Stand dynamics of the shade-tolerant conifers Podocarpus nubigena and Saxegothaea conspicua in Chilean temperate rain forest

Abstract. Population structures were used to infer regeneration patterns of the conifers Podocarpus nubigena and Saxegothaea conspicua and associated angiosperm tree species in six stands in the temperate rainforests of the coastal ranges of southern Chile. On poorly-drained, nutrient-poor and high altitude sites, where most of the associated species were fairly shade-intolerant and light-crowned, dense all-aged populations and the presence of numerous saplings beneath the canopy suggested continuous regeneration of both P. nubigena and S. conspicua. On more favourable sites, where several of the associated angiosperms were highly shade-tolerant and densecrowned, both conifers, particularly P. nubigena, were less abundant, and their regeneration from seed appeared to be sporadic. S. conspicua's great longevity (≥ 750 yr) and frequent vegetative reproduction are probably crucial to its persistence in competition with shade-tolerant broadleaved species in undisturbed stands on favourable sites. Results of this study, and a review of related literature on southern temperate forests are not consistent with the hypothesis that heavily-shaded, infrequently-disturbed habitats are an evolutionary refuge for conifers. Sites likely to have high leaf area indices and infrequent disturbance are better exploited by shade-tolerant angiosperms.     

Importance of the soil seed bank of disturbed sites in Chilean matorral in early secondary succession

Abstract. The dynamics of the seed bank may provide clues to the process of recovery of the vegetation of disturbed sites. The role of the seed bank may be more important in areas with a seasonal climate than in areas where seedling recruitment is not limited to one season. We studied the seed bank and the seed rain in three sites of the Chilean mediterranean-climate region (33° 48'S) which differed in the degree of anthropic disturbance: a closed-canopy, second-growth forest; an open matorral; and an old-field. Additionally, we tested the germination of seeds from the soil and from the current-year seed crop. The seed bank varied considerably between the two years of study, although no change in the vegetation could be observed. Seed density and species richness were lower in 1989 than in 1988. The seed bank of the second-growth forest changed less between years, while the old-field showed the largest change. The highest seed rain occurred under shrub patches in the open matorral, while few seeds fell in the spaces between shrub clumps or in the old-field. In the forest, seed rain was low and correlated with species cover. Germination was low (0 - 15%) in tests using either soil samples or fresh seeds. These results indicate that matorral succession is a very slow process, limited mainly by low germination and low arrival of propagules to open areas. Most woody species have animal-disseminated fleshy propagules. The presence of established shrubs which may serve as perches or refuges for animals increases species richness in the seed rain and the seed bank.     

Nurse effect of Bolax gummifera cushion plants in the alpine vegetation of the Chilean Patagonian Andes

Abstract. It has been proposed that in the harsh arctic and alpine climate zones, small microtopographic variations that can generate more benign conditions than in the surrounding environment could be perceived as safe sites for seedling recruitment. Cushion plants can modify wind pattern, temperature and water availability. Such modifications imply that cushion plants could act as ‘nurse plants’ facilitating the recruitment of other species in the community. This effect should be more evident under stressful conditions. We tested these hypotheses comparing the number of species that grow inside and outside Bolax gummifera cushions at two elevations (700 and 900 m a.s.l.) in the Patagonian Andes of Chile (50°S). At both elevations, and in equivalent areas, the number of species was registered within and outside cushions. A total of 36 and 27 plant species were recorded either within or outside B. gummifera cushions at 700 and 900 m a.s.l., respectively. At 700 m a.s.l., 33 species were recorded growing within cushions and 29 outside them, while at 900 m a.s.l. these numbers were 24 and 13 respectively. At both elevations there were significantly more species growing within than outside cushions, and the proportion of species growing within cushions increased with elevation. Thus there is a nurse effect of cushion plants and it is more evident at higher elevations. Shelter from wind and increased soil water availability seem to be the factors that increase plant recruitment within cushions.     

Variation in vegetation and seed bank in a Chilean semi‐arid community affected by ENSO 1997

Abstract. We followed plant cover and soil seed density of shrubs and herbs in two markedly contrasting years with regard to annual rainfall in two opposite‐facing slopes (mesic vs xeric) and an intervening, relatively wet, ravine of a typical creek in semi‐arid Chile. During the ENSO year 1997 cover of ephemerals increased in all three sites; 43% vs 8% on the xeric slope 75% vs 26% on the mesic slope and 57% vs 32% in the ravine. The number of species was almost three times higher on the xeric slope (27 vs 10 species), increased by 47% on the mesic slope (28 vs 19) and by 14% in the ravine (24 vs 21). Cover of shrubs plus perennial grasses increased from 52% in 1996 to 59% in 1997 on the xeric slope, but no significant cover changes were found on the mesic slope or the ravine. On the xeric slope peak density of seeds was 4500/m² in 1996, while during the ENSO year it was 24000/m². On the mesic slope equivalent values were 3000 and 17000 seeds/m² while in the ravine figures were 8500 and 27000 seeds/m². The strong responses of ephemerals and seed bank to the ENSO‐driven wet year of 1997 demonstrate the importance of this phenomenon in replenishing, recovering and probably maintaining ephemeral vegetation in this region.     

Position on slope,disturbance, and tree species coexistence in a Seasonal Semideciduous Forest in SE Brazil

We investigated the influence of position on a slope (plot relative elevation) and vegetation disturbance (the tallest tree height per plot) on community composition and diversity in a SE Brazilian Seasonal Semideciduous Forest (46°55′ W, 22°50′ S). Trees with dbh ≥5 cm were sampled in one hundred 10 × 10 m plots randomly placed in a 6.5-ha stand. Through partial Mantel test, floristic dissimilarities among plots (Jaccard index computed with species abundance in each plot) were correlated with environmental distances among plots (Euclidian distance index computed with relative elevation and the tallest tree height values in each plot). Relative elevation and the tallest tree per plot height were individually correlated with floristic gradients expressed by PCA axes scores using Pearson’s correlation coefficient. Through resampling, we compared diversity (richness, Berger-Parker D and Shannon H′) among plots in the drier (up) and moister (low) ends of the slope. Floristic dissimilarities were significantly correlated with environmental distances even after geographic distances among plots have been partialled out (r _m = 0.1274, p < 0.001). The first two PCA axes accounted for 22% of the total variance. After Bonferroni and Dutilleul’s corrections, axis 1 showed a marginally significant correlation with plot relative elevation (r = − 0.4097, p = 0.0309), and axis 2 was significantly correlated with the tallest tree height per plot (r = 0.2953, p = 0.0106). Position on the slope and vegetation disturbance were reliable predictors of community composition, thus suggesting the operation of niche assembly organizing processes. Richness and diversity (H′) decreased and dominance (D) increased with elevation on the slope. Dominance increase from D ₍₃₀₀₎ = 0.11 (confidence interval = 0.091–0.131) to D ₍₃₀₀₎ = 0.19 (CI = 0.165–0.210) surpassed the expected dominance increase based on the reduction of richness alone: D ₍₃₀₀₎ = 0.13 (CI = 0.110–0.140), thus highlighting the niche partitioning assembly of the community, especially among abundant species. Given the great amount of floristic variability remaining unexplained, stochastic processes, such as those related to dispersal limitation, may also have influence on the community composition. Therefore, both niche assembly and chance events can operate even on a fine local scale.     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

Distribution of seedlings and saplings of Quercus robur in a grazed deciduous forest

Seedling and sapling distribution of the deciduous tree Quercus robur was, studied in a grazed, partly open forest, ca. 20 km S of Linkoping, Sweden. Numbers of seedlings and saplings were compared between open and shaded parts of the forest. Cover values for the shrub and tree layers, proportion of bare ground and of stones and soil water content were determined. Q. robur was shown to germinate equally well in open and shaded parts of the forest; however, most saplings were found in the shade. No correlation was found between seedlings or saplings and cover of shrub layer or proportion of stones in the soil surface or with soil water content.     

Leaching of dissolved organic carbon,dissolved organic nitrogen,and other solutes from coarse woody debris and litter in a mixed forest in New York State

Coarse woody debris (CWD) may play a role in nutrient cycling in temperate forests through the leaching of solutes, including dissolved organic carbon (DOC) and dissolved organic nitrogen (DON), to the underlying soil. These fluxes need to be considered in element budget calculations, and have the potential to influence microbial activity, soil development, and other processes in the underlying soil, but studies on leaching from CWD are rare. In this study, we collected throughfall, litter leachate, and CWD leachate in situ at a young mixed lowland forest in NY State, USA over one year. We measured the concentrations of DOC, DON, NH₄⁺, NO₃⁻, dissolved organic sulfur, SO₄²⁻, Cl⁻, Al, Ca, K, Mg, Na, and P, estimated the flux of these solutes in throughfall, and measured the cover of CWD to gain some insight into possible fluxes from CWD. Concentrations of DOC were much higher in CWD leachate than in throughfall or litter leachate (15 vs. 0.7 and 1.6 mM, respectively), and greater than reported values for other leachates from within forested ecosystems. Other solutes showed a similar pattern, with inorganic N being an exception. Our results suggest that microsite scale fluxes of DOC from CWD may be An high relative to throughfall and litter leaching fluxes, but since CWD covered a relatively small fraction (2%) of the forest floor in our study, ecosystem scale fluxes from CWD may be negligible for this site. Soil directly beneath CWD may be influenced by CWD leaching, in terms of soil organic matter, microbial activity, and N availability. Concentrations of some metals showed correlations to DOC concentrations, highlighting the possibility of complexation by DOM. Several solute concentrations in throughfall, including DOC, showed positive correlations to mean air temperature, and fewer showed positive correlations in litter leachate, while negative correlations were observed to precipitation, suggesting both biological and hydrologic control of solute concentrations.     

The role of seed dispersal and seedling traits in colonization and coexistence ofSalix species in a seasonally flooded habitat

Regeneration traits of six co-occurringSalix species were studied on a floodplain of the Sorachi River, central Hokkaido, Japan, and their colonization success and coexistence in a local habitat were discussed. MixedSalix communities contained sixSalix species; dominant:S. sachalinensis; four subordinates:S. rorida, S. pet-susu, S. miyabeana andS. subfragilis; rare species:S. jessoensis. Their phenology, falling velocity and longevity of seeds, and the effects of microtopography and soil texture on seedling establishment were studied. The sixSalix species had overlapped seed dispersal periods that coincided with the decrease of water level after a predictable spring flood. This coincidence was crucial for the colonization success because the seedlings were established on wet soils left by the decreasing water level. They showed two types of regeneration trait, specialization and generalization.S. rorida andS. subfragilis showed contrasting regeneration traits; early vs. late seed dispersal, large vs. small seeds, seedling distribution on coarse vs. fine soils, respectively. These two species rarely co-occurred. On the other hand, the dominantS. sachalinensis had an intermediate seed size and dispersal timing, and a wide range of seedling distribution from coarse to fine soils. These results revealed that flooding seasonality influenced the colonization success together with the regeneration traits ofSalix species, and that coexistence of theSalix species was facilitated primarily by regeneration niche separation related to flooding seasonality and soil heterogeneity.     

Regeneration and coexistence of two subalpine conifer species in relation to dwarf bamboo in the understorey

Abstract. Microhabitats for seedling establishment and gap regeneration in subalpine forests of northern Japan were studied for two conifers, Abies sachalinensis and Picea glehnii. The abundance of understorey dwarf bamboo (Sasa spp.) was different for the four plots examined. Two types of micro-habitats were recognized for the two conifers: ground and elevated woody substrates (fallen logs and buttresses). Picea regenerated mostly on elevated sites, while Abies regenerated on both ground and elevated sites. The densities of Picea were independent of those of Sasa, but Abies densities decreased with increasing abundance of Sasa because Sasa reduced regeneration on the ground. Density of Abies on elevated sites was higher than that of Picea, irrespective of Sasa and of the density of adult trees. There was no significant difference in growth in sapling trunk height between the two conifers, but Picea grew more slowly under the canopy than Abies and was aggregated into gaps. Thus, in forests with less Sasa, the recruitment capacity of Abies was greater than that of Picea. The long life span of Picea compensated for its low density on elevated sites. Examination of a dynamic system model showed that Picea was excluded by Abies in forests without Sasa because regeneration on the ground is more advantageous than on elevated sites, but the two conifers could coexist in forests with Sasa because of the increased relative success of regeneration on elevated sites by Picea saplings.     

The herb and dwarf shrubs colonization of decaying logs in subalpine forest in the Polish Tatra Mountains

This is a study of the colonization pattern of herbs and dwarf shrubs on rotten logs in subalpine spruce forests (Plagiothecio Piceetum) in the Tatra Mountains. On four study plots (total area 1.43 ha.) all dead logs were measured and the decomposition stage was estimated using the 8-degree scale. For each log the cover of all vascular species, bryophytes and lichens was determined according to the methods of classical phytosociology. Constancy and an index of coverage were calculated for all vascular species growing on logs. The total volume of logs was relatively high (93 m³ ha^–1) and constituted 22% of the volume of living trees. Logs and stumps covered 411 m² ha^–1. These values are similar to those known from natural spruce forest from Carpathians and Scandinavia. The 8 stages of decomposition were equally represented, which indicates a constant supply of dead wood to the forest floor over time. The colonization of dead wood starts with lichens, followed by bryophytes and finally herbs and tree saplings. The first vascular plant colonists of dead logs appear at decay stage nr. 3 at least 20 years after tree death. The most suitable condition for most of the herb species corresponds to decay stage nr. 6 ca. 50 years after tree death. The herb cover is distinctively dominated by Vaccinium myrtillus. Simultaneously with herb species, tree seedlings colonize the logs. Constancy and abundance of Norway spruce saplings increases with advanced decomposition. It seems that the herb cover of logs does not hinder the regeneration of spruce.     

Response of tree biomass and wood litter to disturbance in a Central Amazon forest

We developed an individual-based stochastic-empirical model to simulate the carbon dynamics of live and dead trees in a Central Amazon forest near Manaus, Brazil. The model is based on analyses of extensive field studies carried out on permanent forest inventory plots, and syntheses of published studies. New analyses included: (1) growth suppression of small trees, (2) maximum size (trunk base diameter) for 220 tree species, (3) the relationship between growth rate and wood density, and (4) the growth response of surviving trees to catastrophic mortality (from logging). The model simulates a forest inventory plot, and tracks recruitment, growth, and mortality of live trees, decomposition of dead trees (coarse litter), and how these processes vary with changing environmental conditions. Model predictions were tested against aggregated field data, and also compared with independent measurements including maximum tree age and coarse litter standing stocks. Spatial analyses demonstrated that a plot size of ~10 ha was required to accurately measure wood (live and dead) carbon balance. With the model accurately predicting relevant pools and fluxes, a number of model experiments were performed to predict forest carbon balance response to perturbations including: (1) increased productivity due to CO₂ fertilization, (2) a single semi-catastrophic (10%) mortality event, (3) increased recruitment and mortality (turnover) rates, and (4) the combined effects of increased turnover, increased tree growth rates, and decreased mean wood density of new recruits. Results demonstrated that carbon accumulation over the past few decades observed on tropical forest inventory plots (~0.5 Mg C ha^–1 year^–1) is not likely caused by CO₂ fertilization. A maximum 25% increase in woody tissue productivity with a doubling of atmospheric CO₂ only resulted in an accumulation rate of 0.05 Mg C ha^–1 year^–1 for the period 1980–2020 for a Central Amazon forest, or an order of magnitude less than observed on the inventory plots. In contrast, model parameterization based on extensive data from a logging experiment demonstrated a rapid increase in tree growth following disturbance, which could be misinterpreted as carbon sequestration if changes in coarse litter stocks were not considered. Combined results demonstrated that predictions of changes in forest carbon balance during the twenty-first century are highly dependent on assumptions of tree response to various perturbations, and underscores the importance of a close coupling of model and field investigations.     

Natural disturbance and tree species coexistence in an old-growth beech - dwarf bamboo forest,southwestern Japan

Abstract. The structure and composition of a cool-temperate old-growth beech (Fagus crenata) - dwarf bamboo (Sasa spp.) forest, partially affected by landslide disturbance, in the Daisen Forest Reserve of southwestern Japan, were investigated in relation to forest floor and canopy conditions. All stems ≥ 4 cm DBH were mapped on a 4-ha plot and analyses were made of population structure, spatial distribution and spatial association of major tree species. The dominant species, F. crenata, which had the maximum DBH among the species present, had the highest stem density. However, for other species, larger-sized species had lower stem density with few smaller stems or saplings, while smaller-sized species had higher stem density with many smaller stems or saplings. Canopy trees of F. crenata were distributed randomly in the plot, while its stems in the other layers and all other species were distributed patchily. Small patches represent gap-phase regeneration. Larger patches correlate with landslide disturbance, difference in soil age, or the presence/absence of Sasa. Cluster analysis for spatial associations among species and stems in the different layers revealed that the forest community consists of several groups. One main group was formed on sites not covered with Sasa. This group contained a successional subgroup (from Betula grossa to Acer mono and/or F. crenata) initiated by landslide disturbance and a subgroup of tree species that avoid Sasa. Another group was formed on sites with mature soils covered largely with Sasa. This contained associations of canopy trees of F. crenata and smaller-sized tree species such as Acanthopanax sciadophylloides and Acer japonicum. It is found that the community of this old-growth beech forest is largely organized by natural disturbance and heterogeneous conditions of the forest floor (difference in soil age and presence/absence of Sasa). The existence of these different factors and the different responses of species to them largely contribute to the maintenance of tree species diversity in this forest.; Keywords: Cluster analysis; Fagus crenata; Forest dynamics; Gap; Landslide; Spatial pattern.     

Seedling establishment in relation to microhabitat variation in a windthrow gap in a boreal Pinus sylvestris forest

Abstract. The characteristics of microhabitats of established Pinus sylvestris and Betula seedlings were studied in a small windthrow gap in a mature P. sylvestris-dominated forest in the Petkeljärvi National Park in eastern Finland. Seedlings were strongly clustered in disturbed microhabitats, particularly uprooting pits and mounds, formed by tree falls. They covered 3% of the 0.3.ha study area consisting of the gap and some of the forest edge. Although Betula occurred only as scattered individuals in the dominant canopy layer of the forest, it accounted for 30% of the seedlings found in the study area. Betula regeneration was almost completely restricted to pits and mounds, where 91% of the seedlings were found. Uprooting spots were also the most important regeneration microhabitats for Pinus, where 60% of the seedlings grew, even though the seedlings were found in other substrates as well, particularly on sufficiently decomposed coarse wood. Undisturbed field- and bottom-layer vegetation had effectively hindered tree seedling establishment, which emphasises the role of soil disturbance for regeneration. While the establishment of seedlings was found to be clearly determined by the availability of favourable regeneration microhabitats, the early growth of seedlings was affected by a complex interaction of environmental variables, including the type of microhabitat, radiation environment and interferences caused by competing seedlings and adjacent trees. In the most important regeneration microhabitats, i.e. in uprooting pits and on mounds, the distributions of the local elevations of Pinus and Betula seedlings were different. Pinus seedlings occurred closer to ground level, i.e. on the fringes of pits and lower on mounds, while Betula seedlings grew deeper in pits and higher on mounds. The position of the Betula seedlings indicate that they may have a competitive advantage over Pinus seedlings in the dense seedling groups occurring in uprooting spots. We suggest that this initial difference in Pinus and Betula establishment may affect the subsequent within-gap tree species succession and can, in part, explain the general occurrence of Betula in conifer-dominated boreal forests.     

Structure of a pristine Picea abies forest in northeastern Europe

Abstract. The forest structure in a large, relatively homogeneous area of pristine Picea abies taiga in the southern boreal region west of the Ural mountains was studied along four 500-m long transects. P. abies dominated the forest in association with Abies sibirica and Betula spp. The mean volume of living trees was 216 m³/ha. This value varied among the four transects, from 138 - 252 m³/ha. Mean density of trees (DBH > 1 cm) (and variation over the transects) was 2 064/ha (1670 - 2710). Living trees classified as dying or seriously damaged made up 2.9 (2.5 - 3.5) % of the living tree volume. Betula was an important canopy component and made up 16% of the living tree volume. The number of dead standing trees varied from 195 - 325/ha, corresponding to a volume of 10.8 - 70.7 m³/ha. The density of trees with a broken stem was 90 - 170/ha and their estimated volume 7.6 - 41.3 m³/ha. Standing dead trees and trees with broken stems represented 10.4 and 8.9% of the total standing tree volume (living + dead), respectively. The mean volume of decaying logs on the forest floor was 117 (84.4–156.2) m³/ha, corresponding to 54 (35 - 113) % of the living tree volume. The canopy-forming trees were present in the understory as abundant saplings and suppressed individuals. The size distribution of the dominant tree species resembled the inverse J-shape. Generally, the forest was characterized by a high small-scale structural variation and a larger-scale relative homogeneity. This pattern is consistent with forest dynamics where the forest consists of a small-scale mosaic of patches in different stages of recovery following disturbance. Our results suggest that the ecology and dynamics of extensive areas of natural boreal forests can be driven by small-scale disturbance.     

Effects of river level fluctuation on plant species richness, diversity, and distribution in a floodplain forest in Central Amazonia

River levels in Central Amazonia fluctuate up to 14 m annually, with the flooding period ranging from 50 to 270 days between the rising and falling phases. Vast areas of forest along the rivers contain plant species that are well adapted to annual flooding. We studied the effect of flooding level on tree species richness, diversity, density, and composition in lake, river, and stream habitats in Jaú National Park, Brazil. 3051 trees >10 cm diameter (at 1.3 m diameter at breast height, dbh) were measured and identified in 25 10 m × 40 m randomly selected plots in each habitat. Ordination methods and analysis of variance results showed that forested areas near lakes had significantly lower species richness of trees than riverine and streamside habitats. Plot species richness and diversity were strongly negatively correlated with the water level and duration of flooding. The drier (stream) habitat had more total species (54 species of trees) and more unique species of trees (6 tree species) than the riverine (52 tree species; 3 unique species) and lake (33 tree species; 3 unique species) habitats. Species composition overlap among habitats was surprisingly high (42.6–60.6% overlap), almost one-third of the species were found in all three habitat types, and few species were unique to each habitat. We conclude that: (1) duration of flooding has a strong impact on species richness, diversity and plant distribution patterns; (2) most species are adapted to a wide range of habitats and flood durations; and (3) while flood duration may decrease local diversity, it also creates and maintains high landscape-scale diversity by increasing landscape heterogeneity. Received: 20 April 1997 / Accepted: 14 January 1999     

Tree competition and species coexistence in a cool-temperate old-growth forest in southwestern Japan

Abstract. The growth dynamics and mode of competition between adult trees ≥ 4 cm in DBH (stem diameter at breast height 1.3 m) of eight abundant species occupying ca. 90 % of the total basal area were investigated in a 4-ha study plot (200 m × 200 m) of a cool-temperate, old-growth forest on Mount Daisen, southwestern Japan. In the study plot, 30 tree species with individuals ≥ 4.0 cm DBH co-occurred. A bimodal DBH distribution showing upper and lower-canopy layers was found for the most dominant and largest species, Fagus crenata (ca. 78 % of the total basal area), whilst other tree species showed unimodal DBH distributions corresponding mostly to the lower-canopy layer. We developed a model for individual growth incorporating both intra and interspecific competition and the degree of competitive asymmetry. Onesided interspecific competition was detected only from Fagus crenata (upper-canopy species) to Acer japonicum and Acanthopanax sciadophylloides (lower-canopy species) on the scale of the 4-ha study plot. Only Acanthopanax sciadophylloides showed symmetric intraspecific competition. However, a positive (non-competitive) interspecific relationship between adult trees prevailed over a competitive relationship; for example, individual DBH growth rate of Fagus crenata (especially lower-canopy trees) was correlated with the abundance of Acer mono. The positive relationship represented a group of species with similar habitat preference [soil type (mature or immature) caused by landslide disturbance and the presence/absence of Sasa dwarf bamboos in the understorey], where tree densities were not so high as to bring about competition. Competitive interactions between adult trees ≥ 4 cm in DBH occurred only locally between a few specific species and were suggested to be almost irrelevant to the variation in species coexistence on the 4-ha scale of cool-temperate forest. Rather, the coexistence of 30 tree species (species diversity) on this large scale was suggested to be governed by the regeneration pattern of each component species (habitat preference, seedling establishment, sapling competition) with respect to landslide disturbance.