长白山阔叶红松林稠李居群遗传结构的空间自相关分析

以长白山阔叶红松林25 hm²样地为平台,选取样地中胸径≥1 cm的稠李个体共计396株,将其分为1～3、3～10和>10 cm 3个径级,通过微卫星标记对这些个体进行了空间遗传结构的空间自相关分析.结果表明: 样地中的稠李种群总体上具有一定的空间特征,在70 m范围内呈显著正相关, 110 m外则呈负相关.不同年龄级的个体空间遗传结构相似, 且主要源于有限的花粉和种子扩散距离.该稠李居群未出现明显的自疏现象.     

Ecological characteristics of Abies georgei population at timberline on the north-facing slope of Baima Snow Mountain, Southwest China

Abies georgei is one of the endemic trees at timberline on Qinghai-Tibetan Plateau, but little research has been done about its timberline population. One plot was set up at timberline above 4400 m on the north-facing slope of Baima Snow Mountain in Southwest China, covering an area of 0.8 hm2. Height, DBH (diameter at breast high) or root-collar diameter of seedlings and saplings, and position were recorded for each individual. The population showed a growing trend and the number of individuals decreased with increase of DBH. The survival curve can be described by Deevy-III. The population had 2 distinct peaks of mortality: age I (seedlings, DBH = 0–5 cm) and age V-VI (DBH = 20–30 cm). The highest mortality of seedlings was due to severe climate at timberline. As the saplings grew up, the interspecific and intraspecific competition resulted in another mortality peak. Through SPPA (spatial point pattern analysis), individuals of all age-classes showed uniform, random or clumped distribution with the change of scales; however, they had mainly clumped distribution. Moreover, the relationships between individuals in different age-classes were all significantly correlated with each other. Seedlings showed the highest clumping intensity and scale.     

白马雪山阴坡林线长苞冷杉(Abies georgei)种群结构特征

长苞冷杉（Abies georgei）是青藏高原特有林线树种。对白马雪山阴坡海拔4 400 m以上林线0.8 hm2样地长苞冷杉种群进行每木调查,分析其种群结构、数量特征及空间分布格局。结果表明：(1）结构呈典型金字塔型,幼苗和幼树在种群中所占比重大,表现为增长型种群,种群个体数随径级的增加而减少,密度为幼苗>幼树>成年树;(2）存活曲线接近Deevy-III型,高径级种群趋于稳定,种群具两个死亡高峰,低径级种群尤其幼苗死亡率高达90％,这是林线区的气候条件如低温、强光照、积雪及冬季冻害等综合作用的结果,种群另一死亡高峰出现于V~VI龄级,种内和种间对空间、光照和养分等生存因子的激烈竞争引起自疏,导致死亡率再度上升;(3）长苞冷杉种群各龄级空间点格局在不同尺度上表现为聚集、随机和均匀分布,以聚集分布为主,由于幼苗来源于种子库且依赖成年树的微生境,幼苗聚集强度和尺度都最大;各龄级关系密切,在不同尺度上表现出显著的相关性。     

水杉人工林建植50年后的分化特征

通过种群统计学、格局分析和树干解析等对50 a生水杉人工林林分的分化特征进行了研究.结果表明:小个体林木占有相对多数(>50.4%);林木分化现象显著,特别是树高与材积的分化十分突出,林分已出现一定强度的自然稀疏;基于TSTRAT程式的林木分级结果表明,亚优势木占绝对优势(45.7%),其树冠构成该林分的主林冠;竞争指数依次为优势木级<亚优势木级<中庸木级<被压木级,各级的基尼系数基本上低于全林总计;优势木绝对生长率和相对生长率在林分成熟期以前均大于标准木,且胸径速生期持续时问较标准木长;林分高度生长受压抑程度低于胸径生长;林木分布格局总体呈均匀型,与造林初始基本一致;优势木和被压木为聚集分布,中庸木和枯死木为随机散布,亚优势木的分布型则介于随机和聚集之间.林木种群分化既表现在个体大小的不均等性上,也表现在空间分布的非匀质性上.目前林分结构特征的主要成因可以归结为密度压力下的种内竞争.     

Structural dynamics of a natural mixed deciduous forest in western Thailand

Abstract. Structural dynamics of a natural tropical seasonal – mixed deciduous – forest were studied over a 4-yr period at Mae Klong Watershed Research Station, Kanchanaburi Province, western Thailand, with particular reference to the role of forest fires and undergrowth bamboos. All trees > 5 cm DBH in a permanent plot of 200 m × 200 m were censused every two years from 1992 to 1996. The forest was characterized by a low stem density and basal area and relatively high species diversity. Both the bamboo undergrowth and frequent forest fires could be dominant factors that prevent continuous regeneration. Recruitment, mortality, gain (growth of survival tree plus ingrowth) and loss in basal area (by tree death) during the four years were 6.70%/yr, 2.91%/yr, 1.22%/yr and 1.34%/yr, respectively. Mortality was size dependent; middle size trees (30–50 cm) had the lowest mortality, while the smallest (5–10 cm) had the highest mortality. Tree recruitment was observed particularly in the first two years, mostly in the area where die-back of undergrowth bamboo occurred. The bias of the spatial distribution of recruitment to the area of bamboo die-back was significant and stronger than that to the forest canopy gaps. Successful regeneration of trees which survive competition with other herbs and trees after dieback of bamboo could occur when repeated forest fires did not occur in subsequent years. It is suggested that both the fire disturbance regime and bamboo life-cycle greatly influence the structure and dynamics of this seasonal tropical forest.     

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。     

Intraspecific Competition in a Natural Stand of Betula ermanii

Density and size of naturally regenerated Betula ermanii weremonitored for six years in a 100 m² plot located in Hokkaido,northern Japan. Natural thinning of 10- to 15-year-old treesoccurred at a negative exponential rate related to tree volume,demonstrating the –3/2 power rule of self-thinning. Diameterdistributions were unimodal at ages 10–13 years and bimodalat 14 and 15 years. Changes in diameter distribution were accompaniedby changes in structure, from a one- to two-storied stand. Mortalitywas highest among smaller trees and where density was greatest.The spatial distribution of trees changed from random to uniformwith increased stand age. Betula ermanii Cham., Self-thinning, mortality, spatial distribution, size distribution     

后河自然保护区珍稀植物群落主要树种6年动态变化

用每木定位监测方法分析了后河自然保护区常绿落叶阔叶混交林1 hm2固定样地主要乔木树种6年的（2001～2007年）动态变化。样地主要分布有14种普通树种和5种珍稀树种。对这19个树种胸径≥5 cm的林木动态进行分析。2001年,样地有998株主要树种,到2007年,其中100株死亡,样地中新增81株更新幼树。19个主要树种基盖度从19.56 m2/hm2增加到20.14 m2/hm2,群落的径级结构没有明显变化,中等径级个体死亡率最高,且死亡植株在小尺度上聚集分布,较大尺度上随机分布,更新幼树几乎在所有尺度聚集分布,活树和死树在所有尺度空间正关联,活树和更新幼树在大部分尺度负关联。稀有濒危植物能长期稳定地在群落中存在,表明干扰、种内和种间竞争共同影响着群落的发展,群落处于动态平衡状态。     

<Emphasis Type="Italic">PRUNUS</Emphasis>: a spatially explicit demographic model to study plant invasions in stochastic,heterogeneous environments

To model the invasion of Prunus serotina invasion within a real forest landscape we built a spatially explicit, non-linear Markov chain which incorporated a stage-structured population matrix and dispersal functions. Sensitivity analyses were subsequently conducted to identify key processes controlling the spatial spread of the invader, testing the hypothesis that the landscape invasion patterns are driven in the most part by disturbance patterns, local demographical processes controlling propagule pressure, habitat suitability, and long-distance dispersal. When offspring emigration was considered as a density-dependent phenomenon, local demographic factors generated invasion patterns at larger spatial scales through three factors: adult longevity; adult fecundity; and the intensity of self-thinning during stand development. Three other factors acted at the landscape scale: habitat quality, which determined the proportion of the landscape mosaic which was potentially invasible; disturbances, which determined when suitable habitats became temporarily invasible; and the existence of long distance dispersal events, which determined how far from the existing source populations new founder populations could be created. As a flexible “all-in-one” model, PRUNUS offers perspectives for generalization to other plant invasions, and the study of interactions between key processes at multiple spatial scales.     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

Mortality and Self-thinning in Postfire Black Spruce

Using age-structure determinations on both living and dead stemsin censused plots, coupled with stem analysis techniques, anhistorical picture of mortality and above-ground tree stem growthwas recreated for ten stands dominated by black spruce in northeasternOntario, Canada. No evidence of mortality was seen in any plot prior to 30 yearsfollowing postfire initiation. Each of the eight oldest standsshowed a linear decline in numbers for a 20–25 year period.The steepness of the mortality slope was proportional to initiallive stem density within and among plots during this phase.The final 10–20 years was marked by a less steep declinein numbers. The log density vs log mean tree volume curves in the eightoldest stands were doubly asymptotic and were fitted to a logisticcurve very tightly in each case. At the point of inflectionthe curves' slopes ranged from –2.14 to –3.89. However,log density vs log mean stem volume among stands at this pointof inflection had a slope of –0.96. Reasons for the inconsistency between within-stand and among-standself-thinning estimates are considered, as well as the poorfit to the –3/2 rule. Ecosystem processes related to thechange in nutrient relations during stand growth are identifiedas a prime influence on self-thinning behaviour in natural blackspruce stands. Mortality, stem analysis, self-thinning, Picea mariana, black spruce     

Long-term dynamics of vegetation and disturbance of a southern boreal spruce swamp forest

Abstract. Analysis of pollen, charcoal and loss-on-ignition in peat cores from a Picea aèies-dominated swamp forest in central Sweden show the vegetation changes and disturbance patterns over 9500 yr. Six major sequences of local vegetation development are identified: (A) Pinus period, ca. 9500–7000 cal. BP; (B) Open mire period (ca. 7000–4500 cal. BP; (C) Betula period, ca. 4500–2300 cal. BP; (D) Picea period (ca. 2300–1000 cal. BP; (E) Human impact period (ca. 1000–100 cal. BP); and (F) Period of human abandonment during the last ca. 100 yr. The swamp forest has been highly dynamic in response to various natural and anthropogenic disturbance agencies. Several fires have heavily influenced the vegetation development. During the last ca. 900 yr human influence has been important, initially from grazing and trampling by domesticated animals (ca. 1000–500 cal. BP), and subsequently small-scale cereal growing (ca. 400–100 BP). Cutting, burning and animal browsing influenced the structure and dynamics of the swamp forest by creating a more open stand and suppressing tree regeneration. Recent cessation of human impact has led to increased tree regeneration and a denser swamp forest stand. The present high biodiversity, and subsequent conservation interest does not result from long-term stability or absence of fire and human impact. However, in spite of repeated disturbances, a continuity of old and senescent trees produced a forest type with abundant dead wood. With the relatively minor importance of fire over long periods of time, the swamp forest developed a structure maintaining a high biological diversity. An important issue for maintaining long-term biodiversity in the boreal landscape must be to create a mosaic where different forest types are present, with a variety of structures, substrates and processes, to provide a certain degree of freedom for species to move around in the landscape.     

Population structure and spatial pattern of predominant tree species in a pine–oak mosaic mixed forest in the Qinling Mountains,China

This study investigated a typical pine-oak mosaic mixed forest in the Qinling Mountains, China. In the sample plot, the population structure and spatial distribution of the stems were analyzed for the predominant species, to identify the mechanisms of species coexistence and successional trends of the forest. The population structures of all species were bimodally distributed, with young trees (DBH <1 cm) more abundant than older trees. The population structures of Quercus aliena var. acuteserrata was bimodal and rather continuous. However, Pinus tabuliformis and Pinus armandii were discontinuously bimodal, with distinct size deficiencies. Q. aliena var. acuteserrata trees were clumped throughout the plot, although those of P. tabuliformis and P. armandii were clumped at small scales. Notable negative spatial associations between Q. aliena var. acuteserrata and P. tabuliformis were found at almost scales. P. armandii and Q. aliena var. acuteserrata were negatively spatially associated at small scales but positively associated at large scales. Our findings suggest that interspecific competition gradually develops among the predominant tree species. The dynamics of the pine-oak mosaic mixed forest formed a mosaic distribution of uniformly mixed types, with the slow infiltration of Q. aliena var. acuteserrata populations that would eventually establish a pure stand.     

SPATIAL DYNAMICS OF A HIMANTHALIA ELONGATA (FUCALES, PHAEOPHYTA) POPULATION

In dense monospecific stands of plants intraspecific competition usually results in self-thinning, the concurrent increase in biomass and decrease in density over time. Self-thinning may also result in a change in the spatial pattern of individuals, but so far the spatial dynamics of marine plants has not been investigated. The brown alga Himanthalia elongata ( L.) S. F. Gray forms dense monospecific stands on many northern temperate rocky shores, and various attributes (including its simple form) facilitated the study of the spatial dynamics of this species .
The spatial pattern of settling zygotes was examined in the laboratory. In the absence of water movement, substratum heterogeneity, and a point source, zygotes usually settled in clumps rather than randomly. Within the clumps zygotes appeared to be regularly distributed at a scale similar to the size of the zygotes themselves. Furthermore, the clumps themselves seemed to be regularly distributed. On the shore, well-established stands of "button-stage" Himanthalia populations were examined during a period of extensive growth and self-thinning. Individual plants were initially highly regular in spatial pattern but became less so over time. The pattern of plants dying during self-thinning was also highly regular and probably reflected existing spatial regularity. However, using a hypothesis of mortality as a random event, I found that smaller plants had a less than average survival potential, while larger plants had a greater than average chance. A consideration of the spatial pattern of plants alive at the end of the study revealed regularity at a scale of 2–7 mm but a random spatial pattern at larger scales, which might indicate a small sphere of influence of competing individuals. The best predictor of mean nearest neighbor distances in the populations was mean plant diameter .     

The effects of density-dependent offspring mortality on the synchrony of reproduction

Mortality rates often depend on the size of a population. Using ideal free theory to model the optimal timing of reproduction in model populations, I considered how the specific relationship between density-dependent offspring mortality and population size affects the optimal temporal distribution of reproduction. The results suggest that the specific form of the relationship between density-dependent mortality and the number of offspring produced determines the degree to which reproduction within a population is synchronous. Specifically, reproductive synchrony decreases as density-dependent mortality becomes increasingly inversely related to the number of offspring produced and is highest when density-dependent mortality is directly density-dependent. These findings support the suggestion that predation pressure selects for greater reproductive synchrony in species where mortality is directly density-dependent, but does not affect the timing of reproduction in species with density-independent rates of mortality. This revised version was published online in July 2006 with corrections to the Cover Date.     

Population structure and spatial patterns for trees in a temperate old-growth evergreen broad-leaved forest in Japan

The population structure and spatial pattern of major tree species in a warm-temperate old-growth evergreen broad-leaved forest in the Tatera Forest Reserve of Japan were investigated. All stems 5 cm in diameter at breast height (DBH) were mapped on a 4 ha plot and analyses were made of population structure and the spatial distribution and spatial association of stems in different vertical layers for nine species. This was done in the context of scale dependency. The plot was located on a very gentle slope and 17.1% of its canopy layer was in gaps. It contained 45 woody plant species and 4570 living stems with a basal area of 63.9 m² ha^–1. Castanopsis cuspidata var. sieboldii, the most dominant species for the basal area, had the maximum DBH among the species present, fewer smaller stems and a lower coefficient of statistical skewness of the DBH distribution. The second most dominant species, Dystilium racemosum, had the highest stem density (410 ha^–1), more abundant smaller stems and a relatively higher coefficient of skewness. Most stems in different vertical layers showed a weakly aggregated distribution with loose colonies as basic units. Gap dependency for the occurrence of stems under the canopy layer was weak. Maximum slope degree of the plot also weakly affected the occurrence of stems. Spatial associations varied among intra- and interspecific cohorts in the different layers and spatial scales examined, and positive associations among cohorts were found more frequently as the scales examined became larger. This tendency suggests that key factors forming observed spatial associations might vary with the spatial scales.     

基于8hm2样地的天山云杉林蒸腾耗水从单株到林分的转换

为准确地估算天山北坡中部雪岭云杉森林的林分蒸腾耗水量,本研究设置了8hm2固定样地,在每木调查的基础上,通过对7株标准木连续晴天液流的观测,分别得到胸径与液流速率和边材面积的拟合方程,最终计算得到了林分的蒸腾耗水量,确定了估算该森林群落蒸腾耗水的最小取样面积,并且讨论了最小取样面积随样地位置的变化。结果表明：①在自然生长系,雪岭云杉林分最大蒸腾耗水量、平均蒸腾耗水量同植株密度呈Extrem函数分布;当种植密度达到1200株/hm2时,林分最大蒸腾耗水量达到最大,为77.27 t/hm2d;②对林分蒸腾耗水量的测算存在明显的最小取样面积,最小取样面积随取样点的不同而略有差异,取样起始点在本样地的海拔上边界和下边界处时最小取样面积为4 hm2,取样起始点在本样地的中部时最小取样面积为2.56hm2。本研究为更精确地估算天山北坡雪岭云杉森林的蒸腾耗水量提供了理论依据。     

Dispersion pattern and mortality of seeds and seedlings ofFagus crenata Blume in a cool temperate forest in western Japan

The seed and seedling mortality ofFagus crenata Blume after a mast year (1993) was examined in relation to density and distance from the nearest conspecific adult tree in a mixed conifer-hardwood forest in Ohdaigahara, western Japan. The mortality of fallen seeds during winter amounted to 93.7%, and 79.2% of the current-year seedlings died in the first growing season. The most important factor of death for both seeds and seedlings was predation by vertebrates. The mortality of seeds during winter was positively correlated with sound seed density. The mortality of seedlings was positively correlated with density but not significantly related to the distance from the nearest crown edge of a conspecific adult tree. Mortality patterns varied with stages and spatial scales due to the behavior of predators; it is thus important to investigate the spatial pattern of seeds and seedling mortality at various temporal and spatial scales. After the first growing season, the difference in seedling density between distance classes was not significant at <4m from the nearest adult trees due to density-dependent mortality. However, seedling density was significantly lower in the ≥4 m class than in the <4 m classes.     

Crowding in clonal seaweeds: Does self-thinning occur in Mastocarpus papillatus shortly before stand biomass peaks?

Fronds from crowded stands of clonal seaweeds, particularly those in which holdfasts are mostly perennial and are the major source of new fronds every year, are thought not to undergo self-thinning during the growth season, unlike those from crowded stands of unitary seaweeds. For clonal seaweeds, it is not known, however, what happens at the very end of the growth season, when crowding is highest for the year. By sampling twice more frequently than previously done for similar species, the possible occurrence of frond self-thinning was tested for Mastocarpus papillatus (Rhodophyta, Gigartinales, Petrocelidaceae) from western Canada during the growth season (spring) of 2003. Initially, stand biomass increased together with frond density, as found previously for similar clonal seaweeds. Shortly before stand biomass peaked for the year (June), frond density remained statistically unchanged. Thus, the increased sampling precision of this study confirms that fronds of these clonal seaweeds do not undergo self-thinning, not even shortly before crowding is highest. Frond size inequality for M. papillatus remained statistically similar during the growth season, which is also consistent with a model of no self-thinning. There are similarities in biomass–density dynamics and in size inequality dynamics between clonal seaweeds and clonal vascular plants.     

Long-term growth dynamics of natural forests in Hokkaido,northern Japan

Abstract. The long-term growth dynamics of natural forest stands on the island of Hokkaido were described on the basis of an analysis of data from 38 permanent plots spanning 15–22 yr. Stand structure was characterized by basal area, stem density and tree size variability. To detect trends in stand structure, regression models for recruitment rate (per ha per yr), mortality rate and the rate of change in stem density and tree size variability were developed by a stepwise method using initial basal area, stem density, tree size variability, species composition summarized by LNMDS ordination, altitude, annual mean temperature, annual precipitation, type of understorey vegetation, topography and slope aspect as candidates for predictor variables. The same analyses were conducted for basal area increment (net growth) and its components: survivor growth = basal area gain by growth of surviving individuals and mortality = basal area loss by death of individuals. Stem density remained generally unchanged; recruitment was relatively low even in very sparse stands. Stand basal area generally increased as survivor growth was approximately double the mortality. Recruitment rate was strongly affected by the presence of dwarf bamboo (Sasa spp.) vegetation on the forest floor which inhibited tree regeneration. Mortality rate was density-dependent; dense stands had higher mortality than sparse stands. Density change rate (recruitment rate - mortality rate) was, therefore, determined by both the type of understorey vegetation and stem density. Survivor growth was high in stands with high stem density and basal area. Mortality was dependent on basal area and altitude. Net basal area increment (net growth) was dependent only on stem density with other factors that influenced survivor growth and mortality omitted. Tree size variability decreased in stands with high tree size variability whereas it increased in stands with low size variability. Based on the obtained models for density change rate and net basal area increment, trajectories of stands were illustrated on a log-log diagram of stem density and basal area. The predicted differences in trajectories as affected by the understorey vegetation type indicated the importance of dwarf bamboo vegetation for forest dynamics on Hokkaido.