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.     

Habitat associations with topography and canopy structure of tree species in a tropical montane forest on Mount Kinabalu,Borneo

Habitat associations with topography and canopy structure of 42 abundant tree species were studied in a 2.74-ha plot of tropical montane forest on Mount Kinabalu, Borneo. Many of these species belong to the same higher taxa including eight families and four genera. Analysis of intraspecific spatial distributions for stems ≥ 10 cm diameter revealed that 28 species (including all six species of Fagaceae) showed aggregated distributions at the 100-m² and/or 400-m² scales, and that 20 species showed habitat associations with topography by torus-translation tests; 17 species showed both characteristics. Species' associations with the local canopy structure were characterized by crown position index (CPI), which was defined relative to neighbour trees. The CPI differed greatly among individual stems at 10–40 cm diameter, and 19 species showed significantly different frequencies of crowns exposed vertically versus those shaded beneath the canopy. Mean growth rates at 10–40 cm diameter and size distributions of species were not related to topographic associations, but were explained by the associations with canopy structure; species with more exposed crowns grew faster and had less positively skewed distributions. Diversity in habitat associations was manifest between two genera (Syzygium and Tristaniopsis) in the family Myrtaceae and among species in these genera, but was less evident in other families and two genera (Garcinia and Lithocarpus). This revised version was published online in June 2006 with corrections to the Cover Date.     

The cupule-carpel theory. A defence

Background: Spatial patterns within forests reflect their formative processes. In Kamchatka, the dominant stone birch (Betula ermanii) forest exists in a primeval state. Betula species can reproduce via clonally produced ramets, leading to a clustered distribution of stems, or from seed on open ground, although subsequent mortality may obscure initial establishment patterns.

Methods: Spatial patterns of all trees and regenerating stems in a fully mapped 0.25 ha plot were analysed using the g(r) function and the pattern of trees modelled using a Matérn cluster process. Mark correlation analysis was used to detect patterns in stem sizes.

Results: Clustering of trees at scales up to 3 m occurred, with model parameters indicating a density of 180 clusters ha^?1, each containing on average three stems >1 cm diameter at breast height (DBH) within a 1.6 m radius. Stem size compensation was detected, with nearby trees smaller than expected by chance. Regenerating stems were strongly clustered at scales below 3.5 m and from 5–8 m, and were aggregated around small trees (<10 cm DBH) but not large trees (≥10 cm DBH), or in gaps.

Conclusions: These patterns are consistent with clonal reproduction followed by competition as the processes determining spatial structure in old-growth stone birch forests.     

Stand structure and regeneration in a Kamchatka mixed boreal forest

Abstract. A 1‐ha plot was established in a Betula platyphylla‐Picea ajanensis mixed boreal forest in the central Kamchatka peninsula in Russia to investigate stand structure and regeneration. This forest was relatively sparse; total density and stand basal area were 1071/ha and 25.8 m²/ha, respectively, for trees > 2.0 cm in trunk diameter at breast height (DBH). 25% of Betula regenerated by sprouting, and its frequency distribution of DBH had a reverse J‐shaped pattern. In contrast, Picea had a bimodal distribution. The growth rates of both species were high, reaching 20 m in ca. 120 yr. The two species had clumped distributions, especially for saplings. Betula saplings were not distributed in canopy gaps. Small Picea saplings were distributed irrespective of the presence/absence of gaps, while larger saplings aggregated in gaps. At the examined spatial scales (6.25–400 m²) the spatial distribution of Betula saplings was positively correlated with living Betula canopy trees and negatively with dead Picea canopy trees. This suggests that Betula saplings regenerated under the crowns of Betula canopy trees and did not invade the gaps created by Picea canopy trees. The spatial distribution of Picea saplings was negatively correlated with living and dead Betula canopy trees and positively with dead Picea canopy trees. Most small Picea seedlings were distributed under the crowns of Picea trees but not under the crowns of Betula trees or in gaps. This suggests that Picea seedlings establish under the crowns of Picea canopy trees and can grow to large sizes after the death of overhead Picea canopy trees. Evidence of competitive exclusion between the two species was not found. At a 20 m × 20 m scale both skewness and the coefficient of variation of DBH frequency distribution of Picea decreased with an increase in total basal area of Picea while those of Betula were unchanged irrespective of the increase in total basal area of Betula. This indicates that the size structure of Picea is more variable with stand development than that of Betula on a small scale. This study suggests that Betula regenerates continuously by sprouting and Picea regenerates discontinuously after gap formation and that the species do not exclude each other.     

Variations in resistance to canopy disturbances and their interactions with the spatial structure of major species in a cool‐temperate forest

Question: How does typhoon‐related disturbance (more specifically, disturbance in the understorey due to tree‐fall and branch‐fall) affect different species mortality rates in a vertically well‐structured forest community? Location: Cool‐temperate, old‐growth forest in the Daisen Forest Reserve, Japan. Methods: We investigated the canopy dynamics and mortality rate trends of trees ≥5 cm diameter at breast height in a 4‐ha study plot, and analysed the effects of tree diameter and spatial structure on the mortality risks for major tree species in the understorey. Results: Significant differences were found in the mortality rates and proportions of injured dead stems between census periods, which were more pronounced in the understorey than in the canopy. Acer micranthum, which showed increased mortality during typhoon disturbance periods, had a clumped distribution. In contrast, Acer japonicum and Viburnum furcatum, which showed similar mortality rates between census periods, had a loosely clumped spatial distribution and a negative association with canopy trees, respectively. In the understorey stems of Acanthopanax sciadophylloides and Fagus crenata, whose spatial distribution patterns depended on canopy gaps, significant increases in mortality rates were observed only during severe typhoon‐related disturbance periods. Conclusions: The sensitivity of trees to typhoon‐related canopy disturbance is more pronounced in the lower layers of vertically structured forest communities. Differences in mortality patterns generated through the combined effects of spatial variation in disturbance regime and species‐specific spatial distribution patterns (spatial aggregation, association with canopy trees, and canopy gap dependency) contribute to the co‐existence of understorey species in forest communities that are subject to typhoon‐related disturbance.     

长白山阔叶红松林中红松与紫椴的空间分布格局及其关联性

以长白山阔叶红松林25hm2样地调查数据为基础,采用点格局分析方法O-ring统计,分析了红松与紫椴两个优势树种在主林层、次林层和林下层的空间分布格局,以及各林层之间的种内和种间关联性.结果表明:在小尺度上,红松和紫椴总体上呈聚集性分布,但不同林层的分布格局各异.物种在较低的林层呈明显的聚集分布,而在较高的林层则呈随机或规则分布,聚集度随林层的增高而降低.主林层与次林层红松在<11m的尺度内呈明显的正相关.主林层与次林层、林下层紫椴之间呈负相关.红松和紫椴在整体上表现为正相关,但不同林层间的关联性各异.主林层红松与3个林层紫椴之间没有明显的关联性;次林层红松与次林层紫椴在>4m的尺度上呈负相关,而与林下层紫椴没有明显的关联性;主林层紫椴与次林层红松在0~100m尺度内都表现为显著正相关.     

海南尖峰岭热带山地雨林不同空间尺度和径级水平的物种丰富度与个体密度关联

以海南尖峰岭热带山地雨林为研究对象, 基于尖峰岭原始林60 hm²大样地, 分析了7个不同空间尺度(5 m × 5 m、10 m × 10 m、20 m × 20 m、40 m × 40 m、60 m × 60 m、80 m × 80 m、100 m × 100 m)、3个不同径级(胸径(DBH) ≥ 1.0 cm、DBH ≥ 2.5 cm、DBH ≥ 7.5 cm)物种丰富度与个体密度之间的关联性, 以期为负密度制约效应等研究的尺度选择提供参考依据。研究结果表明, 物种丰富度和个体密度之间的关联性是基于空间尺度存在的, 并受分析植株径级大小的影响。DBH ≥ 1.0 cm时, 在4个空间尺度下(5 m × 5 m、10 m × 10 m、20 m × 20 m、40 m × 40 m), 物种丰富度随个体密度增加而增加, 但物种丰富度和个体密度两者间的相关性随取样空间尺度的增大而逐渐下降; 在其他3个空间尺度下(60 m × 60 m、80 m × 80 m、100 m × 100 m), 两者间没有显著相关性。当DBH ≥ 2.5 cm和DBH ≥ 7.5 cm时, 物种丰富度和个体密度间的相关性随空间尺度变化的趋势也与上述结果基本一致; 在60 m × 60 m的空间尺度上, DBH ≥ 1.0 cm时物种丰富度和个体密度相关性不显著, DBH ≥ 2.5 cm和DBH ≥ 7.5 cm两个径级的物种丰富度和个体密度表现出弱相关性。DBH ≥ 2.5 cm和DBH ≥ 7.5 cm时, 各空间尺度下物种丰富度和个体密度之间的决定系数几乎一致, 但高于DBH ≥ 1.0 cm时两者之间的决定系数, 说明大径级植株的个体密度和物种丰富度之间有更强的关联性, 负密度制约效应和自疏效应可能是产生这种现象的主要原因。     

太白山牛皮桦-巴山冷杉混交林空间格局及关联性研究

采用改进的点格局分析法——邻域密度函数（Neighborhood Density Function,NDF）对太白山地区桦林-冷杉林交错区内主要树种的空间分布格局、种间空间关联性及其更新空间关联性进行了分析。结果表明：各树种所有个体整体上都呈随机分布,划分不同层级后,则各林层个体分布格局各异,各树种在较低的林层呈明显聚集分布,随林层增高,中、高层内个体聚集分布的空间尺度降低。两个主要竞争种牛皮桦（Betula albo-sinensis var．septentrionalis）和巴山冷杉（Abies fargesii）整体表现出显著空间负相关,巴山冷杉与玉皇柳（Salix yuhuangshanensis）未表现出明显相关性,而牛皮桦与玉皇柳间则有空间负相关的趋势。牛皮桦的更新格局仅依靠自身倒木扰动产生的机会,且被限定在较小尺度上。巴山冷杉在小尺度上会限制自身更新,此外与牛皮桦在小尺度上竞争激烈,可占据利用其空间位置进行更新。从空间格局分析角度认识牛皮桦和巴山冷杉的生态学特性,为进一步研究其共存机制,认识太白山桦林在秦岭植被垂直带中的作用和地位提供参考依据。     

Regeneration processes of a boreal forest in Kamchatka with special reference to the contribution of sprouting to population maintenance

We studied regeneration patterns of three tree species Picea ajanensis, Betula platyphylla and Populus tremula from 1998 to 2000 in the Central Depression of the Kamchatka Peninsula. We paid special attention to the contribution of sprouting to their regeneration. P. ajanensis was the only species that regenerated by seedling. In a 40 × 40 m study plot, the density of P. ajanensis saplings < 2.0 cm in diameter at basal area (DBH) was 1132, and this was the highest among the three species studied. The number of saplings 2 cm in DBH declined sharply with size class. The spatial distribution of P. ajanensis saplings (< 2 cm in DBH) showed a significant positive correlation with that of adult trees and a negative correlation with that of gaps. These trends were not changed after re-measurement in 2000, although nearly half of the juveniles had died or been injured during the two years. These results suggest that small Picea saplings prefer habitats under the canopy of adult trees rather than in gaps for establishment. Most small individuals of B. platyphylla were produced from sprouts. The number of saplings in the smallest size class (< 2 cm in DBH) was much less than that of P. ajanensis, although the number of larger individuals did not decrease remarkably. The spatial distribution of B. platyphylla saplings showed a positive correlation with that of adult trunks and a negative correlation with that of canopy trees of P. ajanensis. These results suggest an effective contribution of sprouts to the regeneration of B. platyphylla. P. tremula was the only species that could invade big gaps and produce many root suckers efficiently. There were 181 suckers of P. tremula in the smallest size class (< 2 cm in DBH) in the study plot, although the number of saplings 2 cm in DBH declined abruptly. The spatial distribution of saplings of this species showed a slight positive correlation with that of gaps, and negative correlation with that of adult trees of B. platyphylla, P. ajanensis, and P. tremula. The root suckering strategy of P. tremula might be adaptive under severe conditions in high-latitude regions. Our data suggest, however, that it does not necessarily contribute to regeneration in mature forests. The three component species in this forest did not seem to utilize canopy gaps for regeneration; we suggest that gap dynamics do not work in this forest. The sparse canopy, which is a typical character of forests in high-latitude regions, might be a consequence of high mortalities of seedlings and root suckers inside gaps.     

Spatial pattern of Quercus regeneration limitation and Acer rubrum invasion in a Piedmont forest

Abstract. Across eastern North America, there is a temporal trend from open Quercus forests to closed forests with increased Acer rubrum in the understory. We used a series of Ripley's K(d) analyses to examine changes in the spatial pattern of Quercus and Acer rubrum stems greater than 2.5 cm DBH over 45 yr in a 2‐ha mapped stand. Specifically, we asked whether changes over time were consistent with the hypothesis that Quercus is being competitively replaced by Acer rubrum. Both Acer rubrum and Quercus stems are spatially clumped, but have become less clumped over time. Stem mortality from Hurricane Fran (1996) was more clumped in all strata of the forest, at all spatial scales, than expected if damage had occurred to stems at random. Acer rubrum ingrowth occurred more often near established trees (all species) in the midstory, whereas Quercus ingrowth occurred less often near established trees in the midstory. The specific hypothesis that stems of Acer rubrum in the midstory of the forest are associated with a lack of Quercus regeneration was strongly supported. This effect occurred at all spatial scales tested, including scales larger than that at which direct competition for light can occur. Edaphic gradients in the plot are correlated with many of the observed trends at large spatial scales, and our results suggest that the presence of such gradients can generate complex spatial patterns over time.     

Interception of photosynthetic photon flux density by woody elements in a hinoki (Chamaecyparis obtusa [Sieb. et Zucc.] Endl.) stand

A hinoki stand was divided into the following two layers: one was a leafed upper layer consisting of leaves and woody elements, such as stems, living branches and dead branches; the other was a leafless layer consisted of woody elements, such as stems and dead branches. Photosynthetic photon flux density (PPFD) penetrating through the stand was measured in relation to the silhouette area of leaves and woody elements. The silhouette area, on a ground area basis, was 11.2 ha ha⁻¹ for leaves (leaf area index), 0.26 ha ha⁻¹ for stems, 0.40 ha ha⁻¹ for living branches and 0.69 ha ha⁻¹ for dead branches, yielding a total wood silhouette area of 1.35 ha ha⁻¹. The apparent extinction coefficient,K, was computed to be 0.420 ha⁻¹ ha for the leafed layer, while the extinction coefficient of woody elements,K _c, was computed to be 1.01 ha⁻¹ ha for the leafless layer. The cumulative wood silhouette area density from the top of the canopy down to a given depth increased with an increase in the corresponding leaf area density within the leafed layer. The extinction coefficient of leaves,K _F, was estimated to be 0.367 ha⁻¹ ha. Of the PPFD extinguished within the canopy, the fraction (K _F/K) due to leaves alone was evaluated to be 87.4%.     

川西米亚罗亚高山暗针叶林岷江冷杉和糙皮桦空间格局及其关联性分析

基于2019年9月完成的川西米亚罗亚高山暗针叶林9.6 hm~2动态样地的调查数据，分析了主要优势种岷江冷杉(Abies fargesii var.faxoniana)与糙皮桦(Betula utilis)的径级结构，并运用O-ring统计中的成对相关函数g(r)和g₁₂(r),研究了两个优势种的空间格局及其种内、种间的关联性。结果表明：岷江冷杉与糙皮桦径级分布连续，均呈倒“J”型。岷江冷杉与糙皮桦的不同径级个体主要呈现由聚集分布向随机分布发展的趋势。岷江冷杉的小树对幼树、中树对幼树、中树对小树以及大树对中树的种内相关性在所有空间尺度上均为显著正相关(P<0.05),而相同径级糙皮桦的种内相关性则表现为由显著正相关(P<0.05)逐渐向无显著相关(P>0.05)变化。随着空间尺度的增大，岷江冷杉的大树对幼树、大树和小树由无显著相关(P>0.05)逐渐向显著正相关(P<0.05)变化，糙皮桦的大树对幼树由显著负相关(P<0.05)逐渐向无显著相关(P>0.05)变化。岷江冷杉大树对糙皮桦所有径级个体的种间相关性主要呈现由为显...     

Composition and structure of a chronosequence of young, mixed-species forests in southeastern Ohio, USA

Clearcutting, a commonly used silvicultural practice in southeastern Ohio, often results in a forest stand with a different species composition than the parent stand. The time frames during which shifts in species composition occur on different sites are unclear. While some studies have documented species composition at specific points in time, none have attempted to examine differences throughout the first decades of stand development. This study focused on the early successional dynamics of young, mixed-species forests of southeastern Ohio. Species compositions were examined across a chronosequence of sixteen stands that developed following clearcutting. Stand ages ranged from six to 26 years. The sample was limited to dry-mesic hardwood forests on southerly aspects and on soils derived from residuum or colluvium. Across the chronosequence, stand density ranged from 17 636 stems ha^-1 at age 6 to 2759 stems ha^-1 at age 26, and basal area ranged from 8.2 m² ha^-1 to 22.1 m² ha^-1. Clumps comprised a substantial portion of the total stand density and basal area. At age 6–8 years after clearcutting, clumped stems accounted for 35.1% of the density and 48.2% of the basal area. At age 26 years, these proportions were 25.7% and 29.4%, respectively. Clumped stems were significantly larger (p<0.05) than non-clumped stems at each age group except 26 years. Total Quercus spp. density was greatest at age 6–8 years (3386 stems ha^-1), and least at age 26 years (581 stems ha^-1). When considered as a proportion of the total stand, however, the proportion was relatively stable, averaging 21.3%. However, importance value (IV=[relative density + relative basal area]/2) of Quercus in the upper canopy (dominant and codominant crown classes) was twice as much (72%) at age 26 years compared to age 6–8 years (35%). Quercus prinus L. was the major species across the chronosequence. For all age groups except 18–20 years. Q. prinus IV was the highest of any individual species in the upper canopy, and it ranged from 27 in the youngest stands (6–8 years) to 69 in the oldest stand (26 years). Within the intermediate crown class, the IV of Q. prinus equaled or exceeded those of all other species, except for the 18–20 year age group where it was second to A. rubrum. Quercus alba L. and Quercus velutina Lam. were minor components at age 26 years, although they dominated a comparison sample of six mature stands of the same ecosystem type. Liriodendron tulipifera L. was abundant 6–8 years after clearcutting, but nearly absent at age 26 years. Acer rubrum L. was the major species in both the intermediate and overtopped crown classes throughout the chronosequence. As gaps in the canopy occur. A. rubrum may become a more common species within the dominant-codominant crown class.     

天山云杉天然林不同林层的空间格局和空间关联性

天山云杉(Picea Schrenkiana)林是天山林区的重要针叶林。研究调查了分别代表天山云杉增长型种群和成熟型种群的两块标准地,应用Ripley's K系数法研究了天山云杉更新层、低林层、中林层和高林层的空间格局和不同林层的空间关联性,并应用Moran's I系数法研究了不同林层的林木高度的空间自相关性。结果表明:所有林层的天山云杉活立木都是显著聚集的,只是发生聚集格局的强度和聚集尺度不同。更新层天山云杉具有最大的聚集强度,从更新层到低林层,聚集强度急速降低。两块标准地中天山桦(Betula tianschanica Rupr.)、天山柳(Salix tianschanica Rgl.)和天山云杉活立木的空间关联性的表现不同。研究结果表明:增长型种群中天山桦、天山柳在所有尺度上都抑制天山云杉的生长;而成熟型种群中天山桦、天山柳在1-10 m尺度内对天山云杉种群的生长没有影响,10-40 m表现出正相关,40 m以上没有表现出明显的关联性;同时天山桦、天山柳基本上在所有尺度都抑制天山云杉的更新。在更新层和低林层间表现出正相关;而更新层和枯死木之间的空间关联性为正相关。     

Mid-successional stand dynamics in a cool-temperate conifer-hardwood forest in northern Japan

Stand dynamics was studied over 13 years in a cool-temperate conifer-hardwood forest, northern Japan. A total 30 hardwood species and one conifer, Abies sachalinensis, larger than 1.5 cm DBH were recorded. The total stand density was 1677 trees ha⁻¹ at the beginning, decreasing to 1184 trees ha⁻¹ (30% reduction) over the study period, but the total stand basal area was almost unchanged (about 49 m² ha⁻¹). This large reduction in total density was mainly due to the death of saplings and infrequent recruitment. Number of recruits gradually decreased with time, while that of dead trees was constant. Cause of death of small trees was mainly due to suppression by tall trees. Skewness of the DBH frequency distribution varied among the species. A less skewed frequency distribution (i.e., few number of saplings) was shown by shade-intolerant species such as Populus maximowiczii and Betula maximowicziana, and a more skewed frequency distribution (i.e., large number of saplings) by shade-tolerant species such as Acer mono and Tilia japonica. DBH frequency distribution changed to less skewed patterns with reduction of density in most species during the census period. Rank of shade tolerance positively correlated with tree density and skewness, and negatively correlated with mean DBH. Skewness also positively correlated with recruitment rates. Furthermore, rank of shade tolerance positively correlated with seed size. These results suggest that shade-intolerant species regenerated immediately after disturbances by wide dispersal of small seeds, but their recruitment was interrupted after that. By contrast, shade-tolerant species were able to recruit even after the ceasation of recruitment of shade-intolerant species, but suffered severe mortality due to the increasing shading with the progress of stand development. This study suggests that the stand is still developing, with changes in species composition and size structure, and that species differences in shade tolerance and seed size are important for the stand structural changes.     

鼎湖山南亚热带常绿阔叶林20公顷样地群落特征研究

 鼎湖山南亚热带常绿阔叶林是北回归线附近保存较完好的地带性植被,是在南亚热带季风湿润型气候条件下发育的植被类型,具有热带向亚热带的过渡性质,群落结构相对复杂,组成种类相对丰富。参照巴拿马巴洛科罗拉多岛（Barro Colorado Island, BCI） 50 hm²热带雨林样地的技术规范,采用中国森林生物多样性监测网络的统一调查研究方法,于2005年在鼎湖山南亚热带常绿阔叶林建立了20 hm² 固定监测样地,调查并鉴定了样方内胸径 （Diameter at breast height, DBH）大于1 cm的木本植物。首次调查结果表明,群落内共有木本植物210种,71 617个活的个体,分属于56科119属。从乔木区系的组成及其特点可以看出,其南亚热带的区系成分占绝对优势,并呈现出由亚热带向热带过渡的趋势。群落垂直结构复杂,地上成层现象较明显,乔木可分为3层,其中重要值最大的锥（Castanopsis chinensis）、木荷（Schima superba）和黄杞（Engelhardtia roxburghiana）均是乔木上层的优势种;中层是群落的主要层,由厚壳桂（Cryptocarya chinensis）、黄叶树（Xanthophyllum hainanense）和华润楠（Machilus chinensis）等中生和耐阴树种组成;下层成分较复杂,物种多样性高,不同地段的物种组成差异较大。样地内物种十分丰富,种-面积曲线拟合显示其物种数量接近于BCI。稀有种比例极高,有110种,占总物种数的52.38%,其中有45%的稀有种源于物种本身的特性,有20%源于区系交汇,人为或自然干扰造成的稀有种占30%以上。样地中所有个体的径级分布(以1 cm等级排列)明显呈倒“J"形,表示群落稳定与正常生长状态。根据对优势种径级结构的分布分析,将各树种的径级结构归纳为4种类型：1）峰型（中径级个体储备型）,此类物种为乔木上层优势种;2）倒“J"型 （正常型）,此类物种为乔木中层的优势种;3）类倒“J"型（偏正常型）,此类物种占据乔木的中、下层; 4）“L”型（灌木型）,此类物种分布于乔木下层和灌木层。点格局分析得出优势种在各个层上的绝大部分尺度都是聚集分布的,而不同径级的空间分布随物种的不同而呈现出明显的差异性,点格局显示了优势种在相同径级的分布（特别是DBH=10～40 cm）具有一定的空间互补性。对大径级（DBH＞40 cm）的504个个体进行空间分布格局分析,结果表明,所有这些个体随机分布于整个样地,但不同的物种在空间分布上存在一定的规律。     

Stand Structure and Maintenance of <Emphasis Type="Italic">Picea jezoensis</Emphasis> in a Northern Temperate Forest,South Korea

Stand structure and spatial distribution of Picea jezoensis (Siebold et Zucc.) Carrière on Mt. Gyebang, Korea was investigated to provide information on the structural characteristics and the maintenance of P. jezoensis population in northern temperate mixed coniferous forests. Height and diameter at breast height (DBH) distribution, age, growth, and spatial distribution patterns of P. jezoensis were examined in thirty nine 100-400 m² quadrats or circular plots. The overall stand structure attributes in the study sites are stem density of 709 trees ha⁻¹, a mean DBH of 12.8 cm, and a mean height of 5.6 m, with reverse J shapes of DBH and height distributions. The stem density of P. jezoensis population was 81 trees ha⁻¹, a mean DBH of 20.7 cm, and a mean height of 9.1 m, showing bimodal-like shapes in age and DBH distributions. Several growth release periods implied that P. jezoensis stands experienced small disturbances. The radius of patches of similar-sized P. jezoensis in the variogram was equivalent with the height of the tallest trees, indicating that patches were established following the fall of trees in the upper canopy layer. Small windthrows in this region contributed to the maintenance of the P. jezoensis stand by releasing sapling growth and providing nursing logs and space for seedlings.     

Structure and regeneration of canopy species in an old-growth evergreen broad-leaved forest in Aya district,southwestern Japan

The population structure and regeneration of canopy species were studied in a 4 ha plot in an old-growth evergreen broad-leaved forest in the Aya district of southwestern Japan. The 200 m × 200 m plot contained 50 tree species, including 22 canopy species, 3,904 trees (dbh5 cm) and a total basal area of 48.3 m²/ha. Forty one gaps occurred within the plot, and both the average gap size (67.3 m²) and the total area of gap to plot area (6.9%) were small. Species found in the canopy in the plot were divided into three groups (A, B, C) based on size and spatial distribution patterns, and density in each tree size. Group A (typical species: Distylium racemosum, Persea japonica) showed a high density, nearly random distribution and an inverse J-shaped size distribution. Species in group B (Quercus salicina, Quercus acuta, Quercus gilva) were distributed contagiously with conspicuous concentration of small trees (<5 cm dbh) around gaps. However, the species in this group included few trees likely to reach the canopy in the near future. Group C included fast-growing pioneer and shade intolerant species (e.g. Cornus controversa, Carpinus tschonoskii, Fagara ailanthoides), which formed large clumps. Most gaps were not characterized by successful regeneration of group B and C but did appear to accelerate the growth of group A. Group B species appear to require long-lived or large gaps while group C species require large, catastrophic disturbances, such as landslides, for regeneration.     

Vegetative Life History of a Dominant Rain Forest Canopy Tree1

Tetramerista glabra has a remarkable combination of life history traits. It is a dense-wooded, large, common canopy tree in primary peat swamp rain forest. Its seedlings, although shade tolerant, can grow rapidly in high light conditions, but frequently lack structural stability. Most juvenile stems (94% in the understory and 38% in canopy gaps) collapse under their own weight or from branchfalls. Fallen stems then ramify into vegetative sprouts, which in turn may collapse, perpetuating a vegetative juvenile cycle. Most recruitment is from sprouts rather than from seed. Structural analysis of stem dimensions shows that stems 2–8 cm DBH (diameter at breast height) are close to the theoretical buckling limit, especially for those dependent on neighboring vegetation to maintain vertical form. Trees > 4 cm DBH persisting as upright stems develop stilt root support and become structurally independent at ca 8 cm DBH. Eventually, as stems thicken, stilt roots anastamose and trees adopt the cylindrical growth form of mature canopy trees (up to 150 cm DBH). Thus, the vegetative life history strategy of the species is to: (i) regenerate a large “ramet bank” from the majority of juveniles that fail structurally while suppressed in the understoty, and (ii) to maximize height growth at the expense of diameter growth in high light conditions. The growth pattern and plastic form of T. glabra shows how a shade tolerant species may adapt to utilize the ephemeral light resource in canopy gaps. The growth strategy of this species allows it to circumvent the normal trade-off between shade tolerance and rapid growth in canopy gaps.     

长白山阔叶红松林样地(CBS)：群落组成与结构

 阔叶红松(Pinus koraiensis)林是中国东北地区的地带性植被,长白山区是阔叶红松林的核心分布区。参照巴拿马Barro Colorado Island (BCI) 50 hm² 热带雨林样地的技术规范,于2004年在长白山自然保护区的阔叶红松林内建立了一块25 hm²的固定样地（简称CBS）,这是目前中国科学院生物多样性委员会中国森林多样性动态研究网络中最北端的一块,也是全球温带地区最大的一块森林样地。2004年夏的第一次调查结果表明,样地内胸径≥1 cm的木本植物有52种,隶属于18科32属。总的独立个体数为38 902,包括分枝的总个体数为59 121。植物组成上属典型的长白山植物区系,同时混有一些亚热带和亚寒带成分。群落优势种明显,个体数最多的前3个种的个体数占到总个体数的60%,前14个种占到95%,而其余38个种只占到5 %。从物种多度、 胸高断面积、平均胸径和重要值来看,群落成层现象显著,具有比较明显的优势种。主要树种的径级结构近似于正态分布或双峰分布,而次林层和林下层树种则表现出倒 “J" 形或 “L"形。红松、紫椴(Tilia amurensis)、蒙古栎(Quercus mongolica)、 水曲柳(Fraxinus mandshurica)、色木槭(Acer mono)和春榆(Ulmus japonica )几个主要树种的空间分布随物种、径级的变化表现出不同的分布格局,其它一些树种的分布格局也表现出一定的空间异质性。