Effects of plant species richness on stand structure and productivity

Aims Aboveground biomass production commonly increases with species richness in plant biodiversity experiments. Little is known about the direct mechanisms that cause this result. We tested if by occupying different heights and depths above and below ground, and by optimizing the vertical distribution of leaf nitrogen, species in mixtures can contribute to increased resource uptake and, thus, increased productivity of the community in comparison with monocultures.Methods We grew 24 grassland plant species, grouped into four nonoverlapping species pools, in monoculture and 3- and 6-species mixture in spatially heterogeneous and uniform soil nutrient conditions. Layered harvests of above- and belowground biomass, as well as leaf nitrogen and light measurements, were taken to assess vertical canopy and root space structure.Important findings The distribution of leaf mass was shifted toward greater heights and light absorption was correspondingly enhanced in mixtures. However, only some mixtures had leaf nitrogen concentration profiles predicted to optimize whole-community carbon gain, whereas in other mixtures species seemed to behave more 'selfish'. Nevertheless, even in these communities, biomass production increased with species richness. The distribution of root biomass below ground did not change from monocultures to three- and six-species mixtures and there was also no indication that mixtures were better than monocultures at extracting heterogeneously as compared to homogeneously distributed soil resources. We conclude that positive biodiversity effect on aboveground biomass production cannot easily be explained by a single or few common mechanisms of differential space use. Rather, it seems that mechanisms vary with the particular set of species combined in a community.     

Light and nitrogen competition limit Lolium perenne in experimental grasslands of increasing plant diversity

Positive species richness effects on aboveground community productivity in experimental grasslands have been reported to correlate with variable responses of individual species. So far, it is largely unknown whether more complete use of resources at the community level correlates with resource limitation of particular species and may explain their decreasing performance with increasing plant diversity. Using the subordinate grass species Lolium perenne L. as a model, we monitored populations in 82 experimental grasslands of different plant diversity (Jena Experiment) from year 2 to 6 after establishment, and measured ecophysiological leaf traits related to light and nutrient acquisition and use. Population and plant individual sizes of L. perenne decreased with increasing species richness. A decrease in transmitted light with increasing species richness and legume proportion correlated with increasing specific leaf area (SLA). Despite this morphological adaptation to lower light availability, decreasing foliar δ(13) C signatures with increasing species richness and low variation in leaf gas exchange and chlorophyll concentrations suggested a low capacity of L. perenne for adjustment to canopy shade. Leaf nitrogen concentrations and foliar δ(15) N signatures indicated a better N supply in communities with legumes and a shift in the uptake of different N forms with increasing species richness. Leaf blade nitrate and carbohydrate concentrations as indicators of plants nutritional status supported that light limitation with increasing species richness and legume proportions, combined with a N limitation in communities with increasing proportions of non-legumes, correlated with the decreasing performance of L. perenne in communities of increasing plant diversity.     

Nutrient concentrations and fibre contents of plant community biomass reflect species richness patterns along a broad range of land-use intensities among agricultural grasslands

Understanding changes in biodiversity in agricultural landscapes in relation to land-use type and intensity is a major issue in current ecological research. In this context nutrient enrichment has been identified as a key mechanism inducing species loss in Central European grassland ecosystems. At the same time, insights into the linkage between agricultural land use and plant nutrient status are largely missing. So far, studies on the relationship between chemical composition of plant community biomass and biodiversity have mainly been restricted to wetlands and all these studies neglected the effects of land use. Therefore, we analyzed aboveground biomass of 145 grassland plots covering a gradient of land-use intensities in three regions across Germany. In particular, we explored relationships between vascular plant species richness and nutrient concentrations as well as fibre contents (neutral and acid detergent fibre and lignin) in the aboveground community biomass.We found the concentrations of several nutrients in the biomass to be closely linked to plant species richness and land use. Whereas phosphorus concentrations increased with land-use intensity and decreased with plant species richness, nitrogen and potassium concentrations showed less clear patterns. Fibre fractions were negatively related to nutrient concentrations in biomass, but hardly to land-use measures and species richness. Only high lignin contents were positively associated with species richness of grasslands. The N:P ratio was strongly positively related to species richness and even more so to the number of endangered plant species, indicating a higher persistence of endangered species under P (co-)limited conditions. Therefore, we stress the importance of low P supply for species-rich grasslands and suggest the N:P ratio in community biomass to be a useful proxy of the conservation value of agriculturally used grasslands.     

典型草原围封后羊草地上功能性状对枯落物累积的响应

植物功能性状反映植物适应环境变化过程中在不同器官形态与功能间的资源权衡与分配策略。典型草原围封后枯落物累积导致群落光照、热量和水分的重新分配并改变微环境特征。在此过程中植物地上功能性状将通过怎样的变化来适应新的环境,目前尚不清楚。2015—2017年每年8月对内蒙古地区3种典型草原共有物种羊草（Leymus chinensis）的植株、叶片和茎干功能性状进行了测量与分析。结果表明：枯落物累积显著增加了羊草的植株高度、单株重量、茎叶比和总叶面积;枯落物累积显著增加了羊草的叶片长度、叶片重量、单叶面积、节间长度和茎干重量,这些性状属于敏感性状;枯落物累积对羊草的叶片数量和节间数量无显著影响,相对而言,这些性状属于惰性性状;羊草的单株重量与植株高度、叶片重量呈极显著的正相关关系（P<0.0001）;羊草的植株高度与节间数量、节间长度呈极显著的正相关关系（P<0.0001）。本研究结果从植物地上功能性状的角度阐明了典型草原植物对环境变化的适应方式,可为围封草原的合理管理提供基础数据与理论依据。     

Using root traits to understand temporal changes in biodiversity effects in grassland mixtures

Biodiversity–ecosystem functioning (BEF) studies typically show that species richness enhances community biomass, but the underlying mechanisms remain debated. Here, we combine metrics from BEF research that distinguish the contribution of dominant species (selection effects, SE) from those due to positive interactions such as resource partitioning (complementarity effects, CE) with a functional trait approach in an attempt to reveal the functional characteristics of species that drive community biomass in species mixtures. In a biodiversity experiment with 16 plant species in monocultures, 4‐species and 16‐species mixtures, we used aboveground biomass to determine the relative contributions of CE and SE to biomass production in mixtures in the second, dry year of the experiment. We also measured root traits (specific root length, root length density, root tissue density and the deep root fraction) of each species in monocultures and linked the calculated community weighted mean (CWM) trait values and trait diversity of mixtures to CE and SE. In the second year of the experiment, community biomass, CE and SE increased compared to the first year. The contribution of SE to this positive effect was greater than that of CE. The increased contribution of SE was associated with root traits: SE increased most in communities with high abundance of species with deep, thick and dense roots. In contrast, changes in CE were not related to trait diversity or CWM trait values. Together, these results suggest that increased positive effects of species richness on community biomass in a dry year were mainly driven by increased dominance of deep‐rooting species, supporting the insurance hypothesis of biodiversity. Positive CE indicates that other positive interactions did occur, but we could not find evidence that belowground resource partitioning or facilitation via root trait diversity was important for community productivity in our biodiversity experiment.     

Increasing native, but not exotic, biodiversity increases aboveground productivity in ungrazed and intensely grazed grasslands

Species-rich native grasslands are frequently converted to species-poor exotic grasslands or pastures; however, the consequences of these changes for ecosystem functioning remain unclear. Cattle grazing (ungrazed or intensely grazed once), plant species origin (native or exotic), and species richness (4-species mixture or monoculture) treatments were fully crossed and randomly assigned to plots of grassland plants. We tested whether (1) native and exotic plots exhibited different responses to grazing for six ecosystem functions (i.e., aboveground productivity, light interception, fine root biomass, tracer nitrogen uptake, biomass consumption, and aboveground biomass recovery), and (2) biodiversity-ecosystem functioning relationships depended on grazing or species origin. We found that native and exotic species exhibited different responses to grazing for three of the ecosystem functions we considered. Intense grazing decreased fine root biomass by 53% in exotic plots, but had no effect on fine root biomass in native plots. The proportion of standing biomass consumed by cattle was 16% less in exotic than in native grazed plots. Aboveground biomass recovery was 30% less in native than in exotic plots. Intense grazing decreased aboveground productivity by 25%, light interception by 14%, and tracer nitrogen uptake by 54%, and these effects were similar in native and exotic plots. Increasing species richness from one to four species increased aboveground productivity by 42%, and light interception by 44%, in both ungrazed and intensely grazed native plots. In contrast, increasing species richness did not influence biomass production or resource uptake in ungrazed or intensely grazed exotic plots. These results suggest that converting native grasslands to exotic grasslands or pastures changes ecosystem structure and processes, and the relationship between biodiversity and ecosystem functioning.     

Density may alter diversity–productivity relationships in experimental plant communities

Contemporary biodiversity experiments, in which plant species richness is manipulated and aboveground productivity of the system measured, generally demonstrate that lowering plant species richness reduces productivity. However, we propose that community density may in part compensate for this reduction of productivity at low diversity. We conducted a factorial experiment in which plant functional group richness was held constant at three, while plant species richness increased from three to six to 12 species and community density from 440 to 1050 to 2525 seedlings m⁻². Response variables included density, evenness and above- and belowground biomass at harvest. The density gradient converged slightly during the course of the experiment due to about 10% mortality at the highest sowing density. Evenness measured in terms of aboveground biomass at harvest significantly declined with density, but the effect was weak. Overall, aboveground, belowground and total biomass increased significantly with species richness and community density. However, a significant interaction between species richness and community density occurred for both total and aboveground biomass, indicating that the diversity–productivity relationship was flatter at higher than at lower density. Thus, high species richness enabled low-density communities to reach productivity levels otherwise seen only at high density. The relative contributions of the three functional groups C₃, C₄ and nitrogen-fixers to aboveground biomass were less influenced by community density at high than at low species richness. We interpret the interaction effects between community density and species richness on community biomass by expanding findings about constant yield and size variation from monocultures to plant mixtures.     

Short‐term harvesting of biomass from conservation grasslands maintains plant diversity

High yields are a priority in managing biomass for renewable energy, but the environmental impacts of various feedstocks and production systems should be equally considered. Mixed‐species, perennial grasslands enrolled in conservation programs are being considered as a source of biomass for renewable energy. Conservation grasslands are crucial in sustaining native biodiversity throughout the US Upper Midwest, and the effects of biomass harvest on biodiversity are largely unknown. We measured the effect of late‐season biomass harvest on plant community composition in conservation grasslands in three regions of Minnesota, USA from 2009 to 2012. Temporal trends in plant species composition within harvested grasslands were compared to unharvested grasslands using mixed effects models. A before‐after control‐impact approach using effect sizes was applied to focus on pre‐ and postharvest conditions. Production‐scale biomass harvest did not affect plant species richness, species or functional group diversity, nor change the relative abundance of the main plant functional groups. Differences in the relative abundances of plant functional groups were observed across locations; and at some locations, changed through time. The proportion of non‐native species remained constant, while the proportion of noxious weeds decreased through time in both harvested and unharvested grasslands at the central location. Ordination revealed patterns in species composition due to location, but not due to harvest treatment. Therefore, habitat and bioenergy characteristics related to grassland plant communities are not expected to change due to short‐term or intermittent late‐season biomass harvest.     

放牧对内蒙古锡林河流域草地群落植物茎叶生物量资源分配的影响

以内蒙古锡林河流域沿水分梯度分布的灰脉苔草(Carex appendiculata)、贝加尔针茅(Stipa baicalensis)、羊草(Leymus chinensis)、大针茅(Stipa grandis)、小叶锦鸡儿(Caragana microphylla)和冷蒿(Artemisia frigida) 6个草地群落为对象, 研究了围封禁牧与放牧样地中144个共有植物种的高度、丛幅面积、茎、叶和株(丛)生物量、茎叶比等性状。结果表明: 1)在个体水平上, 放牧样地中植物的生殖枝高度、营养枝高度、丛幅面积、单株(丛)生物量、茎、叶生物量和茎叶比均显著低于围封禁牧样地, 植物在放牧干扰下表现出明显的小型化现象; 2)在群落水平上, 放牧亦显著降低了群落总生物量和茎、叶生物量; 3)过度放牧显著改变了物种的资源分配策略, 使生物量向叶的分配比例增加, 向茎的分配比例减少。资源优先向同化器官分配可能是植物对长期放牧干扰的一种重要适应对策; 4)轻度放牧对物种的资源分配没有显著影响, 单株(丛)生物量和群落茎、叶及总生物量均表现出增加趋势, 这与过度放牧的影响正好相反。过度放牧引起的植物个体小型化改变了生态系统中物种的资源分配策略, 进而对生态系统功能产生重要的影响。     

Short-term invasibility patterns in burnt and unburnt experimental Mediterranean grassland communities of varying diversities

This paper reports the findings of a short-term natural invasibility field study in constructed Mediterranean herbaceous communities of varying diversities, under a fire treatment. Three components of invasibility, i.e. species richness, density and biomass of invaders, have been monitored in burnt and unburnt experimental plots with resident diversity ranging from monocultures to 18-species mixtures. In general, species richness, density and biomass of invaders decreased significantly with the increase of resident species richness. Furthermore, the density and biomass of invading species were significantly influenced by the species composition of resident communities. Although aboveground biomass, leaf area index, canopy height and percent bare ground of the resident communities explained a significant part of the variation in the success of invading species, these covariates did not fully explain the effects of resident species richness. Fire mainly influenced invasibility via soil nutrient levels. The effect of fire on observed invasibility patterns seems to be less important than the effects of resident species richness. Our results demonstrate the importance of species richness and composition in controlling the initial stages of plant invasions in Mediterranean grasslands but that there was a lack of interaction with the effects of fire disturbance.     

Grassland resistance and resilience after drought depends on management intensity and species richness

The degree to which biodiversity may promote the stability of grasslands in the light of climatic variability, such as prolonged summer drought, has attracted considerable interest. Studies so far yielded inconsistent results and in addition, the effect of different grassland management practices on their response to drought remains an open question. We experimentally combined the manipulation of prolonged summer drought (sheltered vs. unsheltered sites), plant species loss (6 levels of 60 down to 1 species) and management intensity (4 levels varying in mowing frequency and amount of fertilizer application). Stability was measured as resistance and resilience of aboveground biomass production in grasslands against decreased summer precipitation, where resistance is the difference between drought treatments directly after drought induction and resilience is the difference between drought treatments in spring of the following year. We hypothesized that (i) management intensification amplifies biomass decrease under drought, (ii) resistance decreases with increasing species richness and with management intensification and (iii) resilience increases with increasing species richness and with management intensification.We found that resistance and resilience of grasslands to summer drought are highly dependent on management intensity and partly on species richness. Frequent mowing reduced the resistance of grasslands against drought and increasing species richness decreased resistance in one of our two study years. Resilience was positively related to species richness only under the highest management treatment. We conclude that low mowing frequency is more important for high resistance against drought than species richness. Nevertheless, species richness increased aboveground productivity in all management treatments both under drought and ambient conditions and should therefore be maintained under future climates.     

Linking species performance to community structure as affected by UV-B radiation: an attenuation experiment

Aims UV-B radiation is known to affect plant physiology and growth rate in ways that can influence community species composition and structure. Nevertheless, comparatively little is known about how UV-B radiation induced changes in the performance of individual species cascades to affect overall community properties. Because foliage leaves are primarily responsible for photosynthesis and carbon gain and are the major organ that senses and responds to UV-B radiation, we hypothesized that, under reduced UV-B radiation, species with larger leaf areas per plant would manifest higher growth rates and hence tend to improve their community status compared to species with smaller leaf areas per plant in herbaceous plant communities.Methods We tested this hypothesis by examining plant traits (leaf area per plant and plant height), plant growth rate (aboveground biomass per plant and plant biomass per area) and community status (species within-community relative biomass) for 19 common species in a two-year field experiment in an alpine meadow on Tibetan Plateau.Important findings Aboveground biomass per plant, as well as per area, progressively increased in a 39% reduced (relative to ambient) UV-B treatment during the experimental period. At the second year, 11 out of 19 species significantly or marginally significantly increased their plant height, leaf area per plant and aboveground biomass per plant. No species was negatively affected by reducing UV-B. As hypothesized, the increase in aboveground biomass per plant increased with increasing leaf area per plant, as indicated by cross-species regression analysis. Moreover, the change in species within-community status increased with increasing leaf area per plant. Our study demonstrates that UV-B radiation has differential effects on plant growth rate across species and hence significantly affects species composition and plant community structure. We suggest that UV-B radiation is an ecological factor structuring plant communities particularly in alpine and polar areas.     

Effects of trait plasticity on aboveground biomass production depend on species identity in experimental grasslands

Experimental studies with grassland species found a positive relationship between species richness and community biomass production, however the response of individual species was highly variable. The mechanisms behind these patterns are poorly understood. Here we studied aboveground biomass production and plasticity of growth characteristics of four legumes with similar morphology (Lotus corniculatus, Medicago lupulina, Onobrychis viciifolia, Trifolium hybridum) in experimental grasslands varying in species richness (1, 2, 4, 8, 16 and 60) and composition. We identified O. viciifolia and T. hybridum as species that reached higher biomass production in mixtures than expected from monoculture yields, while L. corniculatus and M. lupulina mostly had lower yields than expected. Variation of morphological traits across the species-richness gradient was lowest in the highly competitive O. viciifolia, but increased in the smaller species. The tall-growing O. viciifolia achieved higher biomass production by both, a higher number of plant individuals and an increase in mean mass per individual. Mean shoot number per individual remained constant, but individuals produced heavier shoots. The medium-sized T. hybridum also increased the number of plant individuals, but mean mass per individual did not respond to community species richness. The average mass per shoot was increased in mixtures, but the species developed less shoots per individual. Shoot length and stem weight ratio of T. hybridum increased with community species richness. Morphological changes in the less successful L. corniculatus and M. lupulina with a smaller growth stature were similarly directed as those of T. hybridum. The observed morphological changes are known as typical shade-avoidance mechanisms in dense vegetation. Our study shows that stress responses to changes in resource availability may be a mechanism to enforce higher aboveground biomass production of individual species in mixtures, but it depends on species identity whether trait plasticity is large enough to exceed stress-induced growth limitations.     

Moderate grazing promotes the root biomass in Kobresia meadow on the northern Qinghai–Tibet Plateau

Grazing is an important modulator of both plant productivity and biodiversity in grassland community, yet how to determine a suitable grazing intensity in alpine grassland is still controversy. Here, we explore the effects of different grazing intensities on plant biomass and species composition, both at community level and functional group level, and examines the productivity–species richness relationship under four grazing patterns: no grazing (CK), light grazing (LG), moderate grazing, (MG) and heavy grazing (HG), attempt to determine a suitable grazing intensity in alpine grassland. The results were as follows. The total aboveground biomass (AGB) reduced with increasing grazing intensity, and the response of plant functional groups was different. AGB of both sedges and legumes increased from MG to HG, while the AGB of forbs reduced sharply and the grass AGB remained steady. There was a significant positive relationship between productivity and species richness both at community level and functional group level. In contrast, the belowground biomass (BGB) showed a unimodal relationship from CK to HG, peaking in MG (8,297.72 ± 621.29 g/m²). Interestingly, the grassland community tends to allocate more root biomass to the upper soil layer under increasing grazing intensities. Our results suggesting that moderate levels of disturbance may be the optimal grassland management strategy for alpine meadow in terms of root production.     

Biodiversity influences ecosystem functioning in aquatic angiosperm communities

Knowledge of the connection between aquatic plant diversity and ecosystem processes is still limited. To examine how plant species diversity affects primary productivity, plant nutrient use, functional diversity of secondary producers and population/community stability, we manipulated submerged angiosperm species diversity in a field experiment lasting 15 weeks. Plant richness increased the shoot density for three of four species. Polyculture biomass production was enhanced by increasing richness, with positive complementarity and selection effects causing positive biodiversity effects. Species richness enhanced the community stability for biomass production and shoot density. Sediment ammonium availability decreased with plant diversity, suggesting improved nutrient usage with increasing plant richness. Interestingly, positive multitrophic effects of plant species richness on structural and functional diversity of macrobenthic secondary producers were recorded. The results suggest that mixed seagrass meadows play an important role for ecosystem functioning and thus contribute to the provision of goods and services in coastal areas.     

Community assembly and biomass production in regularly and never weeded experimental grasslands

We studied the natural colonisation of new species in experimental grasslands varying in plant species richness (from 1 to 60) and plant functional group richness (from 1 to 4) in either regularly or never weeded subplots during the first 3 years after establishment. Sown species established successfully, with no differences in species richness or their relative abundances between the regularly and never weeded subplots during the study period. Aboveground biomass of sown species increased with increasing sown species richness in both treatments. While a positive relationship between sown species richness and total aboveground biomass (including colonising species) existed in the 2nd year after sowing in the regularly and never weeded subplots, this positive relationship decayed in the 3rd year in the never weeded subplots because of a higher biomass of colonising species in species-poor mixtures. Total aboveground biomass varied independently of total species richness 3 years after sowing in both treatments. Jaccard similarity of coloniser species composition between regularly and never weeded subplots decreased from the 2nd to the 3rd year, indicating a divergence in coloniser species composition. Coloniser immigration and turnover rates were higher in regularly weeded subplots, confirming that weeding counteracts species saturation and increases the chance that new colonisers would establish. Although our study shows that low diversity plant communities are unstable and converge to higher levels of biodiversity, the effects of initially sown species on community composition persisted 3 years after sowing even when allowing for succession, suggesting that colonising species mainly filled empty niche space.     

Plant community diversity and composition affect individual plant performance

Effects of plant community diversity on ecosystem processes have recently received major attention. In contrast, effects of species richness and functional richness on individual plant performance, and their magnitude relative to effects of community composition, have been largely neglected. Therefore, we examined height, aboveground biomass, and inflorescence production of individual plants of all species present in 82 large plots of the Jena Experiment, a large grassland biodiversity experiment in Germany. These plots differed in species richness (1–60), functional richness (1–4), and community composition. On average, in more species-rich communities, plant individuals grew taller, but weighed less, were less likely to flower, and had fewer inflorescences. In plots containing legumes, non-legumes were higher and weighed more than in plots without legumes. In plots containing grasses, non-grasses were less likely to flower than in plots without grasses. This indicates that legumes positively and grasses negatively affected the performance of other species. Species richness and functional richness effects differed systematically between functional groups. The magnitude of the increase in plant height with increasing species richness was greatest in grasses and was progressively smaller in legumes, small herbs, and tall herbs. Individual aboveground biomass responses to increasing species richness also differed among functional groups and were positive for legumes, less pronouncedly positive for grasses, negative for small herbs, and more pronouncedly negative for tall herbs. Moreover, these effects of species richness differed strongly between species within these functional groups. We conclude that individual plant performance largely depends on the diversity of the surrounding community, and that the direction and magnitude of the effects of species richness and functional richness differs largely between species. Our study suggests that diversity of the surrounding community needs to be taken into account when interpreting drivers of the performance of individual plants.     

Species richness promotes ecosystem carbon storage: evidence from biodiversity-ecosystem functioning experiments

Plant diversity has a strong impact on a plethora of ecosystem functions and services, especially ecosystem carbon (C) storage. However, the potential context-dependency of biodiversity effects across ecosystem types, environmental conditions and carbon pools remains largely unknown. In this study, we performed a meta-analysis by collecting data from 95 biodiversity-ecosystem functioning (BEF) studies across 60 sites to explore the effects of plant diversity on different C pools, including aboveground and belowground plant biomass, soil microbial biomass C and soil C content across different ecosystem types. The results showed that ecosystem C storage was significantly enhanced by plant diversity, with stronger effects on aboveground biomass than on soil C content. Moreover, the response magnitudes of ecosystem C storage increased with the level of species richness and experimental duration across all ecosystems. The effects of plant diversity were more pronounced in grasslands than in forests. Furthermore, the effects of plant diversity on belowground plant biomass increased with aridity index in grasslands and forests, suggesting that climate change might modulate biodiversity effects, which are stronger under wetter conditions but weaker under more arid conditions. Taken together, these results provide novel insights into the important role of plant diversity in ecosystem C storage across critical C pools, ecosystem types and environmental contexts.     

How do leaf trait values change spatially and temporally with light availability in a grassland diversity experiment?

Complementarity in light use might increase light exploitation and could be an important mechanism explaining the coexistence of multiple species in plant communities of increasing diversity. We measured vertical light profiles and leaf traits related to light acquisition and light use in 40 mixtures of varying species richness (SR, 2, 4, 8 and 16) and functional group richness (FR, 1‐4) in a large grassland biodiversity experiment at five different times during the growing season. Light attenuation within the canopy differed significantly among mixtures of varying SR at peak biomass, with 40% in 2‐species mixtures and up to 80% in 16‐species mixtures. In contrast, increasing SR did not affect light attenuation at the beginning of the growing season or during regrowth after mowing, when large fractions of incoming radiation reached the ground level. These patterns suggested the presence of highly variable light niches over space and time. Trait expression differed among functional groups (except specific leaf area (SLA)) and varied within the growing season. However, we found no direct effect of increased SR or FR on the expression of leaf traits, except for positive species richness‐effects on SLA at peak biomass time. SLA and stomatal conductance increased and leaf dry matter content decreased at lower light at leaf height, while leaf greenness was independent of relative light availability. Dissimilarity of leaf traits (except SLA) at the community level increased with increasing SR. Thus, our results suggest that after accounting for light availability, which was driven by SR and time of year, variations in leaf trait expression within the grassland canopies did not depend on SR, but rather on functional group identity and time of year. Consequently, increased complementarity in light use at higher plant diversity is due to presence of more species with different leaf trait expression and trait variation in response to the actual light environment.     

Time course of plant diversity effects on Centaurea jacea establishment and the role of competition and herbivory

Aims Invasion resistance in experimental plant communities is known to increase with increasing diversity and further to depend on the presence of particular functional groups. To test whether these effects also hold true for the invader establishment phase beyond the seedling stage, we studied survival and performance of Centaurea jacea L. (brown knapweed) planted into experimental grassland communities of varying plant biodiversity over three consecutive years. Moreover, we analysed the role of insect herbivory and biomass of the recipient community for mediating diversity effects.Methods In 2005, seedlings of Centaurea were transplanted into experimental grassland communities (the Jena Experiment) covering a species richness (1–60) and functional group richness (1–4) gradient. Half of these transplants and the community surrounding them in each plot were sprayed with insecticide while the other half served as control. In 2006 and 2007 (during the second and third year after transplantation), we recorded survival, growth-related (e.g. transplant biomass, height) and reproduction-related traits (e.g. number of flower heads). Annual data on community aboveground biomass served as covariate to investigate mediating effects of aboveground competition with the recipient community.Important findings Species richness was the most important factor responsible for Centaurea limitation. Higher levels of diversity decreased survival and all performance traits in both years. These diversity effects were partly driven by community biomass, but not fully explained by that covariate, suggesting the importance also of further processes. The influence of functional group richness was strong in the second year after transplantation and weaker in the third year. Among the particular functional groups, only the presence of legumes showed strong negative effects on Centaurea survival and weak negative effects on growth and reproduction, the latter two being mediated by biomass. Insect herbivore reduction considerably benefited Centaurea in sprayed monocultures, where it grew significantly larger than in all other diversity levels and than in the control subplots. We conclude that effects of plant community properties on invading individuals change in the course of establishment, that plant species richness effects are also important during later stages of establishment, and that biomass (especially at high diversity) and herbivory (especially at low diversity) of the recipient community are important in mediating community effects on invaders.