The impacts of rising vapour pressure deficit in natural and managed ecosystems.

An exponential rise in the atmospheric vapour pressure deficit (VPD) is among the most consequential impacts of climate change in terrestrial ecosystems. Rising VPD has negative and cascading effects on nearly all aspects of plant function including photosynthesis, water status, growth and survival. These responses are exacerbated by land-atmosphere interactions that couple VPD to soil water and govern the evolution of drought, affecting a range of ecosystem services including carbon uptake, biodiversity, the provisioning of water resources and crop yields. However, despite the global nature of this phenomenon, research on how to incorporate these impacts into resilient management regimes is largely in its infancy, due in part to the entanglement of VPD trends with those of other co-evolving climate drivers. Here, we review the mechanistic bases of VPD impacts at a range of spatial scales, paying particular attention to the independent and interactive influence of VPD in the context of other environmental changes. We then evaluate the consequences of these impacts within key management contexts, including water resources, croplands, wildfire risk mitigation and management of natural grasslands and forests. We conclude with recommendations describing how management regimes could be altered to mitigate the otherwise highly deleterious consequences of rising VPD.

Clarifying the effect of biodiversity on productivity in natural ecosystems with longitudinal data and methods for causal inference.

Causal effects of biodiversity on ecosystem functions can be estimated using experimental or observational designs - designs that pose a tradeoff between drawing credible causal inferences from correlations and drawing generalizable inferences. Here, we develop a design that reduces this tradeoff and revisits the question of how plant species diversity affects productivity. Our design leverages longitudinal data from 43 grasslands in 11 countries and approaches borrowed from fields outside of ecology to draw causal inferences from observational data. Contrary to many prior studies, we estimate that increases in plot-level species richness caused productivity to decline: a 10% increase in richness decreased productivity by 2.4%, 95% CI [-4.1, -0.74]. This contradiction stems from two sources. First, prior observational studies incompletely control for confounding factors. Second, most experiments plant fewer rare and non-native species than exist in nature. Although increases in native, dominant species increased productivity, increases in rare and non-native species decreased productivity, making the average effect negative in our study. By reducing the tradeoff between experimental and observational designs, our study demonstrates how observational studies can complement prior ecological experiments and inform future ones.

Soil versus atmospheric drought: A test case of plant functional trait responses.

Climate change alters mean global surface temperatures, precipitation regimes, and atmospheric moisture. Resultant drought affects the composition and diversity of terrestrial ecosystems worldwide. To date, there have been no assessments of the combined impacts of reduced precipitation and atmospheric drying on functional trait distributions of any species in an outdoor experiment. Here, we examined whether soil and atmospheric drought affected the functional traits of a focal grass species (Poa secunda) growing in monoculture and eight-species grass communities in outdoor mesocosms. We focused on specific leaf area (SLA), leaf area, stomatal density, root:shoot ratio, and fine root:coarse root ratio responses. Leaf area and overall growth were reduced with soil drying. Root:shoot ratio only increased for P. secunda growing in monoculture under combined atmospheric and soil drought. Plant energy allocation strategy (measured using principal components) differed when P. secunda was grown in combined soil and atmospheric drought conditions compared with soil drought alone. Given a lack of outdoor manipulations of this kind, our results emphasize the importance of atmospheric drying on functional trait responses more broadly. We suggest that drought methods focused purely on soil water inputs may be imprecisely predicting drought effects on other terrestrial organisms as well (other plants, arthropods, and higher trophic levels).

Belowground facilitation and trait matching: two or three to tango?

High biodiversity increases ecosystem functions; however, belowground facilitation remains poorly understood in this context. Here, we explore mechanisms that operate via 'giving-receiving feedbacks' for belowground facilitation. These include direct effects via root exudates, signals, and root trait plasticity, and indirect biotic facilitation via the effects of root exudates on soil biota and feedback from biota to plants. We then highlight that these two- or three-way mechanisms must affect biodiversity-ecosystem function relationships via specific combinations of matching traits. To tango requires a powerful affinity and harmony between well-matched partners, and such matches link belowground facilitation to the effect of biodiversity on function. Such matching underpins applications in intercropping, forestry, and pasture systems, in which diversity contributes to greater productivity and sustainability.

The Future of Complementarity: Disentangling Causes from Consequences.

Evidence suggests that biodiversity supports ecosystem functioning. Yet, the mechanisms driving this relationship remain unclear. Complementarity is one common explanation for these positive biodiversity-ecosystem functioning relationships. Yet, complementarity is often indirectly quantified as overperformance in mixture relative to monoculture (e.g., 'complementarity effect'). This overperformance is then attributed to the intuitive idea of complementarity or, more specifically, to species resource partitioning. Locally, however, several unassociated causes may drive this overperformance. Here, we differentiate complementarity into three types of species differences that may cause enhanced ecosystem functioning in more diverse ecosystems: (i) resource partitioning, (ii) abiotic facilitation, and (iii) biotic feedbacks. We argue that disentangling these three causes is crucial for predicting the response of ecosystems to future biodiversity loss.

Plant species richness sustains higher trophic levels of soil nematode communities after consecutive environmental perturbations.

The magnitude and frequency of extreme weather events are predicted to increase in the future due to ongoing climate change. In particular, floods and droughts resulting from climate change are thought to alter the ecosystem functions and stability. However, knowledge of the effects of these weather events on soil fauna is scarce, although they are key towards functioning of terrestrial ecosystems. Plant species richness has been shown to affect the stability of ecosystem functions and food webs. Here, we used the occurrence of a natural flood in a biodiversity grassland experiment that was followed by a simulated summer drought experiment, to investigate the interactive effects of plant species richness, a natural flood, and a subsequent summer drought on nematode communities. Three and five months after the natural flooding, effects of flooding severity were still detectable in the belowground system. We found that flooding severity decreased soil nematode food-web structure (loss of K-strategists) and the abundance of plant feeding nematodes. However, high plant species richness maintained higher diversity and abundance of higher trophic levels compared to monocultures throughout the flood. The subsequent summer drought seemed to be of lower importance but reversed negative flooding effects in some cases. This probably occurred because the studied grassland system is well adapted to drought, or because drought conditions alleviated the negative impact of long-term soil waterlogging. Using soil nematodes as indicator taxa, this study suggests that high plant species richness can maintain soil food web complexity after consecutive environmental perturbations.

The Overlooked Role of Facilitation in Biodiversity Experiments.

Past research has demonstrated that decreased biodiversity often reduces ecosystem productivity, but variation in the shape of biodiversity-ecosystem function (BEF) relationships begets the need for a deeper mechanistic understanding of what drives these patterns. While mechanisms involving competition are often invoked, the role of facilitation is overlooked, or lumped within several less explicitly defined processes (e.g., complementarity effects). Here, we explore recent advances in understanding how facilitation affects BEF relationships and identify three categories of facilitative mechanisms that can drive variation in those relationships. Species interactions underlying BEF relationships are complex, but the framework we present provides a step toward understanding this complexity and predicting how facilitation contributes to the ecosystem role of biodiversity in a rapidly changing environment.

Plants are less negatively affected by flooding when growing in species-rich plant communities.

Flooding is expected to increase in frequency and severity in the future. The ecological consequences of flooding are the combined result of species-specific plant traits and ecological context. However, the majority of past flooding research has focused on individual model species under highly controlled conditions. An early summer flooding event in a grassland biodiversity experiment in Jena, Germany, provided the opportunity to assess flooding responses of 60 grassland species in monocultures and 16-species mixtures. We examined plant biomass, species-specific traits (plant height, specific leaf area (SLA), root aerenchyma, starch content) and soil porosity. We found that, on average, plant species were less negatively affected by the flood when grown in higher-diversity plots in July 2013. By September 2013, grasses were unaffected by the flood regardless of plant diversity, and legumes were severely negatively affected regardless of plant diversity. Plants with greater SLA and more root aerenchyma performed better in September. Soil porosity was higher in higher-diversity plots and had a positive effect on plant performance. As floods become more frequent and severe in the future, growing flood-sensitive plants in higher-diversity communities and in soil with greater soil aeration may attenuate the most negative effects of flooding.

Plant species richness and functional traits affect community stability after a flood event.

Climate change is expected to increase the frequency and magnitude of extreme weather events. It is therefore of major importance to identify the community attributes that confer stability in ecological communities during such events. In June 2013, a flood event affected a plant diversity experiment in Central Europe (Jena, Germany). We assessed the effects of plant species richness, functional diversity, flooding intensity and community means of functional traits on different measures of stability (resistance, resilience and raw biomass changes from pre-flood conditions). Surprisingly, plant species richness reduced community resistance in response to the flood. This was mostly because more diverse communities grew more immediately following the flood. Raw biomass increased over the previous year; this resulted in decreased absolute value measures of resistance. There was no clear response pattern for resilience. We found that functional traits drove these changes in raw biomass: communities with a high proportion of late-season, short-statured plants with dense, shallow roots and small leaves grew more following the flood. Late-growing species probably avoided the flood, whereas greater root length density might have allowed species to better access soil resources brought from the flood, thus growing more in the aftermath. We conclude that resource inputs following mild floods may favour the importance of traits related to resource acquisition and be less associated with flooding tolerance.

Shifting grassland plant community structure drives positive interactive effects of warming and diversity on aboveground net primary productivity.

Ecosystems worldwide are increasingly impacted by multiple drivers of environmental change, including climate warming and loss of biodiversity. We show, using a long-term factorial experiment, that plant diversity loss alters the effects of warming on productivity. Aboveground primary productivity was increased by both high plant diversity and warming, and, in concert, warming (≈1.5 °C average above and belowground warming over the growing season) and diversity caused a greater than additive increase in aboveground productivity. The aboveground warming effects increased over time, particularly at higher levels of diversity, perhaps because of warming-induced increases in legume and C4 bunch grass abundances, and facilitative feedbacks of these species on productivity. Moreover, higher plant diversity was associated with the amelioration of warming-induced environmental conditions. This led to cooler temperatures, decreased vapor pressure deficit, and increased surface soil moisture in higher diversity communities. Root biomass (0-30 cm) was likewise consistently greater at higher plant diversity and was greater with warming in monocultures and at intermediate diversity, but at high diversity warming had no detectable effect. This may be because warming increased the abundance of legumes, which have lower root : shoot ratios than the other types of plants. In addition, legumes increase soil nitrogen (N) supply, which could make N less limiting to other species and potentially decrease their investment in roots. The negative warming × diversity interaction on root mass led to an overall negative interactive effect of these two global change factors on the sum of above and belowground biomass, and thus likely on total plant carbon stores. In total, plant diversity increased the effect of warming on aboveground net productivity and moderated the effect on root mass. These divergent effects suggest that warming and changes in plant diversity are likely to have both interactive and divergent impacts on various aspects of ecosystem functioning.

Flooding disturbances increase resource availability and productivity but reduce stability in diverse plant communities.

The natural world is increasingly defined by change. Within the next 100 years, rising atmospheric CO2 concentrations will continue to increase the frequency and magnitude of extreme weather events. Simultaneously, human activities are reducing global biodiversity, with current extinction rates at ~1,000 × what they were before human domination of Earth's ecosystems. The co-occurrence of these trends may be of particular concern, as greater biological diversity could help ecosystems resist change during large perturbations. We use data from a 200-year flood event to show that when a disturbance is associated with an increase in resource availability, the opposite may occur. Flooding was associated with increases in productivity and decreases in stability, particularly in the highest diversity communities. Our results undermine the utility of the biodiversity-stability hypothesis during a large number of disturbances where resource availability increases. We propose a conceptual framework that can be widely applied during natural disturbances.