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Aims Silicon (Si) has been shown to beneficially affect plant performance under stressful environmental conditions, such as water or nutrient deficiency. Here we tested the effects of two important plant nutrients, nitrogen (N) and phosphorus (P), on Si content in different plant organs in the grass species Holcus lanatus . Methods We studied trait responses to N limitation, balanced nutrient availability and P limitation. Single plant individuals were grown in sand-filled pots in a greenhouse for 2 months. Nitrogen, phosphorus, carbon and silicon contents were determined in leaves, stems and roots, as were leaf and roots traits, biomass production and root enzyme activity. Results Si content was lowest under balanced nutrient supply in all plant organs. Under P limitation Si content was highest in leaves and stems, in roots it was highest under N limitation. Si:C ratios were lowest under balanced conditions, and highest under nutrient limitation. Root phosphatase activity was highest under P limitation and chlorophyll content was lowest under N limitation. Conclusions Our model species assimilated less ‘high cost C’ and took up more ‘low cost Si’ under nutrient limitation, especially under P deficiency. Si potentially plays an important role in different environments, such as nutrient or light limitation, which in turn may be related to different plant strategies, for example higher stem rigidity in high Si plants versus higher stem flexibility in low Si plants. More research is needed to further elucidate the role of silicon in different concepts of trait-environment relationships.

Plant performance is driven by nutrient availability. So far, studies on plant nutrient relationships mostly focused on nitrogen (N) and phosphorus (P), whereas potassium (K) received less attention. We evaluated whether plant responses to variation in nutrient availability are similar or different for the nutrients N, P and K and whether the same plant traits are decisive for the competitive ability of species along gradients of N, P and K availabilities. We studied plant trait responses of three temperate perennial grass species (Alopecurus pratensis, Agrostis capillaris and Anthoxanthum odoratum, each dominant under N limitation, P limitation and K limitation, respectively). Each gradient consisted of eight treatments (N1–N8, P1–P8, K1–K8), with N:P:K supply rates being 1:0.8:6 (N1), 128:0.8:6 (N8), 12:0.075:6 (P1), 12:9.6:6 (P8), 12:0.8:0.56 (K1) and 12:0.8:72 (K8) (total supply per plant individual in mg). Plant individuals were kept either in single‐species pots (intraspecific competition) or in mixed‐species pots (interspecific competition). After 2 months, plant biomass, leaf and root traits and root enzyme activity were measured. Most below‐ground traits (e.g. total root length, root diameter) responded most strongly to competition as predictor variable and only secondly to the type of nutrient or their supply rates (unlike RGRTotal and SLA). Leaf chlorophyll content and root enzyme activity responded most strongly to supply rate of nutrients and the interaction of nutrient type and supply rate, respectively. In line with our prediction, A. pratensis was the superior competitor under N limitation and A. odoratum was the superior competitor under K limitation (assessed by relative dominance [RD] and relative competitive strength [RCS]). Both species showed high investment in total root length. Opposite to our expectation, A. capillaris was a bad competitor under P limitation, with low root length but higher root phosphatase activity. Our results indicate that below‐ground plant traits are decisive in nutrient‐related competition between plant species. Further, competitive strength is determined by a high total root length per soil volume both under N and K limitation. Under P limitation, the model species A. capillaris showed a weak competitive performance either because it was lacking colonization with mycorrhiza and/or the high investment in phosphatase activity posed a trade‐off with higher N investment in phosphatase production. A free Plain Language Summary can be found within the Supporting Information of this article. A free Plain Language Summary can be found within the Supporting Information of this article.

It is known that the number of limiting nutrients may affect the species richness of plant communities, but it is unclear whether the type of nutrient limitation is also important. I place the results from a study in Patagonia (elsewhere in this issue) in the context of the number and types of nutrients that are limiting. I present four mechanisms through which N or P limitation may potentially influence species richness. These mechanisms are related to: (i) the number of forms in which P or N are present in soil and the plant traits needed to acquire them, (ii) the mechanisms and traits that control species competition and coexistence under N or P limitation, (iii) the regional species pools of plants capable of growing under N- and P-limited conditions, and (iv) the interaction between the type of nutrient limitation and community productivity. It appears likely that P limitation can favour a higher species richness than N limitation, in at least in a variety of low productive plant communities, but evidence to support this conclusion is so far lacking. The four mechanisms proposed here offer a framework for exploring whether the type of nutrient limitation per se, or an interaction with productivity, is a potential driver for variation in species diversity.

The dung microbiome is a complex system that is highly influenced by species and diet. This study characterized the dung bacterial and fungal communities of five herbivore species inhabiting the National Park Zuid‐Kennemerland, the Netherlands. The five selected herbivore species were rabbit (Oryctolagus cuniculus L.), cow (Bos taurus L.), horse (Equus ferus caballus L.), fallow deer (Dama dama L.), and European bison (Bison bonasus L.). We explored the effects of distinct digestive physiology (ruminants vs. non‐ruminants) and diverse dietary preferences on the microbial community composition of herbivore dung. Firmicutes and Bacteroidetes were dominant bacterial phyla in the dung of all five herbivore species, and Ascomycota was the predominant fungal phylum. Verrucomicrobiota and Mucoromycota were more present in horse dung and Proteobacteria were more abundant in rabbit dung than the three ruminant dung types. There were few significant differences in the microbial community structure among the three ruminant dung types. The alpha and beta diversity of dung microbial communities significantly differed between ruminants and non‐ruminants, especially in bacterial communities. Based on MetaCyc pathways, we found that the primary functions of bacteria in herbivore dung were focused on biosynthesis, various super pathways, and degradation, with a few differences between ruminant and non‐ruminant dung. FUNGuild analysis showed that horse dung had more saprotrophic fungi, while the fungi in fallow deer dung had more symbiotrophic properties, with the fungal functions of bison, cow, and rabbit dung somewhere in between. There was also a correlation between microbial community and nutrient composition of the substrate in herbivore dung. Understanding the dung microbial community composition of these herbivore species can enrich the database of mammalian gut microbiomes for studying the mechanisms of microbial community variation while preparing for exploring a new perspective to study the impact of herbivores on ecosystems through dung deposition.

1. Different grass species dominate grasslands fertilized with nitrogen (N) or phosphorus (P), possibly due to the impact of N : P stoichiometry on competitive interactions. How species compete for nutrients, and whether the mechanisms are similar for N and P, is still not fully understood. 2. We investigated whether the outcome of competition between Alopecurus pratensis and Agrostis capillaris depends on N : P stoichiometry, and on the ability to acquire supplied N or P. Monocultures and mixtures of the two species were grown in pots at nine combinations of N and P supply (N : P ratios 1∙7, 15 and 135, all at three supply levels). After 3 months, we determined plant biomass, morphological traits and nutrient concentrations of plant tissues. 3. N : P supply ratios had similar effects on the growth of the two species but contrasting effects on their competitive strength, as assessed by comparing growth in competition to growth in monoculture: Alopecurus was the stronger competitor under N limitation (N : P ratio 1∙7), whereas Agrostis was an equal or stronger competitor under P limitation (N : P ratio 135). This result resembled patterns of species distribution observed in the field. 4. The strong competitive response of Alopecurus at the low N : P supply ratio was associated with a high investment in root biomass and root length, and a high nitrogen productivity. The competitive response of Agrostis at the high N : P supply ratio was associated with low root mortality and high root phosphatase activity. 5. Our results obtained under N limitation support the pre-emption theory in which plants with the highest root length are able to acquire more N from the soil than their competitors, and therefore can suppress their growth. Under P limitation, however, plant investment in root length could not explain competitor suppression. Here, other factors important in competition for P, such as mycorrhizal hyphal length, root longevity, or exudation rates of P releasing compounds, merit investigation.

Worldwide, alien plant invasions have been intensively studied in the past decades, but mechanisms controlling the invasibility of native communities are not fully understood yet. The stochastic niche hypothesis predicts that species-rich plant communities are less prone to alien plant invasions than species-poor communities, which is supported by some but not all field studies, with some very species-rich communities such as the Brazilian Cerrado becoming heavily invaded. However, species-rich communities potentially contain a greater variety of facilitative interactions in resource exploitation than species-poor communities, from which invasive plants might benefit. This alternative hypothetical mechanism might explain why nutrient-poor, species-rich ecosystems are prone to invasion. Here we show that a high species richness both impedes and promotes invasive plants in the Brazilian Cerrado, using structural equation modelling and data from 38 field sites. We found support for the stochastic niche hypothesis through an observed direct negative influence of species richness on abundance of alien invasive species, but an indirect positive effect of species richness on invasive alien plants through soil phosphatase activity that enhances P availability was also found. These field observations were supported with results from a mesocosm experiment. Root phosphatase activity of plants increased with species richness in the mesocosms, which was associated with greater community P and N uptake. The most prominent alien grass species of the region, Melinis minutiflora , benefited most from the higher N and P availability in the species mixtures. Hence, this study provides a novel explanation of why species-richness may sometimes promote rather than impede invasion, and highlights the need to perform facilitation experiments in multi-species communities.

Global change drivers such as eutrophication and plant invasions will create novel environments for many plant species. Through adaptive trait plasticity plants may maintain their performance under these novel conditions and may outcompete those showing low‐adaptive trait plasticity. In a greenhouse study, we determined if plasticity in traits is adaptive or maladaptive in endangered, nonendangered, and invasive plant species in response to variation of nitrogen (N) and phosphorus (P) availability (N:P ratios 1.7, 15, and 135) and whether plastic trait responses are adaptive and/or costly for fitness (i.e., biomass). Species choice comprised 17 species from three functional groups (legumes, nonlegume forbs, and grasses), either classified as endangered, nonendangered, or invasive. After 2 months, plants were harvested and nine traits related to carbon assimilation and nutrient uptake were measured (leaf area, SLA, LDMC, SPAD, RMR, root length, SRL, root surface area, and PME activity). We found more traits responding plastically to variation in P than in N. Plasticity only created costs when P was varied. Plasticity in traits was mostly adaptively neutral toward fitness, with plasticity in three traits being similarly adaptive across all species groups: SPAD (as a measure of chlorophyll content, adaptive to N and P limitation), leaf area, and root surface area (adaptive to P limitation). We found little differences in trait plasticity between endangered, nonendangered, and invasive species. Synthesis. Along a gradient from N limitation, balanced N:P supply, and P limitation, we found that the type of fluctuating nutrient (i.e., if N or P is varied) is decisive for the adaptive value of a trait. Variation in P availability (from balanced supply to P limitation) created both a stronger reduction in fitness as well as created plasticity costs in more traits than variation in N availability (from balanced supply to N limitation). However, the patterns observed in our study may change if nutrient availability is altered, either by nutrient inputs or by a shift in nutrient availabilities, for example, by decreasing N input as foreseen by European Legislation, but without simultaneously decreasing P input. In the context of global change drivers, specifically eutrophication and exotic species invasion, we seek to answer if plasticity in plant traits is adaptive or maladaptive for fitness in endangered, non‐endangered and invasive congeneric plant species in response to variation of nitrogen (N) and phosphorus (P) availability. Our results indicate that although some plant traits responded plastically to nutrient availability, they did not contribute to plant fitness. We further show differences in trait plasticity and their effects on fitness between the type of nutrient that was varied (N and P), but they were similar between endangered, non‐endangered and invasive plant species.

Nutrient availability is important for plant community composition and diversity, but most studies focus on inorganic nutrients. Far less is known about the impact of nutrients in organic forms such as herbivore dung. Here we show that dung of 11 European herbivore species varies widely in nitrogen (N) and phosphorus (P) concentrations, as well as in C:N:P ratios. We demonstrate that variation in dung quality of five herbivore species influences the diversity and composition of a mesocosm plant community. The impact of dung quality was at least as strong as, or stronger than, the effect of manipulating the quantity of dung by a factor six. Our study supports the hypothesis that both nutrient quantity and nutrient imbalances are important controlling factors for plant species diversity, and stresses the important role of herbivores on plant communities, not only via selective foraging, but also via stoichiometric variation of nutrients in their dung.

Anthropogenic nutrient enrichment and shifts in herbivory can lead to dramatic changes in the composition and diversity of aboveground plant communities. In turn, this can alter seed banks in the soil, which are cryptic reservoirs of plant diversity. Here, we use data from seven Nutrient Network grassland sites on four continents, encompassing a range of climatic and environmental conditions, to test the joint effects of fertilization and aboveground mammalian herbivory on seed banks and on the similarity between aboveground plant communities and seed banks. We find that fertilization decreases plant species richness and diversity in seed banks, and homogenizes composition between aboveground and seed bank communities. Fertilization increases seed bank abundance especially in the presence of herbivores, while this effect is smaller in the absence of herbivores. Our findings highlight that nutrient enrichment can weaken a diversity maintaining mechanism in grasslands, and that herbivory needs to be considered when assessing nutrient enrichment effects on seed bank abundance. Seed banks are reservoirs of plant diversity. This study shows that nutrient addition decreases diversity of grassland seed banks, increases their similarity to aboveground communities and interacts with aboveground herbivory to affect their abundance.