Cristina Escolar

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.

Decoupling of soil nutrient cycles as a function of aridity in global drylands

The biogeochemical cycles of carbon (C), nitrogen (N) and phosphorus (P) are interlinked by primary production, respiration and decomposition in terrestrial ecosystems. It has been suggested that the C, N and P cycles could become uncoupled under rapid climate change because of the different degrees of control exerted on the supply of these elements by biological and geochemical processes. Climatic controls on biogeochemical cycles are particularly relevant in arid, semi-arid and dry sub-humid ecosystems (drylands) because their biological activity is mainly driven by water availability. The increase in aridity predicted for the twenty-first century in many drylands worldwide may therefore threaten the balance between these cycles, differentially affecting the availability of essential nutrients. Here we evaluate how aridity affects the balance between C, N and P in soils collected from 224 dryland sites from all continents except Antarctica. We find a negative effect of aridity on the concentration of soil organic C and total N, but a positive effect on the concentration of inorganic P. Aridity is negatively related to plant cover, which may favour the dominance of physical processes such as rock weathering, a major source of P to ecosystems, over biological processes that provide more C and N, such as litter decomposition. Our findings suggest that any predicted increase in aridity with climate change will probably reduce the concentrations of N and C in global drylands, but increase that of P. These changes would uncouple the C, N and P cycles in drylands and could negatively affect the provision of key services provided by these ecosystems.

Changes in biocrust cover drive carbon cycle responses to climate change in drylands.

Dryland ecosystems account for ca. 27% of global soil organic carbon (C) reserves, yet it is largely unknown how climate change will impact C cycling and storage in these areas. In drylands, soil C concentrates at the surface, making it particularly sensitive to the activity of organisms inhabiting the soil uppermost levels, such as communities dominated by lichens, mosses, bacteria and fungi (biocrusts). We conducted a full factorial warming and rainfall exclusion experiment at two semiarid sites in Spain to show how an average increase of air temperature of 2-3 °C promoted a drastic reduction in biocrust cover (ca. 44% in 4 years). Warming significantly increased soil CO2 efflux, and reduced soil net CO2 uptake, in biocrust-dominated microsites. Losses of biocrust cover with warming through time were paralleled by increases in recalcitrant C sources, such as aromatic compounds, and in the abundance of fungi relative to bacteria. The dramatic reduction in biocrust cover with warming will lessen the capacity of drylands to sequester atmospheric CO2 . This decrease may act synergistically with other warming-induced effects, such as the increase in soil CO2 efflux and the changes in microbial communities to alter C cycling in drylands, and to reduce soil C stocks in the mid to long term.

Do biotic interactions modulate ecosystem functioning along stress gradients? Insights from semi-arid plant and biological soil crust communities

Climate change will exacerbate the degree of abiotic stress experienced by semi-arid ecosystems. While abiotic stress profoundly affects biotic interactions, their potential role as modulators of ecosystem responses to climate change is largely unknown. Using plants and biological soil crusts, we tested the relative importance of facilitative–competitive interactions and other community attributes (cover, species richness and species evenness) as drivers of ecosystem functioning along stress gradients in semi-arid Mediterranean ecosystems. Biotic interactions shifted from facilitation to competition along stress gradients driven by water availability and temperature. These changes were, however, dependent on the spatial scale and the community considered. We found little evidence to suggest that biotic interactions are a major direct influence upon indicators of ecosystem functioning (soil respiration, organic carbon, water-holding capacity, compaction and the activity of enzymes related to the carbon, nitrogen and phosphorus cycles) along stress gradients. However, attributes such as cover and species richness showed a direct effect on ecosystem functioning. Our results do not agree with predictions emphasizing that the importance of plant–plant interactions will be increased under climate change in dry environments, and indicate that reductions in the cover of plant and biological soil crust communities will negatively impact ecosystems under future climatic conditions.

Warming reduces the growth and diversity of biological soil crusts in a semi-arid environment: implications for ecosystem structure and functioning

Biological soil crusts (BSCs) are key biotic components of dryland ecosystems worldwide that control many functional processes, including carbon and nitrogen cycling, soil stabilization and infiltration. Regardless of their ecological importance and prevalence in drylands, very few studies have explicitly evaluated how climate change will affect the structure and composition of BSCs, and the functioning of their constituents. Using a manipulative experiment conducted over 3 years in a semi-arid site from central Spain, we evaluated how the composition, structure and performance of lichen-dominated BSCs respond to a 2.4°C increase in temperature, and to an approximately 30 per cent reduction of total annual rainfall. In areas with well-developed BSCs, warming promoted a significant decrease in the richness and diversity of the whole BSC community. This was accompanied by important compositional changes, as the cover of lichens suffered a substantial decrease with warming (from 70 to 40% on average), while that of mosses increased slightly (from 0.3 to 7% on average). The physiological performance of the BSC community, evaluated using chlorophyll fluorescence, increased with warming during the first year of the experiment, but did not respond to rainfall reduction. Our results indicate that ongoing climate change will strongly affect the diversity and composition of BSC communities, as well as their recovery after disturbances. The expected changes in richness and composition under warming could reduce or even reverse the positive effects of BSCs on important soil processes. Thus, these changes are likely to promote an overall reduction in ecosystem processes that sustain and control nutrient cycling, soil stabilization and water dynamics.

Are soil lichen communities structured by biotic interactions? A null model analysis

Abstract Question: Are soil lichen communities structured by biotic interactions? Location: Gypsum outcrops located next to Belmonte del Tajo, central Spain. Methods: We sampled a total of 68 (50 cm × 50 cm) plots in gypsum outcrops from central Spain. Each plot was divided into 100 (5 cm × 5 cm) sampling quadrats, and the presence of all lichen species in every quadrat was recorded (6800 quadrats in total). We used two realistic null models to generate random communities unstructured by biotic interactions, and used them to test the hypothesis that soil lichen species co-occur less often than expected by chance. Results: We found fewer species combinations and less co-occurrence than expected by chance. However, the latter result was dependent on the null model selected. The number of checkerboard pairs did not differ significantly from the null expectation. Conclusions: Overall, our results suggest that gypsiferous soil lichen communities are structured by competitive interactions. They are consistent with studies conducted with a wide variety of taxa, and fill a gap in our knowledge of the factors driving the small-scale distribution of these important organisms. Nomenclature: Tutin et al. (1964–1980) for vascular plants; Hladun & Llimona (2002–2007) and Breuss (1996) for lichens.

Simulated climate change reduced the capacity of lichen-dominated biocrusts to act as carbon sinks in two semi-arid Mediterranean ecosystems

The importance of biological soil crusts (biocrusts) for the biogeochemistry of drylands is widely recognized. However, there are significant gaps in our knowledge about how climate change will affect these organisms and the processes depending on them. We conducted a manipulative full factorial experiment in two representative dryland ecosystems from central (Aranjuez) and southeastern (Sorbas) Spain to evaluate how precipitation, temperature and biocrust cover affected the assimilation and net C balance of biocrusts. Chlorophyll fluorescence, net photosynthesis and dark respiration were measured in situ bimonthly during a year. We also conducted daily cycle measurements of net photosynthesis in winter and at the end of spring. In Sorbas, warming reduced the fixation of atmospheric C in biocrust-dominated microsites throughout the year. In Aranjuez, there was an interaction between the three factors evaluated; during winter, net photosynthesis was significantly greater in high biocrust cover plots under natural conditions and in the rainfall exclusion treatment. During the daily surveys, rainfall exclusion and warming reduced C fixation in Sorbas and in Aranjuez respectively. The effects of the treatments evaluated varied with the rainfall and non-rainfall water inputs (NRWIs) registered before the measurements. Our results suggest that changes in NRWI regimes as consequence of warming could have a greater impact on the C balance of biocrusts than changes in rainfall amounts. They also indicate that climate change may reduce the photosynthetic ability of lichens, with a consequent reduction of their dominance in biocrust communities at the mid to long term. This could reduce the ability of dryland ecosystems to fix atmospheric C.

Direct and indirect impacts of climate change on microbial and biocrust communities alter the resistance of the N cycle in a semiarid grassland

Summary 1. Climate change will raise temperatures and modify precipitation patterns in drylands worldwide, affecting their structure and functioning. Despite the recognized importance of soil communities dominated by mosses, lichens and cyanobacteria (biocrusts) as a driver of nutrient cycling in drylands, little is known on how biocrusts will modulate the resistance (i.e., the amount of change caused by a disturbance) of the N cycle in response to climate change. 2. Here, we evaluate how warming (ambient vs. ~2.5 °C increase), rainfall exclusion (ambient vs. ~30% reduction in total annual rainfall) and biocrust cover (incipient vs. well-developed biocrusts) affect multiple variables linked to soil N availability (inorganic and organic N and potential net N mineralization rate) and its resistance to climate change during 4 years in a field experiment. We also evaluate how climate change-induced modifications in biocrust and microbial communities indirectly affect such resistance. 3. Biocrusts promoted the resistance of soil N availability regardless of the climatic conditions considered. However, the dynamics of N availability diverged progressively from their original conditions with warming and/or rainfall exclusion, as both treatments enhanced N availability and promoted the dominance of inorganic over organic N. In addition, the increase in fungal:bacterial ratio and the decrease in biocrust cover observed under warming had a negative indirect effect on the resistance of N cycle variables. 4. Synthesis. Our results indicate that climate change will have negative direct and indirect (i.e. through changes in biocrust and microbial communities) impacts on the resistance of the N cycle in dryland soils. While biocrusts can play an important role slowing down the impacts of climate change on the N cycle due to their positive and continued effects on the resistance of multiple variables from the N cycle, such change will progressively alter N cycling in biocrust-dominated ecosystems, enhancing both N availability and inorganic N dominance.

Biological soil crusts (biocrusts) as a model system in community, landscape and ecosystem ecology

Model systems have had a profound influence on the development of ecological theory and general principles. Compared to alternatives, the most effective models share some combination of the following characteristics: simpler, smaller, faster, general, idiosyncratic or manipulable. We argue that biological soil crusts (biocrusts) have unique combinations of these features that should be more widely exploited in community, landscape and ecosystem ecology. In community ecology, biocrusts are elucidating the importance of biodiversity and spatial pattern for maintaining ecosystem multifunctionality due to their manipulability in experiments. Due to idiosyncrasies in their modes of facilitation and competition, biocrusts have led to new models on the interplay between environmental stress and biotic interactions and on the maintenance of biodiversity by competitive processes. Biocrusts are perhaps one of the best examples of micro-landscapes—real landscapes that are small in size. Although they exhibit varying patch heterogeneity, aggregation, connectivity and fragmentation, like macro-landscapes, they are also compatible with well-replicated experiments (unlike macro-landscapes). In ecosystem ecology, a number of studies are imposing small-scale, low cost manipulations of global change or state factors in biocrust micro-landscapes. The versatility of biocrusts to inform such disparate lines of inquiry suggests that they are an especially useful model system that can enable researchers to see ecological principles more clearly and quickly.

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.