This differential habitat use is explained by different distributions of high quality food resources during winter. The response of deer to understory cover differed between roe deer and fallow deer at high cover percentages. The amount of browsing damage we observed on coniferous trees was not consistent with the relative deer abundance. Browsing damage was consistently higher on most deciduous trees, except for beech saplings which sustained less damage when roe deer density was low. Because roe deer is a highly selective feeder, it was reported to affect tree diversity by feeding only on trees with high nutritional value. Consequently, we propose that managing the number of all deer species by hunting is necessary to allow successful forest regeneration. Such an adjustment to deer numbers would need to account for both current tree diversity and alternative food resources. Our findings may be applicable to other forest landscapes in northeastern Germany including mature pine stands and differently harvested deciduous forests.     

Effect of natural disturbances on the abundance of Norway spruce (Picea abies (L.) Karst.) regeneration in nemoral forests of the southern boreal zone

The impact of natural disturbances on the canopy (trees ≥14 m high) and sapling stratum (>0.3 and ≤14 m high) composition was studied in nemoral old-growth forests located within the southern boreal zone in Central Russia (Central Forest Reserve, 32°29′–33°01′E, 56°26′–56°31′N). I hypothesized that the current disturbance regime does not allow the maintenance of current spruce abundance in the canopy, and, as a result, there is a continuous shift in the canopy composition towards a greater abundance of deciduous species. Three 300×20 m² transects were established to estimate the proportions of stand under non-closed unexpanded canopy gaps. Data on sapling composition of 49 canopy gaps were used to analyze pattern of gap refuting in these forests. Additionally, data from three forest inventories showed changes in canopy composition over a period from 1972 to 1990.

The current status of nemoral forests is characterized by the high proportion of stand area under treefall gaps (71%). The loss of spruce from the canopy caused by treefalls (53% of the total basal area of gap-makers) was slightly greater than its canopy abundance (45%). Canopy gaps of all sizes encouraged spruce regeneration which might be due to a decrease in sapling mortality and/or more active recruitment of spruce seedlings. After a gap was formed, the presence of spruce in sapling strata increased. However, within both small (<200 m² in size) and large (>200 m²) gaps, tall (>6 m) spruce saplings did not reach the level of its abundance in the tree canopy. In gaps, tall (>6 m) saplings of lime (Tilia cordata) and elm (Ulmus glabra) grew more quickly than those of spruce and maple. These data suggested a decrease in canopy spruce and an increase in deciduous species in the near future which supported the original hypothesis. Analysis of forest inventory records revealed similar changes in the canopy structure over the past two decades. However, the observed high proportion of stand area under gaps implies that for the next few decades large areas of nemoral communities will be occupied by relatively young stands. This may, in turn, decrease the frequency of large-scale treefalls revegetated mainly by deciduous saplings.     

The spreading of the S type of Heterobasidion annosum from Norway spruce stumps to the subsequent tree stand

The occurrence of Heterobasidion annosum in stumps and growing trees was investigated on 15 forest sites in southern Finland where the previous tree stand had been Norway spruce (Picea abies) infected by H. annosum, and the present stand was either Scots pine (Pinus sylvestris), lodgepole pine (Pinus contorta), Siberian larch (Larix siberica), silver birch (Betula pendula) or Norway spruce 8–53 years old. Out of 712 spruce stumps investigated of the previous tree stand, 26.3% were infected by the S group and 0.3% by the P group of H. annosum. The fungus was alive and the fruit bodies were active even in stumps cut 46 years ago. In the subsequent stand, the proportion of trees with root rot increased in spruce stands and decreased in stands of other tree species. On average, one S type genet spreading from an old spruce stump had infected 3.0 trees in the following spruce stand, 0.5 trees in lodgepole pine, 0.3 trees in Siberian larch, 0.05 trees in Scots pine and 0.03 trees in silver birch stand. Although silver birch generally was highly resistant to the S type of H. annosum, infected trees were found on one site that was planted with birch of a very northern provenance.     

油松—白桦混交林种间关系研究

在调查河北隆化县油松－白烨混交林的林木生物量（地上部生物量及根系生物量）、林下草灌木、林地枯落物的总量以及不同混交距离油松地上地下部分生长发育状态的基础上，研究了混交林的种间关系。结果表明，该混交林作为华北地区广泛分布的一种林型，林下草灌木繁茂，林地枯落物分解迅速，有利于土壤理化性质和林分营养状况的改善，种间关系处于基本协调状态。由于林龄及密度原因，混交林生物量略低于油松纯林，在混交林中白桦对油松的生长有一定的抑制作用，抑制作用因混交距离而变化。本文提出了合适的混交距离和混交方式。     

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.     

The nutrient status of Norway spruce in pure and in mixed-species stands

Atmospheric deposition of N and S appears to have caused nutrient imbalance in Norway spruce stands in southern Sweden. This calls for a change of forest management to procedures that promote nutrient balance. Studies have shown lower soil acidity in Norway spruce/deciduous mixed stands than in spruce monocultures, but the tree nutrient status in such mixtures has not been much investigated so far.

The nutrient status of Norway spruce foliage and top mineral soil chemistry in monocultures and in stands mixed with beech, birch, or oak was investigated through paired comparisons on 30 sites in southern Sweden (27 sites) and eastern Denmark (three sites). In total, 45 mixed stands and 34 pure stands were included in the study.

Spruce needles from mixed stands had higher concentrations and ratios to N of K, P, and Zn than needles from pure spruce stands. Among the mixed stands, the K status appeared to be positively correlated with the percentage of deciduous tree basal area. Soil samples from mixed stands had a higher Mg concentration, base saturation, and BC/Al ratio than soil samples from pure stands. The spruce needle nutrient status was comparable in pure stands on fertile sites and in mixed stands on poor sites. We did not detect any differences in spruce tree growth between pure and mixed stands.

This paper discusses possible reasons for a positive effect on the tree nutrient status in mixed-species stands and the possibility of using mixed-species stands as a forest management procedure to avoid nutrient imbalance.