Do species evenness and plant density influence the magnitude of selection and complementarity effects in annual plant species mixtures?

Abstract Plant species richness influences primary productivity via mechanisms that (1) favour species with particular traits (selection effect) and (2) promote niche differentiation between species (complementarity). Influences of species evenness, plant density and other properties of plant communities on productivity are poorly defined, but may depend on whether selection or complementarity prevails in species mixtures. We predicted that selection effects are insensitive to species evenness but increase with plant density, and that the converse is true for complementarity. To test predictions, we grew three species of annuals in monocultures and in three‐species mixtures in which evenness of established plants was varied at each of three plant densities in a cultivated field in Texas, USA. Above‐ground biomass was smaller in mixtures than expected from monocultures because of negative ‘complementarity’ and a negative selection effect. Neither selection nor complementarity varied with species evenness, but selection effects increased at the greatest plant density as predicted.     

Colonization sequence influences selection and complementarity effects on biomass production in experimental algal microcosms

Species extinction and immigration are both common in natural communities and the sequence with which species are lost from or added to communities may be crucial to community structure. We experimentally addressed this issue by growing six green algal species in monocultures and all possible two-species mixtures, with two colonization sequences for each mixture. Both convergence and divergence in community structure were observed. The compositions containing particularly productive species were more likely to converge, while those comprising of species with similar monoculture yields were more likely to diverge. The species mixtures with high-yielding initial and low-yielding invading species produced more biomass than monocultures, but mixtures with the opposite assembly order produced only the same level of biomass as monocultures did. To address the diversity–ecosystem functioning issue, we estimate complementarity effect by relative yield total (RYT) and selection effect by the correlation between species' monoculture yields and their relative yields in mixtures, respectively. We found overall negative complementarity and positive selection effect in mixtures with high-yielding species as initial colonizers, but positive complementarity and negative selection effect in mixtures with low-yielding initial species. Nonetheless, because we used only up to two species in each microcosm, our results are limited in addressing the relationship between species diversity and ecosystem functioning. Future research should study the effects of immigration history with many more species involved in community assembly.     

Effect of functional group richness and species richness in manipulated productivity–diversity studies: a glasshouse pot experiment

Species and functional group (grasses, legumes, creeping nonlegume forbs, rosette nonlegume forbs) richness of species assemblages composed of 16 species from four functional plant groups were manipulated to evaluate the productivity-diversity relationships in a greenhouse pot experiment. Pots were filled with sand, and supplied at two levels of nutrients. The plants were grown in monocultures, two, four, eight and 16 species mixtures. Individual two, four, and eight species mixtures differed in the richness of functional groups. Although the two characteristics of biodiversity, i.e. species and functional group richness, were necessarily correlated, it was shown that it is possible to separate their effect statistically, and also test for their common effect without pronounced loss of test power. There was a pronounced increase of average aboveground biomass and a mild increase in belowground biomass with biodiversity. The effect of functional group richness was more pronounced than the effect of the number of species. By using the method of Loreau and Hector (Nature 411 (2001) 72), selection and complementarity effects were statistically separated, and the overyielding index was calculated as a ratio of the productivity of a mixture to the productivity of its most productive component (to demonstrate transgressive overyielding). Positive values of complementarity and transgressive overyielding were both found, particularly in some rich communities and under high nutrient levels. Complementarity significantly increased only with functional group richness and mainly under high nutrients in the belowground biomass. Some species, when grown in monocultures, had decreased productivity under higher nutrients, and thus were more productive in mixtures than in monocultures. It seems that those species suffered from too high nutrient levels when grown in monocultures, but not in the presence of other species, which were able to use the nutrients in high concentrations and effectively decrease the nutrient levels. As a consequence, mixtures of high diversity were always more productive under high nutrients. The difference in species proportions between high and low nutrients, characterized by chord distance, increased with species richness. The relative change in productivity decreased with the number of functional groups. This suggests that species richness might lead to stabilization of aggregate characteristics (like total productivity) under changing environmental conditions by changing the proportions of individual species.     

Effects of plant diversity on soil carbon in diverse ecosystems: a global meta‐analysis

Soil organic carbon (SOC) is a valuable resource for mediating global climate change and securing food production. Despite an alarming rate of global plant diversity loss, uncertainties concerning the effects of plant diversity on SOC remain, because plant diversity not only stimulates litter inputs via increased productivity, thus enhancing SOC, but also stimulates microbial respiration, thus reducing SOC. By analysing 1001 paired observations of plant mixtures and corresponding monocultures from 121 publications, we show that both SOC content and stock are on average 5 and 8% higher in species mixtures than in monocultures. These positive mixture effects increase over time and are more pronounced in deeper soils. Microbial biomass carbon, an indicator of SOC release and formation, also increases, but the proportion of microbial biomass carbon in SOC is lower in mixtures. Moreover, these species‐mixture effects are consistent across forest, grassland, and cropland systems and are independent of background climates. Our results indicate that converting 50% of global forests from mixtures to monocultures would release an average of 2.70 Pg C from soil annually over a period of 20 years: about 30% of global annual fossil‐fuel emissions. Our study highlights the importance of plant diversity preservation for the maintenance of soil carbon sequestration in discussions of global climate change policy.     

Plant functional diversity mediates the effects of vegetation and soil properties on community-level plant nitrogen use in the restoration of semiarid sandy grassland

Species-rich plant communities use nitrogen (N) more efficiently in grassland ecosystems; however, the role of plant functional diversity in affecting community level plant N-use has received little attention. We examined plant N content, stock and N-use efficiency at community-level along a restoration gradient of sandy grassland (mobile dune, semi-fixed dune, fixed dune and grassland) in Horqin Sand Land, northern China. We used the functional trait-based approach to examine how plant functional diversity, reflected by the most abundant species’ traits (community-weighted mean, CWM) and the dispersion of functional trait values (FDis), affected N-use efficiency in sandy grassland restoration. We further used the structure equation model (SEM) to evaluate the direct or indirect effects of plant species richness, biomass, functional diversity and soil properties on community-level plant N-use efficiency. We found that plant biomass and its N stock increased following sandy grassland restoration, and there were lower plant N content and higher N-use efficiency in semi-fixed dune, fixed dune and grassland as compared with mobile dune. N-use efficiency was positively associated with plant species richness, biomass, CWM plant height, CWM leaf C:N, FDis and soil gradient, but SEM results showed that species richness, CWM leaf C:N, plant biomass and FDis controlled by soil properties were the main factors exerting direct effects. CWM plant height also had a positive effect on N-use efficiency through its indirect effect on plant biomass. Soil gradient increased N-use efficiency through an indirect effect on vegetation rather than a direct effect. Final SEM models based on different plant functional diversity explained over 74% of variances in N-use efficiency. Effects of plant functional diversity on N-use efficiency supported both the mass ratio hypothesis and the complementarity hypothesis. Our results clearly highlight the important role of plant functional diversity in mediating the effects of vegetation and soil properties on community level plant N-use in sandy grassland ecosystems.     

Species ecological strategy and soil phosphorus supply interactively affect plant biomass and phosphorus concentration

Excess soil phosphorus often constrains ecological restoration of degraded semi-natural grasslands in Western-Europe. Slow-growing species, often target of restoration (measures), are at a disadvantage because they are outcompeted by fast-growing species. Gaining insight into the responses of plant species and communities to soil phosphorus availability will help understanding restoration trajectories of grassland ecosystems. We set up two pot experiments using twenty grassland species with contrasting growth forms (i.e. grasses versus forbs) and nutrient use strategies (i.e. acquisitive versus conservative nutrient use). We quantified the nutrient use strategy of a species based on the stress-tolerance value of the CSR framework (StrateFy et al. 2017). We grew these species (1) as monocultures and (2) in mixtures along a soil phosphorus gradient and measured the aboveground biomass and plant phosphorus concentrations. Plant phosphorus concentration generally increased with soil phosphorus supply and biomass increased with soil phosphorus supply only in conservative communities. Forbs had higher plant phosphorus concentrations compared to grasses both in monocultures and mixtures. The species’ nutrient use strategy had contrasting effects on plant tissue phosphorus concentrations, depending on soil phosphorus supply (interaction effect) and vegetation biomass (dilution effect). Our findings contribute to the knowledge required for successful ecological restoration of species-rich grasslands. Our results suggest that under specific conditions (i.e. nitrogen limitation, no dispersal limitation, no light limitation), slow-growing species can survive and even thrive under excess soil phosphorus availability. In the field, competition by fast-growing species may be reduced by increased mowing or grazing management.     

Feedbacks of plant identity and diversity on the diversity and community composition of rhizosphere microbiomes from a long‐term biodiversity experiment

Soil microbes are known to be key drivers of several essential ecosystem processes such as nutrient cycling, plant productivity and the maintenance of plant species diversity. However, how plant species diversity and identity affect soil microbial diversity and community composition in the rhizosphere is largely unknown. We tested whether, over the course of 11 years, distinct soil bacterial communities developed under plant monocultures and mixtures, and if over this time frame plants with a monoculture or mixture history changed in the bacterial communities they associated with. For eight species, we grew offspring of plants that had been grown for 11 years in the same field monocultures or mixtures (plant history in monoculture vs. mixture) in pots inoculated with microbes extracted from the field monoculture and mixture soils attached to the roots of the host plants (soil legacy). After 5 months of growth in the glasshouse, we collected rhizosphere soil from each plant and used 16S rRNA gene sequencing to determine the community composition and diversity of the bacterial communities. Bacterial community structure in the plant rhizosphere was primarily determined by soil legacy and by plant species identity, but not by plant history. In seven of the eight plant species the number of individual operational taxonomic units with increased abundance was larger when inoculated with microbes from mixture soil. We conclude that plant species richness can affect below‐ground community composition and diversity, feeding back to the assemblage of rhizosphere bacterial communities in newly establishing plants via the legacy in soil.     

Belowground biodiversity effects of plant symbionts support aboveground productivity

Soil microbes play key roles in ecosystems, yet the impact of their diversity on plant communities is still poorly understood. Here we demonstrate that the diversity of belowground plant-associated soil fungi promotes plant productivity and plant coexistence. Using additive partitioning of biodiversity effects developed in plant biodiversity studies, we demonstrate that this positive relationship can be driven by complementarity effects among soil fungi in one soil type and by a selection effect resulting from the fungal species that stimulated plant productivity the most in another soil type. Selection and complementarity effects among fungal species contributed to improving plant productivity up to 82% and 85%, respectively, above the average of the respective fungal species monocultures depending on the soil in which they were grown. These results also indicate that belowground diversity may act as insurance for maintaining plant productivity under differing environmental conditions.     

Bacterial endophytes and root hairs

Aims

This study aimed to measure the effect of plant diversity on N uptake in grasslands and to assess the mechanisms contributing to diversity effects.

Methods

Annual N uptake into above- and belowground organs and soil nitrate pools were measured in the Jena experiment on a floodplain soil with mixtures of 2–16 species and 1–4 functional groups, and monocultures. In mixtures, the deviation of measured data from data expected from monoculture performance was calculated to assess the contribution of complementarity/facilitation and selection.

Results

N uptake varied from <1 to 45 g?N m^?2 yr^?1, and was higher in grasslands with than without legumes. On average, N uptake was higher in mixtures (21?±?1 g?N m^?2 yr^?1) than monocultures (13?±?1 g?N m^?2 yr^?1), and increased with species richness in mixtures. However, compared to N uptake expected from biomass proportions of species in mixtures, N uptake of mixtures was only slightly higher and a significant surplus N uptake was confined to mixtures containing legumes and non-legumes.

Conclusions

In our study, high N uptake of species rich mixtures was mainly due to dominance of productive species and facilitation by legumes whereas complementarity among non-legumes was of minor relevance.     

Ecological Assessment of Dune Restorations in the Great Lakes Region

Because of the economic and environmental importance of stabilizing fragile sand dune habitats, restoration of dunes has become a common practice. Restoration efforts in the Great Lakes and East Coast regions of North America often consist of planting monocultures of the dominant native grass species, Ammophila breviligulata. We evaluated 18 dune restoration projects in the Great Lakes region conducted over the past 25 years. We characterized attributes of diversity (plants and insects), vegetation structure (plant biomass and cover), and ecological processes (soil nutrients and mycorrhizal fungi abundance) in each restoration, and we compared these measures to geographically paired natural dune communities. Restoration sites were similar to reference sites in most measured variables. Differences between restorations and reference sites were mostly explained by differences in ages, with the younger sites supporting slightly lower plant diversity and mycorrhizal spore abundance than older sites. Plant community composition varied little between restored and reference sites, with only one native forb species, Artemisia campestris, occurring significantly more often in reference sites than restored sites. Although it remains unclear whether more diverse restoration plantings could accelerate convergence on the ecological conditions of reference dunes, in general, traditional restoration efforts involving monoculture plantings of A. breviligulata in Great Lakes sand dunes appear to achieve ecological conditions found in reference dunes.     

Species diversity and identity effects on the water consumption of tree sapling assemblages under ample and limited water supply

Studies examining the influence of biodiversity on ecosystem functioning have rarely considered water turnover, the quantitatively most important biogeochemical flux in ecosystems and a process with high sensitivity to climate warming. With a tree sapling experiment consisting of three diversity levels (1, 3, 5 species), 11 different species combinations and two soil moisture levels (moist and dry), we examined the influence of tree species diversity and species identity on stand transpiration (T) under ample and restricted water supply. We further asked whether growth in mixture leads to adaptive responses in the hydraulic system and water loss regulation in plants with heterospecific neighbors compared to plants in monoculture. In moist soil, T was on average ～11% higher in the mixtures than in the monocultures (significant net diversity effect), which can mostly be attributed to a selection effect. Overyielding in T was highest in mixtures when Tilia cordata and/or Fraxinus excelsior were present. Both species developed larger leaf areas (LA) and sapwood areas (SA) in monocultures than the other species and furthermore increased LA and SA from the monocultures to the mixtures. Thus, inherent species differences in LA and hydraulics, but also neighbor effects on these traits determined T to a large extend. In dry soil, the diversity effect on T was not larger but slightly smaller, which is not in agreement with other published studies. We conclude that differences between pure and mixed sapling assemblages in stand water consumption and drought response are mainly caused by species identity effects, while species diversity seems to be less influential.     

Native soil microbial amendments generate trade‐offs in plant productivity,diversity, and soil stability in coastal dune restorations

Given the important role that soil microbes play in structuring plant communities and mediating ecosystem functions, there is growing interest in harnessing microbial communities to restore degraded ecosystems. Dune restorations, in particular, may benefit from native soil amendments because microbial diversity and abundance are very low in unvegetated areas. In an outdoor mesocosm experiment simulating Texas Gulf Coast dune restorations, we tested how native soil microbial amendments and restored diversity of foundational grasses influenced three key restoration responses: plant performance, plant diversity (including the colonization of native forbs), and soil stability. We found that native microbial amendments increased plant diversity and have the potential to increase soil stability, but this came at the cost of decreased plant biomass. Our results suggest that soil enemies in the native microbial amendments increased plant diversity by decreasing the performance of the dominant grass species and that arbuscular mycorrhizal fungi in the native microbial amendments increased the density of fungal hyphae in the soil, which can increase soil stability. Depending on the goals of the restoration, native soil microbial amendments may be a simple and inexpensive method to provide restoration benefits.     

Plant diversity loss reduces soil respiration across terrestrial ecosystems

The rapid global biodiversity loss has led to the decline in ecosystem function. Despite the critical importance of soil respiration (Rs) in the global carbon and nutrient cycles, how plant diversity loss affects Rs remains uncertain. Here we present a meta‐analysis using 446 paired observations from 95 published studies to evaluate the effects of plant and litter mixtures on Rs and its components. We found that total Rs and heterotrophic respiration (Rh) were, on average, greater in plant mixtures than expected from those of monocultures. These mixture effects increased with increasing species richness (SR) in both plant and litter mixtures. While the positive effects of species mixtures remained similar over time for total Rs, they increased over time for Rh in plant mixtures but decreased in litter mixtures. Despite the wide range of variations in mean annual temperature, annual aridity index, and ecosystem types, the plant mixture effects on total Rs and Rh did not change geographically, except for a more pronounced increase of total Rs in species mixtures with reduced water availability. Our structural equation models suggested that the positive effects of SR and stand age on total and Rh were driven by increased plant inputs and soil microbial biomass. Our results suggest that plant diversity loss has ubiquitous negative impacts on Rs, one of the fundamental carbon‐cycle processes sustaining terrestrial element cycling and ecosystem function.     

Soil fertility increases with plant species diversity in a long-term biodiversity experiment

Most explanations for the positive effect of plant species diversity on productivity have focused on the efficiency of resource use, implicitly assuming that resource supply is constant. To test this assumption, we grew seedlings of Echinacea purpurea in soil collected beneath 10-year-old, experimental plant communities containing one, two, four, eight, or 16 native grassland species. The results of this greenhouse bioassay challenge the assumption of constant resource supply; we found that bioassay seedlings grown in soil collected from experimental communities containing 16 plant species produced 70% more biomass than seedlings grown in soil collected beneath monocultures. This increase was likely attributable to greater soil N availability, which had increased in higher diversity communities over the 10-year-duration of the experiment. In a distinction akin to the selection/complementarity partition commonly made in studies of diversity and productivity, we further determined whether the additive effects of functional groups or the interactive effects of functional groups explained the increase in fertility with diversity. The increase in bioassay seedling biomass with diversity was largely explained by a concomitant increase in N-fixer, C4 grass, forb, and C3 grass biomass with diversity, suggesting that the additive effects of these four functional groups at higher diversity contributed to enhance N availability and retention. Nevertheless, diversity still explained a significant amount of the residual variation in bioassay seedling biomass after functional group biomass was included in a multiple regression, suggesting that interactions also increased fertility in diverse communities. Our results suggest a mechanism, the fertility effect, by which increased plant species diversity may increase community productivity over time by increasing the supply of nutrients via both greater inputs and greater retention.     

Rapid biodiversity declines in both ungrazed and intensely grazed exotic grasslands

Exotic-dominated ecosystems with low diversity are becoming increasingly common. It remains unclear, though, whether differences between native and exotic species (driver model), or changes in disturbances or resources (passenger model), allow exotics to become competitive dominants. In our field experiment, plant species origin (native or exotic), cattle grazing (ungrazed or intensely grazed once), and species composition treatments were fully crossed and randomly assigned to four-species mixtures and monocultures of grassland plants. We found that biodiversity declined more rapidly in exotic than in native species mixtures, regardless of our grazing disturbance treatment. Early declines in species evenness (i.e., increases in dominance) led to subsequent declines in species richness (i.e., local extinctions) in exotic mixtures. Specifically, Simpson’s diversity was 29% lower after 1 year, and species richness was 15% lower after 3 years, in exotic than in native mixtures. These rapid biodiversity declines in exotic mixtures were partly explained by decreased complementarity (i.e., niche partitioning and facilitation), presumably because exotic species lack the coevolutionary history that can lead to complementarity and coexistence in native communities. Thus, our results suggest that exotic species can drive biodiversity declines in the presence or absence of a grazing disturbance, partly because exotic species interactions differ from native species interactions. This implies that restoring plant biodiversity in grasslands may require removal of exotic species, in addition to disturbance management.     

Plant species identity and diversity effects on different trophic levels of nematodes in the soil food web

Previous studies on biodiversity and soil food web composition have mentioned plant species identity, as well as plant species diversity as the main factors affecting the abundance and diversity of soil organisms. However, most studies have been carried out under limitations of time, space, or appropriate controls. In order to further examine the relation between plant species diversity and the soil food web, we conducted a three-year semi-field experiment in which eight plant species (4 forb and 4 grass species) were grown in monocultures and mixtures of two, four and eight plant species. In addition there were communities with 16 plant species. We analyzed the abundance and identity of the nematodes in soil and roots, including feeding groups from various trophic levels (primary and secondary consumers, carnivores, and omnivores) in the soil food web.
Plant species diversity and plant identity affected the diversity of nematodes. The effect of plant diversity was attributed to the complementarity in resource quality of the component plant species rather than to an increase in total resource quantity. The nematode diversity varied more between the different plant species than between different levels of plant species diversity, so that plant identity is more important than plant diversity. Nevertheless the nematode diversity in plant mixtures was higher than in any of the plant monocultures, due to the reduced dominance of the most abundant nematode taxa in the mixed plant communities. Plant species identity affected the abundances of the lower trophic consumer levels more than the higher trophic levels of nematodes. Plant species diversity and plant biomass did not affect nematode abundance. Our results, therefore, support the hypothesis that resource quality is more important than resource quantity for the diversity of soil food web components and that plant species identity is more important than plant diversity per se.     

Plant functional diversity enhances associations of soil fungal diversity with vegetation and soil in the restoration of semiarid sandy grassland

The trait‐based approach shows that plant functional diversity strongly affects ecosystem properties. However, few empirical studies show the relationship between soil fungal diversity and plant functional diversity in natural ecosystems. We investigated soil fungal diversity along a restoration gradient of sandy grassland (mobile dune, semifixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China, using the denaturing gradient gel electrophoresis of 18S rRNA and gene sequencing. We also examined associations of soil fungal diversity with plant functional diversity reflected by the dominant species' traits in community (community‐weighted mean, CWM) and the dispersion of functional trait values (FD_is). We further used the structure equation model (SEM) to evaluate how plant richness, biomass, functional diversity, and soil properties affect soil fungal diversity in sandy grassland restoration. Soil fungal richness in mobile dune and semifixed dune was markedly lower than those of fixed dune and grassland (P < 0.05). Soil fungal richness was positively associated with plant richness, biomass, CWM plant height, and soil gradient aggregated from the principal component analysis, but SEM results showed that plant richness and CWM plant height determined by soil properties were the main factors exerting direct effects. Soil gradient increased fungal richness through indirect effect on vegetation rather than direct effect. The negative indirect effect of FDis on soil fungal richness was through its effect on plant biomass. Our final SEM model based on plant functional diversity explained nearly 70% variances of soil fungal richness. Strong association of soil fungal richness with the dominant species in the community supported the mass ratio hypothesis. Our results clearly highlight the role of plant functional diversity in enhancing associations of soil fungal diversity with community structure and soil properties in sandy grassland ecosystems.     

Moderate plant–soil feedbacks have small effects on the biodiversity–productivity relationship: A field experiment

Plant–soil feedback (PSF) has gained attention as a mechanism promoting plant growth and coexistence. However, most PSF research has measured monoculture growth in greenhouse conditions. Translating PSFs into effects on plant growth in field communities remains an important frontier for PSF research. Using a 4‐year, factorial field experiment in Jena, Germany, we measured the growth of nine grassland species on soils conditioned by each of the target species (i.e., 72 PSFs). Plant community models were parameterized with or without these PSF effects, and model predictions were compared to plant biomass production in diversity–productivity experiments. Plants created soils that changed subsequent plant biomass by 40%. However, because they were both positive and negative, the average PSF effect was 14% less growth on “home” than on “away” soils. Nine‐species plant communities produced 29 to 37% more biomass for polycultures than for monocultures due primarily to selection effects. With or without PSF, plant community models predicted 28%–29% more biomass for polycultures than for monocultures, again due primarily to selection effects. Synthesis: Despite causing 40% changes in plant biomass, PSFs had little effect on model predictions of plant community biomass across a range of species richness. While somewhat surprising, a lack of a PSF effect was appropriate in this site because species richness effects in this study were caused by selection effects and not complementarity effects (PSFs are a complementarity mechanism). Our plant community models helped us describe several reasons that even large PSF may not affect plant productivity. Notably, we found that dominant species demonstrated small PSF, suggesting there may be selective pressure for plants to create neutral PSF. Broadly, testing PSFs in plant communities in field conditions provided a more realistic understanding of how PSFs affect plant growth in communities in the context of other species traits.     

Spatial aggregation facilitates coexistence and diversity of wild plant species in field margins

European agri-environment schemes encourage farmers to establish sown field margin strips to protect and enhance wild plant diversity. However, plant diversity in such wild plant sowings based on seed mixtures is often low due to the high competitiveness of few, common species. Here we analysed whether intraspecific aggregation could enhance the performance of less competitive species, and how plant performance is influenced by the number of species in a mixture. We focused on inter- and intraspecific competition between six agricultural wild plant species (Centaurea cyanus, Calendula arvensis, Melilotus officinalis, Poa annua, Bromus mollis, Medicago lupulina), and tested (i) two different seeding patterns (intraspecifically aggregated vs. randomly dispersed) and (ii) three different species mixtures (monocultures, three-species, and six-species mixtures). Plant performance was measured in terms of number of individuals, biomass per individual, and biomass per m². Intraspecific aggregation resulted in higher numbers of individuals of all species, while mixtures generated lower numbers of individuals. The performance of plant species differed depending on their position in the competitive hierarchy. Competitively weak species suffered much less from intraspecific than interspecific competition in terms of biomass, and the competitively weakest species became even excluded in the most species rich and randomly dispersed sowings with high interspecific competition. In conclusion, the performance of wild plant species was influenced by both seeding pattern and number of species in a mixture. Intraspecific aggregation enabled the coexistence of competitively weak species by reducing interspecific competitive exclusion processes. Consequently, agri-environmental schemes designed to preserve and enhance biodiversity should consider small-scale processes influencing the distribution and abundance of plants, and develop new agricultural sowing technologies to cultivate competitively weak and endangered wild plant species.     

Functional identity is the main driver of diversity effects in young tree communities

Two main effects are proposed to explain biodiversity–ecosystem functioning relationships: niche complementarity and selection effects. Both can be functionally defined using the functional diversity (FD) and functional identity (FI) of the community respectively. Herein, we present results from the first tree diversity experiment that separated the effect of selection from that of complementarity by varying community composition in high‐density plots along a gradient of FD, independent of species richness and testing for the effects of FD and community weighted means of traits (a proxy for FI) on stem biomass increment (a proxy for productivity). After 4 years of growth, most mixtures did not differ in productivity from the averages of their respective monocultures, but some did overyield significantly. Those positive diversity effects resulted mostly from selection effects, primarily driven by fast‐growing deciduous species and associated traits. Net diversity effect did not increase with time over 4 years.