Intraspecific trait variation increases species diversity in a trait‐based grassland model

Intraspecific trait variation (ITV) is thought to play a significant role in community assembly, but the magnitude and direction of its influence are not well understood. Although it may be critical to better explain population persistence, species interactions, and therefore biodiversity patterns, manipulating ITV in experiments is challenging. We therefore incorporated ITV into a trait‐ and individual‐based model of grassland community assembly by adding variation to the plants’ functional traits, which then drive life‐history tradeoffs. Varying the amount of ITV in the simulation, we examine its influence on pairwise‐coexistence and then on the species diversity in communities of different initial sizes. We find that ITV increases the ability of the weakest species to invade most, but that this effect does not scale to the community level, where the primary effect of ITV is to increase the persistence and abundance of the competitively‐average species. Diversity of the initial community is also of critical importance in determining ITV's efficacy; above a threshold of interspecific diversity, ITV does not increase diversity further. For communities below this threshold, ITV mainly helps to increase diversity in those communities that would otherwise be low‐diversity. These findings suggest that ITV actively maintains diversity by helping the species on the margins of persistence, but mostly in habitats of relatively low alpha and beta diversity.     

长白山温带森林不同演替阶段群落功能性状的空间变化

群落构建机制研究是生态学研究的热点。长白山自然保护区拥有完整的原始阔叶红松林生态系统, 近年来随着物种多样性丧失愈发严重, 对该地区开展群落构建机制研究显得尤为重要。该研究以长白山不同演替阶段的3块5.2 hm ²固定监测样地(次生杨桦林、次生针阔混交林、原始椴树红松林)为研究对象, 通过采集样地内主要树种的6个关键功能性状(叶面积、比叶面积、叶片厚度、叶氮含量、叶磷含量、最大树高), 分析不同空间尺度下(5 m × 5 m, 10 m × 10 m, 20 m × 20 m, 30 m × 30 m, 40 m × 40 m, 50 m × 50 m和60 m × 60 m)及不同演替阶段群落性状空间值的变化, 结合零模型的模拟结果对长白山温带森林演替过程中的群落构建机制进行讨论。结果表明: 种库大小对于研究结果具有重要影响。在较大的种库下, 环境过滤作用影响显著。而在样地水平进行研究时, 演替早期和中期, 群落性状空间值与零模型模拟值无显著差异, 在演替的晚期, 群落性状空间值显著高于零模型模拟值。结合多个群落功能多样性指数分析发现, 环境过滤和竞争作用共同决定该地区顶级群落的物种组成。在演替早期大量物种迁入, 群落内物种间存在强烈的资源竞争, 而随着演替进行, 部分物种逐渐被竞争排除出群落, 群落中的物种呈现明显的生态位分化, 竞争作用是维持物种共存的主要机制。     

Intraspecific trait variability matters

Intraspecific trait variability (ITV) plays a central part in various ecological processes, though using mean trait values may be sufficient in some instances. Ecologists need thus to find under which circumstances. Carlucci et al. (2014, this issue) bring new evidence on the importance of ITV for community assembly across a strong gradient. Sampling design may affect ITV quantification across gradients.     

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta‐analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole‐plant (e.g. plant height) vs. organ‐level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait‐based community and ecosystem studies.     

Seasonality,species richness and poor dispersion mediate intraspecific trait variability in stonefly community responses along an elevational gradient

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Intraspecific trait variation in three common grass species reveals finescale species adjustment to local environmental conditions

Aims Studies along environmental gradients have shown that intraspecific trait variation (ITV) may contribute considerably to community-level trait variation. However, we lack knowledge about how the extent of ITV varies on a local scale and whether a varying extent of ITV is related to differences in local environmental site and plant community characteristics.     

大兴安岭不同演替阶段天然林物种多度格局模拟

为实现大兴安岭地区落叶松天然林的全周期可持续经营,以新林林业局翠岗林场3块100m×100m不同演替阶段(白桦林、落叶松-白桦混交林和落叶松林)的天然林固定调查样地数据为基础,采用统计模型(对数正态模型)、生态位模型(断棍模型、生态位优先占领模型、Zipf模型、Zipf-Mandelbrot模型)和中性模型(群落零和多项式模型、Volkov模型)对不同演替阶段天然林的乔木层、幼苗层和由乔木层划分的不同林层的物种多度格局进行拟合,并用χ² 检验和Kolmogorov-Smirnov检验选择最佳模型。结果表明:(1) 在大兴安岭地区不同演替阶段的落叶松天然林群落中,Zipf-Mandelbrot模型的拟合效果均最好。(2) 对数正态模型对于稳定的演替阶段白桦林和落叶松林的群落相较于过渡阶段群落落叶松-白桦混交林的物种多度分布的拟合效果更好。(3) 在大兴安岭地区落叶松天然林中,生境过滤主导群落构建,随着群落不断演替生境过滤和中性过程减弱。(4) 天然林群落乔木层的群落构建过程与乔木层内不同生长阶段林层的构建过程不同,生境过滤过程在落叶松林乔木层中逐渐减弱,而在落叶松林不同生长阶段的林层和更新层中增强。     

刈割与施肥干扰下高寒草甸植物功能性状的构建模式

针对目前关于植物群落功能性状构建模式及其驱动因素存在的争议, 研究了33个物种10个功能性状的构建模式。研究结果显示: (1)在刈割-施肥复合梯度上, 这些功能性状主要表现为随机构建模式, 发生随机性和确定性构建的样方比例分别为82.7%和17.3%; (2)在10个功能性状中, 生长型、生活周期、单株地上干质量、叶面积和叶干质量5个功能性状为随机构建模式, 不受试验处理和群落特征(地上净初级生产力、刈割生物量损失、群落高度)变化的影响。植物倾斜度、繁殖方式、固氮性、株高和比叶面积5个功能性状的构建模式与试验处理或群落特征变化有关, 表现出趋同或趋异构建响应。其中, 植物倾斜度和比叶面积的构建模式仅受群落特征影响, 而固氮性、繁殖方式和株高3个性状的构建模式同时还受刈割或施肥处理的影响, 其构建模式因功能性状而异; (3)群落的生物量损失和地上净初级生产力是解释植物功能性状构建模式变化较理想的群落特征; (4)刈割和施肥处理对株高的构建模式具有相反效应, 而刈割和施肥的交互作用对其无显著影响。上述结果说明该研究群落植物功能性状的构建存在不同模式, 以随机构建模式为主, 确定性构建模式居次要地位。确定性构建模式与试验处理和群落特征变化有关, 而且是性状依赖的。相反选择力对趋同和趋异构建模式的平衡效应能引起功能性状发生随机构建。     

Individual‐level leaf trait variation and correlation across biological and spatial scales

Even with increasing interest in the ecological importance of intraspecific trait variation (ITV) for better understanding ecological processes, few studies have quantified ITV in seedlings and assessed constraints imposed by trade‐offs and correlations among individual‐level leaf traits. Estimating the amount and role of ITV in seedlings is important to understand tree recruitment and long‐term forest dynamics. We measured ten different size, economics, and whole leaf traits (lamina and petiole) for more than 2,800 seedlings (height ≥ 10 cm and diameter at breast height < 1 cm) in 283 seedling plots and then quantified the amount of ITV and trait correlations across two biological (intraspecific and interspecific) and spatial (within and among plots) scales. Finally, we explored the effects of trait variance and sample size on the strength of trait correlations. We found about 40% (6%–63%) variation in leaf‐level traits was explained by ITV across all traits. Lamina and petiole traits were correlated across biological and spatial scales, whereas leaf size traits (e.g., lamina area) were weakly correlated with economics traits (e.g., specific lamina area); lamina mass ratio was strongly related to the petiole length. Trait correlations varied among species, plots, and different scales but there was no evidence that the strength of trait relationships was stronger at broader than finer biological and spatial scales. While larger trait variance increased the strength of correlations, the sample size was the most important factor that was negatively related to the strength of trait correlations. Our results showed that a large amount of trait variation was explained by ITV, which highlighted the importance of considering ITV when using trait‐based approaches in seedling ecology. In addition, sample size was an important factor that influenced the strength of trait correlations, which suggests that comparing trait correlations across studies should consider the differences in sample size.     

Explaining variation in productivity requires intraspecific variability in plant height among communities

解释生产力的变异需要考虑群落间植物高度的种内变异 已有研究表明,种内性状变异在群落构建过程中具有重要作用,但是迄今为止关于种内性状变异对生态系统功能的贡献仍然知之甚少。我们在青藏高原的高寒草甸进行了为期4年的功能群去 除实验,以研究植物高度的种间和种内变异对生产力的相对重要性。将75个控制群落内的株高总 变异分为种间变异(TV_inter)和种内变异(ITV_within),将群落间的群落加权平均高度分解为固定群落加权平 均值(CWM_fixed)和群落间种内变异(ITV_among)。我们通过广义加性混合模型、模型筛选和结构方程模型评估了性状变异(即TV_inter、ITV_within、CWM_fixed和ITV_among)如何间接介导功能群去除后生产力的变化。研究结果表明,功能群去除不仅会直接引起生产力变化,同时还会通过改变种间和种内株高变异间接影响生产力(即CWM_fixed和ITV_among)。“选择效应”和“避荫综合征”都可能导致更高的CWM_fixed和ITV_among,从而起到调节生产力的作用。该研究结果说明,只考虑种间变异可能会低估植物群落功能结构在驱动生态系统过程中的贡献。     

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...     

Quantifying relationships between traits and explicitly measured gradients of stress and disturbance in early successional plant communities

Questions: How can one explicitly quantify, and separately measure, stress and disturbance gradients? How do these gradients affect functional composition in early successional plant communities and to what extent? Can we accurately predict trait composition from knowledge of these gradients? Location: Southern Quebec, Canada. Methods: Using eight environmental variables measured in 48 early successional plant communities, we estimated stress and disturbance gradients through structural equation modelling. We then measured 10 functional traits on the most abundant species of these 48 communities and calculated their community‐level mean and variance weighted by the relative abundance of each species. Finally, we related these community‐weighted means and variances to the estimated stress and disturbance gradients using general linear models or generalized additive models. Results: We obtained a well‐fitting measurement model of the stress and disturbance gradients existing in our sites. Of the 10 studied traits, only average plant reproductive height was strongly correlated with the stress (r²=0.464) and disturbance (r²=0.543) gradients. Leaf traits were not significantly related to either the stress or disturbance gradients. Conclusions: The well‐fitting measurement model of the stress and disturbance gradients, combined with the generally weak trait–environment linkages, suggests that community assembly in these early successional plant communities is driven primarily by stochastic processes linked to the history of arrival of propagules and not to trait‐based environmental filtering.     

片段化森林群落构建的生态过程及其检验方法

鉴于全球森林均呈现片段化(破碎化)的分布状态, 理解片段化森林群落构建的过程很有必要。该文通过综述群落构建的主要生态过程如生态漂变、扩散、选择和物种形成等在片段化森林群落构建中的相对作用, 发现因片段化森林形成方式的不同, 重构群落(片段化生境中通过次生演替重新形成的森林群落)和解构群落(原有森林被片段化后形成的森林群落)在不同演替阶段所受到的主要生态过程的相对作用有所不同。虽然利用基于群落内物种分布格局推测构建过程(如物种多度分布、零模型结合β多样性的方法、功能特征的收敛和发散等)、人工控制实验、群落结构动态分析等方法对片段化森林中群落构建的过程进行了有效的检验, 但是针对片段化森林群落构建过程的实验性研究仍然不足。未来有待在理论模型、群落构建过程的检验以及理论与物种保护相结合等方面继续开展深入的研究。     

Community assembly during vegetation succession after metal mining is driven by multiple processes with temporal variation

The mechanisms governing community assembly is fundamental to ecological restoration and clarification of the assembly processes associated with severe disturbances (characterized by no biological legacy and serious environmental problems) is essential. However, a systematic understanding of community assembly in the context of severe anthropogenic disturbance remains lacking. Here, we explored community assembly processes after metal mining, which is considered to be a highly destructive activity to provide insight into the assembly rules associated with severe anthropogenic disturbance. Using a chronosequence approach, we selected vegetation patches representing different successional stages and collected data on eight plant functional traits from each stage. The traits were classified as establishment and regenerative traits. Based on these traits, null models were constructed to identify the processes driving assembly at various successional stages. Comparison of our observations with the null models indicated that establishment and regenerative traits converged in the primary stage of succession. As succession progressed, establishment traits shifted to neutral assembly, whereas regeneration traits alternately converged and diverged. The observed establishment traits were equal to expected values, whereas regenerative traits diverged significantly after more than 20 years of succession. Furthermore, the available Cr content was linked strongly to species'' ecological strategies. In the initial stages of vegetation succession in an abandoned metal mine, the plant community was mainly affected by the available metal content and dispersal limitation. It was probably further affected by strong interspecific interaction after the environmental conditions had improved, and stochastic processes became dominant during the stage with a successional age of more than 20 years.     

On the importance of intraspecific variability for the quantification of functional diversity

Functional diversity (FD) is a key facet of biodiversity used to address central questions in ecology. Despite recent methodological advances, FD remains a complex concept and no consensus has been reached either on how to quantify it, or on how it influences ecological processes. Here we define FD as the distribution of trait values within a community. When and how to account for intraspecific trait variability (ITV) when measuring FD remains one of the main current debates. It remains however unclear to what extent accounting for population‐level ITV would modify FD quantification and associated conclusions. In this paper, we address two critical questions: (1) How sensitive are different components of FD to the inclusion of population‐level ITV? (2) Does the omission of ITV obscure the understanding of ecological patterns? Using a mixture of empirical data and simulation experiments, we conducted a sensitivity analysis of four commonly used FD indices (community weighted mean traits, functional richness, Rao's quadratic entropy, Petchey and Gaston's FD index) and their relationships with environmental gradients and species richness, by varying both the extent (plasticity or not) and the structure (contingency to environmental gradient due to local adaptation) of population‐level ITV. Our results suggest that ITV may strongly alter the quantification of FD and the detection of ecological patterns. Our analysis highlights that 1) species trait values distributions within communities are crucial to the sensitivity to ITV, 2) ITV structure plays a major role in this sensitivity and 3) different indices are not evenly sensitive to ITV, the single‐trait FD from Petchey and Gaston being the most sensitive among the four metrics tested. We conclude that the effects of intraspecific variability in trait values should be more systematically tested before drawing central conclusions on FD, and suggest the use of simulation studies for such sensitivity analyses.     

植物群落构建的生态过滤机制研究进展

生物多样性的形成和维持机制,即群落构建机制,一直以来都是群落生态学研究的核心问题。植物群落构建的确定性过程主要是生态过滤机制(包括环境过滤和生物过滤,其中生物过滤包括种间竞争和种内功能性状变异)作用的结果。学者们构建了大量的理论、方法、模型来解释和验证生态过滤机制对群落构建的影响,并取得了显著的成果。然而,关于生态过滤机制在不同尺度的作用、生态过滤机制的各要素分解和量化等方面的研究仍有诸多疑问。重点综述了环境过滤、种间竞争和种内功能性状变异的最新研究进展,并指出了现有研究的不足之处。在未来的研究中,应注重生态过滤机制的各要素分解和量化,加强研究手段的综合运用,关注时空动态变化对植物群落构建的影响,重视对不同植物群落构建机制的共性和个性特征的认识,同时强调与其他生态过程、群落构建机制的整合。通过这些尝试,有助于人们更好的理解植物群落构建过程中的生态过滤机制的作用。     

Species coexistence in temperate, mixed deciduous forests

The response of tree life-history traits to community profiles (horizontal and vertical heterogeneity, disturbances and biotic interactions) determines community assembly rules, which are currently a hot issue in community ecology. Important mechanisms of coexistence differ throughout the developing stages of tree life history. Many processes of niche partitioning and tradeoffs that potentially enable tree coexistence have been reported to be present in temperate forests, although some of these life-history traits are either correlated with each other or are not independent. Not all of the proposed mechanisms explain coexistence equally well; some could predominate in determining the community organization of forest communities. Population studies need to concentrate more on the component species of a target community to detect the ecological assembly rule. These approaches can also address how chance factors contribute to the composition of temperate tree communities, which might be less dependent on chance than are tropical ones.     

Grazing effects on intraspecific trait variability vary with changing precipitation patterns in Mongolian rangelands

Functional traits are proxies for plant physiology and performance, which do not only differ between species but also within species. In this work, we hypothesized that (a) with increasing precipitation, the percentage of focal species which significantly respond to changes in grazing intensity increases, while under dry conditions, climate‐induced stress is so high that plant species hardly respond to any changes in grazing intensity and that (b) the magnitude with which species change their trait values in response to grazing, reflected by coefficients of variation (CVs), increases with increasing precipitation. Chosen plant traits were canopy height, plant width, specific leaf area (SLA), chlorophyll fluorescence, performance index, stomatal pore area index (SPI), and individual aboveground biomass of 15 species along a precipitation gradient with different grazing intensities in Mongolian rangelands. We used linear models for each trait to assess whether the percentage of species that respond to grazing changes along the precipitation gradient. To test the second hypothesis, we assessed the magnitude of intraspecific trait variability (ITV) response to grazing, per species, trait, and precipitation level by calculating CVs across the different grazing intensities. ITV was most prominent for SLA and SPI under highest precipitation, confirming our first hypothesis. Accordingly, CVs of canopy height, SPI, and SLA increased with increasing precipitation, partly confirming our second hypothesis. CVs of the species over all traits increased with increasing precipitation only for three species. This study shows that it remains challenging to predict how plant performance will shift under changing environmental conditions based on their traits alone. In this context, the implications for the use of community‐weighted mean trait values are discussed, as not only species abundances change in response to changing environmental conditions, but also values of traits considerably change. Including this aspect in further studies will improve our understanding of processes acting within and among communities.     

Community assembly in Lake Tanganyika cichlid fish: quantifying the contributions of both niche‐based and neutral processes

The cichlid family features some of the most spectacular examples of adaptive radiation. Evolutionary studies have highlighted the importance of both trophic adaptation and sexual selection in cichlid speciation. However, it is poorly understood what processes drive the composition and diversity of local cichlid species assemblages on relatively short, ecological timescales. Here, we investigate the relative importance of niche‐based and neutral processes in determining the composition and diversity of cichlid communities inhabiting various environmental conditions in the littoral zone of Lake Tanganyika, Zambia. We collected data on cichlid abundance, morphometrics, and local environments. We analyzed relationships between mean trait values, community composition, and environmental variation, and used a recently developed modeling technique (STEPCAM) to estimate the contributions of niche‐based and neutral processes to community assembly. Contrary to our expectations, our results show that stochastic processes, and not niche‐based processes, were responsible for the majority of cichlid community assembly. We also found that the relative importance of niche‐based and neutral processes was constant across environments. However, we found significant relationships between environmental variation, community trait means, and community composition. These relationships were caused by niche‐based processes, as they disappeared in simulated, purely neutrally assembled communities. Importantly, these results can potentially reconcile seemingly contrasting findings in the literature about the importance of either niche‐based or neutral‐based processes in community assembly, as we show that significant trait relationships can already be found in nearly (but not completely) neutrally assembled communities; that is, even a small deviation from neutrality can have major effects on community patterns.