群落构建的中性理论和生态位理论

物种共存和生物多样性维持一直是生态学研究的中心论题。基于物种生态位分化的群落构建理论已经发展了近一个世纪, 但我们对群落构建和生物多样性维持的机理仍然不清楚。近年来, 群落中性理论以其简约性和预测能力成为群落生态学研究的焦点, 但由于其“物种在生态功能上等价”的假设与大量研究结果相悖, 同时对自然群落结构的准确预测也只限于少数的生态系统, 因而饱受质疑。如今, 越来越多的生态学家认为群落构建的生态位理论与中性理论之争的最终归宿应该是二者的整合。 在本文中, 我们在简要回顾生态位理论和群落中性理论发展的基础上, 分析二者之间的主要分歧和互补性, 试图梳理二者整合的途径。我们认为, 尽管中性理论的发展极大地丰富了群落构建理论, 但二者的整合尚处于初级阶段; 群落构建零模型假说、中性—生态位连续体假说、随机生态位假说等都不失为有价值的尝试, 今后需要在其他类型的生态系统中进行实验验证, 以更好地理解确定性过程和随机过程在决定群落构建和生物多样性维持中的作用。     

水淹干扰下水库水滨带植物群落谱系结构与物种多样性格局

群落谱系结构将物种系统发育信息整合到群落中,给生物多样性研究提供了全新视角。为探讨水淹干扰下水库水滨带植物群落物种多样性维持机制,以丹江口水库水滨带为研究区域,以南水北调中线一期工程运行前的原有水滨带植被为研究对象,分析不同水淹干扰强度下群落谱系结构和物种多样性及其与环境因素的关系。结果表明：(1)随着水淹干扰强度增加,群落物种丰富度、物种多样性和谱系多样性显著降低。谱系结构指数虽无显著差异,但随着水淹干扰强度增加呈现由发散到聚集的变化趋势。表明水滨带植物群落物种多样性维持机制是生境过滤和生态位分化共同驱动的结果,并且随着水淹干扰强度增加呈现生境过滤作用增加、物种间竞争排斥作用减弱的生态过程影响规律。(2)坡度、土壤质地、速效氮和全氮含量对整个水滨带植物群落物种丰富度、物种多样性和谱系多样性影响显著。此外,土层厚度和速效磷对物种丰富度和土层厚度对谱系多样性也影响显著。而仅有速效氮含量对谱系结构指数影响显著;轻度水淹干扰下坡向、速效磷含量与多样性指数显著相关;中度水淹干扰下坡度、土层厚度、土壤质地、土壤总氮含量与多样性指数显著相关,土壤pH、全氮和全磷含量与谱系结构指数显著相关;重度水淹...     

基于生态位理论研究群落构建的方法进展

理解群落构建过程可以解释生物多样性格局的形成和维持,对于生物多样性保护起到关键作用。生态位理论是群落构建研究的核心框架之一。该理论认为群落构建是生物作用和非生物作用将区域物种库中的物种选入局域群落的确定过程。近年来,随着该领域受到的关注越来越多,研究者不但从物种、谱系或功能等生物多样性维度来研究群落构建,所使用的多样性指数、零模型算法和物种库界定方式等也多种多样。本文回顾了从生物多样性不同维度研究群落构建的优势与局限,总结了群落构建过程中构建零模型和界定物种库时需要注意的一些问题,介绍了部分群落构建研究的最新方法学进展和研究成果。最后,结合近年来的研究案例提出了对未来群落构建研究的一些建议。     

澳门城市栖息地斑块中鸟类群落功能和谱系多样性

了解群落构建过程是生态学的核心和基本问题,这对于解释物种共存和物种多样性的维持,完善生物多样性保护政策至关重要。与传统的物种多样性相比,功能和谱系多样性能够提供更多的信息,特别是在群落构建过程方面提供深刻见解。2014年5月—2017年1月,采用样线法对澳门地区5个城市栖息地斑块中(生态一区、鹭鸟林、赛马场滩涂、关闸口岸滩涂和莲花桥侧红树林)的鸟类进行繁殖季和越冬季的调查,共调查14次。利用鸟类丰富度和多度来表征群落的物种多样性,其中多度以最大值保留法计;功能和谱系多样性分别采用功能丰富度(functional richness:FRic)和Faith'PD。同时,利用基于多度加权的平均成对功能距离(mean pairwise functional distance,MFD)和平均成对谱系距离(mean pairwise phylogenetic distance,MPD)来表征群落的功能和谱系结构。结果表明:共记录98种鸟类,隶属于14目32科,目、科、种中数量最多的依次为,雀形目鸟类最多为47种,鹭科物种数为11种,白鹭多度为849只;鸟类物种丰富度(68种)和PD在生态一区最高,多度在莲花桥侧红树林最高为2940只,MFD和FRic在赛马场滩涂最高,MPD在生态一区最高;斑块面积在鸟类丰富度、多度、FRic和PD中具有普遍重要性,植被面积和滩涂面积则分别强烈影响着MFD和MPD;关闸口岸滩涂和莲花桥侧红树林的鸟类群落SES.MFD显著低于随机值(P <0. 05),鹭鸟林和赛马场滩涂的鸟类群落SES. MPD显著低于随机值(P<0.05),表明关闸口岸滩涂和莲花桥侧红树林鸟类群落功能聚集(functional clustering),可能与生境过滤驱动群落构建有关。栖息地斑块的特征差异导致了鸟类群落构建机制的不同,这是仅仅考虑物种多样性难以提供的信息,说明了研究群落构建机制时考虑多维度多样性的必要性。     

澳门城市栖息地斑块中鸟类群落功能和谱系多样性

了解群落构建过程是生态学的核心和基本问题,这对于解释物种共存和物种多样性的维持,完善生物多样性保护政策至关重要。与传统的物种多样性相比,功能和谱系多样性能够提供更多的信息,特别是在群落构建过程方面提供深刻见解。2014年5月—2017年1月,采用样线法对澳门地区5个城市栖息地斑块中(生态一区、鹭鸟林、赛马场滩涂、关闸口岸滩涂和莲花桥侧红树林)的鸟类进行繁殖季和越冬季的调查,共调查14次。利用鸟类丰富度和多度来表征群落的物种多样性,其中多度以最大值保留法计;功能和谱系多样性分别采用功能丰富度(functional richness:FRic)和Faith'PD。同时,利用基于多度加权的平均成对功能距离(mean pairwise functional distance,MFD)和平均成对谱系距离(mean pairwise phylogenetic distance,MPD)来表征群落的功能和谱系结构。结果表明:共记录98种鸟类,隶属于14目32科,目、科、种中数量最多的依次为,雀形目鸟类最多为47种,鹭科物种数为11种,白鹭多度为849只;鸟类物种丰富度(68种)和PD在生态一区最高,多度在莲花桥侧红树林最高为2940只,MFD和FRic在赛马场滩涂最高,MPD在生态一区最高;斑块面积在鸟类丰富度、多度、FRic和PD中具有普遍重要性,植被面积和滩涂面积则分别强烈影响着MFD和MPD;关闸口岸滩涂和莲花桥侧红树林的鸟类群落SES.MFD显著低于随机值(P 0. 05),鹭鸟林和赛马场滩涂的鸟类群落SES. MPD显著低于随机值(P0.05),表明关闸口岸滩涂和莲花桥侧红树林鸟类群落功能聚集(functional clustering),可能与生境过滤驱动群落构建有关。栖息地斑块的特征差异导致了鸟类群落构建机制的不同,这是仅仅考虑物种多样性难以提供的信息,说明了研究群落构建机制时考虑多维度多样性的必要性。     

功能多样性对典型阔叶红松林生产力的影响

为比较生物量比率假说与生态位互补假说在解释生产力变异的相对重要性,探讨生物多样性和生产力之间的关系是否受到生物和非生物因素的影响,该研究依托小兴安岭9 hm~2阔叶红松(Pinus koraiensis)林动态监测样地,通过计算群落初始生物量、物种多样性、功能多样性、植物性状的群落加权平均值和测定环境因子,运用线性回归模型、结构方程模型,比较了物种多样性和功能多样性与生产力的相关性。结果表明:(1)物种多样性和功能多样性均对生产力有显著作用,功能多样性比物种多样性与生产力的关系更为密切;(2)功能多样性指数比群落加权平均值能更好地解释生产力变异,说明生态位互补假说更适用于解释阔叶红松林群落内生产力的变异;(3)生物多样性与生产力的关系受生物因素与非生物因素的共同作用,相较于多样性和功能性状组成(植被质量),初始林分生物量(植被数量)能更有效地解释生产力的变异。生物多样性与生产力关系的研究应从植被质量与植被数量同时出发,评估生态系统过程的多种非生物和生物驱动因素,同时维护森林功能多样性,加强植物与土壤环境的保护,对有效增加生产力和维持生物多样性具有重要意义。     

物种共存理论研究进展

群落内的多物种如何共存是群落生态学的核心研究内容之一。经典的物种共存理论强调物种之间的生态位分化,注重具体共存机制的研究。这种以具体共存机制为研究对象的方法一定程度上促进了当代物种共存理论框架的形成。在当代物种共存理论框架下,物种间的差异被划分为两类综合性的抽象差异——生态位差异和平均适合度差异,前者促进物种共存,对应稳定化机制;后者导致竞争排除,对应均等化机制。本文在简要回顾经典物种共存理论的基础上,介绍了当代物种共存理论的框架(包括理论的形成和定义)、基于该理论的部分实验验证工作及其在一些重要生态学问题中的应用。当代物种共存理论不仅揭示了群落内物种是如何共存的这一基本理论问题,更重要的是在全球变化的背景下该理论对生物多样性的保护和管理具有重要的应用价值。期望本文的介绍有助于国内生态学和生物多样性工作者了解当代物种共存理论,并将其应用于群落构建和生物多样性维持机制等方面的研究。     

生物多样性理论最新进展

生物多样性是生态系统复杂性的重要特征, 理解多样性的形成和维持机制一直是理论生态学研究的核心议题。本文从三方面概述了生物多样性理论的最新进展。一是物种共存和群落构建, 总结了现代共存理论和基于过程的群落构建理论的新进展。二是物种相互作用, 综述了利用经验数据推断物种相互作用关系和强度的最新方法。三是生态-进化动态, 介绍了生态-进化模型的一般框架及其在生物多样性研究中的应用。最后对生物多样性理论的发展趋势做了展望, 特别是多尺度整合理论和全球变化下的预测理论。     

功能多样性对典型阔叶红松林生产力的影响

为比较生物量比率假说与生态位互补假说在解释生产力变异的相对重要性, 探讨生物多样性和生产力之间的关系是否受到生物和非生物因素的影响, 该研究依托小兴安岭9 hm ²阔叶红松(Pinus koraiensis)林动态监测样地, 通过计算群落初始生物量、物种多样性、功能多样性、植物性状的群落加权平均值和测定环境因子, 运用线性回归模型、结构方程模型, 比较了物种多样性和功能多样性与生产力的相关性。结果表明: (1)物种多样性和功能多样性均对生产力有显著作用, 功能多样性比物种多样性与生产力的关系更为密切; (2)功能多样性指数比群落加权平均值能更好地解释生产力变异, 说明生态位互补假说更适用于解释阔叶红松林群落内生产力的变异; (3)生物多样性与生产力的关系受生物因素与非生物因素的共同作用, 相较于多样性和功能性状组成(植被质量), 初始林分生物量(植被数量)能更有效地解释生产力的变异。生物多样性与生产力关系的研究应从植被质量与植被数量同时出发, 评估生态系统过程的多种非生物和生物驱动因素, 同时维护森林功能多样性, 加强植物与土壤环境的保护, 对有效增加生产力和维持生物多样性具有重要意义。     

甲虫群落构建研究进展

甲虫作为节肢动物中一个庞大的类群,具有繁简各异的生活习性,在生态系统中发挥极为重要的作用.甲虫群落构建是群落生态研究的重要内容之一,可揭示不同生态环境下生物多样性的形成与维持机制.现阶段对甲虫群落构建的研究主要通过物种的功能性状,或将系统发育结构与物种功能性状相结合的方法,对群落构建过程进行解析,用以判断主要的驱动因素...     

Neutral lipid fatty acid composition as trait and constraint in Collembola evolution

Functional traits determine the occurrence of species along environmental gradients and their coexistence with other species. Understanding how traits evolved among coexisting species helps to infer community assembly processes. We propose fatty acid composition in consumer tissue as a functional trait related to both food resources and physiological functions of species. We measured phylogenetic signal in fatty acid profiles of 13 field‐sampled Collembola (springtail) species and then combined the data with published fatty acid profiles of another 24 species. Collembola fatty acid profiles generally showed phylogenetic signal, with related species resembling each other. Long‐chain polyunsaturated fatty acids, related to physiological functions, demonstrated phylogenetic signal. In contrast, most food resource biomarker fatty acids and the ratios between bacterial, fungal, and plant biomarker fatty acids exhibited no phylogenetic signal. Presumably, fatty acids related to physiological functions have been constrained during Collembola evolutionary history: Species with close phylogenetic affinity experienced similar environments during divergence, while niche partitioning in food resources among closely related species favored species coexistence. Measuring phylogenetic signal in ecologically relevant traits of coexisting species provides an evolutionary perspective to contemporary assembly processes of ecological communities. Integrating phylogenetic comparative methods with community phylogenetic and trait‐based approaches may compensate for the limitations of each method when used alone and improve understanding of processes driving and maintaining assembly patterns.     

The roles of environment,space, and phylogeny in determining functional dispersion of rodents (Rodentia) in the Hengduan Mountains,China

The recently described trait‐based approach is becoming widely popular for a mechanistic understanding of species coexistence. However, the greatest challenge in functional analyses is decomposing the contributions of different ecological and evolutionary processes (e.g., niche‐based process, neutral process, and evolutionary process) in determining trait structure. Taking rodents (Rodentia) in the Hengduan Mountains as our study model, we aim to (1) quantify the vertical patterns of functional structure for head–body length (HL), tail/body ratio (TR), animal component in diet (ACD), and all traits; (2) disentangle the relative importance of different assembly processes (environment, space, and phylogeny) in structuring trait dispersion; and (3) assess the feasibility of Bergmann's rule and Allen's rule along elevational gradient. Our results have suggested that the vertical functional structure pattern varied across these three traits, indicating distinct functional roles in the community assembly process. These nonrandom vertical patterns of HL, TR, and terminal ACD have demonstrated these traits were dominated by different ecological process along environmental gradient. In variance partitioning, high proportion of the spatial variations in trait dispersion was explained by environmental and spatial models, which have provided supporting strong evidence for niche‐based and neutral processes in leading species coexistence. Although the three traits all exhibited apparent phylogenetic signals, phylogenetic relationship within community failed to predict the spatial variations of functional dispersion, confirming the enormous inference of phylogenetic signals in predicting trait structure. By assessing the vertical patterns of HL and TR at order and family levels, we argued that functional adaptation along an environmental gradient is a surrogate of series of complex processes (e.g., environmental filtering, interspecific interaction, and neutral dispersal) acting on multiple functional axes, which results in inconsistence with the empirical rules along elevational gradient.     

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.     

Advances, challenges and a developing synthesis of ecological community assembly theory

Ecological approaches to community assembly have emphasized the interplay between neutral processes, niche-based environmental filtering and niche-based species sorting in an interactive milieu. Recently, progress has been made in terms of aligning our vocabulary with conceptual advances, assessing how trait-based community functional parameters differ from neutral expectation and assessing how traits vary along environmental gradients. Experiments have confirmed the influence of these processes on assembly and have addressed the role of dispersal in shaping local assemblages. Community phylogenetics has forged common ground between ecologists and biogeographers, but it is not a proxy for trait-based approaches. Community assembly theory is in need of a comparative synthesis that addresses how the relative importance of niche and neutral processes varies among taxa, along environmental gradients, and across scales. Towards that goal, we suggest a set of traits that probably confer increasing community neutrality and regionality and review the influences of stress, disturbance and scale on the importance of niche assembly. We advocate increasing the complexity of experiments in order to assess the relative importance of multiple processes. As an example, we provide evidence that dispersal, niche processes and trait interdependencies have about equal influence on trait-based assembly in an experimental grassland.     

The ecology of differences: assessing community assembly with trait and evolutionary distances

Species enter and persist in local communities because of their ecological fit to local conditions, and recently, ecologists have moved from measuring diversity as species richness and evenness, to using measures that reflect species ecological differences. There are two principal approaches for quantifying species ecological differences: functional (trait‐based) and phylogenetic pairwise distances between species. Both approaches have produced new ecological insights, yet at the same time methodological issues and assumptions limit them. Traits and phylogeny may provide different, and perhaps complementary, information about species' differences. To adequately test assembly hypotheses, a framework integrating the information provided by traits and phylogenies is required. We propose an intuitive measure for combining functional and phylogenetic pairwise distances, which provides a useful way to assess how functional and phylogenetic distances contribute to understanding patterns of community assembly. Here, we show that both traits and phylogeny inform community assembly patterns in alpine plant communities across an elevation gradient, because they represent complementary information. Differences in historical selection pressures have produced variation in the strength of the trait‐phylogeny correlation, and as such, integrating traits and phylogeny can enhance the ability to detect assembly patterns across habitats or environmental gradients.     

Incorporating intraspecific variation in tests of trait-based community assembly

Environmental filtering and niche differentiation are processes proposed to drive community assembly, generating nonrandom patterns in community trait distributions. Despite the substantial intraspecific trait variation present in plant communities, most previous studies of trait-based community assembly have used species mean trait values and therefore not accounted for intraspecific variation. Using a null model approach, I tested for environmental filtering and niche differentiation acting on three key functional traits??vegetative height, specific leaf area (SLA), and leaf dry matter content (LDMC)??in old-field plant communities. I also examined how accounting for intraspecific variation at the among-plot and individual levels affected the detection of nonrandom assembly patterns. Tests using fixed species mean trait values provided evidence of environmental filtering acting on height and SLA and niche differentiation acting on SLA. Including plot-level intraspecific variation increased the strength of these patterns, indicating an important role of intraspecific variation in community assembly. Tests using individual trait data indicated strong environmental filtering acting on all traits, but provided no evidence of niche differentiation, although these signals may have been obscured by the effects of dispersal limitation and spatial aggregation of conspecific individuals. There was also strong evidence of nonrandom assembly of individuals within single species, with the strength of environmental filtering varying among species. This study demonstrates that, while analyses using fixed species mean trait values can provide insights into community assembly processes, accounting for intraspecific variation provides a more complete view of communities and the processes driving their assembly.     

Environment differentially affects the functional and phylogenetic structures of plant communities in a dry evergreen Afromontane tropical forest

Testing how local environmental conditions influence plant community assembly is important to understand the underlying mechanisms that promote and/or maintain biodiversity. Functional traits are used to find the broad spectrum of resource use strategies that plants use to respond to environmental variation. The patterns and drivers of plant community assembly through the lens of traits and phylogeny; however, remain to be studied in a uniquely biodiversity rich but poorly known fragmented dry Afromontane forest of Ethiopia. Here, we combined trait and community phylogenetic data from thirty sampling plots of 20 × 20 m size to determine the functional and phylogenetic structures and their drivers in a fragmented, human-dominated dry evergreen Afromontane forest. We found phylogenetic and functional clustering of plants in which the effect of environment was found to be trait specific. A weak phylogenetic signal for traits was detected suggesting that species resource use strategies may not be inferred using species phylogenetic distance. Additionally, we found functional traits to be weak in predicting species abundance distribution. Overall, while this study shows a non-random community assembly pattern, it also highlights the importance of deterministic processes being trait specific.     

不同纬度植物群落系统发育与功能性状结构研究

研究植物群落系统发育和功能性状结构有助于了解植物多样性维持机制及物种间的亲缘关系。甘肃省地理环境复杂,显著而多变的气候梯度形成了区域植被和环境差异,丰富了栖息地类型,具有显著的纵向连通性和纬度隔离性,以甘肃省典型纬度梯度植物群落为研究对象,通过对其进行群落学调查和功能性状测定,计算净亲缘关系指数(Net relatedness index, NRI)和平均成对性状距离(Mean pairwise trait distance, PW)来分析植物群落系统发育结构和功能性状格局对不同纬度的响应。结果表明：(1) Shannon-Weiner多样性指数,物种丰富度,谱系α多样性指数表现出随纬度增加而显著降低的变化趋势(P<0.05),Pielou均匀度指数随纬度的升高没有显著的变化趋势;(2)系统发育结构在高、低纬度上趋于发散状态(NRI<0),在中纬度上又表现出聚集(NRI>0)的谱系结构,表明种间竞争作用减弱,环境过滤作用逐渐增强,随纬度继续升高相似性限制作用在物种聚集过程中占优势;而群落的功能性状结构随着纬度增加表现出与谱系结构相反的状态,因此植物群落的系统发育和功能...     

Integrating trait and phylogenetic distances to assess scale‐dependent community assembly processes

Biodiversity is structured by multiple mechanisms that are dependent, at least in part, on ecological similarities and differences among species. Integrating traits and phylogenies in diversity metrics may provide deeper insight into community assembly processes across spatial scales. However, different traits are influenced by processes at different spatial scales, and it is not clear how trait‐spatial scale mismatches skew our ability to detect assembly patterns. An additional complexity is how phylogenetic distances, which might capture unmeasured traits, reflect spatially dependent processes. Here we analyze a freshwater zooplankton dataset from 91 ponds and show that different traits are associated with processes at different spatial scales. We first assessed the response of individual traits to processes at both α‐ and β‐scales, and then quantified the power of different combinations of traits and phylogenetic distances to reveal environmental and spatial drivers of α‐ and β‐diversity. We found that explanatory power was maximised when we accounted for environmental and spatial drivers with single, but different traits for α‐ and β‐diversity. Using the most appropriate trait for each spatial scale outperformed phylogenetic information, but phylogenetic information outperformed the same traits when these were used at the wrong spatial scale, and all outperformed taxonomic analyses that ignore trait and phylogenetic information. We demonstrate that accounting for species’ similarities and differences provides important information about dominant assembly mechanisms at different spatial scales, and that phylogeny is especially useful when measured traits are uninformative at a given spatial scale or when there is lack of trait data. Our study also indicates, however, that trait‐scale mismatches among phylogenetically conserved traits may affect the performance of phylogenetic indices compared to indices that account only for the best single trait at each spatial scale.     

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.