Priority Effects of Time of Arrival of Plant Functional Groups Override Sowing Interval or Density Effects: A Grassland Experiment

Priority effects occur when species that arrive first in a habitat significantly affect the establishment, growth, or reproduction of species arriving later and thus affect functioning of communities. However, we know little about how the timing of arrival of functionally different species may alter structure and function during assembly. Even less is known about how plant density might interact with initial assembly. In a greenhouse experiment legumes, grasses or forbs were sown a number of weeks before the other two plant functional types were sown (PFT) in combination with a sowing density treatment. Legumes, grasses or non-legume forbs were sown first at three different density levels followed by sowing of the remaining PFTs after three or six-weeks. We found that the order of arrival of different plant functional types had a much stronger influence on aboveground productivity than sowing density or interval between the sowing events. The sowing of legumes before the other PFTs produced the highest aboveground biomass. The larger sowing interval led to higher asymmetric competition, with highest dominance of the PFT sown first. It seems that legumes were better able to get a head-start and be productive before the later groups arrived, but that their traits allowed for better subsequent establishment of non-legume PFTs. Our study indicates that the manipulation of the order of arrival can create priority effects which favour functional groups of plants differently and thus induce different assembly routes and affect community composition and functioning.     

Resource availability mediates the importance of priority effects in plant community assembly and ecosystem function

Assembly history, including the order in which species arrive into a community, can influence long‐term community structure; however we know less about how timing of species arrival may alter assembly especially under varying resource conditions. To explore how the timing of species arrival interacts with resource availability to alter community assembly, we constructed experimental plant communities and manipulated the interval between plantings of groups of seedlings (0, 5, 10, 15 or 20 days) at low and high levels of soil nutrient supply. To see if community changes influenced ecosystem‐scale processes, we measured parameters across the plant–soil continuum (e.g. plant biomass and net ecosystem carbon dioxide exchange). We found that the timing of species arrival had a large impact on community assembly, but the size of the effect depended on soil fertility. As planting interval increased, plant communities diverged further from the control, but the divergence was stronger at high than at low nutrient supply. Our data suggest that at high nutrient supply, early‐planted species preempted light resources more quickly, thus preventing the successful establishment of later arriving species even at short planting intervals. Finally, we found that assembly related divergence in plant communities scaled to impact ecosystem‐level characteristics such as green leaf chemistry, but had little effect on total community biomass and net ecosystem exchange of CO₂ and water vapor. Our data indicate that the effect of a stochastic factor, here the timing of species arrival on community composition, depends on the resource level under which the community assembles.     

Subordinate plant species impact on soil microbial communities and ecosystem functioning in grasslands: Findings from a removal experiment

Despite their low relative abundance, subordinate plant species may have larger impacts on ecosystem functioning than expected, but their role in plant communities remains poorly understood. The aim of this study was to test how subordinate plant species influence the functioning of a species-rich semi-natural grasslands. A plant removal experiment was set-up in the mountain grasslands of the Jura Mountains (Switzerland) to test the impact of subordinate plant species on soil microbial communities and ecosystem functioning. The experiment included three treatments: removal of all subordinate species, partial biomass removal of dominant species, and a no biomass removal control. After 2 years of treatments, we determined soil microbial community (bacteria and mycorrhizal fungi) by T-RFLP analysis and measured litter decomposition, soil respiration, soil inorganic nitrogen (DIN) availability and throughout above-ground biomass production as measures of ecosystem function. The removal of subordinate plant species strongly affected bacterial and weakly influenced mycorrhizal fungi communities and decreased rates of plant litter decomposition, soil respiration and DIN availability with larger effects than the partial loss of dominant biomass. The removal of subordinate plant species did not modify plant community structure, but it did reduce total above-ground biomass production compared to the control plots. Collectively, our findings indicate that the loss of subordinate species can have significant consequences for soil microbial communities and ecosystem functions, suggesting that subordinate species are important drivers of ecosystem properties.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

Biodiversity acts as insurance of productivity of bacterial communities under abiotic perturbations

Anthropogenic disturbances are detrimental to the functioning and stability of natural ecosystems. Critical ecosystem processes driven by microbial communities are subjected to these disturbances. Here, we examine the stabilizing role of bacterial diversity on community biomass in the presence of abiotic perturbations such as addition of heavy metals, NaCl and warming. Bacterial communities with a diversity gradient of 1–12 species were subjected to the different treatments, and community biomass (OD₆₀₀) was measured after 24 h. We found that initial species richness and phylogenetic structure impact the biomass of communities. Under abiotic perturbations, the presence of tolerant species in community largely contributed in community biomass production. Bacterial diversity stabilized the biomass across the treatments, and differential response of bacterial species to different perturbations was the key reason behind these effects. The results suggest that biodiversity is crucial for maintaining the stability of ecosystem functioning and acts as ecological insurance under abiotic perturbations. Biodiversity in natural ecosystems may also uphold the ecosystem functioning under anthropogenic disturbance.     

藏香猪翻拱型放牧对不同类型高寒草甸植物特征的影响及土壤环境驱动力

生态系统脆弱的高寒草甸是生态学研究的热点。藏香猪翻拱土壤并取食地下草根层,是藏族自治区一种特有的放牧类型,关于该放牧类型对藏区高寒草甸的影响尚缺乏研究。以滇西北高原国际重要湿地——纳帕海流域的典型高寒草甸为对象,研究了不同高寒草甸类型(陆生草甸与沼泽化草甸)中的植物群落结构特征对藏香猪翻拱型放牧的响应。结果表明,翻拱型放牧显著降低了植被盖度(63.5%)、地上生物量(84.6%)与地下生物量(97.4%),并促进植物生物量向地上部分的再分配。翻拱型放牧下随着植被盖度与生物量降低,植物竞争强度减小,植株高度与空间利用率提高。然而,不同草甸类型中的植物群落结构对翻拱型放牧表现出差异响应。陆生草甸上的群落组成基本未变,而沼泽化草甸上的优势植物华扁穗草(Blysmus sinocompyessus)向1年生水生植物水蓼(Polygonum hydropiper)演替。土壤含水率是导致植物响应差异的关键驱动力。综上,藏香猪翻拱型放牧对植物生物量及盖度等特征均产生不利影响,导致高寒草甸生物量积累锐减,而植物群落结构的变化是放牧干扰与土壤水分、空间竞争协同作用的结果,反映出植物在不同环境中对放牧干扰的差异化响应。研究区域环境特征与放牧干扰的耦合关系,可以为藏香猪放牧作用下的高寒草地植物多样性保护及恢复提供理论依据。     

Biodiversity effects increase linearly with biotope space

Understanding the influence of environmental variation on the relationship between biodiversity and ecosystem functioning is of theoretical and practical interest. We predicted that the strength of this relationship should increase with available biotope space (the physical space associated with a species’ niche) due to increased niche complementarity between species. In this study, biotope space specifically refers to soil volume which is associated with the niche dimension of nutrient acquisition. We tested our prediction by growing plant communities on a gradient of increasing soil depth and volume, offering increased rooting space to species. Our results provide support for a linear increase of the magnitude of positive biodiversity effects on above‐ and belowground community biomass with increasing biotope space. This increase was caused by complementarity effects between species. Soil erosion may thus reduce intercropping benefits.     

Diversity and community biomass depend on dispersal and disturbance in microalgal communities

The evidence for species diversity effects on ecosystem functions is mainly based on studies not explicitly addressing local or regional processes regulating coexistence or the importance of community structure in terms of species evenness. In experimental communities of marine benthic microalgae, we altered the successional stages and thus the strength of local species interactions by manipulating rates of dispersal and disturbance. The treatments altered realized species richness, evenness and community biomass. For species richness, dispersal mattered only at high disturbance rates; when opening new space, dispersal led to maximized richness at intermediate dispersal rates. Evenness, in contrast, decreased with dispersal at low or no disturbance, i.e. at late successional stages. Community biomass showed a non-linear hump-shaped response to increasing dispersal at all disturbance levels. We found a positive correlation between richness and biomass at early succession, and a strong negative correlation between evenness and biomass at late succession. In early succession both community biomass and richness depend directly on dispersal from the regional pool, whereas the late successional pattern shows that if interactions allow the most productive species to become dominant, diverting resources from this species (i.e. higher evenness) reduces production. Our study emphasizes the difference in biodiversity–function relationships over time, as different mechanisms contribute to the regulation of richness and evenness in early and late successional stages.     

Rare taxa maintain the stability of crop mycobiomes and ecosystem functions

Plants harbour highly diverse mycobiomes which sustain essential functions for host health and productivity. However, ecological processes that govern the plant–mycobiome assembly, interactions and their impact on ecosystem functions remain poorly known. Here we characterized the ecological role and community assembly of both abundant and rare fungal taxa along the soil–plant continuums (rhizosphere, phyllosphere and endosphere) in the maize–wheat/barley rotation system under different fertilization practices at two contrasting sites. Our results indicate that mycobiome assembly is shaped predominantly by compartment niche and host species rather than by environmental factors. Moreover, crop-associated fungal communities are dominated by few abundant taxa mainly belonging to Sordariomycetes and Dothideomycetes, while the majority of diversity within mycobiomes are represented by rare taxa. For plant compartments, the abundant sub-community is mainly determined by stochastic processes. In contrast, the rare sub-community is more sensitive to host selection and mainly governed by deterministic processes. Furthermore, our results demonstrate that rare taxa play an important role in fungal co-occurrence network and ecosystem functioning like crop yield and soil enzyme activities. These results significantly advance our understanding of crop mycobiome assembly and highlight the key role of rare taxa in sustaining the stability of crop mycobiomes and ecosystem functions.     

The factors controlling species density in herbaceous plant communities: an assessment

This paper evaluates both the ideas and empirical evidence pertaining to the control of species density in herbaceous plant communities. While most theoretical discussions of species density have emphasized the importance of habitat productivity and disturbance regimes, many other factors (e.g. species pools, plant litter accumulation, plant morphology) have been proposed to be important. A review of literature presenting observations on the density of species in small plots (in the vicinity of a few square meters or less), as well as experimental studies, suggests several generalizations: (1) Available data are consistent with an underlying unimodal relationship between species density and total community biomass. While variance in species density is often poorly explained by predictor variables, there is strong evidence that high levels of community biomass are antagonistic to high species density. (2) Community biomass is just one of several factors affecting variations in species density. Multivariate analyses typically explain more than twice as much variance in species density as can be explained by community biomass alone. (3) Disturbance has important and sometimes complex effects on species density. In general, the evidence is consistent with the intermediate disturbance hypothesis but exceptions exist and effects can be complex. (4) Gradients in the species pool can have important influences on patterns of species density. Evidence is mounting that a considerable amount of the observed variability in species density within a landscape or region may result from environmental effects on the species pool. (5) Several additional factors deserve greater consideration, including time lags, species composition, plant morphology, plant density and soil microbial effects.Based on the available evidence, a conceptual model of the primary factors controlling species density is presented here. This model suggests that species density is controlled by the effects of disturbance, total community biomass, colonization, the species pool and spatial heterogeneity. The structure of the model leads to two main expectations: (1) while community biomass is important, multivariate approaches will be required to understand patterns of variation in species density, and (2) species density will be more highly correlated with light penetration to the soil surface, than with above-ground biomass, and even less well correlated with plant growth rates (productivity) or habitat fertility. At present, data are insufficient to evaluate the relative importance of the processes controlling species density. Much more work is needed if we are to adequately predict the effects of environmental changes on plant communities and species diversity.     

Do assembly history effects attenuate from species to ecosystem properties? A field test with wood‐inhabiting fungi

Assembly history, or the order of species arrival, can have wide-ranging effects on species, communities and ecosystems. However, it remains unclear whether assembly history primarily affects individual species, with effects attenuating at the level of communities and ecosystems or, alternatively, has consistent effect sizes across increasing levels of ecological organisation. We address this question using a field-based manipulation of assembly history of wood-inhabiting fungi. The largest effect sizes were observed for the frequency of some individual species, and mean effect sizes were lower for community metrics of fungi immigrating from the regional species pool. There was little evidence, however, of attenuation in effect sizes at the ecosystem level (carbon, nitrogen, decomposition) in comparison to the species or community level. These results indicate that assembly history can have strong effects on ecosystem properties even under natural levels of environmental variability.     

Regional adaptation improves the performance of grassland plant communities

Many plant species are adapted locally or regionally. Whether such individual species performance translates into effects at community and ecosystem levels has rarely been tested. Such tests are crucial, however, to predict ecosystem consequences of sowing seed mixtures for grassland restoration or hay production. We compared the performance of replicated sown plant communities of regional origin with the performance of four foreign communities consisting of the same grassland species but originating from distances up to 890 km from our experimental site. The regional communities performed better than foreign communities in plant cover and diversity but not in aboveground biomass production. Additionally, in communities based on regional seeds fewer unsown species occurred and less bare ground was left open for erosion. Variation in community performance among source regions was related to climatic differences rather than to geographic distance to source regions. Individual species performance only partly explained community patterns, highlighting the importance of community level experiments. Our results suggest that the use of regional seeds represents an important approach to improve sown managed grasslands.     

Plant functional traits with particular reference to tropical deciduous forests: a review

Functional traits (FTs) integrate the ecological and evolutionary history of a species, and can potentially be used to predict its response as well as its influence on ecosystem functioning. Study of inter-specific variation in the FTs of plants aids in classifying species into plant functional types (PFTs) and provides insights into fundamental patterns and trade-offs in plant form and functioning and the effect of changing species composition on ecosystem functions. Specifically, this paper focuses on those FTs that make a species successful in the dry tropical environment. Following a brief overview, we discuss plant FTs that may be particularly relevant to tropical deciduous forests (TDFs). We consider the traits under the following categories: leaf traits, stem and root traits, reproductive traits, and traits particularly relevant to water availability. We compile quantitative information on functional traits of dry tropical forest species. We also discuss trait-based grouping of plants into PFTs. We recognize that there is incomplete knowledge about many FTs and their effects on TDFs and point out the need for further research on PFTs of TDF species, which can enable prediction of the dynamics of these forests in the face of disturbance and global climate change. Correlations between structural and ecophysiological traits and ecosystem functioning should also be established which could make it possible to generate predictions of changes in ecosystem services from changes in functional composition.     

Spatial patterns of plant diversity below-ground as revealed by DNA barcoding

Our understanding of the spatial organization of root diversity in plant communities and of the mechanisms of community assembly has been limited by our ability to identify plants based on root tissue, especially in diverse communities. Here, we test the effectiveness of the plastid gene rbcL, a core plant DNA barcoding marker, for investigating spatial patterns of root diversity, and relate observed patterns to above-ground community structure. We collected 3800 root fragments from four randomly positioned, 1-m-deep soil profiles (two vertical transects per plot), located in an old-field community in southern Ontario, Canada, and extracted and sequenced DNA from 1531 subsampled fragments. We identified species by comparing sequences with a DNA barcode reference library developed previously for the local flora. Nearly 85% of sampled root fragments were successfully sequenced and identified as belonging to 29 plant species or species groups. Root abundance and species richness varied in horizontal space and were negatively correlated with soil depth. The relative abundance of taxa below-ground was correlated with their frequency above-ground (r = 0.73, P = 0.0001), but several species detected in root tissue were not observed in above-ground quadrats. Multivariate analyses indicated that diversity was highly structured below-ground, and associated with depth, root morphology, soil chemistry and soil texture, whereas little structure was evident above-ground. Furthermore, analyses of species co-occurrence indicates strong species segregation overall but random co-occurrence among confamilials. Our results provide insights into the role of environmental filtering and competitive interactions in the organization of plant diversity below-ground, and also demonstrate the utility of barcoding for the identification of plant roots.     

Experimental whole‐stream warming alters community size structure

How ecological communities respond to predicted increases in temperature will determine the extent to which Earth's biodiversity and ecosystem functioning can be maintained into a warmer future. Warming is predicted to alter the structure of natural communities, but robust tests of such predictions require appropriate large‐scale manipulations of intact, natural habitat that is open to dispersal processes via exchange with regional species pools. Here, we report results of a two‐year whole‐stream warming experiment that shifted invertebrate assemblage structure via unanticipated mechanisms, while still conforming to community‐level metabolic theory. While warming by 3.8 °C decreased invertebrate abundance in the experimental stream by 60% relative to a reference stream, total invertebrate biomass was unchanged. Associated shifts in invertebrate assemblage structure were driven by the arrival of new taxa and a higher proportion of large, warm‐adapted species (i.e., snails and predatory dipterans) relative to small‐bodied, cold‐adapted taxa (e.g., chironomids and oligochaetes). Experimental warming consequently shifted assemblage size spectra in ways that were unexpected, but consistent with thermal optima of taxa in the regional species pool. Higher temperatures increased community‐level energy demand, which was presumably satisfied by higher primary production after warming. Our experiment demonstrates how warming reassembles communities within the constraints of energy supply via regional exchange of species that differ in thermal physiological traits. Similar responses will likely mediate impacts of anthropogenic warming on biodiversity and ecosystem function across all ecological communities.     

Identifying qualitative effects of different grazing types on below‐ground communities and function in a long‐term field experiment

Herbivory is an important modulator of plant biodiversity and productivity in grasslands, but our understanding of herbivore‐induced changes on below‐ground processes and communities is limited. Using a long‐term (17 years) experimental site, we evaluated impacts of rabbit and invertebrate grazers on some soil functions involved in carbon cycling, microbial diversity, structure and functional composition. Both rabbit and invertebrate grazing impacted soil functions and microbial community structure. All functional community measures (functions, biogeochemical cycling genes, network association between different taxa) were more strongly affected by invertebrate grazers than rabbits. Furthermore, our results suggest that exclusion of invertebrate grazers decreases both microbial biomass and abundance of genes associated with key biogeochemical cycles, and could thus have long‐term consequences for ecosystem functions. The mechanism behind these impacts are likely to be driven by both direct effects of grazing altering the pattern of nutrient inputs and by indirect effects through changes in plant species composition. However, we could not entirely discount that the pesticide used to exclude invertebrates may have affected some microbial community measures. Nevertheless, our work illustrates that human activity that affects grazing intensity may affect ecosystem functioning and sustainability, as regulated by multi‐trophic interactions between above‐ and below‐ground communities.     

Complexity of multitrophic interactions in a grassland ecosystem depends on plant species diversity

1.?We studied the theoretical prediction that a loss of plant species richness has a strong impact on community interactions among all trophic levels and tested whether decreased plant species diversity results in a less complex structure and reduced interactions in ecological networks. 2.?Using plant species-specific biomass and arthropod abundance data from experimental grassland plots (Jena Experiment), we constructed multitrophic functional group interaction webs to compare communities based on 4 and 16 plant species. 427 insect and spider species were classified into 13 functional groups. These functional groups represent the nodes of ecological networks. Direct and indirect interactions among them were assessed using partial Mantel tests. Interaction web complexity was quantified using three measures of network structure: connectance, interaction diversity and interaction strength. 3.?Compared with high plant diversity plots, interaction webs based on low plant diversity plots showed reduced complexity in terms of total connectance, interaction diversity and mean interaction strength. Plant diversity effects obviously cascade up the food web and modify interactions across all trophic levels. The strongest effects occurred in interactions between adjacent trophic levels (i.e. predominantly trophic interactions), while significant interactions among plant and carnivore functional groups, as well as horizontal interactions (i.e. interactions between functional groups of the same trophic level), showed rather inconsistent responses and were generally rarer. 4.?Reduced interaction diversity has the potential to decrease and destabilize ecosystem processes. Therefore, we conclude that the loss of basal producer species leads to more simple structured, less and more loosely connected species assemblages, which in turn are very likely to decrease ecosystem functioning, community robustness and tolerance to disturbance. Our results suggest that the functioning of the entire ecological community is critically linked to the diversity of its component plants species.     

优势种去除对崇明东滩盐沼湿地生态系统的影响

盐沼生态系统环境梯度明显,物种组成较简单,是研究生物多样性与生态系统功能关系的理想对象。本研究以崇明东滩盐沼湿地为研究区域,研究优势种去除对植物群落结构以及底栖动物群落的影响。结果表明:(1)去除处理仅对植物群落分株密度有极显著效应(P0.01)。去除组和对照组物种组成差异随时间增加而减小,处理效应逐渐减弱。(2)去除组底栖动物密度均低于对照组,但差异不显著。(3)盐沼植物群落特征与底栖动物群落有密切关系,植物密度、冠层高度与底栖动物密度相关性极显著。去除优势种后,植物群落分株密度升高,群落内剩余物种占比有所上升,次优势种对群落的补偿效应具有较大贡献;而底栖动物群落密度下降,其生物量和多样性指数的变化趋势与密度并不一致。上述结果表明生物多样性变化影响了盐沼湿地生态系统植物群落和底栖动物群落结构,进而可能影响物质循环和能量流动过程。     

Restoration of Prairie Community Structure and Ecosystem Function in an Abandoned Hayfield: A Sowing Experiment

Using a multispecies seed sowing experiment, we investigated the roles of seed and microsite limitation in constraining the restoration of native prairie diversity and ecosystem function in an abandoned upland hayfield in northeastern Kansas. Seeds of 32 native and naturalized plant species from the regional pool were sown into undisturbed and experimentally disturbed field plots. After six growing seasons, experimental sowing led to major shifts in species and functional group composition, increases in native species abundance and floristic quality, declines in abundance of non‐native species, and increases in plant diversity. These changes in community structure led to significant changes at the ecosystem level including increases in light capture, peak biomass, primary production, litter biomass, root biomass, and C storage in roots. Our findings reveal the importance of seed limitations in constraining the natural recovery of prairie vegetation, biodiversity, and ecosystem functioning in this grassland and confirm broadcast sowing as a useful tool for the restoration of upland hayfield sites.     

养分添加对天山高寒草地植物多样性和地上生物量的影响

为揭示高寒草地物种多样性和地上生物量以及二者之间关系对养分添加的响应模式,该研究以天山高寒草地为对象,通过两年的多重养分添加实验,研究氮(N)、磷(P)、钾(K)3种养分单独和组合添加对天山高寒草地群落物种多样性和地上生物量的影响。结果表明:(1)养分添加使当地植物物种多样性不同程度地减少,其中以N+P、N+K、N+P+K添加的效应最为显著,多重养分添加导致的土壤生态位维度降低是当地物种丧失的重要原因。(2)养分添加能显著提高群落地上生物量,其中N为第一限制养分,解除N限制后P和K成为限制养分, N+P+K复合添加对地上生物量的提高最为显著。(3)养分添加两年后,地上生物量与物种丰富度之间无显著回归关系且地上生物量增加主要是由于禾草类生物量增加导致,说明地上生物量主要由少数优势种决定而非群落物种数。