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.
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.
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.
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.
Soil respiration is often used as an index of fertility because the majority of nutrients are cycled through the microbial biomass. We assessed the role of soil respiration as a measure of resource ...productivity in three long-term grassland experiments near Pietermaritzburg, South Africa. All of these experiments have shown significant changes in grass species composition and productivity. An ongoing Veld (=field) Fertilizer Experiment (VFE) that manipulated the level of nitrogen fertilizer, phosphorus and lime has been running since 1951. A Burning and Mowing Experiment (BME) has been running since 1950. The third experiment is part of the Nutrient Network (NutNet), a global fertilizer experiment that has been manipulating nitrogen, phosphorus, potassium and micronutrients and has been running since 2009. We found that longer-term experiments were more likely to show significant effects on soil respiration. We found several significant effects in the VFE but no significant differences in soil respiration among fertilization treatments in the shortest-term experiment (NutNet). In the VFE, we found significant differences in soil respiration due to levels of nitrogen fertilizer, form of nitrogen fertilizer (limestone ammonium nitrate and ammonium sulphate), phosphorus and lime. We found no significant relationship between above-ground net primary productivity and soil respiration despite the frequent detection of such a pattern due to the link between soil respiration, soil fertility and productivity. We found that, while there was a consistent increase in total soil nitrogen with increasing levels of nitrogen fertilizer applied, there was a consistent decrease in soil microbial respiration. There was a significant positive correlation between soil respiration and pH. Possible mechanisms behind this are unclear but may involve changes in dominant enzymes and possibly switches between dominance of bacteria and fungi. We also found significant effects of the timing of burning in the BME, but not due to the frequency of burning or the occurrence of mowing. Our results suggest that studies may need to be long-term, for example here at least 10 years, before key functional relationships with soil fertility can be reliably understood.
•We studied soil respiration in three adjacent long-term grassland experiments.•Longer experiments showed more significant variation than shorter experiments.•There was a negative correlation between soil respiration and N fertilization.•Soil respiration was positively correlated with soil pH.•Only season of burning was significant in a burning and mowing experiment.
Naked mole-rats (
) live in large colonies with one breeding female (queen), one to three breeding males (BMs) and the remainder are non-reproductive subordinates. The animals have a linear dominance ...rank with the breeders at the top of the hierarchy. We investigated how dominance rank in naked mole-rats differs with exploration (the propensity to explore a novel environment) and related endocrine markers. Exploration behaviour, faecal progestagen metabolite (fPM), faecal glucocorticoid metabolite (fGCM), faecal androgen metabolite (fAM) and plasma prolactin concentrations were quantified in breeding, high-, middle- and low-ranked females and males from five naked mole-rat colonies. There were no significant differences between the dominance rank and exploration behaviour. Interestingly, the queens and high-ranking females had higher fGCM and fAM concentrations compared with middle- and low-ranked females. The queens had significantly higher fPM concentrations than all other ranked females, since they are responsible for procreation. In the males, the BMs had higher fGCM concentrations compared with high- and low-ranked males. In addition, BMs and middle-ranking males had overall higher prolactin levels than all other ranked males, which could be linked to cooperative care. Overall, the results suggest that physiological reproductive suppression is linked to high dominance rank.
Soil stores approximately twice as much carbon as the atmosphere and fluctuations in the size of the soil carbon pool directly influence climate conditions. We used the Nutrient Network global change ...experiment to examine how anthropogenic nutrient enrichment might influence grassland soil carbon storage at a global scale. In isolation, enrichment of nitrogen and phosphorous had minimal impacts on soil carbon storage. However, when these nutrients were added in combination with potassium and micronutrients, soil carbon stocks changed considerably, with an average increase of 0.04 KgCm−2 year−1 (standard deviation 0.18 KgCm−2 year−1). These effects did not correlate with changes in primary productivity, suggesting that soil carbon decomposition may have been restricted. Although nutrient enrichment caused soil carbon gains most dry, sandy regions, considerable absolute losses of soil carbon may occur in high‐latitude regions that store the majority of the world's soil carbon. These mechanistic insights into the sensitivity of grassland carbon stocks to nutrient enrichment can facilitate biochemical modelling efforts to project carbon cycling under future climate scenarios.
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