Elevated temperature may reduce functional but not taxonomic diversity of fungal assemblages on decomposing leaf litter in streams

Mounting evidence points to a linkage between biodiversity and ecosystem functioning (B-EF). Global drivers, such as warming and nutrient enrichment, can alter species richness and composition of aquatic fungal assemblages associated with leaf-litter decomposition, a key ecosystem process in headwater streams. However, effects of biodiversity changes on ecosystem functions might be countered by the presumed high functional redundancy of fungal species. Here, we examined how environmental variables and leaf-litter traits (based on leaf chemistry) affect taxonomic and functional α- and β-diversity of fungal decomposers. We analysed taxonomic diversity (DNA-fingerprinting profiles) and functional diversity (community-level physiological profiles) of fungal communities in four leaf-litter species from four subregions differing in stream-water characteristics and riparian vegetation. We hypothesized that increasing stream-water temperature and nutrients would alter taxonomic diversity more than functional diversity due to the functional redundancy among aquatic fungi. Contrary to our expectations, fungal taxonomic diversity varied little with stream-water characteristics across subregions, and instead taxon replacement occurred. Overall taxonomic β-diversity was fourfold higher than functional diversity, suggesting a high degree of functional redundancy among aquatic fungi. Elevated temperature appeared to boost assemblage uniqueness by increasing β-diversity while the increase in nutrient concentrations appeared to homogenize fungal assemblages. Functional richness showed a negative relationship with temperature. Nonetheless, a positive relationship between leaf-litter decomposition and functional richness suggests higher carbon use efficiency of fungal communities in cold waters.     

Plant functional diversity enhances associations of soil fungal diversity with vegetation and soil in the restoration of semiarid sandy grassland

The trait‐based approach shows that plant functional diversity strongly affects ecosystem properties. However, few empirical studies show the relationship between soil fungal diversity and plant functional diversity in natural ecosystems. We investigated soil fungal diversity along a restoration gradient of sandy grassland (mobile dune, semifixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China, using the denaturing gradient gel electrophoresis of 18S rRNA and gene sequencing. We also examined associations of soil fungal diversity with plant functional diversity reflected by the dominant species' traits in community (community‐weighted mean, CWM) and the dispersion of functional trait values (FD_is). We further used the structure equation model (SEM) to evaluate how plant richness, biomass, functional diversity, and soil properties affect soil fungal diversity in sandy grassland restoration. Soil fungal richness in mobile dune and semifixed dune was markedly lower than those of fixed dune and grassland (P < 0.05). Soil fungal richness was positively associated with plant richness, biomass, CWM plant height, and soil gradient aggregated from the principal component analysis, but SEM results showed that plant richness and CWM plant height determined by soil properties were the main factors exerting direct effects. Soil gradient increased fungal richness through indirect effect on vegetation rather than direct effect. The negative indirect effect of FDis on soil fungal richness was through its effect on plant biomass. Our final SEM model based on plant functional diversity explained nearly 70% variances of soil fungal richness. Strong association of soil fungal richness with the dominant species in the community supported the mass ratio hypothesis. Our results clearly highlight the role of plant functional diversity in enhancing associations of soil fungal diversity with community structure and soil properties in sandy grassland ecosystems.     

农业生态系统中的AM真菌多样性

农业生态系统复杂庞大,是由如麦田生态系统、水稻田生态系统、果园生态系统、草地生态系统、保护地生态系统等组成的一个复合生态系统。重点介绍农业生态系统中丛枝菌根(AM)和AM真菌多样性,探讨农业生态系统中调控AM真菌多样性的途径以及今后研究的动向。     

Soil fertility shifts the relative importance of saprotrophic and mycorrhizal fungi for maintaining ecosystem stability

Soil microbial communities are essential for regulating the dynamics of plant productivity. However, how soil microbes mediate temporal stability of plant productivity at large scales across various soil fertility conditions remains unclear. Here, we combined a regional survey of 51 sites in the temperate grasslands of northern China with a global grassland survey of 120 sites to assess the potential roles of soil microbial diversity in regulating ecosystem stability. The temporal stability of plant productivity was quantified as the ratio of the mean normalized difference vegetation index to its standard deviation. Soil fungal diversity, but not bacterial diversity, was positively associated with ecosystem stability, and particular fungal functional groups determined ecosystem stability under contrasting conditions of soil fertility. The richness of soil fungal saprobes was positively correlated with ecosystem stability under high-fertility conditions, while a positive relationship was observed with the richness of mycorrhizal fungi under low-fertility conditions. These relationships were maintained after accounting for plant diversity and environmental factors. Our findings highlight the essential role of fungal diversity in maintaining stable grassland productivity, and suggest that future studies incorporating fungal functional groups into biodiversity–stability relationships will advance our understanding of their linkages under different fertility conditions.     

Belowground biodiversity effects of plant symbionts support aboveground productivity

Soil microbes play key roles in ecosystems, yet the impact of their diversity on plant communities is still poorly understood. Here we demonstrate that the diversity of belowground plant-associated soil fungi promotes plant productivity and plant coexistence. Using additive partitioning of biodiversity effects developed in plant biodiversity studies, we demonstrate that this positive relationship can be driven by complementarity effects among soil fungi in one soil type and by a selection effect resulting from the fungal species that stimulated plant productivity the most in another soil type. Selection and complementarity effects among fungal species contributed to improving plant productivity up to 82% and 85%, respectively, above the average of the respective fungal species monocultures depending on the soil in which they were grown. These results also indicate that belowground diversity may act as insurance for maintaining plant productivity under differing environmental conditions.     

Functional complementarity in the arbuscular mycorrhizal symbiosis

The causes and consequences of biodiversity are central themes in ecology. Perhaps one reason for much of the current interest in biodiversity is the belief that the loss of species (by extinction) or their gain (by invasion) will significantly influence ecosystem function. Arbuscular mycorrhizal (AM) fungi are components of most terrestrial ecosystems and, while many research programs have shown that variability among species or isolates of AM fungi does occur (Giovannetti & Gianinazzi-Pearson, 1994), the basis for this variability and its consequences to the function of communities and ecosystems remains largely unexplored. Smith et al . (pp. 357–366 in this issue) now show clearly that ecologically significant functional diversity exists among AM fungal species in the regions of the soil from which they absorb phosphate, and their results suggest that such diversity may have significant ecological consequences.     

Plant community composition, not diversity, regulates soil respiration in grasslands

Soil respiration is responsible for recycling considerable quantities of carbon from terrestrial ecosystems to the atmosphere. There is a growing body of evidence that suggests that the richness of plants in a community can have significant impacts on ecosystem functioning, but the specific influences of plant species richness (SR), plant functional-type richness and plant community composition on soil respiration rates are unknown. Here we use 10-year-old model plant communities, comprising mature plants transplanted into natural non-sterile soil, to determine how the diversity and composition of plant communities influence soil respiration rates. Our analysis revealed that soil respiration was driven by plant community composition and that there was no significant effect of biodiversity at the three levels tested (SR, functional group and species per functional group). Above-ground plant biomass and root density were included in the analysis as covariates and found to have no effect on soil respiration. This finding is important, because it suggests that loss of particular species will have the greatest impact on soil respiration, rather than changes in biodiversity per se.     

Peeking through a frosty window: molecular insights into the ecology of Arctic soil fungi

Fungi are ubiquitous in Arctic soils, where they function as symbionts and decomposers and may affect the carbon balance of terrestrial ecosystems subjected to climate change, and yet little is known about soil fungi at high latitudes. Here we review data from recent molecular studies to determine broad patterns in Arctic soil fungal ecology. The data indicate comparatively high fungal diversity in Arctic soils, with currently no evidence for lower species richness at higher latitudes. The dominant fungi, and particularly ectomycorrhizal-forming fungi, appear to be cosmopolitan species. Arctic soil fungi are capable of growth at sub-zero temperatures, melanized forms are frequent, host specificity is low and there is evidence that community composition alters under experimental warming. Future challenges are to determine the drivers of fungal diversity, whether or not diversity alters at higher latitudes and how apparently cosmopolitan fungi are able to survive the extreme environments encountered in Arctic habitats.     

Nearctic earthworm fauna in the southern USA: biodiversity and effects on ecosystem processes

An important aspect of biodiversity is the relative importance of species in the functioning of ecosystems; this is particularly so for the soil biota which regulate organic matter and nutrient dynamics in soil. This paper explores some of the relationships between biodiversity and ecosystem processes, using the example of the nearctic earthworm fauna in the glacial refugium of the southern USA. Competitive exclusion of nearctic earthworm species by exotic species has been postulated but there is little direct evidence of it; habitat alteration is the likely cause of native species decline. Reduced earthworm diversity may or may not strongly affect certain ecosystem processes, but more diverse assemblages may more effectively exploit soil resources and influence a wider array of processes. Nearctic species may be better adapted than exotics to local conditions and thus more strongly influence ecosystem processes. Earthworm communities provide a clear case for the union of functional and taxonomic biodiversity studies, because of the recognized ecological strategies of many species. However, some nearctic taxa may deviate from these strategies. Earthworms utilize course woody debris in forests both as a refuge and a resource, while enhancing the decomposition of wood. Management strategies to maintain or increase biodiversity of soil biota should include residual wood on the forest floor. An important task for ecosystem management is to restore biodiversity in degraded ecosystems; introduction programmes and techniques such as periodic burning may increase the abundance and diversity of native earthworm species. Whole ecosystem conservation and management are probably the most practical ways to conserve biodiversity generally and may be the only ways to maintain soil biodiversity.     

The essential role of biodiversity in the key axes of ecosystem function

Biodiversity is essential for maintaining the terrestrial ecosystem multifunctionality (EMF). Recent studies have revealed that the variations in terrestrial ecosystem functions are captured by three key axes: the maximum productivity, water use efficiency, and carbon use efficiency of the ecosystem. However, the role of biodiversity in supporting these three key axes has not yet been explored. In this study, we combined the (i) data collected from more than 840 vegetation plots across a large climatic gradient in China using standard protocols, (ii) data on plant traits and phylogenetic information for more than 2,500 plant species, and (iii) soil nutrient data measured in each plot. These data were used to systematically assess the contribution of environmental factors, species richness, functional and phylogenetic diversity, and community-weighted mean (CWM) and ecosystem traits (i.e., traits intensity normalized per unit land area) to EMF via hierarchical partitioning and Bayesian structural equation modeling. Multiple biodiversity attributes accounted for 70% of the influence of all the variables on EMF, and ecosystems with high functional diversity had high resource use efficiency. Our study is the first to systematically explore the role of different biodiversity attributes, including species richness, phylogenetic and functional diversity, and CWM and ecosystem traits, in the key axes of ecosystem functions. Our findings underscore that biodiversity conservation is critical for sustaining EMF and ultimately ensuring human well-being.     

Global change, soil biodiversity, and nitrogen cycling in terrestrial ecosystems: three case studies

The relative contribution of different soil organism groups to nutrient cycling has been quantified for a number of ecosystems. Some functions, particularly within the N-cycle, are carried out by very specific organisms. Others, including those of decomposition and nutrient release from organic inputs are, however, mediated by a diverse group of bacteria, protozoa, fungi and invertebrate animals. Many authors have hypothesized that there is a high degree of equivalence and flexibility in function within this decomposer community and thence a substantial extent of redundancy in species richness and resilience in functional capacity. Three case studies are presented to examine the relationship between soil biodiversity and nitrogen cycling under global change in ecosystem types from three latitudes, i.e. tundra, temperate grassland and tropical rainforest. In all three ecosystems evidence exists for the potential impact of global change factors (temperature change, CO₂ enrichment, land-use-change) on the composition and diversity of the soil community as well as on various aspects of the nitrogen and other cycles. There is, however, very little unequivocal evidence of direct causal linkage between species richness and nutrient cycling efficiency. Most of the changes detected are shifts in the influence of major functional groups of the soil biota (e.g. between microflora and fauna in decomposition). There seem to be few data, however, from which to judge the significance of changes in diversity within functional groups. Nonetheless the soil biota are hypothesized to be a sensitive link between plant detritus and the availability of nutrients to plant uptake. Any factors affecting the quantity or quality of plant detritus is likely to change this link. Rigorous experimentation on the relationships between soil species richness and the regulation or resilience of nutrient cycles under global change thus remains a high priority.     

Factors related to diversity of decomposer fungi in tropical forests

Recent studies suggest that host-preferences are common among certain groups of tropical fungal decomposers but rare in others, and sometimes occur where we least expect them. Host preferences among microfungi and ascomycetes that decompose leaf litter are common but usually involve differences in relative frequencies more than presence/absence, so their diversity may be loosely correlated with species richness of host trees. Strong host-specificity appears to be rare among wood decomposer fungi, whereas characteristics of their substrata and habitat are very important for this group. Anthropogenic disturbance predisposed a tropical forest to subsequent hurricane damage, and the resulting direct and indirect effects on host diversity and habitat heterogeneity were reflected in the decomposer fungal community more than sixty years after the original disturbance. While species richness of dictyostelid slime molds and functional diversity of their bacterial prey increased with disturbance, the more diverse microfungi and ascomycetes were apparently negatively affected by disturbance.     

The importance of individuals: intraspecific diversity of mycorrhizal plants and fungi in ecosystems

A key component of biodiversity is the number and abundance of individuals (i.e. genotypes), and yet such intraspecific diversity is rarely considered when investigating the effects of biodiversity of mycorrhizal plants and fungi on ecosystem processes. Within a species, individuals vary considerably in important reproductive and functional attributes, including carbon fixation, mycelial growth and nutrient utilization, but this is driven by both genetic and environmental (including climatic) factors. The interactions between individual plants and mycorrhizal fungi can have important consequences for the maintenance of biodiversity and regulation of resource transfers in ecosystems. There is also emerging evidence that assemblages of genotypes may affect ecosystem processes to a similar extent as assemblages of species. The application of whole-genome sequencing and population genomics to mycorrhizal plants and fungi will be crucial to determine the extent to which individual variation in key functional attributes is genetically based. We argue the need to unravel the importance of the diversity (especially assemblages of different evenness and richness) of individuals of both mycorrhizal plants and fungi, and the need to take a 'community genetics' approach to better understand the functional significance of the biodiversity of mycorrhizal symbioses.     

Intensive agriculture reduces soil biodiversity across Europe

Soil biodiversity plays a key role in regulating the processes that underpin the delivery of ecosystem goods and services in terrestrial ecosystems. Agricultural intensification is known to change the diversity of individual groups of soil biota, but less is known about how intensification affects biodiversity of the soil food web as a whole, and whether or not these effects may be generalized across regions. We examined biodiversity in soil food webs from grasslands, extensive, and intensive rotations in four agricultural regions across Europe: in Sweden, the UK, the Czech Republic and Greece. Effects of land‐use intensity were quantified based on structure and diversity among functional groups in the soil food web, as well as on community‐weighted mean body mass of soil fauna. We also elucidate land‐use intensity effects on diversity of taxonomic units within taxonomic groups of soil fauna. We found that between regions soil food web diversity measures were variable, but that increasing land‐use intensity caused highly consistent responses. In particular, land‐use intensification reduced the complexity in the soil food webs, as well as the community‐weighted mean body mass of soil fauna. In all regions across Europe, species richness of earthworms, Collembolans, and oribatid mites was negatively affected by increased land‐use intensity. The taxonomic distinctness, which is a measure of taxonomic relatedness of species in a community that is independent of species richness, was also reduced by land‐use intensification. We conclude that intensive agriculture reduces soil biodiversity, making soil food webs less diverse and composed of smaller bodied organisms. Land‐use intensification results in fewer functional groups of soil biota with fewer and taxonomically more closely related species. We discuss how these changes in soil biodiversity due to land‐use intensification may threaten the functioning of soil in agricultural production systems.     

Ecosystem type drives soil eukaryotic diversity and composition in Europe

Soil eukaryotes play a crucial role in maintaining ecosystem functions and services, yet the factors driving their diversity and distribution remain poorly understood. While many studies focus on some eukaryotic groups (mostly fungi), they are limited in their spatial scale. Here, we analyzed an unprecedented amount of observational data of soil eukaryomes at continental scale (787 sites across Europe) to gain further insights into the impact of a wide range of environmental conditions (climatic and edaphic) on their community composition and structure. We found that the diversity of fungi, protists, rotifers, tardigrades, nematodes, arthropods, and annelids was predominantly shaped by ecosystem type (annual and permanent croplands, managed and unmanaged grasslands, coniferous and broadleaved woodlands), and higher diversity of fungi, protists, nematodes, arthropods, and annelids was observed in croplands than in less intensively managed systems, such as coniferous and broadleaved woodlands. Also in croplands, we found more specialized eukaryotes, while the composition between croplands was more homogeneous compared to the composition of other ecosystems. The observed high proportion of overlapping taxa between ecosystems also indicates that DNA has accumulated from previous land uses, hence mimicking the land transformations occurring in Europe in the last decades. This strong ecosystem-type influence was linked to soil properties, and particularly, soil pH was driving the richness of fungi, rotifers, and annelids, while plant-available phosphorus drove the richness of protists, tardigrades, and nematodes. Furthermore, the soil organic carbon to total nitrogen ratio crucially explained the richness of fungi, protists, nematodes, and arthropods, possibly linked to decades of agricultural inputs. Our results highlighted the importance of long-term environmental variables rather than variables measured at the time of the sampling in shaping soil eukaryotic communities, which reinforces the need to include those variables in addition to ecosystem type in future monitoring programs and conservation efforts.     

树种多样性和土壤微生物多样性对人工林生产力的影响

林分生产力通常会随着树种多样性增加而增加,但不同营养级生物多样性以及树种和土壤微生物多样性之间的相互作用如何影响生产力目前尚不清楚。以亚热带不同丰富度和树种组成的人工林为研究对象,从物种、功能性状、遗传三个维度的树种多样性以及土壤真菌和细菌系统发育多样性,探究了中国亚热带人工林树种多样性和土壤微生物多样性对林分生产力的影响。研究发现,林分生产力随树种功能多样性(FD)(P<0.001)、比叶面积群落加权均值(CWM-SLA)(P<0.01)、树种系统发育多样性(PD)(P<0.05)和土壤真菌多样性(PDF)(P<0.01)的增加而显著增加,分别解释了林分生产力总变异的12.86%、6.80%、3.67%和3.08%。FD和CWM-SLA可分别通过增加土壤真菌、细菌多样性而间接提高林分生产力。研究结果表明多营养级生物多样性是维持高水平林分生产力的基础,树种多样性和土壤微生物多样性之间的自上而下的级联效应在调节生态系统生产力方面发挥着重要作用。     

Litter and soil biodiversity jointly drive ecosystem functions

The decomposition of litter and the supply of nutrients into and from the soil are two fundamental processes through which the above- and belowground world interact. Microbial biodiversity, and especially that of decomposers, plays a key role in these processes by helping litter decomposition. Yet the relative contribution of litter diversity and soil biodiversity in supporting multiple ecosystem services remains virtually unknown. Here we conducted a mesocosm experiment where leaf litter and soil biodiversity were manipulated to investigate their influence on plant productivity, litter decomposition, soil respiration, and enzymatic activity in the littersphere. We showed that both leaf litter diversity and soil microbial diversity (richness and community composition) independently contributed to explain multiple ecosystem functions. Fungal saprobes community composition was especially important for supporting ecosystem multifunctionality (EMF), plant production, litter decomposition, and activity of soil phosphatase when compared with bacteria or other fungal functional groups and litter species richness. Moreover, leaf litter diversity and soil microbial diversity exerted previously undescribed and significantly interactive effects on EMF and multiple individual ecosystem functions, such as litter decomposition and plant production. Together, our work provides experimental evidence supporting the independent and interactive roles of litter and belowground soil biodiversity to maintain ecosystem functions and multiple services.     

Significant and persistent impact of timber harvesting on soil microbial communities in Northern coniferous forests

Forest ecosystems have integral roles in climate stability, biodiversity and economic development. Soil stewardship is essential for sustainable forest management. Organic matter (OM) removal and soil compaction are key disturbances associated with forest harvesting, but their impacts on forest ecosystems are not well understood. Because microbiological processes regulate soil ecology and biogeochemistry, microbial community structure might serve as indicator of forest ecosystem status, revealing changes in nutrient and energy flow patterns before they have irreversible effects on long-term soil productivity. We applied massively parallel pyrosequencing of over 4.6 million ribosomal marker sequences to assess the impact of OM removal and soil compaction on bacterial and fungal communities in a field experiment replicated at six forest sites in British Columbia, Canada. More than a decade after harvesting, diversity and structure of soil bacterial and fungal communities remained significantly altered by harvesting disturbances, with individual taxonomic groups responding differentially to varied levels of the disturbances. Plant symbionts, like ectomycorrhizal fungi, and saprobic taxa, such as ascomycetes and actinomycetes, were among the most sensitive to harvesting disturbances. Given their significant ecological roles in forest development, the fate of these taxa might be critical for sustainability of forest ecosystems. Although abundant bacterial populations were ubiquitous, abundant fungal populations often revealed a patchy distribution, consistent with their higher sensitivity to the examined soil disturbances. These results establish a comprehensive inventory of bacterial and fungal community composition in northern coniferous forests and demonstrate the long-term response of their structure to key disturbances associated with forest harvesting.     

Relationships between macro-fungal dark diversity and habitat parameters using LiDAR

Despite the important role of fungi for ecosystems, relatively little is known about the factors underlying the dynamics of their diversity. Moreover, studies do not typically consider their dark diversity: the species absent from an otherwise suitable site. Here, we examined potential drivers of local fungal dark diversity in temperate woodland and open habitats using LiDAR and in-situ field measurements, combined with a systematically collected and geographically comprehensive macro-fungi and plant data set. For the first time, we also estimated species pools of fungi by considering both plant and fungi co-occurrences. The most important LiDAR variables for explaining fungal dark diversity were amplitude and echo ratio, which represent vegetation structure. These results suggest that the local fungal dark diversity is highest in production forests like plantations and lowest in more open forests and in open habitats with little woody vegetation. Plant species richness was the strongest explanatory factor overall and negatively correlated with local fungal dark diversity. Soil fertility showed a positive relationship with dark diversity in open habitats. These findings indicate that the local dark diversity of macro-fungi is highest in areas with a relatively high human impact (typically areas with low plant species richness and high soil fertility). Overall, this study brings novel insights into local macro-fungi dark diversity patterns, suggesting that a multitude of drivers related to both soil and vegetation act simultaneously to determine fungal dark diversity.     

树种多样性对土壤微生物群落结构和元素生物地球化学循环的影响研究进展

森林是陆地生态系统的重要组成部分,其巨大的生产力和生态服务功能对人类的生存和发展至关重要。森林树种多样性增加能够显著提高森林生产力,关于树种多样性如何影响地下生物多样性及生态功能逐渐受到国内外学者的广泛关注。从土壤微生物及其介导的元素生物地球化学循环这一视角出发,综述了树种多样性对土壤细菌和真菌多样性、群落结构及功能的影响,提出需要进一步深入研究的方向。总体来说,树种多样性有利于增加土壤细菌生物量和多样性,是预测病原性真菌和菌根真菌多样性及群落结构的重要生物因子。树种多样性能增加土壤有机碳储量,增强森林土壤的甲烷氧化能力,并提高土壤磷周转速率及有效磷含量。关于树种多样性对森林土壤氮循环的影响需考虑多样性假说和质量比假说的相对贡献。今后应加强树种多样性对多个营养级之间相互作用的研究;关注树种多样性对生态系统多功能的影响;加强学科交叉,引入微生物种群动态模型和气候模型等模型预测方法,研究树种多样性对全球气候变化的应对机制,以期促进地上植物多样性与地下生态系统功能关系的研究,增强森林生态系统应对未来全球环境变化的能力。