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1. Plant–soil feedback (PSF), plant trait and functional group concepts advanced our understanding of plant community dynamics, but how they are interlinked is poorly known. 2. To test how plant functional groups (FGs: graminoids, small herbs, tall herbs, legumes) and plant traits relate to PSF, we grew 48 grassland species in sterilized soil, sterilized soil with own species soil inoculum and sterilized soil with soil inoculum from all species, and quantified relative growth rate (RGR), specific leaf area (SLA), specific root length (SRL) and per cent arbuscular mycorrhizal fungi colonization (%AMF). 3. Plant growth response to the plant species' own soil biota relative to sterilized soil (PSFsterilized) reflects net effects of all (generalist + specialized) soil biota. Growth response to the plant species' own soil biota relative to soil biota of all plant species (PSFaway) reveals effects of more specialized soil organisms. 4. PSFsterilized showed that graminoids and small herbs have a negative and tall herbs a positive response to their own soil biota, whereas legumes responded neutrally. However, PSFaway showed that on average, all plant FGs benefitted from growing with other species' soil biota, suggesting that pathogens are more specialized than plant growth-promoting soil biota. Feedback to plant growth from all soil biota (PSFsterilized) was stronger than from more specialized soil biota (PSFaway) and could be predicted by SRL and especially by %AMF colonization. Species with high SRL and low %AMF colonization when grown in away soil experienced most negative soil feedback. 5. Synthesis. Plant species from all plant FGs grow better in soil from other species because of less net negative effects of soil biota (in graminoids), or because of more net positive soil biota effects (in tall herbs). Explorative plant species (high SRL, low %AMF colonization) suffer most from negative feedback of all soil biota, whereas more resource conservative species (low SRL, high %AMF colonization) benefit from soil feedback of all soil biota. These findings help to understand replacement of explorative species during succession. Moreover, we suggest a potentially larger role for species with positive feedback than for species with negative feedback to contribute to maintain plant community productivity of diverse communities over time.

1. Plants influence associated soil biotic communities that in turn can alter the performance of the subsequently growing plants. Although such \"plant-soil feedbacks\" (PSFs) are considered as important drivers of plant community assembly, past PSF studies have mainly addressed plant biomass production. However, plant performance is not only the production of biomass but comprises a sequence of different life stages: from seed germination over vegetative growth up to the production of a viable progeny. 2. Here, we assessed the effects of soil biotic communities that were previously conditioned for 3 years by a focal plant species monoculture or species mixtures on key plant life stages from germination and vegetative growth to flowering and the production of viable seeds. We used three common grassland herb species that were grown in a sterile substrate and inoculated with a sterile control soil or with living soils. Living soils were conditioned either by the focal species in monoculture or a four- or eight-species mixture that included the focal species to represent a decrease in the target plants' conspecific influence on the soil communities. 3. We show that the effect of soil biota changed from positive at the plants' juvenile life stages to neutral or negative at the plants' adult life stages and ultimately decreased plant fitness. A higher conspecific influence on the soil communities pronounced the positive effects at the juvenile life stage but also the negative effects at adult life stages. Further, we observed direct soil biotic effects on flower production and plant fitness that were not mediated by adult biomass production. This suggests that soil biotic effects may alter plant resource allocation and even may have transgenerational effects on plant fitness. 4. Synthesis. We conclude that there is no overarching effect of soil biota that remains consistent at all the life stages of a plant. Thus, our results highlight the importance to consider plant life stage and ultimately plant fitness especially when plant-soil interactions are used to explain plant community dynamics.

1. Understanding the importance of biotic interactions in driving the distribution and abundance of species is a central goal of plant ecology. Early vascular plants likely colonized land occupied by biocrusts — photoautotrophic, surface-dwelling soil communities comprised of cyanobacteria, bryophytes, lichens and fungi — suggesting biotic interactions between biocrusts and plants have been at play for some 2,000 million years. Today, biocrusts coexist with plants in dryland ecosystems worldwide, and have been shown to both facilitate or inhibit plant species performance depending on ecological context. Yet, the factors that drive the direction and magnitude of these effects remain largely unknown. 2. We conducted a meta-analysis of plant responses to biocrusts using a global data-set encompassing 1,004 studies from six continents. 3. Meta-analysis revealed there is no simple positive or negative effect of biocrusts on plants. Rather, plant responses differ by biocrust composition and plant species traits and vary across plant ontogeny. Moss-dominated biocrusts facilitated, while lichen-dominated biocrusts inhibited overall plant performance. Plant responses also varied among plant functional groups: C₄ grasses received greater benefits from biocrusts compared to C₃ grasses, and plants without N-fixing symbionts responded more positively to biocrusts than plants with N-fixing symbionts. Biocrusts decreased germination but facilitated growth of non-native plant species. 4. Synthesis. Results suggest that interspecific variation in plant responses to biocrusts, contingent on biocrust type, plant traits, and ontogeny can have strong impacts on plant species performance. These findings have important implications for understanding biocrust contributions to plant productivity and community assembly processes in ecosystems worldwide.

Enhanced deposition of atmospheric nitrogen (N) has profound impacts on ecosystem processes such as above‐ground productivity and community structure in grasslands across the globe. But how N deposition affects below‐ground processes of grasslands is less well known. Here, we evaluated the effects of chronic N amendment at a relatively low rate (20 kg ha⁻¹ year⁻¹) on root traits (root productivity, root biomass, root/shoot ratio) in Inner Mongolia steppes by rhizotron and ingrowth core and soil monolith techniques at levels of individual species, functional groups and ecosystem. For 8 years, N amendment suppressed above‐ground net primary production (ANPP), photosynthetic rates and root biomass of forbs, but enhanced ANPP and root biomass of grasses. This led to an overall reduction in below‐ground productivity of the grassland by 24–33%, while ANPP remained unchanged. Nitrogen amendment acidified soil and subsequently increased extractable soil manganese (Mn) concentration. Nitrogen amendment increased foliar Mn concentrations in forb, but not grass species, leading to a significant inhibition of photosynthetic rates in forb species. Synthesis. These findings highlight the importance of the differentiating responses of plant functional groups to long‐term N deposition and the important consequences of these responses for below‐ground productivity and long‐term soil C sequestration.

1. The study of coupled green and brown food webs has improved our ability to understand how nutrient cycling and plant communities respond to perturbations. Yet, it is still difficult to predict how rapidly and consistently changes in one food web propagate to the other. An area of particular uncertainty is the response of plants to the changes in leaf litter nitrogen release caused by herbivores and detritivores. 2. We combined a field experiment and theoretical analysis to assess how herbivory, fertilization, and detritivory changed leaf litter nitrogen release and plant growth. We produced leaf litter in greenhouse cages with a factorial combination of grassshopper herbivory and additional nitrogen fertilizer. Our experiment used the resulting ¹⁵N-labelled leaf litter to measure isopod litter consumption, litter nitrogen release, and plant nitrogen uptake in the field. We then used a dynamical systems model to distinguish whether plant growth relied (a) directly on litter nitrogen release or (b) on other nitrogen pathways so that changes in litter nitrogen release have little immediate impact. 3. Our empirical results show that the effect of nitrogen fertilization on isopod processing of leaf litter was stronger than the effect of herbivory. Regardless, the available soil nitrogen and plant growth were independent of litter nitrogen release. Our dynamical model attributes the independence to other nitrogen sources and sinks, which buffer the plant-available nitrogen pool. Isopods and litter with a history of herbivory reduced above-ground plant growth when we accounted for initial field mesocosm conditions, especially the abundance of the competitively dominant goldenrods. Consequently, the impact of isopods and litter traits do propagate to influence plant growth, but do not have a large enough effect to overcome initial differences in plant biomass and community composition in one year. 4. Synthesis. Our results indicate animals in green and brown food webs can alter plant litter nitrogen release without any significant nutrient recycling effect on plant growth. Considering the availability of external and soil-based nitrogen sources, as well as the current plant community and microbial biomass, may be critical to determining when plant growth will respond to animal-mediated changes in litter decomposition.

1. The abundance of nitrogen (N)-fixing plants in ecosystems where phosphorus (P) limits plant productivity poses a paradox because N fixation entails a high P cost. One explanation for this paradox is that the N-fixing strategy allows greater root phosphatase activity to enhance acquisition from organic sources, but evidence to support this contention is limited. 2. We measured root phosphomonoesterase (PME) activity of 10 N-fixing species, including rhizobial legumes and actinorhizal Allocasuarina species, and eight non-N-fixing species across a retrogressive soil chronosequence showing a clear shift from N to P limitation of plant growth and representing a strong natural gradient in P availability. 3. Legumes showed greater root PME activity than non-legumes, with the difference between these two groups increasing markedly as soil P availability declined. By contrast, root PME activity of actinorhizal species was always lower than that of co-occurring legumes and not different from non-N-fixing plants. 4. The difference in root PME activity between legumes and actinorhizal plants was not reflected in a greater or similar reliance on N fixation for N acquisition by actinorhizal species compared to cooccurring legumes. 5. Synthesis. Our results support the idea that N-fixing legumes show high root phosphatase activity, especially at low soil P availability, but suggest that this is a phylogenetically conserved trait rather than one directly linked to their N-fixation capacity.

Divergent hypotheses have been proposed that suggest plant invasions either enhance or degrade the mutualism between plants and arbuscular mycorrhizal (AM) fungi, but their relative support remains unknown. We conducted a meta‐analysis using 67 publications, involving 70 native and 55 invasive plant species to assess support for the enhanced mutualism hypothesis, the degraded mutualism hypothesis and an alternative hypothesis that factors other than invasive status (such as plant functional group) better predict AM function following invasion. We used multiple measurements to test these hypotheses: AM fungal colonization, growth responses to AM fungi and AM fungal‐mediated shifts in competitive interactions among native and invasive plants. Additionally, we assessed whether invasive plants alter AM associations in native plants and whether native and invasive plants host different AM fungal abundances and communities. Arbuscular mycorrhizal fungal colonization (%) and average growth responses did not differ between native and invasive plants. However, growth responses (±) were dampened among invasive plants, and the positive correlation between AM fungal colonization and growth response in native plants was absent in invasive plants. Rather than plant invasive status, plant functional group was a significant explanatory factor; forbs were generally more colonized and exhibited positive growth responses (when grown alone and in competition), whereas grass responses were neutral to negative. Arbuscular mycorrhizal fungal abundance (measured by percentage colonization, extraradical hyphal and spore densities, as well as neutral lipid fatty acid and glomalin concentrations) did not differ between native and invasive plants, but invasive plants hosted different AM fungal communities in 78% of studies. AM fungal colonization of native plants was lower when grown with, or after, invasive plants, likely due to the prevalence of non‐mycorrhizal plants in studies of neighbour and legacy effects. Synthesis. Neither the degraded nor the enhanced mutualism hypothesis was supported, suggesting that invasions do not select for directional shifts in AM associations. Instead, our results indicate that AM fungi are most likely to influence invasion trajectories when native and invasive plants belong to different functional groups.

The storage of carbon (C) and nitrogen (N) in soil is important ecosystem functions. Grassland biodiversity experiments have shown a positive effect of plant diversity on soil C and N storage. However, these experiments all included legumes, which constitute an important N input through N₂‐fixation. Indeed, the results of these experiments suggest that N₂ fixation by legumes is a major driver of soil C and N storage. We studied whether plant diversity affects soil C and N storage in the absence of legumes. In an 11‐year grassland biodiversity experiment without legumes, we measured soil C and N stocks. We further determined above‐ground biomass productivity, standing root biomass, soil organic matter decomposition and N mineralization rates to understand the mechanisms underlying the change in soil C and N stocks in relation to plant diversity and their feedbacks to plant productivity. We found that soil C and N stocks increased by 18% and 16% in eight‐species mixtures compared to the average of monocultures of the same species, respectively. Increased soil C and N stocks were mainly driven by increased C input and N retention, resulting from enhanced plant productivity, which surpassed enhanced C loss from decomposition. Importantly, higher soil C and N stocks were associated with enhanced soil N mineralization rates, which can explain the strengthening of the positive diversity–productivity relationship observed in the last years of the experiment. Synthesis. We demonstrated that also in the absence of legumes, plant species richness promotes soil carbon (C) and nitrogen (N) stocks via increased plant productivity. In turn, enhanced soil C and N stocks showed a positive feedback to plant productivity via enhanced N mineralization, which could further accelerate soil C and N storage in the long term.

There is a fundamental trade-off between leaf traits associated with either resource acquisition or resource conservation. This gradient of trait variation, called the economics spectrum, also applies to fine roots, but whether it is consistent for coarse roots or at the plant community level remains untested. We measured a set of morphological and chemical root traits at a community level (functional parameters; FP) in 20 plant communities located along land-use intensity gradients and across three climatic zones (tropical, mediterranean and montane). We hypothesized (i) the existence of a root economics spectrum in plant communities consistent within root types (fine, < 2 mm; coarse, 2–5 mm), (ii) that variations in root FP occur with soil depths (top 20 cm of soil and 100–150 cm deep) and (iii) along land-use gradients. Root FP covaried, in line with the resource acquisition–conservation trade-off, from communities with root FP associated with resource acquisition (e.g. high specific root length, SRL; thin diameters and low root dry matter contents, RDMC) to root FP associated with resource conservation (e.g. low SRL, thick diameters and high RDMC). This pattern was consistent for both fine and coarse roots indicating a strong consistency of a trade-off between resource acquisition and conservation for plant roots. Roots had different suites of traits at different depths, suggesting a disparity in root function and exploitation capacities. Shallow, fine roots were thinner, richer in nitrogen and with lower lignin concentrations associated with greater exploitation capacities compared to deep, fine roots. Shallow, coarse roots were richer in nitrogen, carbon and soluble concentrations than deep, coarse roots. Fine root parameters of highly disturbed, herbaceous-dominated plant communities in poorer soils were associated with foraging strategies, that is greater SRL and lower RDMC and lignin concentration than those from less disturbed communities. Coarse roots, however, were less sensitive to the land-use gradient. Synthesis. This study demonstrates the existence of a general trade-off in root construction at a community level, which operates within all root types, suggesting that all plant tissues are controlled by the trade-off between resource acquisition and conservation.

Arbuscular mycorrhizal fungi (AMF) mediate plant interspecific competition and community structure. However, the magnitude and direction of AMF effects and underlying mechanisms are not clear. Here, we synthesized the results of 304 studies to evaluate how AMF affect plant competition and community structure and which abiotic and biotic conditions in experimental design modify these AMF effects. The magnitude and direction of AMF effects on plant competitive ability (in terms of competitive response) differed markedly among plant functional groups. When AMF inoculum was added, competitive ability was strongly enhanced in N‐fixing forbs and was significantly suppressed in C₃ grasses, whereas no effect was observed in C₄ grasses, non‐N‐fixing forbs and woody species. Furthermore, AMF inoculation increased competitive ability of perennial species when their competitors were annual species. AMF inoculation differentially influenced separate aspects of plant community structure and species composition. AMF inoculation significantly increased plant diversity but had no effects on plant productivity. Response of dominant plant species to AMF inoculation was the determining factor in explaining variations in how and to what degree plant diversity was influenced by AMF inoculation. When dominant species derived strong benefits from AMF, their dominance level was increased by AMF inoculation, which consequently decreased plant diversity. We did not find stronger AMF effects on plant diversity and productivity when greater numbers of AMF species were used in the inoculation. Synthesis. Despite large variations in AMF effects among studies, a unifying mechanism was observed that the mycorrhizal responsiveness (differences in plant growth between AMF and non‐AMF colonization treatments) of target and neighbouring plant species can determine AMF effects on the competitive outcome among plant species, which in turn influenced plant species diversity and community composition. Given that plant traits, soil nutrient conditions and probably mycorrhizal fungal traits are all factors determining the degree of mycorrhizal response of plant species, future studies should explicitly consider each of these factors in experimental design to better understand AMF effects on plant coexistence, plant community dynamics and ecosystem processes.