后河自然保护区珍稀植物群落谱系结构的时空变化

群落中所有物种对的亲缘关系程度构成群落的谱系结构。研究群落的谱系结构可以有效地揭示环境过滤、竞争排斥、中性过程三大生态过程对群落构建的相对重要性。本文以湖北后河自然保护区1hm。固定样地的珍稀植物群落为研究对象,分别研究了5m×5m、10m×10m、20m×20m、30m×30m、40m×40m、50m×50m6个空间尺度和在10m×10m、20m×20m、30m×30m、40m×40m、50m×50m5个空间尺度下5cm≤DBH〈10cm、10cm≤DBH〈15cm、15cm≤DBH〈20cm、DBH≥20cm4个径级对亚热带常绿落叶阔叶混交林群落谱系结构的影响。结果发现：随着空间尺度的增加,谱系结构逐渐趋向聚集。随着径级的增大,群落谱系聚集程度越来越小。但当DBHI〉20cm时,群落谱系聚集度较高。在25m。的尺度上群落谱系结构表现为随机,而在100m^2及更大的尺度上群落谱系结构都表现为聚集,说明在亚热带常绿阔叶落叶混交林的群落构建过程中,小尺度上中性过程可能起主导作用,而大尺度上生态位过程可能更重要。     

山西霍山植物群落谱系结构的空间格局

以山西霍山七里峪植物群落为研究对象,用R语言分别对9个海拔梯度(1 200~2 000m)、5个空间尺度(100、400、900、1 600和2 500m2)和5个空间尺度下划分的6个径级(Ⅰ.DBH5cm、Ⅱ.5cm≤DBH10cm、Ⅲ.10cm≤DBH15cm、Ⅳ.15cm≤DBH20cm、Ⅴ.20cm≤DBH25cm、Ⅵ.DBH≥25cm)的植物群落谱系结构及其在不同研究方向上的变化规律进行分析,探讨其群落构建成因。结果显示:(1)在所有海拔梯度上,霍山七里峪植物群落都表现出一定的谱系结构,在不同海拔梯度上,群落谱系结构表现出一定的差异性,且在中低海拔群落的谱系结构是发散的,在较高海拔群落的谱系结构是聚集的。(2)5个空间尺度下研究区植物群落都具有谱系结构,且随着空间尺度的增加,谱系聚集程度越来越大。(3)不同径级下的群落都有一定的谱系结构,随着径级的增大,谱系聚集程度逐渐降低,趋向于发散结构,但在Ⅵ径级时群落聚集程度较Ⅳ、Ⅴ径级的高。研究表明,山西霍山七里峪植物群落在不同海拔梯度、不同空间尺度和不同径级下都表现出显著的谱系结构,与中性理论所预测的谱系随机并不相符,证明生态位理论在霍山植物群落构建过程中的作用更加重要。     

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。     

不同时间和空间尺度上台湾水青冈群落谱系结构动态变化

群落谱系结构是了解群落聚群过程的一个基础研究内容。但是现有大部分研究内容集中在群落组成和结构的时空差异,对谱系结构的动态变化研究较少。以浙江省清凉峰国家级自然保护区的国家二级保护植物台湾水青冈(Fagus hayatae)林为研究对象,利用2006年、2011年、2016年3次群落动态调查数据,从10、20、50m的3个空间尺度上研究该群落在10a间的谱系结构动态变化,分析时空尺度对台湾水青冈群落谱系结构的影响,探究调控台湾水青冈群落动态变化的主要因子,为后期台湾水青冈林的保护提供理论基础。研究发现(1)在10a森林动态变化过程中,群落的MPD指数下降,MNTD指数增加。NRI和NTI指数在大尺度上随时间显著增加,但在小尺度上无显著变化。(2)随着空间尺度的增加,上述指数的动态变化趋势均不断增强。以上结果表明,2006—2016年间台湾水青冈群落总体谱系结构表现出不断聚集的趋势,而近缘种的谱系关系则不断疏远;群落谱系结构的聚集趋势随空间尺度的增加而增强。大尺度上的环境过滤和小尺度上的随机过程和种间竞争作用可能是导致该地区台湾水青冈群落谱系结构动态变化的主要生态学过程。     

山西芦芽山针叶林草本层群落谱系结构与多样性的海拔格局

物种多样性格局及其成因是生态学的经典问题之一。谱系方法将演化信息(即物种亲缘关系)整合到群落学研究中,给群落学研究带来了全新视角。该研究以山西芦芽山针叶林冠层下草本群落为研究对象,将群落物种组成数据、厘定有进化时间的谱系树与地形、土壤等生境数据相结合,运用一般线性回归、基于距离矩阵的多元回归和谱系结构主坐标分析(PCPS)等统计方法,探究了草本群落谱系结构与谱系多样性的海拔格局及其与生境因子的关系。结果表明:所有样地的谱系结构指标均落在随机置信带之内,说明中性因素对草本群落构建的影响较为强烈。同时,谱系结构与海拔梯度之间存在显著的正线性关系,暗示确定性过程的作用也不可忽略,种间竞争排斥可能促进了低海拔地区谱系结构呈发散趋势,而高海拔的生境过滤引发了谱系聚集趋势。基于最近谱系距离的beta多样性与海拔差异之间呈极显著正相关关系,同样反映了草本层物种集聚受到了生境过滤的显著影响。PCPS模型前两轴代表了55.9%的谱系-物种组成变异,海拔是引起该变异的关键因子;PCPS二维排序图显示,高海拔地区群落的谱系聚集可能由菊科、禾本科等大科所主导,低海拔则显示出较高的演化分支多样性,从而表现出谱系发散。综上所述,该研究揭示了山西芦芽山山地针叶林冠层下草本群落谱系多样性格局,并初步明晰了主要生态过程对物种集聚的影响规律。     

不同海拔矮嵩草与火绒草种群分布格局及空间关联性

研究植物种群的分布格局及空间关联性是揭示植物种群与环境间关系、自身特性以及种间相互作用的一种重要手段。本研究以甘南高寒草甸为对象,沿海拔梯度(2900、3500、3800 m)设置3个样地,采用Ripley K函数对不同海拔矮嵩草(Kobresia humilis)与火绒草(Leontopodium nanum)种群的空间分布格局及种间关联性进行了分析。结果表明:(1)随着海拔的上升,矮嵩草种群的重要值、盖度、株数等呈递增趋势,火绒草种群的重要值、盖度、株数等则呈递减趋势。(2)矮嵩草种群在不同海拔的研究尺度内均表现为聚集分布,但其聚集的程度有所不同;火绒草种群在海拔2900 m处呈聚集分布状态;海拔3500 m处,火绒草种群在0～2.8 m尺度上呈聚集分布,但其聚集强度随着研究尺度的增加而逐渐减弱,最终转变为随机分布; 3800 m处,火绒草种群在0～2 m尺度内呈聚集分布,在较大尺度(2～5 m)上呈随机分布。(3)海拔2900 m处,矮嵩草与火绒草种群在小尺度(0～1.1 m)内表现为正相关,较大尺度上表现为无相关;海拔3500 m处,在整个研究尺度内表现为负相关;海拔3800 m处,在小尺度范围(0～1.4 m)内表现为无相关,较大尺度上表现为负相关,且随研究尺度的增大,其关联性增强。     

不同程度人为干扰对古田山森林群落谱系结构的影响

群落谱系结构包含群落发育的历史信息,能从新的角度反映群落形成的生态过程.作者在浙江古田山自然保护区亚热带常绿阔叶林中选择了人为干扰强度不同的4种群落类型,以20 m×20 m为研究尺度探讨了不同干扰程度对群落谱系结构的影响.结果表明,人工林(类型Ⅰ)谱系结构发散;但间伐林(类型Ⅱ)、自然恢复林(类型Ⅲ)以及自然老龄林(类型Ⅳ)谱系结构聚集,且以类型Ⅱ和Ⅳ聚集度最高.进一步分析不同径级谱系结构发现,在中小径级(DBH≤5 cm和5 cm10 cm时,除了类型Ⅳ,其他3种群落都是谱系结构发散,这反映了在恢复早期种子扩散对这些林型群落构建影响较大:而皆伐后的演替和间伐增加了群落生境异质性,生境过滤作用增强,使类型Ⅱ、Ⅲ群落中小径级谱系结构表现为聚集;类型Ⅳ群落不同径级谱系结构均表现为聚集,可能与其稳定的生境过滤作用有关.     

长白山蒙古栎次生林群落结构特征及优势树种空间分布格局

群落结构特征研究是揭示植物群落维持机制和演替动态的基础,是合理安排森林经营活动的基本前提.以吉林省汪清林业局2块1 hm²的蒙古栎阔叶混交林固定样地为对象,研究了长白山蒙古栎次生林的群落结构特征,并采用点格局O-ring统计法对群落内优势树种的空间分布格局进行分析.结果表明: 两块样地都是以蒙古栎为优势树种的群落类型,都具有明显的层次结构,主要伴生树种存在差异,样地Ⅰ以大青杨、白桦、红松为主,样地Ⅱ以红松、紫椴和色木槭为主.且样地Ⅰ的树种数量、Shannon多样性指数高于样地Ⅱ.两块样地中所有个体的径级分布呈倒“J”型,蒙古栎径级结构呈近似正态型,红松呈倒J型,其他主要伴生树种径级结构存在一定差异.两块样地0～50 m尺度上蒙古栎呈现小尺度聚集、中大尺度随机分布的格局,红松呈先聚集、后随机的分布格局,但样地Ⅰ内红松聚集的尺度和强度均大于样地Ⅱ.样地Ⅰ白桦和大青杨均在0～17 m尺度上呈聚集分布、且聚集强度显著高于其他树种,而在18～50 m尺度分别呈现随机分布和均匀-随机分布.样地Ⅱ中紫椴基本呈现随机分布,色木槭在中大尺度上呈随机或均匀分布,聚集分布集中在小尺度.两块样地处于演替早期阶段的不同发育阶段,相比样地Ⅰ,样地Ⅱ的演替阶段更高、群落相对更加稳定.     

物种谱系不确定性对群落谱系格局指标的影响

物种谱系关系常被用于衡量群落谱系格局及推断格局背后的生态过程,但多数研究往往忽视谱系关系的不确定性及其可能对群落谱系格局造成的影响.为此,本文以浙江天童20 hm^2样地内150个树种为研究对象,采用这些物种叶绿体DNA的rbcL和matK碱基序列构建1棵一致系统发育树和反映谱系不确定性的999棵系统发育树,然后结合样地物种分布数据计算标准化净亲缘指数(NRI)和最近亲缘指数(NTI),最后运用独立置换零模型衡量样地群落谱系格局.结果表明:物种系统发育树在拓扑结构和物种谱系分支节点年龄上均存在较大的不确定性,谱系不确定性随着谱系分支节点年龄的减小而增大,也随物种间平均谱系距离的增加而增加;在样方尺度上,物种谱系的不确定性增加了标准化NRI和NTI指数的变异,但对两个指数的影响几乎独立;其对两指数的空间分布影响不同,且程度不一,其中标准化NRI受到的影响相对更大;在群落尺度上,物种谱系的不确定性增加了标准化NRI和NTI的变异,平均变异系数分别为0.37和0.077,表明群落水平的标准化NRI更易受到谱系不确定性的影响.这说明物种谱系不确定性会传递到常用的群落谱系格局指标中,且不同指标受影响的程度不同,进而影响对群落谱系格局的衡量及相关生态过程的推断.该结论也暗示以往不考虑谱系不确定性的研究中,非随机的群落谱系格局比例可能被高估.     

群落构建研究的新进展:进化和生态相结合的群落谱系结构研究

群落如何构建足群落生态学中的重要问题.群落谱系结构研究将物种间的亲缘进化关系运用到群落生态学研究中,利用物种的系统发育状况推测历史因素对现有群落的影响,为推断影响群落组成的生态学机制提供了有效方法.群落谱系结构的研究方法是首先建立可代表群落物种库的超级系统进化树,然后计算群落内物种间的谱系距离,最后通过统计方法检测其与随机模型下的谱系距离是否有显著差异来获得谱系结构(如谱系聚集、谱系发散),从而揭示群落构建中的关键生态过程(如生境过滤、竞争作用).群落谱系结构与空间尺度、分类群尺度、时间尺度等不同研究尺度有关.在小的空间尺度下,随着分类群尺度降低、树木年龄级增大,群落谱系结构从聚集逐渐转为发散;而随群落空间尺度的增大,谱系趋向于聚集.谱系结构受到环境因素影响,因此分析集合群落下的谱系可以揭示区域生态过程的影响.另外,群落谱系结构研究还有助于探讨中性理论、密度制约假说等生态学理论,并预测干扰作用下的群落演化趋势.在利用谱系结构深入探讨群落构建成因时,需要基于生态特征和环境变量共同分析,同时考虑小尺度局域过程(群落的微环境或群落内种间相互作用等)和大尺度区域过程(地史过程和物种形成等),并可结合生态控制实验,以确认群落构建的关键因素.在研究方法和手段上,今后需要注重通过选择合适的基因片段建立系统树,然后通过生态特征来加以校正,以更准确地反映物种间的亲缘距离.另外,获得谱系树后还需要寻找更加合理的统计模型和指数,增加统计分析和解决问题的能力.     

Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests

Patterns of phylogenetic relatedness within communities have been widely used to infer the importance of different ecological and evolutionary processes during community assembly, but little is known about the relative ability of community phylogenetics methods and null models to detect the signature of processes such as dispersal, competition and filtering under different models of trait evolution. Using a metacommunity simulation incorporating quantitative models of trait evolution and community assembly, I assessed the performance of different tests that have been used to measure community phylogenetic structure. All tests were sensitive to the relative phylogenetic signal in species metacommunity abundances and traits; methods that were most sensitive to the effects of niche-based processes on community structure were also more likely to find non-random patterns of community phylogenetic structure under dispersal assembly. When used with a null model that maintained species occurrence frequency in random communities, several metrics could detect niche-based assembly when there was strong phylogenetic signal in species traits, when multiple traits were involved in community assembly, and in the presence of environmental heterogeneity. Interpretations of the causes of community phylogenetic structure should be modified to account for the influence of dispersal.     

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。     

Trait evolution, community assembly, and the phylogenetic structure of ecological communities

Taxa co-occurring in communities often represent a nonrandom sample, in phenotypic or phylogenetic terms, of the regional species pool. While heuristic arguments have identified processes that create community phylogenetic patterns, further progress hinges on a more comprehensive understanding of the interactions between underlying ecological and evolutionary processes. We created a simulation framework to model trait evolution, assemble communities (via competition, habitat filtering, or neutral assembly), and test the phylogenetic pattern of the resulting communities. We found that phylogenetic community structure is greatest when traits are highly conserved and when multiple traits influence species membership in communities. Habitat filtering produces stronger phylogenetic structure when taxa with derived (as opposed to ancestral) traits are favored in the community. Nearest-relative tests have greater power to detect patterns due to competition, while total community relatedness tests perform better with habitat filtering. The size of the local community relative to the regional pool strongly influences statistical power; in general, power increases with larger pool sizes for communities created by filtering but decreases for communities created by competition. Our results deepen our understanding of processes that contribute to phylogenetic community structure and provide guidance for the design and interpretation of empirical research.     

Stochastic and deterministic assembly processes in subsurface microbial communities

A major goal of microbial community ecology is to understand the forces that structure community composition. Deterministic selection by specific environmental factors is sometimes important, but in other cases stochastic or ecologically neutral processes dominate. Lacking is a unified conceptual framework aiming to understand why deterministic processes dominate in some contexts but not others. Here we work toward such a framework. By testing predictions derived from general ecological theory we aim to uncover factors that govern the relative influences of deterministic and stochastic processes. We couple spatiotemporal data on subsurface microbial communities and environmental parameters with metrics and null models of within and between community phylogenetic composition. Testing for phylogenetic signal in organismal niches showed that more closely related taxa have more similar habitat associations. Community phylogenetic analyses further showed that ecologically similar taxa coexist to a greater degree than expected by chance. Environmental filtering thus deterministically governs subsurface microbial community composition. More importantly, the influence of deterministic environmental filtering relative to stochastic factors was maximized at both ends of an environmental variation gradient. A stronger role of stochastic factors was, however, supported through analyses of phylogenetic temporal turnover. Although phylogenetic turnover was on average faster than expected, most pairwise comparisons were not themselves significantly non-random. The relative influence of deterministic environmental filtering over community dynamics was elevated, however, in the most temporally and spatially variable environments. Our results point to general rules governing the relative influences of stochastic and deterministic processes across micro- and macro-organisms.     

The merging of community ecology and phylogenetic biology

The increasing availability of phylogenetic data, computing power and informatics tools has facilitated a rapid expansion of studies that apply phylogenetic data and methods to community ecology. Several key areas are reviewed in which phylogenetic information helps to resolve long-standing controversies in community ecology, challenges previous assumptions, and opens new areas of investigation. In particular, studies in phylogenetic community ecology have helped to reveal the multitude of processes driving community assembly and have demonstrated the importance of evolution in the assembly process. Phylogenetic approaches have also increased understanding of the consequences of community interactions for speciation, adaptation and extinction. Finally, phylogenetic community structure and composition holds promise for predicting ecosystem processes and impacts of global change. Major challenges to advancing these areas remain. In particular, determining the extent to which ecologically relevant traits are phylogenetically conserved or convergent, and over what temporal scale, is critical to understanding the causes of community phylogenetic structure and its evolutionary and ecosystem consequences. Harnessing phylogenetic information to understand and forecast changes in diversity and dynamics of communities is a critical step in managing and restoring the Earth's biota in a time of rapid global change.     

Fire modifies the phylogenetic structure of soil bacterial co‐occurrence networks

Fire alters ecosystems by changing the composition and community structure of soil microbes. The phylogenetic structure of a community provides clues about its main assembling mechanisms. While environmental filtering tends to reduce the community phylogenetic diversity by selecting for functionally (and hence phylogenetically) similar species, processes like competitive exclusion by limiting similarity tend to increase it by preventing the coexistence of functionally (and phylogenetically) similar species. We used co‐occurrence networks to detect co‐presence (bacteria that co‐occur) or exclusion (bacteria that do not co‐occur) links indicative of the ecological interactions structuring the community. We propose that inspecting the phylogenetic structure of co‐presence or exclusion links allows to detect the main processes simultaneously assembling the community. We monitored a soil bacterial community after an experimental fire and found that fire altered its composition, richness and phylogenetic diversity. Both co‐presence and exclusion links were more phylogenetically related than expected by chance. We interpret such a phylogenetic clustering in co‐presence links as a result of environmental filtering, while that in exclusion links reflects competitive exclusion by limiting similarity. This suggests that environmental filtering and limiting similarity operate simultaneously to assemble soil bacterial communities, widening the traditional view that only environmental filtering structures bacterial communities.     

Patterns of phylogenetic community structure of sand dune plant communities in the Yucatan Peninsula: the role of deterministic and stochastic processes in community assembly

ABSTRACT

Background: Tropical sand dunes are ideal systems for understanding drivers of community assembly as dunes are subject to both deterministic and stochastic processes. However, studies that evaluate the factors that mediate plant community assembly in these ecosystems are few.

Aims: We evaluated phylogenetic community structure to elucidate the role of deterministic and stochastic processes in mediating the assembly of plant communities along the north of the Yucatan Peninsula, Mexico.

Methods: We used plastid genetic markers to evaluate phylogenetic relationships in 16 sand-dune communities. To evaluate the role of climate in shaping plant community structure we carried out linear regressions between climatic variables and mean phylogenetic distance. We estimated the Net Relatedness Index and Nearest Taxon Index to identify ecological processes mediating community assembly.

Results: Observed phylogenetic structure was not different from random, suggesting that stochastic processes are the major determinants of community assembly. Climate was slightly correlated with phylogenetic diversity suggesting that abiotic environment plays a minimal role in community assembly.

Conclusions: Random assembly appears to be the primary factor structuring the studied sand dune plant communities. Environmental filters may represent a secondary factor contributing to the observed phylogenetic structure. Thus, both processes may act simultaneously to mediate the assembly of sand-dune plant communities.     

柴达木盆地荒漠灌丛群落谱系结构研究

谱系信息是群落生态学和保育生物学研究的主要内容之一。为探究柴达木盆地荒漠灌丛群落谱系结构及其与环境因子的关系,该研究以柴达木盆地荒漠灌丛为对象,基于群落中物种存在与否的物种组成数据,使用R语言中picante软件包计算了灌丛群落谱系多样性指数和谱系结构指数,并且分析了谱系结构指数与年均温度、年均降水以及土壤含水量之间的关系,以揭示柴达木盆地灌丛群落物种之间的亲缘关系和群落生物多样性维持机制。结果表明:(1)柴达木盆地灌丛群落谱系结构与土壤含水量之间存在极显著相关性(P=2.77×10-6),随着土壤含水量的增加,群落谱系结构聚集程度逐渐降低,表现出生境过滤作用逐步减弱,生物间相互作用逐渐加强的变化趋势。(2)群落谱系结构与年均温度、年均降水之间无显著相关性。(3)典型荒漠生境灌丛和河谷(河漫滩)生境灌丛群落的谱系结构差异显著(P0.05),整体上分别表现为谱系聚集状态和谱系发散状态;河谷(河漫滩)生境灌丛群落的谱系多样性显著高于典型荒漠生境灌丛(P0.05)。(4)将群落谱系信息应用在生物多样性保护实践中,发现河谷(河漫滩)生境灌丛群落较典型荒漠生境的灌丛群落可能具有更高的保护价值。研究认为,将群落谱系研究与保育生物学理论结合将会使生物多样性保护策略更加科学有效。