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.     

Influence of slope position, stand type and rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech (Fagus orientalis Lipsky.) and spruce [Picea orientalis (L.) Link]

Oriental beech (Fagus orientalis Lipsky.) and Oriental spruce [Picea orientalis (L.) Link] are the two most common tree species in northeast Turkey. Their distribution, stand type and understorey species are known to be influenced by topographical landforms. However, little information is available as to how these changes affect litter decomposition rates of these two species. Here, we investigated the effects of slope positions (top 1,800 m, middle 1,500 m and bottom 1,200 m), stand type (pure and mixed stands) and purple-flowered rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech and spruce for 4 years using the litterbag technique in the field. Among these three factors, stand type had the strongest influence on litter decomposition (P < 0.001, F = 58.8), followed by rhododendron (P < 0.001, F = 46.8) and slope position (P < 0.05, F = 11.6). Litter decomposition was highest under mixed beech/spruce forest, followed by pure beech and spruce forest. Beech and spruce litter decomposed much faster in mixed bags (beech–spruce) than they did separately under each stand type. Purple-flowered rhododendron significantly reduced litter decomposition of Oriental beech and spruce. Beech and spruce litter decomposed much slower at top slope position than at either bottom or middle position. Differential litter decomposition of Oriental beech and spruce was mainly due to adverse conditions in spruce forest and the presence of rhododendron on the ground which was associated with lower soil pH. Higher elevations (top slope position) slowed down litter decomposition by changing environmental conditions, most probably by decreasing temperature as also other factors are different (pH, precipitation) and no detailed investigations were made to differentiate these factors. The adverse conditions for litter decomposition in spruce forest can be effectively counteracted by admixture of beech to spruce monoculture and by using the clear-cutting method for controlling rhododendron.     

Species mixing effect on Norway spruce response to elevated CO2 and climatic variables: root and radial growth response

A 7-year study was conducted to examine the growth (diameter and root) response of Norway spruce (Picea abies (L.) Karst.) seedlings to elevated CO₂ (CO_2ELV, 770 μmol (CO₂) mol^?1) in different mixture types (monospecific (M): a Norway spruce seedling surrounded by six spruce seedlings, group-admixture (G): a spruce seedling surrounded by three spruce and three European beech seedlings, single-admixture (S): a spruce seedling surrounded by six beech seedlings). After seven years of treatments, no significant effect from elevated CO₂ was found on the root dry mass (p?=?0.90) and radial growth (p?=?0.98) of Norway spruce. Neither did we find a significant interaction between [CO₂]?×?mixing treatments (p?=?0.56), i.e. there was not a significant effect of CO₂ concentrations [CO₂] in all the admixture types. On the contrary, spruce responses to admixture treatments were significant under CO_2AMB (p?=?0.05), which demonstrated that spruce mainly increased its growth (diameter and root) in M and neighbouring with beech was not favourable for spruce seedlings. In particular, spruce growth diminished when growing beside high proportions/numbers of European beech (S). Here, we also evaluated the association between tree-ring formation and climatic variables (precipitation and air temperature) in different admixture types under elevated and ambient CO₂ (CO_2AMB, 385 μmol (CO₂) mol^?1). Overall, our result suggests that spruce responses to climate factors can be affected by tree species mixing and CO₂ concentrations, i.e. the interaction between climatic variables?×?admixture types?×?[CO₂] could alter the response of spruce to climatic variables.

     

Do mature pine plantations resemble deciduous natural forests regarding understory plant diversity and canopy structure in historically modified landscapes?

We compared the structure of the arboreal layer and the diversity and species composition of the understory vegetation of three types of mature forest communities: oak (Quercus pyrenaica) and beech (Fagus sylvatica) forests and Scots pine (Pinus sylvestris) plantations. Our main aim was to determine whether differences in these variables existed and were due to the identity of the dominant tree species. We selected four stands or replicates per forest type located geographically close and with relatively similar conditions. We found no differences in the arboreal structure of oak and beech forests, which were characterised by great variability in tree size, while in case of plantations, this variability was lower at both the intra-stand (estimated by the coefficient of variation) and inter-stand (i.e. the four replicates harboured trees of similar sizes) scales. However, the highest variability in the canopy layer of natural forests was not consistently linked to greater understory species richness. Indeed, the lowest plant species richness was found in beech forests, while oak forests harboured the highest value at either the sampling unit (per m²) or stand scales. The greatest negative correlation between plant diversity and the environmental variables measured was found for litter depth, which was the highest in beech forests. The results obtained by the CCA indicated that the four replicates of each forest type clustered together, due to the presence of characteristic species. We concluded that pine plantations did not approach the environmental conditions of native forests, as plantations were characterised by singular understory species composition and low arboreal layer variability, compared to natural woodlands.     

Stocks and dynamics of soil organic carbon and coarse woody debris in three managed and unmanaged temperate forests

The effect of forest conservation on the organic carbon (C) stock of temperate forest soils is hardly investigated. Coarse woody debris (CWD) represents an important C reservoir in unmanaged forests and potential source of C input to soils. Here, we compared aboveground CWD and soil C stocks at the stand level of three unmanaged and three adjacent managed forests in different geological and climatic regions of Bavaria, Germany. CWD accumulated over 40–100 years and yielded C stocks of 11 Mg C ha^?1 in the unmanaged spruce forest and 23 and 30 Mg C ha^?1 in the two unmanaged beech–oak forests. C stocks of the organic layer were smaller in the beech–oak forests (8 and 19 Mg C ha^?1) and greater in the spruce forest (36 Mg C ha^?1) than the C stock of CWD. Elevated aboveground CWD stocks did not coincide with greater C stocks in the organic layers and the mineral soils of the unmanaged forests. However, radiocarbon signatures of the O _e and O _a horizons differed among unmanaged and managed beech–oak forests. We attributed these differences to partly faster turnover of organic C, stimulated by greater CWD input in the unmanaged forest. Alternatively, the slower turnover of organic C in the managed forests resulted from lower litter quality following thinning or different tree species composition. Radiocarbon signatures of water-extractable dissolved organic carbon (DOC) from the top mineral soils point to CWD as potent DOC source. Our results suggest that 40–100 years of forest protection is too short to generate significant changes in C stocks and radiocarbon signatures of forest soils at the stand level.     

Litter of an alien tree, <Emphasis Type="Italic">Casuarina equisetifolia</Emphasis>, inhibits seed germination and initial growth of a native tree on the Ogasawara Islands (subtropical oceanic islands)

We investigated the effects of the accumulation of litter of an alien tree, Casuarina equisetifolia, on the initial establishment of Schima mertensiana, a tree native to the Ogasawara (Bonin) Islands, in the northwestern Pacific Ocean, in field and greenhouse experiments. The field experiment compared emergence of seedlings in forests dominated by C. equisetifolia with those in native forests, with and without litter removal. The greenhouse experiment compared seedling germination and biomass among 12 treatments that included two soil types (collected from the C. equisetifolia and native forests), two litter types (C. equisetifolia and native litter), and three amounts of litter (5, 15, or 45 g/pot). Significantly fewer S. mertensiana seedlings emerged at sites dominated by C. equisetifolia than at sites of native species in the field experiment. Litter removal tended to increase seedling emergence. Fewer and smaller S. mertensiana seedlings germinated with greater litter cover regardless of soil type or litter type in the greenhouse experiment. Our results demonstrate that C. equisetifolia litter suppresses germination and initial growth of S. mertensiana, and that the main cause of the suppression is the greater amount of litter accumulated under C. equisetifolia forest floor rather than factors such as the nature of the soil or litter between native and alien species.     

Mixed forests and a flexible harvest policy: a problem for conventional risk analysis?

The famous “Faustmann” equation, which allows for identifying the most profitable tree species on a given unstocked piece of land, assumes constant timber prices. In reality, timber prices may fluctuate dramatically. Several authors have proven for monocultures that waiting for an acceptable timber price (reservation price) before harvesting (flexible harvest policy) increases the net present value of forest management. The first part of this paper investigates how efficient a flexible harvest strategy may be applied in mixed forests and whether the optimal species mixture is changed under such harvest policy. Mixtures of the conifer Norway spruce [Picea abies (L.) Karst] and the broadleaf European beech (Fagus sylvatica L.) were investigated. In order to evaluate mixed forests, the risks and the correlation of risks between tree species as well as the attitude towards risk of the decision-maker (risk-aversion is assumed) were considered according to the classical theory of optimal portfolio selection. In the second part we took up a recent critique on modern financial theory by Mandelbrot. Whether or not the assumption of normally distributed financial flows, which are supposed to occur under risk, would be appropriate to evaluate the risk of forest management was investigated. Market and hazard risks as well as their correlation were integrated in the evaluation of mixed forests by means of Monte-Carlo simulations (MCS). The risk of the timber price fluctuation was combined with the natural hazard risk, caused mainly by insects, snow and wind. Applying the μ-σ-rule, the mean net present value (NPV) from 1,000 simulations and their standard deviation were used for the optimisation. Given a low-return, risk-free interest rate to assess potential species mixtures of the Norway spruce and European beech, optimal proportions of European beech increased according to the theory of optimum portfolio selection with growing risk aversion from 0 (ignorance of risk) to 60% (great risk-aversion). In relation to a fixed harvest policy, the net present value of both, Norway spruce and European beech, could be increased significantly. Since the hazard risks of European beech were substantially lower compared with the Norway spruce (relation of susceptibility 1:4) beech benefited more from the flexible harvest policy. A comparison of simulated frequency distributions of the NPV with the expected density functions under the assumption of a normal distribution revealed significant differences. Only in the case of European beech was the general shape of the simulated frequency distribution similar to a normal distribution (bell-shaped curve). However, the density of NPV close to the mean was much greater than expected under the assumption of a normal distribution. Consequently, the frequency of a negative NPV for a European beech forest was greatly overestimated when applying the normal distribution. Though the shape of the simulated frequency distribution was rather different from a normal distribution for Norway spruce the simulated part of negative NPV was quite well approximated by the normal distribution. Therefore the simulated and expected frequencies of negative NPV were similar in case of Norway spruce; only a slight underestimation was seen in the assumption of a normal distribution. It can be concluded that actually simulated frequencies of negative NPV seem to be better measures for risk than computed probabilities of negative NPV, which assume normal distribution. As the risk for European beech was greatly overestimated by the conventional assumption of a normal distribution, the optimal proportions of European beech were surely rather underestimated according to the theory of portfolio. MCS on optimum mixtures derived by the classical portfolio theory seems necessary to test the robustness of such mixtures.

Characterization of Cylindrocarpon‐like species associated with litter in the old‐growth beech forests of Central Europe

Cylindrocarpon‐like fungi are globally distributed plant pathogens and have a wide range of host species. However, very little is known about the species that live in the topsoil of forests and their potential role in reducing the natural regeneration of tree species, particularly of forest trees that produce abundant fruit only once every few years. To enhance our understanding of the species that inhabit the topsoil, we studied the diversity and pathogenicity of Cylindrocarpon‐like fungi inhabiting the litter in old‐growth mixed‐beech forests in the Carpathians (Poland) and in the Alps (Austria), and in a managed beech stand in the Krakowsko‐Cz?stochowska Highland (Poland). The fungi inhabiting the beech litter were investigated using beechnuts and pine seedlings as bait. Isolates were identified based on morphology and DNA sequencing. The pathogenicity of the most common species was investigated by inoculating beech germinants. A wide range of Cylindrocarpon‐like fungi were associated with the beech litter: 718 cultures representing 12 species were isolated. Five taxa were identified down to species level: namely Ilyonectria crassa, I. pseudodestructans, I. rufa, Neonectria candida and N. obtusispora, and seven species were identified to genus level (Neonectria or Ilyonectria species). Ilyonectria destructans, which is considered to be the sexual morph of ‘Cylindrocarpon destructans’, was not found. There were qualitative and quantitative differences between the different forest sites in terms of Neonectria and Ilyonectria species composition in beech litter. The isolation frequency and species richness of Cylindrocarpon‐like fungi were greatest in beech litter taken from old‐growth mixed‐beech forests. Neonectria and Ilyonectria species were capable of killing beech germinants, suggesting that they may play a negative role in natural beech regeneration.     

Spatial and temporal patterns in seed predation as revealed by reciprocal experiments and video surveillance in neighbouring beech and spruce forests

European beech and Norway spruce are late successional and competitive species meeting each other in southern Norway. Beech is predicted to expand northwards with increased temperature, implying increased competition between the two tree species. Seed survival is a key process in this competition and here we estimate detection times and predation rates of beech and spruce seeds in adjacent beech and spruce forests by combining quantitative and qualitative approaches based on reciprocal seed predation experiments and video surveillance. These experiments were repeated over two growing seasons, and by using a video-surveillance technique, we were able to precisely reveal seed detection times and high seed predation rates, up to 15 seeds?min^?1, with bank vole and chaffinch as main predators. Moreover, the two main predators were different in their seed preferences, that is, beech seeds were exclusively removed or predated by the vole, while the finch consumed spruce seeds only. As we show high predation rates, even small variations in seed predator population densities would impact the regeneration success of beech and spruce. Because the highest predation rates were recorded for spruce seeds, it is likely that this will alter the competitive balance between the two tree species in favour of beech.     

Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

Effects of sika deer (Cervus nippon) and dwarf bamboo (Sasamorpha borealis) on seedling emergence and survival in cool-temperate mixed forests in the Kyushu Mountains

Effects of sika deer (Cervus nippon) and dwarf bamboo (“sasa;” Sasamorpha borealis) on seedling emergence and survival were investigated in cool-temperate mixed forests in the Kyushu mountain range, Japan. We compared the effects of sika deer between two sites with different sika deer densities. One site (no-sasa forest) has sparse cover of S. borealis and a high density of sika deer, and the other site (sasa forest) has dense cover of S. borealis with a low density of sika deer. In the no-sasa forest, more seedlings emerged and the survival rate of the seedlings was higher compared with the sasa forest. Compared with the sasa forest, the canopy in the no-sasa forest was more open, the organic layer was shallower and drier, the mean daily soil temperature was higher, and soil temperature fluctuated more widely. Those environmental parameters did not differ between unfenced and fenced plots. The comparison between the sasa and no-sasa forests suggested that removal of S. borealis by sika deer had an indirect positive effect on seedling emergence and survival. However, the fenced-plot experiments indicated that sika deer inhibited seedling emergence and survival. These contradictory conclusions could result from the much larger negative effects of S. borealis on seedling emergence and survival compared with the effects of sika deer browsing.     

A comparison of litterfall and element fluxes in even aged Norway spruce,sitka spruce and beech stands in Denmark

Litterfall was investigated in three even-aged Norway spruce (Picea abies), sitka spruce (Picea sitchensis) and beech (Fagus sylvatica) stands on a nutrient poor-soil in Southern Denmark. Dry weights and N, P, K, S, Mg, Ca, Na, Al, and Fe concentrations and fluxes were examined in litterfall fractions. Foliage litter amounted to 90% of total litterfall. The tree stands showed a similar mean annual litterfall. In the spruce stands, annual litterfall was correlated negatively with the current year increment and positively with the previous year increment. Annual litterfall in beech was constant during the 6 study years whereas Norway spruce and sitka spruce showed large fluctuations between years caused by drought, spruce aphid infestations and probably sea salt stress. Norway spruce responded with a long lasting elevated needle loss. Sitka spruce responded to infestations with premature needle loss during short periods. The presence of a large syrphid (Coccinellidae) population was important in regulating aphid (Elatobium abietinum) population density. The between-year variation in element concentrations of litterfall was small whereas variations during the year were large. Interspecific levels were recognized: Norway spruce>beech>sitka spruce. High concentrations in Norway spruce were ascribed to a combination of drought, sea salt stress and elevated transpiration. In sitka spruce, aphid infestations reduced the litterfall N content. Sitka spruce showed the smallest amount of base cation fluxes with litterfall. In contrast, spruce and beech exhibited even litterfall element fluxes. Litterfall studies revealed reduced vitality in the non-native spruce stands and underlined the perception of a healthy stand of native beech.     

Understory dynamics after disturbance accelerate succession from spruce to beech-dominated forest—the Siggaboda case study

??Context

It is assumed that climate change will favour European beech (Fagus sylvatica L.) to Norway spruce (Picea abies [L.] Karst.) at its northern range margins due to climate change and induced disturbance events.

??Aims

An old-growth mixed forest of spruce and beech, situated near the northern beech margin, was studied to reveal effects of disturbances and response processes on natural forest dynamics, focussing on the understory.

??Methods

We carried out analyses on understory dynamics of beech and spruce in relation to overstory release. This was done based on a sequence of stand and tree vitality inventories after a series of abiotic and biotic disturbances.

??Results

It became apparent that beech (understory) has a larger adaptive capacity to disturbance impacts and overstory release (68 % standing volume loss) than spruce. Understory dynamics can play a key role for forest succession from spruce to beech-dominated forests. Disturbances display an acceleration effect on forest succession in the face of climate change.

??Conclusion

Beech is poised strategically to replace spruce as the dominant tree species at the study area. Due to an increasing productivity and a lower risk of stand failure, beech may raise into the focus of forestry in southern Sweden.     

Seed and microsite limitation for seedling recruitment of <Emphasis Type="Italic">Quercus wutaishanica</Emphasis> on Mt. Ziwuling,Loess Plateau,China

The relative influence of seed and microsite availability in the seedling emergence and recruitment of Liaodong oak (Quercus wutaishanica), which dominates the native hardwood forest in Loess Plateau, was examined by seed sowing experiments. Experiments were performed in 216 plots (30 × 30 cm), located in three typical stands (Pinus tabulaeformis plantation, Liaodong oak forest and grassland, 72 plots per stand). A seed augmentation experiment was performed in an oak forest and a conifer plantation, and a seed introduction experiment was carried out in a grassland. Three main factors were considered: seed addition, disturbance and shrub cover. The seedling number and growth in every plot were recorded continuously over 3 years. GLM multivariate was used to analyze the relationships between seedling emergence/recruitment and the explanatory variables in every stand. The results showed that seed addition in undisturbed or disturbed plots always caused significantly increased seedling emergence and recruitment in oak and conifer forests. It demonstrated that the recruitment of Liaodong oak was limited by seed and microsite availability under closed forest, and litter is one of the factors leading to microsite limitation. However, in the grassland, no treatments improved recruitment, indicating that the grassland was not suitable for seedling recruitment because of intense light and soil drought. In the conifer forest, more recruited seedlings, and lower herbivory on seedling leaves, as well as thicker stem basal diameters, indicated that the P. tabulaeformis plantation was safer for Liaodong oak seedling establishment. Unexpectedly, shrub cover did not affect the recruitment of Liaodong oak in the three stands.     

Stand density and growth of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> (L.) Karst.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.): evidence from long-term experimental plots

On the basis of nine Norway spruce (Picea abies (L.) Karst.) and ten European beech (Fagus sylvatica L.) thinning experiments in Germany, for which both residual and removed stock had been registered first during 1870, I scrutinize how moderate and heavy thinning from below (B-, C-grade) affects the production of merchantable volume compared with light thinning (A-grade). In relation to A-grade, cumulative merchantable volume (CV) of B- and C-grade amounts in average to 103–107% in juvenile and to 97–102% in mature Norway spruce stands. The corresponding findings for European beech are 101–106% and 94–102%. CV of individual stands varies between 89% and 130% for Norway spruce and 73% and 155% for European beech (CV of A-grade = 100%). These findings are substantiated by the relation between stand density (SDI) and periodic annual increment (PAI). On the B- and C-grade plots of spruce and beech, respectively, SDI was reduced down to 41–91% and 31–83% of the A-grade. When SDI is reduced in young stands, PAI follows a unimodal curve. Norway spruce’s PAI culminates in 109% if SDI is reduced to 59%; European beech’s PAI culminates in 123% when density is reduced to 50%. Whereas Norway spruce’s growth reacts most positively on thinning under poor site conditions and with increment reduction on favourable sites, European beech behaves oppositely. With stand development the culmination point of the unimodal relation moves towards maximum density, so that in older stands PAI follows the increasing pattern, which is the left portion of a unimodal curve. A model is presented which apparently unifies contradictory patterns of stand density–growth reactions by integrating relative stand density, average tree size and site fertility effects, and makes the findings operable for forest management.     

Modelling and economic evaluation of forest biome shifts under climate change in Southwest Germany

We evaluated the economic effects of a predicted shift from Norway spruce (Picea abies (Karst) to European beech (Fagus sylvatica (L) for a forest area of 1.3 million ha in southwest Germany. The shift was modelled with a generalised linear model (GLM) by using presence/absence data from the National Forest Inventory in Baden-Württemberg, a digital elevation model, and regionalised climate parameters from the period 1970 to 2000. Two scenarios from the International Panel on Climate Change (IPCC) (B1, A2) for three different time scales (2030, 2065, and 2100) were investigated. The GLM predicted a decrease of the suitable area for growing Norway spruce between 21% (B1, 2030) and 93% (A2, 2100) in comparison to 2000. This corresponds to a reduction in the potential area of Norway spruce from between 190,000 and 860,000 ha. The financial effect of this reduction in area was then evaluated by using a classical Faustmann approach, namely the land expectation value (LEV) as an economic parameter for forests of Norway spruce versus European beech. Underlying cash flows were derived from a distance dependent, single-tree growth simulator (SILVA) based on data for prices and costs of the year 2004. With an interest rate of r = 2%, the predicted loss in the potential area of Norway spruce is related to a decrease of the LEV between 690 million and 3.1 billion Euro. We discuss the sensitivity of these results to changing interest rates, risk levels, and rotation lengths. Results suggest that managing forestland for profitability will be increasingly difficult under both climate scenarios.     

温带森林系统中碳矿化和氮氧化释放的温度依赖性研究

测定森林土壤中的CO2 和N2O 的释放,在评价森林平衡大气中CO2 和N2O的作用方面有着重要的意义.为了量化立地对净碳矿化和N2O氮释放温度依赖性的影响,在德国的索尔森林,调查了三毛举林,挪威云杉和混合林三种相邻的地被物,并进行了3个月不同温度的培养试验.结果表明,3种森林地被物的净碳矿化率和N2O氮的释放量随温度上升呈现指数性增长.在一定温度范围内(1(20oC),利用温度系数函数(Q10)拟合通量率来描述森林地被物的温度敏感性.各试验点的森林地被物的温度敏感性拟合曲线与净碳矿化和N2O氮释放率都显正相关.各试验林的全部数据表明,每个单位的净碳矿化和N2O氮释放的温度系数函数值(Q10)分别为1.73(2.10和2.81(3.58,可用以描述试验地净碳矿化和N2O氮释放率对温度的依赖性.在三毛举和云杉的单一树种和混合种中,净碳矿化率 和N2O 氮释放率的温度依赖性没有明显的差异,表明净净碳矿化率和N2O 氮释放量不受不同树种凋落物质量的影响.     

Litterfall and nutrient return in five tree species in a common garden experiment

Canopy litterfall is a significant pathway for return of nutrients and carbon (C) to the soil in forest ecosystems. Litterfall was studied in five even-aged stands of Norway spruce, Sitka spruce, Douglas-fir, European beech and common oak at three different locations in Denmark; two sandy sites, Ulborg and Lindet in Jutland, and one loamy site, Frederiksborg on Zealand. Litterfall was collected during three years from 1994 to 1996 in all five species and during six years from 1994 to 1999 in Norway spruce, Sitka spruce and European beech. The average total litterfall was in the range of 3200–3700 kg ha⁻¹ yr⁻¹ and did not differ significantly among tree species. There were no significant differences in total litterfall among sites during the short period, but during the longer period the richer site Frederiksborg had significantly higher total and foliar litterfall amounts compared to the more nutrient-poor sites Lindet and Ulborg. There were close relationships between foliar and total litterfall suggesting that foliar litterfall can be reliably estimated from total litterfall. Beech and oak bud scale litter was significantly related to foliar litterfall. The amount of branch and twig litter was significantly higher in oak than in other tree species. The average foliar litterfall was well related to the annual volume increment. The relationship differed markedly from previously reported relationships based on global litterfall data suggesting that such relationships are better evaluated at the regional level. Nutrient concentrations and fluxes in foliar litterfall were not significantly different among the five tree species. However, there was a significant effect of site on most nutrient concentrations of the three litterfall fractions, and foliar fluxes of P, Ca and Mn were all significantly highest at Frederiksborg and lowest at Ulborg. The similarity in litterfall inputs to the forest floor under these five tree species suggested that previous reports of large variability in forest floor accumulation should primarily be attributed to differences in litter decomposition.     

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.