Arbuscular mycorrhizal fungal communities change among three stages of primary sand dune succession but do not alter plant growth

Plant interactions with soil biota could have a significant impact on plant successional trajectory by benefiting plants in a particular successional stage over others. The influence of soil mutualists such as mycorrhizal fungi is thought to be an important feedback component, yet they have shown benefits to both early and late successional plants that could either retard or accelerate succession. Here we first determine if arbuscular mycorrhizal (AM) fungi differ among three stages of primary sand dune succession and then if they alter growth of plants from particular successional stages. We isolated AM fungal inoculum from early, intermediate or late stages of a primary dune succession and compared them using cloning and sequencing. We then grew eight plant species that dominate within each of these successional stages with each AM fungal inoculum. We measured fungal growth to assess potential AM functional differences and plant growth to determine if AM fungi positively or negatively affect plants. AM fungi isolated from early succession were more phylogenetically diverse relative to intermediate and late succession while late successional fungi consistently produced more soil hyphae and arbuscules. Despite these differences, inocula from different successional stages had similar effects on the growth of all plant species. Host plant biomass was not affected by mycorrhizal inoculation relative to un‐inoculated controls. Although mycorrhizal communities differ among primary dune successional stages and formed different fungal structures, these differences did not directly affect the growth of plants from different dune successional stages in our experiment and therefore may be less likely to directly contribute to plant succession in sand dunes.     

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.     

Phylogenetic turnover during subtropical forest succession across environmental and phylogenetic scales

Although spatial and temporal patterns of phylogenetic community structure during succession are inherently interlinked and assembly processes vary with environmental and phylogenetic scales, successional studies of community assembly have yet to integrate spatial and temporal components of community structure, while accounting for scaling issues. To gain insight into the processes that generate biodiversity after disturbance, we combine analyses of spatial and temporal phylogenetic turnover across phylogenetic scales, accounting for covariation with environmental differences. We compared phylogenetic turnover, at the species‐ and individual‐level, within and between five successional stages, representing woody plant communities in a subtropical forest chronosequence. We decomposed turnover at different phylogenetic depths and assessed its covariation with between‐plot abiotic differences. Phylogenetic turnover between stages was low relative to species turnover and was not explained by abiotic differences. However, within the late‐successional stages, there was high presence‐/absence‐based turnover (clustering) that occurred deep in the phylogeny and covaried with environmental differentiation. Our results support a deterministic model of community assembly where (i) phylogenetic composition is constrained through successional time, but (ii) toward late succession, species sorting into preferred habitats according to niche traits that are conserved deep in phylogeny, becomes increasingly important.     

系统发育多样性与系统发育结构在岛屿植物群落保护中的意义——以蜈支洲岛为例

为探讨系统发育多样性和系统发育结构在岛屿植物群落保护决策制定中的作用,以海南三亚蜈支洲岛被子植物群系为例,分析物种丰富度和系统发育多样性的相关性。结果表明,蜈支洲岛植被可分为10种群系类型,22个样方的物种丰富度与系统发育多样性呈显著正相关,但系统发育多样性与物种丰富度对植物多样性保护优先顺序的指示有差异。森林群系的物种丰富度和系统发育多样性普遍高于灌木和草本群系,4个森林群系样方表现为系统发育结构发散,3个森林群系样方表现为系统发育结构聚集。灌木群系中露兜树(Pandanus tectorius)灌丛和草海桐(Scaevola sericea)灌丛系统发育多样性较高,草本群系中厚藤(Vitex trifolia)灌草丛系统发育多样性较高,灌木和草本群系大多倾向于系统发育结构聚集。距海或人类活动区较近的群系可能受环境过滤的作用,而远离海和人类活动区的群系主要受竞争排斥作用影响。因此,在岛屿植物多样性保护策略制定中应当综合考虑物种和系统发育维度,以及环境、演替阶段及用岛方式等因素的影响。     

Convergence of community composition during secondary succession on Zokor rodent mounds on the Tibetan Plateau

Aims The community succession theory is much debated in ecology. We studied succession on Zokor rodent mounds on the Tibetan Plateau to address several fundamental questions, among them: (i) During secondary succession, does the community composition converge towards one community state or multiple states depending on the initial colonization? (ii) Do mound communities located in different background communities exhibit different assembly trajectories?Methods In a sub-alpine meadow, we investigated a total of 80 mound communities at several successional stages in three different background communities resulting from different management histories and compared their changes in species composition. The distribution of plant communities over time was analyzed with quantitative classification and ordination methods. The co-occurrence patterns of species were evaluated at each successional stage, and the degree of convergence/divergence among communities was obtained by calculating two beta-diversity indices.Important findings During secondary succession, species richness of mound communities changed over time, and this change was dependent on the background community. Five life-form groups exhibited different dynamic patterns in species richness and plant cover. Community composition and the degree of species co-occurrence between communities increased over time since disturbance. There was much variation in species composition at earlier stages of succession, but communities on older mounds became more similar to each other and to their surrounding vegetation over the course of secondary succession. Post-disturbance succession of Zokor mound communities transitioned from 'multiple alternative states' to 'background-based deterministic community assembly' over time. Tradeoffs between competition and colonization, as well as the characteristics of different life-forms and mass effects within a limited species pool are the mechanisms responsible for convergence of mound communities.     

Mycorrhizal fungal growth responds to soil characteristics,but not host plant identity,during a primary lacustrine dune succession

Soil factors and host plant identity can both affect the growth and functioning of mycorrhizal fungi. Both components change during primary succession, but it is unknown if their relative importance to mycorrhizas also changes. This research tested how soil type and host plant differences among primary successional stages determine the growth and plant effects of arbuscular mycorrhizal (AM) fungal communities. Mycorrhizal fungal community, plant identity, and soil conditions were manipulated among three stages of a lacustrine sand dune successional series in a fully factorial greenhouse experiment. Late succession AM fungi produced more arbuscules and soil hyphae when grown in late succession soils, although the community was from the same narrow phylogenetic group as those in intermediate succession. AM fungal growth did not differ between host species, and plant growth was similarly unaffected by different AM fungal communities. These results indicate that though ecological filtering and/or adaptation of AM fungi occurs during this primary dune succession, it more strongly reflects matching between fungi and soils, rather than interactions between fungi and host plants. Thus, AM fungal performance during this succession may not depend directly on the sequence of plant community succession.     

放牧干扰梯度下川西亚高山植物群落的组合机理

为了阐明放牧干扰对川西亚高山区域植物群落的组合过程以及群落结构的影响, 研究了放牧干扰梯度下的功能群均匀度和群落谱系结构的变化趋势。结果显示: 在干扰较轻的阔叶林与针叶林样地, 部分样方的功能群均匀度显著高于无效模型, 随着干扰梯度的增强, 功能群均匀度呈线性下降, 样方平均值从0.930降至0.840, 其高于无效模型的次数也逐渐降低, 干扰程度较大的草甸中出现部分样方的功能群均匀度显著低于无效模型。随着干扰程度的增强, 群落的谱系结构指数也呈逐渐上升趋势, 净关联指数平均值由-0.634逐渐增加至2.360, 邻近类群指数由-0.158上升至2.179。草甸与低矮灌丛受干扰较为严重, 其大部分样方的谱系结构指数显著高于随机群落, 表明干扰群落的谱系结构呈聚集分布。功能群均匀度与谱系结构的变化趋势一致, 表明生境筛滤效应与种间竞争作用的平衡决定着群落的组合过程。干扰降低了竞争作用, 促进了少数耐干扰功能群的优势地位, 造成功能群均匀度下降, 同时通过生境筛滤作用, 使群落的谱系结构呈现出聚集分布; 而未干扰的群落中由于竞争作用的效应, 功能群均匀度较高, 谱系结构也更加分散。研究区域植物群落的功能群均匀度与物种丰富度呈负相关, 表明物种间特别是相似物种间的竞争限制了群落的物种多样性。研究结果说明, 生态位分化和物种间的相互竞争在物种共存与群落组合中具有重要作用。     

Primary succession of Bistorta vivipara (L.) Delabre (Polygonaceae) root‐associated fungi mirrors plant succession in two glacial chronosequences

Glacier chronosequences are important sites for primary succession studies and have yielded well‐defined primary succession models for plants that identify environmental resistance as an important determinant of the successional trajectory. Whether plant‐associated fungal communities follow those same successional trajectories and also respond to environmental resistance is an open question. In this study, 454 amplicon pyrosequencing was used to compare the root‐associated fungal communities of the ectomycorrhizal (ECM) herb Bistorta vivipara along two primary succession gradients with different environmental resistance (alpine versus arctic) and different successional trajectories in the vascular plant communities (directional replacement versus directional non‐replacement). At both sites, the root‐associated fungal communities were dominated by ECM basidiomycetes and community composition shifted with increasing time since deglaciation. However, the fungal community's successional trajectory mirrored the pattern observed in the surrounding plant community at both sites: the alpine site displayed a directional‐replacement successional trajectory, and the arctic site displayed a directional‐non‐replacement successional trajectory. This suggests that, like in plant communities, environmental resistance is key in determining succession patterns in root‐associated fungi. The need for further replicated study, including in other host species, is emphasized.     

High specialization and limited structural change in plant‐herbivore networks along a successional chronosequence in tropical montane forest

Secondary succession is well‐understood, to the point of being predictable for plant communities, but the successional changes in plant‐herbivore interactions remains poorly explored. This is particularly true for tropical forests despite the increasing importance of early successional stages in tropical landscapes. Deriving expectations from successional theory, we examine properties of plant‐herbivore interaction networks while accounting for host phylogenetic structure along a succession chronosequence in montane rainforest in Papua New Guinea. We present one of the most comprehensive successional investigations of interaction networks, equating to > 40 person years of field sampling, and one of the few focused on montane tropical forests. We use a series of nine 0.2 ha forest plots across young secondary, mature secondary and primary montane forest, sampled almost completely for woody plants and larval leaf chewers (Lepidoptera) using forest felling. These networks comprised of 12 357 plant‐herbivore interactions and were analysed using quantitative network metrics, a phylogenetically controlled host‐use index and a qualitative network beta diversity measure. Network structural changes were low and specialisation metrics surprisingly similar throughout succession, despite high network beta diversity. Herbivore abundance was greatest in the earliest stages, and hosts here had more species‐rich herbivore assemblages, presumably reflecting higher palatability due to lower defensive investment. All herbivore communities were highly specialised, using a phylogenetically narrow set of hosts, while host phylogenetic diversity itself decreased throughout the chronosequence. Relatively high phylogenetic diversity, and thus high diversity of plant defenses, in early succession forest may result in herbivores feeding on fewer hosts than expected. Successional theory, derived primarily from temperate systems, is limited in predicting tropical host‐herbivore interactions. All succession stages harbour diverse and unique interaction networks, which together with largely similar network structures and consistent host use patterns, suggests general rules of assembly may apply to these systems.     

Effects of earthworms and arbuscular mycorrhizal fungi depend on the successional stage of a grassland plant community

Earthworms and arbuscular mycorrhizal fungi (AMF) have profound impacts on plant performance. However, it is largely unknown if and how earthworms and AMF may affect plant succession. We planted mesocosms with an early-mid successional and a mid-late successional grassland plant community and added endogeic earthworms and commercial AMF in a full-factorial way to natural background soil. Earthworms had a positive effect on the total root and shoot biomass of both plant communities, with the effect on the shoots being slightly enhanced by co-inoculation with AMF. Surprisingly, the earthworm effect on the mid successional plant species depended on the successional stage of the plant community. Earthworms had a positive effect on the mid successional plant species when they were growing in the mid-late successional plant community, but no effect when the same plant species were growing in the early-mid successional plant community. Addition of AMF alone tended to reduce the shoot biomass of the early successional plant species, while the addition of earthworms in the presence or absence of AMF increased their shoot biomass. We conclude that the impacts of earthworms on plant species may depend on the successional stage of the plant community, while the effect of AMF addition depends on the successional stage of the plant community and may be changed by the presence of earthworms. Earthworms and AMF addition affect plants and plant communities of different successional stages differently with potential effects on plant succession.     

Examining differences in phylogenetic composition enhances understanding of the phylogenetic structure of the shrub community in the northeastern Qinghai‐Tibetan Plateau

Periodic climatic oscillations and species dispersal during the postglacial period are two important causes of plant assemblage and distribution on the Qinghai‐Tibet Plateau (QTP). To improve our understanding of the bio‐geological histories of shrub communities on the QTP, we tested two hypotheses. First, the intensity of climatic oscillations played a filtering role during community structuring. Second, species dispersal during the postglacial period contributed to the recovery of species and phylogenetic diversity and the emergence of phylogenetic overdispersion. To test these hypotheses, we investigated and compared the shrub communities in the alpine and desert habitats of the northeastern QTP. Notably, we observed higher levels of species and phylogenetic diversity in the alpine habitat than in the desert habitat, leading to phylogenetic overdispersion in the alpine shrub communities versus phylogenetic clustering in the desert shrub communities. This phylogenetic overdispersion increased with greater climate anomalies. These results suggest that (a) although climate anomalies strongly affect shrub communities, these phenomena do not act as a filter for shrub community structuring, and (b) species dispersal increases phylogenetic diversity and overdispersion in a community. Moreover, our investigation of the phylogenetic community composition revealed a larger number of plant clades in the alpine shrub communities than in the desert shrub communities, which provided insights into plant clade‐level differences in the phylogenetic structures of alpine and desert shrub communities in the northeastern QTP.     

Plant succession and rhizosphere microbial communities in a recently deglaciated alpine terrain

This study describes how early and late successional plant species affect soil microorganisms in alpine ecosystems. We quantify the relative importance of plant species and soil properties as determinants of belowground microbial communities. Sixteen plant species were selected from six successional stages (4–14–20–43–75–135 years) within the foreland of the Rotmoosferner glacier, Austria, and at one (reference) site outside the foreland. The size, composition and function of the communities of microorganism in the bulk soil and the rhizosphere were characterized by ninhydrin-reactive nitrogen, phospholipid fatty acids and enzyme activities (β-glucosidase, β-xylosidase, N-acetyl-β-glucosaminidase, leucine aminopeptidase, acid phosphatase, sulphatase). The results show that the microbial data could be grouped according to early (up to 43 years) and late-colonizing plant species (75 or more years). In early succession, no plant species or soil age effect was detected on the microbial biomass, phospholipid fatty acids, or enzyme activity. The rhizosphere microbial community was similar to that in the bulk soil, which in turn was determined by the abiotic environmental conditions. In late succession, improved soil conditions probably mediated plant species effects on the belowground microbial community. The most pronounced rhizosphere effects were attributed to plant species of the 75- and 135-year-old sites. The microbial colonization (size, composition, activity) of the bulk soil predominantly followed changes in vegetation cover, plant life forms and soil organic matter. In summary, the observed successional pattern of the above- and belowground communities provides an example of the facilitation models of primary succession.     

Impact of alien species on dune systems: a multifaceted approach

We applied a multifaceted approach, in terms of taxonomic, phylogenetic and functional diversity, to study at fine scale how three plant communities occurring in a Mediterranean dune have been affected by the encroachment of alien species. We sampled 81 sites in a Site of Community Importance in Central Italy. Past and present land use/cover data have been derived using GIS and remote sensing tools. Information on plants phylogenesis and functional traits has been gathered from several databases. Ecological variables have been collected. GLMs in conjunction with an Information Based approach were used to model species composition, richness and phylogenetic diversity. Multivariate analysis has been used to study functional diversity. The results outlined how total species richness is related to recent land transformations and to a set of environmental factors. The analyses of functional and phylogenetic diversity support the idea that alien species significantly affect the functional and phylogenetic characteristics of the native plant communities. Habitat filtering seems to be predominant in not-invaded plots, whereas limiting similarity/niche differentiation is predominant in driving community assembly of invaded communities. The attained scenario depicts the spread of a reduced group of alien species phylogenetically and functionally well-differentiated, able to reduce the abundance of native species, not to exclude them though. Ultimately, the multifaceted approach assisted in understanding the community assembly of dune vegetation, and to discern the relative impact of alien species on native plant communities. Such approach represents a crucial step to achieve an efficient management of dune habitats, as useful tool to monitor and to effectively protect their biodiversity and functioning.     

Multi-scale phylogenetic structure in coastal dune plant communities across the globe

Aims Studies integrating phylogenetic history and large-scale community assembly are few, and many questions remain unanswered. Here, we use a global coastal dune plant data set to uncover the important factors in community assembly across scales from the local filtering processes to the global long-term diversification and dispersal dynamics. Coastal dune plant communities occur worldwide under a wide range of climatic and geologic conditions as well as in all biogeographic regions. However, global patterns in the phylogenetic composition of coastal dune plant communities have not previously been studied.Methods The data set comprised vegetation data from 18463 plots in New Zealand, South Africa, South America, North America and Europe. The phylogenetic tree comprised 2241 plant species from 149 families. We calculated phylogenetic clustering (Net Relatedness Index, NRI, and Nearest Taxon Index, NTI) of regional dune floras to estimate the amount of in situ diversification relative to the global dune species pool and evaluated the relative importance of land and climate barriers for these diversification patterns by geographic analyses of phylogenetic similarity. We then tested whether dune plant communities exhibit similar patterns of phylogenetic structure within regions. Finally, we calculated NRI for local communities relative to the regional species pool and tested for an association with functional traits (plant height and seed mass) thought to vary along sea–inland gradients.Important findings Regional species pools were phylogenetically clustered relative to the global pool, indicating regional diversification. NTI showed stronger clustering than NRI pointing to the importance of especially recent diversifications within regions. The species pools grouped phylogenetically into two clusters on either side of the tropics suggesting greater dispersal rates within hemispheres than between hemispheres. Local NRI plot values confirmed that most communities were also phylogenetically clustered within regions. NRI values decreased with increasing plant height and seed mass, indicating greater phylogenetic clustering in communities with short maximum height and good dispersers prone to wind and tidal disturbance as well as salt spray, consistent with environmental filtering along sea–inland gradients. Height and seed mass both showed significant phylogenetic signal, and NRI tended to correlate negatively with both at the plot level. Low NRI plots tended to represent coastal scrub and forest, whereas high NRI plots tended to represent herb-dominated vegetation. We conclude that regional diversification processes play a role in dune plant community assembly, with convergence in local phylogenetic community structure and local variation in community structure probably reflecting consistent coastal-inland gradients. Our study contributes to a better understanding of the globally distributed dynamic coastal ecosystems and the structuring factors working on dune plant communities across spatial scales and regions.     

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.     

Phylogenetic community structure in Minnesota oak savanna is influenced by spatial extent and environmental variation

The relative importance of environmental filtering, biotic interactions and neutral processes in community assembly remains an openly debated question and one that is increasingly addressed using phylogenetic approaches. Closely related species may occur together more frequently than expected (phylogenetic clustering) if environmental filtering operates on traits with significant phylogenetic signal. Recent studies show that phylogenetic clustering tends to increase with spatial scale, presumably because greater environmental variation is encompassed at larger spatial scales, providing opportunities for species to sort across environmental gradients. However, if environmental filtering is the cause of species sorting along environmental gradients, then environmental variation rather than spatial scale per se should drive the processes governing community assembly. Using species abundance and light availability data from a long‐term experiment in Minnesota oak savanna understory communities, we explicitly test the hypothesis that greater environmental variation results in greater phylogenetic clustering when spatial scale is held constant. Concordant with previous studies, we found that phylogenetic community structure varied with spatial extent. At the landscape scale (~1000 ha), communities were phylogenetically clustered. At the local scale (0.375ha), phylogenetic community structure varied among plots. As hypothesized, plots encompassing the greatest environmental variation in light availability exhibited the strongest phylogenetic clustering. We also found strong correlations between species functional traits, particularly specific leaf area (SLA) and perimeter per area (PA), and species light availability niche. There was also a phylogenetic signal in both functional traits and species light availability niche, providing a mechanistic explanation for phylogenetic clustering in relation to light availability. We conclude that the pattern of increased phylogenetic clustering with increased environmental variation is a consequence of environmental filtering acting on phylogenetically conserved functional traits. These results indicate that the importance of environmental filtering in community assembly depends not on spatial scale per se, but on the steepness of the environmental gradient.     

陕西子午岭生态因素对植物群落的影响

为探讨生境对植被格局分布的影响,对黄土高原马栏林区60块样地进行植被学调查的基础上,采用17个环境指标刻画植物群落的空间位置、地形和土壤特征;利用双向指示种分析(TWINSPAN)划分了该区不同演替阶段的植物群落类型;利用前向选择法(forward selection)及Monte Carlo检验对不同演替阶段植物群落物种组成影响显著(p＜0.05)的环境因子进行筛选;利用去势典范对应分析(DCCA)排序方法分析不同演替阶段植物群落分布格局与环境的关系;利用偏典范对应分析(partial CCA)定量分离环境、空间及其交互作用对植被格局总体变异的影响.结果表明:(1)马栏林区的植物群落可划分为13个类型,分别属于4个不同的演替阶段;(2)对演替初期阶段群落影响显著的环境因子是土壤含盐量和碱解氮,对演替过渡阶段群落影响显著的环境因子是海拔和腐殖质厚度,对演替亚顶级阶段群落影响显著的因子是海拔、坡向、枯落物厚度、腐殖质厚度和pH值,而对演替顶级阶段群落分异影响显著的因子是海拔、坡向、pH值和速效磷;(3)不同演替阶段群落的生态学特性和分布规律与环境空间的生态梯度格局吻合较好;(4)随着演替的进行,环境因子单独对植物群落的影响越来越大,而样地位置单独作用和样地位置与环境因子的交互作用之和越来越小.