Spatial pattern and process in forest stands within the Virginia piedmont

Abstract. Question: Underlying ecological processes have often been inferred from the analysis of spatial patterns in ecosystems. Using an individual‐based model, we evaluate whether basic assumptions of species’life‐history, drought‐susceptibility, and shade tolerance generate dynamics that replicate patterns between and within forest stands. Location: Virginia piedmont, USA. Method: Model verification examines the transition in forest composition and stand structure between mesic, intermediate and xeric sites. At each site, tree location, diameter, and status were recorded in square plots ranging from 0.25 to 1.0 ha. Model validation examines the simulated spatial pattern of individual trees at scales of 1–25 m within each forest site using a univariate Ripley's K function. Results: 7512 live and dead trees were surveyed across all sites. All sites exhibit a consistent, significant shift in pattern for live trees by size, progressing from a clumped understorey (trees ± 0.1 m in diameter) to a uniform overstorey (trees > 0.25 m). Simulation results reflect not only the general shift in pattern of trees at appropriate scales within sites, but also the general transition in species composition and stand structure between sites. Conclusions: This shift has been observed in other forest ecosystems and interpreted as a result of competition; however, this hypothesis has seldom been evaluated using simulation models. These results support the hypothesis that forest pattern in the Virginia piedmont results from competition involving species’life‐history attributes driven by soil moisture availability between sites and light availability within sites.     

Spatial patterns and associations of four species in an old-growth temperate forest

The spatial pattern of a tree species is an important characteristic of plant communities, providing critical information to explain species coexistence. The spatial distribution and association of four different successional species were analyzed among different life-history stages in an old-temperate forest. Significant aggregation patterns were found, and the degree of aggregation decreased with the scales and life-history stages. Significant interspecific spatial associations were detected. In comparing the relationships among the different life-history stages, positive associations were found at small scales in all of the juvenile species pairs. In the adult stage, negative associations were detected in coniferous vs. deciduous species pairs, while the deciduous species pairs, which have identical resource requirements, showed a positive association in this study. The coniferous species pairs showed a positive association at small scales. We infer that seed dispersal, competitive ability, or the requirement for specific topographic and light environments may contribute to the coexistence of these species.     

庐山森林景观空间分布格局及多尺度特征

森林景观是区域整体景观的重要组成部分,研究其空间分布格局对于优化区域景观整体结构和发挥生态经济效益具有重要意义。基于庐山2010年植被斑块数据,将森林景观划分为常绿阔叶林、落叶阔叶林、松类、杉类和竹类等5类景观,从森林景观的不同发育阶段和林分类型角度出发,运用点格局分析法分析5类森林景观空间分布格局特征。最邻近距离分析表明:5类森林景观空间分布类型均服从集聚分布但聚集强弱有变化;不同发育阶段的森林景观空间分布类型以集聚分布为主,随机分布为辅,尤其幼龄林比较显著,中龄林和老龄林次之;5类森林景观的天然林均服从集聚分布,人工林大多趋于随机分布,只有松类和杉类呈显著集聚分布。Ripley's K函数揭示了不同发育阶段和林分类型的森林景观的多尺度集聚特征,即在小尺度范围内服从随机分布,随着空间距离的增大,以空间特征尺度为分界线,空间聚集强度先逐渐增强,随后不断减弱。总体来看,庐山森林景观的发育阶段主要处于幼年时期,原始植被遭到人类大肆破坏,幼龄林大片分布,属于典型的恢复性植被,未来要重点保护好天然林,减少人为干扰,实现森林景观适度集聚。研究庐山森林景观的空间分布和多尺度特征可以为生态环境保护和实现森林可持续经营提供理论指导。     

Neighbourhood interactions and environmental factors influencing old‐pasture succession in the Central Pyrenees

Abstract. The shrub Buxus sempervirens and the trees Abies alba and Fagus sylvatica have recently recolonized old‐pastures in the Central Pyrenees. We mapped all live and dead individuals (> 1.3 m tall) in a large forest plot in Ordesa Valley to examine the importance of density‐dependent processes during recolonization. Biotic interactions were inferred from changes in horizontal structure and the influences of neighbours on tree survival. Buxus differentially influenced establishment and survival of tree species, thereby controlling future canopy composition and spatial structure. The rapidly invading Abies formed denser patches on elevated sites less occupied by Buxus, whereas Fagus preferentially established within shrubs. Abies reached densities which led to intense intraspecific competition and high mortality rates among saplings. Self‐thinning in Abies led to smaller numbers of regularly spaced survivors, and greater relative dominance of Fagus. Disregarding intraspecific effects and abiotic environment, Abies survival was significantly lower under Buxus shrubs, which suggests that the spatial location and abundance of Abies was constrained by the location of Buxus. Fagus survival was not related to Buxus density, but remained significantly lower in denser Abies patches. The higher mortality of Fagus in denser Abies patches, and the resulting spatial segregation of the species, reflects asymmetric interspecific competition. Inhibition from Buxus shrubs and intraspecific competition prevent invading Abies from dominating and may therefore help in maintaining a mixed Abies‐Fagus stand.     

Spatial distribution patterns of trees at different life stages in a warm temperate forest

We have investigated tree distributions in relation to topography between different tree life history stages, from the seed-dispersal stage to the adult stage in a warm temperate evergreen broadleaved forest on Yakushima Island, Japan, to clarify the critical stages in determining adult tree distributions. We conducted a census of all living trees > or =30 cm tall and collected seed falls over three years using 25 seed traps in a 50 m x 50 m quadrat. Four life stages were defined: stage 1, dispersed seed; stage 2, individuals taller than 30 cm and diameter at breast height (DBH) < 1 cm; stage 3, trunks 1 cm < or = DBH < 10 cm; stage 4, trunks with DBH > or = 10 cm. We classified 17 common tree species into three groups; group A was distributed mainly on the upper slope, group B on the lower slope, and group C on both. Most of group A and B trees at stages 2-4 showed an aggregated distribution along the topographical gradient. The densities at stage 1 showed weaker aggregations according to slope. Topography-specific tree distribution was probably determined at the regeneration stage, and later survival was less effective as a mechanism of vegetation differentiation.     

小兴安岭天然针阔混交林主要树种空间格局及其关联性

在小兴安岭天然针阔混交林1.1 hm~2的固定样地内,本文调查和统计了其物种组成和林分径级结构。运用空间点格局分析方法中的O-ring函数,对种群的空间分布格局及其关联性进行分析。结果表明:样地内共有乔木树种15种,重要值排在前四位的依次为红皮云杉(Picea koraiensis)、枫桦(Betula costata)、春榆(Ulmus davidiana)和色木槭(Acer mono),红皮云杉和枫桦属于样地内的优势种,重要值分别为15.81%和11.10%。林下更新状态良好,枫桦的径级结构有呈正态分布的趋势,其余三种均呈现倒"J"形。O-ring函数结果证实,四个主要树种的种群空间分布格局相似,均在0—5 m小尺度上呈聚集分布,随着尺度的增大,逐渐变为随机分布;红皮云杉与其他三个主要种群之间在较大尺度上均呈现显著正相关或无相关,在小尺度上仅与枫桦呈极显著正相关;色木槭在较小尺度上与其他三个种群呈弱负相关性,随着尺度的增大,逐渐变为无相关性。     

华北山地典型天然次生林群落的树种空间分布格局及其关联性

关帝山华北落叶松、云杉、杨桦天然次生林植被类型是华北高海拔地带代表性的森林类型。以该地区上述3种典型的天然次生林群落为例,运用点格局分析中的单变量O-ring统计方法,分析各群落主要树种的种群空间分布格局;用双变量O-ring统计方法,分析各群落中树种间的空间关联性;并对同一树种在不同森林群落类型中种群空间分布格局和种间关联性的变化进行比较,以探讨华北山地次生林群落空间格局形成和种群维持与动态机制。研究结果表明,(1)各种群在相对小的尺度上聚集分布特征明显,随尺度增加树木种群的聚集性减弱并逐渐表现出随机分布的格局特征;(2)同一树种在不同森林群落类型中空间分布格局差异很大。红桦、白桦和山杨在阔叶林中呈明显的小尺度聚集分布格局,而在针叶林群落呈随机分布。在阔叶次生林中混生的小龄级华北落叶松呈现聚集性空间分布格局。在华北落叶松为优势树种的针叶林中,大龄级的华北落叶松表现出小尺度上均匀分布和较大尺度上随机分布的格局特征;(3)各群落中树种间的空间关联性主要表现在小尺度范围,随尺度加大空间关联性逐渐不明显;(4)在阔叶树占优势的次生林,小龄级华北落叶松与桦木的分布有较明显的空间正关联性,而在华北落叶松为优势树种的针叶林,桦与华北落叶松并未表现明显的空间关联性。种群分布格局和种群间的空间关联性随群落结构、空间尺度和龄级结构不同而有较大变化。     

山西灵空山刺五加种群空间分布格局及种间空间关联性

以山西灵空山自然保护区刺五加(Acanthopanax senticosus(Rupr.Maxim.)Harms)种群为对象,采用空间点格局方法中的Ripley’s K函数,对刺五加种群不同龄级的空间分布格局及其与主要灌木物种间的空间关联性进行研究,并对种群空间分布格局进行可视化解析。结果显示,刺五加种群中幼年个体数量多,径级结构呈金字塔型,为增长型种群,能够实现持续更新;刺五加种群的径级Ⅰ在小尺度上呈聚集分布,随着尺度增大出现随机分布;径级Ⅱ、Ⅲ、Ⅳ在全尺度上呈聚集分布,径级Ⅴ、Ⅵ在小尺度上为聚集分布,随着龄级和空间尺度的增大,刺五加种群聚集强度渐弱,呈现随机分布;刺五加种内关联在全尺度上呈正相关,但与群落中主要灌木在小尺度上呈不相关或负相关,在大尺度上呈负相关。研究表明刺五加的生长条件较为适宜,并可以通过相互之间的庇护作用,减弱种内竞争,但与其他树种间存在强烈竞争,生存现状较差,在群落中处于劣势。     

Spatial patterns and interspecific associations of three canopy species at different life stages in a subtropical forest, China

Spatial patterns of species at different life stages are an important aspect for understanding causal mechanisms that facilitate species co-existence.Using Ripley's univariate L(t) and bivariate L12(t) functions,we analyzed the spatial patterns and interspecific associations of three canopy species at different life history stages in a 20-ha subtropical forest plot in Dinghushan Nature Reserve.Based on diameter at breast height (DBH),four life stages were distinguished.Castanopsis chinensis and Schima superba showed a unimodal DBH distribution.Engelhardtia roxburghiana showed a bimodal curve.L(t) function analysis showed significantly aggregated distributions of all three species at later life stages and random distribution at early life stages at some scales.From the analysis of L12(t) function,the results showed the positive association was a dominant pattern for most species pairs at most scales but the intensity of association decreases with the increase of life stages.Juveniles of the three species had no negative intra- and interspecific associations with the older life stages.Only premature trees were suppressed by overmature trees at some scales.Considering these results,we found three canopy-dominant species that lacked regeneration.There was no direct competition occurring between understorey individuals.Young trees can grow well under conspecific species with two other species.Longevity and lack of regeneration led to a large number of trees stored in mature and overmature stages,therefore,intra-and inter-competition can be strong at later life stages.     

神农架天然针阔混交林乔木更新的空间格局

为深入了解神农架天然针阔混交林群落乔木树种的更新特征,于神农架国家公园设置1 hm²森林动态监测样地对树种更新进行调查,利用Ripley的L函数分析优势更新树种的空间分布格局、种间空间关联性以及与大树的空间分布关系。结果显示：（1）更新个体共有35 752株,隶属于19科29属45种。更新优势种基本是乔木层优势树种,物种更新数量表现为：巴山冷杉（Abies fargesii Franch.） > 华山松（Pinus armandii Franch.） > 四蕊槭（Acer stachyophyllum subsp.tetramerum（Pax）A.E.） > 华中山楂（Crataegus wilsonii Sarg.）,混交林群落更新良好。（2）随着尺度增加,巴山冷杉、华山松更新空间格局由聚集分布向随机分布和均匀分布发展;华中山楂、四蕊槭更新的空间格局在整个尺度上均为聚集分布,表明针叶、阔叶树种更新在空间生态利用策略上出现分化。（3）巴山冷杉与其余3个优势更新树种、华山松与华中山楂更新呈显著正相关,表现出对微生境的共同喜好;四蕊槭分别与华山松、华中山楂更新的空间关系表现为显著负相关,种间竞争激烈。（4）巴山冷杉、华中山楂、四蕊槭的更新个体与大树的空间分布总体为显著正相关,这可能与种子传播限制相关;华山松幼苗、幼树与大树的空间分布在小尺度上呈负相关,存在资源不对称竞争。     

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.     

A practical approach to the study of spatial structure in simple cases of heterogeneous vegetation

Abstract. Spatial heterogeneity is a characteristic of most natural ecosystems which is difficult to handle analytically, particularly in the absence of knowledge about the exogenous factors responsible for this heterogeneity. While classical methods for analysis of spatial point patterns usually require the hypothesis of homogeneity, we present a practical approach for partitioning heterogeneous vegetation plots into homogeneous subplots in simple cases of heterogeneity without drastically reducing the data. It is based on the detection of endogenous variations of the pattern using local density and second‐order local neighbour density functions that allow delineation of irregularly shaped subplots that could be considered as internally homogeneous. Spatial statistics, such as Ripley's K‐function adapted to analyse plots of irregular shape, can then be computed for each of the homogeneous subplots. Two applications to forest ecological field data demonstrate that the method, addressed to ecologists, can avoid misinterpretations of the spatial structure of heterogeneous vegetation stands.     

Spatial dynamics of regeneration in a conifer/broad‐leaved forest in northern Japan

Abstract. This study deals with stand dynamics over a 6‐yr period in a conifer/broad‐leaved mixed forest in Hokkaido, northern Japan. The annual rates of gap formation and recovery were 81.3 m²/ha and 66.7 m²/ha, respectively and turnover time of the canopy was 125 yr. The recruitment processes of the component species in this cool‐temperate forest were governed by different canopy types: gap, canopy edge and closed canopy. Magnolia obovata regenerated in canopy edges, and Acer mono and Prunus ssiori regenerated in canopy edges and gaps. The results suggested that the mosaic structure made up of closed canopy, canopy edge and gap was related to various regeneration niches. Abies sachalinensis had high mortality rates, initiating gap expansion. The transition probabilities from closed canopy or canopy edge to gap for deciduous broad‐leaved trees were lower than for A. sachalinensis, which implies that the difference in degeneration patterns of conifer and broad‐leaved canopies contributes to the heterogeneity of spatial structure in the mixed forests. Spatial dynamics were determined by a combination of gap expansion by A. sachalinensis (neighbour‐dependent disturbance) and gap formation by deciduous broad‐leaved trees (random disturbance).     

吉林蛟河针阔混交林树木生长的空间关联格局

以吉林蛟河21.12hm2(660m×320m)针阔混交林样地为对象,利用2009年和2014年森林生长观测数据,研究树木生长的空间自相关格局及其生境影响机制。在样地生境型划分结果的基础上,采用Ripley's L(r)函数分析不同生境型中树木种群空间分布特征;利用标记相关函数分析不同生境型中树木生长特征的空间关联格局。研究结果表明:(1)红松(生境型3:1—5m)、蒙古栎(生境型3:1—3m)、胡桃楸(生境型2:1—2m;生境型3:1—7m)、黄檗(生境型2:1—3m;生境型4:1—5m)、水曲柳(生境型3:1—2m;生境型4:1—2m)、瘤枝卫矛(生境型2:1—15m)在特定生境和空间尺度上呈随机分布,但空间格局仍以聚集性分布为主;其余10个物种则在全部0—30m尺度上呈聚集分布。(2)标记相关函数分析显示春榆、毛榛、色木槭、瘤枝卫矛和千金榆的径向生长至少在一个生境中表现出正相关格局;暴马丁香、胡桃楸、裂叶榆、瘤枝卫矛、水曲柳、紫椴、糠椴、毛榛、色木槭和白牛槭的径向生长至少在一个生境中表现出负相关格局;红松、黄檗、蒙古栎和簇毛槭的径向生长在全部尺度上均未检测到显著的空间关联格局。因此,不同树种径向生长的空间自相关特征不同,树种生长特征的空间关联格局具有明显的生境依赖性。     

不同干扰强度下针阔混交林树种空间格局及种间关联性

为了探究不同干扰程度下针阔混交林树种空间格局变化及树种种间关联性,本研究以关帝山庞泉沟自然保护区不同干扰强度的针阔混交林为对象,选取郁闭度、林分密度、伐桩数量作为划分干扰程度的因子,采用相对影响法,将林分划分为未干扰、中度干扰和严重干扰3个等级.利用空间点格局K2函数分析了林分的空间分布格局及其不同树种的种间关联.结果...