Intraspecific traits change biodiversity effects on ecosystem functioning under metal stress

Studies investigating the impacts of biodiversity loss on ecosystem processes have often reached different conclusions, probably because insufficient attention has been paid to some aspects including (1) which biodiversity measure (e.g., species number, species identity or trait) better explains ecosystem functioning, (2) the mechanisms underpinning biodiversity effects, and (3) how can environmental context modulates biodiversity effects. Here, we investigated how species number (one to three species) and traits of aquatic fungal decomposers (by replacement of a functional type from an unpolluted site by another from a metal-polluted site) affect fungal production (biomass acumulation) and plant litter decomposition in the presence and absence of metal stress. To examine the putative mechanisms that explain biodiversity effects, we determined the contribution of each fungal species to the total biomass produced in multicultures by real-time PCR. In the absence of metal, positive diversity effects were observed for fungal production and leaf decomposition as a result of species complementarity. Metal stress decreased diversity effects on leaf decomposition in assemblages containing the functional type from the unpolluted site, probably due to competitive interactions between fungi. However, dominance effect maintained positive diversity effects under metal stress in assemblages containing the functional type from the metal-polluted site. These findings emphasize the importance of intraspecific diversity in modulating diversity effects under metal stress, providing evidence that trait-based diversity measures should be incorporated when examining biodiversity effects.     

Functional structure of biological communities predicts ecosystem multifunctionality

The accelerating rate of change in biodiversity patterns, mediated by ever increasing human pressures and global warming, demands a better understanding of the relationship between the structure of biological communities and ecosystem functioning (BEF). Recent investigations suggest that the functional structure of communities, i.e. the composition and diversity of functional traits, is the main driver of ecological processes. However, the predictive power of BEF research is still low, the integration of all components of functional community structure as predictors is still lacking, and the multifunctionality of ecosystems (i.e. rates of multiple processes) must be considered. Here, using a multiple-processes framework from grassland biodiversity experiments, we show that functional identity of species and functional divergence among species, rather than species diversity per se, together promote the level of ecosystem multifunctionality with a predictive power of 80%. Our results suggest that primary productivity and decomposition rates, two key ecosystem processes upon which the global carbon cycle depends, are primarily sustained by specialist species, i.e. those that hold specialized combinations of traits and perform particular functions. Contrary to studies focusing on single ecosystem functions and considering species richness as the sole measure of biodiversity, we found a linear and non-saturating effect of the functional structure of communities on ecosystem multifunctionality. Thus, sustaining multiple ecological processes would require focusing on trait dominance and on the degree of community specialization, even in species-rich assemblages.     

菌根真菌多样性与植物多样性的相互作用研究进展

菌根共生双方多样性影响着生态系统的过程与功能。菌根真菌-寄主植物之间的共生组合存在偏好性或特异性,这导致菌根真菌对寄主植物的效益差异和寄主植物对菌根真菌的利益差别：两者在互利共生过程中不仅相互选择,还存在相互促进与制约的关系(如互补与选择效应、竞争),从而在一定程度上决定生态系统的演化与发展。本文概述了植物多样性与菌根真菌多样性的相互影响,探讨了两者互作可能存在的调控因素与机制,对存在的问题和争议进行了总结,并提出了进一步研究的方向。深入阐明植物多样性与菌根真菌多样性之间的互作关系,将丰富生物共生学理论,增强菌根应用潜力及生物多样性的维持。     

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.     

Does chemical composition of individual Scots pine trees determine the biodiversity of their associated ground vegetation?

Despite plant secondary metabolites being major determinants of species interactions and ecosystem processes, their role in the maintenance of biodiversity has received little attention. In order to investigate the relationship between chemical and biological diversity in a natural ecosystem, we considered the impact of chemical diversity in individual Scots pine trees (Pinus sylvestris) on species richness of associated ground vegetation. Scots pine trees show substantial genetically determined constitutive variation between individuals in concentrations of a group of secondary metabolites, the monoterpenes. When the monoterpenes of particular trees were assessed individually, there was no relationship with species richness of associated ground flora. However, the chemical diversity of monoterpenes of individual trees was significantly positively associated with the species richness of the ground vegetation beneath each tree, mainly the result of an effect among the non‐woody vascular plants. This correlation suggests that the chemical diversity of the ecosystem dominant species has an important role in shaping the biodiversity of the associated plant community. The extent and significance of this effect, and its underlying processes require further investigation.     

AM真菌在草原生态系统中的功能

AM真菌是土壤生态系统中重要的微生物类群,能与陆地生态系统中80%以上的高等植物建立共生体系。目前,AM真菌在维持草原生态系统稳定性中的功能已经成为生态学研究的热点问题之一。基于此,从植物个体、种群、群落和生态系统等不同层次探究AM真菌在维持植物群落多样性和草原生态系统稳定性中的功能。分析发现在个体水平上,AM真菌对宿主植物具有促生效应、抑制效应或中性效应。在种群水平上,分析AM真菌对不同宿主植物吸收土壤矿质营养的分配和调控策略,围绕构成草原植被的两大组成成分:牧草和有毒植物,论述AM真菌对植物种群增长和衰败的调控机制,并从草原植物群落的物种多样性和稳定性角度,探讨AM真菌与植物群落之间的相关性。在生态系统水平上,围绕AM真菌对草原生态系统的演替和退化草原的修复等展开论述,以期为利用AM真菌开展草原生态系统保护和恢复治理提供理论依据,并对草原菌根生态学领域未来的研究进行展望。     

A single European aspen (Populus tremula) tree individual may potentially harbour dozens of Cenococcum geophilum ITS genotypes and hundreds of species of ectomycorrhizal fungi

Ectomycorrhizal fungi (EcMF) form diverse communities and link different host plants into mycorrhizal networks, yet little is known about the magnitude of mycobiont diversity of a single tree individual. This study addresses species richness and spatial structure of EcMF in the root system of a single European aspen (Populus tremula) individual in an old-growth boreal mixed forest ecosystem in Estonia. Combining morphological and molecular identification methods for both plant and fungi, 122 species of EcMF were recovered from 103 root samples of the single tree. Richness estimators predicted the total EcMF richness to range from 182 to 207 species, reflecting the observation of 62.3% singletons and doubletons within the community. Fine-scale genetic diversity in Cenococcum geophilum indicates the presence of 23 internal transcribed spacer genotypes. EcMF community was significantly spatially autocorrelated only at the lineage level up to 3 m distance, but not at the species level. Proximity of other hosts had a significant effect on the spatial distribution of EcMF lineages. This study demonstrates that a single tree may host as many EcMF species and individuals as recovered on multiple hosts in diverse communities over larger areas.     

The Soil FungiLog procedure: method and analytical approaches toward understanding fungal functional diversity

Conservation methods often are focused on preserving the biodiversity of a particular landscape or ecosystem. Scientists frequently employ species richness as an indicator of biodiversity. However, species richness data are problematic when attempts are made to enumerate microfungi, particularly those from the soil. Many soil fungi fail to sporulate, making identification difficult. Other means of assessing the importance of fungi to ecosystem preservation must be developed. Otherwise, microfungi might be overlooked in discussions of ecosystem management and conservation issues. Herein, we have described a procedure (Soil FungiLog) and analytical techniques that will let investigators examine the functional role that soil fungi play in providing structure and stability to ecosystems. Ecosystem function in many cases might be more important than species diversity in gaining an understanding of ecosystem dynamics. Functional attributes are critical for maintaining ecosystem structure and stability. The preservation of the functions associated with the extant biota, particularly from soil microbes, might be just as important as species diversity in the conservation of ecosystems and biodiversity. The Soil FungiLog procedure was used to assess functional diversity of soil fungi in a Georgia forest disturbed by human activity and along an elevational gradient in the Chihuahuan Desert. Sites within each location were separated on the basis of fungal carbon substrate utilization profiles. These profiles were analyzed to provide information regarding the functional diversity of soil fungal assemblages at each site. The effects of disturbance and elevation were evaluated with respect to soil fungal functional diversity.     

A multi-faceted framework of diversity for prioritizing the conservation of fish assemblages

Floodplain waterbodies and their biodiversity are increasingly threatened by human activities. Given the limited resources available to protect them, methods to identify the most valuable areas for biodiversity conservation are urgently needed. In this study, we used freshwater fish assemblages in floodplain waterbodies to propose an innovative method for selecting priority areas based on four aspects of their diversity: taxonomic (i.e. according to species classification), functional (i.e. relationship between species and ecosystem processes), natural heritage (i.e. species threat level), and socio-economic (i.e. species interest to anglers and fishermen) diversity. To quantitatively evaluate those aspects, we selected nine indices derived either from metrics computed at the species level and then combined for each assemblage (species rarity, origin, biodiversity conservation concern, functional uniqueness, functional originality, fishing interest), or from metrics directly computed at the assemblage level (species richness, assemblage rarity, diversity of biological traits). Each of these indices belongs to one of the four aspects of diversity. A synthetic index defined as the sum of the standardized aspects of diversity was used to assess the multi-faceted diversity of fish assemblages. We also investigated whether the two main environmental gradients at the catchment (distance from the sea) and at the floodplain (lateral connectivity of the waterbodies) scales influenced the diversity of fish assemblages, and consequently their potential conservation value. Finally, we propose that the floodplain waterbodies that should be conserved as a priority are those located in the downstream part of the catchment and which have a substantial lateral connectivity with the main channel.     

Species and individual replacements contribute more than nestedness to shape vertebrate scavenger metacommunities

Understanding the mechanisms that organize biodiversity is central in ecology and conservation. Beta diversity links local (alfa) and regional (gamma) diversity, giving insight into how communities organize spatially. Metacommunity ecology provides the framework to interpret regional and local processes interacting to shape communities. However, the lack of metacommunity studies for large vertebrates may limit the understanding and compromise the preservation of ecosystem functions and services. We aim to understand the mechanisms underlying differences in species composition among vertebrate scavenger communities ? which provide key ecosystem functions, e.g. carrion consumption ? within a metacommunity context. We obtained species richness and abundances at scavenger communities consuming ungulate carcasses monitored through motion‐triggered remote cameras in seven terrestrial ecosystems in Spain. We partitioned beta diversity to decompose incidence‐based (species presence/absence) and abundance‐based dissimilarities into their components (turnover/balanced variation and nestedness/abundance gradient, respectively). We identified the environmental factors explaining the observed patterns. The vertebrate scavenger metacommunity consisted of 3101 individuals from 30 species. Changes in composition among ecosystems were mostly (> 84%) due to species or individual replacement (i.e. turnover or balanced variation). Species or individual loss/gain (i.e. nestedness or abundance gradient) accounted for 13–16% of these changes. Mean carcass weight, elevation and habitat diversity were the main factors explaining species/individual replacement. Our findings suggest that local processes such as species‐sorting through habitat heterogeneity would dominate scavenger metacommunity dynamics together with stochastic forces (i.e. related to carrion unpredictability and scavenging being a widespread strategy among vertebrates). The presence of structured patterns (i.e. nestedness) in beta diversity could reflect a role of deterministic processes: mass‐effects through dispersal and defaunation. Vultures are long‐distance foragers and functionally dominant species, which would connect local assemblages within the metacommunity, supporting scavenger diversity and functions across space. These results highlight the importance of managing vertebrate scavenger assemblages within a metacommunity context.     

Disentangling community functional components in a litter‐macrodetritivore model system reveals the predominance of the mass ratio hypothesis

Recent investigations have shown that two components of community trait composition are important for key ecosystem processes: (i) the community‐weighted mean trait value (CWM), related to the mass ratio hypothesis and dominant trait values in the community, and (ii) functional diversity (FD), related to the complementarity hypothesis and the divergence of trait values. However, no experiments controlling for the inherent dependence between CWM and FD have been conducted so far. We used a novel experimental framework to disentangle the unique and shared effects of CWM and FD in a leaf litter‐macrodetritivore model system. We manipulated isopod assemblages varying in species number, CWM and FD of litter consumption rate to test the relative contribution of these community parameters in the decomposition process. We showed that CWM, but also the combination of CWM and FD, is a main factor controlling litter decomposition. When we tested individual biodiversity components separately, CWM of litter consumption rate showed a significant effect on decomposition, while FD and species richness alone did not. Our study demonstrated that (i) trait composition rather than species diversity drives litter decomposition, (ii) dominant trait values in the community (CWM) play a chief role in driving ecosystem processes, corroborating the mass ratio hypothesis, and (iii) trait dissimilarity can contribute in modulating the overall biodiversity effects. Future challenge is to assess whether the generality of our finding, that is, that dominant trait values (CWM) predominate over trait dissimilarity (FD), holds for other ecosystem processes, environmental conditions and different spatial and temporal scales.     

Intraspecific Diversity Regulates Fungal Productivity and Respiration

Individuals and not just species are key components of biodiversity, yet the relationship between intraspecific diversity and ecosystem functioning in microbial systems remains largely untested. This limits our ability to understand and predict the effects of altered genetic diversity in regulating key ecosystem processes and functions. Here, we use a model fungal system to test the hypothesis that intraspecific genotypic richness of Paxillus obscurosporus stimulates biomass and CO₂ efflux, but that this is dependent on nitrogen supply. Using controlled experimental microcosms, we show that populations containing several genotypes (maximum 8) of the fungus had greater productivity and produced significantly more CO₂ than those with fewer genotypes. Moreover, intraspecific diversity had a much stronger effect than a four-fold manipulation of the carbon:nitrogen ratio of the growth medium. The effects of intraspecific diversity were underpinned by strong roles of individuals, but overall intraspecific diversity increased the propensity of populations to over-yield, indicating that both complementarity and selection effects can operate within species. Our data demonstrate the importance of intraspecific diversity over a range of nitrogen concentrations, and the need to consider fine scale phylogenetic information of microbial communities in understanding their contribution to ecosystem processes.     

Interactive effects of species richness and species traits on functional diversity and redundancy

The importance of species diversity for ecosystem function has emerged as a key question for conservation biology. Recently, there has been a shift from examining the role of species richness in isolation towards understanding how species interact to effect ecosystem function. Here, we briefly review theoretical predictions regarding species contributions to functional diversity and redundancy and further use simulated data to test combined effects of species richness, number of functional traits, and species differences within these traits on unique species contributions to functional diversity and redundancy, as well as on the overall functional diversity and redundancy within species assemblages. Our results highlighted that species richness and species functional attributes interact in their effects on functional diversity. Moreover, our simulations suggested that functional differences among species have limited effects on the proportion of redundancy of species contributions as well as on the overall redundancy within species assemblages, but that redundancy rather was determined by number of traits and species richness. Our simulations finally indicated scale dependence in the relative effects of species richness and functional attributes, which suggest that the relative influence of these factors may affect individual contributions differently compared to the overall ecosystem function of species assemblages. We suggest that studies on the relationship between biological diversity and ecosystem function will benefit from focusing on multiple processes and ecological interactions, and that the relative functional attributes of species will have pivotal roles for the ecosystem function of a given species assembly.     

The effects of road building on arbuscular mycorrhizal fungal diversity in Huangshan Scenic Area

Arbuscular mycorrhizal (AM) fungi are vital soil microbes that connect many individual plants into a large functional organism via a vast mycelial network under the ground. In this study, the changes of soil AM fungal community in response to road-building disturbance caused by tourism development in Huangshan (Yellow Mountain) Scenic Area are assessed. Road building have brought negative effects on AM fungal community, inducing lower diversity parameters, including species number, spore density and diversity indices. However, the dominant genus and species of AM fungi which play key roles in the AM fungal community composition are quite similar before and after road building. Moreover, there are no significant differences in species richness of AM fungi associated with plants, suggesting the tolerance of AM fungal community to the disturbance of road building.     

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.     

Patterns and regulation of mycorrhizal plant and fungal diversity

The diversity of mycorrhizal fungi does not follow patterns of plant diversity, and the type of mycorrhiza may regulate plant species diversity. For instance, coniferous forests of northern latitudes may have more than 1000 species of ectomycorrhizal (EM) fungi where only a few ectomycorrhizal plant species dominate, but there are fewer than 25 species of arbuscular mycorrhizal (AM) fungi in tropical deciduous forest in Mexico with 1000 plant species. AM and EM fungi are distributed according to biome, with AM fungi predominant in arid and semiarid biomes, and EM fungi predominant in mesic biomes. In addition, AM fungi tend to be more abundant in soils of low organic matter, perhaps explaining their predominance in moist tropical forest, and EM fungi generally occur in soils with higher surface organic matter.EM fungi are relatively selective of host plant species, while AM tend to be generalists. Similar morphotypes of AM fungi collected from different sites confer different physiological benefits to the same plant species. While the EM fungi have taxonomic diversity, the AM fungi must have physiological diversity for individual species to be so widespread, as supported by existing studies. The environmental adaptations of mycorrhizal fungi are often thought to be determined by their host plant, but we suggest that the physiology and genetics of the fungi themselves, along with their responses to the plant and the environment, regulates their diversity. We observed that one AM plant species,Artemisia tridentata, was associated with different fungal species across its range, indicating that the fungi can respond to the environment directly and must not do so indirectly via the host. Different species of fungi were also active during different times of the growing season on the same host, again suggesting a direct response to the environment.These patterns suggest that even within a single functional group of microorganisms, mycorrhizal fungi, considerable diversity exists. A number of researchers have expressed the concept of functional redundancy within functional groups of microorganisms, implying that the loss of a few species would not be detectable in ecosystem functioning. However, there may be high functional diversity of AM fungi within and across habitats, and high species diversity as well for EM fungi. If one species of mycorrhizal fungus becomes extinct in a habitat, field experimental data on AM fungi suggest there may be significant shifts in how plants acquire resources and grown in that habitat.     

An ecological view of the formation of VA mycorrhizas

In spite of the major advances in understanding the functioning of symbioses between plants and arbuscular mycorrhizal fungi, details of the ecology of mycorrhizal fungi are not well documented. The benefits of the association are related to the timing and extent of colonization of roots, and fungi differ in their contribution to plant growth and presumably to soil aggregation. Knowledge of the processes that lead to successful colonization of roots by beneficial fungi at appropriate times for the host plants will form the basis of guidelines for soil management to maximize the benefits from the symbiosis. Fungi differ in the manner and extent to which they colonize roots. They also differ in their capacity to form propagules. The importance of hyphae, spores and propagules within living or dead mycorrhizal roots also differs among species and for the same species in different habitats. The relationships between colonization of roots and propagule formation, and between propagule distribution and abundance and subsequent mycorrhiza formation, for different fungi in field environments, are not well understood. Methods for quantifying mycorrhizal fungi are not especially suitable for distinguishing among different fungi within roots. Consequenctly, the dynamics of colonization of roots by different fungi, within and between seasons, have been little studied. Research is required that focuses on the dynamics of fungi within roots as well as on changes in the abundance of propagules of different fungi within soil. Interactions between fungi during the colonization of roots, the colonization of soil by hyphae and sporulation are all poorly understood. Without knowledge of these processes, it will by difficult to predict the likely success of inoculation with introduced fungi. Such knowledge is also required for selecting soil management procedures to enhance growth and survival of key species within the population. The relative tolerance of various fungi to perturbations in their surroundings will provide a basis for identifying those fungi that are likely to persist in specific environments. The processes that influence mycorrhizal fungi in field soils can be identified in controlled studies. However, greater emphasis is required on studying these processes with mixed populations of fungi. The role played by diversity within populations of mycorrhizal fungi is virtually unexplored.     

Functional diversity of plant-pollinator interaction webs enhances the persistence of plant communities

Pollination is exclusively or mainly animal mediated for 70% to 90% of angiosperm species. Thus, pollinators provide an essential ecosystem service to humankind. However, the impact of human-induced biodiversity loss on the functioning of plant–pollinator interactions has not been tested experimentally. To understand how plant communities respond to diversity changes in their pollinating fauna, we manipulated the functional diversity of both plants and pollinators under natural conditions. Increasing the functional diversity of both plants and pollinators led to the recruitment of more diverse plant communities. After two years the plant communities pollinated by the most functionally diverse pollinator assemblage contained about 50% more plant species than did plant communities pollinated by less-diverse pollinator assemblages. Moreover, the positive effect of functional diversity was explained by a complementarity between functional groups of pollinators and plants. Thus, the functional diversity of pollination networks may be critical to ecosystem sustainability.     

Functional Diversity of Plant–Pollinator Interaction Webs Enhances the Persistence of Plant Communities

Pollination is exclusively or mainly animal mediated for 70% to 90% of angiosperm species. Thus, pollinators provide an essential ecosystem service to humankind. However, the impact of human-induced biodiversity loss on the functioning of plant–pollinator interactions has not been tested experimentally. To understand how plant communities respond to diversity changes in their pollinating fauna, we manipulated the functional diversity of both plants and pollinators under natural conditions. Increasing the functional diversity of both plants and pollinators led to the recruitment of more diverse plant communities. After two years the plant communities pollinated by the most functionally diverse pollinator assemblage contained about 50% more plant species than did plant communities pollinated by less-diverse pollinator assemblages. Moreover, the positive effect of functional diversity was explained by a complementarity between functional groups of pollinators and plants. Thus, the functional diversity of pollination networks may be critical to ecosystem sustainability.     

Forest structure and fungal endophytes

Sufficient biodiversity is required for ecosystem functions. The question is how we can assess required biodiversity if we are able to recognize only a fraction of diversity, and/or unable to place a known species into a trophic level or into their niche dimensions. The species diversity of higher plants and animals usually can be assessed in most terrestrial environments. In contrast, microbial diversity is often ignored although the number and genetic diversity of microbes is enormous, and are profoundly important as plant and animal mutualists, pathogens, parasites and saprobes. Thus, one of the biggest challenges when disentangling relevant diversity to ecosystem functions is to reveal composition of focal microbial assemblage and the place of the key groups of them in the food web. In this review I focus on ubiquitous but poorly understood group of foliar fungi, asymptomatic endophytic fungi, of woody plants emphasizing how geographic, age and genetic structure of forest might affect endophyte-plant interactions.