A phylogenetic and trait‐based analysis of community assembly in a subtropical forest in central China

Despite several decades of study in community ecology, the relative importance of the ecological processes that determine species co‐occurrence across spatial scales remains uncertain. Some of this uncertainty may be reduced by studying the scale dependency of community assembly in the light of environmental variation. Phylogenetic information and functional trait information are often used to provide potentially valuable insights into the drivers of community assembly. Here, we combined phylogenetic and trait‐based tests to gain insights into community processes at four spatial scales in a large stem‐mapped subtropical forest dynamics plot in central China. We found that all of the six leaf economic traits measured in this study had weak, but significant, phylogenetic signal. Nonrandom phylogenetic and trait‐based patterns associated with topographic variables indicate that deterministic processes tend to dominate community assembly in this plot. Specifically, we found that, on average, co‐occurring species were more phylogenetically and functionally similar than expected throughout the plot at most spatial scales and assemblages of less similar than expected species could only be found on finer spatial scales. In sum, our results suggest that the trait‐based effects on community assembly change with spatial scale in a predictable manner and the association of these patterns with topographic variables, indicates the importance of deterministic processes in community assembly relatively to random processes.     

Phylogenetic turnover during subtropical forest succession across environmental and phylogenetic scales

Although spatial and temporal patterns of phylogenetic community structure during succession are inherently interlinked and assembly processes vary with environmental and phylogenetic scales, successional studies of community assembly have yet to integrate spatial and temporal components of community structure, while accounting for scaling issues. To gain insight into the processes that generate biodiversity after disturbance, we combine analyses of spatial and temporal phylogenetic turnover across phylogenetic scales, accounting for covariation with environmental differences. We compared phylogenetic turnover, at the species‐ and individual‐level, within and between five successional stages, representing woody plant communities in a subtropical forest chronosequence. We decomposed turnover at different phylogenetic depths and assessed its covariation with between‐plot abiotic differences. Phylogenetic turnover between stages was low relative to species turnover and was not explained by abiotic differences. However, within the late‐successional stages, there was high presence‐/absence‐based turnover (clustering) that occurred deep in the phylogeny and covaried with environmental differentiation. Our results support a deterministic model of community assembly where (i) phylogenetic composition is constrained through successional time, but (ii) toward late succession, species sorting into preferred habitats according to niche traits that are conserved deep in phylogeny, becomes increasingly important.     

功能多样性和谱系多样性对热带云雾林群落构建的影响

近年来, 功能多样性和谱系多样性为探究群落构建机制提供了新方法。为了更准确地了解海南岛高海拔热带云雾林群落构建机制, 该研究以海南岛霸王岭热带云雾林为对象, 测定7个环境因子和13个植物功能性状。利用主成分分析(PCA)筛选环境因子, 以霸王岭、尖峰岭和黎母山热带云雾林分布物种建立区域物种库, 结合模型, 分析Rao二次熵(RaoQ)和平均成对谱系距离(MPD)变化对植物群落构建的影响。结果表明: 林冠开阔度、土壤全磷含量和坡度是影响植物群落构建的关键环境因子。多数功能性状的系统发育信号很低且不显著, 说明热带云雾林群落的系统发育关系与功能性状随历史进程变化不一致。RaoQ和MPD的实际观测值都显著低于期望值, 且其标准效应值与土壤磷含量显著相关, 说明生境过滤是驱动热带云雾林群落构建的关键因子, 土壤磷含量是群落构建的关键环境筛。     

Phylogenetic structure of angiosperm communities during tropical forest succession

The phylogenetic structure of ecological communities can shed light on assembly processes, but the focus of phylogenetic structure research thus far has been on mature ecosystems. Here, I present the first investigation of phylogenetic community structure during succession. In a replicated chronosequence of 30 sites in northeastern Costa Rica, I found strong phylogenetic overdispersion at multiple scales: species present at local sites were a non-random assemblage, more distantly related than chance would predict. Phylogenetic overdispersion was evident when comparing the species present at each site with the regional species pool, the species pool found in each age category to the regional pool or the species present at each site to the pool of species found in sites of that age category. Comparing stem size classes within each age category, I found that during early succession, phylogenetic overdispersion is strongest in small stems. Overdispersion strengthens and spreads into larger size classes as succession proceeds, corroborating an existing model of forest succession. This study is the first evidence that succession leaves a distinct signature in the phylogenetic structure of communities.     

Exploring the community assembly of subalpine meadow communities based on functional traits and community phylogeny

Aims The community assembly has been a prominent issue in community ecology. This work was intended to explore the mechanisms of the species coexistence and biodiversity in communities. Our objective was to explore the mechanisms of community assembly in subalpine meadow plant communities along slope gradients in Gannan Tibetan Autonomous Prefecture, Gansu Province, Northwest China.Methods We selected five slope-oriented plots to construct a super-tree representing the species pool. We surveyed the leaf functional traits and soil environmental factors in different slopes. Then we tested the phylogenetic signal of leaf dry matter content (LDMC), specific leaf area (SLA), leaf nitrogen content (LNC) and leaf phosphorus content (LPC).Important findings The changes of slope aspect had significant influence on soil water content (SWC) and soil nutrient content. Most of the plants leaf functional traits had significant difference along different slope aspects. The LDMC was higher in south and southwest slope than north slope, while SLA, LNC and LPC were relatively high in north and northwest slope. The LPC showed feeble phylogenetic signal, while LDMC, SLA, LNC did not have a significant phylogenetic signal. With changes in the slope aspect from south to north, community phylogenetic structure shifted from over-dispersion to clustered dispersion. In south and southwest slope, habitat filtering was the driving force for community assembly. Interspecific competition was the main driving factor for community assembly in north and northwest slope aspects. But in west slope, two indices showed contrary consequence. This means the process of community assembly in west slope was more complicated and its phylogenetic index may be the result of several mechanisms working together.     

放牧干扰梯度下川西亚高山植物群落的组合机理

为了阐明放牧干扰对川西亚高山区域植物群落的组合过程以及群落结构的影响, 研究了放牧干扰梯度下的功能群均匀度和群落谱系结构的变化趋势。结果显示: 在干扰较轻的阔叶林与针叶林样地, 部分样方的功能群均匀度显著高于无效模型, 随着干扰梯度的增强, 功能群均匀度呈线性下降, 样方平均值从0.930降至0.840, 其高于无效模型的次数也逐渐降低, 干扰程度较大的草甸中出现部分样方的功能群均匀度显著低于无效模型。随着干扰程度的增强, 群落的谱系结构指数也呈逐渐上升趋势, 净关联指数平均值由-0.634逐渐增加至2.360, 邻近类群指数由-0.158上升至2.179。草甸与低矮灌丛受干扰较为严重, 其大部分样方的谱系结构指数显著高于随机群落, 表明干扰群落的谱系结构呈聚集分布。功能群均匀度与谱系结构的变化趋势一致, 表明生境筛滤效应与种间竞争作用的平衡决定着群落的组合过程。干扰降低了竞争作用, 促进了少数耐干扰功能群的优势地位, 造成功能群均匀度下降, 同时通过生境筛滤作用, 使群落的谱系结构呈现出聚集分布; 而未干扰的群落中由于竞争作用的效应, 功能群均匀度较高, 谱系结构也更加分散。研究区域植物群落的功能群均匀度与物种丰富度呈负相关, 表明物种间特别是相似物种间的竞争限制了群落的物种多样性。研究结果说明, 生态位分化和物种间的相互竞争在物种共存与群落组合中具有重要作用。     

植物群落构建机制研究进展

群落构建研究对于解释物种共存和物种多样性的维持是至关重要的,因此一直是生态学研究的中心论题。尽管近年来关于生态位和中性理论的验证研究已经取得了显著的成果,但对于局域群落构建机制的认识仍存在很大争议。随着统计和理论上的进步使得用功能性状和群落谱系结构解释群落构建机制变为可能,主要是通过验证共存物种的性状和谱系距离分布模式来实现。然而,谱系和功能性状不能相互替代,多种生物和非生物因子同时控制着群落构建,基于中性理论的扩散限制、基于生态位的环境过滤和竞争排斥等多个过程可能同时影响着群落的构建。所以,综合考虑多种方法和影响因素探讨植物群落的构建机制,对于预测和解释植被对干扰的响应,理解生物多样性维持机制有重要意义。试图在简要回顾群落构建理论及研究方法发展的基础上,梳理其最新研究进展,并探讨整合功能性状及群落谱系结构的研究方法,解释群落构建和物种多样性维持机制的可能途径。在结合功能性状和谱系结构研究群落构建时,除了考虑空间尺度、环境因子、植被类型外,还应该关注时间尺度、选择性状的种类和数量、性状的种内变异、以及人为干扰等因素对群落构建的影响。     

Plastome-based phylogeny improves community phylogenetics of subtropical forests in China

Phylogenetic trees have been extensively used in community ecology. However, how the phylogeny construction affects ecological inferences is poorly understood. In this study, we constructed three different types of phylogenetic trees (a synthetic-tree generated using V.PhyloMaker, a barcode-tree generated using rbcL+matK+trnH-psbA, and a plastome-tree generated from plastid genomes) that represented an increasing level of phylogenetic resolution among 580 woody plant species from six forest dynamic plots in subtropical evergreen broadleaved forests of China. We then evaluated the performance of each phylogeny in estimations of community phylogenetic structure, turnover and phylogenetic signal in functional traits. As expected, the plastome-tree was most resolved and most supported for relationships among species. For local phylogenetic structure, the three trees showed consistent results with Faith's PD and MPD; however, only the synthetic-tree produced significant clustering patterns using MNTD for some plots. For phylogenetic turnover, contrasting results between the molecular trees and the synthetic-tree occurred only with nearest neighbor distance. The barcode-tree agreed more with the plastome-tree than the synthetic-tree for both phylogenetic structure and turnover. For functional traits, both the barcode-tree and plastome-tree detected phylogenetic signal in maximum height, but only the plastome-tree detected signal in leaf width. This is the first study that uses plastid genomes in large-scale community phylogenetics. Our results highlight the improvement of plastome-trees over barcode-trees and synthetic-trees for the analyses studied here. Our results also point to the possibility of type I and II errors in estimation of phylogenetic structure and turnover and detection of phylogenetic signal when using synthetic-trees.     

A taxonomic and phylogenetic perspective on plant community assembly along an elevational gradient in subtropical forests

亚热带森林植物群落沿海拔梯度的分类与系统发育研究 生物多样性沿海拔梯度的分布格局已受到广泛关注。然而,生物多样性格局沿海拔梯度的变异及其潜在机制尚不清楚。整合生物多样性的多维度信息为理解群落构建机制提供了新思路。本研究在我国东部亚热带森林沿海拔270–1470 m的梯度上设置了17个木本植物固定样地,分析了沿海拔梯度植物群落 构建的生态和进化驱动力。基于样地内物种出现(0–1数据)和多度信息,计算群落内被子植物的物种和系统发育alpha和beta多样性、系统发育结构等,并量化多样性指标与微气候和地形之间的关系。研究发现,不论多度加权与否,物种alpha多样性均沿海拔升高而增加,物种和系统发育的相似性随海拔距离的增加而呈衰减趋势。然而,多度加权与否会形成不同的系统发育alpha多样性格局。对于系统发育结构而言,沿海拔增加并无明显趋势。地形和微气候是多样性格局和系统发育结构的主要驱动力。与未考虑物种多度的多样性指标相比,多度加权的指标与坡度和胸高断面积相关性更高。这些结果表明,由局域物种多度介导的确定性过程对沿海拔梯度的植物群落构建具有一定影响。     

Key characteristics for facilitating Leucaena leucocephala to successfully invade pioneer communities of tropical rain forests

Aims Due to fast-growing and high drought stress tolerance, Leucaena leucocephala has been widely used for afforestation in degraded tropical forests worldwide, but it is also a global invasive exotic species. Studies have shown that fast-growing can help L. leucocephala successfully invade subtropical forests. In this study, we aimed to investigate whether fast-growing and high drought stress tolerance can help L. leucocephala invade tropical rain forests.Methods The pioneer community of tropical rain forest which had been invaded by L. leucocephala in the Baopoling Mountain, Sanya, China was the research object. Through the t-test, we compared the differences in key functional traits that were related to growth rate (photosynthesis rate, stomatal conductance and transpiration rate) and drought stress tolerance (leaf turgor loss point) in both wet and dry seasons between L. leucocephala and eight dominant native species of pioneer community of tropical rain forest. And the principal component analysis (PCA) was used to investigate whether these functional traits can best discriminate between Leucaena leucocephala and the eight dominant native species.Important findings Leucaena leucocephala could be invariably growing fast (photosynthesis rate, stomatal conductance and transpiration rate much higher than native species) from wet to dry seasons and had higher drought stress tolerance (leaf turgor loss point much lower than native species) in the dry season. The results of PCA showed that these functional traits could significantly discriminate between L. leucocephala and the eight dominant native species. Therefore, invariable fast-growing from wet to dry season and high drought stress tolerance in the dry season make L. leucocephala successfully invade pioneer communities of tropical rain forests. In the future, these functional traits can be used to select many native species to perform biological control of L. leucocephala in other tropical forests.     

银合欢成功入侵热带雨林先锋群落的关键因素

由于生长速率高, 耐旱性强, 银合欢(Leucaena leucocephala)被广泛应用于世界各地退化热带亚热带森林的修复, 但它也是一种全球性的外来入侵植物。已经有研究发现高生长速率可以帮助银合欢成功入侵亚热带森林, 但是目前还不清楚高生长速率和强耐旱性是否能帮助银合欢成功入侵热带森林。该研究以位于中国三亚抱坡岭被银合欢入侵的热带雨林先锋群落为研究对象, 通过t检验比较干季和湿季银合欢和8个热带雨林先锋群落的本地优势种与快速生长(光合速率、气孔导度和蒸腾速率)和耐旱性(叶片膨压丧失点)紧密相关的功能性状的差异, 并利用主成分分析(PCA)研究这些功能性状是否能很好地区分银合欢和其他8个本地优势种。结果表明: 银合欢在干湿季均能快速地生长(比本地物种显著更高的光合速率、气孔导度和蒸腾速率), 且在干季拥有更强的耐旱性(比本地物种显著更低的叶片膨压丧失点)。PCA结果表明这些功能性状能够显著区分银合欢和其他8个本地优势种。因此干湿季的稳定的快速生长和干季的强耐旱性使银合欢成功入侵热带雨林先锋群落。未来可利用这些功能性状筛选合适的本地物种对入侵其他热带森林的银合欢进行有效的生物防治。     

不同空间尺度西藏色季拉山急尖长苞冷杉群落谱系结构特征

基于西藏色季拉山4 hm²亚高山暗针叶林固定样地调查数据,分析了急尖长苞冷杉群落内6个径级(1～2、2～4、4～7、7～11、11～16、>16 cm)的植物在5个空间尺度(5 m×5 m、10 m×10 m、20 m×20 m、50 m×50 m、100 m×100 m)上的谱系结构。结果表明: 不同空间尺度上,该群落的净谱系亲缘关系指数(NRI)和净最近种间亲缘关系指数(NTI)值均大于零,整体上表现为谱系聚集。在小空间尺度(5 m×5 m)上,谱系结构聚集强度最大。在所有空间尺度上,群落内小径级个体表现为谱系聚集,随着径级的增大(DBH>7 cm),逐渐表现为谱系发散,且发散程度随径级增加而增大。小径级个体的谱系聚集强度随空间尺度而增大;而大径级个体(DBH>7 cm)的谱系发散程度随空间尺度增加而增大。小径级群落的生境过滤和大径级群落的竞争排斥作用可能是驱动该区域急尖长苞冷杉群落构建的主要生态学过程。     

Australian Tropical and Subtropical Rain Forest Community Assembly: Phylogeny,Functional Biogeography,and Environmental Gradients

To compare community assemblage patterns in tropical northeastern and subtropical central eastern Australia across selected gradients and scales, we tested the relationship of species traits with phylogenetic structure, and niche breadth. We considered phylogenetic relationships across current‐day species in assemblages in relation to rain forest species pool sizes, and trait values along gradients including elevation and latitude. Trait values were quantified across scales for seed size, leaf area, wood density and maximum height at maturity for 1137 species and 596 assemblages using trait gradient analysis (TGA). Local assemblages of subtropical species had narrower trait ranges, and higher niche breadth values than corresponding assemblages of tropical species. Leaf size and seed size increased at low latitudes, and community phylogenetic structure was most strongly correlated with seed traits in the subtropics, reflecting dispersal and re‐colonization processes. Elevation accounted for little of the variance in community phylogenetic structure or trait variation across local and regional scales. Stable moist forest areas retained many species from ancestral rain forest lineages across a range of temporally conserved habitats; species within assemblages were less related; and rain forest assemblages had higher functional diversity, but lower niche breadth. This suggests that on average, assemblages of species in stable areas had greater trait variation and narrower distributions. Historic and recent rain forest contraction and re‐expansion can result in recolonized areas that are dominated by species that are more related (phylogenetically) than by chance, have smaller, widely dispersed seeds, and greater niche breadth (broader distributions).     

Response of recruitment to light availability across a tropical lowland rain forest community

1.  Many hypotheses about species coexistence involve differential resource use and trade-offs in species' life-history traits. Quantifying resource use across most species in diverse communities, although, has seldom been attempted.
2.  We use a hierarchical Bayesian approach to quantify the light dependence of recruitment in 263 woody species in a 50-ha long-term forest census plot in Panama. Data on sapling recruitment were obtained using the 1985–1990 and 1990–1995 census intervals. Available light was estimated for each recruit from yearly censuses of canopy density.
3.  We use a power function (linear log–log relationship) to model the light effect on recruitment. Different responses of recruitment to light are expressed by the light effect parameter b . The distribution of b had a central mode at 0.8, suggesting that recruitment of many species responds nearly linearly to increasing light.
4.  Nearly every species showed increases in recruitment with increasing light. Just nine species (3%) had recruitment declining with light, while 198 species (75%) showed increasing recruitment in both census intervals. Most of the increases in recruitment were decelerating, i.e. the increase was less at higher light ( b  < 1). In the remaining species, the response to light varied between census intervals (24 species) or species did not have recruits in both intervals (41 species).
5.   Synthesis. Nearly all species regenerate better in higher light, and recruitment responses to light are spread along a continuum ranging from modest increase with light to a rather strict requirement for high light. These results support the hypothesis that spatio-temporal variation in light availability may contribute to the diversity of tropical tree species by providing opportunities for niche differentiation with respect to light requirements for regeneration.     

Using phylogeny and functional traits for assessing community assembly along environmental gradients: A deterministic process driven by elevation

Community assembly processes is the primary focus of community ecology. Using phylogenetic‐based and functional trait‐based methods jointly to explore these processes along environmental gradients are useful ways to explain the change of assembly mechanisms under changing world. Our study combined these methods to test assembly processes in wide range gradients of elevation and other habitat environmental factors. We collected our data at 40 plots in Taibai Mountain, China, with more than 2,300 m altitude difference in study area and then measured traits and environmental factors. Variance partitioning was used to distinguish the main environment factors leading to phylogeny and traits change among 40 plots. Principal component analysis (PCA) was applied to colligate other environment factors. Community assembly patterns along environmental gradients based on phylogenetic and functional methods were studied for exploring assembly mechanisms. Phylogenetic signal was calculated for each community along environmental gradients in order to detect the variation of trait performance on phylogeny. Elevation showed a better explanatory power than other environment factors for phylogenetic and most traits’ variance. Phylogenetic and several functional structure clustered at high elevation while some conserved traits overdispersed. Convergent tendency which might be caused by filtering or competition along elevation was detected based on functional traits. Leaf dry matter content (LDMC) and leaf nitrogen content along PCA 1 axis showed conflicting patterns comparing to patterns showed on elevation. LDMC exhibited the strongest phylogenetic signal. Only the phylogenetic signal of maximum plant height showed explicable change along environmental gradients. Synthesis. Elevation is the best environment factors for predicting phylogeny and traits change. Plant's phylogenetic and some functional structures show environmental filtering in alpine region while it shows different assembly processes in middle‐ and low‐altitude region by different trait/phylogeny. The results highlight deterministic processes dominate community assembly in large‐scale environmental gradients. Performance of phylogeny and traits along gradients may be independent with each other. The novel method for calculating functional structure which we used in this study and the focus of phylogenetic signal change along gradients may provide more useful ways to detect community assembly mechanisms.     

Disentangling environmental and spatial processes of community assembly in tropical forests from local to regional scales

Understanding patterns and mechanisms of variation in the compositional structure of communities across spatial scales is one of the fundamental challenges in ecology and biogeography. In this study, we evaluated the effects of spatial extent (i.e. size of study region) on: 1) whether community composition can be better explained by environmental (i.e. niche‐based) or spatial (e.g. dispersal‐based) processes ; and 2) how climate and soils contribute to the influence of environment on plant community composition. We surveyed community composition across a network of 398 forest plots spanning a ～4000 m elevational gradient in the Madidi region in northwestern Bolivia. Using redundancy analyses and hierarchical variation partitioning, we disentangled the effects of environmental and spatial predictors on species composition, further decomposing the environmental effect between its climatic and soil components. We repeated analyses for 200 sub‐regions ranging in spatial extent from ～250 to ～17 500 km². Our analyses show a high degree of idiosyncrasy in results that come from different sub‐regions. Despite this variability, we were able to identify various important patterns in the structure of tropical plant communities in our study system. First, even though sub‐regions varied in size by nearly two orders of magnitude, the total amount of explained variation in community composition was scale independent; at all spatial scales, environment and space accounted for about 25% of the differences in community composition among plots. Second, the measured environmental effect was higher than the spatial effect on average and in the vast majority of sub‐regions. This was true regardless of the spatial extent of analysis. Finally, we found that both climatic and soil variables accounted for significant fractions of variation, but climate was always more important than soils.     

The community ecology of Asian rain forests,in relation to catastrophic events

Although natural catastrophic disturbance of tropical forests in Asia can be caused by volcanism and earthquakes climate-induced catastrophes are most widespread. These are prevalent and most diverse at high tropical latitudes because of the single annual dry and wet monsoon. Comparative studies indicate that periodicity of catastrophes may influence forest physiognomy, structure and species richness but long-term research in sample plots suggests that a variety of other factors are locally influential. The importance of accounting for natural catastrophes in silvicultural protocols is stressed, and research priorities identified.