Bacterial genome size and gene functional diversity negatively correlate with taxonomic diversity along a pH gradient.

Bacterial gene repertoires reflect adaptive strategies, contribute to ecosystem functioning and are limited by genome size. However, gene functional diversity does not necessarily correlate with taxonomic diversity because average genome size may vary by community. Here, we analyse gene functional diversity (by shotgun metagenomics) and taxonomic diversity (by 16S rRNA gene amplicon sequencing) to investigate soil bacterial communities along a natural pH gradient in 12 tropical, subtropical, and temperate forests. We find that bacterial average genome size and gene functional diversity decrease, whereas taxonomic diversity increases, as soil pH rises from acid to neutral; as a result, bacterial taxonomic and functional diversity are negatively correlated. The gene repertoire of acid-adapted oligotrophs is enriched in functions of signal transduction, cell motility, secretion system, and degradation of complex compounds, while that of neutral pH-adapted copiotrophs is enriched in functions of energy metabolism and membrane transport. Our results indicate that a mismatch between taxonomic and functional diversity can arise when environmental factors (such as pH) select for adaptive strategies that affect genome size distributions.

Assembly processes lead to divergent soil fungal communities within and among 12 forest ecosystems along a latitudinal gradient.

Latitudinal gradients provide opportunities to better understand soil fungal community assembly and its relationship with vegetation, climate, soil and ecosystem function. Understanding the mechanisms underlying community assembly is essential for predicting compositional responses to changing environments. We quantified the relative importance of stochastic and deterministic processes in structuring soil fungal communities using patterns of community dissimilarity observed within and between 12 natural forests and related these to environmental variation within and among sites. The results revealed that whole fungal communities and communities of arbuscular and ectomycorrhizal fungi consistently exhibited divergent patterns but with less divergence for ectomycorrhizal fungi at most sites. Within those forests, no clear relationships were observed between the degree of divergence within fungal and plant communities. When comparing communities at larger spatial scales, among the 12 forests, we observed distinct separation in all three fungal groups among tropical, subtropical and temperate climatic zones. Soil fungal ß-diversity patterns between forests were also greater when comparing forests exhibiting high environmental heterogeneity. Taken together, although large-scale community turnover could be attributed to specific environmental drivers, the differences among fungal communities in soils within forests was high even at local scales.

Late Quaternary climate change explains soil fungal community composition rather than fungal richness in forest ecosystems.

The dramatic climate fluctuations of the late Quaternary have influenced the diversity and composition of macroorganism communities, but how they structure belowground microbial communities is less well known. Fungi constitute an important component of soil microorganism communities. They play an important role in biodiversity maintenance, community assembly, and ecosystem functioning, and differ from many macroorganisms in many traits. Here, we examined soil fungal communities in Chinese temperate, subtropical, and tropic forests using Illumina MiSeq sequencing of the fungal ITS1 region. The relative effect of late Quaternary climate change and contemporary environment (plant, soil, current climate, and geographic distance) on the soil fungal community was analyzed. The richness of the total fungal community, along with saprotrophic, ectomycorrhizal (EM), and pathogenic fungal communities, was influenced primarily by the contemporary environment (plant and/or soil) but not by late Quaternary climate change. Late Quaternary climate change acted in concert with the contemporary environment to shape total, saprotrophic, EM, and pathogenic fungal community compositions and with a stronger effect in temperate forest than in tropic-subtropical forest ecosystems. Some contemporary environmental factors influencing total, saprotrophic, EM, and pathogenic fungal communities in temperate and tropic-subtropical forests were different. We demonstrate that late Quaternary climate change can help to explain current soil fungal community composition and argue that climatic legacies can help to predict soil fungal responses to climate change.

Host plant phylogeny and geographic distance strongly structure Betulaceae-associated ectomycorrhizal fungal communities in Chinese secondary forest ecosystems.

Environmental filtering and dispersal limitation are two of the primary drivers of community assembly in ecosystems, but their effects on ectomycorrhizal (EM) fungal communities associated with wide ranges of Betulaceae taxa at a large scale are poorly documented. In this study, we examined EM fungal communities associated with 23 species from four genera (Alnus, Betula, Carpinus and Corylus) of Betulaceae in Chinese secondary forest ecosystems, using Illumina MiSeq sequencing of the ITS2 region. Effects of host plant phylogeny, soil, climate and geographic distance on EM fungal community were explored. In total, we distinguished 1738 EM fungal operational taxonomic units (OTUs) at a 97% sequence similarity level. The EM fungal communities of Alnus had significantly lower OTU richness than those associated with the other three plant genera. The EM fungal OTU richness was significantly affected by geographic distance, host plant phylogeny, soil and climate. The EM fungal community composition was significantly influenced by host plant phylogeny (12.1% of variation explained in EM fungal community), geographic distance (7.7%), soil (4.6%) and climate (1.1%). This finding highlights that environmental filtering linked to host plant phylogeny and dispersal limitation strongly influence EM fungal communities associated with Betulaceae plants in Chinese secondary forest ecosystems.

Community Assembly of Endophytic Fungi in Ectomycorrhizae of Betulaceae Plants at a Regional Scale.

The interaction between aboveground and belowground biotic communities drives community assembly of plants and soil microbiota. As an important component of belowground microorganisms, root-associated fungi play pivotal roles in biodiversity maintenance and community assembly of host plants. The Betulaceae plants form ectomycorrhizae with soil fungi and widely distribute in various ecosystems. However, the community assembly of endophytic fungi in ectomycorrhizae is less investigated at a large spatial scale. Here, we examined the endophytic fungal communities in ectomycorrhizae of 22 species in four genera belonging to Betulaceae in Chinese forest ecosystems, using Illumina Miseq sequencing of internal transcribed spacer 2 amplicons. The relative contribution of host phylogeny, climate and soil (environmental filtering) and geographic distance (dispersal limitation) on endophytic fungal community was disentangled. In total, 2,106 endophytic fungal operational taxonomic units (OTUs) were obtained at a 97% sequence similarity level, dominated by Leotiomycetes, Agaricomycetes, Eurotiomycetes, and Sordariomycetes. The endophytic fungal OTU richness was significantly related with host phylogeny, geographic distance, soil and climate. The endophytic fungal community composition was significantly affected by host phylogeny (19.5% of variation explained in fungal community), geographic distance (11.2%), soil (6.1%), and climate (1.4%). This finding suggests that environmental filtering by plant and abiotic variables coupled with dispersal limitation linked to geographic distance determines endophytic fungal community assembly in ectomycorrhizae of Betulaceae plants, with host phylogeny being a stronger determinant than other predictor variables at the regional scale.

Host Phylogeny Is a Major Determinant of Fagaceae-Associated Ectomycorrhizal Fungal Community Assembly at a Regional Scale.

Environmental filtering (niche process) and dispersal limitation (neutral process) are two of the primary forces driving community assembly in ecosystems, but how these processes affect the Fagaceae-associated ectomycorrhizal (EM) fungal community at regional scales is so far poorly documented. We examined the EM fungal communities of 61 plant species in six genera belonging to the Fagaceae distributed across Chinese forest ecosystems (geographic distance up to ∼3,757 km) using Illumina Miseq sequencing of ITS2 sequences. The relative effects of environmental filtering (e.g., host plant phylogeny, soil and climate) and dispersal limitation (e.g., spatial distance) on the EM fungal community were distinguished using multiple models. In total, 2,706 operational taxonomic units (OTUs) of EM fungi, corresponding to 54 fungal lineages, were recovered at a 97% sequence similarity level. The EM fungal OTU richness was significantly affected by soil pH and nutrients and by host phylogeny. The EM fungal community composition was significantly influenced by combinations of host phylogeny, spatial distance, soil and climate. Furthermore, host phylogeny had the greatest effect on EM fungal community. The study suggests that the assembly of the EM fungal community is governed by both environmental filtering and dispersal limitation, with host effect being the most important determinant at the regional scale.

Relationships between soil fungal and woody plant assemblages differ between ridge and valley habitats in a subtropical mountain forest.

Elucidating interactions of above-ground and below-ground communities in different habitat types is essential for understanding biodiversity maintenance and ecosystem functioning. Using 454 pyrosequencing of ITS2 sequences we examined the relationship between subtropical mountain forest soil fungal communities, abiotic conditions, and plant communities using correlation and partial models. Ridge and valley habitats with differing fungal communities were delineated. Total, saprotrophic and pathogenic fungal richness were significantly correlated with plant species richness and/or soil nutrients and moisture in the ridge habitat, but with habitat convexity or basal area of Castanopsis eyrei in the valley habitat. Ectomycorrhizal (EM) fungal richness was significantly correlated with basal area of C. eyrei and total EM plants in the ridge and valley habitats, respectively. Total, saprotrophic, pathogenic and EM fungal compositions were significantly correlated with plant species composition and geographic distance in the ridge habitat, but with various combinations of plant species composition, plant species richness, soil C : N ratio and pH or no variables in the valley habitat. Our findings suggest that mechanisms influencing soil fungal diversity and community composition differ between ridge and valley habitats, and relationships between fungal and woody plant assemblages depend on habitat types in the subtropical forest ecosystem.