Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation

Understanding processes underlying spatial distribution of tree species is fundamental to studying species coexistence and diversity. This study modeled point patterns of tree distribution, expressed by Cartesian coordinates of individual trees within a mapped forest stand, for the purpose of identifying processes that may generate spatial patterns of tree communities. We used four primary point pattern processes (homogeneous Poisson process, inhomogeneous Poisson process, homogeneous Thomas process, and inhomogeneous Thomas process) to model tree distribution in two stem-mapped forests in Taiwan, Republic of China. These four models simulate spatial processes of habitat association and seed dispersal, allowing us to evaluate the potential contribution of habitat heterogeneity and dispersal limitation to the formation of spatial patterns of tree species. The results showed that the inhomogeneous Thomas process was the best fit model and described most of the species studied, suggesting that spatial patterns of tree species might be formed by the joint effects of habitat associations and dispersal limitation. The homogeneous Thomas process that models the effect of dispersal limitation was the second best model. We also found that the best fit models could be predicted by species attributes, including species abundance and dispersal mode. The significant traits, however, differed between the two study plots and demonstrated site-specific patterns. This study indicated that the interactive operation of niche-based (habitat heterogeneity) and neutral-based (dispersal limitation) may be important in generating spatial patterns of tree species in forest communities.     

喀斯特常绿落叶阔叶林死亡个体空间分布格局及生境关联

树木死亡在森林动态中发挥着重要作用, 环境因子是影响树木死亡的重要因素。为了阐明喀斯特常绿落叶阔叶林死亡个体空间分布格局及其与环境的相关性, 本研究以木论25 ha森林动态监测样地的两次木本植物调查数据为基础, 分析了死亡个体的物种组成、径级结构、分布格局及其与生境的关联。结果表明, 样地内共有死亡个体17,306株, 隶属57科130属194种; 其中, 死亡率排名前三的科和属分别为八角枫科、蔷薇科、大风子科以及山麻杆属(Alchornea)、火棘属(Pyracantha)、八角枫属(Alangium); 在物种水平上, 山麻杆(Alchornea trewioides)、毛桐(Mallotus barbatus)、野桐(Mallotus japonicus)等死亡率较高。样地内死亡个体的平均胸径为3.83 cm, 最大47.11 cm; 径级分布呈倒“J”型。从分布格局来看, 样地内大部分死亡个体在0-50 m的尺度上呈聚集分布, 部分大径级树木死亡个体趋向于随机分布。生境关联分析发现, 个体死亡率与山顶、陡坡、缓坡、洼地呈正关联的物种数分别为41种、13种、41种和38种, 呈负关联的物种数分别为38种、67种、33种和10种, 与各生境无关联的物种数分别为4种、3种、9种和35种; 多度排名前20的物种其个体死亡率均与4种生境类型呈现出正关联或者负关联; 不同径级个体死亡率都与山顶生境呈正关联, 与缓坡生境呈负关联, 其中5 cm ≤ DBH < 10 cm和10 cm ≤ DBH < 15 cm个体死亡率均与洼地生境呈正关联; DBH ≥ 15 cm个体死亡率与陡坡生境呈正关联, 与洼地生境呈负关联。在喀斯特亚热带常绿落叶阔叶林中, 死亡个体空间分布格局总体表现为聚集分布, 树木死亡受生境作用的影响较大。     

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。     

小兴安岭阔叶红松林粗木质残体空间分布的点格局分析

采用点格局分析方法对小兴安岭典型阔叶红松林9hm2(300m×300m)固定样地内粗木质残体(CWD)的空间分布格局进行了研究。结果表明:固定样地内CWD的总密度为368.8株/hm2,径级结构呈现明显的正态分布,各径级密度差别较大。花楷槭(27.8株/hm2)和枫桦(26.1株/hm2)是阔叶CWD的主要组成树种。红松(41.6株/hm2)是针叶CWD的主要组成树种,针叶树种CWD的数量随着径级的增大而增加,呈典型的J型分布。在150m的空间尺度内,CWD总体在较小尺度上表现为集群分布,在40m尺度上聚集强度最大(0.40),随着尺度的增加,CWD趋于均匀分布。不同径级组在不同规模尺度聚集,随着径级的增加,聚集强度呈下降趋势。不同物种CWD在各径级下的空间分布格局有所不同,具体表现为集群分布和由集群分布向随机分布发展两种形式。不同存在形式的CWD在研究尺度内随着尺度的增加,由集群分布向随机分布发展。随着CWD腐烂等级的增高,其聚集强度和聚集尺度均增大。CWD的空间分布格局是阔叶红松林群落与其自然环境长期作用的结果,从某种程度上也反映了该林型天然更新的格局和机制。     

Landscape patterns of species-level association between ground-beetles and overstory trees in boreal forests of western Canada (Coleoptera, Carabidae)

Spatial associations between species of trees and ground-beetles (Coleoptera: Carabidae) involve many indirect ecological processes, likely reflecting the function of numerous forest ecosystem components. Describing and quantifying these associations at the landscape scale is basic to the development of a surrogate-based framework for biodiversity monitoring and conservation. In this study, we used a systematic sampling grid covering 84 km(2) of boreal mixedwood forest to characterize the ground-beetle assemblage associated with each tree species occurring on this landscape. Projecting the distribution of relative basal area of each tree species on the beetle ordination diagram suggests that the carabid community is structured by the same environmental factors that affects the distribution of trees, or perhaps even by trees per se. Interestingly beetle species are associated with tree species of the same rank order of abundance on this landscape, suggesting that conservation of less abundant trees will concomitantly foster conservation of less abundant beetle species. Landscape patterns of association described here are based on characteristics that can be directly linked to provincial forest inventories, providing a basis that is already available for use of tree species as biodiversity surrogates in boreal forest land management.     

Point Processes for Fine‐Scale Spatial Genetics and Molecular Ecology

This paper introduces point processes into fine‐scale spatial genetics and molecular ecology. Datasets given in the form of a complete map of individuals and their genotypes can be analyzed by means of the theory of marked or multivariate point processes. Beginning with reformulation of conventional spatial autocorrelation statistics in genetics by the language of point processes, this paper first shows an example of point process models that describe spatial patterns of both tree locations and their genotypes, on the assumption of limited seed dispersal and long pollen movement. The results show that isolation‐by‐distance slightly occurs from the assumption above, and more importantly, an increment of the degree of clustering of trees reduces the degree of genetic clustering. Next, the point process model is applied to field data of secondary forest regenerated after seed tree harvesting, and tests the hypothesis that the current population was formed only from a small number of seed trees. The hypothesis was not acceptable, instead, the alternative assuming advance reproduction conducted prior to the harvesting is supported. The results of this first trial of point process models suggest that point processes can provide a useful mathematical methodology in fine‐scale spatial genetics and molecular ecology.     

Size-structure-based models of forest dynamics to interpret population- and community-level mechanisms

Functional size-structure-based models of forest tree population dynamics present a unifying explanation for population-level patterns and tree community organization. Density-dependent regulation can be explicitly replaced by the effect of size-structure-dependent suppression on demographic processes in functional size-structure models. This suppression effect sufficiently explains various patterns reported for crowded evenaged populations. Further, it stabilizes natural forest populations of overlapping generations at a stationary state with balanced recruitment and mortality. The spatial heterogeneity of light resources created by tree size structure offers an opportunity for multiple species to coexist by means of trade-offs between demographic parameters. The energy correlation of tree species diversity at a geographic scale is also attributable to the architectural feature of forests. Recipient of the Botanical Society Award of Young Scientists, 1992.     

Impacts of gene flow and logging history on the local genetic structure of a scattered tree species, Sorbus torminalis L. Crantz

Sorbus torminalis L. Crantz is a colonizing tree species usually found at low density in managed European forests. Using six microsatellite markers, we investigated spatial and temporal patterns of genetic structure within a 472-ha population of 185 individuals to infer processes shaping the distribution of genetic diversity. Only eight young stems were found to be the result of vegetative reproduction. Despite high levels of gene flow (standard deviation of gene dispersal = 360 m), marked patterns of isolation by distance were detected, associated with an aggregated distribution of individuals in approximately 100-m patches. This spatial structure of both genes and individuals is likely to result from patterns of seedling recruitment combined with low tree density. Our results suggest that landscape factors and logging cycles markedly shape the distribution of favourable sites for seedling establishment, which are then colonized by sibling cohorts as a result of joint seed transportation by frugivores. These combined genetic and demographic processes result in similar genetic structure both within and among logging units. However, conversion to high forest may enhance genetic structuring.     

黑龙江省大兴安岭林区火烧迹地森林更新及其影响因子

林火干扰是大兴安岭森林更新的影响因子之一,研究火烧迹地森林更新的影响因子(立地条件、火前植被、火干扰特征)对理解生态系统的结构、功能和火后演替轨迹具有重要意义。选取呼中及新林林业局55个代表性火烧样地,利用增强回归树分析法分析了火烧迹地森林更新的影响因素。结果表明:(1)立地条件是影响针、阔叶树更新苗密度的主要因素;海拔对针叶树更新苗密度的影响最大;坡度对阔叶树更新苗密度影响最大;(2)距上次火烧时间对针叶树更新苗比重影响最大,其次是林型;(3)中度林火干扰后森林更新状况好于轻度和重度火烧迹地。根据火烧迹地森林更新调查分析可知:林型影响火后演替模式,火前为针叶树或阔叶树纯林,火后易发生自我更新(火后树种更新组成与火前林型相同),而针阔混交林在火干扰影响下易于发生序列演替(火后初期以早期演替树种更新为主)。     

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.     

Distributions of tree species along point bars of 10 rivers in the south‐eastern US Coastal Plain

Aim To determine the degree to which rivers within the south‐eastern US Coastal Plain show a predictable spatial distribution of floodplain tree species along each point bar of river bends in relation to elevation and/or soil texture, as seen on the Bogue Chitto River, Louisiana, USA. Also, to understand spatial patterns of tree species on land created during river‐bend migration, and to interpret which physical characteristics of rivers predict this pattern of vegetation. Location The south‐eastern US Coastal Plain. Methods Ten randomly selected rivers within a portion of the region were studied. At each of 10 river bends per river, a census of trees and shrubs was taken and elevation and soil texture were measured at upstream, mid‐ and downstream locations along the forest–point bar margin. To identify physical characteristics of rivers that are predictive of patterns of tree species along point bars, aerial photographs, hydrographs and field data were analysed. Results Tree species composition varied predictably among the three point bar locations, corresponding to an elevation gradient on each bar, on seven of 10 rivers. Species occupying a given point bar location on one river usually occupied the same location on other rivers, in accordance with species‐elevation associations identified in past studies of floodplain forests. Multivariate analysis of river characteristics suggested that rivers failing to show the expected pattern were those with relatively low stream energy and geomorphic dynamics and/or those with hydrological regimes altered by upstream dams. Main conclusions A distinct pattern of streamside forest community structure is related to fluvial geomorphic processes characterizing many rivers within the south‐eastern US Coastal Plain. Characteristics of rivers required to promote the predicted pattern of tree species include a single, meandering channel with point bars; an intermediate level of stream energy; a natural hydrological regime; and location in a biome where a large number of tree species are capable of colonizing point bars.     

云南兰坪云南红豆杉种群年龄结构与空间分布格局分析

利用点格局分析法对滇西北兰坪县4个云南红豆杉(Taxus yunnanensis)种群的年龄结构、不同生长阶段空间分布格局及其关联性进行分析。结果表明:(1)林窗与人为干扰影响4个云南红豆杉种群的龄级结构,小林窗的种群呈稳定型;较大林窗的种群呈进展型;群落结构完整未出现林窗和有较大人为干扰的种群年龄结构均呈衰退型,幼苗、幼树与小树个体缺乏,种群存在一定程度的更新困难,林窗的大小是种群更新的重要因素。(2)人为干扰、自然生境与本身生物学特性影响4个种群空间分布格局,种群1整体及其不同生长阶段个体均呈聚集分布;种群2和种群4在小尺度呈聚集分布,而在大尺度上呈随机分布;种群3随着空间尺度增大分布格局表现为聚集→随机→聚集趋势。(3)种群2中幼苗、小树和中树在小尺度下表现为聚集分布,其他尺度下表现为随机分布;种群3的小树阶段聚集分布和随机分布皆有出现,中树与大树在不同尺度上表现为聚集分布;种群4聚集分布的强度随生长阶段的增加在减弱。(4)4个种群各发育阶段间皆呈或接近显著正相关,种内竞争弱,有利于种群维持。     

Negative density dependence and environmental heterogeneity effects on tree ferns across succession in a tropical montane forest

Although tree ferns are an important component of temperate and tropical forests, very little is known about their ecology. Their peculiar biology (e.g., dispersal by spores and two-phase life cycle) makes it difficult to extrapolate current knowledge on the ecology of other tree species to tree ferns. In this paper, we studied the effects of negative density dependence (NDD) and environmental heterogeneity on populations of two abundant tree fern species, Cyathea caracasana and Alsophila engelii, and how these effects change across a successional gradient. Species patterns harbor information on processes such as competition that can be easily revealed using point pattern analysis techniques. However, its detection may be difficult due to the confounded effects of habitat heterogeneity. Here, we mapped three forest plots along a successional gradient in the montane forests of Southern Ecuador. We employed homogeneous and inhomogeneous K and pair correlation functions to quantify the change in the spatial pattern of different size classes and a case–control design to study associations between juvenile and adult tree ferns. Using spatial estimates of the biomass of four functional tree types (short- and long-lived pioneer, shade- and partial shade-tolerant) as covariates, we fitted heterogeneous Poisson models to the point pattern of juvenile and adult tree ferns and explored the existence of habitat dependencies on these patterns. Our study revealed NDD effects for C. caracasana and strong environmental filtering underlying the pattern of A. engelii. We found that adult and juvenile populations of both species responded differently to habitat heterogeneity and in most cases this heterogeneity was associated with the spatial distribution of biomass of the four functional tree types. These findings show the effectiveness of factoring out environmental heterogeneity to avoid confounding factors when studying NDD and demonstrate the usefulness of covariate maps derived from mapped communities.     

Functional role of forest diversity: Pros and cons of synthetic stands and across-site comparisons in established forests

In contrast to grasslands, forests have only rarely been the subject of functional biodiversity research. This paper addresses specific problems, which arise from applying the synthetic stand approach to long-lived plants, which can be taken to simulate the loss or addition of tree species or tree functional groups in forest stands. Planted synthetic stands often deviate from natural forests in several properties including trophic structure, tree age distribution, and horizontal and vertical canopy structures. Moreover, edge effects and the absence of a quasi-steady state in soil development limit the spectrum of processes which can be analysed for biodiversity effects in synthetic tree stands. We propose that these shortcomings can partly be overcome by combining the synthetic stand approach with observational studies in existing forest stands of contrasting tree diversity. The potentials and limitations of comparisons among existing stands for functional biodiversity research are briefly discussed using the example of the Hainich Tree Diversity Matrix, a species-rich temperate broad-leaved forest consisting of a small-scale mosaic of stands differing in tree diversity, which originated from a variety of historic forest-use practices.     

LANDIS PRO: a landscape model that predicts forest composition and structure changes at regional scales

LANDIS PRO predicts forest composition and structure changes incorporating species‐, stand‐, and landscape‐scales processes at regional scales. Species‐scale processes include tree growth, establishment, and mortality. Stand‐scale processes contain density‐ and size‐related resource competition that regulates self‐thinning and seedling establishment. Landscape‐scale processes include seed dispersal and disturbances. LANDIS PRO is designed to be compatible with forest inventory data, thus extensive inventory data can be directly utilized to initialize and calibrate model parameters before predicting future forest changes. LANDIS PRO allows for exploring the effects of disturbances, management, climate change, and modeling the spread of invasive species. We demonstrate that LANDIS PRO successfully predicts forest successional trajectories and stand development patterns in the Central Hardwood Forest region in U.S.     

An analytical model of stand dynamics as a function of tree growth, mortality and recruitment: the shade tolerance-stand structure hypothesis revisited

Light competition and interspecific differences in shade tolerance are considered key determinants of forest stand structure and dynamics. Specifically two main stand diameter distribution types as a function of shade tolerance have been proposed based on empirical observations. All-aged stands of shade tolerant species tend to have steeply descending, monotonic diameter distributions (inverse J-shaped curves). Shade intolerant species in contrast typically exhibit normal (unimodal) tree diameter distributions due to high mortality rates of smaller suppressed trees. In this study we explore the generality of this hypothesis which implies a causal relationship between light competition or shade tolerance and stand structure. For this purpose we formulate a partial differential equation system of stand dynamics as a function of individual tree growth, recruitment and mortality which allows us to explore possible individual-based mechanisms--e.g. light competition-underlying observed patterns of stand structure--e.g. unimodal or inverse J-shaped equilibrium diameter curves. We find that contrary to expectations interspecific differences in growth patterns can result alone in any of the two diameter distributions types observed in the field. In particular, slow growing species can present unimodal equilibrium curves even in the absence of light competition. Moreover, light competition and shade intolerance evaluated both at the tree growth and mortality stages did not have a significant impact on stand structure that tended to converge systematically towards an inverse J-shaped curves for most tree growth scenarios. Realistic transient stand dynamics for even aged stands of shade intolerant species (unimodal curves) were only obtained when recruitment was completely suppressed, providing further evidence on the critical role played by juvenile stages of tree development (e.g. the sampling stage) on final forest structure and composition. The results also point out the relevance of partial differential equations systems as a tool for exploring the individual-level mechanisms underpinning forest structure, particularly in relation to more complex forest simulation models that are more difficult to analyze and to interpret from a biological point of view.     

吉林蛟河针阔混交林功能性状分布格局及其对地形因素的响应

森林群落的构建过程及其内在机制是生态学研究的热点问题。植物功能性状是指能够代表植物的生活史策略,反映植物对环境变化响应的一系列植物属性。通过植物功能性状的分布格局及其对环境因素的响应有助于推测群落的构建过程及其内在作用机制。以吉林蛟河21.12hm²温带针阔混交林样地为研究对象,采集并测量了样地内34种木本植物的6种不同的功能性状。以20m×20m的样方为研究单元,通过计算平均成对性状距离指数（mean pairwise trait distance;PW）和平均最近邻体性状距离指数（mean nearest neighbor trait distance;NN）来探讨群落中单个性状和综合性状的分布格局。同时结合地形因子采用回归分析探讨功能性状的分布格局对局域生境变化的响应。基于PW的结果显示：单个性状中除叶面积外,其余性状的分布格局均为聚集分布多于离散分布;基于NN的结果显示：除叶面积和最大树高外,其余性状的分布格局为聚集分布多于离散分布。此外,由6种单个性状组成的综合性状的分布格局同样为聚集分布多于离散分布。基于回归分析的结果显示：森林群落中功能性状的分布格局受到海拔、坡度和坡向等因素的显著影响,而凹凸度的影响则不显著。研究结果表明包括环境过滤和生物相互作用的非随机过程能够影响温带针阔混交林的群落构建过程,中性过程对该区域群落构建过程的影响不显著。     

Strong plant-soil associations in a heterogeneous subtropical broad-leaved forest

The relative importance of deterministic and neutral processes on community assembly is currently a topic of much debate among ecologists. Analyzing species-environment associations is an effective way to assess the importance of deterministic process such as niche differentiation, but both habitat association and dispersal limitation can produce similar patterns of spatial aggregation in species. Therefore, it is crucial to control for the impact of dispersal limitation on species distributions when analyzing species-environment associations. We sampled soil with high resolutions in a 24 ha stem-mapped subtropical forest and tested plant-soil associations. We controlled for the influence of dispersal limitation by employing the homogeneous Thomas process to simulate the effect of dispersal limitation on the aggregation of tree species. After controlling for the effect of dispersal limitation, we found that the spatial heterogeneity of soil properties was associated with distributions of 88.2% (90 of 102 species) of tree species in this subtropical forest. Furthermore, not only did soil properties influence the distribution of tree species, but also tree species tended to affect properties of the soil around them. The soil factors most strongly influencing species distributions were TC, TN, TP, K, Mg, Si, soil moisture, and bulk density. We found the spatial heterogeneity of soil properties to be strongly associated with tree species distributions. Niche partitioning of soil gradients contributed substantially to species coexistence in this subtropical forest.     

The Glocal Forest

Spatial ecological patterns reflect the underlying processes that shape the structure of species and communities. Mechanisms like intra- and inter-specific competition, dispersal and host-pathogen interactions can act over a wide range of scales. Yet, the inference of such processes from patterns is a challenging task. Here we call attention to a quite unexpected phenomenon in the extensively studied tropical forest at the Barro-Colorado Island (BCI): the spatial deployment of (almost) all tree species is statistically equivalent, once distances are normalized by ℓ₀, the typical distance between neighboring conspecific trees. Correlation function, cluster statistics and nearest-neighbor distance distribution become species-independent after this rescaling. Global observables (species frequencies) and local spatial structure appear to be interrelated. This "glocality" suggests a radical interpretation of recent experiments that show a correlation between species'' abundance and the negative feedback among conspecifics. For the forest to be glocal, the negative feedback must govern spatial patterns over all scales.