Phenological phases of common beech (Fagus sylvatica L.) and their dependence on region and altitude in Southern Germany

Long-term observational data of the phenological network of the German Weather Service (DWD) were used to derive the altitudinal dependence of leaf unfolding (LU), leaf discolouration (LD) and the length of the vegetation period (LD-LU) of common beech (Fagus sylvatica L.) in different regions of Southern Germany. Depending on altitude, the vegetation period starts in Bavaria between the 2nd and 24th May with a delay of around 2 days per 100 m altitude increase (averaged over the last 30 years). In Southern Bavaria, LU starts a few days earlier as well as increases slightly faster into higher altitudes than in the northern regions. This can be explained by a more favourable supply of warmth and radiation in the south. The end of the vegetation period, defined here as the beginning of LD, showed no altitudinal dependence, which reflects an interaction of endogenous and exogenous factors as triggers for LD. In the Bavarian Forest region, altitudinal dependences are strongly influenced by sites which are affected by cold-air tailback (‘Kaltluftstau’) was obtained. Hence, phenological gradients should only be derived and applied above a phenological inversion zone. The calculated vertical gradients for LU can be used together with available observational data to estimate the beginning of the forest vegetation period of beech for different altitudes of Bavaria.     

Genetic structure of populations of beech (Fagus sylvatica L.) in Denmark

The genetic structure in Danish indigenous and introduced provenances of beech (Fagus sylvatica L.) is described. Ten stands are analysed based on variation revealed by seven enzyme systems. The present study reveals significant allele frequency differences between populations with a small absolute amount of genetic differentiation, with no pattern related to geographic distance. The genetic differentiation between indigenous populations measured by Wright's F _ST‐statistic does not deviate significantly from zero, and only 2.9% of the total genetic variation in the foreign populations was due to interpopulation differences, The allele frequencies in relatively small and isolated populations do not differ from the frequencies found in larger areas with beech forest. The low genetic differentiation of isozyme level between indigenous and foreign populations was opposed by significant differences in foliation performance measured on three pairs of neighbouring stands. Foliation performance in adjacent stands of indigenous and introduced beech indicates that quantitative characteristics must be considered separately from neutral variation at single loci.     

Economic evaluation of cyclic regimes in beech (fagus sylvatica L.)

A general model for comparing the profitability of natural regeneration in beech (Fagus sylvatica L.) with planting of Norway spruce (Picea abies L.) is developed. The model is basically a method of comparing a series of net present values. An example of using the model is given: BEECH (90–110) denotes a cyclic regime in which regeneration is initiated at age 90 of a stand, and the canopy trees are cleared at age 110. This regime is compared with the alternative of planting Norway spruce. Economic data used for the comparison are representative of the southeastern regions of Denmark. The conclusion of the comparison is as follows: BEECH (90–110) is superior to planting of Norway spruce for required real rates of return up to 3–4%. For higher required rates of return planting of Norway spruce is preferable.     

Gap formation in Danish beech (Fagus sylvatica) forests of low management intensity: soil moisture and nitrate in soil solution

Soil moisture content (0–90 cm depth) and nitrate-nitrogen (NO₃-N) concentrations in soil solution (90 cm depth) were monitored after gap formation (diameter 15–18 m) in three Danish beech-dominated forests on nutrient-rich till soils. NO₃-N drainage losses were estimated by the water balance model WATBAL for one of the sites. Two forests were non-intervention forests (semi-natural and unmanaged), the third was subject to nature-based management. The study was intended to assess the range of effects of gap formation in forests of low management intensity. In the unmanaged and the nature-based managed forest, soil solution was collected for 5 years and soil moisture measured in the fourth year after gap formation. Average NO₃-N concentrations were significantly higher in the gaps (9.9 and 8.1 mg NO₃-N l⁻¹, respectively) than under closed canopy (0.2 mg l⁻¹). In the semi-natural forest, measurements were carried out up to 29 months after gap formation. Average NO₃-N concentrations in the gap were 19.3 mg NO₃-N l⁻¹. Gap formation alone did not account for this high level, as concentrations were high also under closed canopy (average 12.4 mg NO₃-N l⁻¹). However, the gap had significantly higher N concentrations when trees were in full leaf, and NO₃-N drainage losses were significantly increased in the gap. No losses occurred under closed canopy in growing seasons. Soil moisture was close to field capacity in all three gaps, but decreased under closed canopy in growing seasons. In the semi-natural forest, advanced regeneration and lateral closure of the gap affected soil moisture levels in the gap in the last year of the study.     

Dominant radial and height growth reveal comparable historical variations for common beech in north-eastern France

While forest productivity is usually inferred from height growth indices, retrospective analyses of tree rings have been intensively used to assess long-term trends in forest productivity. However, radial growth is sensitive to the degree of competition between trees and influenced by management practices or local disturbances. Whether radial growth is accurate for diagnosing and quantifying productivity changes remains a debated question.     

Notes on beech (Fagus crenata Blume) seed and seedling mortality due to rodent herbivory in a northernmost beech forest, Utasai, Hokkaido

We investigated the mortality of post-dispersal seeds and current-year seedlings of a beech species (Fagus crenata Blume) at the northernmost extent of its geographical range at Utasai (Kuromatsunai), Hokkaido, Japan. Introduced seeds that were protected from rodent predation by wire cages had a higher survival rate from winter to the following spring (mean ± SD: 84 ± 16.7%,n = 10) than controls without cages (mean ± SD: 12 ± 17.9%,n = 10). Unprotected transplanted current-year seedlings with almost fully opened primary leaves never survived more than three days (0%,n = 30), while more than 80% (n = 30) of seedlings within cages survived to the end of the growing season. These results indicate that rodent herbivory is a major mortality factor that strongly prevents the establishment of beech seedlings. In a natural population, we observed that 90% of seedlings (n = 197) disappeared within 10 days after their emergence and rodent herbivory caused this heavy mortality. A herbivorous rodent, the gray-backed vole,Clethrionomys rufocanus bedfordiae, was remarkably abundant in late June (101.5 voles/ha), suggesting that this species strongly affects the formation of the seedling bank. The presence ofC. rufocanus bedfordiae may be one of the reasons for the scarcity of beech seedlings and saplings and the rarity of recruitment in this northernmost beech forest.     

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.     

Development of oak and beech seedlings planted under varying shelterwood densities and with different site preparation methods in southern Sweden

Effects of site preparation, shelterwood density and planting depth on the survival and growth of planted beech and oak seedlings were studied. Experiments were performed in one oak and one beech stand in southern Sweden. Two areas with different densities of shelterwood and one clearcut were established in each stand. Growth, damage and survival of the planted seedlings were observed for three years. Soil water potential was recorded weekly and radiation and soil temperatures were recorded continuously during the growing seasons.Neither site preparation methods nor planting depth affected oak seedling growth, when planting was carried out on fresh clearcuts or in shelterwoods, while growth of beech seedlings was positively affected by mounding. Growth of oak seedlings was inhibited by the shelterwood treatments. In beech seedlings, growth was lowest in dense shelterwood, while there was no difference in growth between seedlings on the clearcut area and in the shelterwood of low density. Differences in growth may be explained by differences in radiation and soil water potential.When planting was carried out on a one-year-old clearcut, site preparation improved the subsequent growth of oak and beech seedlings.     

The influences of log storage and kiln drying climate on the colour of non-steamed beech (Fagus sylvatica L.) wood

Abstract

Discolouration during the drying of non-steamed beech is a major industrial concern. The generally preferred yellow-white colour can easily develop a reddish or dull grey appearance during drying. In this study, the influences of log storage time and kiln drying climate on the colour of non-steamed sawn beech have been investigated and quantified. Samples 27×87 mm in cross section were dried in laboratory kilns and the average colour, in CIELAB colour space was measured on dry planed surfaces using a photoelectric colorimeter. Log storage for 13 weeks under low-temperature conditions had no visible effect. The reddish discolouration is mainly temperature related while the greyish discolouration is mainly controlled by the equilibrium moisture content (EMC) during the initial drying. Within the investigated climate interval, the EMC was twice as important as temperature for the final colour. Regression models developed show that, as long as the EMC is kept below 15%, a temperature of up to 37°C can be allowed without any visually detectable discolouration.     

Drought matters - Declining precipitation influences growth of Fagus sylvatica L. and Quercus robur L. in north-eastern Germany

For north-eastern Germany regional climate models project rising temperatures in combination with decreasing summer and increasing winter precipitation. The resulting overall drier conditions during the growing season will considerably impact forest growth there. We evaluate the consequences of increasing drought on the growth of the two locally most important broadleaf tree species common beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.). Three mixed forests of beech and oak were sampled along a west-east gradient of declining precipitation. In total we used 257 ring-width samples from 133 trees to build six species and site specific chronologies. Additionally, we modelled the soil water budget for each site. We performed continuous and discontinuous (pointer year) analysis of climate-tree-growth relationships with particular emphasis on inter-annual-variations and their dependence upon climatic factors (temperature, precipitation, soil moisture) and on the stability of the obtained relationships. Results of climate-growth correlations together with pointer year analysis indicate a strong dependency of growth of both species from water availability, especially during early summer (June and July). General correlation pattern between growth and climate are similar for both species, but climate sensitivity of beech is generally higher. We identified drought as the main driver of negative growth depressions in both species. Increasing drought stress along the gradient is expressed in higher correlations to climatic variables, higher sensitivity (variance) of growth, and a higher number of negative pointer years for both species. For beech we also found a significant trend of decreasing average growth rates along the gradient. Growth superiority of beech compared to oak declines with decreasing precipitation. The relationships were generally stable throughout the 20th century. A rise of sensitivity together with a higher frequency of negative pointer years during the last decades suggests that increasing climatic variability together with rising temperatures might be influencing growth of Fagus at the more humid sites. If we substitute space by time it seems that already small changes in precipitation regime can have considerable impact, especially on the growth of beech. Other, more drought tolerant species like oak might gain competitive advantages under the projected climatic changes.     

Canopy recruitment in the beech (<Emphasis Type="Italic">Fagus engleriana</Emphasis>) forest of Mt. Shennongjia,Central China

A dendrochronological technique was used to investigate canopy recruitment of beech (Fagus engleriana Seem.) trees in western Hubei Province of Central China by identifying growth releases through gap formation. The results indicated that 83% and 17% of the canopy trees regenerate in gaps and the understory, respectively. Radial growth analyses demonstrated that there are three modes for beech regeneration and canopy recruitment: (1) gap origin without suppression; (2) gap origin with period(s) of suppression; and (3) understory origin. Multiple periods of release and suppression imply that beech regeneration and canopy recruitment are very sensitive to gap formation.     

65—70mm厚欧洲山毛榉锯材干燥基准的研究

对65－70mm厚的欧洲山毛榉锯材干燥生产进行了初步试验研究，提出了可供企业参考的木材加工生产用干燥基准。     

Soluble N compound profiles and concentrations in European beech (Fagus sylvatica L.) are influenced by local climate and thinning

We assessed seasonal changes of total soluble nonprotein nitrogen compounds (TSNN) in adult European beech trees (Fagus sylvatica, L.) growing under different local climate during the growing season immediately following a thinning treatment and 3 years later. In both years, samples of leaves, xylem sap and phloem exudates from beech trees growing in thinned and unthinned (control) stands on a dry, warm SW exposed and a cooler, moist NE exposed site were collected in May, July and September. In May of both years, asparagine (Asn) and glutamine (Gln) were most abundant in leaves and xylem, respectively, whereas arginine (Arg) dominated in the phloem. In July, TSNN concentrations decreased in all tissues and sites, but differences in water availability between aspects were reflected in TSNN concentrations. In September, differences in the increase of Arg concentration in the phloem were related to differences in the onset of senescence between treatments. Thinning treatment increased amino compound concentrations of beech tissues in July on both aspects, particularly at the NE thinned site. It is supposed that, the N balance of adult beech is favoured by both, the thinning treatments as well as the cool-moist climate prevailing at the NE aspect.     

Spatial and seasonal variations in photosynthetic properties within a beech (Fagus crenata Blume) crown

The light response curve and the intercellular CO₂ concentration response curve of CO₂ assimilation rate were investigated together with the light conditions at the four different heights within the beech crown from 1995 to 1997 on Mt. Fuji in Japan. On the seasonal fluctuation, the CO₂ assimilation rate at light saturated condition increased rapidly in May, and attained to the maximum between the end of June and July, thereafter, slightly decreased until the middle of August and rapidly decreased in September and October. The daily sum of photosynthetic photon flux density attenuated with deeping within the crown, and particularly, the relative value on 2nd position dropped to only 30%. TheA _max decreased from 10 to 5μmol m⁻² s⁻¹, approximately, with deeping within the crown. The light saturation point, quantum yield, light compensation point and dark respiration rate also varied with deeping. These results suggest that the photosynthetic properties vary gradually from sun to shade leaves along the light attenuation within a beech crown. At light saturated condition, the stomatal conductance and mesophyll conductance were strongly correlated withA _max among the four different heights (r > 0.96, respectively). TheC _i/C _a ratio was around 0.8, and there were no remarkable differences among the four different heights. These results suggest that the vertical gradient ofA _max depends on the variation of mesophyll conductance. The stomatal conductance may be also one of the major factor in the vertical gradient ofA _max. However the intercellular CO₂ concentration doesn’t influence the vertical gradient ofA _max within the crown. This work is supported by the Sasagawa Scientific Research Grant from The Japan Science Society and Grant-in-Aid for Scientific Research (C).     

Recent changes in forest productivity: An analysis of national forest inventory data for common beech (Fagus sylvatica L.) in north-eastern France

Changes in forest growth have been found in European forests and worldwide. However most observations have been derived from samples of restricted size, whose representativeness at a regional forest scale is questionable. National forest inventories provide an interesting perspective for both regional scale assessment of these trends and the investigation of their variations over environmental gradients, but have been little used.     

Investigations on viruses from declining beech trees (Fagus sylvatica L.) in Rhineland and Westfalia,Federal Republic of Germany

Viruses of the Poty-, Potex- and Tobamo-virus groups, and presumably Nepoviruses could be isolated from degenerating beech trees (Fagus sylvatica L.) in West-Germany. They are discussed as predisposing factors of tree decline.     

Effects of selective thinning on growth and development of beech (Fagus sylvatica L.) forest stands in south-eastern Slovenia

We studied the effects of two types of selective thinning on beech stands formed by a shelterwood cut in 1910 — with lower number of crop trees and higher thinning intensity (T1) and higher number of crop trees with lower thinning intensity (T2). The stands were thinned in 1980, 1991 and 2001. Despite a lower stand density after thinning, the annual basal area increments of thinned stands in both thinning periods (1980–1991 and 1991–2002) were around 20% higher compared to those of the control (unthinned) stands. The mean annual basal area increment of dominant trees was 30–56% larger in the thinned plots compared to the control plots. Of 176 initial crop trees in the T1, 72% were chosen again during the last thinning. In the T2, 258 crop trees were chosen in the first thinning, and only 62% of these trees were chosen again during the last thinning. Only crown suppression and diameter classes of crop trees significantly influenced their basal area increment when diameter classes, crown size, crown suppression, and social status were tested. In the thinned stands, the dominant trees are more uniformly distributed if compared to the dominant trees in the control plots. Finally, the herbaceous cover and the species diversity were higher in the thinned plots.     

Frost damage on the terminal shoot as a risk factor of fork incidence on common beech (Fagus sylvatica L.)

Forking is a frequent flaw in beech which can adversely affect the formation of high quality logs. Even though the hypothesis that late frosts increase forking is widely accepted by forest managers, it remains to be proved and quantified experimentally. Damage caused by a late frost on 15 May 1995 to young beech trees in an experimental site at Lyons-La-Forêt (France) and the consequences on the development of forking were analysed. The beech trees were 17 years old with heights between 1.5 and 9 m. Of all the plantation densities tested only the lowest density (625/ha) showed various and severe frost damage. Three levels of immediate damage to the growing terminal shoot were observed, with increasing intensity from: (1) no trace of frost damage, (2) reddened leaves and (3) frost-nipped shoots. The intensity of visible damage to the terminal shoot appeared to be very dependent on the height of the plant which justified our original experimental approach, based on an evaluation of the damage for each individual. Almost all of the beech trees less than 4 m high suffered frost damage and all those taller than 7 m were unaffected. Three-year-old forks due to frost which occured in May 1995 were recorded at the end of 1997 using an objective definition based on dendrometric and age criteria. The worst frost damage resulted in a significant increase in the risk of fork emergence (71.7% of the plants were forked). With a risk related to forking in the presence of frost-nipped shoots equal to 2.15 and a fraction of the attributed risk of 0.36 it was concluded that the late frost observed played a significant role in the degradation of the shape of the beech affected by frost and the overall quality of the stand.     

Effects of weather conditions on mast year frequency in beech (Fagus sylvatica L.) in Sweden

During the last two decades, an increase of the frequency ofnaturally regenerated beech seedlings has been reported. Thismay be due to an increased masting. In this investigation insouthern Sweden, mast year frequency, mast crop size and therelationships between mast year and climatic variables werestudied. Our analysis shows that while the average mast yearinterval was 4–6 years from the end of the seventeenthcentury up to the 1960s, the mean interval has decreased to2.5 years during the most recent 30 years, and there have beentwo consecutive mast years on two occasions during this laterperiod. Mast years have often followed years in which the temperaturein July and September was higher than the 30-year mean. However,there were significant variations in the amounts of beechnutsproduced between different mast years, and beechnut productionincreased with increasing site index. Climatic changes, especiallyincreases in temperature, may have been responsible for thehigher frequency of mast years, but increased atmospheric nitrogendeposition may also have been a contributory factor.     

The effect of forest management on trace gas exchange at the pedosphere–atmosphere interface in beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) forests stocking on calcareous soils

The effect of forest management (thinning) on in situ carbon dioxide (CO₂), methane (CH₄) and nitrous oxide (N₂O) trace gas exchange between soil and atmosphere was studied in three consecutive years at three beech forest sites, which differ in aspect [southwest (SW), northeast (NE), northwest (NW)]. At all sites adjacent thinning plots (“T”) and untreated control plots (“C”) were established. Measurements at the SW and NE sites covered the years 4–6 after thinning while at the NW site measurements covered the year before and the first 2 years after thinning. Mean N₂O fluxes were <3 μg N₂O–N m⁻² h⁻¹ at all plots except for the newly thinned NWT plot. CH₄ uptake was rather low, too. Very low CH₄ oxidation rates during dry periods are explained by physiological drought stress for CH₄ oxidizers. Heterotrophic litter decomposition constitutes the largest part of total soil respiration. On the whole, no significant positive or negative effects of the silvicultural treatment on the magnitude of CO₂-, CH₄- and N₂O-trace gas exchange could be observed at the SW site 4–6 years after thinning. Also at the NE site, no effects of thinning on CO₂ and N₂O fluxes could be demonstrated. However, at this site a significant moisture-induced lower CH₄ uptake could be shown. At the NW site forest management led to a dramatic increase in N₂O emissions in the first two summers after thinning and to distinct effects on CO₂ emissions and CH₄ uptake in the first year after the felling. The unambiguous effects of thinning at the NW site are mainly related to higher C input by dead residues leading to enhanced mineralization activity, to a shift in the competition for nutrients favoring microorganisms as compared to trees and to changes in the soil water availability at the thinned plot. Considering the data obtained from the NE and SW site we expect that with the development of an understorey vegetation at the NW site the observed effects on the magnitude of trace gas exchange due to thinning will continue to decline in the following years. Our results implicate that it is indispensable to take account of the effects of forest management in order to accurately calculate trace gas emission inventories for the investigated forest ecosystem in case thinning took place immediately before.