The loss of biodiversity can compromise the functioning of ecosystems and services that the society depends upon (Tilman and Downing, 1994; Hector and Bagchi, 2007; Cardinale, 2012; Hooper et al., 2012). For example, evidence is mounting that the reduction in species diversity can reduce multiple ecosystem functions, such as processes that drive nutrient and carbon cycling and plant productivity (Zavaleta et al., 2010; Maestre et al., 2012; Hautier et al., 2018). Reduced biodiversity has also been shown to destabilize ecosystem functioning over time, and thus the maintenance of biodiversity is critical to long-term ecosystem sustainability (Tilman et al., 2006; Hector et al., 2010; Isbell, 2011; Hautier, 2015). Experiments with plant communities have shown that species diversity can support multiple ecosystem functions simultaneously because plant species that do not contribute to that function at a particular time may contribute at another time, or to another function at the same time. Thus, the asynchronous temporal fluctuations among species in their abundance and contribution to various ecosystem functions in more diverse communities contribute to a greater overall ecosystem functioning over time (Hautier et al., 2018; Isbell, 2011; Yachi and Loreau, 1999).

While the reduction of plant diversity is well known to destabilize ecosystem functioning, effects of soil biodiversity loss on stabilizing ecosystem functioning are still poorly understood. A number of studies have shown that soil biodiversity is rapidly declining in intensively managed soils (Helgason et al., 1998; Birkhofer, 2008; Verbruggen, 2010; Tsiafouli et al., 2015), and one-quarter of soils worldwide are now degraded, causing a reduction of biological productivity (Stavi and Lal, 2015). Thus, understanding the consequences of soil biodiversity loss is crucial, as there is a growing consensus that soil biota play key roles in ecosystems (Bardgett and van der Putten, 2014; van der Heijden et al., 2008; Delgado-Baquerizo et al., 2016) and support a number of ecosystem services, including food production and nutrient cycling (Brussaard et al., 2007; Bender et al., 2016). Several studies have demonstrated that the composition and richness of soil microbiota can predict multiple ecosystem functions, such as plant diversity and productivity, soil carbon assimilation, and nutrient cycling (Bradford, 2014; Wagg et al., 2014; Jing et al., 2015; Delgado-Baquerizo et al., 2020; Mori et al., 2016; Wagg et al., 2019). However, few studies have linked soil microbiota communities to the temporal stability of plant community composition and productivity (Eisenhauer et al., 2012; Yang et al., 2014; Pellkofer et al., 2016).

The compositions of soil bacterial and fungal communities are highly dynamic over short time scales, such as months and growing seasons, and temporal changes in composition are further altered by land management practices such as tillage and crop rotation (Lauber et al., 2013; Coudrain et al., 2016; Wagg et al., 2018). Whether these temporal dynamics in microbial communities lead to temporal fluctuations in ecosystem functions and if microbial community dynamics are inhibited due to soil microbial biodiversity loss have not been assessed yet (Bardgett and van der Putten, 2014). Past studies have shown that some soil functions do not track soil microbial loss, or they recover rapidly after perturbations of soil microbial communities. This suggests that such functions may either be highly resilient or contain many functionally redundant taxa (Griffiths et al., 2000; Fitter et al., 2005; Allison and Martiny, 2008). However, functional redundancy is likely to fade as multiple time points and ecosystem functions are considered (Hector and Bagchi, 2007; Isbell, 2011; Hautier et al., 2018). Considering that soil biodiversity supports numerous ecosystem functions, we hypothesize that greater soil biodiversity would also maintain a greater and less variable ecosystem functioning through time and would thus stabilize multiple ecosystem functions. Specifically, microbial taxa may fluctuate asynchronously in abundance through time such that different taxa support different functions at different times. This should provide insurance that some microbial taxa that support an ecosystem function will be present at any given time to stabilize the functioning of the ecosystem (Isbell, 2011; Yachi and Loreau, 1999; Mori et al., 2016). Therefore, we predict that temporal asynchrony among taxa in diverse soil microbial communities should relate to greater temporal stability in multiple ecosystem functions (Loreau and de Mazancourt, 2008; Thibaut and Connolly, 2013; Gross et al., 2014).

Here, we decreased soil diversity by filtering out soil organisms based on size along a gradient to assess the effects of soil microbial communities on stabilizing four ecosystem functions. The exclusion of organisms based on size can lead to a functional simplification of soil communities because body size is directly associated with trophic guilds, metabolic rates, population density, and generational turnover (Coudrain et al., 2016; Bradford, 2002; Yodzis and Innes, 1992; Wall and Moore, 1999; Woodward, 2005). Furthermore, the size-based reduction of soil organisms parallels the impact of land management practices, such as soil tillage, that physically damage soil organisms depending on their size and, thus, also disrupts the community structure of soil biota (Wagg et al., 2018; Jansa et al., 2003; Köhl et al., 2014; Postma-Blaauw et al., 2010).

Using soil sieving gradient (5000, 100, 25, and 0 µm [sterile]), we created a monotonically declining gradient in fungal and bacterial richness among replicated soil-plant ecosystems established in self-contained mesocosms (F_{1, 46} = 172.0, p<0.001; and F_{1, 45.7} = 753.9, p<0.001, respectively, for fungal and bacterial richness decline; Figure 1—figure supplement 1, Supplementary file 1 – table 1). The decline in diversity over our experimental gradient was maintained throughout the experimental duration (time [factor] by sieve gradient (log-linear) interaction: F_{4, 170.1} = 0.75, p=0.561; and F_{4, 177.1} = 1.60, p=0.174, respectively, for fungi and bacteria; Figure 1—figure supplement 1, Supplementary file 1- table 1). This decline in bacterial richness along the four-step gradient was steepest for the final step from smallest sieve size to the sterile soil treatment), which resulted in a 60% loss of bacterial and 55% loss of fungal richness, compared to the highest soil biodiversity treatment. Community composition differed among the soil diversity treatments and the different time points, for both fungal and bacterial communities (Figure 1—figure supplement 1 and 2, Supplementary file 1 – table 2). All four ecosystem functions (plant biomass production, plant diversity, litter decomposition, and soil carbon assimilation) changed significantly over our soil biodiversity treatment gradient and varied significantly through time depending on the soil biodiversity level (Figure 1a–d, Supplementary file 1 – tables 3 and 4). Bacterial and fungal richness and community composition were significantly related to all four ecosystem functions, depending on the time, and differences in microbial community composition were all correlated with differences in ecosystem functions at all time points (Figure 1—figure supplement 3–5). These results demonstrate that we successfully established a broad gradient of soil microbial diversity, composition, and ecosystem functioning. This enabled us to assess effects of the soil microbial community on the temporal stability of ecosystem functioning, which we calculated as the inverse of the coefficient of variation of ecosystem functions across all five time points (Tilman et al., 2006; Pimm, 1984; Tilman, 1999).

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We found that greater bacterial and fungal richness supported higher stability of two aboveground ecosystem functions, plant productivity and plant diversity (Figure 1e). Plant biomass production was particularly strongly destabilized in our sterile treatment with the lowest soil microbial diversity, exhibiting a ‘boom and bust’ temporal response, where plant biomass was the lowest of all treatments after 11 weeks, then the highest at 22 weeks and steadily declined afterwards to substantially lower values than in all other treatments by 55 weeks (Figure 1a). The effects of microbial diversity on the stability of plant biomass production were also significant when the sterile treatment was not considered, showing that microbial diversity is also important for plant community stability even at higher levels of microbial diversity. All mesocosms were initially planted with the same plant community and plant diversity increased steadily over time as communities developed. This occurred in all but the sterile soil treatment, which remained low throughout the experiment. This resulted in the positive effect of microbial diversity on stabilizing plant diversity (Figure 1b).

The effect of soil microbial diversity on stabilizing plant biomass and diversity resulted from its influence on the changes in plant community composition. In the sterile and low soil biodiversity treatments, plant communities were dominated by the grass Lolium perenne (Figure 1—figure supplement 6) which drove the ‘boom and bust’ trend in biomass production (Figure 1a). In contrast, the biomass production by the legume Trifolium pratense increased steadily over time and was highest in the highest soil biodiversity treatment, contributing to the greater stability in plant diversity and productivity. The sterile treatment could account for the effect of microbial diversity on supporting more stable plant diversity, as well as the bacterial richness-biomass stability relationship, but not the stabilizing effect that fungal richness had on plant biomass production (Supplementary file 1 – table 5).

Fungal richness was significantly related to greater stability in litter decomposition (Figure 1e), and soil carbon assimilation was positively related to greater bacterial richness (Figure 1e). Using an averaging multifunctionality approach, where multifunctionality is calculated as the standardized average of all ecosystem functions (Maestre et al., 2012; Byrnes, 2014), we found that greater microbial diversity (richness of bacteria, fungi, and their scaled average) all showed strong positive relationships with the ability to maintain greater and more stable ecosystem functioning on average (Figure 1f). Using a multiple threshold approach (Byrnes, 2014) to characterize how many functions achieve an increasing threshold of stabilization with greater diversity, we found that soil microbial diversity had a significant stabilizing effect on multiple functions at thresholds above 30% of the maximum stability value for each function (Figure 2a). At this threshold, more than three functions had high level of temporal stability (Figure 2b), showing that high soil biodiversity can promote the stability of multiple functions simultaneously. This stabilizing effect increased to become strongest at a threshold of 72%. Soil bacterial richness had a significant stabilizing effect on multiple functions at low threshold values of 29% and was strongest at 68% (Figure 2c). Similarly, soil fungal richness had a positive effect on the stability of multiple functions at 30% and had the strongest effect on supporting stability in multiple functions at 82% of the maximum observed stability (Figure 2d).

Figure 2

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Since greater plant diversity is often associated with greater plant biomass production (Tilman and Downing, 1994; Hector and Bagchi, 2007; Cardinale, 2012; Hooper et al., 2012), and biomass production with soil carbon assimilation (Lange et al., 2015; Yang et al., 2019), we assessed how soil microbial diversity may have indirectly influenced the temporal stability of plant biomass production through its strong relationship with plant diversity using multi-model comparisons and structural equation modeling (SEM). We found that soil microbial diversity alone was the most parsimonious explanatory variable of plant biomass stability compared with models that included plant diversity (Supplementary file 1 – table 6). Further, using SEM we found that soil microbial diversity reduced the temporal variation and increased the temporal mean in biomass production, while plant diversity had no detectable effect (Figure 3a). We also found that the temporal variation had a much stronger influence compared with the temporal mean on the stability of biomass production (Figure 3b). Comparing multiple models using soil microbial diversity, plant diversity, plant biomass, and the stability in plant biomass to predict the stability in soil carbon assimilation, we found that stability in soil carbon assimilation was best explained by the stability in plant biomass production alone (Supplementary file 1 – table 7). Therefore, the effect of soil microbial diversity on stabilized soil carbon assimilation was likely indirect through its stabilizing effect on biomass production (Figure 3a). Specifically, microbial diversity reduced the temporal variation in biomass production, thus increasing its stability. In turn, greater stability in biomass production reduced the temporal variation in soil carbon assimilation, thus stabilizing soil carbon assimilation (Figure 3b).

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To identify microbial taxa that may influence an ecosystem function at a given time, we used a randomization approach (Mori et al., 2016; Gotelli et al., 2011) to calculate the standardized effect size (SES) of each fungal and bacterial taxon on each ecosystem function. This method generates a distribution for the null hypothesis that the effect of the presence of a taxon on a given ecosystem function at any given time occurs by chance. The influence of a taxon on an ecosystem service at a given time was considered to be significant when the effect of the presence of the taxon on an ecosystem function was greater than 1.96 (the 95 percentile of the t-distribution) times the SD of the null distribution. As we considered more time points, we detected a greater proportion of microbial taxa that were positively, or negatively, related to supporting an ecosystem function as more time points were considered (Figure 4a, see Supplementary file 2 for a summary for their taxonomic assignment). In addition, we found that the proportion of soil taxa with positive or negative associations with ecosystem functioning increased as more functions as well as more time points were independently considered (Figure 4), demonstrating that different microbial taxa are important for different ecosystem functions at different times. These relationships were found for bacterial and fungal taxa combined (Figure 4a,b), as well as for bacterial (Figure 4c,e) and fungal taxa (Figure 4d,f) considered separately. Moreover, we found a larger cumulative proportion of soil taxa with positive than negative effects on ecosystem functioning with increasing number of functions and times considered (black versus grey lines in Figure 4).

Figure 4

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Of the microbes that were found to have a significant positive association with a function at any given time, we found that the average temporal stability in the abundance of these fungi and bacteria (species stability CV_species⁻¹) was positively related to the temporal stability of that function, with the exception of soil carbon assimilation (Figure 5). In other words, the more stable the abundance of the taxa that support an ecosystem function at any time, the greater the stability of that ecosystem function (see Supplementary file 1–tables 9 and 10 for the taxonomic assignment of the taxa that were significantly related to functions at more than one time point). These species stability-ecosystem function stability relationships could be explained by the most extreme soil biodiversity level (the ‘sterile treatment’, Supplementary file 1 – table 8). We also found that for all four ecosystem functions, a lower temporal synchrony among those taxa that had a positive association with a given function at any given time point also significantly related to a greater stability of that ecosystem function (Figure 5). These species asynchrony-ecosystem function stability relationships were largely independent of the most extreme sterile soil biodiversity level. After first accounting for the sterile soil biodiversity level, the residual effect of microbial asynchrony on plant diversity and litter decomposition stability remained significant (p=0.036 and 0.048, respectively) and the effect on litter decomposition stability was marginally non-significant (p=0.058, Supplementary file 1 – table 9).

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Here, we demonstrate that that the maintenance of greater soil microbial diversity not only supports ecosystem functions but also stabilizes multiple ecosystem functions simultaneously. This is supported by our analysis of individual functions, the average response in stability of all four functions (Maestre et al., 2012), and the multiple threshold approach (Byrnes, 2014). Our results show that both fungal and bacterial richness support greater stability in multiple functions simultaneously with the strongest effect of microbial richness on stability occurring when functions need to achieve at least 70% of their maximum observed stability. The effect of soil biodiversity loss on multifunctional stability was strongest in our most extreme ‘sterile’ treatment, suggesting a loss of microbial richness over 50% as a potential tipping point over which the stabilizing effect of soil microbial diversity might drop abruptly.

We found that the stabilizing effect of a diverse soil microbial community is attributed to the asynchronous temporal dynamics in the soil microbial community composition. Despite the strong contribution of the sterile soil treatment to the results, we still found the effects of the composition and temporal asynchrony to be important for stabilizing ecosystem functions even after accounting for our most extreme treatment. To our knowledge, our results demonstrate for the first time that the maintenance of a more diverse soil microbial community allows for greater asynchronous temporal dynamics in the microbial communities to result in the maintenance of higher nutrient and carbon cycling processes through time (reflected here by four key ecosystem functions). This implies that different microbes can complement each other by functioning at different times to provide a stable support for ecosystem functions.

Asynchrony in microbial communities may arise as microbial community compositional changes track the phenology of the associated plant communities through plant nutrient uptake, root exudates, and litter inputs that generate changes in soil chemistry and resources for soil microbes (Marschner et al., 2002; Bray et al., 2012; Buscardo et al., 2018; Upton and Hofmockel, 2019). For instance, different N-cycling microbes exhibit temporal seasonal dynamics as they are active at different times to contribute to nitrification and denitrification processes that are also linked to plant community development (Prosser and Nicol, 2012; Regan, 2017). Additionally, microbial community composition changes during decomposition as microbes have different resource acquisition strategies where some fast-cycling microbes acquire readily available carbon early, while slow-cycling microbes decompose recalcitrant carbon at later stages (Herzog, 2019; Alavoine and Bertrand, 2020; Su et al., 2020). Thus, by having a greater diversity of microbes there is a greater opportunity for different microbes to perform specific functions associated with the different stages of litter decomposition, nutrient and carbon cycling that together maintain the broad ecosystem functions of plant productivity, diversity, decomposition, and carbon assimilation through time.

Past studies assessing soil microbial diversity-ecosystem functioning relationships have shown that the suppression of soil microbial diversity and abundance can have little impact on particular functions measured at a single time point, possibly due to high redundancy among taxa (Griffiths et al., 2000; Fitter et al., 2005; Allison and Martiny, 2008; Louca et al., 2018). However, here we show that as more functions and time points are considered, a greater proportion of the microbial community is required to support these functions and correspondingly redundancy, or the proportion of microbes not associated with ecosystem functions, fades away. This is supported by the consistent trend that a greater number of taxa were found to positively support ecosystem functions when more functions and time points were considered, highlighting that increasing the number of taxa ensures the maintenance of the functioning of the overall ecosystem over time (Isbell, 2011; Hautier, 2015; Yachi and Loreau, 1999). This suggests that whereas a large number of soil microbial taxa may overlap in supporting a single function at a single time point, fewer and fewer taxa may overlap in supporting multiple functions over multiple time points. Similar to studies assessing the functional redundancy of plant communities (Hautier et al., 2018; Isbell, 2011), as more ecosystem functions and times were considered in the present study, more soil taxa were found to affect the overall functioning of the ecosystem. While high soil microbial diversity may appear functionally redundant for a given broad function, such as microbial respiration and decomposition (Schimel and Schaeffer, 2012), there are studies demonstrating low functional redundancy in soils in both broad functions (soil respiration) and specialized functions (degradation of specific organic compounds such as cellulose and microbial toxins)(Wohl et al., 2004; DelgadoBaquerizo, 2016). Here we found, for instance, that the mycorrhizal fungi Diversispora was frequently associated with positive effects on biomass production, but not other functions, while the plant pathogen Fusarium was frequently found to positively influence plant diversity and no other functions (see Supplementary file 2 – table 2). Such findings are logical as mycorrhiza are well known to drive plant biomass production (e.g. van der Heijden et al., 2008) and pathogens can maintain plant diversity (Petermann et al., 2008; Schnitzer et al., 2011).

It is important to note that the composition and diversity of other key groups of soil biota, including protozoa and nematodes not assessed in our study were likely also affected by the sieving treatments and may also have contributed to the observed effect of our soil biodiversity measures on ecosystem functioning and stability either directly or through trophic interactions with soil microbes (de Vries and Shade, 2013; Bradford, 2002; Trap et al., 2016; Thakur and Geisen, 2019). Assuming this was the case, this would suggest that the broader spectrum of soil biodiversity beyond the soil microbiome may also hold a key role in promoting multiple ecosystem functions (Delgado-Baquerizo et al., 2016; Bradford, 2014; Wagg et al., 2014; Jing et al., 2015; Delgado-Baquerizo et al., 2020; Mori et al., 2016; Wagg et al., 2019), their temporal stability and by extension possibly their resistance and resilience to environmental perturbation (Griffiths et al., 2000; Delgado-Baquerizo et al., 2017; Griffiths and Philippot, 2013). Nevertheless, the effects of soil biodiversity loss and the role of functional redundancy in the soil microbiome for stabilizing soil functions have yet to be fully elucidated in the context of the complexity of interactions among the various trophic levels of soil organisms (Soliveres et al., 2016).

Studies focussing on plant communities have demonstrated the importance of plant diversity for promoting biomass production (Cardinale, 2012; Hooper et al., 2012) and the temporal stability in biomass production (Tilman and Downing, 1994; Hector and Bagchi, 2007; Hautier et al., 2018), as well as building soil carbon pools (Lange et al., 2015; Yang et al., 2019). In our model system, we were able to reveal that soil microbial diversity had a direct positive effect on the stability of biomass production that in turn stabilized soil carbon assimilation. Specifically, soil microbial diversity supported a higher stability in biomass production due to its strong effect on lowering the temporal variation in plant biomass production, which was independent of plant diversity. While experimental manipulations of plant diversity are well known to enhance plant productivity (Tilman and Downing, 1994; Cardinale, 2012; Hooper et al., 2012; Tilman et al., 2006; Hector et al., 2010; Isbell, 2011; Hautier, 2015), the lack of a direct effect of plant diversity on the stability of biomass production and carbon in our experiment may be not surprising, because all our treatments started off with the same plant diversity while soil communities were experimentally manipulated. At the lowest levels of soil biodiversity, we found that plant productivity initially increased over time and was the most productive. This was driven by the rapid growth of L. perenne grass, which is highly competitive for soil nitrogen and is a plant species that is hardly affected by changes in arbuscular mycorrhizal diversity when grown alone (Wagg et al., 2011a; Wagg et al., 2011b). Its initial high productivity may be due to initial release from pathogens and plant competition at low levels of soil biodiversity. As time progressed, the productivity of this highly productive grass was unsustainable, and the productivity crashed compared with our more diverse treatments, resulting in the ‘boom and bust’ temporal trend. We attribute this decline in productivity at the lowest level of soil biodiversity to the inability of the soil community to maintain the presence and growth of legumes and impaired nutrient availability and reduced nutrient cycling processes needed to maintain higher plant community productivity (Wagg et al., 2019; Roscher, 2013; Marquard, 2009; Schmidtke et al., 2010).

In contrast to the stability in biomass production, we found that the effect of soil microbial diversity on stabilizing soil carbon assimilation was indirect through its effect on promoting the stability in plant biomass production. This is logical since the primary path for atmospheric carbon to become assimilated into soils is through photosynthesis, upon which plant biomass production is based, that is then transferred to soil microbes symbiotically or through the microbial accumulation of plant-derived carbon through decomposition processes (De Deyn et al., 2008; De Deyn et al., 2011; Merino et al., 2015). Together with our findings, these results provide support for the concept that there is a tight link between the diversity and composition of the soil microbiome belowground and the biomass production of plants aboveground that together promote and stabilize soil carbon assimilation.

Earlier studies showed that the diversity and composition of the soil microbiome has a big impact on ecosystem functioning. Yet it had not been shown previously whether altered temporal variations in community composition and biodiversity in the soil microbiome impairs the stability of soil functions. Our study provides a first proof of concept that severe degradation of belowground biological communities reduces multiple ecosystem functions and their stability over time with a likely tipping point (>50% microbial biodiversity loss). Moreover, our work highlights that asynchrony of different microbes can improve the stability of ecosystem functioning over time. Our result is concerning as soil biodiversity is being lost in many parts of the world in response to anthropogenic disturbances and increasing land-use intensification compounded by effects of climate change (Helgason et al., 1998; Birkhofer, 2008; Verbruggen, 2010; Tsiafouli et al., 2015; Stavi and Lal, 2015). Such biological degradation of soils may likely have cascading effects on their ability to maintain diverse productive plant communities that promote carbon sequestration. Our study shows that this is because there are fewer microbes that are needed to support the cycling of soil nutrients and plant growth at different times throughout the development and growth of plant communities. Consequently, anthropogenic activities that simplify soil diversity are likely to diminish the reliable provisioning of essential ecosystem services.

All data generated or analysed during this study are available online at: https://figshare.com/s/790d5ba1fb7f8e670a27.

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Author details

1. Cameron Wagg

   1. Department of Evolutionary Biology and Environmental Studies, University of Zürich, Zürich, Switzerland
   2. Plant-Soil Interactions, Research Division Agroecology and Environment, Agroscope, Zürich, Switzerland
   3. Fredericton Research and Development Centre, Agriculture and Agri-Food Canada, Fredericton, Canada

   Contribution

   Conceptualization, Data curation, Formal analysis, Supervision, Visualization, Writing - original draft, Writing - review and editing

   Contributed equally with

   Yann Hautier and Sarah Pellkofer

   For correspondence

   cameron.wagg@canada.ca

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-9738-6901

2. Yann Hautier

   Ecology and Biodiversity Group, Department of Biology, Utrecht University, Padualaan, Netherlands

   Contribution

   Conceptualization, Formal analysis, Methodology, Writing - original draft, Writing - review and editing

   Contributed equally with

   Cameron Wagg and Sarah Pellkofer

   Competing interests

   No competing interests declared

3. Sarah Pellkofer

   1. Department of Evolutionary Biology and Environmental Studies, University of Zürich, Zürich, Switzerland
   2. Plant-Soil Interactions, Research Division Agroecology and Environment, Agroscope, Zürich, Switzerland

   Contribution

   Data curation, Investigation, Methodology, Writing - review and editing

   Contributed equally with

   Cameron Wagg and Yann Hautier

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-0208-0350

4. Samiran Banerjee

   1. Plant-Soil Interactions, Research Division Agroecology and Environment, Agroscope, Zürich, Switzerland
   2. Department of Microbiological Sciences, North Dakota State University, Fargo, United States

   Contribution

   Formal analysis, Methodology, Writing - review and editing

   Competing interests

   No competing interests declared

5. Bernhard Schmid

   1. Department of Evolutionary Biology and Environmental Studies, University of Zürich, Zürich, Switzerland
   2. Department of Geography, Remote Sensing Laboratories, University of Zürich, Zürich, Switzerland

   Contribution

   Supervision, Funding acquisition, Writing - review and editing

   Competing interests

   Reviewing editor, eLife

   "This ORCID iD identifies the author of this article:" 0000-0002-8430-3214

6. Marcel GA van der Heijden

   1. Department of Evolutionary Biology and Environmental Studies, University of Zürich, Zürich, Switzerland
   2. Plant-Soil Interactions, Research Division Agroecology and Environment, Agroscope, Zürich, Switzerland
   3. Department of Plant and Microbial Biology, University of Zürich, Zürich, Switzerland

   Contribution

   Conceptualization, Supervision, Funding acquisition, Writing - review and editing

   Competing interests

   No competing interests declared

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