Diazotrophic activity and denitrification in two long-term chronosequences of maritime Antarctica.

The main goals of this study were to identify whether key processes involved in microbial soil nitrogen transformations, such as diazotrophic activity and denitrification, the chemical properties of limiting elements in the soil, and microbial community structure, differ in the different successional stages of two long term chronosequences in maritime Antarctica. Moreover, we expect the rates of diazotrophic activity and denitrification to be stimulated by increases in air temperature and moisture. To answer these questions, we selected three stages in the succession (early, mid and late) in each of two well established chronosequences: three raised beaches in Ardley Island; and the Barton Peninsula, which includes two cosmogenically dated sites and the forefield of the Fourcade glacier. In the Ardley chronosequence, higher diazotrophic activity was found in the older successional stages, concomitant with an increase in the abundance of Cyanobacteria. In the Barton chronosequence, Cyanobacteria were only present and abundant (Microcoleus) in the early successional stage, coinciding with the highest diazotrophic activity. Denitrification in the Barton chronosequence tended to be highest at the mid successional sites, associated with the highest abundance of Rhodanobacter. In the Ardley chronosequence, the lowest abundance of Rhodanobacter was linked to lower denitrification rates in the mid successional stage. In the Ardley chronosequence, significant positive effects of passive warming and water addition on diazotrophic activity were detected in the first and the second years of the study respectively. In the Barton chronosequence on the other hand, there was no response to either passive warming or water addition, probably a manifestation of the higher nutrient limitation in this site. Denitrification showed no response to either warming or water addition. Thus, the response of microbial nitrogen transformations to global change is highly dependent on the environmental setting, such as soil origin, age and climate regime.

The influence of soil age on ecosystem structure and function across biomes.

The importance of soil age as an ecosystem driver across biomes remains largely unresolved. By combining a cross-biome global field survey, including data for 32 soil, plant, and microbial properties in 16 soil chronosequences, with a global meta-analysis, we show that soil age is a significant ecosystem driver, but only accounts for a relatively small proportion of the cross-biome variation in multiple ecosystem properties. Parent material, climate, vegetation and topography predict, collectively, 24 times more variation in ecosystem properties than soil age alone. Soil age is an important local-scale ecosystem driver; however, environmental context, rather than soil age, determines the rates and trajectories of ecosystem development in structure and function across biomes. Our work provides insights into the natural history of terrestrial ecosystems. We propose that, regardless of soil age, changes in the environmental context, such as those associated with global climatic and land-use changes, will have important long-term impacts on the structure and function of terrestrial ecosystems across biomes.

Multiple elements of soil biodiversity drive ecosystem functions across biomes.

The role of soil biodiversity in regulating multiple ecosystem functions is poorly understood, limiting our ability to predict how soil biodiversity loss might affect human wellbeing and ecosystem sustainability. Here, combining a global observational study with an experimental microcosm study, we provide evidence that soil biodiversity (bacteria, fungi, protists and invertebrates) is significantly and positively associated with multiple ecosystem functions. These functions include nutrient cycling, decomposition, plant production, and reduced potential for pathogenicity and belowground biological warfare. Our findings also reveal the context dependency of such relationships and the importance of the connectedness, biodiversity and nature of the globally distributed dominant phylotypes within the soil network in maintaining multiple functions. Moreover, our results suggest that the positive association between plant diversity and multifunctionality across biomes is indirectly driven by soil biodiversity. Together, our results provide insights into the importance of soil biodiversity for maintaining soil functionality locally and across biomes, as well as providing strong support for the inclusion of soil biodiversity in conservation and management programmes.

Climatic vulnerabilities and ecological preferences of soil invertebrates across biomes.

Unlike plants and vertebrates, the ecological preferences, and potential vulnerabilities of soil invertebrates to environmental change, remain poorly understood in terrestrial ecosystems globally. We conducted a cross-biome survey including 83 locations across six continents to advance our understanding of the ecological preferences and vulnerabilities of the diversity of dominant and functionally important soil invertebrate taxa, including nematodes, arachnids and rotifers. The diversity of invertebrates was analyzed through amplicon sequencing. Vegetation and climate drove the diversity and dominant taxa of soil invertebrates. Our results suggest that declines in forest cover and plant diversity, and reductions in plant production associated with increases in aridity, can result in reductions of the diversity of soil invertebrates in a drier and more managed world. We further developed global atlases of the diversity of these important soil invertebrates, which were cross-validated using an independent database. Our study advances the current knowledge of the ecological preferences and vulnerabilities of the diversity and presence of functionally important soil invertebrates in soils from across the globe. This information is fundamental for improving and prioritizing conservation efforts of soil genetic resources and management policies.

Global ecological predictors of the soil priming effect.

Identifying the global drivers of soil priming is essential to understanding C cycling in terrestrial ecosystems. We conducted a survey of soils across 86 globally-distributed locations, spanning a wide range of climates, biotic communities, and soil conditions, and evaluated the apparent soil priming effect using 13C-glucose labeling. Here we show that the magnitude of the positive apparent priming effect (increase in CO2 release through accelerated microbial biomass turnover) was negatively associated with SOC content and microbial respiration rates. Our statistical modeling suggests that apparent priming effects tend to be negative in more mesic sites associated with higher SOC contents. In contrast, a single-input of labile C causes positive apparent priming effects in more arid locations with low SOC contents. Our results provide solid evidence that SOC content plays a critical role in regulating apparent priming effects, with important implications for the improvement of C cycling models under global change scenarios.

Changes in belowground biodiversity during ecosystem development.

Belowground organisms play critical roles in maintaining multiple ecosystem processes, including plant productivity, decomposition, and nutrient cycling. Despite their importance, however, we have a limited understanding of how and why belowground biodiversity (bacteria, fungi, protists, and invertebrates) may change as soils develop over centuries to millennia (pedogenesis). Moreover, it is unclear whether belowground biodiversity changes during pedogenesis are similar to the patterns observed for aboveground plant diversity. Here we evaluated the roles of resource availability, nutrient stoichiometry, and soil abiotic factors in driving belowground biodiversity across 16 soil chronosequences (from centuries to millennia) spanning a wide range of globally distributed ecosystem types. Changes in belowground biodiversity during pedogenesis followed two main patterns. In lower-productivity ecosystems (i.e., drier and colder), increases in belowground biodiversity tracked increases in plant cover. In more productive ecosystems (i.e., wetter and warmer), increased acidification during pedogenesis was associated with declines in belowground biodiversity. Changes in the diversity of bacteria, fungi, protists, and invertebrates with pedogenesis were strongly and positively correlated worldwide, highlighting that belowground biodiversity shares similar ecological drivers as soils and ecosystems develop. In general, temporal changes in aboveground plant diversity and belowground biodiversity were not correlated, challenging the common perception that belowground biodiversity should follow similar patterns to those of plant diversity during ecosystem development. Taken together, our findings provide evidence that ecological patterns in belowground biodiversity are predictable across major globally distributed ecosystem types and suggest that shifts in plant cover and soil acidification during ecosystem development are associated with changes in belowground biodiversity over centuries to millennia.

Coupling of microbial nitrogen transformations and climate in sclerophyll forest soils from the Mediterranean Region of central Chile.

The Mediterranean region of central Chile is experiencing extensive "mega-droughts" with detrimental effects for the environment and economy of the region. In the northern hemisphere, nitrogen (N) limitation of Mediterranean ecosystems has been explained by the decoupling between N inputs and plant uptake during the dormant season. In central Chile, soils have often been considered N-rich in comparison to other Mediterranean ecosystems of the world, yet the impacts of expected intensification of seasonal drought remain unknown. In this work, we seek to disentangle patterns of microbial N transformations and their seasonal coupling with climate in the Chilean sclerophyll forest-type. We aim to assess how water limitation affects microbial N transformations, thus addressing the impact of ongoing regional climate trends on soil N status. We studied four stands of the sclerophyll forest-type in Chile. Field measurements in surface soils showed a 67% decline of free-living diazotrophic activity (DA) and 59% decrease of net N mineralization rates during the summer rainless and dormant season, accompanied by a stimulation of in-situ denitrification rates to values 70% higher than in wetter winter. Higher rates of both free-living DA and net N mineralization found during spring, provided evidence for strong coupling of these two processes during the growing season. Overall, the experimental addition of water in the field to litter samples almost doubled DA but had no effect on denitrification rates. We conclude that coupling of microbial mediated soil N transformations during the wetter growing season explains the N enrichment of sclerophyll forest soils. Expected increases in the length and intensity of the dry period, according to climate change models, reflected in the current mega-droughts may drastically reduce biological N fixation and net N mineralization, increasing at the same time denitrification rates, thereby potentially reducing long-term soil N capital.

Successional changes in soil nitrogen availability, non-symbiotic nitrogen fixation and carbon/nitrogen ratios in southern Chilean forest ecosystems.

Vast areas of southern Chile are now covered by second-growth forests because of fire and logging. To study successional patterns after moderate-intensity, anthropogenic fire disturbance, we assessed differences in soil properties and N fluxes across a chronosequence of seven successional stands (2-130 years old). We examined current predictions of successional theory concerning changes in the N cycle in forest ecosystems. Seasonal fluctuations of net N mineralization (N(min)) in surface soil and N availability (N(a); N(a)=NH4+-N+NO3--N) in upper and deep soil horizons were positively correlated with monthly precipitation. In accordance with theoretical predictions, stand age was positively, but weakly related to both N(a) ( r(2)=0.282, P<0.001) and total N (N(tot); r(2)=0.192, P<0.01), and negatively related to soil C/N ratios ( r(2)=0.187, P<0.01) in surface soils. A weak linear increase in soil N(min) (upper plus deep soil horizons) was found across the chronosequence ( r(2)=0.124, P<0.022). N(min) occurred at modest rates in early successional stands, suggesting that soil disturbance did not impair microbial processes. The relationship between N fixation (N(fix)) in the litter layer and stand age best fitted a quadratic model ( r(2)=0.228, P<0.01). In contrast to documented successional trends for most temperate, tropical and Mediterranean forests, non-symbiotic N(fix) in the litter layer is a steady N input to unpolluted southern temperate forests during mid and late succession, which may compensate for hydrological losses of organic N from old-growth ecosystems.