Plant diversity plays key ecological roles in forest ecosystems, including influencing succession, resilience and nutrient cycling. This study was conducted to investigate the effect of livestock grazing and human use...Plant diversity plays key ecological roles in forest ecosystems, including influencing succession, resilience and nutrient cycling. This study was conducted to investigate the effect of livestock grazing and human uses on herbaceous species diversity. We surveyed 50 ha of protected area and 50 ha of unprotected area to evaluate herbaceous species diversity in oriental beech （Fagus orientalis Lipsky） forests in northern Iran. We calculated and compared three indices each of diversity and evenness, and species richness between the two areas. Herbaceous cover was higher in the unprotected area while leaf litter depth and tree canopy cover were higher in the protected area. The diversity indices, H （Shan- non-Wiener index ）, N~ （McArthur index）, N2 （Hill＇s index）, EQ （Modified Nee index）, E,ar （Smith-Wilson index）, E5 （modified index of Hill） and R=S （species richness） and species richness R=S were greater in the protected area than in the unprotected area, suggesting that protection from grazing results in increased numbers of plants and species. The effect of land protection on plant diversity was more pronounced for evenness than for species richness and the positive correlation between diversity and evenness indices was higher than that between diversity and richness.展开更多
We studied relationships between stand structure and stand stability according to thinning intensity in an afforested oriental beech stand. Various thinning intensities were applied in sample stands. We sampled eight ...We studied relationships between stand structure and stand stability according to thinning intensity in an afforested oriental beech stand. Various thinning intensities were applied in sample stands. We sampled eight plots in stands that were lightly thinned, eight plots in heavily thinned stands and eight plots in unthinned stands as a control. Height and diameter distributions of the stands were measured to assess stand structure. We quantified individual tree stability and collective stability. Heavy thinning during the first thinning operation damaged the storied structure of the stand in thicket stage and affected collective structuring ability. While most control plots had multi-storied stands, after light and heavy thinning two-storied structure became more common.Large gaps occurred in the canopy after heavy thinning. On average, nine tree collectives were formed per sampling plot in the untreated stand, seven collectives after thinning in 2008 and four collectives after thinning in 2009. Stable trees accounted for 17 % of trees in control plots, 24 % in lightly thinned plots, and 15 % in heavily thinned plots. Collective stability values were 83 % in control plots, 82 % in lightly thinned plots and 36 % in heavily thinned plots. We conclude that it is necessary to retain collective structuring capacity during thinning operations for sustaining stand stability.展开更多
For developing nature-based silvicultural practices in a beech forest to promote, abundance, height, vitality, and preferred growth form, regenerated trees growing in gaps were compared with those under closed canopie...For developing nature-based silvicultural practices in a beech forest to promote, abundance, height, vitality, and preferred growth form, regenerated trees growing in gaps were compared with those under closed canopies. A systematic 50 ×50 m grid was plotted in a beech stand in the Kheyrud Experimental Forest for selecting trees to measure variables in gaps and under closed canopies. Abundance and mean height of regenerated beech trees were significantly higher in closed canopies than in canopy gaps. Beech seedlings with excurrent growth were significantly taller within regeneration patches under closed canopy. Moreover, regenerated trees with high vitality were more abundant in closed canopy areas than in gaps. Thus, beech regeneration should improve under closed canopies; hence, gaps in a near natural forest should be created only after adequate regeneration and appropriate growth under the parent tree in a closed canopy area is ensured. The results of this research have profound implications for the sustainable management of the forest and for ensuring sustainable beech regeneration. The presence of a closed canopy cover likely will reduce potential stresses on oriental beech regeneration.展开更多
Oriental Beech is the most important commercial tree species in northern Iran. In recent years wood production companies interested in felling large beech trees for profit have challenged advocates of close-to-nature ...Oriental Beech is the most important commercial tree species in northern Iran. In recent years wood production companies interested in felling large beech trees for profit have challenged advocates of close-to-nature silviculture who favor conservation. Our study objective was to assess the economic value of over-mature beech trees by relating tree diameter （DBH） to amount of decay. Based on the location of onset of decay, we categorized three types of decay as stump, stem, and crown decay. Trees of greater diameter （age） typically showed greater decay in the stem. Percent of decayed volume, diameter of decayed tissue, and length of decay in tree stems varied between 0.5%-64.3%, 15 cm-75 cm, and 2.0-19.5 m, respectively. With increasing trunk diameter, the propor- tion of truck decay increased. Red heart and dark red heart constituted 25% and 14.3% of sampled trees, respectively. However, we found no correlation between intensity of stem decay and morphological charac- teristics of trees. Seedlings were not abundant around the bases of over-mature trees, suggesting that the trees did not contribute to regeneration of the stand. Beech trees of diameter 〉1 m do not provide valu- able round wood for industries and cause to raise wood production costs. We recommend that these trees 〉1 m DBH should be retained in forest stands because of their low commercial value but high ecological and conservational values such as maintaining biodiversity in forest ecosystems.展开更多
We studied the long-term impacts of natural burning on herbaceous species diversity 37 years after a fire occurred in the Roudbar forests of Guilan Province, northern Iran. Numerous studies have examined short-term ch...We studied the long-term impacts of natural burning on herbaceous species diversity 37 years after a fire occurred in the Roudbar forests of Guilan Province, northern Iran. Numerous studies have examined short-term changes in understory vegetation following wildfire; however, very few long-term studies are available or changes inferred from retrospective studies based on chro- nosequences. For this study, 170 ha of forest （85 ha, burned areas; 85 ha, unburned areas） were surveyed. Because the 1000 m2 plots were too large for detailed measurements of herbaceous species, we determined a sub-sample size according to the Whittaker＇s nested plot sampling protocol and minimal areas method. Hence, sub-plots of 32 m2 were used for herbaceous species measurements, which consisted of percent cover of each species based on the Domin criterion. We measured plant diversity （Shannon-Wiener index）, spe- cies richness （Margalef＇s index）, and evenness （Smith-Wilson index）. Mean percent cover, together with diversity, richness and even- ness, increased markedly in burned areas compared to unburned controls. This suggests that the biodiversity of these forests could be restored within 37 years after fire. However, the abundance of invasive species such as Rubus fruticosus and Bromus benekenii increased significantly in burned areas, but these could be controlled by relevant silvicultural operations.展开更多
Background:Higher exportation of harvest residues from forests due to increased demand for woody biomass,has reportedly diminished soil mineral resources and may lead to degraded tree nutrition as well as growth.Howev...Background:Higher exportation of harvest residues from forests due to increased demand for woody biomass,has reportedly diminished soil mineral resources and may lead to degraded tree nutrition as well as growth.However,as nutrients become less available in the soil,the remobilization of nutrients in biomass tissues(plant internal cycling)helps sustain tree nutrition.Our study aims to quantify the impact of Removing Harvest Residues and Litter(RHRL)during five years on tree growth,wood density,and stem wood nutrient concentrations in young beech and oak forest stands.Result:Our study found that,RHRL significantly decreased tree growth ring width by 14%,and wood density by 3%,in beech trees,in near bark rings.RHRL also significantly reduced nutrient concentration in near bark and near pith areas of both studied species.Mg,Na and S were found lower by 44%,76%,and 56%,respectively,in near bark area of beech trees.In near bark area of oak trees,K,Ca,Mg,Na,S,and Fe were lower by 20%,25%,41%,48%,41%,and 16%,respectively.K and Mg concentrations decreased more strongly in near pith area compared to near bark area suggesting internal translocation of these two elements.Conclusion:In beech trees,wood density proved to be an important factor while quantifying the effect of removing harvest residuals on tree growth and biomass.Soil nutrient loss intensified the remobilization of nutrients con-tained in older tree rings(close to the pith)towards newly formed rings(close to bark).In our study,in beech trees,K was found to be the most recycled major nutrient.These results demonstrate the potential of such analysis for providing valuable insight into the effect of RHRL in premature stands on the physiological adaptive strategies of trees and an indication of soil fertility status.展开更多
Plant species diversity is one of the most widely used indicators in ecosystem management.The relation of species diversity with the size of the sample plot has not been fully determined for Oriental beech forests(Fag...Plant species diversity is one of the most widely used indicators in ecosystem management.The relation of species diversity with the size of the sample plot has not been fully determined for Oriental beech forests(Fagus orientalis Lipsky),a widespread species in the Hyrcanian region.Assessing the impacts of plot size on species diversity is fundamental for an ecosystem-based approach to forest management.This study determined the relation of species diversity and plot size by investigating species richness and abundance of both canopy and forest floor.Two hundred and fifty-six sample plots of 625 m^(2) each were layout in a grid pattern across 16 ha.Base plots(25 m×25 m)were integrated in different scales to investigate the effect of plot size on species diversity.The total included nine plots of 0.063,0.125,0.188,0.250,0.375,0.500,0.563,0.750 and 1 ha.Ten biodiversity indices were calculated.The results show that species richness in the different plot sizes was less than the actual value.The estimated value of the Simpson species diversity index was not significantly different from actual values for both canopy and forest floor diversity.The coefficient of variation of this index for the 1-ha sample plot showed the lowest amount across different plot sizes.Inverse Hill species diversity was insignificant difference across different plot sizes with an area greater than 0.500 ha.The modified Hill evenness index for the 1-ha sample size was a correct estimation of the 16-ha for both canopy and forest floor;however,the precision estimation was higher for the canopy layer.All plots greater than 0.250-ha provided an accurate estimation of the Camargo evenness index for forest floor species,but was inaccurate across different plot sizes for the canopy layer.The results indicate that the same plot size did not have the same effect across species diversity measurements.Our results show that correct estimation of species diversity measurements is related to the selection of appropriate indicators and plot size to increase the accuracy of the estimate so that the cost and time of biodiversity management may be reduced.展开更多
Foliar resorption is a principal nutrient conservation mechanism in terrestrial vegetation that could be sensitive to ongoing changes in climate and atmospheric nitrogen(N)deposition.We quantified N resorption in nort...Foliar resorption is a principal nutrient conservation mechanism in terrestrial vegetation that could be sensitive to ongoing changes in climate and atmospheric nitrogen(N)deposition.We quantified N resorption in northern hardwood forests along an elevation gradient of decreasing temperature and increasing soil N availability to evaluate how this critical nutrient cycling process can be expected to respond to global and regional environmental changes.Foliar N resorption proficiency(NRP)increased significantly at lower elevations for both sugar maple and American beech,the dominant species in these forests.Foliar N resorption efficiency(NRE)also decreased with increasing elevation,but only in one year.Both species exhibited strong negative relationships between NRP and soil N availability.Thus,we anticipate that with climate warming and decreasing N inputs,northern hardwood forests can be expected to exhibit stronger N conservation via foliar resorption.Both species also exhibited strong correlations between resorption efficiency of N and C,but resorption of both elements was much greater for beech than sugar maple,suggesting contrasting mechanisms of nutrient conservation between these two widespread species.展开更多
In this study,using a dendrological and isotopic approaches,we investigated the responses to climate of two pure Fagus sylvatica L.stands(Campobraca and Falode)in the southernmost part of the distribution range in sou...In this study,using a dendrological and isotopic approaches,we investigated the responses to climate of two pure Fagus sylvatica L.stands(Campobraca and Falode)in the southernmost part of the distribution range in southern Italy.Theδ^(13)C data were used for calculating the intrinsic water use efficiency(iWUE)as a proxy of the balance between the water and carbon cycles.The results showed that the iWUE of both stands was sensitive to the amount of precipitation during the summer months(negative,significant effect)and to atmospheric CO_(2)concentration.Growth was sensitive to climate only in the Campo Braca site;the most influential variables were the VPD(vapour pressure deficit)and precipitation of the summer months that had a negative and a positive effect,respectively.The iWUE showed a negative correlation with growth in Campo Braca and a non-significant one in Falode.Water availability was the most influential variable on F.sylvatica growth and physiology.The iWUE increase was mainly driven by atmospheric CO_(2)concentration,and by decreased precipitation,as a response of the trees to drought.Our results highlight the importance of understanding the hydrological changes due to climate change for forecasting/modelling forest responses.CO_(2)increase does not compensate for the effect of adverse climate on F.sylvatica in the forests of southern Italy,while local conditions play an important role in determining tree growth.展开更多
The consequences of climate change continue to threaten European forests,particularly for species located at the edges of their latitudinal and altitudinal ranges.While extensively studied in Central Europe,European b...The consequences of climate change continue to threaten European forests,particularly for species located at the edges of their latitudinal and altitudinal ranges.While extensively studied in Central Europe,European beech forests require further investigation to understand how climate change will affect these ecosystems in Mediterranean areas.Proposed silvicultural options increasingly aim at sustainable management to reduce biotic and abiotic stresses and enhance these forest ecosystems'resistance and resilience mechanisms.Process-based models(PBMs)can help us to simulate such phenomena and capture early stress signals while considering the effect of different management approaches.In this study,we focus on estimating sensitivity of two state-of-the-art PBMs forest models by simulating carbon and water fluxes at the stand level to assess productivity changes and feedback resulting from different climatic forcings as well as different management regimes.We applied the 3D-CMCC-FEM and MEDFATE forest models for carbon(C)and water(H_(2)O)fluxes in two sites of the Italian peninsula,Cansiglio in the north and Mongiana in the south,under managed vs.unmanaged scenarios and under current climate and different climatic scenarios(RCP4.5 and RCP8.5).To ensure confidence in the models’results,we preliminary evaluated their performance in simulating C and H_(2)O flux in three additional beech forests of the FLUXNET network along a latitudinal gradient spanning from Denmark to central Italy.The 3D-CMCC-FEM model achieved R^(2)values of 0.83 and 0.86 with RMSEs of 2.53 and 2.05 for C and H_(2)O fluxes,respectively.MEDFATE showed R^(2)values of 0.76 and 0.69 with RMSEs of 2.54 and 3.01.At the Cansiglio site in northern Italy,both models simulated a general increase in C and H_(2)O fluxes under the RCP8.5 climate scenario compared to the current climate.Still,no benefit in managed plots compared to unmanaged ones,as the site does not have water availability limitations,and thus,competition for water is low.At the Mongiana site in southern Italy,both models predict a decrease in C and H_(2)O fluxes and sensitivity to the different climatic forcing compared to the current climate;and an increase in C and H_(2)O fluxes when considering specific management regimes compared to unmanaged scenarios.Conversely,under unmanaged scenarios plots are simulated to experience first signals of mortality prematurely due to water stress(MEDFATE)and carbon starvation(3D-CMCC-FEM)scenarios.In conclusion,while management interventions may be considered a viable solution for the conservation of beech forests under future climate conditions at moister sites like Cansiglio,in drier sites like Mongiana conservation may not lie in management interventions alone.展开更多
This study investigated biomass allocation in young stands of European beech(Fagus sylvatica L.)and Norway spruce(Picea abies(L.)Karst.)across 31 forest sites in the Western Carpathians,Slovakia.A total of 541 trees a...This study investigated biomass allocation in young stands of European beech(Fagus sylvatica L.)and Norway spruce(Picea abies(L.)Karst.)across 31 forest sites in the Western Carpathians,Slovakia.A total of 541 trees aged 2–10 years,originating from natural regeneration and planting,were destructively sampled to quantify biomass in four components:foliage,branches,stems,and roots.Generalized non-linear least squares(GNLS)models with a weighing variance function outperformed log-transformed seemingly unrelated regression(SUR)models in terms of accuracy and robustness,especially for foliage and branch biomass.When using height as the predictor,SUR models tended to underestimate biomass in planted beech,leading to notable underprediction of aboveground and total biomass.Biomass allocation patterns varied significantly by species and regeneration origin.Using a non-linear system of equations and component ratio modelling,we found out that planted spruce displayed low variability and a consistent dominance of needle biomass,while naturally regenerated beech showed greater variability and a higher proportion of stem biomass,reflecting stronger competition-driven vertical growth.Interspecific differences in total biomass were more pronounced when using tree height,with spruce generally exhibiting greater biomass than beech at equivalent heights.Overall,stem base diameter marginally outperformed tree height as a predictor of biomass.However,tree height-based models showed strong performance and are particularly suitable for integration with remote sensing applications.These findings can directly support forest managers and modellers in comparing regeneration methods and biomass estimation approaches for early-stage stand development,carbon accounting,and remote sensing calibration.展开更多
We studied the species diversity of the herb layer and ecological factors in harvest-created gaps in beech stands under a single-tree selection system in Northern Iran. To determine diversity, the number of beech seed...We studied the species diversity of the herb layer and ecological factors in harvest-created gaps in beech stands under a single-tree selection system in Northern Iran. To determine diversity, the number of beech seedlings, and other ecological factors, 16 gaps were selected and subplots of 5 m2 were positioned at the centre and at the cardinal points of each gap. Species richness and Simpson diversity index increased with increasing gap area as did numbers of seedlings. With increasing humus layer thickness, species richness declined but the Hill evenness index increased. Species richness increased with increasing light availability. There was no relationship between crown radii of beech trees and diversity indices. Correlations between environmental factors and numbers of individuals of some species in the herb layer were not significant except in a few cases. The results help explain the effects of man-made gaps on the dynamics of managed beech stands and this benefits evaluation of silvicultural operating plans.展开更多
Height–diameter relationships are essential elements of forest assessment and modeling efforts.In this work,two linear and eighteen nonlinear height–diameter equations were evaluated to find a local model for Orient...Height–diameter relationships are essential elements of forest assessment and modeling efforts.In this work,two linear and eighteen nonlinear height–diameter equations were evaluated to find a local model for Oriental beech（Fagus orientalis Lipsky） in the Hyrcanian Forest in Iran.The predictive performance of these models was first assessed by different evaluation criteria： adjusted R^2（R^2_（adj））,root mean square error（RMSE）,relative RMSE（%RMSE）,bias,and relative bias（%bias） criteria.The best model was selected for use as the base mixed-effects model.Random parameters for test plots were estimated with different tree selection options.Results show that the Chapman–Richards model had better predictive ability in terms of adj R^2（0.81）,RMSE（3.7 m）,%RMSE（12.9）,bias（0.8）,%Bias（2.79） than the other models.Furthermore,the calibration response,based on a selection of four trees from the sample plots,resulted in a reduction percentage for bias and RMSE of about 1.6–2.7%.Our results indicate that the calibrated model produced the most accurate results.展开更多
In a reserved forest parcel in a virgin eastern Hyrcanian mixed beech forest,80 ha was surveyed to determine the pit and mound topography,canopy gaps and dead trees.The aim was to investigate the spatial patterns and ...In a reserved forest parcel in a virgin eastern Hyrcanian mixed beech forest,80 ha was surveyed to determine the pit and mound topography,canopy gaps and dead trees.The aim was to investigate the spatial patterns and correlation of pit and mound features with canopy gaps.Seventy-five canopy gaps and 61 pit and mound features were identified.The univariate first order nearest neighbor(R_(CE))and bivariate second order test(Ripley’s K)statistic were applied.R_(CE) statistics highlighted a general aggregation pattern for canopy gaps and pits and mounds,while pits and mounds alone were more clumped.Distances between canopy gaps were 130 m average,whereas distances between pit and mound features and dead trees were 60 and 78 m,respectively.Spatial positive correlation of canopy gaps with pits and mounds were observed with all distances.The result of spatial correlations between canopy gaps with pits and mounds confirmed that windthrows cause micro successions in fallen tree ecosystem-scale correlated with gap phase dynamics in the forest community-scale.展开更多
Beech （Fagus orientalis Lipsky） forest covers about 565,000 ha of land in Guilan province, north of Iran and forms a major carbon pool. It is an important economic, soil protection and recreation resource. We studie...Beech （Fagus orientalis Lipsky） forest covers about 565,000 ha of land in Guilan province, north of Iran and forms a major carbon pool. It is an important economic, soil protection and recreation resource. We studied long-term effects of fire on the structure and composition 37 years after fire occurrence in these forests. To do this research, we selected 85 ha burned and 85 ha unburned beech forests）. The results indicated that the fire had not changed the overall uneven-aged structure, but it changed forest composition from pure stands to mixed stands that now include species such as Carpinus betulus, Acer cappadocicum and Alnus subcordata. The density of trees and regeneration was significantly increased, while the density of shrubs significantly decreased. The main reasons for increased tree regeneration were attributed to （1） reduction of litter depth, and （2） increase in available light from opening of the canopy and reduction in shrub competition. It is apparent that the forest is on a path to return to its natural state before the fire after 37 years.展开更多
Background: Climate change triggered many studies showing that trends and events of environmental conditions can reduce but also accelerate growth at the stand and individual tree level. However, it is still rather un...Background: Climate change triggered many studies showing that trends and events of environmental conditions can reduce but also accelerate growth at the stand and individual tree level. However, it is still rather unknown how climate change modifies the growth partitioning between the trees in forest stands.Methods: Based on long-term girth-tape measurements in mature monospecific and mixed-species stands of Norway spruce(Picea abies(L.) Karst.) and European beech(Fagus sylvatica L.) we traced the effect of the severe droughts in 2003 and 2015 from the stand down to the tree level.Results: Stand growth of Norway spruce decreased by about 30% in the once-in-a-century drought 2015, while European beech was much more drought resistant. Water availability generally amplified size-asymmetric growth partitioning. Especially in case of Norway spruce water availability primarily fostered the growth of predominant trees, whereas drought favoured the growth of small trees at the expense of the predominant ones. We could not detect significant differences between mixed and monospecific stands in this regard.Conclusions: The drought-induced reallocation of growth in favour of small trees in case of spruce may result from its isohydric character. We hypothesize that as small trees are shaded, they can benefit from the reduced water consumption of their sun-exposed taller neighbours. In case of beech, as an anisohydric species, tall trees suffer less and smaller trees benefit less under drought. The discussion elaborates the consequences of the water dependent growth allocation for forest monitoring, growth modelling, and silviculture.展开更多
Allocation of biomass and nutrient elements including Nitrogen to above and belowground compartments of beech seedlings （Fagus sylvatica L.） treated by labeled nitrogen fertilizer in the form of 15NH4 and 15NO3 were...Allocation of biomass and nutrient elements including Nitrogen to above and belowground compartments of beech seedlings （Fagus sylvatica L.） treated by labeled nitrogen fertilizer in the form of 15NH4 and 15NO3 were investigated at the end of two successive growing seasons. Pot cultured beech seedlings were grown at a green house on intact soil cores sampled from three adjacent stands including beech, Norway spruce and mixed beech-spruce cultures of Soiling forest, Germany. Comparing biomass allocation and nutrients concentrations of the seedlings between the control and 15N-fertilized treatments revealed no significant effect of N fertilization on nutrients uptake by seedlings over the experiment. The form of N input influenced its movement into plant pools. It was demonstrated that beech seedlings take up nitrogen mainly in the form of nitrate, which is then reduced in the leaves, although the differences between the retention of NO3^--N and NH4^＋-N in plants were not statistically significant. Percent recoveries of 15N in trees were typically greater after 15NO3 than after 15NH4 additions. It was indicated that immobilization of ~SN tracer in fine roots was a slower process comparing other plant compartments such as stem and coarse roots, but a powerful sink for N during the course of study.展开更多
Background： Wood density is considered to be the most important predictor of wood quality but despite its importance, diffuse-porous tree species have been the subject of only a limited number of studies. The importa...Background： Wood density is considered to be the most important predictor of wood quality but despite its importance, diffuse-porous tree species have been the subject of only a limited number of studies. The importance of European beech forests for Central Europe calls for profound research to examine the potential impact of a warmer climate on the quality of beech timber. Methods： In this study we analysed the influence of tree-ring width and tree-ring age on the wood density of beech, and whether the wood density response to these two parameters is modified by aspect. A linear mixed-effects model for wood density was constructed for mean density data measured with high frequency densitometry on stem discs from 72 beech trees sampled from two different aspects （northeast -NE and southwest -SW） of a valley in southwestern Germany. Results： Part of the variability of mean annual wood density was explained by cambial age： an increase in cambial age resulted in an increase in mean wood density. Tree-ring width and aspect had only a small influence on wood density, Wood density on the SW aspect was lower than on the NE with a difference of approximately 0.006 g/cm3. The between-tree variability was very high. Conclusions： The significant interaction between cambial age and aspect reflects the importance of site conditions at older tree ages： with increasing cambial age the difference between aspects becomes stronger. Our results give a better understanding of the importance of site conditions on the wood quality of beech.展开更多
文摘Plant diversity plays key ecological roles in forest ecosystems, including influencing succession, resilience and nutrient cycling. This study was conducted to investigate the effect of livestock grazing and human uses on herbaceous species diversity. We surveyed 50 ha of protected area and 50 ha of unprotected area to evaluate herbaceous species diversity in oriental beech （Fagus orientalis Lipsky） forests in northern Iran. We calculated and compared three indices each of diversity and evenness, and species richness between the two areas. Herbaceous cover was higher in the unprotected area while leaf litter depth and tree canopy cover were higher in the protected area. The diversity indices, H （Shan- non-Wiener index ）, N~ （McArthur index）, N2 （Hill＇s index）, EQ （Modified Nee index）, E,ar （Smith-Wilson index）, E5 （modified index of Hill） and R=S （species richness） and species richness R=S were greater in the protected area than in the unprotected area, suggesting that protection from grazing results in increased numbers of plants and species. The effect of land protection on plant diversity was more pronounced for evenness than for species richness and the positive correlation between diversity and evenness indices was higher than that between diversity and richness.
基金supported by Karadeniz Technical University Research Fund,Project number 2010.113.001.11
文摘We studied relationships between stand structure and stand stability according to thinning intensity in an afforested oriental beech stand. Various thinning intensities were applied in sample stands. We sampled eight plots in stands that were lightly thinned, eight plots in heavily thinned stands and eight plots in unthinned stands as a control. Height and diameter distributions of the stands were measured to assess stand structure. We quantified individual tree stability and collective stability. Heavy thinning during the first thinning operation damaged the storied structure of the stand in thicket stage and affected collective structuring ability. While most control plots had multi-storied stands, after light and heavy thinning two-storied structure became more common.Large gaps occurred in the canopy after heavy thinning. On average, nine tree collectives were formed per sampling plot in the untreated stand, seven collectives after thinning in 2008 and four collectives after thinning in 2009. Stable trees accounted for 17 % of trees in control plots, 24 % in lightly thinned plots, and 15 % in heavily thinned plots. Collective stability values were 83 % in control plots, 82 % in lightly thinned plots and 36 % in heavily thinned plots. We conclude that it is necessary to retain collective structuring capacity during thinning operations for sustaining stand stability.
文摘For developing nature-based silvicultural practices in a beech forest to promote, abundance, height, vitality, and preferred growth form, regenerated trees growing in gaps were compared with those under closed canopies. A systematic 50 ×50 m grid was plotted in a beech stand in the Kheyrud Experimental Forest for selecting trees to measure variables in gaps and under closed canopies. Abundance and mean height of regenerated beech trees were significantly higher in closed canopies than in canopy gaps. Beech seedlings with excurrent growth were significantly taller within regeneration patches under closed canopy. Moreover, regenerated trees with high vitality were more abundant in closed canopy areas than in gaps. Thus, beech regeneration should improve under closed canopies; hence, gaps in a near natural forest should be created only after adequate regeneration and appropriate growth under the parent tree in a closed canopy area is ensured. The results of this research have profound implications for the sustainable management of the forest and for ensuring sustainable beech regeneration. The presence of a closed canopy cover likely will reduce potential stresses on oriental beech regeneration.
文摘Oriental Beech is the most important commercial tree species in northern Iran. In recent years wood production companies interested in felling large beech trees for profit have challenged advocates of close-to-nature silviculture who favor conservation. Our study objective was to assess the economic value of over-mature beech trees by relating tree diameter （DBH） to amount of decay. Based on the location of onset of decay, we categorized three types of decay as stump, stem, and crown decay. Trees of greater diameter （age） typically showed greater decay in the stem. Percent of decayed volume, diameter of decayed tissue, and length of decay in tree stems varied between 0.5%-64.3%, 15 cm-75 cm, and 2.0-19.5 m, respectively. With increasing trunk diameter, the propor- tion of truck decay increased. Red heart and dark red heart constituted 25% and 14.3% of sampled trees, respectively. However, we found no correlation between intensity of stem decay and morphological charac- teristics of trees. Seedlings were not abundant around the bases of over-mature trees, suggesting that the trees did not contribute to regeneration of the stand. Beech trees of diameter 〉1 m do not provide valu- able round wood for industries and cause to raise wood production costs. We recommend that these trees 〉1 m DBH should be retained in forest stands because of their low commercial value but high ecological and conservational values such as maintaining biodiversity in forest ecosystems.
文摘We studied the long-term impacts of natural burning on herbaceous species diversity 37 years after a fire occurred in the Roudbar forests of Guilan Province, northern Iran. Numerous studies have examined short-term changes in understory vegetation following wildfire; however, very few long-term studies are available or changes inferred from retrospective studies based on chro- nosequences. For this study, 170 ha of forest （85 ha, burned areas; 85 ha, unburned areas） were surveyed. Because the 1000 m2 plots were too large for detailed measurements of herbaceous species, we determined a sub-sample size according to the Whittaker＇s nested plot sampling protocol and minimal areas method. Hence, sub-plots of 32 m2 were used for herbaceous species measurements, which consisted of percent cover of each species based on the Domin criterion. We measured plant diversity （Shannon-Wiener index）, spe- cies richness （Margalef＇s index）, and evenness （Smith-Wilson index）. Mean percent cover, together with diversity, richness and even- ness, increased markedly in burned areas compared to unburned controls. This suggests that the biodiversity of these forests could be restored within 37 years after fire. However, the abundance of invasive species such as Rubus fruticosus and Bromus benekenii increased significantly in burned areas, but these could be controlled by relevant silvicultural operations.
基金This work was supported within the XyloDensMap project,INRAE funded by the French Ministry of Agriculture under the convention n°A6.01/2017.
文摘Background:Higher exportation of harvest residues from forests due to increased demand for woody biomass,has reportedly diminished soil mineral resources and may lead to degraded tree nutrition as well as growth.However,as nutrients become less available in the soil,the remobilization of nutrients in biomass tissues(plant internal cycling)helps sustain tree nutrition.Our study aims to quantify the impact of Removing Harvest Residues and Litter(RHRL)during five years on tree growth,wood density,and stem wood nutrient concentrations in young beech and oak forest stands.Result:Our study found that,RHRL significantly decreased tree growth ring width by 14%,and wood density by 3%,in beech trees,in near bark rings.RHRL also significantly reduced nutrient concentration in near bark and near pith areas of both studied species.Mg,Na and S were found lower by 44%,76%,and 56%,respectively,in near bark area of beech trees.In near bark area of oak trees,K,Ca,Mg,Na,S,and Fe were lower by 20%,25%,41%,48%,41%,and 16%,respectively.K and Mg concentrations decreased more strongly in near pith area compared to near bark area suggesting internal translocation of these two elements.Conclusion:In beech trees,wood density proved to be an important factor while quantifying the effect of removing harvest residuals on tree growth and biomass.Soil nutrient loss intensified the remobilization of nutrients con-tained in older tree rings(close to the pith)towards newly formed rings(close to bark).In our study,in beech trees,K was found to be the most recycled major nutrient.These results demonstrate the potential of such analysis for providing valuable insight into the effect of RHRL in premature stands on the physiological adaptive strategies of trees and an indication of soil fertility status.
基金funded by Gorgan University of Agricultural Sciences and Natural Resources(grant number 9318124503).
文摘Plant species diversity is one of the most widely used indicators in ecosystem management.The relation of species diversity with the size of the sample plot has not been fully determined for Oriental beech forests(Fagus orientalis Lipsky),a widespread species in the Hyrcanian region.Assessing the impacts of plot size on species diversity is fundamental for an ecosystem-based approach to forest management.This study determined the relation of species diversity and plot size by investigating species richness and abundance of both canopy and forest floor.Two hundred and fifty-six sample plots of 625 m^(2) each were layout in a grid pattern across 16 ha.Base plots(25 m×25 m)were integrated in different scales to investigate the effect of plot size on species diversity.The total included nine plots of 0.063,0.125,0.188,0.250,0.375,0.500,0.563,0.750 and 1 ha.Ten biodiversity indices were calculated.The results show that species richness in the different plot sizes was less than the actual value.The estimated value of the Simpson species diversity index was not significantly different from actual values for both canopy and forest floor diversity.The coefficient of variation of this index for the 1-ha sample plot showed the lowest amount across different plot sizes.Inverse Hill species diversity was insignificant difference across different plot sizes with an area greater than 0.500 ha.The modified Hill evenness index for the 1-ha sample size was a correct estimation of the 16-ha for both canopy and forest floor;however,the precision estimation was higher for the canopy layer.All plots greater than 0.250-ha provided an accurate estimation of the Camargo evenness index for forest floor species,but was inaccurate across different plot sizes for the canopy layer.The results indicate that the same plot size did not have the same effect across species diversity measurements.Our results show that correct estimation of species diversity measurements is related to the selection of appropriate indicators and plot size to increase the accuracy of the estimate so that the cost and time of biodiversity management may be reduced.
基金National Science Foundation supported this research through the Long Term Ecological Research(LTER)Grant to Hubbard Brook(NSF DEB1114804,1637685,and 2224545)by the project grant(NSF DEB,2020397).
文摘Foliar resorption is a principal nutrient conservation mechanism in terrestrial vegetation that could be sensitive to ongoing changes in climate and atmospheric nitrogen(N)deposition.We quantified N resorption in northern hardwood forests along an elevation gradient of decreasing temperature and increasing soil N availability to evaluate how this critical nutrient cycling process can be expected to respond to global and regional environmental changes.Foliar N resorption proficiency(NRP)increased significantly at lower elevations for both sugar maple and American beech,the dominant species in these forests.Foliar N resorption efficiency(NRE)also decreased with increasing elevation,but only in one year.Both species exhibited strong negative relationships between NRP and soil N availability.Thus,we anticipate that with climate warming and decreasing N inputs,northern hardwood forests can be expected to exhibit stronger N conservation via foliar resorption.Both species also exhibited strong correlations between resorption efficiency of N and C,but resorption of both elements was much greater for beech than sugar maple,suggesting contrasting mechanisms of nutrient conservation between these two widespread species.
基金partially supported by MIUR Project (PRIN 2020) "Unravelling interactions between WATER and carbon cycles during drought and their impact on water resources and forest and grassland ecosySTEMs in the Mediterranean climate (WATERSTEM)" (protocol code:20202WF53Z)"WAFER" at CNR (Consiglio Nazionale delle Ricerche)MIUR Project (PRIN 2017) "The Italian TREETALKER NETWORK:continuous large scale monitoring of tree functional traits and vulnerabilities to climate change"
文摘In this study,using a dendrological and isotopic approaches,we investigated the responses to climate of two pure Fagus sylvatica L.stands(Campobraca and Falode)in the southernmost part of the distribution range in southern Italy.Theδ^(13)C data were used for calculating the intrinsic water use efficiency(iWUE)as a proxy of the balance between the water and carbon cycles.The results showed that the iWUE of both stands was sensitive to the amount of precipitation during the summer months(negative,significant effect)and to atmospheric CO_(2)concentration.Growth was sensitive to climate only in the Campo Braca site;the most influential variables were the VPD(vapour pressure deficit)and precipitation of the summer months that had a negative and a positive effect,respectively.The iWUE showed a negative correlation with growth in Campo Braca and a non-significant one in Falode.Water availability was the most influential variable on F.sylvatica growth and physiology.The iWUE increase was mainly driven by atmospheric CO_(2)concentration,and by decreased precipitation,as a response of the trees to drought.Our results highlight the importance of understanding the hydrological changes due to climate change for forecasting/modelling forest responses.CO_(2)increase does not compensate for the effect of adverse climate on F.sylvatica in the forests of southern Italy,while local conditions play an important role in determining tree growth.
基金the Institute Research Centre for Ecological and Forestry Applications (CREAF) of Barcelona that supported the research by the Spanish “Ministerio de Ciencia e Innovacio'n”(MCIN/AEI/ 10.13039/501100011033) (grant agreement No. PID 2021-126679OBI00)partially supported by MIUR Project (PRIN 2020) between WATER and carbon cycles during droug“Unraveling interactionsht and their impact on water resources and forest and grassland ecosySTEMs in the Mediterranean climate (WATERSTEM)”(Project number: 20202WF53Z),“WAFER”at CNR (Consiglio Nazionale delle Ricerche)+3 种基金Priwitzer et al. (2014) (cod. 2020E52THS)-Research Projects of National Relevance funded by the Italian Ministry of University and Research entitled: “Multi-scale observations to predict Forest response to pollution and climate change”(MULTIFOR, project number: 2020E52THS)funding by the project OptForEU Horizon Europe research and innovation programme under grant agreement No. 101060554the project funded under the National Recovery and Resilience Plan (NRRP), Mission 4 Component 2 Investment 1.4-Call for tender No. 3138 of December 16, 2021, rectified by Decree n.3175 of December 18, 2021 of Italian Ministry of UniversityResearch funded by the European UnionationEU under award Number: Project code CN_00000033–Next Gener, Concession Decree No. 1034 of June 17, 2022 adopted by the Italian Ministry of University and Research, CUP B83C22002930006, Project title“National Biodiversity Future Centre-NBFC”
文摘The consequences of climate change continue to threaten European forests,particularly for species located at the edges of their latitudinal and altitudinal ranges.While extensively studied in Central Europe,European beech forests require further investigation to understand how climate change will affect these ecosystems in Mediterranean areas.Proposed silvicultural options increasingly aim at sustainable management to reduce biotic and abiotic stresses and enhance these forest ecosystems'resistance and resilience mechanisms.Process-based models(PBMs)can help us to simulate such phenomena and capture early stress signals while considering the effect of different management approaches.In this study,we focus on estimating sensitivity of two state-of-the-art PBMs forest models by simulating carbon and water fluxes at the stand level to assess productivity changes and feedback resulting from different climatic forcings as well as different management regimes.We applied the 3D-CMCC-FEM and MEDFATE forest models for carbon(C)and water(H_(2)O)fluxes in two sites of the Italian peninsula,Cansiglio in the north and Mongiana in the south,under managed vs.unmanaged scenarios and under current climate and different climatic scenarios(RCP4.5 and RCP8.5).To ensure confidence in the models’results,we preliminary evaluated their performance in simulating C and H_(2)O flux in three additional beech forests of the FLUXNET network along a latitudinal gradient spanning from Denmark to central Italy.The 3D-CMCC-FEM model achieved R^(2)values of 0.83 and 0.86 with RMSEs of 2.53 and 2.05 for C and H_(2)O fluxes,respectively.MEDFATE showed R^(2)values of 0.76 and 0.69 with RMSEs of 2.54 and 3.01.At the Cansiglio site in northern Italy,both models simulated a general increase in C and H_(2)O fluxes under the RCP8.5 climate scenario compared to the current climate.Still,no benefit in managed plots compared to unmanaged ones,as the site does not have water availability limitations,and thus,competition for water is low.At the Mongiana site in southern Italy,both models predict a decrease in C and H_(2)O fluxes and sensitivity to the different climatic forcing compared to the current climate;and an increase in C and H_(2)O fluxes when considering specific management regimes compared to unmanaged scenarios.Conversely,under unmanaged scenarios plots are simulated to experience first signals of mortality prematurely due to water stress(MEDFATE)and carbon starvation(3D-CMCC-FEM)scenarios.In conclusion,while management interventions may be considered a viable solution for the conservation of beech forests under future climate conditions at moister sites like Cansiglio,in drier sites like Mongiana conservation may not lie in management interventions alone.
基金funded by the grant“EVA4.0”,No.Z.02.1.01/0.0/0.0/16_019/0000803 supported by OP RDE as well as by the projects APVV-19-0387,APVV-22-0056,and APVV-23-0293 from the Slovak Research and Development Agencyco-funded by the European Commission under the Horizon Europe Teaming for Excellence action+1 种基金project Ligno Silvagrant agreement No.101059552。
文摘This study investigated biomass allocation in young stands of European beech(Fagus sylvatica L.)and Norway spruce(Picea abies(L.)Karst.)across 31 forest sites in the Western Carpathians,Slovakia.A total of 541 trees aged 2–10 years,originating from natural regeneration and planting,were destructively sampled to quantify biomass in four components:foliage,branches,stems,and roots.Generalized non-linear least squares(GNLS)models with a weighing variance function outperformed log-transformed seemingly unrelated regression(SUR)models in terms of accuracy and robustness,especially for foliage and branch biomass.When using height as the predictor,SUR models tended to underestimate biomass in planted beech,leading to notable underprediction of aboveground and total biomass.Biomass allocation patterns varied significantly by species and regeneration origin.Using a non-linear system of equations and component ratio modelling,we found out that planted spruce displayed low variability and a consistent dominance of needle biomass,while naturally regenerated beech showed greater variability and a higher proportion of stem biomass,reflecting stronger competition-driven vertical growth.Interspecific differences in total biomass were more pronounced when using tree height,with spruce generally exhibiting greater biomass than beech at equivalent heights.Overall,stem base diameter marginally outperformed tree height as a predictor of biomass.However,tree height-based models showed strong performance and are particularly suitable for integration with remote sensing applications.These findings can directly support forest managers and modellers in comparing regeneration methods and biomass estimation approaches for early-stage stand development,carbon accounting,and remote sensing calibration.
文摘We studied the species diversity of the herb layer and ecological factors in harvest-created gaps in beech stands under a single-tree selection system in Northern Iran. To determine diversity, the number of beech seedlings, and other ecological factors, 16 gaps were selected and subplots of 5 m2 were positioned at the centre and at the cardinal points of each gap. Species richness and Simpson diversity index increased with increasing gap area as did numbers of seedlings. With increasing humus layer thickness, species richness declined but the Hill evenness index increased. Species richness increased with increasing light availability. There was no relationship between crown radii of beech trees and diversity indices. Correlations between environmental factors and numbers of individuals of some species in the herb layer were not significant except in a few cases. The results help explain the effects of man-made gaps on the dynamics of managed beech stands and this benefits evaluation of silvicultural operating plans.
基金This research received no specific grant from any funding agency in the public,commercial,or not-for-profit sectors
文摘Height–diameter relationships are essential elements of forest assessment and modeling efforts.In this work,two linear and eighteen nonlinear height–diameter equations were evaluated to find a local model for Oriental beech（Fagus orientalis Lipsky） in the Hyrcanian Forest in Iran.The predictive performance of these models was first assessed by different evaluation criteria： adjusted R^2（R^2_（adj））,root mean square error（RMSE）,relative RMSE（%RMSE）,bias,and relative bias（%bias） criteria.The best model was selected for use as the base mixed-effects model.Random parameters for test plots were estimated with different tree selection options.Results show that the Chapman–Richards model had better predictive ability in terms of adj R^2（0.81）,RMSE（3.7 m）,%RMSE（12.9）,bias（0.8）,%Bias（2.79） than the other models.Furthermore,the calibration response,based on a selection of four trees from the sample plots,resulted in a reduction percentage for bias and RMSE of about 1.6–2.7%.Our results indicate that the calibrated model produced the most accurate results.
基金supported by Gorgan University of Agricultural Sciences and Natural Resources
文摘In a reserved forest parcel in a virgin eastern Hyrcanian mixed beech forest,80 ha was surveyed to determine the pit and mound topography,canopy gaps and dead trees.The aim was to investigate the spatial patterns and correlation of pit and mound features with canopy gaps.Seventy-five canopy gaps and 61 pit and mound features were identified.The univariate first order nearest neighbor(R_(CE))and bivariate second order test(Ripley’s K)statistic were applied.R_(CE) statistics highlighted a general aggregation pattern for canopy gaps and pits and mounds,while pits and mounds alone were more clumped.Distances between canopy gaps were 130 m average,whereas distances between pit and mound features and dead trees were 60 and 78 m,respectively.Spatial positive correlation of canopy gaps with pits and mounds were observed with all distances.The result of spatial correlations between canopy gaps with pits and mounds confirmed that windthrows cause micro successions in fallen tree ecosystem-scale correlated with gap phase dynamics in the forest community-scale.
文摘Beech （Fagus orientalis Lipsky） forest covers about 565,000 ha of land in Guilan province, north of Iran and forms a major carbon pool. It is an important economic, soil protection and recreation resource. We studied long-term effects of fire on the structure and composition 37 years after fire occurrence in these forests. To do this research, we selected 85 ha burned and 85 ha unburned beech forests）. The results indicated that the fire had not changed the overall uneven-aged structure, but it changed forest composition from pure stands to mixed stands that now include species such as Carpinus betulus, Acer cappadocicum and Alnus subcordata. The density of trees and regeneration was significantly increased, while the density of shrubs significantly decreased. The main reasons for increased tree regeneration were attributed to （1） reduction of litter depth, and （2） increase in available light from opening of the canopy and reduction in shrub competition. It is apparent that the forest is on a path to return to its natural state before the fire after 37 years.
基金European UnionAward Number 281116ERA02S+9 种基金Recipient:Hans PretzschProject Title:“Management of mixedspecies stands.Options for a low-risk forest management(REFORM)”Bayerisches Staatsministerium fur Ernahrung,Landwirtschaft und Forsten (Bavarian State Ministry for Nutrition,Agriculture,and Forestry)Award Number W07 7831–22209-2013Recipient:Hans PretzschProject Title:“Long-term experimental plots for forest growth and yield research”Deutsche Forschungsgemeinschaft (German Science Foundation)Award Number PR 292/12–1Recipient:Hans PretzschProject Title:“Tree and stand-level growth reactions on drought in mixed versus pure forests of Norway spruce and European beech”
文摘Background: Climate change triggered many studies showing that trends and events of environmental conditions can reduce but also accelerate growth at the stand and individual tree level. However, it is still rather unknown how climate change modifies the growth partitioning between the trees in forest stands.Methods: Based on long-term girth-tape measurements in mature monospecific and mixed-species stands of Norway spruce(Picea abies(L.) Karst.) and European beech(Fagus sylvatica L.) we traced the effect of the severe droughts in 2003 and 2015 from the stand down to the tree level.Results: Stand growth of Norway spruce decreased by about 30% in the once-in-a-century drought 2015, while European beech was much more drought resistant. Water availability generally amplified size-asymmetric growth partitioning. Especially in case of Norway spruce water availability primarily fostered the growth of predominant trees, whereas drought favoured the growth of small trees at the expense of the predominant ones. We could not detect significant differences between mixed and monospecific stands in this regard.Conclusions: The drought-induced reallocation of growth in favour of small trees in case of spruce may result from its isohydric character. We hypothesize that as small trees are shaded, they can benefit from the reduced water consumption of their sun-exposed taller neighbours. In case of beech, as an anisohydric species, tall trees suffer less and smaller trees benefit less under drought. The discussion elaborates the consequences of the water dependent growth allocation for forest monitoring, growth modelling, and silviculture.
基金supported by Institut für Bodenkunde und Waldernhrung,Georg-August-Universitt,Gttingen,Germany.
文摘Allocation of biomass and nutrient elements including Nitrogen to above and belowground compartments of beech seedlings （Fagus sylvatica L.） treated by labeled nitrogen fertilizer in the form of 15NH4 and 15NO3 were investigated at the end of two successive growing seasons. Pot cultured beech seedlings were grown at a green house on intact soil cores sampled from three adjacent stands including beech, Norway spruce and mixed beech-spruce cultures of Soiling forest, Germany. Comparing biomass allocation and nutrients concentrations of the seedlings between the control and 15N-fertilized treatments revealed no significant effect of N fertilization on nutrients uptake by seedlings over the experiment. The form of N input influenced its movement into plant pools. It was demonstrated that beech seedlings take up nitrogen mainly in the form of nitrate, which is then reduced in the leaves, although the differences between the retention of NO3^--N and NH4^＋-N in plants were not statistically significant. Percent recoveries of 15N in trees were typically greater after 15NO3 than after 15NH4 additions. It was indicated that immobilization of ~SN tracer in fine roots was a slower process comparing other plant compartments such as stem and coarse roots, but a powerful sink for N during the course of study.
基金the collaborative research project SFB 433("Buchendominierte Laubwalder unter dem Einfluβ von Klima und Bewirtschaftung:Okologische,waldbauliche und sozialwissenschaftliche Analysen"-Beech dominated deciduous forests under the influence of climate and forest management)funded by the project Bu Klim within Waldklimafods program of BMEL/BMUB("Gefordert durch Bundesministerium für Ernahrung und Landwirtschaft und das Bundesministerium für Umwelt,Naturschutz,Bau und Reaktorsicherheit aufgrund eines Beschlusses des Deutschen Bundestages")
文摘Background： Wood density is considered to be the most important predictor of wood quality but despite its importance, diffuse-porous tree species have been the subject of only a limited number of studies. The importance of European beech forests for Central Europe calls for profound research to examine the potential impact of a warmer climate on the quality of beech timber. Methods： In this study we analysed the influence of tree-ring width and tree-ring age on the wood density of beech, and whether the wood density response to these two parameters is modified by aspect. A linear mixed-effects model for wood density was constructed for mean density data measured with high frequency densitometry on stem discs from 72 beech trees sampled from two different aspects （northeast -NE and southwest -SW） of a valley in southwestern Germany. Results： Part of the variability of mean annual wood density was explained by cambial age： an increase in cambial age resulted in an increase in mean wood density. Tree-ring width and aspect had only a small influence on wood density, Wood density on the SW aspect was lower than on the NE with a difference of approximately 0.006 g/cm3. The between-tree variability was very high. Conclusions： The significant interaction between cambial age and aspect reflects the importance of site conditions at older tree ages： with increasing cambial age the difference between aspects becomes stronger. Our results give a better understanding of the importance of site conditions on the wood quality of beech.
