Rapid climate change-related growth decline at the southern range edge of Fagus sylvatica

Studies on Fagus sylvatica show that growth in populations toward the southern limit of this species' distribution is limited strongly by drought. Warming temperatures in the Mediterranean region are expected to exacerbate drought where they are not accompanied by increases in precipitation. We studied levels of annual growth in mature F. sylvatica trees over the last half‐century in the Montseny Mountains in Catalonia (northeast Spain). Our results show significantly lower growth of mature trees at the lower limit of this species' distribution when compared with trees at higher altitudes. Growth at the lower Fagus limit is characterized by a rapid recent decline starting in approximately 1975. By 2003, growth of mature trees had fallen by 49% when compared with predecline levels. This is not an age‐related phenomenon, nor is it seen in comparable populations at higher altitudes. Analysis of climate‐growth relationships suggests that the observed decline in growth is a result of warming temperatures and that, as precipitation in the region has not increased, precipitation is now insufficient to ameliorate the negative effects of increased temperatures on tree growth. As the climate‐response of the studied forest is comparable with that of F. sylvatica forests in other southern European regions, it is possible that this growth decline is a more widespread phenomenon. Warming temperatures may lead to a rapid decline in the growth of range‐edge populations and a consequent retreat of the species distribution in southern Europe. Assessment of long‐term growth trends across the southern range edge of F. sylvatica therefore merits further attention.     

Growth of adult Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) under free-air ozone fumigation

This study attempted to detect the impact of ozone on adult trees of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) in an experimental mixed stand in Southern Bavaria, Germany. The aim was to examine whether there is a decrease in growth when trees are exposed to higher than atmospheric concentrations of ozone. This exposure was put into effect using a free-air fumigation system at tree crown level. Growth analysis was carried out on a group of 47 spruce and 36 beech trees, where radial stem increment at breast height - a sensitive index for stress - was measured. The ozone monitoring system allowed values to be obtained for the accumulated ozone exposure (SUM00) of each individual tree, so that their radial increment over three years could be correlated with the corresponding ozone exposure for the same time period. Correlation and regression analysis were then carried out to test the influence of ozone on diameter increment. In both spruce and beech, the initial stem diameter was the most influential factor on radial increment in the following year. A linear model was applied, including the diameter of the preceding year and the ozone exposure of the current year as predicting factors. For spruce trees, a significant negative influence of ozone exposure was found. In contrast, no significant ozone effect on diameter increment of beech was detected. The effect of ozone stress on a large spruce tree can lead to a decrease in potential radial increment of 22 %. The results are discussed in relation to other stress factors such as drought and lack of light.     

Bioclimatology of beech (Fagus sylvatica L.) in the Eastern Alps: spatial and altitudinal climatic signals identified through a tree-ring network

Aim To identify the dominant spatial patterns of Fagus sylvatica radial growth in the Eastern Alps, and to understand their relationships to climate variation and bioclimatic gradients. Location Fourteen beech stands in the Eastern Alps, growing between 200 and 1500 m a.s.l. in Italy, Slovenia and Austria. Methods At each site, trees were sampled using increment borers or by taking discs from felled trees. Cores and discs were processed by measuring and crossdating ring width. Ring width series were standardized, averaged, and prewhitened to obtain site chronologies. Hierarchical Cluster Analysis (HCA) and Principal Components Analysis of prewhitened site chronologies were used to identify spatial and altitudinal growth patterns, related to the bioclimatic position of each stand. Bootstrap correlation and response functions were computed between monthly climatic variables and either principal component scores or composite chronologies from stands associated by HCA. The stability of dendroclimatic signals was analyzed by moving correlation functions (MCF). Correlation analysis (teleconnections) based on a data base of 37 Italian and Slovenian beech tree‐ring chronologies revealed the spatial extent of principal component scores. Results Sampled trees were 200–400 years old, representing the oldest beech trees that have been crossdated for the Alps to date. Maximum age was directly related to altitude and to the presence of historical forms of conservation. Tree‐ring parameters varied according to geographic patterns and the age of sampled trees. Stands were bioclimatically organized according to their location, and with reference to their elevation and distance from the Adriatic Sea. A direct response to winter temperature was found at all elevations. The altitudinal gradient ranged from low‐elevation stands, characterized by a Mediterranean‐type, late spring–summer drought signal, to mountain and high‐elevation stands, characterized by a direct response to growing season temperature plus an inverse response to the previous year’s July temperature. The mountain and high‐elevation signal was evident in Austria, the Central Alps and Slovenia, while the low‐elevation signal was confined to mountains adjacent to the Adriatic Sea. MCF revealed trends in the response to climatic factors affecting tree‐ring formation in mountain and high‐mountain stands linked to climatic warming. Main conclusions Dendroclimatic networks can be used for bioclimatic studies of tree populations. A biogeographical separation emerged between the Alps and the Apennines at the upper elevations, while different degrees of mediterraneity distinguished sites at lower elevations. This information will be useful in assessing any future climate‐related bioclimatic shifts, especially for forests at ecotones and along altitudinal gradients.     

Growth responses to climate in a tree-ring network of European beech (Fagus sylvatica L.) from the eastern limit of its natural distribution area

The first dendrochronological network of European beech (Fagus sylvatica L.) from eastern limit of its distribution was established covering the species’ altitudinal range (210–1160 m a.s.l.) and three eco-regions. Statistical analysis revealed spatial trends among the 14 chronologies, with increasing tree-ring variability and synchronicity towards the east, particularly expressed by rBAR (mean interseries correlation). By means of HCA (Hierarchical Cluster Analysis) and PCA (Principal Component Analysis), the chronologies were divided into three main groups: a Mountain, Plateau and Eastern class. These correspond to the Alpine South, Continental and Pannonian bioclimatological zones. Dendroclimatological analysis revealed a general climate response pattern, with summer temperatures, April–May precipitation and April–June drought being the main limiting factors in the study area. The intensity of the correlations increases towards the eastern distribution limits and decreases towards the upper treeline of European beech. Similarly, the length of the climate-sensitive period changes, with the precipitation window increasing from April in the Mountains region to March–May in the East region and the scPDSI (self-calibrated Palmer drought severity index) window increasing from April–June in the Mountains to the previous July to the current September in the East. The western and uppermost site shows signs of being limited by low July temperatures. This chronology network represents a sensitive monitoring instrument to quantify the impact and adaptability of European beech to actual and future climatic change.     

Carbon,nitrogen, phosphorus,and sulphur concentration and partitioning in beech ecotypes (Fagus sylvatica L.): phosphorus most affected by drought

Beech seedlings of different drought sensitivity originating from 11 German provenances were grown in pots and cultivated in a greenhouse. The present paper aims to give insights on uptake, transport and use of macronutrients, since the knowledge of drought effects on the nutrition of trees is low compared to water relations. Therefore, the elemental composition, the ratio of inorganic to total content, and the partitioning between roots and shoots of carbon, nitrogen, phosphorus, and sulphur were investigated as affected by provenance and drought treatment. Phosphorus and phosphate concentrations decreased in all tissues after a 3 week drought treatment simulating a summer drought period. In roots carbon increased and nitrate decreased, in stems nitrogen decreased but nitrate increased following drought. The observed effects on phosphorus and phosphate are discussed in terms of lower phosphate mobility in the substrate due to lower water availability. The decrease in the ratio of phosphate to phosphorus in the tissues suggests the use of vacuolar phosphate pools for maintaining organic phosphorus homeostasis. The partitioning of all macronutrients was not affected by drought, although phosphorus and phosphate were significantly lowered in tissues. In most of the parameters studied significant differences between provenances were found. The recently observed drought sensitivity of provenances was not reflected in the strength of concentration changes or partitioning of macronutrients by drought over provenances.     

A unigene set for European beech (Fagus sylvatica L.) and its use to decipher the molecular mechanisms involved in dormancy regulation

Systematic sequencing is the method of choice for generating genomic resources for molecular marker development and candidate gene identification in nonmodel species. We generated 47 357 Sanger ESTs and 2.2M Roche‐454 reads from five cDNA libraries for European beech (Fagus sylvatica L.). This tree species of high ecological and economic value in Europe is among the most representative trees of deciduous broadleaf forests. The sequences generated were assembled into 21 057 contigs with MIRA software. Functional annotations were obtained for 85% of these contigs, from the proteomes of four plant species, Swissprot accessions and the Gene Ontology database. We were able to identify 28 079 in silico SNPs for future marker development. Moreover, RNAseq and qPCR approaches identified genes and gene networks regulated differentially between two critical phenological stages preceding vegetative bud burst (the quiescent and swelling buds stages). According to climatic model‐based projection, some European beech populations may be endangered, particularly at the southern and eastern edges of the European distribution range, which are strongly affected by current climate change. This first genomic resource for the genus Fagus should facilitate the identification of key genes for beech adaptation and management strategies for preserving beech adaptability.     

Genetic variation of European beech (Fagus sylvatica L.) along an altitudinal transect at Mount Vogelsberg in Hesse, Germany

Allelic and genotypic variation at 13 different enzyme loci of autochthonous European beech (Fagus sylvatica L.) was investigated in six 110-160-year-old stands growing at elevations between 150 and 660 m above sea level on the western slope of mount Vogelsberg in central Germany. The highest elevated population showed the highest number of effective alleles (Ne), the highest total heterozygosity (He) and the highest population differentiation deltaT. Also, the genotype SKD-A2A3 of shikimate dehydrogenase was significantly more frequent at the two highest elevated stands (P = 11%) than at the three lowest elevated stands (P = 1%). Further differences in genotype frequencies between 11 of 15 stand pairs were elevation independent.     

Carbon and nitrogen dynamics in acid detergent fibre lignins of beech (Fagus sylvatica L.) during the growth phase

To study the incorporation of carbon and nitrogen in different plant fractions, 3‐year‐old‐beech (Fagus sylvatica L.) seedlings were exposed in microcosms to a dual‐labelling experiment employing ¹³C and ¹⁵N throughout one season. Leaves, stems, coarse and fine roots were harvested 6, 12 and 18 weeks after bud break (June to September) and used to isolate acid‐detergent fibre lignins (ADF lignin) for the determination of carbon and nitrogen and their isotope ratios. Lignin concentrations were also determined with the thioglycolic acid method. The highest lignin concentrations were found in fine roots. ADF lignins of all tissues analysed, especially those of leaves, also contained significant concentrations of nitrogen. This suggests that lignin‐bound proteins constitute an important cell wall fraction and shows that the ADF method is not suitable to determine genuine lignin. ADF lignin should be re‐named as ligno‐protein fraction. Whole‐leaf biomass was composed of 50 to 70% newly assimilated carbon and about 7% newly assimilated nitrogen; net changes in the isotope ratios were not observed during the experimental period. In the other tissues analysed, the fraction of new carbon and nitrogen was initially low and increased significantly during the time‐course of the experiment, whereas the total tissue concentrations of carbon remained almost unaffected and nitrogen declined. At the end of the experiment, the whole‐tissue biomass and ADF lignins of fine roots contained about 65 and 50% new carbon and about 50 and 40% new nitrogen, respectively. These results indicate that significant metabolic activity was related to the formation of structural biopolymers after leaf growth, especially below‐ground and that this activity also led to a substantial binding of nitrogen to structural compounds.     

Physiological performance of beech (Fagus sylvatica L.) at its southeastern distribution limit in Europe: seasonal changes in nitrogen, carbon and water balance

To assess the physiological performance of drought-sensitive European beech ( Fagus sylvatica L.) under the dry Mediterranean climate prevailing at its southeastern distribution limit in Europe, we analyzed seasonal changes in carbon, nitrogen and water balance of naturally grown adult trees. We determined the foliar C and N contents, delta13C and delta18O signatures, total soluble non-protein nitrogen compounds (TSNN) in xylem, leaves, and phloem, as well as leaf water potential and photosynthetic quantum yield in northern Greece during 2003. Tissue sampling was performed in May, July, and September, while field measurements were conducted regularly. Climatic conditions for the 2003 growing season fall within the typical range of the studied area. The N- and C-related parameters displayed distinct seasonal courses. TSNN was highest in May in all tissues, and asparagine (Asn) was then the most abundant compound. Thereafter, TSNN decreased significantly in all tissues and both its concentration and composition remained constant in July and September. In both months, glutamate (Glu) prevailed in leaves, gamma-aminobutyric acid (GABA) in phloem exudates from twigs and trunks, and arginine (Arg) in the xylem sap, where loading with amino acids was rather low during that period, amounting to only 0.8 micromol N ml-1 in September. Highest total foliar N and C contents were detected in May, and the elevated abundance of nutrients as well as an increased foliar delta13C signature at the beginning of the growing season is attributed to remobilization processes. The signatures of delta18O, quantum yield and leaf water potentials varied only slightly throughout the growing season. Although summer precipitation at the study site was considerably lower compared to what is usual for typical central European beech forests, no intensive drought responses of the physiological apparatus were detected in the studied beech trees. This suggests efficient internal regulation mechanisms, constantly ensuring a favourable physiological status under the relatively dry Mediterranean climate.     

Winter North Atlantic oscillation effects on the tree rings of the Italian beech (Fagus sylvatica L.)

Climatic signals in beech tree-ring width series from Central Italy have been studied over different periods of time. Prewhitened tree-ring chronologies respond mainly to summer precipitation and they do not correlate in a significant manner with the winter North Atlantic oscillation (NAO) index. In this high-frequency pattern the NAO signs are only found on a small number of rings characterized by being very narrow or wide. By contrast, tree-ring width chronologies in which all the frequency components are conserved were significantly related to the NAO. The significant inverse correlation between actual measurements of ring width and NAO is a consequence of the availability of water in the soil at the beginning of the growing season. In fact, in the Mediterranean area the recharging of soil moisture depends on the amount of winter precipitation, which is inversely correlated with the NAO. Strong signals of winter precipitation and NAO are found in the low-frequency components of tree-ring growth. Received: 18 March 1999 / Revised: 29 February 2000 / Accepted: 1 March 2000     

Antioxidants in the apoplast and symplast of beech (Fagus sylvatica L.) leaves: seasonal variations and responses to changing ozone concentrations in air

Concentrations of the antioxidants ascorbate and glutathione were measured in the apoplast of beech (Fagus sylvatica L.) leaves and in leaf tissue. During early leaf development, reduced ascorbate (ASC) was almost absent from the apoplast, whereas levels of oxidized ascorbate (DHA) were high. Less than 20% of the apoplastic ascorbate was reduced. ASC increased towards midsummer, reaching top levels of about 4molm^?3 apoplast volume in July and August. Reduction increased to 60–75% in summer. Neither DHA reductase nor glutathione was detected in the apoplast of beech leaves. Levels of apoplastic ascorbate were compared with ambient concentrations of ozone in air. Statistical analysis indicated a significant interrelation between atmospheric ozone and apoplastic ascorbate. In midsummer of 1993, contents of DHA were increased in the apoplast when ozone concentrations were high. Apoplastic ASC was also positively correlated with ambient ozone concentrations, but with a delay of 3 to 7d. In leaf tissue, levels of ascorbate were between 17 and 21 μmolg^?1 FW in summer. Except for late April and November, more than 95% of the intracellular ascorbate was reduced. Glutathione contents were lowest during the summer. Oxidation was increased in spring and autumn, when apoplastic ascorbate was also largely oxidized. Usually, 80 to 90% of the glutathione was reduced. During the summer, intracellular concentrations of oxidized glutathione (GSSG) were increased, with a delay of about 1d following periods of high ambient ozone concentrations. The transitory accumulation of GSSG may be explained by slow enzymatic regeneration of glutathione.     

Crown allometry and growing space efficiency of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) in pure and mixed stands

In pure and mixed stands of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) we have analyzed crown allometry and growing space efficiency at the tree level and have scaled this from tree level to stand level production. Allometry is quantified by the ratio A between the relative growth rates of laterally and vertically oriented tree dimensions. Efficiency parameters, EOC for efficiency in space occupation, EEX for efficiency in space exploitation, and EBI for efficiency in biomass investment, were evaluated, based on quantity and quality of growing space and were measured using crown size and competition index. The evaluation reveals why pure stands of spruce are preferred by foresters, even though the natural vegetation would be dominated by beech. Spruce occupies its share of resources intensively by means of tightly packed pillar-like crowns, whereas beech seizes resources extensively by means of a multi-layered, veil-like canopy. With a given relative biomass increment, beech achieves a 57 % higher increment in crown projection area and a 127 % higher increment in height due to its particular capacity of lateral and vertical expansion. Beech trees are approximately 60 % more efficient in space occupation than spruce trees, however, on average, they are about 70 % less efficient in space exploitation. As a vertical fast growing tree, spruce is efficient in space exploitation under constant conditions, but far more susceptible to disturbances and less well equipped to overcome them when compared with beech. Beech is weaker in terms of space exploitation, while being superior in space occupation, where it encircles competitors and fills gaps after disturbances, which is a successful long-term strategy. A mixture of the two species reduces stand level production by 24 % in comparison to a pure spruce stand, however, when considering enhanced stabilization of the whole stand and risk distribution in the long term, the mixed stand may exceed the production level of pure spruce stands. EEX reflects a strong ontogenetic drift and competition effect that should be considered when scaling from tree to stand level production.     

Climate-tree-growth relationships of European beech (Fagus sylvatica L.) in the French Permanent Plot Network (RENECOFOR)

The influence of climate on the radial growth of Fagus sylvatica was investigated using 15 chronologies developed from mature stands of the French Permanent Plot Network (RENECOFOR) growing under different climatic and soil conditions. The relationships between climate and ring widths were analyzed using extreme growth years, simple correlations and response functions analysis. Monthly climatic regressors were derived by a physiological water balance model that used daily climatic data and stand parameters to estimate soil water deficits. The three most frequent negative pointer years (1959, 1989, 1976) result from a particularly intense and durable drought, whereas positive years (1977, 1958) coincide with wet conditions. The total ring chronology variance attributable to climate averages 34.1% (15.8% –57%). Current early-summer soil water deficit enters in 10 models and the deficit in June explains alone a large part of the radial growth variability (mean value: 26.6%). Temperature or soil water deficit for the other months and weather conditions during the previous season were of little consistency across stands. The response pattern of earlywood is very similar and the percentage of variance explained is higher (16.2% –57.8%). Latewood widths present a different response pattern. High minimum temperature in August and/or September often favour wide latewood widths and monthly water deficits play a secondary role. The percentage of variance explained ranges from 8.8% to 67.4%. Soil water capacity strongly modulates ring characteristics and climate-growth relationships. Mean sensitivity, expressed population signal, signal-to-noise ratio and the strength of growth-climate correlations increase with decreasing soil water capacity.     

Is the spatial distribution of European beech (Fagus sylvatica L.) limited by its potential height growth?

Aim To improve our understanding of species range limits by studying how height growth, a trait related to plant survival, varies throughout the geographic range of Fagus sylvatica L. in France. Location The geographic range of beech in France, representing the western area of its European distribution, within which this species exhibits range distribution limits in both plains and mountainous areas. Methods A generalized linear regression model was used to link beech growth performance to environmental variables using data from 819 plots of the French National Forest Inventory (IFN) database. This model was applied to predict potential growth on 97,281 IFN plots covering the geographic range of beech in France. A kriging technique was used to interpolate estimated growth potential. Finally, the performance of plot‐based predictions of potential growth from the map (i.e. map quality) was evaluated against an independent data set. Results The beech growth performance model highlighted the major impact of climate on potential tree growth at a broad spatial scale. The relevant climatic factors were related mainly to spring cold, summer heat, and winter temperatures and rainfall. The study also revealed the predictive power of soil parameters, which explained a large proportion of the variation in potential beech growth (c. 30%). Analyses of height growth patterns near the boundary of the species range in France showed that the limit only partly coincides with the growth decline caused by climatic and soil factors. Along parts of the range limit, the predicted potential for growth was high, suggesting that in these areas the limit of the range could be explained by other factors, such as competition or constraints on reproduction. Main conclusions The spatial variation in the potential height growth of Fagus sylvatica can be explained by environmental factors and is partly correlated with its regional range limits. By identifying areas where growth potential constrains the geographic range of species, environmental growth models can help to improve our knowledge of the spatial drivers of species geographic range limits and shed light on their response to future environmental changes.     

Twentieth century changes of tree-ring δ13C at the southern range-edge of Fagus sylvatica: increasing water-use efficiency does not avoid the growth decline induced by warming at low altitudes

We aimed to gain knowledge on the changes in intrinsic water use efficiency (iWUE) in response to increasing atmospheric CO₂ concentrations and climate change over the last century. We investigated the variation in the iWUE of mature Fagus sylvatica trees located in the higher, central and lower altitudinal forest limits (HFL, CFA and LFL) of one of the southernmost sites of beech distribution in Europe, the Montseny Mountains in Catalonia (northeast Spain), during the last century by analysing the δ¹³C of their tree rings. Pre‐ and post‐maturation phases of the trees presented different trends in δ¹³C, Δ¹³C, Ci (internal CO₂ concentration), iWUE and basal area increment (BAI). Moreover, these variables showed different trends and absolute values in the LFL than in the other altitudinal sites, CFA and HFL. Our results show the existence of an age effect on δ¹³C in the CFA and HFL (values increased by ca. 1.25‰ coinciding with the BAI suppression and release phases, previous to maturation). These age‐related changes were not found in the LFL, whose beech trees arrived to maturation earlier and experienced drier conditions during the suppression phase. In the last 26 years of comparable mature trees, the increase of iWUE deduced from the Δ¹³C analyses was ca. 10% in LFL, ca. 6% in CFA and not significant in HFL. These results show that climate change towards more arid conditions accounted for these higher Δ¹³C‐values and increases in the LFL more than the continuous increase in atmospheric CO₂ concentrations. This increased iWUE in the LFL did not avoid a decline in growth in these lowest altitudes of this beech southern range‐edge as a result of warming. Furthermore, since there was no apparent change in iWUE and growth in the beech forests growing in the more standard‐adequate environments of higher altitudes in the last 26 years, the rate of sequestration of C into temperate ecosystems may not increase with increasing atmospheric CO₂ concentrations as predicted by most models based on short‐term small scale experiments.     

Critical forest age thresholds for the diversity of lichens,molluscs and birds in beech (Fagus sylvatica L.) dominated forests

Forest age is one of the most simple but ecologically effective key values that may be controlled by forest management. Young and mature but managed forests differ significantly from old-growth forests in species composition, structure and socio-ecological function. Human land-use has already caused the loss or dramatic reduction in occurrence of some entire species assemblages, especially of logging-sensitive species, in Central European forests. These general statements also apply to beech forests, beech (Fagus sylvatica) being the naturally dominating tree species in Central Europe. Based on data for breeding birds (from 258 sampling plots in a sub-montane and 228 plots in a montane area), molluscs (36 plots in the sub-montane and 79 plots in the montane area) and lichens (84 plots in the montane forest), this paper aims at identifying significant forest age threshold ranges for the occurrence of these old-growth sensitive taxa. The sampling plots in the sub-montane zone (420–520 m a.s.l.) are in beech-oak forests, plots in the montane zone (650–1150 m a.s.l.) are in beech-spruce-fir forests. Stand ages in both areas range up to around 350–400 years. Threshold values for the total number of species related to stand age were calculated by recursive partitioning.In all three taxonomic groups the number of species per plot significantly increases with forest age. The same analysis was run for red-listed lichen and mollusc species as well as hole-nesting bird species. The threshold values obtained are very similar to those for the whole species assemblages, except for molluscs where considerably lower threshold values are computed with red-listed species assemblages. Regarding the confidence intervals, the difference pattern between the whole species datasets and the more sensitive species subsets is inconsistent. Threshold values in sub-montane beech forests range from 100 to 170 years and in mixed montane forests from 160 to 220 years.These threshold levels are clearly incompatible with economic interests that aim on reducing the rotation period in beech stands to less than 140 years to avoid formation of red heartwood. It would therefore seem to be essential to establish a network of trees and stands that are never logged and may thus act as areas for retreat and dispersion for logging-sensitive species.     

Structure and dynamics of Japanese beech (Fagus japonica Maxim.) stools and sprouts in the regeneration of the natural forests

Structure and spatial distribution of stools and root-collar sprouts of Japanese beech (Fagus japonica) were studied to clarify the regeneration processes of the stool and the population, and the ecological importance of this stool formation in five quadrats of the natural forests with different forest floor vegetation on the Pacific side of Japan. F. japonica dominates in the canopy of each quadrat. Most of sprouts formed a circle around the root-collar and lowest parts of the parent stems of the stool with the youngest sprouts at the periphery. The regeneration by seedlings was slight especially on the forest floor vegetation of the dwarf bamboo Sasa. The variety of size structure of stems and the existence of dead traces and/or dead center in each stool, the apparent difference in stool size, and the aggregations of stools in the forests suggest that stool expansion and long persistence of the stool at a given location may contribute to compensate for the scarcity of regeneration by seedlings inhibited by dwarf bamboo, and by the other shrubs and herbs.