Species proportions by area in mixtures of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) dominate many of the European forest stands. Also, mixtures of European beech and Scots pine more or less occur over all European countries, but have been scarcely investigated. The area occupied by each species is of high relevance, especially for growth evaluation and comparison of different species in mixed and monospecific stands. Thus, we studied different methods to describe species proportions and their definition as proportion by area. 25 triplets consisting of mixed and monospecific stands were established across Europe ranging from Lithuania to Spain in northern to southern direction and from Bulgaria to Belgium in eastern to western direction. On stand level, the conclusive method for estimating the species proportion as a fraction of the stand area relates the observed density (tree number or basal area) to its potential. This stand-level estimation makes use of the potential from comparable neighboring monospecific stands or from maximum density lines derived from other data, e.g. forest inventories or permanent observations plots. At tree level, the fraction of the stand area occupied by a species can be derived from the proportions of their crown projection area or of their leaf area. The estimates of the potentials obtained from neighboring monospecific stands, especially in older stands, were poorer than those from the maximum density line depending on the Martonne aridity index. Therefore, the stand-level method in combination with the Martonne aridity index for potential densities can be highly recommended. The species’ proportions estimated with this method are best approximated by the proportions of the species’ leaf areas. In forest practice, the most commonly applied method is an ocular estimation of the proportions by crown projection area. Even though the proportions of pine were calculated here by measuring crown projection areas in the field, we found this method to underestimate the proportion by 25% compared to the stand-level approach.     

Spatial pattern of beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) and oak (<Emphasis Type="Italic">Quercus pubescens</Emphasis> Mill.) seedlings in natural pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) woodlands

Spatial pattern of recruitment is an important factor influencing population dynamics of plant communities. The spatial pattern is determined by seed dispersal and by the spatial variability of germination and initial survival. In the process of forest expansion following farmland abandonment, mid- and late-successional species are often dispersed in pioneer forests by birds. This could result in an aggregated spatial pattern of seeds that could influence the dynamics of these species in mixed pioneer forests. In the sub-Mediterranean area, mid- and late-successional species such as Quercus pubescens (downy oak) and Fagus sylvatica (European beech) are expected to replace pioneer Pinus species. Using a point sampling method we demonstrated that beech and oak seedlings (height <2 m) have a clumped distribution in the understorey of pine. This could result from an aggregated dispersal by jays (dispersal effect) or from preferential recruitment in particular habitats (habitat effect). To test these hypotheses we proposed a statistical analysis of spatial patterns of regeneration of beech and oak. Ground cover variables (i.e. cover by rock outcrops, herbs, box shrubs, mosses or pine) did not differ significantly around seedlings as compared with random sample plots. Likewise, clumped seedlings had growth similar to isolated seedlings, thus refuting the hypothesis of preferential location in the most favourable microsites. Aggregated dispersal seems to be involved in the process of regeneration. Since beech and oak seedlings have contrasting ecological demands, we discuss the implication of this pattern for the replacement dynamics of pine by these species.     

Reaction wood drying kinetics: tension wood in <Emphasis Type="Italic">Fagus sylvatica</Emphasis> and compression wood in <Emphasis Type="Italic">Picea abies</Emphasis>

The drying kinetics of reaction woods in Picea abies (compression wood) and Fagus sylvatica (tension wood) in comparison with their corresponding normal woods was investigated under constant convective drying conditions. Moisture profiles along the thickness of small flat-sawn boards taken from reaction and opposite wood zones were evaluated using a polychromatic X-ray system, a non-destructive method. The results revealed substantial differences in the drying behavior between the reaction and opposite woods. Both reaction woods represented slower drying rate than their matching normal woods mainly during the period of free water loss. However, the reaction and opposite woods reached the final moisture content (MC) of about 12% at the same time due to higher initial MC in the opposite woods. In the case of reaction wood, it took a longer time for the moisture profile to become approximately uniform. Overall, a more striking difference was observed in the drying behavior of compression and opposite wood in P. abies. Some important anatomical differences like the cell and pit dimensions and their proportion give some explanations for these drying behaviors.     

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.     

Nucleotide polymorphisms associated with climate,phenology and physiological traits in European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

Although European beech (Fagus sylvatica L.) is one of the most widespread and ecologically and commercially most important deciduous trees in Europe, little is known about its adaptive genetic variation. We explored single-nucleotide polymorphism (SNP) variation in candidate genes for budburst and drought resistance in beech populations sampled across most of the distribution range, represented in an international provenance trial. SNP variation was monitored for six candidate genes, in 114 individuals from 19 natural populations. Population structure was deduced from the analysis of 20 nuclear microsatellite markers. Different methods to detect imprints of natural selection were used (F _ST-outlier, SNP-climate regression, association tests). The F _ST-outlier approach identified the COV gene with unambiguous signal of selection, which is an orthologue of Arabidopsis gene for a membrane protein previously reported as phenology-related. Based on environmental association analysis at the population level, the dehydrin gene was found associated with drought-related climatic variables. At the individual level, dehydrin gene also showed a significant association with chlorophyll fluorescence parameters, which are considered stress markers. The importance of the knowledge of physiological variation and geographical patterns of adaptive genetic variation for guiding reproductive materials transfer under climate change is stressed.     

Trade-off between height growth and spring flushing in common beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

• Introduction   

The annual development cycle of boreal and temperate trees results from an evolutionary trade-off between two opposing forces. These are namely, the adjustment of leaf phenology to the timing of frost occurrence at the beginning and/or the end of the growth season countered by an effective adjustment to the duration of the growth season to maximise photosynthesis and biomass production during the growing season.     

NIR-spectroscopic investigation of foliage of ozone-stressed <Emphasis Type="Italic">Fagus sylvatica</Emphasis> trees

Near infrared (NIR) reflectance spectroscopy was tested as a fast method for characterizing the toxic effects of air pollution on trees. Fagus sylvatica was exposed to known, different levels of ozone during summer of 2003 in plant chambers. Leaves were taken from the plants and NIR spectra were recorded. In order to derive calibration models, reference analyses were carried out and soluble carbohydrates were determined by an enzymatic method. Minor components, like α-tocopherol, were measured after solvent extraction using gas chromatography-mass spectrometry. A partial least squares (PLS) algorithm was used to perform the chemometric analyses. Good or in some cases very good calibration statistics expressed in terms of root mean square error of cross validation and R² were obtained for glucose, fructose, sucrose, and α-tocopherol. In addition, a better than expected correlation between the NIR spectra of the leaves and the ozone concentration in the plant chambers was observed. NIR spectroscopy appears to be a very useful method to simultaneously determine many components in leaves that are related to the health of trees. It is obviously able to quantitatively describe the changing patterns of constituents in leaves of trees caused by toxic substances. Thus, NIR spectroscopy can be a very effective tool for environmental biomonitoring, especially for observation of forests.     

Towards a DNA marker assisted seed source identification: a pilot study in European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

The widespread European forest tree Fagus sylvatica L. is of great importance for forest management. However, information about seed dispersal is still very rare, though important for harvesting strategies and later on seed source identification. We refined a DNA fingerprinting method for beech nut shells in order to directly assign dispersed seeds to their mother trees. A pilot study was conducted in two beech stands in Germany where leaves of the adult trees and the exocarp of dispersed seeds were fingerprinted at six nSSR loci. While one stand was randomly analysed for adults and dispersed seeds the other was systematically investigated following common harvesting procedures. Imitating the typical net harvesting strategy, seeds were collected beneath 19 adult trees. Exocarp genotyping revealed that on average three different mother trees contributed to a sample of five or six seeds collected beneath a single adult tree. Of the identified mother trees most were located within a radius of 15 m from the sampling point. The repeated pattern of seed dispersal within a short distance constitutes the basis for a straightforward strategy for the assignment of seed lots to a seed source stand. This strategy is based on the matching of individual genotypes without the necessity for a full inventory of the putative source stand. Additionally, we provide allelic ladders of five nSSR loci for standardization among laboratories.     

Spatial patterns and genetic structures within beech populations (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) of forked and non-forked individuals

Analyses of distribution patterns and genetic structures of forest stands can address distinct family structures and provide insights into the association of genetic and phenotypic variation patterns. In this study, point pattern analysis and spatial autocorrelation were used to examine the spatial and genetic structures in two naturally generated beech stands, which differ in age, trunk morphology, and stand management. Significant tree clumping was observed at distances up to 20 m in the young forest stand, whereas dispersion at distances under 10 m was observed in the old stand. The spatial analysis based on Ripley’s k function of the two different groups of trees showed that the non-forked trees match in both stands the spatial pattern of all trees while the forked were randomly distributed. Additionally, according to the bivariate analysis, forked trees in both stands were randomly distributed as related to non-forked tree positions. Finally, Moran’s I values were not very high, though significant genetic autocorrelation was identified at distances up to 20 m in the young stand, suggesting the existence of distinct family structures. However, no significant genetic structuring was observed in the old stand. Our findings suggest that spatial genetic patterns are impacted by stand age, environmental factors and human activities. The spatial distribution of forked trees was not clearly associated to family structures. Random effects and also micro-environmental variation could be additional factors explaining forking of beech individuals.     

Ectomycorrhizal communities within beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) forests that naturally regenerate from clear-cutting in northern Spain

The objectives of the work described here were to evaluate the diversity of ectomycorrhizal (ECM) fungi within Spanish beech (Fagus sylvatica L.) forests subjected to clear-cutting and natural regeneration and to elucidate the extent to which the carbohydrate concentration in roots of trees of different ages and sizes is related to ECM colonization. The study concerned an unmanaged forest, a stand clear-cut in 1996 and another clear-cut in 2001. ECM colonization of beech roots showed seasonal dynamics in the disturbed areas, but the percentage of roots colonized by ECM was not always related to the accumulation of non-structural carbohydrates. The composition of ECM communities differed between different stands and 40% of ECM morphotypes only occurred in disturbed ecosystems. However, comparable numbers of different ECM morphotypes (24, 25) were found in the three beech stands. This finding indicates that ECM diversity was quite high and similar within disturbed and unmanaged areas. This finding suggests that (1) ECM diversity was not affected by the size and age of trees and (2) the potential of ECM inocula remained high within clear-cut areas. Consequently, the introduction of ECM inocula by silvicultural practices would not be needed to improve the regeneration of clear-cut areas described in our study.     

Fine root dynamics in 60-year-old stands of <Emphasis Type="Italic">Fagus sylvatica</Emphasis> and <Emphasis Type="Italic">Picea abies</Emphasis> growing on haplic luvisol soil

Fine root dynamics in mono-specific stands of mature Fagus sylvatica L. and Picea abies Karst. was studied from December 2003 to December 2004 in a stand in Southern Germany. Minirhizotrons were used to draw between species comparisons concerning fine root (≤1 mm) longevity and temporal patterns of fine root dynamics (growth and mortality) as related to seasonal changes in soil water content and soil temperature. In F. sylvatica, median fine root longevity from early seasonal to late-seasonal cohorts was low (77 days). Fine root dynamics scaled positively with seasonal changes in soil water and temperature indicating accelerated fine root turnover during favourable soil conditions. In contrast, fine root longevity in P. abies (273 days) was significantly higher when compared to F. sylvatica and increased from early seasonal to late-seasonal cohorts. Fine root dynamics in P. abies did not correlate with soil environmental conditions. Rather a large proportion of new fine roots occurred during the dry season in superficial soil layers. The data suggest species inherent patterns of fine root longevity and temporal patterns of fine root dynamics.     

Decay dynamic of coarse and fine woody debris of a beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) forest in Central Germany

Decay rates of woody debris were estimated and used to model the decay of various diameters of branches and stems in a beech stand in Central Germany. In addition, use of wood density, volume and mass loss to quantitatively describe the degree of decay was tested. The mass loss during decay could be described by a simple exponential function. Under the presented climatic conditions, beech coarse woody debris (CWD) with a diameter >10 cm decays completely in about 35 years. In the first 8 years of decay the mass loss is determined by the decrease in wood density, and subsequently by the loss in volume. Estimation of wood density allows the first three of the four classes of decay to be distinguished, while trees in the last two decay classes could be distinguished using wood volume. Beech fine woody debris with a diameter between 1 and 10 cm decays within about 18 years. The litter fraction of <1 cm is part of the humus layer after 4 years. If there are goals for the amounts, types and dimensions of woody debris to be provided for conservation of biological biodiversity and other ecological functions in managed beech forests, this study offer indications for how long existing woody debris can meet its functions and how frequent new input of CWD is required.     

Influence of umbrella pine (<Emphasis Type="Italic">Pinus</Emphasis><Emphasis Type="Italic">pinea</Emphasis> L.) stand type and tree characteristics on cone production

Most umbrella pine (Pinus pinea L.) stands are managed as agroforestry systems, whose main production is fruit, due to the edible and highly nutritious kernels, and are frequently associated to natural or seeded pastures and grazing. The stands have low density, in order to enhance crown growth and fruit production. Nevertheless, cone production, both with regard to number and weight, varies greatly between stands, trees and years. In this study were selected three agroforestry systems, representative of umbrella pine stands whose main production is fruit, and one stand representative of the timber production system, where fruit is the secondary production. It was evaluated the variability in cone production as a function of the tree’s diameter at breast height and crown diameter and the individual tree’s competition status. The results indicate that stands managed in agroforestry systems with lower competition and individuals with larger diameter at breast height and crown diameter tend to produce more and heavier cones per tree. The first two principal components of the principal component analysis explain 84 % of the variance in cone production, trees’ dimensions and competition index. Tree competition status has a negative impact on production per tree.     

Leaf area of beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) from different stands in eastern Austria studied by randomized branch sampling

Few tree size/leaf area correlations have been produced for hardwoods, where the extrapolation from individual branches to the whole tree is less straightforward than in conifers with more regular branching patterns. We used randomized branch sampling to estimate leaf area of European beech (Fagus sylvatica L.) trees of different stands, ages and areas in Austria. Cross-sectional areas (CSA) predicted 87–92% of leaf area variation, the best predictor being the sum of branch CSA. Leaf area was somewhat better correlated with CSA at breast height than at the base of the crown, and using sapwood instead of total CSA made little difference. While there was no effect of growth area, a stepwise regression model showed that dominant trees in pole-stage had, for unclear reasons, significantly higher leaf area/CSA relationships. A comparison with regressions produced from smaller beech trees in other parts of Europe suggests that the leaf area/basal area regression is generally valid for beech in central Europe.     

Biological and other alternative control methods against the woolly beech aphid <Emphasis Type="Italic">Phyllaphis fagi</Emphasis> L. on beech <Emphasis Type="Italic">Fagus sylvatica</Emphasis> seedlings in forest nurseries

Biological and other alternative control methods were tested against the woolly beech aphid (Phyllaphis fagi). Field applications of mineral oil to the egg stage reduced initial aphid population by 75%, but only when the eggs were exposed to oil close to the time of hatching. Earlier oil treatments had no effect. Bioassays with the insect pathogenic fungus Lecanicillium lecanii (Verticillium lecanii) in the commercial formulation Vertalec^® were conducted using different dosages, i.e. 1 × 10⁶ ml^?1 (recommended dosage) and 2 × 10⁷ ml^?1. Both nymphs and adults were susceptible to fungal infection at both dosages. The existence of a dense wax-covering in adult P. fagi had no protective effect against fungal infection. In bioassays where leaves were treated with the recommended dosage of Vertalec, there was no difference in mortality measured after 14 days between adult P. fagi with an intact wax-layer and adult P. fagi where the wax-layer had been removed. In semi-field trials with two L. lecanii treatments at the recommended dosage, the aphid population was reduced. There was no enhanced effect with the addition of an additive to the fungal suspension or from covering the plants with a polypropylene cover. The results reveal potential alternative control methods against P. fagi populations. However, adequate control with L. lecanii probably requires several treatments as opposed to the two that were tested in the present experiment. Furthermore, efficiency may depend on summer temperatures and humidity.     

Pathogenic effects of <Emphasis Type="Italic">Xenorhabdus nematophilus</Emphasis> and <Emphasis Type="Italic">Photorhabdus luminescens</Emphasis> (Enterobacteriaceae) against pupae of the Diamondback Moth, <Emphasis Type="Italic">Plutella xylostella</Emphasis> (L.)

The relationship between Xenorhabdus nematophilus and Photorhabdus luminescens, the bacterial symbionts of the entomopathogenic nematodes Steinernema carpocapsae and Heterorhabditis bacteriophora, against the diamondback moth, P. xylostella (L.), pupae showed the pathogenic capability of P. luminescens to be over that of X. nematophilus. They gave 60 and 40?% mortality with LC₅₀ values of 5?×?10⁴ and 5.5?×?10⁵ cells/ml, respectively.The number of bacterial cells influences the rate of killing the pupae of P. xylostella and a direct correlation between mortality of the pupae, percentage of deformed adults and the spraying dose with significant differences were observed. An inverse correlation found between the no. of eggs laid/female, percentage of hatching, adult survival and the spraying dose. These bacterial effects have been attributed to different toxic bacterial enzymes which damage the haemocytes and inhibit activation of the humoral immune system of the insect.     

Modelling the risks of natural stand closure release with ageing in pure beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis>) and spruce (<Emphasis Type="Italic">Picea abies</Emphasis>) stands

• Background  

In the development of pure beech and pure spruce pristine forests, we assume there should be a continuous transition from full stocking to successive disintegration and, finally, total replacement of the first generational wave at the end of its lifetime, dependent on mortality due to overcrowding as well as ageing, leading newly recruited cohorts to grow up from below.     

Bioremediation of CCA-treated wood by brown-rot fungi <Emphasis Type="Italic">Fomitopsis palustris</Emphasis>, <Emphasis Type="Italic">Coniophora puteana</Emphasis>, and <Emphasis Type="Italic">Laetiporus sulphureus</Emphasis>

This study evaluated oxalic acid accumulation and bioremediation of chromated copper arsenate (CCA)-treated wood by three brown-rot fungi Fomitopsis palustris, Coniophora puteana, and Laetiporus sulphureus. The fungi were first cultivated in a fermentation broth to accumulate oxalic acid. Bioremediation of CCA-treated wood was then carried out by leaching of heavy metals with oxalic acid over a 10-day fermentation period. Higher amounts of oxalic acid were produced by F. palustris and L. sulphureus compared with C. puteana. After 10-day fermentation, oxalic acid accumulation reached 4.2 g/l and 3.2 g/l for these fungi, respectively. Fomitopsis palustris and L. sulphureus exposed to CCA-treated sawdust for 10 days showed a decrease in arsenic of 100% and 85%, respectively; however, C. puteana remediation removed only 18% arsenic from CCA-treated sawdust. Likewise, chromium removal in F. palustris and L. sulphureus remediation processes was higher than those for C. puteana. This was attributed to low oxalic acid accumulation. These results suggest that F. palustris and L. sulphureus remediation processes can remove inorganic metal compounds via oxalic acid production by increasing the acidity of the substrate and increasing the solubility of the metals.An erratum to this article can be found at