It is increasingly acknowledged that climate change is influencing terrestrial ecosystems by increased drought and rainfall intensities. Soil microbes are key drivers of many processes in terrestrial systems and rely on water in soil pores to fulfill their life cycles and functions. However, little is known on how drought and rainfall fluctuations, which affect the composition and structure of microbial communities, persist once original moisture conditions have been restored. Here, we study how simulated short-term drying and re-wetting events shape the community composition of soil fungi and prokaryotes. In a mesocosm experiment, soil was exposed to an extreme drought, then re-wetted to optimal moisture (50% WHC, water holding capacity) or to saturation level (100% WHC). Composition, community structure and diversity of microbes were measured by sequencing ITS and 16S rRNA gene amplicons 3 weeks after original moisture content had been restored. Drying and extreme re-wetting decreased richness of microbial communities, but not evenness. Abundance changes were observed in only 8% of prokaryote OTUs, and 25% of fungal OTUs, whereas all other OTUs did not differ between drying and re-wetting treatments. Two specific legacy response groups (LRGs) were observed for both prokaryotes and fungi. OTUs belonging to the first LRG decreased in relative abundance in soil with a history of drought, whereas OTUs that increased in soil with a history of drought formed a second LRG. These microbial responses were spread among different phyla. Drought appeared to be more important for the microbial community composition than the following extreme re-wetting. 16S profiles were correlated with both inorganic N concentration and basal respiration and ITS profiles correlated with fungal biomass. We conclude that a drying and/or an extreme re-wetting history can persist in soil microbial communities via specific response groups composed of members with broad phylogenetic origins, with possible functional consequences on soil processes and plant species. As a large fraction of OTUs responding to drying and re-wetting belonged to the rare biosphere, our results suggest that low abundant microbial species are potentially important for ecosystem responses to extreme weather events.

Most studies focusing on plant-soil feedbacks (PSFs) have considered direct interactions between plants, abiotic conditions (e. g., soil nutrients) and rhizosphere communities (e.g., pathogens, mutualists). However, few studies have addressed the role of indirect interactions mediated by plant litter inputs. This is problematic because it has left a major gap in our understanding of PSFs in natural ecosystems, where plant litter is a key component of feedback effects. Here, we propose a new conceptual framework that integrates rhizosphere-and litter-mediated PSF effects. Our framework provides insights into the relative contribution of direct effects mediated by interactions between plants and soil rhizosphere organisms, and indirect effects between plants and decomposer organisms mediated by plant root and shoot litter. We distinguish between three pathways through which senesced root and shoot litter may influence PSFs. Specifically, we examine: (1) physical effects of litter (layer) traits on seed germination, soil structure, and plant growth; (2) chemical effects of litter on concentrations of soil nutrients and secondary metabolites (e.g., allelopathic chemicals); and (3) biotic effects of saprotrophic soil communities that can perform different functional roles in the soil food web, or that may have specialized interactions with litter types, thereby altering soil nutrient cycling. We assess the role of litter in PSF effects via physical, chemical and biotic pathways to address how litter-mediated feedbacks may play out relative to, and in interaction with, feedbacks mediated through the plant rhizosphere. We also present one of the first experimental studies to show the occurrence and species-specificity of litter-mediated feedbacks and we identify critical research gaps. By formally incorporating the plant-litter feedback pathway into PSF experiments, we will further our understanding of PSFs under natural conditions.

Abstract1. Below-ground nematodes are important for soil functioning, as they are ubiquitous and operate at various trophic levels in the soil food web. However, morphological nematode community analysis is time consuming and requires ample training. qPCR-based nematode identification techniques are well available, but high-throughput sequencing (HTS) might be more suitable for non-targeted nematode community analyses.2. We compared effectiveness of qPCR-and HTS-based approaches with morphological nematode identification while examining how climate warming-induced plant range expansion may influence below-ground nematode assemblages. We extracted nematodes from soil of Centaurea stoebe and C. jacea populations in Slovenia, where both plant species are native, and Germany, where C. stoebe is a range expander and C. jacea is native. Half of each nematode sample was identified morphologically and the other half was analysed using targeted qPCR and a novel HTS approach.3. HTS produced the highest taxonomic resolution of the nematode community.Nematode taxa abundances correlated between the methods. Therefore, especially relative HTS and relative morphological data revealed nearly identical ecological patterns. All methods showed lower numbers of plant-feeding nematodes in rhizosphere soils of C. stoebe compared to C. jacea. However, a profound difference was observed between absolute and relative abundance data; both sampling origin and plant species affected relative abundances of bacterivorous nematodes, whereas there was no effect on absolute abundances.4. Taken together, as HTS correlates with relative analyses of soil nematode communities, while providing highest taxonomic resolution and sample throughput, we propose a combination of HTS with microscopic counting to supplement important quantitative data on soil nematode communities. This provides the most cost-effective, in-depth methodology to study soil nematode communityThis is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Current climate change has led to latitudinal and altitudinal range expansions of numerous species. During such range expansions, plant species are expected to experience changes in interactions with other organisms, especially with belowground biota that have a limited dispersal capacity. Nematodes form a key component of the belowground food web as they include bacterivores, fungivores, omnivores and root herbivores. However, their community composition under climate change‐driven intracontinental range‐expanding plants has been studied almost exclusively under controlled conditions, whereas little is known about actual patterns in the field. Here, we use novel molecular sequencing techniques combined with morphological quantification in order to examine nematode communities in the rhizospheres of four range‐expanding and four congeneric native species along a 2,000 km latitudinal transect from South‐Eastern to North‐Western Europe. We tested the hypotheses that latitudinal shifts in nematode community composition are stronger in range‐expanding plant species than in congeneric natives and that in their new range, range‐expanding plant species accumulate fewest root‐feeding nematodes. Our results show latitudinal variation in nematode community composition of both range expanders and native plant species, while operational taxonomic unit richness remained the same across ranges. Therefore, range‐expanding plant species face different nematode communities at higher latitudes, but this is also the case for widespread native plant species. Only one of the four range‐expanding plant species showed a stronger shift in nematode community composition than its congeneric native and accumulated fewer root‐feeding nematodes in its new range. We conclude that variation in nematode community composition with increasing latitude occurs for both range‐expanding and native plant species and that some range‐expanding plant species may become released from root‐feeding nematodes in the new range.

Climate change is accelerating the spread of plants and their associated species to new ranges. The differences in range shift capacity of the various types of species may disrupt long-term co-evolved relationships especially those belowground, however, this may be less so for seed-borne endophytic microbes. We collected seeds and soil of the range-expanding Centaurea stoebe and the congeneric Centaurea jacea from three populations growing in Slovenia (native range of both Centaurea species) and the Netherlands (expanded range of C. stoebe, native range of C. jacea). We isolated and identified endophytic fungi directly from seeds, as well as from roots of the plants grown in Slovenian, Dutch or sterilized soil to compare fungal endophyte composition. Furthermore, we investigated whether C. stoebe hosts a reduced community composition of endophytes in the expanded range due to release from plant-species specific fungi while endophyte communities in C. jacea in both ranges are similar. We cultivated 46 unique and phylogenetically diverse endophytes. A majority of the seed endophytes resembled potential pathogens, while most root endophytes were not likely to be pathogenic. Only one endophyte was found in both roots and seeds, but was isolated from different plant species. Unexpectedly, seed endophyte diversity of southern C. stoebe populations was lower than of populations from the north, while the seed endophyte community composition of northern C. stoebe populations was significantly different southern C. stoebe as well as northern and southern C. jacea populations. Root endophyte diversity was considerably lower in C. stoebe than in C. jacea independent of plant and soil origin, but this difference disappeared when plants were grown in sterile soils. We conclude that the community composition of fungal endophytes not only differs between related plant species but also between populations of plants that expand their range compared to their native habitat. Our results suggest that fungal endophytes of two Centaurea species are not able to systemically infect plants. We highlight that endophytes remain poorly studied and further work should investigate the functional importance of endophytes.

Current climate warming enables plant species and soil organisms to expand their range to higher latitudes and altitudes. At the same time, climate change increases the incidence of extreme weather events such as drought. While it is expected that plants and soil organisms originating from the south are better able to cope with drought, little is known about the consequences of their range shifts on soil functioning under drought events.Here, we test how range‐expanding plant species and soil communities may influence soil functioning under drought. We performed a full‐factorial outdoor mesocosm experiment with plant communities of range expanders or related natives, with soil inocula from the novel or the original range, with or without summer drought. We measured litter decomposition, carbon mineralization and enzyme activities, substrate‐induced respiration and the relative abundance of soil saprophytic fungi immediately after drought and at 6 and 12 weeks after rewetting.Drought decreased all soil functions regardless of plant and soil origin except one; soil respiration was less reduced in soils of range‐expanding plant communities, suggesting stronger resistance to drought. After rewetting, soil functioning responses depended on plant and soil origin. Soils of native plant communities with a history of drought had more litter mass loss and higher relative abundance of saprophytic fungi than soils without drought and soils of range expanders. Functions of soil from range expanders recovered in a more conservative manner than soils of natives, as litter mass loss did not exceed the control rates. At the end of the experiment, after rewetting, most soil functions in mesocosms with drought history did not differ anymore from the control.We conclude that functional consequences of range‐expanding plants and soil biota may interact with effects of drought and that these effects are most prominent during the first weeks after rewetting of the soil. A free http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13453/suppinfo can be found within the Supporting Information of this article.

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Microbial communities drive plant litter breakdown. Litters originating from different plant species are often associated with specialised microbiomes that accelerate the breakdown of that litter, known as home‐field advantage. Yet, how and how fast microbial communities specialise towards litter inputs is not known. Here we study effects of repeated litter additions on soil microbial community structure and functioning. We set up a 9‐month, full‐factorial, reciprocal litter transplant experiment with soils and litters from six plant species (three grasses and three trees). We measured fungal and bacterial community composition, litter mass loss and home‐field effects. We found that repeated litter additions resulted in convergence in fungal community composition driven by litter functional group (trees vs. grasses). Grasses enriched Sordariomycetes, while Tremellomycetes, Eurotiomycetes and Leotiomycetes were favoured by tree litter. Bacterial community composition, litter mass loss and home‐field effects were not affected by litter incubation, but there was a relationship between fungal community composition and mass loss. We conclude that repeated litter incubations can result in directional shifts in fungal community composition, while 9 months of litter addition did not change bacterial community composition and the functioning and specialisation of microbial communities. Testing further how repeated litter inputs affect microbial functioning is essential for steering decomposer communities for optimal soil carbon and nutrient cycling.