Evaluation of secondary dispersal in a large-seeded tree Aesculus turbinata: a test of directed dispersal

Gap characteristics and gap phase replacement of major tree species were examined in two primary old-growth (mean DBHs of the canopy trees were 45.2 and 56.1 cm) and four secondary developing (range of mean DBH of the canopy trees was 23.5–39.9 cm) beech (Fagus crenata) stands in the Daisen Forest Reserve, southwestern Japan, and these were analyzed in relation to stand development as expressed by the difference of mean DBH of canopy trees. Percentage gap area (percentage of total gap area to total surveyed area) and mean and maximum gap size varied widely and ranged from 1.7 to 20.0%, from 19.4 to 162.8 m2, and from 35.7 to 585.1 m2, respectively. Mean percentage gap area and mean gap size were significantly greater in old-growth than in developing stands. However, they and maximum gap size might not increase linearly with stand development, and the gap feature of less developed stands was greater than that of later stages in developing stands. The cause was a higher formation rate, in younger developing stands, of gaps formed by simultaneous death (multiple trees falling down in domino fashion) which tends to produce larger gaps. In developing stands mean DBH of gapmakers (canopy trees that formed a gap) was smaller than that of canopy trees, though the inverse trend might be found in old-growth stands. Three typical types of death or injury states of gapmakers such as standing dead, trunk broken and uprooted were found in every study stand and the difference in stand development may not cause stand-to-stand variations for them. Importance of F. crenata (the most dominant species) in the canopy layer increased and its importance in the understory layer decreased with stand development. Shade-intolerant Quercus mongolica in the canopy layer was more important in younger than in old-growth stands, and there was no Quercus regeneration in old-growth stands. Acer mono consistently appeared, though in much less abundance than other species, in both canopy and understory layers of all study stands. Sub-canopy layers, which are mainly formed by sub-canopy tree species such as Acanthopanax sciadophylloides and Acer japonicum, may gradually develop with stand development.p>     

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.     

长白山暗针叶林林隙一般特征及干扰状况

对长白山暗针叶林林隙一般特征和干扰状况进行了研究。结果表明：长白山暗针叶林林隙的线状密度为２１．１５个／ｋｍ,扩展林隙所占的面积比例为２９．４５％,冠空隙所占的面积比例为１５．８１％;冠空隙的年干扰频率为０．２４％,干扰轮回期为４１６．７ａ左右;冠空隙的大小变化在１７．９—３４０．３ｍ＾２之间,＜１００ｍ＾２的冠空隙个数较多,冠空隙的平均面积为９３．６０ｍ＾２;扩展林隙的大小变化在４３．６—４８２．３ｍ＾２之间,５０一２００ｍ＾２之间的扩展林隙个数较多,扩展林隙的平均面积为１７４．３４ｍ＾２;暗针叶林林隙形成的主要方式是风倒;在长白山暗针叶林中,大多数林隙是由２—６株形成木形成,其中由３株形成木形成的林隙员多,单株或７株以上形成木形成的林隙数量很少;在暗针叶林中,１０一４０ａ前这段时间形成的林隙较多,特别是２０一３０ａ期间形成的林隙员多。其它阶段形成的林隙较少;暗针叶林的林隙大多是由臭冷杉、落叶松和鱼鳞云杉形成。径级在１０一３０ｃｍ之间,高度在２５—３０ｍ之间的主林层树木形成林隙的可能性最大。暗针叶林林分组成、林隙干扰方式和程度随海拔高度的变化而变化。     

Effects of disturbance and size structure on the regeneration process in a sub-boreal coniferous forest,northern Japan

The stand structure and disturbance history in a sub-boreal coniferous forest dominated byPicea jezoensis, Picea glehnii andAbies sachalinensis were investigated in four study plots set up in Taisetsuzan National Park, Japan. The effect of stand characteristics on the growth and mortality rates of understory trees was examined. Although all the stands showed inverse J-shape d.b.h. (diameter at breast height) distributions, the age structure and disturbance history differed amongst the stands. The stands with wide d.b.h. distribution (i.e. large CV and skewness) were more uneven-aged than those with narrow d.b.h. distribution (i.e. small CV and skewness). The disturbance-return interval based on the model of Hett and Loucks was 31 to 65 years. The gap ratio in the canopy was also different among the stands. These suggest that the variations in stand structure represent different occurrences of natural disturbances. Furthermore, the structural features such as size structure, canopy gap ratio and density of canopy trees also affected the growth dynamics of understory trees (≥2 m in height and <10 cm in diameter at breast height). The growth and mortality rates of understory trees changed with the canopy gap ratio and canopy tree density. The understory trees of stands with wide canopy d.b.h. distribution had higher growth and canopy recruitment rates than those of stands with narrow canopy d.b.h. distribution, contributing to the maintenance of continuous stand stratification. The understory trees of stands with narrow canopy d.b.h. distribution showed lower growth and higher mortality rates than those of stands with narrow canopy d.b.h. distribution, leading to the formation of a single-canopy structure. It is suggested that natural disturbance governs the regeneration process in the future by affecting the growth and mortality patterns of understory trees through the stand structure (size and age structure, canopy tree density, canopy gap ratio).     

Development processes and growth pattern of Pinus densiflora stands in central eastern Korea

Stand growth and developmental processes were investigated in Pinus densiflora Siebold et Zucc. stands of different ages in the central eastern region of Korea. Stands were inventoried and five trees per stand were sampled for stem analysis, age estimation, and growth analysis. More than 80% of sampled trees in a stand were established within 3–5 years, and most stands had a single cohort structure. The initial growth of pine seedlings was slow, but the height growth accelerated beyond 2–3 m height, 5–10 years after establishment. Linear growth was maintained until 10–12 m height, at which suppressed trees fell behind and might die out. The young stand was composed of pure pines, while few pine seedlings and saplings were found in the understory of older stands. The peak of diameter growth rate occurred around 5–15 years after tree establishment, implying that competition begins during that period. The pine stand development follows four stages: (1) the young stage when the growth rate increases and peaks; (2) the height competition stage when trees focus on height growth for light while maintaining a narrow DBH and height distribution; (3) the differentiation stage when suppressed trees die out, and the DBH distribution becomes wider; and (4) the mature stage when stands have a multi-canopy structure with a wide DBH and height distribution, while the understory is dominated by other tree species. The changes in growth rates and stand structure through forest development would be implemented to predict alterations of above-ground carbon sequestration rates.     

Dendroecological study of a subalpine fir (<Emphasis Type="Italic">Abies fargesii</Emphasis>) forest in the Qinling Mountains,China

Dendroecological techniques were used to investigate the stand dynamics and the disturbance history of the subalpine fir forest in the Qinling Mountains of Shaanxi Province, China. The results indicated that 68% of the fir trees experienced 1–2 release events for a total of 10–29 (an average of 15.8) years, and 1–2 suppression events for a total of 10–27 (an average of 13.4) years before they reached canopy. Large number of Abies fargesii and Betula albo-sinensis recruitment coincided temporally with larger increases in the ring-width index from the 1830s to 1880s, suggesting occurrence of a major stand-wide disturbance during this time period. Few seedlings and saplings were found in the forest, and there was a dramatic decline in recruitment after 1890, probably because of the intensive cover of understory umbrella bamboo (Fargesia spathacea). Radial growth analyses indicated frequent canopy opening resulting from small-scale disturbances in the forest. Thus, the subalpine fir forest experienced frequent small-scale disturbances and infrequent large-scale disturbances in its developmental history, and these disturbances coupled with the understory umbrella bamboo might have influenced tree growth and species recruitment.     

长白山阔叶红松林不同强度择伐后关键树种的竞争关系

天然林择伐改变了林分的树种组成和结构,导致林木竞争关系发生变化,进而影响树木的生长和种群的动态.关键树种在维持群落结构和生态系统功能方面具有至关重要的作用.为了弄清长白山区阔叶红松林典型林型关键树种竞争关系的特点,本研究以红松、紫椴和水曲柳为研究对象,运用Hegyi竞争指数分析了受不同程度采伐干扰后形成的原始林(未受干扰)、轻度择伐林、中度择伐林和重度择伐林的林分结构和竞争关系.结果表明: 与原始林相比,轻度择伐林中关键树种的种群结构没有显著变化;中度择伐林和重度择伐林中3个关键树种大树(胸径DBH≥20 cm)的密度和平均胸径均显著减少,但幼树(2 cm≤DBH<10 cm)的数量增加.所有样地中关键树种的竞争指数均随径级的增加而减小,且二者服从幂函数分布,而林木胸径生长到20 cm后,竞争指数进入稳定状态.在原始林、轻度和中度择伐林中,3个树种的幼树的竞争指数受到非冠层树种的影响最大,而红松小树(10 cm≤DBH<20 cm)和大树主要受红松种内和非冠层树种的影响;水曲柳主要受红松和非冠层树种的影响,紫椴主要受种内和红松的影响.重度择伐林中,白桦是3个树种的主要竞争来源,贡献率均超过50%.根据以上关键树种竞争关系的特点,抚育非冠层树种有利于3个关键种幼树的更新和生长;对于小树来说,还需要根据树种类型来采取相应措施,而大树不需要采用抚育措施.本研究对关键树种培育和天然林择伐后快速恢复具有指导意义.     

Gap formation and dynamics after long‐term steady state in an old‐growth Picea abies stand in Norway: Above‐ and belowground interactions

Stand dynamics and the gap initiation prior to gap formation are not well‐understood because of its long‐term nature and the scarcity of late‐successional stands. Reconstruction of such disturbance is normally based on historical records and dendroecological methods. We investigated gap initiation and formation at the fine‐scale stand level in the old‐growth reserve of Karlshaugen in Norway. Given its long‐term conservation history, and thorough mapping in permanent marked plots with spatially referenced trees, it provides an opportunity to present stand development before, during, and after gap formation. Late‐successional decline in biomass was recorded after more than 50 years of close to steady state. Gaps in the canopy were mainly created by large old trees that had been killed by spruce bark beetles. Snapping by wind was the main reason for treefall. Long‐term dominance of Norway spruce excluded downy birch and Scots pine from the stand. Comparisons of the forest floor soil properties between the gap and nongap area showed significantly higher concentrations of plant available Ca within the gap area. Plant root simulator (PRS?) probes showed significantly higher supply rates for Ca and Mg, but significantly lower K for the gap compared to the nongap area. Soil water from the gap area had significantly higher C:N ratios compared to the nongap area. Fine‐scale variation with increasing distance to logs indicated that CWD is important for leaking of DOC and Ca. Our long‐term study from Karlshaugen documents gap dynamics after more than 50 years of steady state and a multiscale disturbance regime in an old‐growth forest. The observed disturbance dynamic caused higher aboveground and belowground heterogeneity in plots, coarse woody debris, and nutrients. Our study of the nutrient levels of the forest floor suggest that natural gaps of old‐growth forest provide a long‐lasting biogeochemical feedback system particularly with respect to Ca and probably also N. Norway spruce trees near the gap edge responded with high plasticity to reduced competition, showing the importance of the edge zone as hot spots for establishing heterogeneity, but also the potential for carbon sequestration in old‐growth forest.     

玉龙雪山自然保护区丽江云杉林林窗特征研究

研究了玉龙雪山自然保护区云杉坪典型丽江云杉林林窗干扰的基本特征．结果表明,滇西北亚高山丽江云杉林林窗占林分面积比例冠层林窗和扩展林窗分别为28．8％和42．5％,干扰频率以林窗密度计算,林窗出现的平均密度为35个·hm^-2左右．林窗大小分布为面积大于100m^2的大林窗约占25％,面积为50～100m^2的中等林窗占41％,面积小于50m。的小林窗占34％．形成林窗最重要的方式是干基和干中折断,其次是枯立滇西北亚高山丽江云杉林林窗形成木80％以上是丽江云杉,通常直径为40～50cm、高度为15～25m．每个林窗形成木数量多为1～2株,约占68．8％,5株以上很少,最多为6株．随着林窗面积由大变小,林窗中更新苗木的密度逐渐变大,小林窗中更新木密度约为大林窗的5倍．     

Treefall gap disturbance in an old‐growth beech forest in southwestern Japan by a catastrophic typhoon

Abstract. In 1991, the catastrophic Typhoon 9119 created many treefall gaps in an old‐growth beech (Fagus crenata) forest on Mt. Garyu, southwestern Japan. In a 3.3‐ha plot, the density and basal area of fatally damaged trunks (DBH ≥ 20 cm) were 29.1 trunks/ha (16.9% of all pre‐typhoon trunks) and 9.1 m²/ha (24.4% of total pre‐typhoon basal area), respectively. Many of the dominant beech were lost and larger trunks were damaged. The pre‐ and post‐typhoon DBH distribution both had an inverse J‐shape. Spatial distribution of living trunks was not random but clustered irrespective of typhoon damage or DBH size, suggesting that the stand structure of the study plot would be an unusual one as compared with the previous study stands in Japanese beech forests. The death of pre‐typhoon trunks, seemed to die standing or to be killed by snap‐off, occurred singly; however, the typhoon fatally damaged many clustered trunks at the same time. Uprooting was the most frequent cause of gaps created by the typhoon. The intermediate DBH size class (40–70 cm) would be more likely to be uprooted than snapped‐off. The DBH distribution of snap‐off may reflect the total pre‐typhoon trunk distribution. Branch‐fall had a flat DBH distribution. Uprooting might have a clustered distribution, while snap‐off and branch‐fall both had random distributions. The smaller trunks of snap‐off and branch‐fall seemed to result from injuries caused by the fall of larger trunks.     

Initial establishment and regeneration processes of an outlying isolated <Emphasis Type="Italic">Fagus crenata</Emphasis> Blume forest stand in the northernmost boundary of its range in Hokkaido,northern Japan

We investigated initial establishment and regeneration of an outlying isolated Fagus crenata forest stand at the northernmost boundary of its range in Hokkaido, northern Japan. The study site was located in the Sannosuke beech forest (42°46′48″N, 140°23′43″E), a representative outlying beech stand beyond its continuous range. A rectangular 0.75 ha plot was established on a southwest-facing slope and divided into 300 square sub-plots of 25 m². Within each sub-plot, stems over 5 cm in diameter at breast height (DBH) were identified and measured. Furthermore, the location of stems over 10 cm in DBH (canopy stem) was recorded within each sub-plot, and their increment core samples were extracted. Wood from fallen logs was sampled to estimate the species composition of the coarse woody debris. Micro-relief of the plot was investigated by leveling with compasses for a 2.5 m × 2.5 m grid system. In the plot, the population of F. crenata was divided into three sub-populations by their frequency distribution of age. The oldest sub-population, over 121 years old, had been established in small-localized gap in the plot. The sub-population between 81 and 120 years old and the sub-population less than 80 years old were regenerated after a landslide and windthrow in a 1954 typhoon, respectively. Furthermore, dominant species in the plot shifted from Quercus mongolica var. grosseserrata to F. crenata. Consequently, regeneration of F. crenata, i.e., expansion of forest stands, at the northernmost boundary of its range was primarily dependent on episodic natural disturbance, which may be responsible for the reduction of their migration rate in Hokkaido, northern Japan.     

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.     

Tree mortality and effects of release from competition in an old‐growth Fagus‐Abies‐Picea stand

Abstract. In a montane mixed Fagus‐Abies‐Picea forest in Babia Gora National Park (southern Poland), the dynamics of an old‐growth stand were studied by combining an 8‐yr annual census of trees in a 1‐ha permanent sample plot with radial increments of Abies and Picea growing in the central part of the plot. The mortality among the canopy trees was relatively high (10% in 8 yr), but the basal area increment of surviving trees slightly exceeded the losses caused by tree death. DBH increment was positively correlated with initial diameter in Abies and Picea, but not in Fagus. For individual trees smaller than the median height, basal area increment was positively related to the basal area of old snags and the basal area of recently deceased trees in their neighbourhood, but negatively related to the basal area of live trees. Dendrochronological analysis of the past growth patterns revealed numerous periods of release and suppression, which were usually not synchronized among the trees within a 0.3 ha plot. The almost normal distribution of canopy tree DBH and the small number of young individuals in the plot indicated that stand dynamics were synchronized over a relatively large area and, hence, were consistent with the developmental phase concept. On the other hand, the lack of synchronization among periods of growth acceleration in individual mature Abies and Picea trees conforms more closely to the gap‐dynamics paradigm.     

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.     

Effects of population density on forest structure and species richness and diversity of trees in Kampong Thom Province, Cambodia

This study examined differences in stand structure, tree species richness, and tree species diversity in relation to population density in Kampong Thom Province, Cambodia. Tree data were obtained from a 1997 forest inventory involving 60 clusters (540 plots) systematically distributed over 30% of the provincial forest area. Spatially referenced population data were obtained from the 1998 national population census. The average number of trees per cluster was 356/ha, the average basal area, 23 m²/ha, the average stand volume, 217 m³/ha, and the average aboveground biomass, 273 Mg/ha for all trees of DBH 10 cm and larger. The average species richness per cluster was 37 species, while average species diversity was measured as 0.916 using Simpson’s index and 2.98 by Shannon’s index. Significant negative correlations were generally found between population density surrounding clusters and tree density, basal area, stand volume, aboveground biomass, and species richness and diversity for three examined diameter classes (DBH of 10–30, ≥30, and ≥10 cm). As the distance from clusters for calculating population density increased, the correlation levels increased up to 5 or 7 km, depending on the variables and diameter class, and then stayed relatively constant for stand structure variables and decreased for species richness and diversity. The results indicate that evidence of disturbance was more pronounced at higher population density up to around 5 to 7 km. We suggest that introduction of greater controls on human disturbance should be a high priority for resource management and conservation in Kampong Thom Province and, presumably, Cambodia as a whole.     

Gap characteristics and gap regeneration in a subalpine coniferous forest on Mt Ontake,central Honshu,Japan

Gap characteristics and gap regeneration were studied in three mature stands belonging to different community types in a subalpine coniferous forest on Mt Ontake, central Honshu, Japan. Gap disturbance regimes were remarkably similar among stands studied; percentage gap area to surveyed area, gap density and mean gap size were 7.3–8.5%, 17.8–20.0 ha⁻¹ and 40.8–42.5 m², respectively. The gap size class distributions were also similar and showed a strong positive skewness with a few large and many small gaps; gaps <40m² were most frequent and gaps >200m² were rare. Forty-five to 66% of gaps were due to the death of single canopy trees. Canopy trees more often died leaving standing dead wood (40–5.7%) or broken trunks (43–49%). Shade tolerantAbies mariesii andAbies veitchii, frequently, andTsuga diversifolia, less frequently, regenerated in gaps, from advance regenerations recruited before gap formation.Picea jezoensis var.hondoensis may regenerate in gaps, from new individuals recruited after gap formation. The breakage of denseSasa coverage and the mineral soils exposed by the uprooted plants that form gaps might provide regeneration opportunities for shade intolerantBetula. Of the important species limited to the ridge site,Chamaecyparis obtusa, frequently, andThuja standishii, infrequently, regenerated from plants alreadyin situ. Regeneration ofPinus parviflora was not seen. Based on the gap characteristics and gap regeneration behaviour of each species described, stand dynamics in each stand are discussed.     

Growth and mortality of trees in Amazonian tropical rain forest in Ecuador

Abstract. Structural changes are analysed in four samples representing 4 ha, two line transects and two hectare plots, of Amazonian tropical lowland rain forest in northern Ecuador. Only trees with a DBH ≥ 10 cm were included. A sample of floodplain forest in Añangu represents the largest turnover found in tropical forests (stand half-life = 23 yr). The line transect and hectare plot both of tierra firme forest in Añangu have the same turnover (37 yr) and were balanced for death and in-growth of both individuals and wood (basal area). The 1-ha tierra firme sample in Cuyabeno had a turnover of 67 yr and was in a growing phase. The floodplain line transect in Añangu was in a phase of structural breakdown. However, the floodplain line transect had the largest growth of basal area per tree (23.4 cm²/yr). The tierra firme samples had a growth of 9.6, 10.1, and 13.6 cm²/yr. Most of the dead trees fell with some uprooting in three of the four samples. However, no significant difference in the distribution of mode of death was found between the four samples. Death was independent of topography and the dead trees were randomly distributed. As the trees grow up they occupy more space and larger trees (DBH ≥ 15 cm) become more uniformly distributed, whereas smaller trees (DBH ≤ 15 cm) were randomly distributed. Our study confirms that plots of 1 ha are not sufficient to include representative samples of different stages of forest structure.     

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.     

Establishment of the evergreen broad-leaved tree species Castanopsis cuspidata in an abandoned secondary forest in western Japan

Recently, populations of Castanopsis cuspidata have often expanded into secondary forests in western Japan. To determine the establishment processes of this species, we analyzed its spatial distribution in a secondary forest dominated by Quercus variabilis and Quercus serrata that is located adjacent to a stand dominated by C. cuspidata. Saplings, defined as ≥30 cm stem length (SL) and <5 cm diameter at breast height (DBH), were abundant and their size distribution was inversely J-shaped, indicating continuous recruitment. Although seedlings (SL < 30 cm) and small saplings (30 ≤ SL < 50 cm) of C. cuspidata were aggregated near flowering trees of this species, some were found ≥40 m from the nearest adults, suggesting that there is animal-aided dispersal of acorns. The distribution of larger-sized individuals (≥100 cm SL) of C. cuspidata was unrelated to distance from the nearest flowering C. cuspidata or dominant Quercus species (≥5 cm DBH), but was associated with dead Pinus densiflora trees, which were abundant at the site. Thus, the establishment of C. cuspidata in this forest is controlled mainly by two factors, viz. patterns of acorn dispersal by animals, and forest disturbance regime (i.e., deaths of pine trees).     

秦岭优势乔木锐齿槲栎的空间分布格局及种内关联

为探究锐齿槲栎种群的空间分布特征及关联性,本研究以秦岭皇冠暖温性落叶阔叶林25 hm²森林样地内优势树种锐齿槲栎为对象,采用成对相关函数g(r)对其空间格局及其种内关联性进行了研究。结果表明: 锐齿槲栎径级结构呈“双峰”型,幼树(1 cm≤胸径DBH＜5 cm)个体数较多,种群呈增长型结构,更新良好;中树(15 cm≤DBH＜25 cm)比大树(25 cm≤DBH＜35 cm)、老树(DBH≥35 cm)的个体数略多,但远少于幼树和小树(5 cm≤DBH＜15 cm)。锐齿槲栎空间分布具有明显的海拔依赖性,主要分布在中高海拔地区。完全空间随机零模型分析表明,各径级个体在＜60 m的大尺度范围聚集分布。使用异质泊松模型剔除生境异质性分析表明,各径级的个体转变为大尺度的随机分布,表明树种的分布明显受到生境变化的影响。在＜40 m的小尺度范围内,径级差距小的个体间的空间关联性为正关联,径级差距大的个体间的空间关联性转变为负关联和无关联;在＞40 m的大尺度范围内,大径级个体间的空间关联性为正关联,而幼树和其他径级个体间的空间关联性为负关联或无关联。锐齿槲栎自身生物学特性和环境异质性是种群空间格局形成的重要原因。