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Winter catch crops are grown to scavenge nutrients over a period of unfavorable growth conditions and to conserve nutrients for subsequent release to the following main crop. Since environmental conditions have a strong impact on the growth and nutrient capture in roots and shoots of individual catch crop species, we anticipated that mixtures will be more durable and efficient in nutrient capture due to compensatory effects among component species. We tested this hypothesis and determined the nitrogen and phosphorus accumulation in the shoots and roots of four catch crop species grown in pure vs. mixed stands at two sites for two or three years. Element concentrations were determined in the root and shoot biomass of each species and used to calculate the nutrient pool fixed in the root or shoot biomass. A qPCR-based technique was applied to quantify the root biomass of individual species based on species-specific DNA sequences. Despite considerable variation across environments, the overall plant biomass of white mustard (Sinapis alba), lacy phacelia (Phacelia tanacetifolia) and bristle oat (Avena strigosa) was similar and higher than that of Egyptian clover (Trifolium alexandrinum). While pure stands varied 6- to 24-fold in shoot biomass depending on environmental conditions, the variation was only ~3-fold for catch crop mixtures, with less pronounced variation in the root biomass. In general, the root biomass was comparable to the shoot biomass in each species. Roots contributed 26–46% of the nitrogen and 36–48% of the phosphorus to the total accumulation of these nutrients in the catch crop biomass, thus emphasizing the importance of plant roots as belowground nutrient pool for potential carry-over of nutrients to the subsequent crop. Although the mixture was mostly dominated by two of the four species, namely mustard and phacelia, it captured similar or even larger amounts of nutrients than the best-performing pure stand under any growth condition. This was the case for shoot- and for root-bound nutrients. Our results indicate that catch crop mixtures have higher durability than pure cultures to environmental variations. The amount of nitrogen captured by the mixture meets the average postharvest nitrogen that is left over by a wide range of cash crops, thus emphasizing that catch crop mixtures represent an efficient nutrient management tool in crop rotations. © 2022

Permafrost-affected soils in the northern circumpolar region store more than 1,000 Pg soil organic carbon (OC), and are strongly vulnerable to climatic warming. However, the extent to which changing soil environmental conditions with permafrost thaw affects different compounds of soil organic matter (OM) is poorly understood. Here, we assessed the fate of lignin and non-cellulosic carbohydrates in density fractionated soils (light fraction, LF vs. heavy fraction, HF) from three permafrost regions with decreasing continentality, expanding from east to west of northern Siberia (Cherskiy, Logata, Tazovskiy, respectively). In soils at the Tazovskiy site with thicker active layers, the LF showed smaller OC-normalized contents of lignin-derived phenols and plant-derived sugars and a decrease of these compounds with soil depth, while a constant or even increasing trend was observed in soils with shallower active layers (Cherskiy and Logata). Also in the HF, soils at the Tazovskiy site had smaller contents of OC-normalized lignin-derived phenols and plant-derived sugars along with more pronounced indicators of oxidative lignin decomposition and production of microbial-derived sugars. Active layer deepening, thus, likely favors the decomposition of lignin and plant-derived sugars, that is, lignocelluloses, by increasing water drainage and aeration. Our study suggests that climate-induced degradation of permafrost soils may promote carbon losses from lignin and associated polysaccharides by abolishing context-specific preservation mechanisms. However, relations of OC-based lignin-derived phenols and sugars in the HF with mineralogical properties suggest that future OM transformation and carbon losses will be modulated in addition by reactive soil minerals. publishedVersion

Improving N cycling in agroecosystems is one of the key challenges in reducing the environmental footprint of agriculture. Further, uncertainty in precipitation makes crop water management relevant in regions where it has not been necessary thus far. Here, we focus on the potential of winter-killed catch crops (CCs) to reduce N leaching losses from N mineralization over the winter and from soil water management. We compared four single CCs (white mustard, phacelia, Egyptian clover and bristle oat) and two CC mixtures with 4 and 12 plant species (Mix4 and Mix12) with a fallow treatment. High-resolution soil mineral N (Nmin) monitoring in combination with the modelling of spatiotemporal dynamics served to assess N cycling under winter-killed CCs, while soil water was continuously monitored in the rooting zone. Catch crops depleted the residual Nmin pools by between 40 % and 72 % compared to the fallow. The amount of residual N uptake was lowest for clover and not significantly different among the other CCs. Catch crops that produce high N litter materials, such as clover and mustard leaves, showed an early N mineralization flush immediately after their termination and the highest leaching losses from litter mineralization over the winter. Except for clover, all CCs showed Nmin values between 18 % and 92 % higher on the sowing date of the following maize crop. However, only Mix12 was statistically significant. Catch crops depleted the soil water storage in the rooting zone during their growth in autumn and early winter, but preserved water later on when their residues covered the ground. The shallow incorporation of CC residues increased water storage capacity during the cropping season of the main crop even under reduced soil water availability. Hence, catch cropping is not just a simple plant cover for the winter but improves the growth conditions for the following crop with decreased N losses. Mixtures have been shown to compensate for the weaknesses of individual CC species in terms of nutrient capture, mineralization and transfer to the following main crop as well as for soil water management. Detailed knowledge about plant performance during growth and litter mineralization patterns is necessary to make optimal use of their potential.

Improving N cycling in agroecosystems is one of the key challenges in reducing the environmental footprint of agriculture. Further, uncertainty in precipitation makes crop water management relevant in regions where it has not been necessary thus far. Here, we focus on the potential of winter-killed catch crops to reduce N leaching losses from N mineralization over the winter and soil water management. We compared four single catch crops (white mustard, phacelia, Egyptian clover and bristle oat) and a fallow treatment with two catch crop mixtures with 4 and 12 plant species (Mix4 and Mix12). High-resolution soil mineral N (Nmin) monitoring in combination with modelling of spatiotemporal dynamics served to assess N cycling under winter-killed catch crops, while soil water was continuously monitored in the rooting zone. Catch crops depleted the residual Nmin pools by between 40 and 72 % compared to the fallow. The amount of residual N uptake was lowest for clover and not significantly different among the other catch crops. Catch crops that produce high N litter materials, such as clover and mustard leaves, showed an early N mineralization flush immediately after their termination and the highest leaching losses from litter mineralization over the winter. Except for clover, all catch crops showed Nmin values between 18 and 92 % higher on the sowing date of the following maize crop. However, only Mix12 was statistically significant. Catch crops depleted the soil water storage in the rooting zone during their growth in autumn and early winter, but preserved water later on when their residues cover the ground. The shallow incorporation of catch crop residues increased water storage capacity during the cropping season of the main crop even under drought conditions. Hence, catch cropping is not just a simple plant cover during the winter but improved the growth conditions for the following crop at decreased N losses. Mixtures have been shown to compensate for the weaknesses of individual catch crop species in terms of nutrient capture, mineralization and transfer to the following main crop as well as for soil water management. Detailed knowledge about plant performance during growth and litter mineralization patterns is necessary to make optimal use of their full potential.

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Agricultural management is a key force affecting soil processes and functions. Triggered by biophysical constraints as well as rapid structural and technological developments, new management practices are emerging with largely unknown impacts on soil processes and functions. This impedes assessments of the potential of such emerging practices for sustainable intensification, a paradigm coined to address the growing demand for food and nonfood products. In terms of soil management, sustainable intensification means that soil productivity is increased while other soil functions and services, such as carbon storage and habitat for organisms, are simultaneously maintained or even improved. In this paper we provide an overview of research challenges to better understand how emerging soil management practices affect soil processes and functions. We distinguish four categories of soil management practices: spatial arrangements of cropping systems, crops and rotations, mechanical pressures, and inputs into the soil. Key research needs identified for each include nutrient efficiency in agroforestry versus conventional cropping systems, soil-rhizosphere microbiome elucidation to understand the interacting roles of crops and rotations, the effects of soil compaction on soil–plant–atmosphere interactions, and the ecotoxicity of plastics, pharmaceuticals and other pollutants that are introduced into the soil. We establish an interdisciplinary, systemic approach to soil science and include cross-cutting research activities related to process modeling, data management, stakeholder interaction, sustainability assessment and governance. The identification of soil research challenges from the perspective of agricultural management facilitates cooperation between different scientific disciplines in the field of sustainable agricultural production.

Catch crops increase plant species richness in crop rotations, but are most often grown as pure stands. Here, we investigate the impacts of increasing plant diversity in catch crop rotations on rhizosphere C input and microbial utilization. Mustard (Sinapis alba L.) planted as a single cultivar was compared to diversified catch crop mixtures of four (Mix4) or 12 species (Mix12). We traced the C transfer from shoots to roots towards the soil microbial community and the soil respiration in a 13C pulse labelling field experiment. Net CO2-C uptake from the atmosphere increased by two times in mix 4 and more than three times in mix 12. Higher net ecosystem C production was linked to increasing catch crop diversity and increased belowground transfer rates of recently fixed photoassimilates. The higher rhizosphere C input stimulated the growth and activity of the soil microbiome, which was investigated by phospholipid fatty acid (PLFA) analyses. Total microbial biomass increased from 14 to 22 g m−2 as compared to the fallow and was 18 and 8% higher for mix 12 and mix 4 as compared to mustard. In particular, the fungal and actinobacterial communities profited the most from the higher belowground C input and their biomass increased by 3.4 and 1.3 times as compared to the fallow. The residence time of the 13C pulse, traced in the CO2 flux from the soil environment, increased with plant diversity by up to 1.8 times. The results of this study suggest positive impacts of plant diversity on C cycling by higher atmospheric C uptake, higher transport rates towards the rhizosphere, higher microbial incorporation and prolonged residence time in the soil environment. We conclude that diversified catch crop mixtures improve the efficiency of C cycling in cropping systems and provide a promising tool for sustainable soil management.

© 2018 Elsevier Ltd A substantial portion of soil organic matter (SOM) is of microbial origin. The efficiency with which soil microorganisms can convert their substrate carbon (C) into biomass, compared to how much is lost as respiration, thus co-determines the carbon storage potential of soils. Despite increasing insight into soil microbial C cycling, empirical measurements of microbial C processing across biomes and across soil horizons remain sparse. The theory of ecological stoichiometry predicts that microbial carbon use efficiency (CUE), i.e. growth over uptake of organic C, strongly depends on the relative availability of C and nutrients, particularly N, as microorganisms will either respire excess C or conserve C while mineralising excess nutrients. Microbial CUE is thus expected to increase from high to low latitudes and from topsoil to subsoil as the soil C:N and the stoichiometric imbalance between SOM and the microbial biomass decrease. To test these hypotheses, we collected soil samples from the organic topsoil, mineral topsoil, and mineral subsoil of seven sites along a 1500-km latitudinal transect in Western Siberia. As a proxy for CUE, we measured the microbial substrate use efficiency (SUE) of added substrates by incubating soil samples with a mixture of13C labelled sugars, amino sugars, amino acids, and organic acids and tracing13C into microbial biomass and released CO2. In addition to soil and microbial C:N stoichiometry, we also determined the potential extracellular enzyme activities of cellobiohydrolase (CBH) and phenol oxidase (POX) and used the CBH:POX ratio as an indicator of SOM substrate quality. We found an overall decrease of SUE with latitude, corresponding to a decrease in mean annual temperature, in mineral soil horizons. SUE decreased with decreasing stoichiometric imbalance in the organic and mineral topsoil, while a relationship of SUE with soil C:N was only found in the mineral topsoil. However, contrary to our hypothesis, SUE did not increase with soil depth and mineral subsoils displayed lower average SUE than mineral topsoils. Both within individual horizons and across all horizons SUE was strongly correlated with CBH:POX ratio as well as with climate variables. Since enzyme activities likely reflect the chemical properties of SOM, our results indicate that SOM quality exerts a stronger control on SUE than SOM stoichiometry, particularly in subsoils were SOM has been turned over repeatedly and there is little variation in SOM elemental ratios.

The occurrence of dark fixation of CO2 by heterotrophic microorganisms in soil is generally accepted, but its importance for microbial metabolism and soil organic carbon (C) sequestration is unknown, especially under C-limiting conditions. To fill this knowledge gap, we measured dark 13CO2 incorporation into soil organic matter and conducted a 13C-labelling experiment to follow the 13C incorporation into phospholipid fatty acids as microbial biomass markers across soil profiles of four tundra ecosystems in the northern circumpolar region, where net primary productivity and thus soil C inputs are low. We further determined the abundance of various carboxylase genes and identified their microbial origin with metagenomics. The microbial capacity for heterotrophic CO2 fixation was determined by measuring the abundance of carboxylase genes and the incorporation of 13C into soil C following the augmentation of bioavailable C sources. We demonstrate that dark CO2 fixation occurred ubiquitously in arctic tundra soils, with increasing importance in deeper soil horizons, presumably due to increasing C limitation with soil depth. Dark CO2 fixation accounted on average for 0.4, 1.0, 1.1, and 16% of net respiration in the organic, cryoturbated organic, mineral and permafrost horizons, respectively. Genes encoding anaplerotic enzymes of heterotrophic microorganisms comprised the majority of identified carboxylase genes. The genetic potential for dark CO2 fixation was spread over a broad taxonomic range. The results suggest important regulatory function of CO2 fixation in C limited conditions. The measurements were corroborated by modeling the long-term impact of dark CO2 fixation on soil organic matter. Our results suggest that increasing relative CO2 fixation rates in deeper soil horizons play an important role for soil internal C cycling and can, at least in part, explain the isotopic enrichment with soil depth.

The potential of a plant species to acquire nutrients depends on its ability to explore the soil by its root system. Co-cultivation of different species is anticipated to lead to vertical root niche differentiation and thus to higher soil nutrient depletion. Using a qPCR-based method we quantified root biomass distribution of four catch crop species in vertical soil profiles in pure vs. mixed stands. Pure stands of mustard and phacelia robustly reached 70 cm soil depth, while oat preferably colonized upper soil layers, and clover developed the shallowest and smallest root system. Analysis of residual nitrate pools in different soil depths and correlation with root biomass showed that, besides rooting depth also root biomass determines soil nitrogen depletion. While occupying the same vertical niches as in pure stands, mustard and phacelia dominated total root biomass of the mix. In contrast, root biomass of clover and oat was severely suppressed in presence of the other species. Below-ground biomass profiling indicated low niche complementarity among the root systems of the examined species. Nonetheless, the mixture mostly overyielded root biomass of the pure stands and thus shows higher potential for efficient soil exploration by roots.

Formation of mineral-organic associations is a key process in the global carbon cycle. Recent concepts propose litter quality-controlled microbial assimilation and direct sorption processes as main factors in transferring carbon from plant litter into mineral-organic associations. We explored the pathways of the formation of mineral-associated organic matter (MOM) in soil profiles along a 120-ky ecosystem gradient that developed under humid climate from the retreating Franz Josef Glacier in New Zealand. We determined the stocks of particulate and mineral-associated carbon, the isotope signature and microbial decomposability of organic matter, and plant and microbial biomarkers (lignin phenols, amino sugars and acids) in MOM. Results revealed that litter quality had little effect on the accumulation of mineral-associated carbon and that plant-derived carbon bypassed microbial assimilation at all soil depths. Seemingly, MOM forms by sorption of microbial as well as plant-derived compounds to minerals. The MOM in carbon-saturated topsoil was characterized by the steady exchange of older for recent carbon, while subsoil MOM arises from retention of organic matter transported with percolating water. Overall, MOM formation is not monocausal but involves various mechanisms and processes, with reactive minerals being effective filters capable of erasing chemical differences in organic matter inputs.

Arctic permafrost soils contain large stocks of organic carbon (OC). Extensive cryogenic processes in these soils cause subduction of a significant part of OC-rich topsoil down into mineral soil through the process of cryoturbation. Currently, one-fourth of total permafrost OC is stored in subducted organic horizons. Predicted climate change is believed to reduce the amount of OC in permafrost soils as rising temperatures will increase decomposition of OC by soil microorganisms. To estimate the sensitivity of OC decomposition to soil temperature and oxygen levels we performed a 4-month incubation experiment in which we manipulated temperature (4–20 °C) and oxygen level of topsoil organic, subducted organic and mineral soil horizons. Carbon loss (CLOSS) was monitored and its potential biotic and abiotic drivers, including concentrations of available nutrients, microbial activity, biomass and stoichiometry, and extracellular oxidative and hydrolytic enzyme pools, were measured. We found that independently of the incubation temperature, CLOSS from subducted organic and mineral soil horizons was one to two orders of magnitude lower than in the organic topsoil horizon, both under aerobic and anaerobic conditions. This corresponds to the microbial biomass being lower by one to two orders of magnitude. We argue that enzymatic degradation of autochthonous subducted OC does not provide sufficient amounts of carbon and nutrients to sustain greater microbial biomass. The resident microbial biomass relies on allochthonous fluxes of nutrients, enzymes and carbon from the OC-rich topsoil. This results in a “negative priming effect”, which protects autochthonous subducted OC from decomposition at present. The vulnerability of subducted organic carbon in cryoturbated arctic soils under future climate conditions will largely depend on the amount of allochthonous carbon and nutrient fluxes from the topsoil.

In permafrost soils, the temperature regime and the resulting cryogenic processes are important determinants of the storage of organic carbon (OC) and its small-scale spatial variability. For cryoturbated soils, there is a lack of research assessing pedon-scale heterogeneity in OC stocks and the transformation of functionally different organic matter (OM) fractions, such as particulate and mineral-associated OM. Therefore, pedons of 28 Turbels were sampled in 5 m wide soil trenches across the Siberian Arctic to calculate OC and total nitrogen (TN) stocks based on digital profile mapping. Density fractionation of soil samples was performed to distinguish between particulate OM (light fraction, LF, < 1.6 g cm−3), mineral associated OM (heavy fraction, HF, > 1.6 g cm−3), and a mobilizable dissolved pool (mobilizable fraction, MoF). Across all investigated soil profiles, the total OC storage was 20.2 ± 8.0 kg m−2 (mean ± SD) to 100 cm soil depth. Fifty-four percent of this OC was located in the horizons of the active layer (annual summer thawing layer), showing evidence of cryoturbation, and another 35 % was present in the upper permafrost. The HF-OC dominated the overall OC stocks (55 %), followed by LF-OC (19 % in mineral and 13 % in organic horizons). During fractionation, approximately 13 % of the OC was released as MoF, which likely represents a readily bioavailable OM pool. Cryogenic activity in combination with cold and wet conditions was the principle mechanism through which large OC stocks were sequestered in the subsoil (16.4 ± 8.1 kg m−2; all mineral B, C, and permafrost horizons). Approximately 22 % of the subsoil OC stock can be attributed to LF material subducted by cryoturbation, whereas migration of soluble OM along freezing gradients appeared to be the principle source of the dominant HF (63 %) in the subsoil. Despite the unfavourable abiotic conditions, low C / N ratios and high δ13C values indicated substantial microbial OM transformation in the subsoil, but this was not reflected in altered LF and HF pool sizes. Partial least-squares regression analyses suggest that OC accumulates in the HF fraction due to co-precipitation with multivalent cations (Al, Fe) and association with poorly crystalline iron oxides and clay minerals. Our data show that, across all permafrost pedons, the mineral-associated OM represents the dominant OM fraction, suggesting that the HF-OC is the OM pool in permafrost soils on which changing soil conditions will have the largest impact. Russian Ministry of Education and Science/14.B25.31.0031 German Federal Ministry of Education and Research/03F0616A Evangelisches Studienwerk Villigst DFG

Cryoturbation, the burial of topsoil material into deeper soil horizons by repeated freeze–thaw events, is an important storage mechanism for soil organic matter (SOM) in permafrost-affected soils. Besides abiotic conditions, microbial community structure and the accessibility of SOM to the decomposer community are hypothesized to control SOM decomposition and thus have a crucial role in SOM accumulation in buried soils. We surveyed the microbial community structure in cryoturbated soils from nine soil profiles in the northeastern Siberian tundra using high-throughput sequencing and quantification of bacterial, archaeal and fungal marker genes. We found that bacterial abundances in buried topsoils were as high as in unburied topsoils. In contrast, fungal abundances decreased with depth and were significantly lower in buried than in unburied topsoils resulting in remarkably low fungal to bacterial ratios in buried topsoils. Fungal community profiling revealed an associated decrease in presumably ectomycorrhizal (ECM) fungi. The abiotic conditions (low to subzero temperatures, anoxia) and the reduced abundance of fungi likely provide a niche for bacterial, facultative anaerobic decomposers of SOM such as members of the Actinobacteria, which were found in significantly higher relative abundances in buried than in unburied topsoils. Our study expands the knowledge on the microbial community structure in soils of Northern latitude permafrost regions, and attributes the delayed decomposition of SOM in buried soils to specific microbial taxa, and particularly to a decrease in abundance and activity of ECM fungi, and to the extent to which bacterial decomposers are able to act as their functional substitutes.

Arctic plant productivity is often limited by low soil N availability. This has been attributed to slow breakdown of N-containing polymers in litter and soil organic matter (SOM) into smaller, available units, and to shallow plant rooting constrained by permafrost and high soil moisture. Using N-15 pool dilution assays, we here quantified gross amino acid and ammonium production rates in 97 active layer samples from four sites across the Siberian Arctic. We found that amino acid production in organic layers alone exceeded literature-based estimates of maximum plant N uptake 17-fold and therefore reject the hypothesis that arctic plant N limitation results from slow SOM breakdown. High microbial N use efficiency in organic layers rather suggests strong competition of microorganisms and plants in the dominant rooting zone. Deeper horizons showed lower amino acid production rates per volume, but also lower microbial N use efficiency. Permafrost thaw together with soil drainage might facilitate deeper plant rooting and uptake of previously inaccessible subsoil N, and thereby promote plant productivity in arctic ecosystems. We conclude that changes in microbial decomposer activity, microbial N utilization and plant root density with soil depth interactively control N availability for plants in the Arctic.

Soil horizons below 30 cm depth contain about 60% of the organic carbon stored in soils. Although insight into the physical and chemical stabilization of soil organic matter (SOM) and into microbial community composition in these horizons is being gained, information on microbial functions of subsoil microbial communities and on associated microbially-mediated processes remains sparse. To identify possible controls on enzyme patterns, we correlated enzyme patterns with biotic and abiotic soil parameters, as well as with microbial community composition, estimated using phospholipid fatty acid profiles. Enzyme patterns (i.e. distance-matrixes calculated from these enzyme activities) were calculated from the activities of six extracellular enzymes (cellobiohydrolase, leucine-amino-peptidase, N-acetylglucosaminidase, chitotriosidase, phosphatase and phenoloxidase), which had been measured in soil samples from organic topsoil horizons, mineral topsoil horizons, and mineral subsoil horizons from seven ecosystems along a 1500 km latitudinal transect in Western Siberia. We found that hydrolytic enzyme activities decreased rapidly with depth, whereas oxidative enzyme activities in mineral horizons were as high as, or higher than in organic topsoil horizons. Enzyme patterns varied more strongly between ecosystems in mineral subsoil horizons than in organic topsoils. The enzyme patterns in topsoil horizons were correlated with SOM content (i.e., C and N content) and microbial community composition. In contrast, the enzyme patterns in mineral subsoil horizons were related to water content, soil pH and microbial community composition. The lack of correlation between enzyme patterns and SOM quantity in the mineral subsoils suggests that SOM chemistry, spatial separation or physical stabilization of SOM rather than SOM content might determine substrate availability for enzymatic breakdown. The correlation of microbial community composition and enzyme patterns in all horizons, suggests that microbial community composition shapes enzyme patterns and might act as a modifier for the usual dependency of decomposition rates on SOM content or C/N ratios., Highlights • Enzyme patterns differed more strongly between horizons than between ecosystems. • Enzyme patterns in subsoils vary more strongly than enzyme patterns in topsoils. • Enzyme patterns in topsoil are related to SOM content and mic. community composition. • Enzyme patterns in mineral subsoil were not related to SOM content or SOM C/N.

openaire: EC/H2020/819202/EU//SOS.aquaterra As global temperatures continue to rise, a key uncertainty of climate projections is the microbial decomposition of vast organic carbon stocks in thawing permafrost soils. Decomposition rates can accelerate up to fourfold in the presence of plant roots, and this mechanism—termed the rhizosphere priming effect—may be especially relevant to thawing permafrost soils as rising temperatures also stimulate plant productivity in the Arctic. However, priming is currently not explicitly included in any model projections of future carbon losses from the permafrost area. Here, we combine high-resolution spatial and depth-resolved datasets of key plant and permafrost properties with empirical relationships of priming effects from living plants on microbial respiration. We show that rhizosphere priming amplifies overall soil respiration in permafrost-affected ecosystems by ~12%, which translates to a priming-induced absolute loss of ~40 Pg soil carbon from the northern permafrost area by 2100. Our findings highlight the need to include fine-scale ecological interactions in order to accurately predict large-scale greenhouse gas emissions, and suggest even tighter restrictions on the estimated 200 Pg anthropogenic carbon emission budget to keep global warming below 1.5 °C.