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.     

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.     

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.     

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.     

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.     

Impacts of drought on mineral macro- and microelements in provenances of beech (Fagus sylvatica L.) seedlings

Beech seedlings originating from 11 German provenances with different climatic conditions were grown in pots and cultivated in a greenhouse. The composition of macro- and microelements in roots, axes and leaves was measured after half of the seedlings were subjected to a simulated summer drought. The recently described sensitivity of these provenances to drought was compared with drought-mediated changes in the elemental and ionic composition in organs of the seedlings; in addition, partitioning between roots and shoots was evaluated. A number of element concentrations were decreased in roots due to drought (K 94% of control, Mg 94%, Mn 75% and Zn 85%). However, chloride concentration increased in all organs (115-125%) and was the only element affected in leaves. Some changes in ionome can be related to sensitivity of provenances, but it is difficult to decide whether these changes are a result of, or a reason for, drought tolerance or sensitivity. Observed increases in chloride concentration in all plant parts of drought-treated beech seedlings can be explained by its function in charge balance, in particular since the level of phosphate was reduced. As a result of chloride accumulation, the sum of added charges of anions (and cations) in water extracts of leaf and root material was similar between drought and control plants. Since only the partitioning of Ca and Al (both only in axis) as well as Mn was affected and other elements (together with previously observed effects on C, N, S and P) remained unaffected by drought in all provenances, it can be concluded that direct effects by means of mass flow inhibition in xylem and phloem are unlikely. Secondary effects, for example on the pH of transport sap and the apoplastic space, cannot be excluded from the present study. These effects may affect partitioning between the apoplast and symplast and therefore may be significant for drought sensitivity.     

Genetic variation of beech (Fagus sylvatica L.) in Rodopi (N.E. Greece)

The recent taxonomic classification of beech in Europe considers existence of one species (Fagus sylvatica L.) with two subspecies: F. sylvatica ssp. sylvatica and F. sylvatica ssp. orientalis. Four beech populations growing on the Greek part of the Rodopi Mountains were studied using morphological traits as well as DNA molecular markers (AFLPs and chloroplast DNA SSR). The aim of the study was to describe the variation patterns of beech in the Rodopi Mountains and to test the hypothesis of possible introgression between the beech subspecies’ sylvatica and orientalis in this area. Both morphological traits and gene markers revealed a possible influence of F. orientalis on the east side of Rodopi and at the low elevations, while characters resembling F. sylvatica were observed mainly on the western part of the mountains and in higher altitudes. There was a clinal increase of genetic diversity from the west to the east, reaching a level firstly reported for beech populations. These results can be explained either by the existence of a main refugial area for beech during the last glaciation or by the occurrence of a recent hybridization among the subspecies, which were spatially isolated during the last glaciation and came into reproductive contact during their postglacial remigration. These two scenarios are not necessarily mutually exclusive.     

Influence of different forest conversion strategies on ground vegetation and tree regeneration in pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) stands: a case study in NE Germany

The conversion of anthropogenic into more natural, self-regenerating forests is one of the major objectives of forestry throughout Europe. In this study, we present investigations on permanent plots with different silvicultural treatment in NE German pine stands. Management of old-growth pine stands on acidic and nutrient-poor sandy sites differs in fencing, thinning, and planting of certain tree species. The investigations were carried out on the community, population, and individual level of the pine forest ecosystems. Thus, vegetation changes, size and height of tree populations, and height increment of tree individuals were observed over a time span of 6 years. Special attention was paid to short-lived tree species such as, e.g., Frangula alnus and Sorbus aucuparia, as well as to Fagus sylvatica as one of the most typical forest tree species of Central Europe. Vegetation changes are interpreted as a consequence of natural regeneration of formerly degraded forest sites, involving an increase in nutrient availability. High browsing pressure can be considered as a key factor for the inhibition of tree seedlings and growth of saplings. Some Sorbus aucuparia individuals, however, succeeded in growing out of the browsing height also in unfenced stands. Few found specimens of Fagus sylvatica proved that this species is able to establish spontaneously on these relatively dry, acidic sites under continental climate influence. Such natural regeneration processes, also including spontaneous rejuvenation of trees, can be integrated into silviculture as passive forest conversion management. An active management like thinning of stands, planting of trees, and fencing can accelerate forest conversion with regard to height growth and species number of trees.     

Effects of litter quality and parent material on organic matter characteristics and N-dynamics in Luxembourg beech and hornbeam forests

To test effects of litter quality and soil conditions on N-dynamics, we selected seven forests in Luxembourg dominated by beech (Fagus sylvatica, L.) and hornbeam (Carpinus betulus L.), and located on acid loam, decalcified marl or limestone, and measured organic matter characteristics, microbial C and N and net N-mineralization in a laboratory incubation experiment. Organic layer characteristics were significantly affected by species, with lower litter decay and higher accumulation under the less palatable beech, even on limestone. However, beech and hornbeam did not show any differences in N-cycling at all. Instead of species, N-cycling was affected by site conditions, albeit different than expected. Microbial N generally increased from acid loam to limestone, but acid loam showed higher net N-mineralization, especially in the organic layer. Also, acid loam showed high instead of low efficiency of N-mineralization per unit microbe, in both organic layer and mineral topsoil. In addition, acid loam showed net consumption of DOC instead of release in both soil layers, which suggests that not N, but C was a limiting factor to decomposition. In contrast, limestone showed low net N-mineralization in the organic layer, despite high mass and well-decomposed organic matter, and low efficiency of N-mineralization per unit microbe in both organic layer and mineral topsoil. DOC was net released instead of consumed, which supported that not C, but N was a limiting factor. The general lack of differences in net N-cycling between species, but relatively clear site effects, is discussed in relation to different microbial strategies. Acid soil may have high net N-release despite low biological activity, because N-requirements of fungi are also low, while in calcareous soil, high bacterial N-demand may counteract high gross N-release. Thus, species producing litter that decomposes rapidly may be planted to improve soil conditions and plant biodiversity, but litter quality effects on N-availability may be less important than soil conditions.     

Responses of a beech (Fagus crenata Blume) stand to late spring frost damage in Morioka,Japan

Following two weeks of a warm period, beech trees at the study site began leafing. However, a spring frost occurred in the cold morning of April 21, 2001 (Day111, Julian day) and killed about 80% of the fresh leaves on a 40-year-old Siebold's beech (Fagus crenata Blume) stand in Morioka, Japan. Leaves emerged on sprouts from latent buds (latent leaves) 36 days after the frost. Field measurements between 1999 and 2003 revealed the following: (1) Leaf area index was about 4 in normal years and about 1.2 in 2001. (2) The elongation of latent leaves took 55 days about 40 days longer, and they matured about 30 days faster than normal leaves in regular years. (3) Photosynthetic capacities were similar between the latent leaves in 2001 and the normal leaves in 2002 at the top of the canopy. A longer leafing period, faster maturation and sustained photosynthetic capacity of the latent leaves would be helpful to reduce the impacts of frost damage on growth. (4) The frost damage greatly reduced NPP from about 2100 DW g m⁻² y⁻¹ in the previous two years to about 1000 DW g m⁻² y⁻¹ in 2001. However, NPP recovered almost completely (about 1800 DW g m⁻² y⁻¹) in 2002. (5) Seasonal NPP increased similarly in 2001 with other years before Day150 for 40 days after the frost, and decreased afterwards. Shoot elongation also stopped at approximately Day150 in normal years. These findings suggested that radial and height growth in the early growing season was supported by the storage of previous year photosynthesis. (6) A regression growth model was created using meteorological data from 1999 and 2000, and the model was applied to data from 2001 to estimate the potential growth under normal conditions. The ratio of measured and estimated NPP changed seasonally, greater than 0.4 between Day150 and Day210, but below 0.25 between Day210 and Day250. This suggests that beech allocated more photosynthetic products for the next year's growth rather than the current growth in the latter growing season, thereby achieving the same initial growth with other years and amazing recovery in 2002.     

Impact of defoliation caused by the sawfly Cephalcia lariciphila (Hymenoptera: Pamphilidae) on radial growth of larch (Larix decidua Mill.)

The period between 2000 and 2002 repeated outbreaks of the web spinning sawfly Cephalcia lariciphila (Wachtl 1898) around the village of Větrny Jeníkov, Czech Republic. The impact of defoliation, caused by C. lariciphila feeding, on tree-ring formation of European larch (Larix decidua Mill.) was studied using dendrochronological methods. Heavy defoliation resulted in much less growth in the years of insect attack, the average incremental loss being 67% for the 2000–2002 period. Also, defoliation resulted in the formation of latewood with fewer cells and reduced cell-wall thickness.