Miguel García-Gómez

Plant species richness and ecosystem multifunctionality in global drylands

Global Ecosystem Analysis The relationship between species richness and the functional properties of their ecosystems has often been studied at small scales in experimental plots. Maestre et al. (p. 214; see the Perspective by Midgley) performed field measurements at 224 dryland sites from six continents and assessed 14 ecosystem functions related to carbon, nitrogen, and phosphorus cycling. Positive relationships were observed between perennial plant species richness and ecosystem functionality. The relative importance of biodiversity was found to be as large as, or larger than, many key abiotic variables. Thus, preservation of plant biodiversity is important to buffer negative effects of climate change and desertification in drylands, which collectively cover 41% of Earths land surface and support over 38% of the human population. Plant species richness is positively related to ecosystem multifunctionality in drylands at a global scale. Experiments suggest that biodiversity enhances the ability of ecosystems to maintain multiple functions, such as carbon storage, productivity, and the buildup of nutrient pools (multifunctionality). However, the relationship between biodiversity and multifunctionality has never been assessed globally in natural ecosystems. We report here on a global empirical study relating plant species richness and abiotic factors to multifunctionality in drylands, which collectively cover 41% of Earth’s land surface and support over 38% of the human population. Multifunctionality was positively and significantly related to species richness. The best-fitting models accounted for over 55% of the variation in multifunctionality and always included species richness as a predictor variable. Our results suggest that the preservation of plant biodiversity is crucial to buffer negative effects of climate change and desertification in drylands.

Increasing aridity reduces soil microbial diversity and abundance in global drylands.

Significance Climate change is increasing the degree of aridity in drylands, which occupy 41% of Earth’s surface and support 38% of its population. Soil bacteria and fungi are largely responsible for key ecosystem services, including soil fertility and climate regulation, yet their responses to changes in aridity are poorly understood. Using a field survey conducted in drylands worldwide and DNA-sequencing approaches, we found that increases in aridity reduce the diversity and abundance of soil bacteria and fungi. This study represents an important advancement in our understanding of soil microbial communities and their likely responses to ongoing climate change. Soil bacteria and fungi play key roles in the functioning of terrestrial ecosystems, yet our understanding of their responses to climate change lags significantly behind that of other organisms. This gap in our understanding is particularly true for drylands, which occupy ∼41% of Earth´s surface, because no global, systematic assessments of the joint diversity of soil bacteria and fungi have been conducted in these environments to date. Here we present results from a study conducted across 80 dryland sites from all continents, except Antarctica, to assess how changes in aridity affect the composition, abundance, and diversity of soil bacteria and fungi. The diversity and abundance of soil bacteria and fungi was reduced as aridity increased. These results were largely driven by the negative impacts of aridity on soil organic carbon content, which positively affected the abundance and diversity of both bacteria and fungi. Aridity promoted shifts in the composition of soil bacteria, with increases in the relative abundance of Chloroflexi and α-Proteobacteria and decreases in Acidobacteria and Verrucomicrobia. Contrary to what has been reported by previous continental and global-scale studies, soil pH was not a major driver of bacterial diversity, and fungal communities were dominated by Ascomycota. Our results fill a critical gap in our understanding of soil microbial communities in terrestrial ecosystems. They suggest that changes in aridity, such as those predicted by climate-change models, may reduce microbial abundance and diversity, a response that will likely impact the provision of key ecosystem services by global drylands.

Uncovering multiscale effects of aridity and biotic interactions on the functional structure of Mediterranean shrublands

Summary 1. Habitat filtering (HF, trait convergence) and niche differentiation (ND, trait divergence) are known to impact upon plant community structure. Both processes integrate individual responses to the abiotic environment and biotic interactions. Thus, it is difficult to clearly identify the underlying abiotic and biotic factors that ultimately impact community structure by looking at community-level patterns of trait divergence or convergence alone. 2. We used a functional trait-based and multiscale approach to assess how biotic interactions and aridity determine the functional structure of semi-arid shrublands sampled along a large aridity gradient in Spain. At the regional scale, we investigated functional differences among species (axes of specialization) to identify important traits for community assembly. At the community scale, we evaluated the relative impact of HF and ND on community structure using a null model approach. Finally, at the plant neighbourhood scale, we evaluated the impact of biotic interactions on community structure by investigating the spatial patterns of trait aggregation. 3. The shrub species surveyed can be separated along four axes of specialization based on their above-ground architecture and leaf morphology. Our community scale analysis suggested that the functional structure of semi-arid communities was clearly non-random, HF and ND acting independently on different traits to determine community structure along the aridity gradient. At the plant neighbourhood scale, the spatial distribution of species was also clearly not random, suggesting that competition and facilitation impacted on the observed changes in the functional diversity of shrubland communities along the aridity gradient. 4. Synthesis: Our results demonstrated that HF and ND acted simultaneously on independent traits to jointly determine community structure. Most importantly, our multiscale approach suggested that competition and facilitation interplayed with aridity to determine this structure. Competition appeared to be constant along the aridity gradient and explained the high functional diversity observed in semi-arid shrublands. Facilitation affected subordinate and rare species and, thus, may act to enhance the biodiversity of these ecosystems. Finally, the framework employed in our study allows moving forward from the examination of patterns to the development of mechanistic traitbased approaches to study plant community assembly.

Climate and soil attributes determine plant species turnover in global drylands

AIM Geographic, climatic, and soil factors are major drivers of plant beta diversity, but their importance for dryland plant communities is poorly known. This study aims to: i) characterize patterns of beta diversity in global drylands, ii) detect common environmental drivers of beta diversity, and iii) test for thresholds in environmental conditions driving potential shifts in plant species composition. LOCATION 224 sites in diverse dryland plant communities from 22 geographical regions in six continents. METHODS Beta diversity was quantified with four complementary measures: the percentage of singletons (species occurring at only one site), Whittakes beta diversity (β(W)), a directional beta diversity metric based on the correlation in species occurrences among spatially contiguous sites (β(R2)), and a multivariate abundance-based metric (β(MV)). We used linear modelling to quantify the relationships between these metrics of beta diversity and geographic, climatic, and soil variables. RESULTS Soil fertility and variability in temperature and rainfall, and to a lesser extent latitude, were the most important environmental predictors of beta diversity. Metrics related to species identity (percentage of singletons and β(W)) were most sensitive to soil fertility, whereas those metrics related to environmental gradients and abundance ((β(R2)) and β(MV)) were more associated with climate variability. Interactions among soil variables, climatic factors, and plant cover were not important determinants of beta diversity. Sites receiving less than 178 mm of annual rainfall differed sharply in species composition from more mesic sites (> 200 mm). MAIN CONCLUSIONS Soil fertility and variability in temperature and rainfall are the most important environmental predictors of variation in plant beta diversity in global drylands. Our results suggest that those sites annually receiving ~ 178 mm of rainfall will be especially sensitive to future climate changes. These findings may help to define appropriate conservation strategies for mitigating effects of climate change on dryland vegetation.

Aridity Modulates N Availability in Arid and Semiarid Mediterranean Grasslands

While much is known about the factors that control each component of the terrestrial nitrogen (N) cycle, it is less clear how these factors affect total N availability, the sum of organic and inorganic forms potentially available to microorganisms and plants. This is particularly true for N-poor ecosystems such as drylands, which are highly sensitive to climate change and desertification processes that can lead to the loss of soil nutrients such as N. We evaluated how different climatic, abiotic, plant and nutrient related factors correlate with N availability in semiarid Stipa tenacissima grasslands along a broad aridity gradient from Spain to Tunisia. Aridity had the strongest relationship with N availability, suggesting the importance of abiotic controls on the N cycle in drylands. Aridity appeared to modulate the effects of pH, plant cover and organic C (OC) on N availability. Our results suggest that N transformation rates, which are largely driven by variations in soil moisture, are not the direct drivers of N availability in the studied grasslands. Rather, the strong relationship between aridity and N availability could be driven by indirect effects that operate over long time scales (decades to millennia), including both biotic (e.g. plant cover) and abiotic (e.g. soil OC and pH). If these factors are in fact more important than short-term effects of precipitation on N transformation rates, then we might expect to observe a lagged decrease in N availability in response to increasing aridity. Nevertheless, our results suggest that the increase in aridity predicted with ongoing climate change will reduce N availability in the Mediterranean basin, impacting plant nutrient uptake and net primary production in semiarid grasslands throughout this region.

Traits of neighbouring plants and space limitation determine intraspecific trait variability in semi‐arid shrublands

Understanding how intraspecific trait variability (ITV) responds to both abiotic and biotic constraints is crucial to predict how individuals are assembled in plant communities, and how they will be impacted by ongoing global environmental change. Three key functional traits [plant height, leaf area (LA) and specific leaf area (SLA)] were assessed to quantify the range of ITV of four dominant plant species along a rainfall gradient in semi-arid Mediterranean shrublands. Variance partitioning and confirmatory multilevel path analyses were used to assess the direct and indirect effects of rainfall, space limitation (crowding) and neighbouring plant traits on ITV. The direct effect of the local neighbourhood on the trait values of subordinate individuals was as strong as the effect of rainfall. The indirect effect of rainfall, however, mediated by the effect of the local neighbourhood on the trait values of subordinate individuals, was weak. Rainfall decreased the height and SLA of subordinate individuals, but increased their LA. Neighbouring plant traits were just as strong predictors as crowding in explaining changes in ITV.Synthesis. Our study provides a framework to disentangle the direct effects of abiotic factors and their indirect effects on ITV mediated by the local neighbourhood. Our results highlight that abiotic and biotic constraints are both substantial sources of trait variations at the individual level, and can blur processes underlying changes in ITV. Considering and disentangling combined sources with an individual perspective would help to refine our predictions for community assembly and functional ecology.

Soil fungal abundance and plant functional traits drive fertile island formation in global drylands

Dryland vegetation is characterized by discrete plant patches that accumulate and capture soil resources under their canopies. These “fertile islands” are major drivers of dryland ecosystem structure and functioning, yet we lack an integrated understanding of the factors controlling their magnitude and variability at the global scale. We conducted a standardized field survey across 236 drylands from five continents. At each site, we measured the composition, diversity and cover of perennial plants. Fertile island effects were estimated at each site by comparing composite soil samples obtained under the canopy of the dominant plants and in open areas devoid of perennial vegetation. For each sample, we measured 15 soil variables (functions) associated with carbon, nitrogen and phosphorus cycling and used the relative interaction index to quantify the magnitude of the fertile island effect for each function. In 80 sites, we also measured fungal and bacterial abundance (quantitative PCR) and diversity (Illumina MiSeq). The most fertile islands, i.e. those where a higher number of functions were simultaneously enhanced, were found at lower elevation sites with greater soil pH values and sand content under semiarid climates, particularly at locations where the presence of tall woody species with a low-specific leaf area increased fungal abundance beneath plant canopies, the main direct biotic controller of the fertile island effect in the drylands studied. Positive effects of fungal abundance were particularly associated with greater nutrient contents and microbial activity (soil extracellular enzymes) under plant canopies. Synthesis. Our results show that the formation of fertile islands in global drylands largely depends on: (1) local climatic, topographic and edaphic characteristics, (2) the structure and traits of local plant communities and (3) soil microbial communities. Our study also has broad implications for the management and restoration of dryland ecosystems worldwide, where woody plants are commonly used as nurse plants to enhance the establishment and survival of beneficiary species. Finally, our results suggest that forecasted increases in aridity may enhance the formation of fertile islands in drylands worldwide.