Masting behaviour and dendrochronology of European beech (Fagus sylvatica L.) in southern Sweden

To identify weather controls of beech diameter growth and masting in southern Sweden, we analyze records of monthly weather, regional masting record, and tree-ring chronologies from five beech-dominated stands. The results indicate a strong weather control of temporal pattern of masting events in southern Sweden over the second half of the 20th century. Negative summer temperature anomaly 2 years prior to a mast year, coupled with positive temperature anomaly in the year immediately preceding the same mast year, is a characteristic weather pattern associated with known mast years. Strong dependence of beech masting behaviour on temperature explains the high degree of regional synchronization of masting events. Growth of beech in southern Sweden is strongly and negatively correlated with previous year's summer temperature and positively – with previous year's October temperature. The present study does not provide a conclusive answer in identifying a full set of direct and indirect effects of climatic variables controlling tree-ring growth, since the negative effect of previous year's summer temperature may be a result of a temperature-controlled increase in the beech nut production in the current year. Consistent and significant negative departures of ring-width index during mast years support the hypothesis about a trade-off between investment of bioassimilates into production of beech nuts and tree-ring growth. Alternative explanation of growth anomalies in mast years, relating such anomaly to a negative impact of previous year's growing season, was not supported by the data. We found a limited effect of masting on diameter growth in the following years, indicating that decline in the overall wood production, associated with heavy masting, is short term and typically occurs in the year of actual masting.     

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.     

On the radial-growth variations of Japanese beech <Emphasis Type="Italic">(Fagus crenata)</Emphasis> on the northernmost part of Honshu Island,Japan

We developed ring-width chronologies for living trees of Japanese beech at two forest sites on the northern-most part of Honshu Island, Japan. A statistical threshold (running expressed population signal) yielded these site chronologies spanning 1853–1994 (142 years) and 1867–1994 (128 years). We examined two factors, climate and masting, that could affect the variations of radial growth. The response function analysis revealed that the ring width correlated positively with July and August temperatures of the previous growth year. The optimal radial growth of Japanese beech may largely depend on a warm previous summer with above-average temperatures. The years of good masting coincided mostly with those showing abrupt growth depression, although only the short-term records of masting were available. Part of this work was presented at the 6th Pacific Regional Wood Anatomy Conference, Kyoto, December 2005, and the 7th International Conference on Dendrochronology, Beijing, June 2006     

Tree Seed Production and Food Conditions for Rodents in an Oak Wood in Southern England

Seed production and seed losses were monitored in an oak (Quercusrobur) wood in southern England between 1975 and 1988. In additionto acorns, seeds from several large beech (Fagus sylvatica)and sweet chestnut (Castanea sativa) trees which occurred withinthe wood, and from the understorey of hazel (Corylus avellana)trees, were also monitored. The seeds from these four tree speciesare all subject to heavy predation by rodents, as well as otherseed-eaters such as birds and insects. Seed fall tended to be irregular and three years, 1976, 1985and 1987 were considered oak mast years. There were too fewyears with large acorn crops to examine whether resource depletionin these years resulted in poor crops in subsequent years. Therewas no correlation in seed production between the four treespecies which does not agree with one of the predictions ofthe predator satiation theory of masting behaviour. The effectsof weather on seed production were investigated but few significantcorrelations were found. In general the results tended to supportprevious findings. Between 1980 and 1982 a serious outbreakof Tortrix moth caterpillars occurred within the wood and mayhave affected acorn production. A negative relationship wasfound between oak defoliation and acorn production in the subsequentyear. The loss of seeds from the forest floor was initially rapidand in 9 years no seeds survived into the following year andfood conditions were poor for granivores. In 3 of the remaining4 years acorns survived well into spring and early summer providinggood food conditions for forest rodents through to the timeswhen alternative food supplies became available. Although oaktrees contributed most seed energy in good years, the othertree species, especially beech, became important when acornproduction was poor. Predispersal losses due to predators werefrequently high and averaged 80 per cent, 40 per cent, 38 percent and 33 per cent in hazel, sweet chestnut, oak and beechrespectively. On average, hazel nuts tended to fall first, some6 weeks before acorns which tended to be the last seeds to fallof the four tree species. Knopper galls and weeviled acornsfell earlier than sound acorns. A key-factor analysis indicatedthat a failure to mature, predispersal predation and insectinfestation from weevils and Knopper galls contributed equallyto changes in numbers of acorns among years.     

A twig blight of understorey European beech (Fagus sylvatica) caused by soilborne Phytophthora spp.

During and after prolonged periods of rainfall in late spring, blighted young twigs of European beech (Fagus sylvatica) were frequently observed in several beech stands in south‐western and southern Germany. Long and short shoots of young understorey trees or lower branches up to 1.5 m above the soil level were affected. Symptoms also occurred regularly on twigs in heights up to 2 m and more above the ground. Necroses usually expanded within the current year’s tissue and often also reached into the previous year’s wood. Ponding rain water in the stands or along forest roads or open soil seemed to promote the disease. Of a total of 54 symptomatic twigs collected in four stands, 37 revealed Phytophthora isolates, of which 33 were P. plurivora and four were P. cambivora. Both species caused extensive lesions on beech twigs in laboratory pathogenicity tests. Patterns of the disease indicated that these pathogens, generally considered soilborne species, in most cases are transmitted from the soil to above‐ground parts of the trees via rain splash. In larger heights, however, other vectors such as snails might be responsible for transmission. Although Phytophthora spp. are well known as causal agents of seedling blight as well as root and cambium rot and aerial bleeding cankers of mature beech, to our knowledge this is the first report of a twig blight in beech associated with soilborne Phytophthora spp. In particular in periods of high precipitation, this disease might pose an additional threat to Central European beech forests, especially endangering the success of artificial and natural regeneration of beech in affected stands.     

Masting in Fagus crenata and its influence on the nitrogen content and dry mass of winter buds

In Fagus, full-mast seeding years are invariably followed by at least one non-mast year. Both flower and leaf primordia develop during the summer within the same winter buds. Flower bud initiation occurs when the N content of developing seeds is increasing rapidly. We hypothesized that competition for nitrogen (N) between developing seeds and buds limits flower primordium formation in mast years and, hence, limits seed production in years following mast years. We tested this hypothesis in three Fagus crenata Blume forests at elevations of 550, 900 and 1500 m. Bud N concentration (N con), amount of N per bud (N bud) and dry mass per bud (DM) were compared between a mast year (2005) and the following non-mast year (2006), and between winter buds containing both leaf and flower primoridia (BF), which were formed during the non-mast year, and winter buds containing leaf primordia only (BL), which were formed in both mast and non-mast years. In addition, leaf numbers per shoot corresponding to the analyzed buds were counted, and the effect of masting on litter production was analyzed by quantifying the amounts of litter that fell in the years 2004 to 2007. The dry mass and N content of BF formed in 2006 by trees at both 550 and 1500 m were 2.1-3.4-fold higher than the corresponding amounts in BL, although the numbers of leaves per current-year shoot in 2007 that developed from the two bud types in the same individuals did not differ significantly. These results indicate that more N and carbohydrate are expended in producing BF than in producing BL. The amount of litter from reproductive organs produced in the mast year was similar to the amount of leaf litter at 900 and 1500 m, but three times as much at 550 m. Leaf numbers per shoot were significantly lower at all elevations in the mast year than in the non-mast years (and the amount of leaf litter at 550 and 1500 m tended to be lower in the mast year than in the non-mast years. In conclusion, preferential allocation of resources to seeds in the mast year reduced the availability of resources for flower primordium formation, and this may have accounted for the poor seed production in the following non-mast year.     

Influence of fertilizer timing on seasonal nutrient uptake and dry-matter production by young Pinus radiata in southern western Australia

Seasonal changes in above ground dry-matter, nitrogen (N), and phosphorus (P) accumulation were measured following application of N and P in autumn or spring to 1-year-old Pinus radiata (D. Don). Dry-matter production and nutrient accumulation were measured eight times over two years following fertilization.

All trees produced dry-matter throughout the year, but during the summer, fertilized trees produced more dry-matter than unfertilized trees. In contrast to dry-matter production, nutrient accumulation showed a distinctly seasonal pattern with maximum accumulation of N and P occurring in winter and spring, when rainfall and soil moisture were highest. Accumulation of N and P either slowed markedly or ceased during summer depending on fertilizer treatment. Continued dry-matter production during summer, when nutrient accumulation was low, resulted in the decline of N and P concentrations in needles, branches and stems of all trees. This indicated that nutrients required for new growth during summer were mobilized from existing foliage and wood. Fertilization increased the concentrations of N and P in foliage and wood, and these higher concentrations persisted through summer. Spring fertilization increased N accumulation to a greater extent than autumn fertilization, this effect lasting two years. The greater dry-matter production by fertilized trees during summer indicated that growth during summer was limited by nutrient supply.     

Observations on the Condition of Beech (Fagus sylvatica L.) in Britain in 1990

In 1990, the Forestry Commission's main monitoring programmeassessed the condition of 864 beech trees across 36 sites inBritain. Sites were located throughout the country, with thenorthernmost site being near Inverness and the southernmostnear Exeter. The initial crown density data suggested that asmall deterioration had occurred. However, further investigationrevealed that the spatial pattern of crown density changed in1990, with trees in southern sites deteriorating markedly. Otherindices of crown condition also changed, with the amount ofdieback, the amount of mast, the incidence of small leaves andthe amount of leaf-rolling all being high in the south of thecountry. In addition, the radial growth of many trees was depressed.Most of the changes in 1990 are thought to have been causedprimarily by the 1989 drought, with the 1990 drought only havingan effect on parameters such as leaf-rolling. Further effectsin 1991 and possibly later years are anticipated.     

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

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

Influence of tree shelters,irrigation and branch pruning on early field performance of micropropagated wild cherry cv. F12/1

The growth potential of micropropagated wild cherry cv. F12/1 under field conditions was assessed. Relatively short trees (19–22 cm tall) were planted in the field just seven months after transfer from tissue culture to soil. Irrigation and shelters promoted shoot heights in the first (establishment) year. If planting techniques can be developed which reduce branch production, then the cost of corrective pruning to achieve clean stems is also reduced. Shelters, but not irrigation almost totally inhibited outgrowth of branches produced in spring. Outgrowth of spring branches by unsheltered trees was strongly influenced by tree height at planting, with shorter trees producing fewer branches. In the second and third years of the trial, pruning branches from trees that had been planted in shelters, resulted in taller trees. By the end of the third year, pruned trees had smaller stem diameters than unpruned trees. These results are discussed with respect to using cv. F12/1 as a productive timber tree.     

利用杨树人工林剩余物生产刨花板可行性研究

The composite board industry in Iran is obliged to use residues from forest operation as well as wood industry for competing with paper industry because of shortage of wood. In the present study we investigated the residues from poplar plantation used for particleboard production. Three kinds of wood materials, poplar branches, small diameter poplar wood (3–8 cm) and beech wood, were used in the experiment of particleboard production. The results demonstrated that the characteristic of particleboard made from poplar branches and small diameter wood is comparable to that made from mature beech wood. To avoid too much residual acid in the final board, the properties of boards produced with 1.5% hardener at 175°C press temperature are acceptable, although the properties of particleboard produced with 2% hardener were higher than were higher than that of the board produced with lower hardener (1% or 1.5%).. The MOR, MOE and IB of particleboard made from branches were measured as 14.57, 2015, and 1.32 MPa, respectively, while The MOR, MOE and IB of particleboard produced from small diameter poplar wood were 19.90, 2199, and 1.86 MPa, respectively. The thickness swelling of boards made from branches after 2 and 24 h immersion in water was 20.14% and 31.26%. The utilization of branches and very small diameter wood of poplar is recommended for the survival and developments of particleboard industry in Iran.     

Induced response of the Siebold's beech (Fagus crenata Blume) to manual defoliation

Siebold's beech (Fagus crenata) was manually defoliated for two successive years. The beech caterpillar (Quadricalcarifera punctatella) was used in a bioassay to determine insect performance. Survival and body size were low on foliage from defoliated trees. Reduced foliar nitrogen and increased tannin content were probably the main causes of the low insect performance. Leaves were less tough on defoliated trees than in controls. Two sucessive years of manual defoliation caused stronger induced resistance than one year defoliation. The quality, as well as the quality of the foliage, decreased the year following manual defoliation; total weight of leaves on a tree was less than one half of that before treatment. Severe defoliation may cause a decrease of leaves the following year and starvation may limit populations. Delayed induced resistance of beech trees is proposed as a possible cause of the cyclical population dynamics ofQ. punctatella. The delayed induced response also affected folivorous insects other thanQ. punctatella.     

泡核桃老龄低产树更新复壮技术

泡核桃树龄达到70~80年以后逐渐进入老龄期,树势开始衰弱,大、中枝条逐渐枯死,结果量逐年减少。通过多年研究,摸索出分3年逐渐更新根系、主枝和侧枝,并加强土肥管理及病虫害防治等综合更新复壮技术,第4年树势明显恢复并大量开花结实,获得了可观的经济收益,从而为泡核桃老产区巩固产业持续发展、提高泡核桃产量和质量提供了技术支撑。     

Interactions between climate,growth and seed production in Spanish black pine (<Emphasis Type="Italic">Pinus nigra</Emphasis> Arn. ssp. <Emphasis Type="Italic">salzmannii</Emphasis>) forests in Cuenca Mountains (Spain)

Tree growth is regulated by a combination of exogenous and endogenous factors. Such factors also interact with each other, complicating the understanding of causal links. IN particular, resource allocation is sensitive to reproductive investment, especially in masting species, which in turn is regulated by climatic variables. Both resource allocation and seed production patterns are also sensitive to tree age. This study aims to (1) evaluate the effects of tree age and local and regional climate on tree ring width and seed production by Spanish black pine (Pinus nigra Arn. ssp. salzmannii) forest in Cuenca Mountains (Spain), and (2) assess the relationship between seed production and secondary growth of Spanish black pine. Seed fall was estimated using 60 rectangular seed traps (40 × 50 × 15 cm) from 2000 to 2014, randomly distributed across the study area. Standardized tree-ring chronologies were calculated using a random sample of 106 trees stratified into three age classes (> 80 years; 26–80 years, and ≤ 25 years). Local climate data was obtained from a meteorological station, and regional climate data from the CRU-TS 3.1 dataset. Average seed production ranged over time from 2 to 189 seeds m^?2 (coefficient of variation = 157%). We identified four masting years (2000, 2003, 2006, and 2014) using a classification based on percentile seed production. Seed production was regulated by climate of the previous 2–3 years, while tree growth responded to precipitation and temperature in the previous and current year. Independent of climate, high seed production had a negative effect on tree ring width and weakened climate growth relationships, indicating resource depletion. Tree age modulated climate sensitivity, increasing correlations between climate and tree-ring index in older trees. P. nigra has been showed to be a climate sensitive species with a bimodal masting behaviour, which should be taking into account for management purposes and silvicultural guidelines under climate change scenarios.     

Feinwurzelmasse und Mykorrhiza von Altbuchen (Fagus sylvatica L.) in Waldschadensgebieten Bayerns

Feeder root biomass and mycorrhizae of old beech trees (Fagus sylvatica L.) in bavarian forest die-back areas . The feeder root biomass (roots = 2 mm diameter) and the quantity of mycorrhizae of apparently healthy and diseased old beech trees were determined in 7 stands in Bavaria. Canopy die-back correlated with reduction of the feeder root biomass and of mycorrhizae in the topsoil. In the deeper layers of the soil (below 50 cm) there were nearly no differences between the decline classes lor these parameters. The feeder root damage described were encountered in all stand types. For that reason, such damage can be considered, at least for comparable stands, as a general symptom of above ground diseased old beech trees.     

Genetic diversity and bark physiology of the European beech (Fagus sylvatica): a coevolutionary relationship with the beech scale (Cryptococcus fagisuga)

In 1994 and 1995, the degree of infestation by the beech scale (Cryptococcus fagisuga Lind.) was recorded on 120 beech trees (Fagus sylvatica L.) in the Pless Forest near G?ttingen, Germany. Simultaneously, the trees were characterized genetically and compounds of primary and secondary metabolism of beech bark were analyzed. A correlation was established between beech scale infestation and the genotype of the host trees, based on gene locus A of isocitrate dehydrogenase (IDH-A). The fraction of infested beech trees was higher in the heterozygous genotype A2A3 group than in the homozygous genotype A2A2 and A3A3 groups, whereas the fraction of beech trees with decreasing infestation from one year to the next was lower in the heterozygous genotype A2A3 group than in the homozygous genotype A2A2 and A3A3 groups. Concentrations of soluble carbohydrates, protein amino acids and proanthocyanidins were determined in the inner and outer bark of trees with differing degrees of infestation. The results indicate that the defense reaction of beech against infestation by beech scale comprises multiple processes in which nutrient availability to the beech scale is reduced by concentration shifts and by the formation of inhibiting compounds.     

Crown plasticity and neighborhood interactions of European beech (Fagus sylvatica L.) in an old-growth forest

Competition for canopy space is a process of major importance in forest dynamics. Although virgin and old-growth European beech (Fagus sylvatica L.) forests in Europe have been studied for many years, there are to date no studies of individual-tree crown plasticity and the way this is influenced by local neighborhood interactions in these forests. In this study, we analyzed crown plasticity and local neighborhood interactions of individual trees in the upper canopy of the old-growth beech forests of Serrahn, northeast Germany. In a 2.8-ha sample plot, we measured crown radii of all upper canopy trees and analyzed the direction and extent of crown asymmetry. Size, relative position, and distance of neighboring trees were used to construct vectors of neighborhood asymmetry within different distances from target trees. The crowns of beech trees showed strong morphological plasticity. Mean absolute and relative displacement of crown centers from the stem base were 1.95 m and 0.37, respectively. Circular–circular rank correlation coefficients between the direction of crown displacement and the direction of neighborhood pressure showed that trees strongly positioned their crowns away from local neighbors. Highest correlation coefficients were obtained when basal area and relative position of neighboring trees within a radial distance of 12 m were considered. Clark and Evans index and Ripley’s K-function showed that crowns were more regularly distributed than stems. Projected canopy cover was about 10% higher than canopy cover with simulated circular crowns. We conclude that the crowns of older beech trees have a high ability to plastically respond to changes in the local canopy conditions, enabling very effective exploitation of canopy space.     

Nutritional effects triggered by the extreme summer 2003 in the free air ozone fumigation experiment at the Kranzberger Forst

Since the year 2000 mature beech and spruce trees were treated in a field experiment with double ambient ozone concentrations. Elevated ozone had no influence on average single leaf biomass and there were also no ozone effects on leaf nutrient concentrations in climatic normal years. However, the extraordinary dry summer 2003 triggered significant differences between the fumigated and control trees. For beech in the year after the drought event the control trees surprisingly had significantly lower foliar levels of K and P than in former years, whereas the ozone exposed trees showed no significant nutritional effects. There are indications, that the trees exposed to double ambient ozone were already adapted to higher ozone values, whereas the control trees experienced extraordinary high ambient ozone concentrations in the dry and sunny summer 2003. For spruce in autumn 2003 and 2004 ozone treated trees had significantly higher foliar levels of K in current year needles than control trees, an effect which cannot be thoroughly interpreted yet on the basis of the dataset available. This article belongs to the special issue „Growth and defence of Norway spruce and European beech in pure and mixed stands“.     

Formation and growth of stem cankers caused by Gremmeniella abietina on young Pinus contorta

From 1990 to 1995, the formation and growth of stem cankers caused by Gremmeniella abietina on Pinus contorta var. latifolia was studied in three stands in northern Sweden. The stands were planted in 1976–80. The total number of cankers on 756 trees that were individually followed increased from 233 to 477 during the 5‐year period. With 42.0% of the cankers, the pathogen entered through or from the base of diseased branches, and 33.6% through visually undamaged bark. Most of the cankers were within 100 cm of the ground. In one of the three areas, the cankers were evenly distributed within 180 cm of the ground. The frequency of cankers facing north exceeded those facing south. The average vertical length of cankers had increased, 55.6% of cankers had increased their percentage coverage of the stem girth; 13.8% had fully girdled the stem. At two of the sites, there was a negative correlation between canker coverage of the stem circumference and tree height increment.     

Annual and geographic variation in seed traits of Argentinean populations of southern beech Nothofagus nervosa (Phil.) Dim. et Mil.

Nothofagus nervosa (=N. alpina) is one of the most important native forest tree species of Argentina, with a very small natural distribution area, fast growth and a high wood quality. Its overexploitation in the past seriously affected some populations and made others disappear, making it necessary to implement conservation policies and breeding programs.In order to study the genetic and geographic variation of the species, seeds from a total of 29 populations from the entire natural distribution area in Argentina were harvested, 11 of them during four consecutive years. Annual and geographic covariation of seed production, seed weight and proportion of seeds damaged by insects were analyzed.Significant differences among populations and among years for all studied traits were found. Seed production was highly variable, both among populations and among years. `Partial mast years' seem to occur every two years, whereas the occurrence of `full mast years' has a lower frequency. A significant population × year interaction was found both for the proportion of damaged seeds and for the seed weight. The proportion of seeds damaged by insects was very important, reaching >80% of the total seed produced in some populations. The differences in the quantity of attacked seeds among years are discussed in relation to seed production, insect's life cycle and climatic factors. Seed weight was also variable among populations and years, despite the small natural distribution area of the species in Argentina. Best seed quality, according to lower proportion of damaged seeds and higher seed weight, was observed in the analyzed `full mast year'. This would also imply a better genetic representation of populations with seed samples harvested during these years. A lower proportion of damaged seeds as well as the heaviest seeds were also found among those seeds fallen during the first part of the seed-fall period.The found covariation is discussed in relation to patterns of genetic variation previously found in this species and to the implementation of genetic studies and conservation and breeding programs.