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Implementing precision fertilization to maximize crop yield while minimizing economic and environmental impacts has become critical for agriculture. Variability in biomass response to fertilization within fields, among regions, and over time creates simultaneous risks of under-yielding and overfertilization. We quantify factors determining fertilization responsiveness (i.e., biomass increases with fertilization) up to 15 years in 61 unfertilized rangelands on six continents. We demonstrate widespread multi-year variability in responsiveness, with fertilization increasing average yield by 43% but failing to improve biomass 26% of the time. All sites were responsive at least once, but only four of 61 responded in all plots and years. Modelled management scenarios highlighted that fertilizer cessation is likely to generate sizable economic savings but always reduces yield because of the difficulty in predicting when and where biomass will be unresponsive. This work reveals substantial scale-dependent variability in fertilization responsiveness globally, while clarifying the prospects and pitfalls of managing more spatially and temporally precise nutrient application.

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining ecosystem functioning remains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis of eight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands—those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)—had higher levels of multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities. Analysis of 65 grasslands worldwide from the Nutrient Network experiment reveals that plant communities with higher α- and β-diversity have higher levels of ecosystem multifunctionality, and that this effect is amplified across scales.

Nutrient enrichment typically causes local plant diversity declines. A common but untested expectation is that nutrient enrichment also reduces variation in nutrient conditions among localities and selects for a smaller pool of species, causing greater diversity declines at larger than local scales and thus biotic homogenization. Here we apply a framework that links changes in species richness across scales to changes in the numbers of spatially restricted and widespread species for a standardized nutrient addition experiment across 72 grasslands on six continents. Overall, we find proportionally similar species loss at local and larger scales, suggesting similar declines of spatially restricted and widespread species, and no biotic homogenization after 4 years and up to 14 years of treatment. These patterns of diversity changes are generally consistent across species groups. Thus, nutrient enrichment poses threats to plant diversity, including for widespread species that are often critical for ecosystem functions.

The distribution of flowering across the growing season is governed by each species' evolutionary history and climatic variability. However, global change factors, such as eutrophication and invasion, can alter plant community composition and thus change the distribution of flowering across the growing season. We examined three ecoregions (tall-, mixed, and short-grass prairie) across the U.S. Central Plains to determine how nutrient (nitrogen (N), phosphorus, and potassium (+micronutrient)) addition alters the temporal patterns of plant flowering traits. We calculated total community flowering potential (FP) by distributing peak-season plant cover values across the growing season, allocating each species' cover to only those months in which it typically flowers. We also generated separate FP profiles for exotic and native species and functional group. We compared the ability of the added nutrients to shift the distribution of these FP profiles (total and sub-groups) across the growing season. In all ecoregions, N increased the relative cover of both exotic species and C3 graminoids that flower in May through August. The cover of C4 graminoids decreased with added N, but the response varied by ecoregion and month. However, these functional changes only aggregated to shift the entire community's FP profile in the tall-grass prairie, where the relative cover of plants expected to flower in May and June increased and those that flower in September and October decreased with added N. The relatively low native cover in May and June may leave this ecoregion vulnerable to disturbance-induced invasion by exotic species that occupy this temporal niche. There was no change in the FP profile of the mixed and short-grass prairies with N addition as increased abundance of exotic species and C3 graminoids replaced other species that flower at the same time. In these communities a disturbance other than nutrient addition may be required to disrupt phenological patterns.

Herbivory and fire shape plant community structure in grass-dominated ecosystems, but these disturbance regimes are being altered around the world. To assess the consequences of such alterations, we excluded large herbivores for seven years from mesic savanna grasslands sites burned at different frequencies in North America (Konza Prairie Biological Station, Kansas, USA) and South Africa (Kruger National Park). We hypothesized that the removal of a single grass-feeding herbivore from Konza would decrease plant community richness and shift community composition due to increased dominance by grasses. Similarly, we expected grass dominance to increase at Kruger when removing large herbivores, but because large herbivores are more diverse, targeting both grasses and forbs, at this study site, the changes due to herbivore removal would be muted. After seven years of large-herbivore exclusion, richness strongly decreased and community composition changed at Konza, whereas little change was evident at Kruger. We found that this divergence in response was largely due to differences in the traits and numbers of dominant grasses between the study sites rather than the predicted differences in herbivore assemblages. Thus, the diversity of large herbivores lost may be less important in determining plant community dynamics than the functional traits of the grasses that dominate mesic, disturbance-maintained savanna grasslands.

The naked mole-rat (Heterocephalus glaber) occurs in colonies with a distinct dominance hierarchy, including one dominant, breeding female (the queen), 1–3 breeding males, and non-reproductive subordinates of both sexes that are reproductively suppressed while in the colony. To non-invasively evaluate reproductive capacity in the species, we first had to examine the suitability of enzyme immunoassays (EIAs) for determining progestogen and androgen metabolite concentrations in the naked mole-rat, using urine and faeces. A saline control and gonadotrophin-releasing hormone (GnRH) were administered to twelve (six males and six females) naked mole-rats which were previously identified as dispersers and housed singly. The results revealed that urine is possibly not an ideal matrix for progestogen and androgen metabolite quantification in naked mole-rats as no signal was detected in the matrix post GnRH administration. A 5α-Progesterone EIA and an Epiandrosterone EIA were identified as suitable for quantifying faecal progesterone metabolites (fPMs) and faecal androgen metabolites (fAMs) in males and females, respectively. The results suggest that there are individual variations in baseline fPM and fAM concentrations, and only two out of six females and no males exhibited an increase in fPM concentrations greater than 100% (−20% SD) post GnRH administration. Conversely, only four out of six females and three out of six males had an increase in fAM concentrations greater than 100% (−20% SD) following GnRH administration. These results imply that some naked mole-rat individuals have a reduced reproductive capacity even when they are separated from the queen.

Numerous studies show that increasing species richness leads to higher ecosystem productivity. This effect is often attributed to more efficient portioning of multiple resources in communities with higher numbers of competing species, indicating the role of resource supply and stoichiometry for biodiversity–ecosystem functioning relationships. Here, we merged theory on ecological stoichiometry with a framework of biodiversity–ecosystem functioning to understand how resource use transfers into primary production. We applied a structural equation model to define patterns of diversity–productivity relationships with respect to available resources. Meta-analysis was used to summarize the findings across ecosystem types ranging from aquatic ecosystems to grasslands and forests. As hypothesized, resource supply increased realized productivity and richness, but we found significant differences between ecosystems and study types. Increased richness was associated with increased productivity, although this effect was not seen in experiments. More even communities had lower productivity, indicating that biomass production is often maintained by a few dominant species, and reduced dominance generally reduced ecosystem productivity. This synthesis, which integrates observational and experimental studies in a variety of ecosystems and geographical regions, exposes common patterns and differences in biodiversity–functioning relationships, and increases the mechanistic understanding of changes in ecosystems productivity.

In areas with diverse herbivore communities such as African savannas, the frequency of disturbance by fire may alter the top–down role of different herbivore species on plant community dynamics. In a seven year experiment in the Kruger National Park, South Africa, we examined the habitat use of nine common herbivore species across annually burned, triennially burned and unburned areas. We also used two types of exclosures (plus open access controls) to examine the impacts of different herbivores on plant community dynamics across fire disturbance regimes. Full exclosures excluded all herbivores > 0.5 kg (e.g. elephant, zebra, impala) while partial exclosures allowed access only to animals with shoulder heights ≤ 0.85 m (e.g. impala, steenbok). Annual burns attracted a diverse suite of herbivores, and exclusion of larger herbivores (e.g. elephant, zebra, wildebeest) increased plant abundance. When smaller species, mainly impala, were also excluded there were declines in plant diversity, likely mediated by a decline in open space available for colonization of uncommon plant species. Unburned areas attracted the least diverse suite of herbivores, dominated by impala. Here, herbivore exclusion, especially of impala, led to strong declines in plant richness and diversity. With no fire disturbance, herbivore exclusion led to competitive exclusion via increases in plant dominance and light limitation. In contrast, on triennial burns, herbivore exclusion had no effect on plant richness or diversity, potentially due to relatively little open space for colonization across exclosure treatments but also little competitive exclusion due to the intermediate fire disturbance. Further, the diverse suite of grazers and browsers on triennial burns may have had a compensating effect of on the diversity of grasses and forbs. Ultimately, our work shows that differential disturbance regimes can result in differential consumer pressure across a landscape and result in heterogeneous patterns in top–down control of community dynamics.

Grasslands cover approximately a third of the Earth’s land surface and account for about a third of terrestrial carbon storage. Yet, we lack strong predictive models of grassland plant biomass, the primary source of carbon in grasslands. This lack of predictive ability may arise from the assumption of linear relationships between plant biomass and the environment and an underestimation of interactions of environmental variables. Using data from 116 grasslands on six continents, we show unimodal relationships between plant biomass and ecosystem characteristics, such as mean annual precipitation and soil nitrogen. Further, we found that soil nitrogen and plant diversity interacted in their relationships with plant biomass, such that plant diversity and biomass were positively related at low levels of nitrogen and negatively at elevated levels of nitrogen. Our results show that it is critical to account for the interactive and unimodal relationships between plant biomass and several environmental variables to accurately include plant biomass in global vegetation and carbon models. A study using data from grasslands on six continents finds unimodal relationships between plant biomass and abiotic ecosystem characteristics and also shows that soil nitrogen and plant diversity interact in their relationships with plant biomass.

Little is currently known about how climate modulates the relationship between plant diversity and soil organic carbon and the mechanisms involved. Yet, this knowledge is of crucial importance in times of climate change and biodiversity loss. Here, we show that plant diversity is positively correlated with soil carbon content and soil carbon-to-nitrogen ratio across 84 grasslands on six continents that span wide climate gradients. The relationships between plant diversity and soil carbon as well as plant diversity and soil organic matter quality (carbon-to-nitrogen ratio) are particularly strong in warm and arid climates. While plant biomass is positively correlated with soil carbon, plant biomass is not significantly correlated with plant diversity. Our results indicate that plant diversity influences soil carbon storage not via the quantity of organic matter (plant biomass) inputs to soil, but through the quality of organic matter. The study implies that ecosystem management that restores plant diversity likely enhances soil carbon sequestration, particularly in warm and arid climates.