Rewilding by large ungulates contributes to organic carbon storage in soils.

The concept of rewilding, which focuses on managing ecosystem functions through self-regulation by restoring trophic interactions through introduced animal species with little human intervention, has gained increasing attention as a proactive and efficient approach to restoring ecosystems quickly and on a large scale. However, the science of rewilding has been criticized for being largely theory-based rather than evidence-based, with available data being geographically biased towards the Netherlands and Scandinavian countries, and a lack of objective data on rewilding effects on soil processes and C sequestration. In response to a call for data-driven experimental rewilding projects focused on national contexts, we collected unique data on the effects of large herbivore rewilding on soil properties from eight sites in the Czech Republic. These include sites with a wide range of edaphic characteristics that were grazed by Exmoor ponies, European bison, and back-bred Bos primigenius cattle (singly or in combination) for 2-6 years on areas ranging from ≈30 to ≈250 ha. Despite the relatively short duration of rewilding actions and considerable variability in the response rate of soil properties to grazing, our results indicate improved nutrient availability (evidenced by higher nitrification rate or higher soluble nitrogen concentration) and accelerated ecosystem metabolism (higher soil microbial biomass and dissolved carbon content). On longer-grazed pastures, rewilding contributed to soil carbon sequestration associated with increased water holding capacity and improved soil structure. However, other soil properties (reduced dissolved P concentration or total P content) showed signs of low P availability in the soils of the rewilding sites. Therefore, carcass retention should be considered where possible. Our data, although limited in number and geographic coverage, allow us to conclude that large ungulate rewilding has the potential to enhance soil carbon sequestration and related ecosystem services in rewilding areas. At the same time, we urge similar monitoring as an essential part of other rewilding projects, which will ultimately allow much more robust conclusions about the effects of this management on soils.

Ammonia oxidation and nitrate reduction marker genes are key indicators of nitrogen losses in temperate forest catchments.

Chronic nitrogen inputs can alleviate N limitation and potentially impose N losses in forests, indicated by soil enrichment in 15 N over 14 N. However, the complexity of the nitrogen cycle hinders accurate quantification of N fluxes. Simultaneously, soil ecologists are striving to find meaningful indicators to characterise the "openness" of the nitrogen cycle. We integrate soil δ15 N with constrained ecosystem N losses and the functional gene potential of the soil microbiome in 14 temperate forest catchments. We show that N losses are associated with soil δ15 N and that δ15 N scales with the abundance of soil bacteria. The abundance of the archaeal amoA gene, representing the first step in nitrification (ammonia oxidation to nitrite), followed by the abundance of narG and napA genes, associated with the first step in denitrification (nitrate reduction to nitrite), explains most of the variability in soil δ15 N. These genes are more informative than the denitrification genes nirS and nirK, which are directly linked to N2 O production. Nitrite formation thus appears to be the critical step associated with N losses. Furthermore, we show that the genetic potential for ammonia oxidation and nitrate reduction is representative of forest soil 15 N enrichment and thus indicative of ecosystem N losses.

Undeveloped till soils in scree areas are an overlooked important phosphorus source for waters in alpine catchments.

Scree deposits in alpine catchments contain undeveloped till soils that are "hidden" between and under stones. These scree areas have no vegetation except for sparse lichen patches on stone surfaces, but the soils exhibit biological activity and active cycling of nitrogen (N), phosphorus (P), and organic carbon (C). We compared the chemical and biochemical properties of till soils in the scree areas (scree soils) with developed soils in alpine meadows (meadow soils) of 14 catchments in the alpine zone of the Tatra Mountains. The data showed that scree soils served as an important source of mobile P forms for waters in high elevation catchments. We then conducted a detailed soil survey focused on four selected alpine catchments with scree cover proportions > 30%. This study confirmed that scree soils have significantly higher concentrations of mobile P forms compared to meadow soils, and a high specific microbial activity directed towards the extraction of P with rapid turnover in the microbial biomass. The combination of these properties and the amounts of scree soils in high-elevation areas highlight their importance in overall biogeochemical P cycling in alpine catchments, and the terrestrial P export to receiving waters.

Surviving trees are key elements in the fate of ectomycorrhizal community after severe bark-beetle forest disturbance.

Bark beetle disturbances are a critical event in the life cycle of Norway spruce forests. However, our knowledge of their effects on ectomycorrhizal fungi (EMF), which play a key role in forest productivity and nutrient cycling, is still incomplete. Special attention has been paid to the dynamics and diversity of EMF communities in managed forests, but studies dealing with disturbed natural stands are underrepresented. We conducted a study in an unmanaged natural spruce forest in the Bohemian Forest (Czech Republic), which suffered severe forest dieback caused by bark beetle. Approximately a decade after the disturbance, the character of the forest structure in the study area (∼60 ha, 41 study plots) ranged from sites with open canopy and sparse tree cover to areas with dense spruce regeneration to patches of closed-canopy forest. We found that relative EMF abundance in soils was positively related to surviving tree and regeneration density. The number of surviving trees also positively affected species EMF richness and tended to support preservation of late-successional EMF species. Our results suggest that trees that survive bark beetle disturbance are key for the fate of the EMF community in natural forests.

Soil warming during winter period enhanced soil N and P availability and leaching in alpine grasslands: A transplant study.

Alpine meadows are strongly affected by climate change. Increasing air temperature prolongs the growing season and together with changing precipitation patterns alters soil temperature during winter. To estimate the effect of climate change on soil nutrient cycling, we conducted a field experiment. We transferred undisturbed plant-soil mesocosms from two wind-exposed alpine meadows at ~2100 m a.s.l. to more sheltered plots, situated ~300-400 m lower in the same valleys. The annual mean air temperature was 2°C higher at the lower plots and soils that were normally frozen at the original plots throughout winters were warmed to ~0°C due to the insulation provided by continuous snow cover. After two years of exposure, we analyzed the nutrient content in plants, and changes in soil bacterial community, decomposition, mineralization, and nutrient availability. Leaching of N and P from the soils was continuously measured using ion-exchange resin traps. Warming of soils to ~0°C during the winter allowed the microorganisms to remain active, their metabolic processes were not restricted by soil freezing. This change accelerated nutrient cycling, as evidenced by increased soil N and P availability, their higher levels in plants, and elevated leaching. In addition, root exudation and preferential enzymatic mining of P over C increased. However, any significant changes in microbial biomass, bacterial community composition, decomposition rates, and mineralization during the growing season were not observed, suggesting considerable structural and functional resilience of the microbial community. In summary, our data suggest that changes in soil temperature and snow cover duration during winter periods are critical for altering microbially-mediated processes (even at unchanged soil microbial community and biomass) and may enhance nutrient availability in alpine meadows. Consequently, ongoing climate change, which leads to soil warming and decreasing snow insulation, has a potential to significantly alter nutrient cycling in alpine and subalpine meadows compared to the current situation and increase the year-on-year variability in nutrient availability and leaching.

Decomposition of peatland DOC affected by root exudates is driven by specific r and K strategic bacterial taxa.

In peatlands, decomposition of organic matter is limited by harsh environmental conditions and low decomposability of the plant material. Shifting vegetation composition from Sphagnum towards vascular plants is expected in response to climate change, which will lead to increased root exudate flux to the soil and stimulation of microbial growth and activity. We aimed to evaluate the effect of root exudates on the decomposition of recalcitrant dissolved organic carbon (DOC) and to identify microorganisms involved in this process. The exudation was mimicked by an addition of a mixture of 13C labelled compounds into the recalcitrant DOC in two realistic levels; 2% and 5% of total DOC and peatland porewater with added root exudates was incubated under controlled conditions in the lab. The early stage of incubation was characterized by a relative increase of r-strategic bacteria mainly from Gammaproteobacteria and Bacteriodetes phyla within the microbial community and their preferential use of the added compounds. At the later stage, Alphaproteobacteria and Acidobacteria members were the dominating phyla, which metabolized both the transformed 13C compounds and the recalcitrant DOC. Only higher exudate input (5% of total DOC) stimulated decomposition of recalcitrant DOC compared to non-amended control. The most important taxa with a potential to decompose complex DOC compounds were identified as: Mucilaginibacter (Bacteriodetes), Burkholderia and Pseudomonas (Gammaproteobacteria) among r-strategists and Bryocella and Candidatus Solibacter (Acidobacteria) among K-strategists. We conclude that increased root exudate inputs and their increasing C/N ratio stimulate growth and degradation potential of both r-strategic and K-strategic bacteria, which make the system more dynamic and may accelerate decomposition of peatland recalcitrant DOC.

Bacteria but not fungi respond to soil acidification rapidly and consistently in both a spruce and beech forest.

Anthropogenically enhanced atmospheric sulphur (S) and nitrogen (N) deposition has acidified and eutrophied forest ecosystems worldwide. However, both S and N mechanisms have an impact on microbial communities and the consequences for microbially driven soil functioning differ. We conducted a two-forest stand (Norway spruce and European beech) field experiment involving acidification (sulphuric acid addition) and N (ammonium nitrate) loading and their combination. For 4 years, we monitored separate responses of soil microbial communities to the treatments and investigated the relationship to changes in the activity of extracellular enzymes. We observed that acidification selected for acidotolerant and oligotrophic taxa of Acidobacteria and Actinobacteria decreased bacterial community richness and diversity in both stands in parallel, disregarding their original dissimilarities in soil chemistry and composition of microbial communities. The shifts in bacterial community influenced the stoichiometry and magnitude of enzymatic activity. The bacterial response to experimental N addition was much weaker, likely due to historically enhanced N availability. Fungi were not influenced by any treatment during 4-year manipulation. We suggest that in the onset of acidification when fungi remain irresponsive, bacterial reaction might govern the changes in soil enzymatic activity.

Surplus Carbon Drives Allocation and Plant-Soil Interactions.

Plant growth is usually constrained by the availability of nutrients, water, or temperature, rather than photosynthetic carbon (C) fixation. Under these conditions leaf growth is curtailed more than C fixation, and the surplus photosynthates are exported from the leaf. In plants limited by nitrogen (N) or phosphorus (P), photosynthates are converted into sugars and secondary metabolites. Some surplus C is translocated to roots and released as root exudates or transferred to root-associated microorganisms. Surplus C is also produced under low moisture availability, low temperature, and high atmospheric CO2 concentrations, with similar below-ground effects. Many interactions among above- and below-ground ecosystem components can be parsimoniously explained by the production, distribution, and release of surplus C under conditions that limit plant growth.

Spatial heterogeneity of belowground microbial communities linked to peatland microhabitats with different plant dominants.

Peatland vegetation is composed mostly of mosses, graminoids and ericoid shrubs, and these have a distinct impact on peat biogeochemistry. We studied variation in soil microbial communities related to natural peatland microhabitats dominated by Sphagnum, cotton-grass and blueberry. We hypothesized that such microhabitats will be occupied by structurally and functionally different microbial communities, which will vary further during the vegetation season due to changes in temperature and photosynthetic activity of plant dominants. This was addressed using amplicon-based sequencing of prokaryotic and fungal rDNA and qPCR with respect to methane-cycling communities. Fungal communities were highly microhabitat-specific, while prokaryotic communities were additionally directed by soil pH and total N content. Seasonal alternations in microbial community composition were less important; however, they influenced the abundance of methane-cycling communities. Cotton-grass and blueberry bacterial communities contained relatively more α-Proteobacteria but less Chloroflexi, Fibrobacteres, Firmicutes, NC10, OD1 and Spirochaetes than in Sphagnum. Methanogens, syntrophic and anaerobic bacteria (i.e. Clostridiales, Bacteroidales, Opitutae, Chloroflexi and Syntrophorhabdaceae) were suppressed in blueberry indicating greater aeration that enhanced abundance of fungi (mainly Archaeorhizomycetes) and resulted in the highest fungi-to-bacteria ratio. Thus, microhabitats dominated by different vascular plants are inhabited by unique microbial communities, contributing greatly to spatial functional diversity within peatlands.

A plant-microbe interaction framework explaining nutrient effects on primary production.

In most terrestrial ecosystems, plant growth is limited by nitrogen and phosphorus. Adding either nutrient to soil usually affects primary production, but their effects can be positive or negative. Here we provide a general stoichiometric framework for interpreting these contrasting effects. First, we identify nitrogen and phosphorus limitations on plants and soil microorganisms using their respective nitrogen to phosphorus critical ratios. Second, we use these ratios to show how soil microorganisms mediate the response of primary production to limiting and non-limiting nutrient addition along a wide gradient of soil nutrient availability. Using a meta-analysis of 51 factorial nitrogen-phosphorus fertilization experiments conducted across multiple ecosystems, we demonstrate that the response of primary production to nitrogen and phosphorus additions is accurately predicted by our stoichiometric framework. The only pattern that could not be predicted by our original framework suggests that nitrogen has not only a structural function in growing organisms, but also a key role in promoting plant and microbial nutrient acquisition. We conclude that this stoichiometric framework offers the most parsimonious way to interpret contrasting and, until now, unresolved responses of primary production to nutrient addition in terrestrial ecosystems.