Niche dimensionality provides a general theoretical explanation for biodiversity-more niches, defined by more limiting factors, allow for more ways that species can coexist. Because plant species compete for the same set of limiting resources, theory predicts that addition of a limiting resource eliminates potential trade-offs, reducing the number of species that can coexist. Multiple nutrient limitation of plant production is common and therefore fertilization may reduce diversity by reducing the number or dimensionality of belowground limiting factors. At the same time, nutrient addition, by increasing biomass, should ultimately shift competition from belowground nutrients towards a one-dimensional competitive trade-off for light. Here we show that plant species diversity decreased when a greater number of limiting nutrients were added across 45 grassland sites from a multi-continent experimental network. The number of added nutrients predicted diversity loss, even after controlling for effects of plant biomass, and even where biomass production was not nutrient-limited. We found that elevated resource supply reduced niche dimensionality and diversity and increased both productivity and compositional turnover. Our results point to the importance of understanding dimensionality in ecological systems that are undergoing diversity loss in response to multiple global change factors.

Symbiotic microorganisms that live intimately associated with terrestrial plants affect both the quantity and quality of resources, and thus the energy supply to consumer populations at higher levels in the food chain. Empirical evidence on resource limitation of food webs points to primary productivity as a major determinant of consumer abundance and trophic structure. Prey quality plays a critical role in community regulation. Plants infected by endophytic fungi are known to be chemically protected against herbivore consumption. However, the influence of this microbe-plant association on multi-trophic interactions remains largely unexplored. Here we present the effects of fungal endophytes on insect food webs that reflect limited energy transfer to consumers as a result of low plant quality, rather than low productivity. Herbivore-parasite webs on endophyte-free grasses show enhanced insect abundance at alternate trophic levels, higher rates of parasitism, and increased dominance by a few trophic links. These results mirror predicted effects of increased productivity on food-web dynamics. Thus 'hidden' microbial symbionts can have community-wide impacts on the pattern and strength of resource-consumer interactions.

2000. Enemy-mediated apparent competition: empirical patterns and the evidence. -Oikos 88: 380 -394.Apparent competition arises when two victim species negatively affect each other (− , − ) by enhancing the equilibrium density or changing the foraging behaviour of a shared natural enemy. Shared enemies can also mediate non-reciprocal ( −, 0) indirect effects, i.e. indirect amensalism, whenever one prey species is not affected by the presence of alternative prey. We review 34 studies on terrestrial and freshwater systems to evaluate the extent to which apparent competition has been perceived as a reciprocal ( −, −) or non-reciprocal ( − , 0) interaction. We found only three studies showing reciprocal effects between apparent competitors. Indirect amensalism was documented in 10 studies and could be inferred for 16 other cases (76% in total). The remaining five studies provided insufficient data to determine the form of indirect interaction. The apparent prevalence of non-reciprocal enemy-mediated interactions resembles that observed for resource-based interspecific competition. Amensal indirect effects via shared predation may result from differences in population size, nutritional value, susceptibility to attack, or asynchronous dynamics of alternative prey, or the predator's feeding preferences. Moreover, experimental protocols may confound the actual form of apparent competition through short-term observations, incomplete designs, or biased consideration of conspicuous interactions, leading to reciprocal effects being overlooked. We conclude that, at present, it is still difficult to determine the relative role of apparent competition vs indirect amensalism in natural food webs because most published studies have failed to document in full interactions via shared enemies.

Isolating the single effects and net balance of negative and positive species effects in complex interaction networks is a necessary step for understanding community dynamics. Facilitation and competition have both been found to operate in harsh environments, but their relative strength may be predicted to change along gradients of herbivory. Moreover, facilitation effects through habitat amelioration and protection from herbivory may act together determining the outcome of neighborhood plant-plant interactions. We tested the hypothesis that grazing pressure alters the balance of positive and negative interactions between palatable and unpalatable species by increasing the strength of positive indirect effects mediated by associational resistance to herbivory. We conducted a two-year factorial experiment in which distance (i.e., spatial association) from the nearest unpalatable neighbor (Stipa speciosa) and root competition were manipulated for two palatable grasses (Poa ligularis and Bromus pictus), at three levels of sheep grazing (none, moderate, and high) in a Patagonian steppe community. We found that grazing shifted the effect of Stipa on both palatable grasses, from negative (competition) in the absence of grazing to positive (facilitation) under increasing herbivore pressure. In ungrazed sites, belowground competition was the dominant interaction, as shown by a significant reduction in performance of palatable grasses transplanted near to Stipa tussocks. In grazed sites, biomass of palatable plants was greater near than far from Stipa regardless of competition treatment. Proximity to Stipa reduced the amount of herbivory suffered by palatable grasses, an indirect effect that was stronger under moderate than under intense grazing. Our results demonstrate that facilitation, resulting mainly from protection against herbivory, is the overriding effect produced by unpalatable neighbors on palatable grasses in this rangeland community. This finding challenges the common view that abiotic stress amelioration should be the predominant type of facilitation in arid environments and highlights the role of herbivory in modulating complex neighborhood plant interactions in grazing systems.

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.

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Questions Does the magnitude of grazing‐induced changes in species composition vary with habitat productivity? How does the sign and magnitude of grazing effects on species richness and beta‐diversity change with increasing productivity? Do major life forms exhibit consistent responses to grazing along productivity gradients? Location Steppes and grasslands of southern South America in Argentina and Uruguay. Methods We evaluated grazing effects on plant composition, species richness, beta‐diversity and life‐form abundances along a ten‐fold, regional productivity gradient and within subregions of contrasting productivity, using a common sampling protocol for 23 paired grazed vs ungrazed plots. The annual integral of the normalized difference vegetation index was used as a surrogate for above‐ground net primary productivity. Results Compositional dissimilarity between grazed and ungrazed plots, as well as grazing‐induced differences in plant richness and beta‐diversity all increased with habitat productivity. Grazing decreased species richness in low‐productive steppes but enhanced the richness of high‐productive grasslands. On average, grazing reduced beta‐diversity in high‐productive sites but not in low‐productive sites. Dominant species were more strongly suppressed by grazing towards productive grasslands. Grazing generally decreased shrub species cover, whereas graminoid and forb cover did not consistently change with grazing through the productivity gradient. Conclusions Our results indicate that the overall grazing effects on vegetation structure increased along a regional productivity gradient. Yet the sign of grazing impacts on species richness and beta‐diversity shifted with habitat productivity, in agreement with models of herbivore‐mediated co‐existence and species colonization in productive systems. Further, we found that narrowing the spatial extent of analysis to the subregion generally obscured grazing–productivity relationships. Biodiversity conservation programmes should carefully weigh the varied impacts of livestock grazing across productivity gradients.

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity–ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.