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.     

Changes in plant species richness induce functional shifts in soil nematode communities in experimental grassland

Background

Changes in plant diversity may induce distinct changes in soil food web structure and accompanying soil feedbacks to plants. However, knowledge of the long-term consequences of plant community simplification for soil animal food webs and functioning is scarce. Nematodes, the most abundant and diverse soil Metazoa, represent the complexity of soil food webs as they comprise all major trophic groups and allow calculation of a number of functional indices.

Methodology/Principal Findings

We studied the functional composition of nematode communities three and five years after establishment of a grassland plant diversity experiment (Jena Experiment). In response to plant community simplification common nematode species disappeared and pronounced functional shifts in community structure occurred. The relevance of the fungal energy channel was higher in spring 2007 than in autumn 2005, particularly in species-rich plant assemblages. This resulted in a significant positive relationship between plant species richness and the ratio of fungal-to-bacterial feeders. Moreover, the density of predators increased significantly with plant diversity after five years, pointing to increased soil food web complexity in species-rich plant assemblages. Remarkably, in complex plant communities the nematode community shifted in favour of microbivores and predators, thereby reducing the relative abundance of plant feeders after five years.

Conclusions/Significance

The results suggest that species-poor plant assemblages may suffer from nematode communities detrimental to plants, whereas species-rich plant assemblages support a higher proportion of microbivorous nematodes stimulating nutrient cycling and hence plant performance; i.e. effects of nematodes on plants may switch from negative to positive. Overall, food web complexity is likely to decrease in response to plant community simplification and results of this study suggest that this results mainly from the loss of common species which likely alter plant – nematode interactions.     

Permanent seed banks in chalk grassland under various management regimes: their role in the restoration of species-rich plant communities

A survey was carried out on four stands of a chalk grassland (open, intensively grazed, tall and scrub) in order to monitor the permanent soil seed bank under various management regimes responsible for the decline of the ecological interest of this ecosystem. There was no real similarity between the seed banks and the associated vegetation, except in the intensively grazed stand. When vegetation changes, seed banks appear to be quite stable and are mostly dominated by three species (Centaurium erythraea, Origanum vulgare, Hypericum perforatum). The seed bank seems to be quite useless in order to restore a species-rich community after both grazing intensification or abandonment. Because recolonization processes are the critical factor, the management of species-rich grassland has to be designed with a view to preserving a given plant community as well as to improving dispersion processes from a source to target areas.     

Diversity reduces invasibility in experimental plant communities: the role of plant species

Several studies have presented experimental evidence that diversity reduces invasibility in grassland communities. The interpretation of these results has been disputed recently and it was proposed that sampling effects were responsible for the observed decrease of invasibility with diversity. The experiments performed to date were not designed to adequately separate sampling from diversity effects. Using the establishment of native plant species in experimental plant communities as a model of invasibility, we show that the number of invaders decreased with increasing diversity. When the presence of particular species is included, their effects are dominant. Centaurea jacea showed a strong effect at low diversity, whereas Leucanthemum vulgare showed a very strong negative impact at each diversity level. The negative effect of the latter might be related to root‐feeding nematodes that showed far higher abundance in plots with Leucanthemum. However, diversity remained a significant factor in determining the number of invading species and the numbers of an abundant invader.     

How plant diversity and legumes affect nitrogen dynamics in experimental grassland communities

Positive relationships between species richness and ecosystem processes such as productivity or nitrogen cycling can be the result of a number of mechanisms. We examined how species richness, biomass, and legume presence, diversity, and abundance explained nitrogen dynamics in experimental grassland plots in northern Sweden. Nitrogen concentrations and '¹⁵N values were measured in plants grown in 28 mixtures (58 plots) including 1, 2, 4, 8 or 12 local grassland species over four years. Values for '¹⁵N declined over time for all three functional groups (grasses, legumes, and non-leguminous forbs), suggesting greater reliance on N fixed by legumes over time by all species. Above ground percent nitrogen (%N) also declined over time but root %N and total N did not. Path analysis of above ground data suggested that two main factors affected %N and the size of the N pool. First, higher plant diversity (species richness) increased total N through increased biomass in the plot. Although in the first two years of the experiment this was the result of a greater probability of inclusion of at least one legume, in the last two years diversity had a significant effect on biomass beyond this effect. Second, percent legumes planted in the plots had a strong effect on above ground %N and '¹⁵N, but a much smaller effect on above ground biomass. In contrast, greater plant diversity affected N in roots both by increasing biomass and by decreasing %N (after controlling for effects mediated by root biomass and legume biomass). Increased legume biomass resulted in higher %N and lower '¹⁵N for both non-legume forbs and grasses in the first year, but only for grasses in the third year. We conclude that a sampling effect (greater probability of including a legume) contributed towards greater biomass and total N in high-diversity communities early on in the experiment, but that over time this effect weakened and other positive effects of diversity became more important.     

Effects of experimental rainfall manipulations on Chihuahuan Desert grassland and shrubland plant communities

Aridland ecosystems are predicted to be responsive to both increases and decreases in precipitation. In addition, chronic droughts may contribute to encroachment of native C₃ shrubs into C₄-dominated grasslands. We conducted a long-term rainfall manipulation experiment in native grassland, shrubland and the grass–shrub ecotone in the northern Chihuahuan Desert, USA. We evaluated the effects of 5 years of experimental drought and 4 years of water addition on plant community structure and dynamics. We assessed the effects of altered rainfall regimes on the abundance of dominant species as well as on species richness and subdominant grasses, forbs and shrubs. Nonmetric multidimensional scaling and MANOVA were used to quantify changes in species composition in response to chronic addition or reduction of rainfall. We found that drought consistently and strongly decreased cover of Bouteloua eriopoda, the dominant C₄ grass in this system, whereas water addition slightly increased cover, with little variation between years. In contrast, neither chronic drought nor increased rainfall had consistent effects on the cover of Larrea tridentata, the dominant C₃ shrub. Species richness declined in shrub-dominated vegetation in response to drought whereas richness increased or was unaffected by water addition or drought in mixed- and grass-dominated vegetation. Cover of subdominant shrubs, grasses and forbs changed significantly over time, primarily in response to interannual rainfall variability more so than to our experimental rainfall treatments. Nevertheless, drought and water addition shifted the species composition of plant communities in all three vegetation types. Overall, we found that B. eriopoda responded strongly to drought and less so to irrigation, whereas L. tridentata showed limited response to either treatment. The strong decline in grass cover and the resistance of shrub cover to rainfall reduction suggest that chronic drought may be a key factor promoting shrub dominance during encroachment into desert grassland.     

A comparative study of seed number, seed size, seedling size and recruitment in grassland plants

In this study we analyse relationships between seed number, seed size, seedling size and recruitment success in grassland plants. The often hypothesised trade-off between seed size and seed number was supported by a cross-species analysis and by an analysis of 35 phylogenetically independent contrasts, derived from a data-set of 72 species. Apart from among-species relatedness, we also controlled for possible confounding effect of plant size that may influence both seed size and seed number. A sowing experiment with 50 species was performed in the field. The seeds were sown in a grassland and subjected to two treatments, disturbance and undisturbed sward. Evidence for seed-limited recruitment was obtained for 45 of the species. Disturbance had a significant, or nearly significant, positive effect on recruitment for 16 of the 45 species. The relative recruitment in undisturbed sward increased with increased seed size, and both recruitment success and seedling size were positively related to seed size. We suggest that a trade-off between competitive ability and number of recruitment opportunities follows from the trade-off between seed size and seed number, through a causal chain from seed size via seedling size to recruitment success. The relationships between seed size, seed number and recruitment may be an important underlying mechanism for abundance and dynamics of plant species in grassland vegetation. This is an example of a direct link between evolutionary life-history theory, and theory of plant community structure.     

The role of geologic grazing refuges in structuring Mediterranean grassland plant communities

We tested the hypothesis that small rock-enclosed geologic refuges have an important role in maintaining grazing-sensitive species in grassland with a long history of intense grazing. The study was carried out in Mediterranean grassland in a basalt landscape in northern Israel. Community composition was compared in 63 sites between samples of 1 m² quadrats in two microhabitats: (1) rock-enclosed, presumed “refuges” and (2) rock-adjacent, “near refuges” but just outside them, accessible to cattle grazing. Median refuge area was 2.75 m², median rock height and diameter around refuges were 1.10 m and 1.50 m. Median height of residual dry herbage was 1.41 m in refuges, compared to 0.38 m outside, indicating the difference in grazing intensity. Species richness at three scales (quadrat, site, all sites) was significantly greater in the near-refuge than in the refuge habitat. In the latter, many annuals were excluded by dominance of tall perennials. Twelve species (of 103) had significantly higher cover in refuges, including tall perennial grasses, tall annuals, climbers, and a shrub. A total of 53 species with a strong significant negative response to refuges were mostly small and medium height annuals. The intermediate group of 38 species with weak or non-significant responses to refuges included, among others, dominant tall grasses that were abundant both in refuges and just outside them. The latter, as well as most refuge-positive species had shown a positive response to protection in exclosures. The results support the hypothesis that small rock-enclosed habitats—more so than artificial exclosures—are effective grazing refuges for rare, grazing-susceptible species. The contribution of refuges to species richness at the landscape scale is much greater than their proportion of the area. Dispersion from refuges maintains small populations of rare species near refuges and can initiate expansion into the landscape when grazing pressure is lowered.     

Differential responses of soil bacteria,fungi, archaea and protists to plant species richness and plant functional group identity

Plants are known to influence belowground microbial community structure along their roots, but the impacts of plant species richness and plant functional group (FG) identity on microbial communities in the bulk soil are still not well understood. Here, we used 454‐pyrosequencing to analyse the soil microbial community composition in a long‐term biodiversity experiment at Jena, Germany. We examined responses of bacteria, fungi, archaea, and protists to plant species richness (communities varying from 1 to 60 sown species) and plant FG identity (grasses, legumes, small herbs, tall herbs) in bulk soil. We hypothesized that plant species richness and FG identity would alter microbial community composition and have a positive impact on microbial species richness. Plant species richness had a marginal positive effect on the richness of fungi, but we observed no such effect on bacteria, archaea and protists. Plant species richness also did not have a large impact on microbial community composition. Rather, abiotic soil properties partially explained the community composition of bacteria, fungi, arbuscular mycorrhizal fungi (AMF), archaea and protists. Plant FG richness did not impact microbial community composition; however, plant FG identity was more effective. Bacterial richness was highest in legume plots and lowest in small herb plots, and AMF and archaeal community composition in legume plant communities was distinct from that in communities composed of other plant FGs. We conclude that soil microbial community composition in bulk soil is influenced more by changes in plant FG composition and abiotic soil properties, than by changes in plant species richness per se.     

Impact of plant functional group, plant species, and sampling time on the composition of nirK-type denitrifier communities in soil

We studied the influence of eight nonleguminous grassland plant species belonging to two functional groups (grasses and forbs) on the composition of soil denitrifier communities in experimental microcosms over two consecutive years. Denitrifier community composition was analyzed by terminal restriction fragment length polymorphism (T-RFLP) of PCR-amplified nirK gene fragments coding for the copper-containing nitrite reductase. The impact of experimental factors (plant functional group, plant species, sampling time, and interactions between them) on the structure of soil denitrifier communities (i.e., T-RFLP patterns) was analyzed by canonical correspondence analysis. While the functional group of a plant did not affect nirK-type denitrifier communities, plant species identity did influence their composition. This effect changed with sampling time, indicating community changes due to seasonal conditions and a development of the plants in the microcosms. Differences in total soil nitrogen and carbon, soil pH, and root biomass were observed at the end of the experiment. However, statistical analysis revealed that the plants affected the nirK-type denitrifier community composition directly, e.g., through root exudates. Assignment of abundant T-RFs to cloned nirK sequences from the soil and subsequent phylogenetic analysis indicated a dominance of yet-unknown nirK genotypes and of genes related to nirK from denitrifiers of the order Rhizobiales. In conclusion, individual species of nonleguminous plants directly influenced the composition of denitrifier communities in soil, but environmental conditions had additional significant effects.     

The relationship between seed size and abundance in plant communities: model predictions and observed patterns

Recent studies have suggested that seed size and plant abundance in communities are associated. However, inconsistent patterns have emerged from these studies, with varying mechanisms proposed to explain emergent relationships. We employ a theoretical framework, based on key theory lineages of vegetation dynamics and species coexistence, to examine relationships between species abundance and seed size. From these theory lineages, we identified four models and their predictions: the Seed size/number trade‐off model (SSNTM), the Succession model (SM), the Spatial competition model (SCM), and the Lottery model (LM). We then explored empirical evidence from ten diverse plant communities for seed size and abundance patterns, and related these patterns to model predictions. The SSNTM predicts a negative correlation between seed size and abundance. The SM predicts either a negative, positive or no correlation dependent on time since disturbance, while the SCM and LM make no predictions for a relationship between seed size and abundance. We found no evidence for consistent relationships between seed size and abundance across the ten communities. There were no consistent differences in seed size and abundance relationships between communities dominated by annuals compared to perennials. In three of the ten communities a significant positive seed size and abundance correlation emerged, which falsified the SSNTM as an important determinant of abundance structure in these communities. For sites in coastal woodland, the relationships between seed size and abundance were consistent with the predictions of the SM (although generally not significant), with fire being the disturbance. We suggest that the significant positive seed size and abundance correlations found may be driven by the association between large seeds and large growth forms, as large growth forms tend to be dominant. It seems likely that patterns of seed size and abundance in communities are determined by a complex interaction between environmental factors and correlations of plant attributes that determine a species’ strategy.     

Shifts in seed size across experimental nitrogen enrichment and plant density gradients

We tested the hypotheses that increased soil resource availability selects for larger seeded plants by: (1) increasing community seed density and seedling competition, (2) increasing light competition, which favours larger seeded species if their additional seed resources are allocated to shoots. We also tested the hypothesis that plants respond to increased nutrient availability by producing larger seeds. In a controlled environment experiment, we grew monocultures of five species of co-occurring annual plant species at two N addition rates and measured seed size and the number of seeds produced. In a second experiment, we sowed seed of the five species together, manipulated nitrogen (N) addition rate and community seed density and measured species performance. In the first experiment, elevated N addition increased seed size, but only in larger seeded species. In the second experiment, high N addition increased community seed production. However, contrary to our hypothesis small seeded species were selected under high seed density, possibly because they germinated and grew more rapidly thus enabling them to pre-empt limiting resources. Larger seeded species were favoured by high N addition, as hypothesised. However, our data suggest that this was due to their seed reserves boosting survival below the denser canopies of fertile conditions rather than via competitive advantages. Our results point to a largely overlooked role of recruitment in determining community response to fertilisation. Although our results may not generalise to all plant communities they suggest that seed traits play a role in community response to both the direct effect of N addition and the stimulation of seed density caused by increased productivity. These results are also consistent with the view that the advantage of large seed size is not a competitive one, but greater survival in the face of environmental hazards.     

Regional adaptation improves the performance of grassland plant communities

Many plant species are adapted locally or regionally. Whether such individual species performance translates into effects at community and ecosystem levels has rarely been tested. Such tests are crucial, however, to predict ecosystem consequences of sowing seed mixtures for grassland restoration or hay production. We compared the performance of replicated sown plant communities of regional origin with the performance of four foreign communities consisting of the same grassland species but originating from distances up to 890 km from our experimental site. The regional communities performed better than foreign communities in plant cover and diversity but not in aboveground biomass production. Additionally, in communities based on regional seeds fewer unsown species occurred and less bare ground was left open for erosion. Variation in community performance among source regions was related to climatic differences rather than to geographic distance to source regions. Individual species performance only partly explained community patterns, highlighting the importance of community level experiments. Our results suggest that the use of regional seeds represents an important approach to improve sown managed grasslands.     

The role of seed traits in determining the phylogenetic structure of temperate plant communities

Background and Aims

Phylogenetic clustering of species within plant communities can be expected to result from environmental filtering acting on an evolutionary-conserved plant trait. One such a candidate trait is the embryo to seed-size ratio (E:S). A high E:S may allow faster germination immediately after imbibition, and is therefore assumed to be advantageous in dry habitats. In this study the hypothesis was tested that habitat filtering driven by soil moisture conditions and acting on seed germination and seedling establishment is an important ecological mechanism in structuring temperate plant communities.

Methods

Vegetation samplings were performed in three habitats located within 200 km of each other in western Europe: Ellenberg indicator values showed that the habitats selected differed substantially in terms of soil moisture and light availability. E.S ratio and seed mass data for all genera were obtained from literature. Data were analysed using recently developed phylogenetic methods.

Key Results

Genera with a similar E:S tend to co-occur, as low and high E:S genera dominate in moist and dry habitats, respectively. A phylogenetically clustered pattern of community structure was evident, and dispersion of E:S was positively related to phylogenetic dispersion.

Conclusions

The phenotypically and phylogenetically clustered pattern indicates that E:S-mediated habitat filtering is an important assembly process structuring the plant community of the temperate climate habitats studied.