停止人为去除植物功能群后的高寒草甸多样性恢复过程与群落构建

已有大量研究利用功能性状或系统发育来推断群落构建机制, 然而不同过程可能会导致相似的格局。本文基于对甘南高寒草甸植物功能群去除处理后群落恢复过程的跟踪调查, 对比了物种多样性、功能多样性和系统发育多样性的动态变化, 并分析了物种定殖与消失过程对功能多样性和系统发育多样性变化的影响。结果表明: 去除不同数量功能群的群落中: (1)包括物种丰富度(SR)、Shannon-Wiener指数(H°)和Simpson指数(D)在内的传统物种多样性均随时间快速上升并与自然群落趋同, 不同群落的均匀度指数(J)随时间呈增加趋势并趋于相似; 功能多样性(FD)与系统发育多样性(PD)呈现出与物种多样性相似的动态变化趋势, 而平均配对距离(MPD/MPD_a、MFD/MFD_a)则向中等程度聚集。(2)不同群落的功能群和物种组成在短期内均恢复到与自然群落非常相似的程度。(3)物种定殖与消失过程的功能格局是群落恢复过程中趋同效应的主要驱动力。本研究揭示了高寒草甸植物功能群去除停止后群落短期内快速恢复的过程, 说明在小尺度且周边具有大范围未退化草甸的情况下, 无论物种多样性、功能多样性还是系统发育多样性都具有较快的恢复能力, 同时说明了利用群落系统发育多样性格局来推断群落构建机制的局限性。     

功能多样性和功能冗余对高寒草甸群落稳定性的影响

功能多样性和功能冗余是影响群落稳定性的重要因素,但它们对稳定性影响的相对强弱尚有争论。通过在青藏高原高寒矮嵩草(Kobresia humilis)草甸为期6a的刈割(不刈割、留茬3cm、留茬1cm)和施肥(施肥、不施肥)控制实验对此进行了探讨。研究结果显示,群落稳定性随着功能多样性和功能冗余的增大而单调增加,但由于功能多样性的变异性大于功能冗余的变异性,功能多样性与物种多样性间的相关性强于功能冗余与物种多样性间的相关性,功能多样性与群落稳定性间的相关性也强于功能冗余与群落稳定性间的相关性,因此,尽管功能冗余产生的保险效应对维持群落稳定性具有促进作用,但功能多样性的互补效应对维持群落稳定性的作用更大。结果表明,功能多样性和功能冗余对群落稳定性的相对影响与其变化程度正相关,功能多样性与群落稳定性的变化具有更紧密的联系,能更好地预测生态系统稳定性的变化。研究为草原生态系统的可持续管理提供了重要启示。     

A dimmer shade of pale: revealing the faint signature of local assembly processes on the structure of strongly filtered plant communities

Trait‐based ecology suggests that abiotic filtering is the main mechanism structuring the regional species pool in different subsets of habitat‐specific species. At more local spatial scales, other ecological processes may add on giving rise to complex patterns of functional diversity (FD). Understanding how assembly processes operating on the habitat‐specific species pools produce the locally observed plant assemblages is an ongoing challenge. Here, we evaluated the importance of different processes to community assembly in an alpine fellfield, assessing its effects on local plant trait FD. Using classical randomization tests and linear mixed models, we compared the observed FD with expectations from three null models that hierarchically incorporate additional assembly constraints: stochastic null models (random assembly), independence null models (each species responding individual and independently to abiotic environment), and co‐occurrence null models (species responding to environmental variation and to the presence of other species). We sampled species composition in 115 quadrats across 24 locations in the central Pyrenees (Spain) that differed in soil conditions, solar radiation and elevation. Overall, the classical randomization tests were unable to find differences between the observed and expected functional patterns, suggesting that the strong abiotic filters that sort out the flora of extreme regional environments blur any signal of other local processes. However, our approach based on linear mixed models revealed the signature of different ecological processes. In the case of seed mass and leaf thickness, observed FD significantly deviated from the expectations of the stochastic model, suggesting that fine‐scale abiotic filtering and facilitation can be behind these patterns. Our study highlights how the hierarchical incorporation of ecological additional constraints may shed light on the dim signal left by local assembly processes in alpine environments.     

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。     

植物功能群去除对高寒草甸群落结构、多样性及生产力的影响

群落中物种的丧失在干扰下普遍存在,但对生态系统过程和功能的影响仍存在较大不确定性。选取青藏高原东缘典型高寒草甸为对象,开展优势植物功能群的梯度去除试验,以模拟长期过牧干扰下物种的损失。经过连续两个生长季的功能群去除,我们对群落的物种组成、结构、多样性和生物量等特征进行了分析,探讨了上述指标的响应过程和机制。研究结果表明：（1）功能群的去除降低了群落高度,增加了物种均匀度,并显著影响了禾草、杂草优势比以及功能群多样性和优势度;（2）同时,去除操作显著减小了凋落物量与禾草生物量,并显著影响了群落地上生物量;（3）进一步分析还发现,禾草、莎草和杂草功能群之间存在显著的竞争关系,群落生产力主要取决于禾草功能群并随物种均匀度的增大而显著减小。上述结果表明,禾草在高寒草甸群落中占据竞争优势地位,植物功能群的损失主要通过改变种间竞争关系、引起有机物质丢失影响群落过程和功能。     

微生物物种多样性、群落构建与功能性状研究进展

微生物主要包括细菌、真菌、古菌、病毒等类群, 是地球上出现时间最早、分布最广泛、个体数量最多, 以及物种和基因多样性十分丰富的生物类群。为了适应各种生境, 微生物衍生出腐生、寄生、共生等多样的生存策略, 在生物地球化学循环、生态系统演替与稳定性、环境修复以及人类健康等方面发挥着重要作用。传统的微生物监测方法限制了我们对微生物多样性的认知; 但是, 近年来高通量测序技术和生物信息学的发展极大推动了微生物多样性的研究进展。本文概述了近年来在微生物多样性分布格局与维持、群落构建以及功能属性多样性的最新进展; 总结分析了细菌、古菌、真菌的多样性纬度分布格局及其驱动因子, 选择、扩散、成种、漂变等过程对细菌、古菌、真菌的群落构建的贡献, 以及细菌和真菌的形态、生理生化、生长繁殖、扩散、基因组等功能性状的多样性; 提出了未来微生物多样性研究的重要领域: 环境宏真菌组研究, 微生物多样性与生态系统多功能性的关系研究, 以及微生物互作网络的生态功能研究。     

高寒草甸不同草地类型功能群多样性及组成对植物群落生产力的影响

对不同类型草地功能群多样性和组成与植物群落生产力之间的关系进行了探讨。结果表明：(1)在矮嵩草(Kobresia humlis)草甸和金露梅(Potentilla froticosa)灌丛中,豆科植物的作用比较明显,而其他功能群植物的作用较弱。(2)在藏嵩草(Kobresia tibetica)沼泽化草甸和小嵩草(K．pygmaca)草甸中,虽然杂类草、C3植物和莎草科植物功能群的生产力占群落初级生产力的比例较大,但二者在统计上没有显著性差异,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响,更主要的是受到功能群内物种密度和均匀度的影响,即功能群组成比功能群多样性更能说明对生态系统过程的影响。(3)不同类型草地群落植物功能群盖度与群落初级生产力呈显著的线性相关。(4)不同类型草地群落生产力与功能群内物种数的变化均表现为单峰曲线关系,即功能群内物种数处于中间水平时,群落生产力最高。     

Changes in plant taxonomic and functional diversity patterns following treeline advances in the South Urals

Background: Treeline ecotones represent environmental boundaries that fluctuate in space and time and thus induce changes in plant taxonomic and functional diversity.

Aims: To study changes through time in taxonomic and functional plant diversity patterns along the treeline ecotone.

Methods: In 2002, vegetation was sampled along a gradient from upper montane forest to the treeline–alpine transition in the South Ural Mountains, Russia. In 2014, vegetation was resampled and plant functional traits were collected. We studied spatial and temporal changes in plant species composition, functional composition and functional diversity.

Results: Species composition and diversity changed along the elevational gradient. The functional composition in height, leaf area, specific leaf area and leaf nitrogen content decreased with elevation, whereas functional composition of leaf carbon content increased. We found a temporal shift towards shorter plants with smaller leaves in treeline sites. Functional richness varied in several traits along the elevational gradient, while functional dispersion showed a trend towards increased functional dispersion in height, specific leaf area and leaf nitrogen in the treeline–tundra transition.

Conclusions: Tree encroachment across the treeline ecotone has resulted in a shift in plant species relative abundances and functional diversity, possibly affecting plant community assembly patterns.     

高寒草甸种间性状差异和物种均匀度对物种多样性与功能多样性关系的影响

物种多样性(SD)与功能多样性(FD)之间存在多种关系,但由于生态系统功能主要由物种的功能属性决定,因而功能多样性对生态系统功能的影响大于物种多样性的影响。但在种间性状差异和物种均匀度这两个构成功能多样性的基本成分中,何者对功能多样性影响更大,并进而决定SD-FD关系尚不明确。通过在高寒矮嵩草(Kobresia humilis)草甸为期6a的刈割(留茬1 cm、3 cm及不刈割)和施肥(尿素7.5 g m~(-2)a~(-1)+磷酸二胺1.8 g m~(-2)a~(-1)、不施肥)控制实验,研究了种间性状差异(33个物种13个性状)和物种均匀度(所有物种)对物种多样性(所有物种)与功能多样性(33个物种13个性状)之间关系的影响。研究结果显示:(1)物种多样性与功能多样性正相关,它们与多性状种间差异负相关,而与物种均匀度正相关。物种均匀度是导致功能多样性变化的主要因素,也是导致SD-FD正相关的原因,这是因为随着物种多样性增加,物种均匀度的增加程度大于多性状种间差异的减少程度,因而功能多样性增加,SD-FD正相关;(2)功能多样性指数(FD_(Rao)和FDis)随物种多样性指数(H')减速递增,表明群落存在一定的功能冗余,且功能冗余随物种多样性的增大而增大,但尚未达到产生SD-FD无相关性的极限H'值;(3)功能多样性对高寒草甸生态系统地上净初级生产力(ANPP)的影响大于物种多样性的影响,二元线性回归显示在同时考虑二者对ANPP的影响时,可排除物种多样性的作用。但由于物种多样性下降或物种丧失引起的物种功能性状丢失或性状空间维度减小将导致功能多样性降低,表明它们之间存在一定互补性,在研究生物多样性与生态系统功能关系时,同时考虑物种多样性和功能多样性的影响仍十分必要。     

A predictive model of community assembly that incorporates intraspecific trait variation

Community assembly involves two antagonistic processes that select functional traits in opposite directions. Environmental filtering tends to increase the functional similarity of species within communities leading to trait convergence, whereas competition tends to limit the functional similarity of species within communities leading to trait divergence. Here, we introduce a new hierarchical Bayesian model that incorporates intraspecific trait variation into a predictive framework to unify classic coexistence theory and evolutionary biology with recent trait‐based approaches. Model predictions exhibited a significant positive correlation (r = 0.66) with observed relative abundances along a 10 °C gradient in mean annual temperature. The model predicted the correct dominant species in half of the plots, and accurately reproduced species' temperature optimums. The framework is generalizable to any ecosystem as it can accommodate any species pool, any set of functional traits and multiple environmental gradients, and it eliminates some of the criticisms associated with recent trait‐based community assembly models.     

Competitive interactions between forest trees are driven by species' trait hierarchy, not phylogenetic or functional similarity: implications for forest community assembly

The relative importance of competition vs. environmental filtering in the assembly of communities is commonly inferred from their functional and phylogenetic structure, on the grounds that similar species compete most strongly for resources and are therefore less likely to coexist locally. This approach ignores the possibility that competitive effects can be determined by relative positions of species on a hierarchy of competitive ability. Using growth data, we estimated 275 interaction coefficients between tree species in the French mountains. We show that interaction strengths are mainly driven by trait hierarchy and not by functional or phylogenetic similarity. On the basis of this result, we thus propose that functional and phylogenetic convergence in local tree community might be due to competition-sorting species with different competitive abilities and not only environmental filtering as commonly assumed. We then show a functional and phylogenetic convergence of forest structure with increasing plot age, which supports this view.     

Phylogenetic diversity,trait diversity and niches: species assembly of ferns along a tropical elevational gradient

Aim To analyse the structure of pteridophyte assemblages, based on phylogenetic relatedness and trait properties, along an elevational gradient. Ecological theory predicts that co‐occurring species may be: randomly selected from a regional pool; ecologically sorted so that they are functionally different hence resulting in reduced competition (overdispersion); or functionally similar as an adaptation to specific ecological conditions (clustering). Location Braulio Carrillo National Park and Cerro de la Muerte, Costa Rica, Central America. Methods We used an empirical dataset of the quantitative pattern of species occurrences and individual numbers of ferns within 156 plots along a tropical elevational gradient to test whether directed ecological sorting might cause deviations in patterns of trait and phylogenetic diversity. Mean pairwise distances of species based on phylogenetic and trait properties were compared with two different sets of null assemblages, one maintaining species frequency distributions (constrained) and one not (unconstrained). Results Applying different null models resulted in varying degrees of overdispersion and clustering, but overall patterns of deviation from random expectations remained the same. Contrary to theoretical predictions, phylogenetic and trait diversity were relatively independent from one another. Phylogenetic diversity showed no patterns along the elevational gradient, whereas trait diversity showed significant trends for epiphytes. Main conclusions Under stressful environmental conditions (drought at low elevations and frost at high elevations), epiphytic fern assemblages tended to be clustered with respect to trait characteristics, which suggests environmental filtering. Conversely, under less extreme environmental conditions (middle of the transect), the sorting was biased towards high differentiation (overdispersion), presumably because of interspecific competition and trait shifts among closely related species (character displacement).     

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.     

Linking species performance to community structure as affected by UV-B radiation: an attenuation experiment

Aims UV-B radiation is known to affect plant physiology and growth rate in ways that can influence community species composition and structure. Nevertheless, comparatively little is known about how UV-B radiation induced changes in the performance of individual species cascades to affect overall community properties. Because foliage leaves are primarily responsible for photosynthesis and carbon gain and are the major organ that senses and responds to UV-B radiation, we hypothesized that, under reduced UV-B radiation, species with larger leaf areas per plant would manifest higher growth rates and hence tend to improve their community status compared to species with smaller leaf areas per plant in herbaceous plant communities.Methods We tested this hypothesis by examining plant traits (leaf area per plant and plant height), plant growth rate (aboveground biomass per plant and plant biomass per area) and community status (species within-community relative biomass) for 19 common species in a two-year field experiment in an alpine meadow on Tibetan Plateau.Important findings Aboveground biomass per plant, as well as per area, progressively increased in a 39% reduced (relative to ambient) UV-B treatment during the experimental period. At the second year, 11 out of 19 species significantly or marginally significantly increased their plant height, leaf area per plant and aboveground biomass per plant. No species was negatively affected by reducing UV-B. As hypothesized, the increase in aboveground biomass per plant increased with increasing leaf area per plant, as indicated by cross-species regression analysis. Moreover, the change in species within-community status increased with increasing leaf area per plant. Our study demonstrates that UV-B radiation has differential effects on plant growth rate across species and hence significantly affects species composition and plant community structure. We suggest that UV-B radiation is an ecological factor structuring plant communities particularly in alpine and polar areas.     

An index for measuring functional extension and evenness in trait space

Most existing functional diversity indices focus on a single facet of functional diversity. Although these indices are useful for quantifying specific aspects of functional diversity, they often present some conceptual or practical limitations in estimating functional diversity. Here, we present a new functional extension and evenness (FEE) index that encompasses two important aspects of functional diversity. This new index is based on the straightforward notion that a community has high diversity when its species are distant from each other in trait space. The index quantifies functional diversity by evaluating the overall extension of species traits and the interspecific differences of a species assemblage in trait space. The concept of minimum spanning tree (MST) of points was adopted to obtain the essential distribution properties for a species assembly in trait space. We combined the total length of MST branches (extension) and the variation of branch lengths (evenness) into a raw FEE₀ metric and then translated FEE₀ to a species richness‐independent FEE index using a null model approach. We assessed the properties of FEE and used multiple approaches to evaluate its performance. The results show that the FEE index performs well in quantifying functional diversity and presents the following desired properties: (a) It allows a fair comparison of functional diversity across different species richness levels; (b) it preserves the essence of single‐facet indices while overcoming some of their limitations; (c) it standardizes comparisons among communities by taking into consideration the trait space of the shared species pool; and (d) it has the potential to distinguish among different community assembly processes. With these attributes, we suggest that the FEE index is a promising metric to inform biodiversity conservation policy and management, especially in applications at large spatial and/or temporal scales.