Overyielding in grassland communities: testing the sampling effect hypothesis with replicated biodiversity experiments

We derive and test some assumptions and predictions of the Sampling Effect Hypothesis (SEH) by examining the relationship between the traits of species in monoculture and their relative abundance in mixture, and by comparing polyculture performance with single-species plots. Although we found a positive relationship between production in monoculture and dominance in mixtures as predicted by the SEH, the relationship had low explanatory power. Counter to predictions, the species with the highest monoculture biomass were not able to strongly dominate all mixtures; instead the dominance of these species decreased with increasing species richness. On average, polycultures did not achieve greater biomass than (transgressively overyield) the species in each mixture, or at each site, that was most productive in monoculture. However, mixture yields did transgressively overyield both the monoculture biomass of the dominant species in the mixtures, and the weighted average of all monocultures (non-transgressive overyielding), both of which were positively related to increasing species richness. The varying responses of different overyielding tests resulted because resource partitioning and positive interactions were often counter-balanced by selection for species with lower biomass than the highest-yielding monocultures. Judging whether or not mixtures overyield therefore depends in part upon which species is the basis for comparison. We present a new general framework for overyielding analysis where every monoculture provides a potential comparison and from which the most relevant tests can be selected.     

Biodiversity effects and transgressive overyielding

Aims The potential for mixtures of plant species to produce more biomass than every one of their constituent species in monoculture is still controversially discussed in the literature. Here we tested how this so-called transgressive overyielding is affected by variation between and within species in monoculture yields in biodiversity experiments.Methods We use basic statistical principles to calculate expected maximum monoculture yield in a species pool used for a biodiversity experiment. Using a real example we show how between- and within-species variance components in monoculture yields can be obtained. Combining the two components we estimate the importance of sampling bias in transgressive overyielding analysis.Important findings The net biodiversity effect (difference between mixture and average monoculture yield) needed to achieve transgressive overyielding increases with the number of species in a mixture and with the variation between constituent species in monoculture yields. If there is no significant variation between species, transgressive overyielding should not be calculated using the best monoculture, because in this case the difference between this species and the other species could exclusively reflect a sampling bias. The sampling bias decreases with increasing variation between species. Tests for transgressive overyielding require replicated species' monocultures. However, it can be doubted whether such an emphasis on monocultures in biodiversity experiments is justified if an analysis of transgressive overyielding is not the major goal.     

Biodiversity maintenance mechanisms differ between native and novel exotic-dominated communities

In many systems, native communities are being replaced by novel exotic-dominated ones. We experimentally compared species diversity decline between nine-species grassland communities under field conditions to test whether diversity maintenance mechanisms differed between communities containing all exotic or all native species using a pool of 40 species. Aboveground biomass was greater in exotic than native plots, and this difference was larger in mixtures than in monocultures. Species diversity declined more in exotic than native communities and declines were explained by different mechanisms. In exotic communities, overyielding species had high biomass in monoculture and diversity declined linearly as this selection effect increased. In native communities, however, overyielding species had low biomass in monoculture and there was no relationship between the selection effect and diversity decline. This suggests that, for this system, yielding behaviour is fundamentally different between presumably co-evolved natives and coevolutionarily naive exotic species, and that native-exotic status is important to consider.     

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.     

Differences in light interception in grass monocultures predict short-term competitive outcomes under productive conditions

Due to its inherent asymmetry, competition for light is thought to cause loss of diversity from eutrophied systems. However, most of the work on this topic in grasslands has been phenomenological and has not measured light directly. We present the results of one of the few mechanistic experiments investigating the outcome of short-term competition using measurements of light interception from monocultures of five perennial grass species grown under fertilized and irrigated conditions. We found that the level of incident light intercepted by each species in monoculture, a direct measure of resource-reduction ability, was an excellent predictor of the relative competitive effect in pairwise mixtures. Competition for light was asymmetric in relation to differences in light intercepting ability. Our results are consistent with the idea that when light is a limiting resource, competition between species for this resource can be asymmetric, contributing to high dominance and low diversity.     

Early-season effects of supplemented solar UV-B radiation on seedling emergence, canopy structure, simulated stand photosynthesis and competition for light

Mixtures and monocultures of wheat (Triticum aestivum) and wild oat (Avena fatua), a common weedy competitor of wheat, were exposed to enhanced solar UV-B radiation simulating a 20% reduction in stratospheric ozone to assess the timing and seasonal development of the UV-B effects on light competition in these species. Results from two years of field study revealed that UV-B enhancement had no detectable effect on the magnitude or timing of seedling emergence in either species. End-of-season measurements showed significant UV-B inhibition of leaf insertion height in wild oat in mixture and monoculture in the second year (irrigated year) but not in the first year (drought year). Leaf insertion height of wheat was not affected by UV-B in either year. The UV-B treatment had no detectable effect on monoculture or total (combined species) mixture LAI but did significantly increase (5–7%) the fractional contribution of wheat to the mixture LAI after four weeks of growth in both years. In addition, the UV-B treatment had subtle effects on LAI height profiles with early season mixtures showing significant reductions in wild oat LAI in lower canopy layers in both years while midseason Year 2 mixtures showed significant reductions in wild oat LAI in upper canopy layers. The changes in canopy structure were found to significantly increase (6–7%) the proportional simulated clear sky canopy photosynthesis and light interception of wheat in mixture. These findings, and others, indicate that the effects of UV-B enhancement on competition are realized very early in canopy development and provide additional support for the hypothesis that UV-B enhancement may shift the balance of competition between these species indirectly by altering competitive interactions for light.     

The role of plant species size in invasibility: a field experiment

Large plant species self-thin to disproportionately lower densities than smaller plant species, and therefore may leave more patches of unused space suitable for invasion. Using experimental monocultures of 11 old-field perennial plant species differing in maximum size, as well as mixtures composed of all monoculture species, we tested our primary hypothesis that monocultures of larger species will be more susceptible to natural invasion. After 3 years, monocultures of larger species were invaded by a significantly greater number of species, and more ramets, from the surrounding vegetation. Invading plant species were significantly smaller than the monoculture species being invaded, suggesting that smaller plant species may be better invaders. Thus, we quantified a trade-off between species size, which is frequently associated with increased competitive ability for light, and invasibility, suggesting one reason why large and small species coexist in virtually all plant communities. Although we expected that invasion would enhance biomass production by more fully capturing available resources, we found that the most highly invaded plots of each species produced significantly less biomass. This suggests that increased diversity resulting from invasion did not result in complementary resource use. Mixture plots containing all experimental species did not admit a significantly different number of invading ramets or species than most monocultures, indicating no obvious role for diversity in resistance to invasion, or complementary resource use. Our results suggest that relatively large species may be limited in their capacity to competitively exclude other, smaller species from communities because pure stands of the former are more susceptible to invasion by the latter.     

Overyielding and species diversity: what should we expect?

Recent empirical studies have found evidence of increased biomass production ('overyielding') in species mixtures relative to monoculture, but the interpretation of these results remains controversial, in part, because of the lack of a theoretical expectation. Here, we examined the expected frequency and stability of overyielding species mixtures using Lotka-Volterra models of species dynamics in two- and four-species systems in conjunction with community, population, and specific rate of biomass production (SRP) definitions of overyielding. Overyielding plant mixtures represented > 55% of potential species assemblages under community definitions and approximately 100% of species were either overyielding or underyielding under the population definition. Our species simulations approached their equilibria in 1-2 yr, supporting the relevancy of an equilibrial analysis. The range of parameter space that we explored produced realistic values of plot biomass, supporting their biological relevance. We show that overyielding is expected to be common under community definitions and population definitions. Overyielding, under community or population definitions, does not imply an actual increase in the specific rate of biomass production. In addition, assemblages of overyielding and underyielding species under all three definitions can be stable over time with underyielding species persisting in the presence of overyielding species.     

Exotic tree seedlings are much more competitive than natives but show underyielding when growing together

Aims Invasive species continue to be a worldwide threat to ecosystems mainly as a cause for biodiversity loss. Forest ecosystems, for example, are subject to a change in species composition due to the invasion of exotic species. Specifying the attributes that cause the strong competitiveness of several exotic species may improve the ability to understand and effectively manage plant invasions in the future. In this study the following hypotheses were tested: (1) biomass production of below- and aboveground plant components of the exotic tree species is higher than that of the natives, resulting in a higher competitiveness of the exotics; (2) the exclusion of root competition has a positive effect on the biomass production of the inferior native species; and (3) mixtures of native and exotic species yield a higher biomass production than the respective monocultures.Methods A pot experiment, containing about 2000 tree seedlings, was established. We investigated the biomass productivity and growth reactions of two native (Quercus robur L., Carpinus betulus L.) and two exotic tree species (Prunus serotina Ehrh., Robinia pseudoacacia L.) in different intra- and interspecific, competitive situations with and without the influence of root competition.Important findings The biomass production of both exotic species was significantly higher and led to a strong competitive advantage, resulting in a biomass decrease of the less competitive native species. The high belowground biomass of both exotic species had a negative effect on the biomass production. The competitive pressure of exotic tree seedlings on the native ones was largely driven by root competition. Furthermore, mixtures of native and exotic tree species had a higher productivity than their growth in monocultures would have predicted. Competition was lower for exotic species in mixtures with the less productive native species compared to the competition in monocultures or in mixture with the other highly productive exotic species. Accordingly, both highly competitive exotic species produced less biomass in mixture with each other compared to monocultures. Despite the significantly higher biomass of P. serotina in all mixtures and in monoculture, R. pseudoacacia seemed to be the dominating species. Due to its strong root competition, R. pseudoacacia significantly reduced the biomass production of P. serotina .     

Species richness, temporal variability and resistance of biomass production in a Mediterranean grassland

We studied the temporal variability and resistance to perturbation of the biomass production of grassland communities from an experimental diversity gradient (the Portuguese BIODEPTH project site). With increasing species richness relative temporal variability (CV) of plant populations increased but that of communities decreased, supporting the insurance hypothesis and related theory. Species‐rich communities were more productive than species‐poor communities in all three years although a natural climatic perturbation in the third year (frequent frost and low precipitation) caused an overall decrease in biomass production. Resistance to this perturbation was constant across the experimental species richness gradient in relative terms, supporting a similar response from the Swiss BIODEPTH experiment. The positive biomass response was generated by different combinations of the complementarity and selection effects in different years. Complementarity effects were positive across mixtures on average in all three years and positively related to diversity in one season. The complementarity effect declined following perturbation in line with total biomass but, counter to predictions, in relative terms overyielding was maintained in all years. Selection effects were positively related to diversity in one year and negative overall in the other two years. The response to perturbation varied among species and for the same species growing in monoculture and mixture, but following the frost communities were more strongly dominated by species with lower monoculture biomass and the selection effect was more negative. In total, our results support previous findings of a positive relationship between diversity and productivity and between diversity and the temporal stability of production, but of no effect of diversity on the resistance to perturbation. We demonstrate for the first time that the relative strength of overyielding remained constant during an exceptional natural environmental perturbation.     

Plant competition for light analyzed with a multispecies canopy model

Summary A multispecies canopy photosynthesis simulation model was used to examine the importance of canopy structure in influencing light interception and carbon gain in mixed and pure stands of wheat (Triticum aestivum L.) and wild oat (Avena fatua L.), a common weedy competitor of wheat. In the mixtures, the fraction of the simulated canopy photosynthesis contributed by wheat was found to decline during the growing season and this decline was closely related to reductions in the amount of leaf area in upper canopy layers. For both species in mixture and in monoculture, simulated photosynthesis was greatest in the middle or upper-middle canopy layers and sensitivity analyses revealed that canopy photosynthesis was most sensitive to changes in leaf area and leaf inclination in these layers. Changes in LAI and leaf inclination affected canopy carbon gain differently for mixtures and monocultures, but the responses were not the same for the two species. Results from simulations where the structural characteristics of the two species were substituted indicated that species differences in leaf inclination, sheath area and the fraction of leaf area alive were of minor consequence compared with the differences in total leaf area in influencing relative canopy carbon gain in mixtures. Competition for light in these species mixtures appears to be influenced most by differences in the positioning of leaf area in upper canopy layers which determines, to a great extent, the amount of light intercepted.     

Procedure for separating the selection effect from other effects in diversity–productivity relationship

In a greenhouse pot experiment we cultivated six meadow species in a replacement series design. The plants were grown at two sowing densities in monocultures and all possible species combinations. Our aim was to separate the selection effect from other diversity effects. This distinction is based on the notion that true overyielding is not a consequence of the selection effect. We suggest a hierarchical procedure, which is based on a repeated division of samples into the pots with the most productive species present and missing. Overyielding can be then demonstrated by a positive dependence of productivity on species richness in the subsets with the most productive species present. Although we found a strong dependence of biomass on species richness in the entire data set, the hierarchical method revealed no evidence of overyielding. Above-ground biomass in a monoculture was a good predictor of species success in a species mix.     

More plant biomass results in more offspring production in annuals,or does it?

Competitive ability in plants has been previously measured almost exclusively in terms of traits related to growth (biomass) or plant size. In this study, however, we used a multi‐species competition experiment with six annuals to measure relative competitive ability in terms of reproductive output, i.e. the number of offspring produced for the next generation. Under greenhouse conditions, plants of each species were started in pots from germinating seeds and were grown singly (free of competition) and at high density in both monocultures and in mixtures with all study species. Several traits traditionally regarded as determinants of competitive ability in plants were recorded for each species grown singly, including: seed mass, germination time, early growth rate and potential plant size (biomass and height). Under competition, several traits were recorded as indicators of relative performance in both monocultures and mixtures, including: biomass of survivors, total number of survivors, number of reproductive survivors, and reproductive output (total seed production) of the survivors. As expected, species that grew to a larger biomass in isolation had higher seed production in isolation. However, none of the traditional plant growth/size‐related traits, measured either in isolation or under competition, could predict between species variation in reproductive output under competition in either monocultures or mixtures. In mixtures, 97% of this variation in reproductive output could be explained by between‐species variation in the number of reproductive survivors. The results indicate that traits measured on plants grown singly may be poor predictors of reproductive output under competition, and that species’ rank order of competitive ability in terms of the biomass of survivors may bear no relationship to their rank order in terms of the number of offspring produced by these survivors. This has important implications for the interpretation of mechanisms of species coexistence and community assembly within vegetation.     

The effect of tree species diversity on fine-root production in a young temperate forest

The phenomenon of overyielding in species-diverse plant communities is mainly attributed to complementary resource use. Vertical niche differentiation belowground might be one potential mechanism for such complementarity. However, most studies that have analysed the diversity/productivity relationship and belowground niche differentiation have done so for fully occupied sites, not very young tree communities that are in the process of occupying belowground space. Here we used a 5–6 year old forest diversity experiment to analyse how fine-root (<2 mm) production in ingrowth cores (0–30 cm) was influenced by tree species identity, as well as the species diversity and richness of tree neighbourhoods. Fine-root production during the first growing season after the installation of ingrowth cores increased slightly with tree species diversity, and four-species combinations produced on average 94.8% more fine-root biomass than monocultures. During the second growing season, fine-root mortality increased with tree species diversity, indicating an increased fine-root turnover in species-rich communities. The initial overyielding was attributable to the response to mixing by the dominant species, Pseudotsuga menziesii and Picea abies, which produced more fine roots in mixtures than could be expected from monocultures. In species-rich neighbourhoods, P. abies allocated more fine roots to the upper soil layer (0–15 cm), whereas P. menziesii produced more fine roots in the deeper layer (15–30 cm) than in species-poor neighbourhoods. Our results indicate that, although there may be no lasting overyielding in the fine-root production of species-diverse tree communities, increasing species diversity can lead to substantial changes in the production, vertical distribution, and turnover of fine roots of individual species.     

Grassland plant species diversity decreases invasion by increasing resource use

Species richness of plant communities has been demonstrated to provide resistance to invasion by unsown species, though the relationship with resource availability varies between studies. The present work involved five grassland species grown in monocultures and in four-species mixtures sown in accordance with a simplex design. The species used represented different functional groups (i.e. grasses, legumes and non-N(2)-fixing species), each of which differed internally in terms of competitiveness. I hypothesized that sown diversity would negatively affect invader performance by decreasing the availability of light and soil nitrogen (N) for invading species, and that functional composition of the sown diversity would affect the functional composition of the invading flora. The experimental plots were harvested for two years, and were fertilized with 100 kg N ha(-1) each year. The number of unsown species (classified into four functional groups) invading each plot and their proportion of the biomass harvested were recorded. The penetration of incoming light through the canopy, the apparent N uptake by the sown species from the soil, and the mineral N content in the soil were measured. I found that diverse communities captured more resources both above- and belowground, and the number of invading species and their biomass production were smaller in mixed than in monoculture plots. However, the sampling effect of one grass was also strong. These results suggest that increased resource use in diverse communities can reduce invasion.     

Genotypic diversity effects on biomass production in native perennial bioenergy cropping systems

The perennial grass species that are being developed as biomass feedstock crops harbor extensive genotypic diversity, but the effects of this diversity on biomass production are not well understood. We investigated the effects of genotypic diversity in switchgrass (Panicum virgatum) and big bluestem (Andropogon gerardii) on perennial biomass cropping systems in two experiments conducted over 2008–2014 at a 5.4‐ha fertile field site in northeastern Illinois, USA. We varied levels of switchgrass and big bluestem genotypic diversity using various local and nonlocal cultivars – under low or high species diversity, with or without nitrogen inputs – and quantified establishment, biomass yield, and biomass composition. In one experiment (‘agronomic trial’), we compared three switchgrass cultivars in monoculture to a switchgrass cultivar mixture and three different species mixtures, with or without N fertilization. In another experiment (‘diversity gradient’), we varied diversity levels in switchgrass and big bluestem (1, 2, 4, or 6 cultivars per plot), with one or two species per plot. In both experiments, cultivar mixtures produced yields equivalent to or greater than the best cultivars. In the agronomic trial, the three switchgrass mixture showed the highest production overall, though not significantly different than best cultivar monoculture. In the diversity gradient, genotypic mixtures had one‐third higher biomass production than the average monoculture, and none of the monocultures were significantly higher yielding than the average mixture. Year‐to‐year variation in yields was lowest in the three‐cultivar switchgrass mixtures and Cave‐In‐Rock (the southern Illinois cultivar) and also reduced in the mixture of switchgrass and big bluestem relative to the species monocultures. The effects of genotypic diversity on biomass composition were modest relative to the differences among species and genotypes. Our findings suggest that local genotypes can be included in biomass cropping systems without compromising yields and that genotypic mixtures could help provide high, stable yields of high‐quality biomass feedstocks.     

Plant competition for light analyzed with a multispecies canopy model

Summary Competition for light among species in a mixed canopy can be assessed quantitatively by a simulation model which evaluates the importance of different morphological and photosynthetic characteristics of each species. A model was developed that simulates how the foliage of all species attenuate radiation in the canopy and how much radiation is received by foliage of each species. The model can account for different kinds of foliage (leaf blades, stems, etc.) for each species. The photosynthesis and transpiration for sunlit and shaded foliage of each species is also computed for different layers in the canopy. The model is an extension of previously described single-species canopy photosynthesis simulation models. Model predictions of the fraction of foliage sunlit and interception of light by sunlit and shaded foliage for monoculture and mixed canopies of wheat (Triticum aestivum) and wild oat (Avena fatua) in the field compared very well with measured values. The model was used to calculate light interception and canopy photosynthesis for both species of wheat/wild oat mixtures grown under normal solar and enhanced ultraviolet-B (290–320 nm) radiation (UV-B) in a glasshouse experiment with no root competition. In these experiments, measurements showed that the mixtures receiving enhanced UV-B radiation had a greater proportion of the total foliage area composed of wheat compared to mixtures in the control treatments. The difference in species foliage area and its position in the canopy resulted in a calculated increase in the portion of total canopy radiation interception and photosynthesis by wheat. This, in turn, is consistent with greater canopy biomass of wheat reported in canopies irradiated with supplemental UV-B.     

Interspecific interactions and biomass allocation among grassland plant species

Although a handful of studies have shown how interspecific interactions may influence plant shoot to root ratios, the issue of how these interactions influence biomass partitioning among coexisting plant species remains largely unexplored. In this study, we determined whether a given plant species could induce other plant species to allocate relative biomass to each of four zones (aboveground, and three soil depth layers) in a different manner to what they would otherwise, and whether this may influence the nature of competitive or facilitative interactions amongst coexisting plant species. We used a glasshouse study in which mixtures and monocultures of ten grassland plant species were grown in cylindrical pots to determine the effects of plant species mixtures versus monocultures on the production of shoots and of roots of other species for each of three soil depths. Across all experiments, stimulation of production in mixtures was far less common than suppression of production. Different plant species shifted their allocation to shoots or roots at different depths, suggesting that interspecific interactions can either: (1) increase the ratio of deep to shallow roots, perhaps because competition reduces root growth in the uppermost part of the soil profile; or (2) decrease this ratio by reducing plant vigour to such an extent that the plant cannot produce roots that can reach deep enough to exploit resources at lower depths. Further, these results suggest that there are instances in which competition may have the potential to enforce resource partitioning between coexisting plant species by inducing different species to root at different depths to each other.     

Cadmium-induced ammonium ion accumulation of rice seedlings at high temperature is mediated through abscisic acid

Significant increases in aboveground biomass production have been observed in mixed plantations of Eucalyptus globulus and Acacia mearnsii when compared to monocultures. However, this positive growth response may be enhanced or lost with changes in resource availability. Therefore this study examined the effect of the commonly limiting resources soil N, P and moisture on the growth of E. globulus and A. mearnsii mixtures in a pot trial. Pots containing two E. globulus plants, two A. mearnsii plants or one of each species were treated with high and low levels of N and P fertiliser. After 50 weeks, E. globulus plants grew more aboveground biomass in mixtures than monocultures. A. mearnsii were larger in mixtures only at low N, where both species were similar in size and the combined aboveground biomass of both species in mixture was greater than that of monocultures. At high N and both high and low levels of P fertiliser E. globulus appeared to dominate and suppress A. mearnsii. In these treatments, the faster growth of E. globulus in mixture did not compensate the reduced growth of A. mearnsii, so mixtures were less productive than (or not significantly different from) E. globulus monocultures. The greater competitiveness of E. globulus in these situations may have resulted from its higher N and P use efficiency and greater growth response to N and P fertilisers compared to A. mearnsii. This trial indicates that the complex interactions between species in mixtures, and thus the success of mixed plantations, can be strongly influenced by site factors such as the availability of N and P.     

Simple plant traits explain functional group diversity decline in novel grassland communities of Texas

Previous research has found that plant diversity declines more quickly in exotic than native grassland plots, which offers a model system for testing whether diversity decline is associated with specific plant traits. In a common garden experiment in the Southern Great Plains in central Texas, USA, we studied monocultures and 9-species mixtures of either all exotic or all native grassland species. A total of 36 native and exotic species were paired by phylogeny and functional group. We used community-level measures (relative abundance in mixture) and whole-plant (height, aboveground biomass, and light capture) and leaf-level traits (area, specific leaf area, and C:N ratio) to determine whether trait differences explained native-exotic differences in functional group diversity. Increases in species’ relative abundance in mixture were correlated with high biomass, height, and light capture in both native and exotic communities. However, increasing exotic species were all C₄ grasses, whereas, increasing native species included forb, C₃ grass and C₄ grass species. Exotic C₄ grasses had traits associated with relatively high resource capture: greater leaf area, specific leaf area, height, biomass, and light capture, but similar leaf C:N ratios compared to native C₄ grasses. Leaf C:N was consistently higher for native than exotic C₃ species, implying that resource use efficiency was greater in natives than exotics. Our results suggest that functional diversity will differ between grasslands restored to native assemblages and those dominated by novel collections of exotic species, and that simple plant traits can help to explain diversity decline.