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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.

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.

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.

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.

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.

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.

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.

Long‐term pedogenesis leads to important changes in the availability of soil nutrients, especially nitrogen (N) and phosphorus (P). Changes in the availability of micronutrients can also occur, but are less well understood. We explored whether changes in leaf nutrient concentrations and resorption were consistent with a shift from N to P limitation of plant productivity with soil age along a > 2‐million‐year dune chronosequence in south‐western Australia. We also compared these traits among plants of contrasting nutrient‐acquisition strategies, focusing on N, P and micronutrients. The range in leaf [P] for individual species along the chronosequence was exceptionally large for both green (103–3000 μg P g⁻¹) and senesced (19–5600 μg P g⁻¹) leaves, almost equalling that found globally. From the youngest to the oldest soil, cover‐weighted mean leaf [P] declined from 1840 to 228 μg P g⁻¹, while P‐resorption efficiency increased from 0% to 79%. All species converged towards a highly conservative P‐use strategy on the oldest soils. Declines in cover‐weighted mean leaf [N] with soil age were less strong than for leaf [P], ranging from 13.4 mg N g⁻¹ on the youngest soil to 9.5 mg N g⁻¹ on the oldest soil. However, mean leaf N‐resorption efficiency was greatest (45%) on the youngest, N‐poor soils. Leaf N:P ratio increased from 8 on the youngest soil to 42 on the oldest soil. Leaf zinc (Zn) concentrations were low across all chronosequence stages, but mean Zn‐resorption efficiency was greatest (55–74%) on the youngest calcareous dunes, reflecting low Zn availability at high pH. N₂‐fixing species had high leaf [N] compared with other species. Non‐mycorrhizal species had very low leaf [P] and accumulated Mn across all soils. We surmise that this reflects Mn solubilization by organic acids released for P acquisition. Synthesis. Our results show community‐wide variation in leaf nutrient concentrations and resorption that is consistent with a shift from N to P limitation during long‐term ecosystem development. High Zn resorption on young calcareous dunes supports the possibility of micronutrient co‐limitation. High leaf [Mn] on older dunes suggests the importance of carboxylate release for P acquisition. Our results show a strong effect of soil nutrient availability on nutrient‐use efficiency and reveal considerable differences among plants of contrasting nutrient‐acquisition strategies.

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.

Fine roots and mycorrhiza often represent the largest input of carbon (C) into soils and are therefore of primary relevance to the soil C balance. Arbuscular mycorrhizal (AM) fungi have previously been found to increase litter decomposition which may lead to reduced soil C stocks, but these studies have focused on immediate decomposition of relatively high amounts of high‐quality litter and may therefore not hold in many ecological settings over longer terms. Here, we assessed the effect of mycorrhizal fungi on the fate of C and nitrogen (N) contained within a realistic amount of highly ¹³C‐/¹⁵N‐labelled root litter in soil. This litter was either added fresh or after a 3‐month incubation period under field conditions to a hyphal in‐growth core where mycorrhizal abundance was either reduced or not through rotation. After 3 months of incubation with a plant under glasshouse conditions, the effect of turning cores on residual ¹³C and ¹⁵N inside the cores was measured, as well as ¹³C incorporation in microbial signature fatty acids and ¹⁵N incorporation of plants. Turning of cores increased the abundance of fungal decomposers and ¹³C loss from cores, while ¹⁵N content of cores and plants was unaffected. Despite the difference in disturbance that turning the cores could have caused, the results suggest that mycorrhizal fungi and field incubation of litter acted to additively increase the proportion of ¹³C left in cores. Synthesis. Apart from stimulating litter decomposition as previously shown, mycorrhizas can also stabilize C during litter decomposition and this effect is persistent through time.