The Association of Dispersal and Persistence Traits of Plants with Different Stages of Succession in Central European Man-Made Habitats

Traits related to seed dispersal, clonality and bud bank affect the success or failure of plant species. Using data from 13 successional seres in various human-made habitats the spectra of traits associated with dispersal and persistence were compared to determine the traits that can be used to predict the occurrence of particular plant species at each stage in a succession and how the importance of these traits changes over time. Differences in the traits of species associated with primary and secondary successions were also studied. Species with seeds that are dispersed by water (hydrochory) decreased in abundance during the course of succession. Species with a splitting main root, monocyclic and dicyclic shoots also decreased in abundance. Species capable of forming a potential below-ground bud bank, hypogeogenous rhizome and retaining a long-term connection with clonal offspring increased in abundance. The results indicate that seed dispersal is more important in determining the species composition in the early stages of succession whereas bud banks and clonal traits are more important in the later stages and for colonizing a locality. Primary and secondary seres did not remarkably differ in the trait spectra of the species present indicating that these trends occur in both types of succession.     

How do species dominating in succession differ from others?

Abstract. Data on succession were collected from 15 seres starting on bare ground in man-made habitats (i.e. sites disturbed by various mining activities, bulldozed sites, ruderal urban sites, the exposed bottom of a destroyed water reservoir, and abandoned fields), all in the western part of the Czech Republic, Central Europe. The period for which the data on succession were available ranged from 12 to 60 yr. 56 species reaching dominance in some period of succession were selected (the criterion being: at least 25 % cover in at least one year in any sere); they were compared for biological and ecological traits with other species participating in the seres (167 species with at least 1 % cover in at least one year in any sere), and with the Central European flora as a whole. Significant differences between the species dominating in succession and others were found for the following traits: life form, life strategy, pollination mode, and ability of lateral spread. Dominant species differed from the regional flora in distribution of life strategies, pollination mode and immigration status. The results suggest that an ‘ideal successional dominant’ is a tall, wind-pollinated plant, often a geophyte capable of intensive lateral spread, requiring high nutrient supply and sufficient site moisture. The set of traits contributing to achieving dominance by a species in human-made habitats includes both features occurring independently of phylogenies (life strategy, pollination mode, plant height, moisture demands) and those which dominant species probably share due to their common ancestors (nutrient demands, capability of extensive lateral spread).     

Spontaneous Establishment of Woody Plants in Central European Derelict Sites and their Potential for Reclamation

The performance of woody plants was analyzed in 15 successional seres starting at bare ground in central European manmade habitats. The total cover of woody species after 10 years of succession was significantly related neither to initial soil moisture nor to nitrogen (expressed using Ellenberg indicator values). But the comparison of seres indicates that establishment of woody plants was easier under moderate environmental conditions and retarded in extreme habitats (dry, nutrient-poor, or acid). The arrival of the first woody plants was delayed in dry sites. No significant differences were found between primary and secondary seres, either with respect to the total cover of woody plants reached after 10 years of succession or considering the time of their arrival. In total, 24 woody species (10 shrubs and 14 trees) appeared in the series investigated. Their successional performance (in terms of the number of seres in which the species occurred and maximum cover reached in any sere) was not related to species traits (life strategy, type of mycorrhizae, mode of dispersal, diaspore weight), except for the regeneration strategy, species with seasonal regeneration by seeds were capable of creating higher cover. Betula pendula (European birch) was the most successful species in spontaneous succession, especially on moist sites. Practical suggestions for the management of particular habitats (sites disturbed by mining, sites reclaimed after acid rain deforestation, urban sites, abandoned fields) are provided regarding the establishment of woody plants.     

黄土高原自然植被演替过程中的植物特征与土壤元素动态

在3～149年的时间尺度上,对黄土高原植被次生演替过程中植物特征和土壤养分元素动态进行了研究.结果表明:1) 随着演替时间尺度的延伸,土壤全C、全N含量呈增加趋势,而土壤全K、全Na和土壤pH值呈下降趋势,土壤全P变化趋势不明显;此外,表层(0～10 cm)土壤CaO 含量在演替系列上呈下降趋势,深层(20～30 cm,40～50 cm)则呈增加趋势.演替过程对几种土壤化学元素含量影响的程度随着土壤深度的增加而减弱.2)植物群落物种丰富度在演替的中间阶段最高.3) 在演替的早期阶段,植物群落优势种往往具有稳定的土壤种子库、CR-生活对策和S-繁殖对策,在贫瘠的土壤上具有较强的竞争能力,且具有较强的水平扩展能力和克隆繁殖能力,C-生活对策、在土壤全N含量较高的生境中具有较强竞争能力的多年生植物,在演替中后期占据群落的优势地位.此外,在所涉及的植物特征中,多年生生活史,C-、CR-、SC-、SR-、S-生活对策,以及R-、W-、Bs-、VBs- 和V-繁殖对策等特征在非优势物种中出现的频率较高.4) C-、 SC-生活对策, 克隆能力, 多年生生活史, 水平扩展能力, 种子的动物传播方式,秋季开花,荚果、坚果等特征出现的比例在一定程度上与土壤全C、全N 和全K含量正相关; 而 S-、SR-、R-、CR-生活对策,一、二年生生活史, 种子繁殖, S-繁殖对策,以及胞果、蒴果等特征的比例与土壤全Na、CaO含量和土壤pH正相关.5)在演替过程中出现的植物均属草本植物生活型,因此,草原可能是黄土高原上受制于大尺度环境条件(显域生境)下的优势植被类型(特别是降雨量不超过550 mm的地区).     

Dispersal traits as indicators of vegetation dynamics in long-term old-field succession

Succession is a key ecological process that supports our understanding of community assembly and biotic interactions. Dispersal potential and dispersal strategies, such as wind- or animal-dispersal, have been assumed to be highly relevant for the success of plant species during succession. However, research yielded varying results on changes in dispersal modes between successional stages. Here, we test the hypotheses that (a) vascular plant species that use a number of dispersal modes dominate in early stages of succession while species specialized on one/few dispersal modes increase in abundance towards later stages of succession; (b) species well adapted to wind-dispersal (anemochory) will peak in abundance in early successional stages and (c) species well adapted to adhesive dispersal (epizoochory) will increase with proceeding succession. We test these hypotheses in four sites within agriculturally dominated landscapes in Germany. Agricultural use in these sites was abandoned 20–28 years ago, leaving them to secondary succession. Sites have been monitored for plant biodiversity ever since. We analyze changes in plant species richness and abundance, number of dispersal modes and two ranking indices for wind- and adhesive dispersal by applying generalized linear mixed-effect models. We used both abundance-weighted and unweighted dispersal traits in order to gain a comprehensive picture of successional developments. Hypothesis (a) was supported by unweighted but not abundance-weighted data. Anemochory showed no consistent changes across sites. In contrast, epizoochory (especially when not weighted by abundance) turned out to be an indicator of the transition from early to mid-successional stages. It increased for the first 9–16 years of succession but declined afterwards. Species richness showed an opposing pattern, while species abundance increased asymptotically. We suggest that plant-animal interactions play a key role in mediating these processes: By importing seeds of highly competitive plant species, animals are likely to promote the increasing abundance of a few dominant, highly epizoochorous species. These species outcompete weak competitors and species richness decreases. However, animals should as well promote the subsequent increase of species richness by disturbing the sites and creating small open patches. These patches are colonized by weaker competitors that are not necessarily dispersed by animals. The changes in the presence of epizoochorous species indicate the importance of plant traits and related plant–animal interactions in the succession of plant communities.     

Relationship between light availability and species richness during fen grassland succession

The complex relation between standing crop and species richness in herbaceous plant communities is still one of the most intensively discussed topics in vegetation ecology. In this study, we focus on the extent to which light availability is able to explain this relationship in fen grasslands. We analyse changes of light availability (measured as relative irradiance, RI), standing crop and species richness during fen grassland succession. Our study include communities representing differences in drainage intensity and nutrient status (‘land use intensification sere’) and successional stages developing after abandonment (‘abandonment seres’). Both in the land use intensification sere and in the abandonment seres, we recorded an increase in standing crop reaching highest values of approx. 800 g m^‐2. RI, species richness and number of small‐growing species declined in the successional seres. RI was highest in stands of low‐productive communities, where light compensation points (5% RI) were already attained at soil surface and light saturation points (30% RI) at soil surface or at 15 cm height. Horizontal heterogeneity of RI at soil surface and at 30 cm height decreased in the abandonment seres, but not at 30 cm height in the land use intensification sere. Furthermore, we detected a signiñcant negative relation between standing crop and RI. Species richness declined with increasing standing crop and thus consequently also with decreasing RI. This result points out that light competition might be of importance for this pattern. The number of small‐growing species decreased exponentially with an increase in standing crop (and decrease in RI). It can be assumed that low light availability negatively affects small‐growing species at standing crop values higher than 400–500 g m². As the maintenance of small‐growing species and the improvement of habitat conditions for their establishment are important aims of nature conservation in fen grasslands, management strategies should be orientated at maintaining standing crops that are not higher than these values.     

黄土高原自然植被演替过程中的植物特征与土壤元素动态

在 3～ 14 9年的时间尺度上 ,对黄土高原植被次生演替过程中植物特征和土壤养分元素动态进行了研究。结果表明 :1)随着演替时间尺度的延伸 ,土壤全C、全N含量呈增加趋势 ,而土壤全K、全Na和土壤pH值呈下降趋势 ,土壤全P变化趋势不明显 ;此外 ,表层 (0～ 10cm)土壤CaO含量在演替系列上呈下降趋势 ,深层 (2 0～ 30cm ,4 0～5 0cm)则呈增加趋势。演替过程对几种土壤化学元素含量影响的程度随着土壤深度的增加而减弱。 2 )植物群落物种丰富度在演替的中间阶段最高。 3)在演替的早期阶段 ,植物群落优势种往往具有稳定的土壤种子库、CR_生活对策和S_繁殖对策 ,在贫瘠的土壤上具有较强的竞争能力 ,且具有较强的水平扩展能力和克隆繁殖能力 ,C_生活对策、在土壤全N含量较高的生境中具有较强竞争能力的多年生植物 ,在演替中后期占据群落的优势地位。此外 ,在所涉及的植物特征中 ,多年生生活史 ,C_、CR_、SC_、SR_、S_生活对策 ,以及R_、W_、Bs_、VBs_和V_繁殖对策等特征在非优势物种中出现的频率较高。 4 )C_、SC_生活对策 ,克隆能力 ,多年生生活史 ,水平扩展能力 ,种子的动物传播方式 ,秋季开花 ,荚果、坚果等特征出现的比例在一定程度上与土壤全C、全N和全K含量正相关 ;而S_、SR_、R_、CR_生活对策 ,一、二年生     

Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats

Changes in traditional agricultural systems in Europe in recent decades have led to widespread abandonment and colonization of various habitats by shrubs and trees. We combined several vegetation databases to test whether patterns of changes in plant diversity after land abandonment in different habitats followed similar pathways. The impacts of land abandonment and subsequent woody colonization on vegetation composition and plant traits were studied in five semi-natural open habitats and two arable habitats in six regions of Europe. For each habitat, vegetation surveys were carried out in different stages of succession using either permanent or non-permanent plots. Consecutive stages of succession were defined on a physiognomic basis from initial open stages to late woody stages. Changes in vegetation composition, species richness, numbers of species on Red Lists, plant strategy types, Ellenberg indicator values of the vegetation, Grime CSR strategy types and seven ecological traits were assessed for each stage of the successional pathway. Abandonment of agro-pastoral land-use and subsequent woody colonization were associated with changes in floristic composition. Plant richness varied according to the different habitats and stages of succession, but semi-natural habitats differed from arable fields in several ecological traits and vegetation responses. Nevertheless, succession occurred along broadly predictable pathways. Vegetation in abandoned arable fields was characterized by a decreasing importance of R-strategists, annuals, seed plants with overwintering green leaves, insect-pollinated plants with hemi-rosette morphology and plants thriving in nutrient-rich conditions, but an increase in species considered as endangered according to the Red Lists. Conversely, changes in plant traits with succession within the initially-open semi-natural habitats showed an increase in plants thriving in nutrient-rich conditions, stress-tolerant plants and plants with sexual and vegetative reproduction, but a sharp decrease in protected species. In conclusion, our study showed a set of similarities in responses of the vegetation in plant traits after land abandonment, but we also highlighted differences between arable fields and semi-natural habitats, emphasizing the importance of land-use legacy.     

高寒草甸不同演替阶段植物叶片功能性状研究

以甘南高寒草甸演替过程中5个阶段的典型群落为研究对象,对围封样地内不同演替阶段群落叶片功能性状进行比较,分析了围封地内各演替阶段群落水平上主要物种叶性状与环境因子的关系。结果显示:(1)此围封地形成了一个从草本到灌木的演替过程。随着演替的进行,Margalef指数显著增加,Shannon-Wiener指数呈先增加后降低的趋势。(2)从演替前期到演替后期,土壤有机碳(SOC)、土壤全氮(STN)、土壤含水量(SWC)逐渐升高,光照度(LI)、土壤温度(ST)逐渐降低,土壤全磷(STP)呈先降低后增加趋势;叶片有机碳(LCC)、全氮(LNC)、含水量(LWC)逐渐升高;比叶面积(SLA)、磷利用效率(PUPE)、稳定碳同位素(δ13C)逐渐下降,叶片全磷(LPC)先降低后升高,而氮利用效率(PUNE)先升高后降低。(3)RDA冗余分析表明,在此围封样地内,演替前期植物群落叶性状主要受到LI和ST的限制作用。而在演替的中后期SWC[WTBZ]是主要影响因子。此研究有助于我们认识高寒草甸生态系统的退化过程所导致的生态环境问题,进而寻求更好的草地恢复和重建方法。     

Getting the Right Traits: Reproductive and Dispersal Characteristics Predict the Invasiveness of Herbaceous Plant Species

To better understand the effect of species traits on plant invasion, we collected comparative data on 20 reproductive and dispersal traits of 93 herbaceous alien species in the Czech Republic, central Europe, introduced after 1500 A. D. We explain plant invasion success, expressed by two measures: invasiveness, i.e. whether the species is naturalized but non-invasive, or invasive; and dominance in plant communities expressed as the mean cover in vegetation plots. We also tested how important reproductive and dispersal traits are in models including other characteristics generally known to predict invasion outcome, such as plant height, life history and residence time. By using regression/classification trees we show that the biological traits affect invasion success at all life stages, from reproduction (seed production) to dispersal (propagule properties), and the ability to compete with resident species (height). By including species traits information not usually available in multispecies analyses, we provide evidence that traits do play important role in determining the outcome of invasion and can be used to distinguish between alien species that reach the final stage of the invasion process and dominate the local communities from those that do not. No effect of taxonomy ascertained in regression and classification trees indicates that the role of traits in invasiveness should be assessed primarily at the species level.     

海岛植物不同演替阶段植物功能性状与环境因子的变化规律

植物功能性状与环境之间的关系是功能性状研究的重点,环境因子驱使植物功能性状发生变化,进而推动群落发生演替。以平潭岛4个不同演替阶段的森林植被(灌草丛、针叶林、针阔混交林、常绿阔叶林)为研究对象,结合不同群落演替阶段的物种特征和群落结构,分析海岛不同演替阶段茎、叶功能性状的变化规律,以及功能性状与环境因子的关系。结果表明:(1)随着演替的进行,土壤养分和水分逐渐增加,土壤pH逐渐下降。比叶面积(SLA)、叶片氮含量(LNC)、叶片磷含量(LPC)、茎氮含量(SNC)、茎磷含量(SPC)下降后上升,叶厚度(LT)、叶片碳含量(LCC)、茎碳含量(SCC)与之相反,叶干物质含量(LDMC)、茎组织密度(STD)逐渐上升。(2)冗余分析表明,演替早期植物主要分布在土壤pH、容重高的贫瘠环境,拥有较高SLA、SNC、SPC、LPC的性状组合;演替后期植物主要分布在土壤养分和水分高的肥沃环境,拥有较高的STD、LDMC、LCC、LNC的性状组合。其中,土壤有机质和全氮含量是影响海岛植物演替过程中功能性状变化的关键环境因子。研究海岛植物功能性状与环境之间的关系随演替的变化规律,探讨各演替阶段功能性状和环境特征,以及功能性状如何响应环境变化。旨在为今后选择合适的树种进行海岛植被修复和重建提供依据。     

武汉东湖水生植物群落演替的研究

 本文根据1992～1993年调查结果并结合前人研究资料,讨论了东湖水生植物群落30多年来的动态变化及其与富营养化和渔业养殖等因素的关系,提出了东湖水生植物群落的演替系列是从微齿眼子菜阶段→微齿眼子菜+大茨藻+金鱼藻+狐尾藻阶段→微齿眼子菜消失阶段→大茨藻阶段→大茨藻+狐尾藻+苦草阶段。同时,还从物种生理生态、补偿和再生能力、生活史及生殖对策、种间关系等方面探讨了水生植物群落演替的物种替代机制及其演替模式,为湖泊水生植被恢复、人工调控和优化提供理论依据。     

Spontaneous vegetation succession in human‐disturbed habitats: A pattern across seres

Abstract. Vegetation samples from 15 successional seres in various disturbed habitats in the western part of the Czech Republic were analysed to detect possible trends. For particular seres, data on species cover were available from the onset to 10–76 yr of succession. All seres started on bare ground. Species which attained at least 1% cover in any sere in any year were used as input data for Canonical Correspondence Analysis, assessing the effect of time as the environmental variable, for Detrended Correspondence Analysis and TWINSPAN classification. Two distinct groups ofseres were distinguished: ‘ruderal’, occurring in agricultural, industrial or urban landscapes altered by men, usually on fertile sites; and ‘non‐ruderul’, occurring in less altered, mostly forested landscapes, usually on acid, nutrient‐poor and wetter soils. The former type of succession starts with ruderal annuals, being followed by ruderal perennials. In the latter case non‐ruderal clonal perennials prevail from the onset of succession. The landscape frame is emphasized, beside site environmental conditions, as influencing the type of succession. The character of species attaining dominance in succession, participation of dominant woody plants and the character of late successional stages, i.e. features important from the point of view of potential restoration of human‐disturbed habitats, are discussed.     

黔东南州白云岩喀斯特区植物群落演化特征

该研究采用时空互代法、连续带指数法等,对贵州省黔东南州的白云岩喀斯特区60个具有代表性的典型样地的植物种组组成、以及各演替阶段植物群落高度、密度、生物量、生物多样性指数、均匀度指数、生态优势度指数等进行调查分析,以探讨白云岩喀斯特区植物群落的演化特征,为深入研究白云岩喀斯特区植物群落自然恢复规律奠定基础。结果表明:(1)各种组的优势种组成不同,种组替代规律依次为先锋种、次先锋种、过渡种,最终被顶极种替代的过程。(2)群落的高度、密度、生物量等结构特征随植物群落演替发展呈逐渐增大趋势,变化范围分别为0.58～9.54m、585～3 145株·hm-2、8.45～128.56t·hm-2。(3)植物群落的物种丰富度随演替阶段的发展呈下降趋势,从草本群落阶段到顶极群落阶段物种数由48种降低到10种。(4)随着植物群落演替阶段的发展,生物多样性指数呈先升高后降低的趋势,即由草本群落的3.48升高到草灌群落的4.73,后降到顶极群落的3.46;均匀度指数呈降低趋势,最高为草本群落阶段的0.95,最低为常绿阔叶林阶段的0.78;生态优势度指数逐渐升高,顶极群落达到最大,为0.10。     

Changes in diversity and importance of clonal plants during sand dune succession in northeastern China

Many major biomes throughout the world are dominated by plants with clonal growth forms. While many recent studies have examined the effects of clonality on the growth of individual plants, relatively few studies have tested the community level effects of clonality as a function of environmental characteristics. By investigating six sand dune sites that have undergone different numbers of years of natural restoration constituting a succession sere, we quantified if the abundance and importance of clonal plants was related to successional age in the stressful environment of a semi-arid sand dune region in northeastern China. We expected that clonal plants would dominate at every stage of the succession sere. We also predicted that species diversity would decrease in later stages of the succession sere due to the extremely high proportion of clonal plants in the community. Our results showed that, through 45 years of succession, the total plant species richness and Shannon–Wiener diversity index continually increased. While the species number of clonal plants was consistently low during the succession, the importance of clonal plants increased gradually from none at 3 years to 49 % of the total, approximately equal to that of aclonal plants, at the 45-year site. Clonal plants with phalanx strategies were more important than guerillas at all ages in sand dune succession. At the beginning and early stages of sand dune succession, aclonal plants were more important than clonal plants, perhaps due to greater seed propagation. The distribution or arrangement of aclonal and clonal plants in the whole process of sand dune complemented each other. The results presented give new perceptions on the function of biodiversity in maintaining ecosystems.     

基于功能性状的毛乌素沙地不同演替阶段适生植物筛选

沙地生态系统修复是恢复生态学研究的热点问题,适生植物筛选是修复的关键。植物功能性状反映了植物在不同环境中的生存策略,探究沙地植物功能性状及其与环境之间的关系,有助于筛选用于植被恢复的物种,为保护沙地生态系统提供理论依据。以毛乌素沙地为研究区,分析了1983-2015年间沙地典型飞播样地群落演替特征及其对环境因子的响应,建立基于10个植物功能性状的毛乌素沙地潜在种库,进一步筛选飞播恢复下沙地不同演替阶段的适生植物。研究表明:(1)飞播恢复下的毛乌素沙地植物群落分为三个演替阶段:固沙先锋物种群落、沙生植物为主的杂类草群落、中生植物为主的杂类草群落。(2)土壤因子是群落演替的主要驱动力,其中土壤全氮、土壤总有机碳、土壤硝态氮是影响群落演替的关键因素。(3)基于功能性状筛选出29种适生物种用于植被恢复,演替第一阶段可用雾冰藜、猪毛菜等,演替第二阶段可用拂子茅、无芒隐子草等,演替第三阶段可用草地风毛菊、猪毛蒿等。通过物种功能性状特征可以快速选择适合沙地退化生态系统修复的候选物种,为植被恢复提供了一定的理论支持。     

Does the sequence of plant dominants affect mycorrhiza development in simulated succession on spoil banks?

The aim of this field study was to examine how the development of arbuscular mycorrhizal fungi (AMF) on coal mine spoil banks is affected by the presence of plants with different mycorrhizal status. A 3-year trial was conducted on the freshly created spoil bank Vršany, North-Bohemian coal basin, the Czech Republic. Three plant species – non-mycotrophic annual Atriplex sagittata, highly mycotrophic annual Tripleurospermum inodorum (both dominants of early stages of succession) and facultatively mycotrophic Arrhenatherum elatius (a perennial grass species of the later stage of succession) – were planted on 1 m² plots over 3 years in different sequences that simulated the progress of succession on spoil banks. The development of AMF populations was monitored by evaluation of mycorrhizal colonization of plant roots and by measurement of the mycorrhizal inoculation potential (MIP) of soil. These two parameters were compared between plots inoculated with the mixture of three AMF isolates – Glomus mosseae BEG95, G. claroideum BEG96 and G. intraradices BEG140 – (“inoculated plots”) and plots exposed only to natural dispersal of AMF propagules (“uninoculated plots”). Highly colonized roots of plants together with a high MIP of soil in uninoculated plots were already found at the end of the first season, indicating rapid natural dispersal of AMF propagules. Root colonization of facultatively mycotrophic and non-mycotrophic plants in later years was affected by the mycorrhizal status of the previous plant species. The MIP of soil continuously increased throughout the experiment; in uninoculated plots, the MIP was temporarily decreased if plant species of higher mycotrophy were replaced by species of lower mycotrophy. The results lead to the conclusion that AMF colonize freshly formed sites very quickly and reproduce or accumulate in the soil, which leads to increasing MIP values. However, this infective potential can be decreased if non-mycotrophic plants predominate on the site.     

Changing assembly rules during secondary succession: evidence for non-random patterns

Describing the rules of community assembly is a central topic of ecology. Studying successional processes through a trait-based null model approach can help to better understand the rules of community assembly.According to theoretical considerations, at the beginning of succession - after getting over the dispersal limitation stage - community composition is primarily shaped by environmental filters (generating functional convergence), while in later stages limiting similarity (generating functional divergence) will be dominant. However, empirical evidence does not clearly support theoretical expectations.Our aim was to detect the presence and changes of trait-based assembly processes during old-field succession based on twelve traits. Changes in vegetation composition were evaluated by a combination of time series and space-for-time substitution: conducting three resurveys of permanent plots on four old-field age-groups. The individual dispersion of traits was transformed into effect size (i.e. departure from null model expectation). The impact of time since abandonment on effect sizes was tested by generalized additive mixed effect models.We detected a non-random pattern for each trait in at least some part of the succession. Departure from randomness did not change significantly over time for six traits: seed mass, lateral spread and pollination type were divergent, while leaf size, generative height and length of flowering were convergent. Six traits had changing patterns along the succession. Four of them showed increasing divergence (e.g. dispersal type, LDMC), which supports our hypothesis. While two (SLA, life form) displayed increasing convergence, contrary to expectations.We confirmed the general hypothesis that convergence is predominant initially and that divergence can be detected later in succession for four traits. However, the large variation found in trait dispersion indicates that complex processes operate during succession.     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.     

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.