Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species corre-spond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

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.     

尖峰岭热带山地雨林优势树种白颜树空间分布格局

优势树种对群落的构建和形成起主要作用。该文以海南岛尖峰岭60 hm²样地内重要值第二大的树种白颜树(Gironniera subaequalis)为研究对象, 探索其种群分布格局特征, 讨论环境异质性、密度依赖死亡、扩散限制等机制在格局形成过程中所起的作用。将白颜树10022个植株分为6个径级, 分别归属于幼树、中龄树、成年树三个生活史阶段, 采用成对相关函数分析各径级的空间分布特征; 双变量成对相关函数和标签关联函数分析不同生活史阶段之间的空间关系; Berman检验方法检验3个地形因子分别对幼树、中龄树、成年树分布影响的显著度。结果表明: 白颜树种群内I、II、III、IV径级呈现聚集分布, 聚集程度随径级的增加减弱; V和VI径级小尺度上均匀分布, 大尺度上以随机分布为主。幼树与中龄树空间正关联; 幼树与成年树空间负关联; 成年树与中龄树在较小的尺度上负关联, 大尺度上微弱正关联。但是不同生活史阶段的个体之间彼此分离, 个体间无直接的促进作用。地形因子中, 坡度、海拔、凹凸度对幼树的分布影响显著; 坡度、凹凸度对中龄树的分布影响显著; 仅坡度对成年树的分布影响显著。从现有的空间格局可以推断出环境异质性和密度依赖死亡对格局形成起作用, 但是种子的扩散限制对空间格局的影响没有明确地表现出来。     

中亚热带喀斯特常绿落叶阔叶林空间结构特征

喀斯特常绿落叶阔叶林是中国西南喀斯特地区非地带性顶极森林群落,该研究基于相邻木关系的角尺度、混交度和大小比数3个结构参数,对木论国家级自然保护区常绿落叶阔叶林25 hm^(2)大型固定监测样地整体、上木层、下木层及其优势物种的空间结构特征进行了分析,以期揭示其森林空间结构现状,为喀斯特植被恢复重建提供依据。结果表明:(1)样地群落整体为轻度聚集分布格局,接近随机分布,物种高度混交且个体间大小分化程度相近,上木层呈现随机分布、强度混交和中庸偏亚优势状态,下木层呈现聚集分布、强度混交和中庸偏劣势状态。(2)样地内优势物种大多处于轻度聚集分布、强度混交且中庸偏劣势的状态。(3)林木大小比数与胸径、树高均呈显著负相关关系,混交度与胸径、树高均呈显著正相关关系,随林木的生长发育,树种优势度逐渐增大,物种多样性逐渐增强。研究认为,木论常绿落叶阔叶林空间结构尚未完全稳定,正处于演替中后期,具有发育成顶极群落的潜力。     

Aggregated spatial distributions of species in a subtropical karst forest,southwestern China

Aims Spatial distribution patterns of species reflect not only the ecological processes but also the habitat features that are related to species distribution. In karst topography, species distribution patterns provide more specific information about their environments. The objectives of this study are as follows: (i) to analyse and explain the spatial distribution patterns of conspecific trees in an old-growth subtropical karst forest; (ii) to investigate pattern changes at different spatial scales; (iii) to test the spatial pattern similarity (or dissimilarity) between trees at different abundances, diameter at breast height classes, canopy layers and different functional groups (shade tolerance and seed dispersal mode); (iv) to examine whether habitat heterogeneity has an important effect on the species spatial distribution.Methods The spatial distributions of woody species with ≥20 individuals in a 1-ha subtropical karst forest plot at Maolan in southwestern China were quantified using the relative neighbourhood density Ω based on the average density of conspecific species in a circular neighbourhood around each species.Important findings Aggregated distribution is the dominant pattern in the karst forest, but the ratio of aggregated species in total species number decreases with an increase in spatial scale. Less abundant species are more aggregated than most abundant species. Aggregation is weaker in larger diameter classes, which is consistent with the prediction of self-thinning. Seed dispersal mode influences spatial patterns, with species dispersed by animals being less aggregated than those dispersed by wind and gravity. Other species functional traits (e.g. shade tolerance) also influence the species spatial distributions. Moreover, differences among species habitat associations, e.g. with rocky outcrops, play a significant role in species spatial distributions. These results indicate that habitat heterogeneity, seed dispersal limitation and self-thinning primarily contribute to the species spatial distributions in this subtropical karst forest.     

基于无人机的昆明海口林场宽地坝林区植物群落建群种格局研究

建群种是植物群落优势层中的主要物种,对群落环境有显著控制作用。空间格局是植物种群的基本特征之一,探讨建群种的空间格局是理解生物与生境之间关系和认识群落生态过程的有效途径。空间格局特征包括种群空间关联性及其空间分布型。利用无人机技术,获取昆明海口林场半湿润常绿阔叶林和落叶栎类林的多光谱遥感影像,采用ArcGIS Pro软件进行遥感影像分类并提取滇青冈(Cyclobalanopsis glaucoides)、光叶石栎(Lithocarpus mairei)、滇石栎(L.dealbatus)、栓皮栎(Quercus variabilis)、槲栎(Q.aliena)植株树冠中心点空间坐标。在Programita Febrero 2014软件中进点格局分析。结果表明:各物种10m以下呈均匀分布或随机,随尺度增加,渐渐表现为聚集分布。光叶石栎种群和滇石栎种群空间关联性表现为负相关。     

江西九连山亚热带常绿阔叶林优势种空间分布格局

于江西九连山国家自然保护区设置4 hm2亚热带常绿阔叶林固定监测样地,用样方法对亚热带常绿阔叶林进行调查分析。应用点格局方法分析生境异质性是否影响树木分布并分析优势种红钩栲(Castanopsis lamontii)、米槠(Castanopsis carlesii)、罗浮柿(Diospyros morrisiana)和细枝柃(Eurya loquaiana)分布格局,对比存在生境异质性和排除生境异质性两种情况下优势树种不同生长阶段(幼树、小树、成年树)的空间分布格局以及不同生长阶段之间的空间关联性,探讨生境异质性之外的其它种群分布影响因子。结果表明:1)生境异质性效应显著影响九连山树木分布,4个优势树种在大尺度上存在明显的生境偏好;2)用完全随机零模型不排除生境异质性时,4个优势种总体及其不同生长阶段在0—30 m所有尺度上主要呈现聚集分布;3)用异质性随机零模型排除生境异质性后,4个优势种及其不同生长阶段的聚集程度显著下降,只在小尺度上(0—5 m)呈现聚集分布;4)4个优势种的幼树与小树均表现显著的正相关。红钩栲的成年树与幼树及成年树与小树总体表现不相关。米槠的成年树与幼树总体以无相关为主;成年树与小树之间总体呈正相关。小乔木罗浮柿的成年树与幼树之间在小尺度上(0—5 m)表现负相关或无相关,在较大尺度范围上总体表现正相关关系;小树与成年树在研究尺度上表现出正相关。灌木细枝柃的成年树与幼树,成年树与小树及小树与幼树之间在研究尺度范围内均呈现正相关关系;5)研究发现九连山优势乔木物种通过密度制约和Janzen-Connell效应释放空间,为其他物种共存提供条件,而优势灌木物种细枝柃没有表现出这两个效应。     

天目山常绿阔叶林空间结构动态变化特征

采用固定样地重复监测的方法,2005-2015年每5年1次对天目山国家级自然保护区内1hm²常绿阔叶林固定样地进行3次样地调查,按优势度分析法确定群落优势树种,根据树种类别将优势树种分为常绿阔叶树种、常绿针叶树种和落叶阔叶树种3个优势树种组,利用聚集指数、树种多样性混交度和Hegyi竞争指数3个空间结构指数,分析群落和优势树种在2005-2015年期间的空间结构动态变化特征。结果表明：（1）群落表现为低度聚集、中度混交和中度竞争状态。群落聚集程度和竞争强度降低,混交程度保持稳定。群落空间结构动态变化总体特征是混交度稳定性最高,其次是分布格局,竞争态势稳定性最低。（2）群落优势种组成变化不大,3个优势树种组均呈低度聚集分布;常绿阔叶树种表现为低度混交、中度竞争状态;常绿针叶和落叶阔叶树种均表现为中度混交、低度竞争状态。常绿阔叶树种的聚集程度和竞争强度降低,混交程度保持稳定,与群落空间结构动态变化特征基本一致。常绿针叶和落叶阔叶树种的3种空间结构均保持稳定。在常绿阔叶林经营或人工林近自然改造过程中,应重视群落和优势树种的空间结构变化规律,确定阶段性空间结构优化目标,逐步促进群落进展演替。     

海南岛霸王岭热带低地雨林树木的空间格局

树木空间格局及其形成过程是物种共存及生物多样性维持机制研究的一个重要方面。该文以海南岛两个1 hm ²的典型热带低地雨林老龄林森林动态样地为基础, 通过4个点格局模型(均质Poisson过程、异质Poisson过程、均质Thomas过程和异质Thomas过程)模拟扩散限制和生境异质性作用对树木空间分布格局的影响, 并分析不同空间尺度下(< 2 m, 2-5 m, 5-10 m, 10-15 m, 15-20 m和20-25 m)不同作用的相对重要性。结果表明: 热带低地雨林的所有树木总体上呈现聚集分布的空间格局, 随着尺度的增大, 聚集强度逐渐减小。树种在模拟空间分布格局最优模型中的比例由高到低分别是: 均质Thomas过程, 均质Poisson过程、异质Thomas过程和异质Poisson过程。扩散限制作用是形成热带低地雨林树木空间分布格局最重要的生态过程, 其次是完全随机作用以及生境异质性和扩散限制的联合作用, 而生境异质性的作用最小。不同空间尺度上模拟各树种空间分布格局的最优模型比例差异显著, 扩散限制作用能够在多数空间尺度上模拟多个树种的空间分布格局, 其次为随机作用; 生境异质性和扩散限制的联合作用主要在小尺度(0-5 m)影响树种分布, 而生境异质性在较大尺度(15-25 m)上影响树种的空间分布格局。     

贵州赤水常绿阔叶林不同层次树木空间分布格局和竞争的关系

在贵州赤水桫椤国家级自然保护区的亚热带常绿阔叶林中设置0.96 hm~2的固定样地进行调查,将2299株乔木按树高分级(林木下层、中层、上层),进行空间定位并记录胸径和冠幅等个体指标。采用基于个体间距离的对数相关函数(Paircorrelation functions,PCF)和基于个体胸径、冠幅的标记点格局分析(Marked correlation functions,MCF)空间统计方法检测不同高度层内和层间的空间分布格局和潜在的竞争,结果表明:(1)样地内上层树种呈现随机分布,中下层树种在0—6 m的小尺度内呈聚集分布,在层间关系上表现为0—1 m尺度下显著的正相关关系。(2)在群落内上层个体中没有检测到显著的层内竞争,也未发现其与中、下层树种存在层间竞争关系。(3)群落中、下层树种的层内和层间竞争对两者间空间分布格局和个体大小均产生了一定影响,其相邻个体的胸径和冠幅呈现显著负相关关系。这说明在贵州赤水地区常绿阔叶林中群落中不同层次个体处于不同演替阶段,群落内层内和层间竞争对个体较大的林木上层物种的影响已经非常微弱,而主要发生在中、下层树种内,导致了个体较小的中下层更新树种在层间和层内都出现了小尺度下聚集分布格局,这解释了研究地内不同高度级别树种间竞争关系与共存策略,也验证了标记点格局分析方法在检测植物群落竞争中的敏感性。     

Spatial distributions of tree species in a subtropical forest of China

The spatial dispersion of individuals in a species is an important pattern that is controlled by many mechanisms. In this study we analyzed spatial distributions of tree species in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. O-ring statistic was used to measure spatial patterns of species with abundance >10. Ω_0–10, the mean conspecific density within 10 m of a tree, was used as a measure of the intensity of aggregation of a species. Our results showed: (1) aggregated distribution was the dominant pattern in the plot. The percentage of aggregated species decreased with increased spatial scale. (2) The percentages of significantly aggregated species decreased from abundant to intermediate and to rare species. Rare species was more strongly aggregated than common species. Aggregation was weaker in larger diameter classes. (3) Seed traits determined the spatial patterns of trees. Seed dispersal mode can influence spatial patterns of species, with species dispersed by both modes being less clumped than species dispersed by animal or wind, respectively. Considering these results, we concluded that seed dispersal limitation, self-thinning and habitat heterogeneity primarily contributed to spatial patterns and species coexistence in the forest.     

What causes conspecific plant aggregation? Disentangling the role of dispersal,habitat heterogeneity and plant–plant interactions

Spatial patterns of plant species are determined by an array of ecologica factors including biotic and abiotic environmental constraints and intrinsic species traits. Thus, an observed aggregated pattern may be the result of short‐distance dispersal, the presence of habitat heterogeneity, plant–plant interactions or a combination of the above. Here, we studied the spatial pattern of Mediterranean alpine plant Silene ciliata (Caryophyllaceae) in five populations and assessed the contribution of dispersal, habitat heterogeneity and conspecific plant interactions to observed patterns. For this purpose, we used spatial point pattern analysis combined with specific a priori hypotheses linked to spatial pattern creation. The spatial pattern of S. ciliata recruits was not homogeneous and showed small‐scale aggregation. This is consistent with the species’ short‐distance seed dispersal and the heterogeneous distribution of suitable sites for germination and establishment. Furthermore, the spatial pattern of recruits was independent of the spatial pattern of adults. This suggests a low relevance of adult‐recruits interactions in the spatial pattern creation. The difference in aggregation between recruits and adults suggests that once established, recruits are subjected to self‐thinning. However, seedling mortality did not erase the spatial pattern generated by seed dispersal, as S. ciliata adults were still aggregated. Thus, the spatial aggregation of adults is probably due to seed dispersal limitation and the heterogeneous distribution of suitable sites at seedling establishment rather than the presence of positive plant–plant interactions at the adult stage. In fact, a negative density‐dependent effect of the conspecific neighbourhood was found on adult reproductive performance. Overall, results provide empirical evidence of the lack of a simple and direct relationship between the spatial structure of plant populations and the sign of plant–plant interactions and outline the importance of considering dispersal and habitat heterogeneity when performing spatial analysis assessments.     

长白山阔叶红松林物种多度和空间分布格局的关系

应用随机分布多度模型和聚集分布多度模型,探讨不同研究尺度下物种多度和空间分布格局的关系.结果表明,预测的物种多度不仅受物种分布面积大小的影响,还受其聚集程度的影响.物种多度和空间分布格局的关系存在着明显的尺度效应,即随着研究尺度的增加,无论是随机分布多度模型还是聚集分布多度模型,通过物种空间分布格局来预测物种多度的准确度都在下降.聚集分布多度模型预测物种多度的结果要好于随机分布多度模型,这表明该区大多数物种是聚集分布的.由于物种的空间分布格局不同,不同物种多度的预测值和真实值之间的差异也不同.因此,为了进一步提高模型预测的准确性,进一步考虑不同物种的生活史特性是必要的.     

辽东山区次生林木本植物空间分布

森林木本植物的空间格局有助于揭示群落结构的形成机制与潜在的生态学过程,且对林分经营具有一定指导意义。在0—50 m尺度范围内综合分析了辽东山区4 hm2温带次生林样地多度10的树种空间格局。研究发现:(1)在完全随机零模型下,大部分树种呈现聚集格局,聚集格局树种的比例随尺度增加而降低;在32 m的较大尺度下,随尺度增加,随机和规则格局成为树种分布的主要形式;(2)在异质性泊松过程零模型下,55.9%的树种呈现随机格局,其余大部分树种在10 m的尺度下呈现聚集格局,且随尺度增加,规则格局成为主要形式;(3)在完全随机零模型下,树种属性(林层、径级和多度)显著地影响种群聚集度,而在异质性泊松过程零模型下,树种属性对种群聚集度不存在显著影响。综上,生境异质性、扩散限制和树种属性部分解释了辽东山区次生林木本植物空间分布格局,相对而言,生境异质性的效应更为突出。研究结果有助于揭示次生林群落生物多样性的维持机制。     

Disturbance-dependent spatial distribution of sexes in a gynodioecious understory shrub

Several ecological conditions and processes occurring naturally in plant populations may lead to spatial aggregation of sexes within populations of sexually polymorphic species. In addition, ecological disturbances such as forest management or fire could also affect the spatial distribution of sexes within populations. Spatial aggregation of sexes can have important consequences for the fitness of the individuals in sex-biased patches through increased pollen limitation and/or variation in the male fitness of hermaphrodites. Therefore, spatial aggregation of sexes could be relevant for the maintenance of the sexual polymorphism in plant species. Here, we used point pattern analysis to study the spatial distribution of female and hermaphrodite individuals within a single population of the gynodioecious understory shrub Daphne laureola, inhabiting a young pine reforestation and the adjacent natural undisturbed area. In the undisturbed area, we found that hermaphrodites were distributed randomly whereas females were spatially aggregated at short distances. Such aggregation could result from narrower ecological amplitude, cyto-nuclear determination of sex and limited dispersal and/or increased cloning in females. In the reforested area, females were slightly more abundant and the two sexes were randomly distributed among all plants. Recolonization processes after certain forest disturbances could thus favour the establishment of females and alter the spatial distribution of sexes in this understory species.     

古田山中亚热带常绿阔叶林动态监测样地——群落组成与结构

 亚热带常绿阔叶林是世界上主要植被类型之一,集中分布于我国,其中以中亚热带的常绿阔叶林最为典型。为了更好地研究常绿阔叶林森林生 物多样性维持机理,按照CTFS (Centre for Tropical Forest Science)样地建设的标准,于2004年11月～2005年9月在浙江开化古田山国家级 自然保护区建立了常绿阔叶林24 hm²永久样地。该文对样地内胸径≥1 cm的木本植物进行了统计,初步分析了古田山森林样地(Gutianshan (GTS) forest plot)的群落组成与空间结构。 群落的区系类型以热带成分比较多,在属的水平上53个是热带分布,44个是温带分布。共有159 种,隶属于49科103属,总计140 700株,以常绿树种为主（91个物种,占总优势度的90.6%,重要值为85.6%,占样地总个体数的85.9%）;样地 群落有明显的优势物种和大量稀有种(Rare species,每hm2个体数小于1)。甜槠(Castanopsis eyrei )、木荷(Schima superba)和马尾松 (Pinus massoniana)在群落中占有主要优势; 稀有种占总物种数的37.1％（59/159）;具有温带落叶阔叶林和热带雨林的相关特征,较好地代 表了中亚热带常绿阔叶林群落。垂直结构由林冠层（63个种）、亚乔木层（70个种）和灌木层 （26个种）组成。样地所有木本植物物种总径级 分布呈倒“J"形,群落更新良好。几个优势种空间分布随着径级增大,聚集程度降低;并且它们的分布表现出与生境紧密关联,不同的物种表 现出对生境不同的偏好。该文最后系统地比较了大样地方法与传统取样方法。     

Does spatial heterogeneity blur the signature of dispersal syndromes on spatial patterns of woody species? A test in a tropical dry forest

Spatial patterns of adult plants are a consequence of several ecological processes related to seed dispersal and recruitment. Dispersal limitation, mediated by dispersal syndrome, is considered a key factor in the formation of adult plant spatial patterns. Although this initial pattern determined by dispersal has been thoroughly studied, the subsequently modification by the effect of additional ecological factors, such as habitat heterogeneity is less understood. We explored the relative importance of dispersal syndrome and spatial heterogeneity on the realization of spatial patterns of adult trees in an Ecuadorian tropical dry forest. The spatial distribution of 28 species was modeled with four different spatial point processes each: homogeneous Poisson (HPP), inhomogeneous Poisson (IPP), homogeneous Poisson cluster (HPCP), and inhomogeneous Poisson cluster process (IPCP). These models allowed us to discern between effects of random processes, habitat heterogeneity, limited dispersal, and joint effects of habitat heterogeneity and limited dispersal. We employed Akaike's information criterion (AIC) to select the model which best fit the spatial pattern of each species. The best model of each species was used to analyze differences in cluster size and degree of aggregation, between dispersal syndromes. Seventy‐five percent of the species showed inhomogeneous patterns. IPCP yielded the best fit for the spatial distribution of 50% of species in the studied forest and was the prevalent model for the three dispersal syndromes. Thus, the effect of spatial heterogeneity was prevalent in the distribution of most species in this dry tropical forest. Only 21% of species had spatial patterns compatible with random mechanisms associated to limited dispersal around parent sources. Clearly, ignoring habitat heterogeneity could bias the analysis of relationships between dispersal syndrome and species patterns.     

Aggregated recruitment patterns under adult crowns in Photinia glabra,a bird‐dispersed tree species

To clarify recruitment patterns of Photinia glabra, which is an evergreen, broad‐leaved, bird‐dispersed tree species, we analyzed spatial distribution in P. glabra recruits at each growth stage and demography of current‐year seedlings with respect to distributions of adults in a warm‐temperate secondary forest, western Japan. Although individuals ≥ 5 cm diameter at breast height (DBH) that had nearly produced fruits showed a random distribution, seedlings (≥ 1 year old, < 10‐cm stem length [SL]), small saplings (10 ≤ SL < 30 cm) and large saplings (≥ 30‐cm SL, < 5‐cm DBH) were clumped and associated with reproductive adults at approximately 2–3‐m scales, nearly equal to their average crown radius. Based on monitoring the demography of current‐year seedlings, emerged seedling density profoundly decreased, and no seedlings survived at longer than an adult's crown scales, with distance‐dependent mortality as a result of disease and herbivory not greatly affecting the current‐year seedling mortality. Thus, aggregated seed dispersal under the crown of adult P. glabra would directly influence the distribution of recruits for P. glabra in this forest. Of the bird‐dispersed tree species in this forest, P. glabra produced the highest amount of fruits during large crop years, and their fruits ripened during the late seasonal period (early January), suggesting that birds might be strongly attracted to these species, in turn leading to seeds being deposited mostly under the tree crowns. We propose that dispersal limitation would occur, even in a bird‐dispersed tree species such as P. glabra, owing to plant–bird interactions in the forest.     

Seed dispersal and seedling recruitment of trees at different successional stages in a temperate forest in northeastern China

Aims Spatial distribution of adult trees in a forest community is determined by patterns of both seed dispersal and seedling recruitment. The objectives of our study were to understand the processes of seed dispersal and seedling recruitment of dominant tree species in a temperate forest of northeastern China and to identify the factors constraining seed dispersal and seedling establishment at different stages of forest succession.Methods During three summer and autumn sessions between 2006 and 2008, altogether 113080 seeds from 22 different tree species were collected in three large field plots representing different forest types in the Changbai Mountain region of northeastern China. The spatial distribution of seed abundance was analyzed using a Syrjala test. Regeneration success of nine major tree species was assessed using variables defining 'limitations' in 'seeds' and 'seedling establishment'.Important findings We found that seed production fluctuated between years and varied greatly with forest types. Four tree species, Acer spp., Fraxinus mandshurica, Tilia amurensis and Betula spp., had the greatest seed production and the widest range of seed dispersal, whereas Quercus mongolica showed the most sustained seed production pattern. The spatial patterns of seed abundance differed significantly among forest types and years. The tree species investigated in this study differed in the degree of seed limitation, as well as in limitation of seedling establishment. There were both negative and positive correlations between seed density and seedling density, depending on site and parental tree density. Seeds of 16 tree species were found in the Populus davidiana–Betula platyphylla forest (PBF) plot, 11 in the conifer and broad-leaved mixed forest (CBF) plot but only 8 in the broad-leaved-Korean pine mixed forest (BKF) plot. The number of seed-contributing species was not only greater in the secondary forests (CBF and PBF plots) than in the primary forest (BKF plot) but was also more variable during the 3 years of assessment. Results from the correlations between seed density and seedling occurrence and that between parental tree density or seed weight and dispersal limitation confirm our intuitive expectations, i.e. heavy seeds had greater dispersal limitation but higher establishment success than light seeds.     

密度制约对宝天曼落叶阔叶林锐齿栎死亡前后分布格局的影响

为了在温带和亚热带的过渡带中验证森林树木死亡是否受密度制约的影响, 我们在宝天曼国家级自然保护区选择1个100 m×100 m的暖温带落叶阔叶林样地, 用双变量函数g(r)(the pair-correlation function)研究了锐齿栎(Quercus aliena var. acuteserrata)死亡前(活树和枯木统称为死亡前树木)和死亡后(活树为死亡后树木)的分布格局。把样地中的锐齿栎分为幼树(1 cm ≤ DBH<10 cm)、小树(10 cm ≤ DBH<20 cm)和成年树(DBH≥20 cm)3个不同的生长阶段来研究密度制约对空间分布格局的影响机制。结果表明: (1)死亡前锐齿栎在r>5 m尺度呈聚集分布, 死亡后幼树(1 cm≤DBH<10 cm)和成年树(DBH≥20 cm)在1-25 m尺度呈现随机分布, 死亡后小树(10 cm≤DBH< 20 cm)在r<1.5 m和2.5-4.5 m的尺度为随机分布, 在r>5 m的尺度呈聚集分布; (2)采用随机标签零模型和案例-对照设计的方法, 排除生境异质性影响后, 将幼树和小树的分布格局作为案例, 将成年树的分布格局作为对照, 并代表生境异质性的作用, 通过小径级树木与成年树分布格局的对比发现, 密度制约效应对死亡前后的锐齿栎分布格局均具有影响; (3)幼树和小树在成年树周围的分布死亡前为显著聚集分布格局, 死亡后剩余树木的聚集强度下降; 随着与成年树之间距离的增加, 死亡后的幼树(仅包括现存的活树)逐渐向随机分布格局演替。本研究初步表明锐齿栎空间格局受生境异质性的影响并呈现出显著的聚集效应, 排除生境异质性影响后, 锐齿栎死亡前后的空间格局受到密度制约的影响, 这一结果为Janzen-Connell假说提供了支持。