Cyanobacteria and chloroflexi-dominated hypolithic colonization of quartz at the hyper-arid core of the Atacama Desert,Chile

Quartz stones are ubiquitous in deserts and are a substrate for hypoliths, microbial colonists of the underside of such stones. These hypoliths thrive where extreme temperature and moisture stress limit the occurrence of higher plant and animal life. Several studies have reported the occurrence of green hypolithic colonization dominated by cyanobacteria. Here, we describe a novel red hypolithic colonization from Yungay, at the hyper-arid core of the Atacama Desert in Chile. Comparative analysis of green and red hypoliths from this site revealed markedly different microbial community structure as revealed by 16S rRNA gene clone libraries. Green hypoliths were dominated by cyanobacteria (Chroococcidiopsis and Nostocales phylotypes), whilst the red hypolith was dominated by a taxonomically diverse group of chloroflexi. Heterotrophic phylotypes common to all hypoliths were affiliated largely to desiccation-tolerant taxa within the Actinobacteria and Deinococci. Alphaproteobacterial phylotypes that affiliated with nitrogen-fixing taxa were unique to green hypoliths, whilst Gemmatimonadetes phylotypes occurred only on red hypolithon. Other heterotrophic phyla recovered with very low frequency were assumed to represent functionally relatively unimportant taxa.     

The spatial structures of hypolithic communities in the Dry Valleys of East Antarctica

Hypolithic communities represent important reservoirs of microbial life in hyper-arid deserts. A number of studies on the diversity and ecology of these communities from different geographic areas have been reported in the past decade, but the spatial distribution of the different components of these communities is still not understood. Moss- and cyanobacteria-dominated hypolithic community morphotypes from Miers Valley (McMurdo Dry Valleys, East Antarctica) were analyzed by electron microscopy in order to characterize the microscale spatial structure. The two communities showed a high degree of internal organization, but differing according to the biological composition. In moss-dominated hypoliths, the moss plantlets are intermixed with mineral fragments of soil origin. However, in cyanobacteria-dominated hypoliths, a layered spatial organization was structured by filamentous cyanobacteria and associated extracellular polymeric components. While moss cells were lacking in cyanobacteria-dominated communities, biofilms formed by cyanobacteria and heterotrophic bacteria were observed in both community morphotypes. The water-holding capacity of both live and dead moss cells and the associated organic matrix, together with the protective properties of the extracellular polymeric substances, could facilitate the survival and activity of these communities. Similar structural strategies can favor the survival of microbial communities in different extreme environments.     

Hypolithic microbial communities: between a rock and a hard place

Drylands are the largest terrestrial biome on Earth and a ubiquitous feature is desert pavement terrain, comprising rocks embedded in the mineral soil surface. Quartz and other translucent rocks are common and microbial communities termed hypoliths develop as biofilms on their ventral surfaces. In extreme deserts these represent major concentrations of biomass, and are emerging as key to geobiological processes and soil stabilization. These highly specialized communities are dominated by cyanobacteria that support diverse heterotrophic assemblages. Here we identify global-scale trends in the ecology of hypoliths that are strongly related to climate, particularly with regard to shifts in cyanobacterial assemblages. A synthesis of available data revealed a linear trend for colonization with regard to climate, and we suggest potential application for hypoliths as 'biomarkers' of aridity on a landscape scale. The potential to exploit the soil-stabilizing properties of hypolithic colonization in environmental engineering on dryland soils is also discussed.     

Hypolithic microbial communities of quartz rocks from Miers Valley, McMurdo Dry Valleys, Antarctica

The McMurdo Dry Valleys region of eastern Antarctica is a cold desert that presents extreme challenges to life. Hypolithic microbial colonisation of the subsoil surfaces of translucent quartz rocks represent a significant source of terrestrial biomass and productivity in this region. Previous studies have described hypoliths as dominated by cyanobacteria. However, hypoliths that occur in the lower Dry Valleys such as the Miers, Garwood and Marshall Valleys are unusual as they are not necessarily cyanobacteria-dominated. These hypoliths support significant eukaryal colonisation by fungi and mosses in addition to cyanobacteria-dominated bacterial assemblages and so have considerable ecological value in this barren landscape. Here, we characterise these novel hypoliths by analysis of environmental rRNA gene sequences. The hypolithic community was demonstrated to be distinct from the surrounding soil and non-translucent rocks. Hypoliths supported cyanobacterial signatures from the Oscillatoriales and Nostocales. Other heterotrophic bacterial signatures were also recovered, and these were phylogenetically diverse and spanned 8 other bacterial phyla. Archaeal phylotypes recovered were phylogenetically affiliated with the large group of unclassified, uncultured Crenarcheota. Eukaryal phylotypes indicated that free-living ascomycetous fungi, chlorophytes and mosses (Bryum sp.) were all supported by these hypoliths, and these are thought to be responsible for the extensive eukaryotic biomass that develops around quartz rocks.     

Local factors control the community composition of cyanobacteria in lakes while heterotrophic bacteria follow a neutral model

1. Neutral community models are derived from the proposition that basic probabilities of species loss (extinction, emigration) and gain (immigration, speciation) explain biological community structure, such that species with many individuals are very likely to be widespread. Niche models on the other hand assume that interactions between species and differential resource use mediate species coexistence, thus invoking environmental factors to explain community patterns. 2. In this study, we compared neutral and niche models to test how much of the spatial variability of assemblages of heterotrophic bacteria and phytoplankton in 13 lakes they could explain. Analysis of phytoplankton was restricted to cyanobacteria, so that they could be studied with the same molecular fingerprinting method, automated ribosomal intergenic spaces analysis (ARISA), as heterotrophic bacteria. We determined local biotic and abiotic lake variables as well as lake age, glacial history and distance between sites. 3. The neutral community model had a good fit to the community composition of heterotrophic bacteria (R² = 0.69), whereas it could not produce a significant model for the community composition of cyanobacteria. 4. The community composition of cyanobacteria was instead correlated to environmental variables. The best model, a combination of total organic carbon, biomass of eukaryotic phytoplankton, pH and conductivity, could explain 8% of the variation. In contrast, variation in the community composition of heterotrophic bacteria was not predicted by any of the environmental variables. Historical and spatial variables were not correlated to the community composition of either group. 5. The pattern found for heterotrophic bacteria suggests that stochastic processes are important. The correlation of cyanobacteria with local environmental variables alone is consistent with the niche model. We suggest that cyanobacteria, a group of organisms containing bloom‐forming species, may be less likely to fit a neutral community model, since these blooms are usually triggered by a particular combination of environmental conditions.     

中央戈壁石下生物土壤结皮固氮细菌群落结构和多样性

【背景】戈壁荒漠石下生物土壤结皮(Biological Soil Crusts,BSCs)由石下生物定殖繁衍而成,广泛存在于石英石下方,在相关生态系统的物质循环中起重要作用;其细菌群落结构受空间和土壤环境因子影响而变化较大。固氮细菌是石下BSCs的形成和发育主要驱动力;中央戈壁面积较大,为温带的代表戈壁之一,但目前其石下BSCs中固氮细菌群落结构和多样性尚未有研究报道。【目的】阐释中央戈壁石下BSCs中固氮细菌的群落结构、多样性及其影响因素。【方法】应用Mi Seq对nif H基因进行高通量测序,并使用生物信息学方法基于nif H序列分析固氮细菌的群落结构和多样性及其影响因素。使用CoNet软件绘制物种共现性图,以期揭示石下固氮细菌群落结构的关键物种。【结果】石下固氮细菌的优势菌门有Cyanobacteria (47.20%-69.90%)和Proteobacteria (27.47%-48.91%);优势菌属为Scytonema (45.05%-69.09%)、Skermanella (10.26%-20.48%)和未知属(13.72%-22.00%);9月份物种丰富度较5月份高,但这2个月份的多样性无明显差异;在土壤理化因子中,速效氮对石下固氮细菌群落组成的影响最大;其中各微生物之间均存在较强的互作关系,以共存关系为主(约占66.98%),点度中心性、接近中心性和中介中心性均较高的节点属于Alphaproteobacteria。【结论】中央戈壁石下BSCs中固氮细菌以Cyanobacteria和Proteobacteria为最优势菌群;Alphaproteobacteria为稳定石下BSCs固氮细菌群落的关键类群,可能是主要固氮者,这为认识和利用石下生物土壤结皮固氮细菌提供了基础依据。     

Hypolithic Cyanobacteria Supported Mainly by Fog in the Coastal Range of the Atacama Desert

The Atacama Desert is one of the driest places on Earth, with an arid core highly adverse to the development of hypolithic cyanobacteria. Previous work has shown that when rain levels fall below ~1 mm per year, colonization of suitable quartz stones falls to virtually zero. Here, we report that along the coast in these arid regions, complex associations of cyanobacteria, archaea, and heterotrophic bacteria inhabit the undersides of translucent quartz stones. Colonization rates in these areas, which receive virtually no rain but mainly fog, are significantly higher than those reported inland in the hyperarid zone at the same latitude. Here, hypolithic colonization rates can be up to 80%, with all quartz rocks over 20 g being colonized. This finding strongly suggests that hypolithic microbial communities thriving in the seaward face of the Coastal Range can survive with fog as the main regular source of moisture. A model is advanced where the development of the hypolithic communities under quartz stones relies on a positive feedback between fog availability and the higher thermal conductivity of the quartz rocks, which results in lower daytime temperatures at the quartz–soil interface microenvironment.     

Ecosystem productivity is associated with bacterial phylogenetic distance in surface marine waters

Understanding the link between community diversity and ecosystem function is a fundamental aspect of ecology. Systematic losses in biodiversity are widely acknowledged but the impact this may exert on ecosystem functioning remains ambiguous. There is growing evidence of a positive relationship between species richness and ecosystem productivity for terrestrial macro‐organisms, but similar links for marine micro‐organisms, which help drive global climate, are unclear. Community manipulation experiments show both positive and negative relationships for microbes. These previous studies rely, however, on artificial communities and any links between the full diversity of active bacterial communities in the environment, their phylogenetic relatedness and ecosystem function remain hitherto unexplored. Here, we test the hypothesis that productivity is associated with diversity in the metabolically active fraction of microbial communities. We show in natural assemblages of active bacteria that communities containing more distantly related members were associated with higher bacterial production. The positive phylogenetic diversity–productivity relationship was independent of community diversity calculated as the Shannon index. From our long‐term (7‐year) survey of surface marine bacterial communities, we also found that similarly, productive communities had greater phylogenetic similarity to each other, further suggesting that the traits of active bacteria are an important predictor of ecosystem productivity. Our findings demonstrate that the evolutionary history of the active fraction of a microbial community is critical for understanding their role in ecosystem functioning.     

Disentangling the ‘brown world’ faecal–detritus interaction web: dung beetle effects on soil microbial properties

Many ecosystem services are sustained by the combined action of microscopic and macroscopic organisms, and shaped by interactions between the two. However, studies tend to focus on only one of these two components. We combined the two by investigating the impact of macrofauna on microbial community composition and functioning in the context of a major ecosystem process: the decomposition of dung. We compared bacterial communities of pasture soil and experimental dung pats inhabited by one (Aphodius), two (Aphodius and Geotrupes), or no dung beetle genera. Overall, we found distinct microbial communities in soil and dung samples, and that the communities converged over the course of the experiment. Characterising the soil microbial communities underlying the dung pats revealed a significant interactive effect between the microflora and macrofauna, where the diversity and composition of microbial communities was significantly affected by the presence or absence of dung beetles. The specific identity of the beetles had no detectable impact, but the microbial evenness was lower in the presence of both Aphodius and Geotrupes than in the presence of Aphodius alone. Differences in microbial community composition were associated with differences in substrate usage as measured by Ecoplates. Moreover, microbial communities with similar compositions showed more similar substrate usage. Our study suggests that the presence of macrofauna (dung beetles) will modify the microflora (bacteria) of both dung pats and pasture soil, including community diversity and functioning. In particular, the presence of dung beetles promotes the transfer of bacteria across the soil–dung interface, resulting in increased similarity in community structure and functioning. The results demonstrate that to understand how microbes contribute to the ecosystem process of dung decomposition, there is a need to understand their interactions with larger co‐occurring fauna.     

Microclimate and limits to photosynthesis in a diverse community of hypolithic cyanobacteria in northern Australia

Hypolithic microbes, primarily cyanobacteria, inhabit the highly specialized microhabitats under translucent rocks in extreme environments. Here we report findings from hypolithic cyanobacteria found under three types of translucent rocks (quartz, prehnite, agate) in a semiarid region of tropical Australia. We investigated the photosynthetic responses of the cyanobacterial communities to light, temperature and moisture in the laboratory, and we measured the microclimatic variables of temperature and soil moisture under rocks in the field over an annual cycle. We also used molecular techniques to explore the diversity of hypolithic cyanobacteria in this community and their phylogenetic relationships within the context of hypolithic cyanobacteria from other continents. Based on the laboratory experiments, photosynthetic activity required a minimum soil moisture of 15% (by mass). Peak photosynthetic activity occurred between approximately 8°C and 42°C, though some photosynthesis occurred between ?1°C and 51°C. Maximum photosynthesis rates also occurred at light levels of approximately 150–550 μmol m^?2 s^?1. We used the field microclimatic data in conjunction with these measurements of photosynthetic efficiency to estimate the amount of time the hypolithic cyanobacteria could be photosynthetically active in the field. Based on these data, we estimated that conditions were appropriate for photosynthetic activity for approximately 942 h (～75 days) during the year. The hypolithic cyanobacteria community under quartz, prehnite and agate rocks was quite diverse both within and between rock types. We identified 115 operational taxonomic units (OTUs), with each rock hosting 8–24 OTUs. A third of the cyanobacteria OTUs from northern Australia grouped with Chroococcidiopsis, a genus that has been identified from hypolithic and endolithic communities from the Gobi, Mojave, Atacama and Antarctic deserts. Several OTUs identified from northern Australia have not been reported to be associated with hypolithic communities previously.     

Evidence for successional development in Antarctic hypolithic bacterial communities

Hypoliths (cryptic microbial assemblages that develop on the undersides of translucent rocks) are significant contributors to regional C and N budgets in both hot and cold deserts. Previous studies in the Dry Valleys of Eastern Antarctica have reported three morphologically distinct hypolithic community types: cyanobacteria dominated (type I), fungus dominated (type II) and moss dominated (type III). Here we present terminal-restriction fragment length polymorphism analyses to elucidate the bacterial community structure in hypolithons and the surrounding soils. We show clear and robust distinction in bacterial composition between bulk surface soils and hypolithons. Moreover, the bacterial assemblages were similar in types II and III hypolithons and clearly distinct from those found in type I. Through 16S rRNA gene 454 pyrosequencing, we show that Proteobacteria dominated all three types of hypolithic communities. As expected, Cyanobacteria were more abundant in type I hypolithons, whereas Actinobacteria were relatively more abundant in types II and III hypolithons, and were the dominant group in soils. Using a probabilistic dissimilarity metric and random sampling, we demonstrate that deterministic processes are more important in shaping the structure of the bacterial community found in types II and III hypolithons. Most notably, the data presented in this study suggest that hypolithic bacterial communities establish via a successional model, with the type I hypolithons acting as the basal development state.     

Structure of microbial communities in <Emphasis Type="Italic">Sphagnum</Emphasis> peatlands and effect of atmospheric carbon dioxide enrichment

Little is known about the structure of microbial communities in Sphagnum peatlands, and the potential effects of the increasing atmospheric CO2 concentration on these communities are not known. We analyzed the structure of microbial communities in five Sphagnum-dominated peatlands across Europe and their response to CO2 enrichment using miniFACE systems. After three growing seasons, Sphagnum samples were analyzed for heterotrophic bacteria, cyanobacteria, microalgae, heterotrophic flagellates, ciliates, testate amoebae, fungi, nematodes, and rotifers. Heterotrophic organisms dominated the microbial communities and together represented 78% to 97% of the total microbial biomass. Testate amoebae dominated the protozoan biomass. A canonical correspondence analysis revealed a significant correlation between the microbial community data and four environmental variables (Na+, DOC, water table depth, and DIN), reflecting continentality, hydrology, and nitrogen deposition gradients. Carbon dioxide enrichment modified the structure of microbial communities, but total microbial biomass was unaffected. The biomass of heterotrophic bacteria increased by 48%, and the biomass of testate amoebae decreased by 13%. These results contrast with the absence of overall effect on methane production or on the vegetation, but are in line with an increased below-ground vascular plant biomass at the same sites. We interpret the increase in bacterial biomass as a response to a CO2-induced enhancement of Sphagnum exudation. The causes for the decrease of testate amoebae are unclear but could indicate a top-down rather than a bottom-up control on their density.     

Genotypic richness and dissimilarity opposingly affect ecosystem functioning

Biodiversity is an essential determinant of ecosystem functioning. Numerous studies described positive effects of diversity on the functioning of communities arising from complementary resource use and facilitation. However, high biodiversity may also increase competitive interactions, fostering antagonism and negatively affecting community performance. Using experimental bacterial communities we differentiated diversity effects based on genotypic richness and dissimilarity. We show that these diversity characteristics have opposite effects on ecosystem functioning. Genotypic dissimilarity governed complementary resource use, improving ecosystem functioning in complex resource environments. Contrastingly, genotypic richness drove allelopathic interactions, mostly reducing ecosystem functioning. The net biodiversity effect on community performance resulted from the interplay between the genetic structure of the community and resource complexity. These results demonstrate that increasing richness, without concomitantly increasing dissimilarity, can decrease ecosystem functioning in simple environments due to antagonistic interactions, an effect insufficiently considered so far in mechanistic models of the biodiversity-ecosystem functioning relationship.     

Species traits as drivers of food web structure

The use of functional traits to describe community structure is a promising approach to reveal generalities across organisms and ecosystems. Plant ecologists have demonstrated the importance of traits in explaining community structure, competitive interactions as well as ecosystem functioning. The application of trait‐based methods to more complex communities such as food webs is however more challenging owing to the diversity of animal characteristics and of interactions. The objective of this study was to determine how functional structure is related to food web structure. We consider that food web structure is the result of 1) the match between consumer and resource traits, which determine the occurence of a trophic interaction between them, and 2) the distribution of functional traits in the community. We implemented a statistical approach to assess whether or not 35 466 pairwise interactions between soil organisms are constrained by trait‐matching and then used a Procrustes analysis to investigate correlations between functional indices and network properties across 48 sites. We found that the occurrence of trophic interactions is well predicted by matching the traits of the resource with those of the consumer. Taxonomy and body mass of both species were the most important traits for the determination of an interaction. As a consequence, functional evenness and the variance of certain traits in the community were correlated to trophic complementarity between species, while trait identity, more than diversity, was related to network topology. The analysis was however limited by trait data availability, and a coarse resolution of certain taxonomic groups in our dataset. These limitations explain the importance of taxonomy, as well as the complexity of the statistical model needed. Our results outline the important implications of trait composition on ecological networks, opening promising avenues of research into the relationship between functional diversity and ecosystem functioning in multi‐trophic systems.     

Resource availability influences the diversity of a functional group of heterotrophic soil bacteria

Resource availability is a key factor regulating biodiversity and ecosystem functioning, but the relationship between resource availability and diversity has only been rarely investigated in microbial communities. The aim of this study was to determine how diversity and community structure of a functional group of soil bacteria are influenced by resource concentration. To achieve this, we used soil microcosms to investigate degradation of benzoate, which served as a model compound, by soil bacterial communities. Microcosms were supplied with ¹³C-labelled benzoate at four concentrations and RNA-stable isotope probing followed by molecular fingerprinting analysis of 16S rRNA genes was employed to identify bacteria able to assimilate benzoate at different concentrations. The composition of the benzoate degrader community differed at different concentrations and there was a significant decrease in taxa evenness at the highest substrate concentration. Active organisms could be grouped into generalists, occurring at all substrate concentrations, specialists, active at one particular benzoate concentration only, and taxa that were active at either the two lowest or two highest concentrations. The study comprises the first explicit demonstration that resource availability has an effect on the diversity of a functional group of heterotrophic soil bacteria.     

土壤细菌网络互作调控线虫肠道细菌群落

动物肠道细菌群落在联系宿主与生态系统功能中发挥着至关重要的作用。【目的】本研究旨在评估绿肥翻压和水稻生长不同时期对土壤细菌和线虫肠道细菌群落组成和结构的影响,并探究土壤细菌和线虫肠道细菌群落间的潜在关联关系。【方法】基于盆栽试验,结合16S rRNA基因高通量测序技术,分析黑麦草翻压和对照处理下水稻生长的前期(返青期)和后期(收获期)土壤细菌和线虫肠道细菌群落,结合网络分析研究土壤细菌网络互作对线虫肠道细菌群落的潜在影响。【结果】黑麦草翻压对土壤细菌和线虫肠道细菌群落组成和结构没有显著影响(P>0.05);水稻生长后期样品比前期样品具有更高的α多样性。基于随机森林机器学习法获得的土壤细菌和线虫肠道细菌生物标志物之间存在广泛的显著相关关系,为土壤细菌群落变化调控线虫肠道细菌群落组成提供了有力的证据。共现网络分析表明土壤细菌之间的正相互作用显著促进了土壤细菌和线虫肠道细菌之间的正相互作用(P<0.01),进而影响了线虫肠道细菌之间的网络互作。结构方程模型进一步表明土壤养分含量的降低主要通过降低土壤细菌之间正相互作用,从而间接影响线虫肠道细菌之间的互作。【结论】土壤细菌互作可能在...     

Structural Diversity of Bacterial Communities Associated with Bloom-Forming Freshwater Cyanobacteria Differs According to the Cyanobacterial Genus

The factors and processes driving cyanobacterial blooms in eutrophic freshwater ecosystems have been extensively studied in the past decade. A growing number of these studies concern the direct or indirect interactions between cyanobacteria and heterotrophic bacteria. The presence of bacteria that are directly attached or immediately adjacent to cyanobacterial cells suggests that intense nutrient exchanges occur between these microorganisms. In order to determine if there is a specific association between cyanobacteria and bacteria, we compared the bacterial community composition during two cyanobacteria blooms of Anabaena (filamentous and N₂-fixing) and Microcystis (colonial and non-N₂ fixing) that occurred successively within the same lake. Using high-throughput sequencing, we revealed a clear distinction between associated and free-living communities and between cyanobacterial genera. The interactions between cyanobacteria and bacteria appeared to be based on dissolved organic matter degradation and on N recycling, both for N₂-fixing and non N₂-fixing cyanobacteria. Thus, the genus and potentially the species of cyanobacteria and its metabolic capacities appeared to select for the bacterial community in the phycosphere.     

Evidence of microbial rhodopsins in Antarctic Dry Valley edaphic systems

Microorganisms able to synthesize rhodopsins have the capacity to translocate ions through their membranes, using solar energy to generate a proton motive force. Rhodopsins are the most abundant phototrophic proteins in oceanic surface waters and are key constituents in marine bacterial ecology. However, it remains unclear how rhodopsins are used in most microorganisms. Despite their abundance in marine and fresh‐water systems, the presence of functional rhodopsin systems in edaphic habitats has never been reported. Here, we show the presence of several new putative H⁺, Na⁺ and Cl⁺ pumping rhodopsins identified by metagenomic analysis of Antarctic desert hypolithic communities. Reconstruction of two Proteobacteria genomes harboring xanthorhodopsin‐like proteins and one Bacteroidetes genome with a Na‐pumping‐like rhodopsin indicated that these bacteria were aerobic heterotrophs possessing the apparent capacity for the functional expression of rhodopsins. The existence of these protein systems in hypolithic bacteria expands the known role of rhodopsins to include terrestrial environments and suggests a possible predominant function as heterotrophic energy supply proteins, a feasible microbial adaptation to the harsh conditions prevalent in Antarctic edaphic systems.     

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.