Global decomposition experiment shows soil animal impacts on decomposition are climate‐dependent

Climate and litter quality are primary drivers of terrestrial decomposition and, based on evidence from multisite experiments at regional and global scales, are universally factored into global decomposition models. In contrast, soil animals are considered key regulators of decomposition at local scales but their role at larger scales is unresolved. Soil animals are consequently excluded from global models of organic mineralization processes. Incomplete assessment of the roles of soil animals stems from the difficulties of manipulating invertebrate animals experimentally across large geographic gradients. This is compounded by deficient or inconsistent taxonomy. We report a global decomposition experiment to assess the importance of soil animals in C mineralization, in which a common grass litter substrate was exposed to natural decomposition in either control or reduced animal treatments across 30 sites distributed from 43°S to 68°N on six continents. Animals in the mesofaunal size range were recovered from the litter by Tullgren extraction and identified to common specifications, mostly at the ordinal level. The design of the trials enabled faunal contribution to be evaluated against abiotic parameters between sites. Soil animals increase decomposition rates in temperate and wet tropical climates, but have neutral effects where temperature or moisture constrain biological activity. Our findings highlight that faunal influences on decomposition are dependent on prevailing climatic conditions. We conclude that (1) inclusion of soil animals will improve the predictive capabilities of region‐ or biome‐scale decomposition models, (2) soil animal influences on decomposition are important at the regional scale when attempting to predict global change scenarios, and (3) the statistical relationship between decomposition rates and climate, at the global scale, is robust against changes in soil faunal abundance and diversity.     

Magnitude and variability of process rates in fungal diversity-litter decomposition relationships

There is compelling evidence that losses in plant diversity can alter ecosystem functioning, particularly by reducing primary production. However, impacts of biodiversity loss on decomposition, the complementary process in the carbon cycle, are highly uncertain. By manipulating fungal decomposer diversity in stream microcosm experiments we found that rates of litter decomposition and associated fungal spore production are unaffected by changes in decomposer diversity under benign and harsher environmental conditions. This result calls for caution when generalizing outcomes of biodiversity experiments across systems. In contrast to their magnitude, the variability of process rates among communities increased when species numbers were reduced. This was most likely caused by a portfolio effect (i.e. statistical averaging), with the uneven species distribution typical of natural communities tending to weaken that effect. Curbing species extinctions to maintain ecosystem functioning thus can be important even in situations where process rates are unaffected.     

Effects of historical legacies on soil nematode communities are mediated by contemporary environmental conditions

Both contemporary and historical factors are documented to be crucial in regulating species diversity and distribution. Soil fauna contribute substantially to global biodiversity and ecosystem functioning, while it is unclear whether and to what extent historical factors shape their diversity patterns. Here, we used soil nematodes as a model organism to test historical effects on soil fauna and to investigate the relative importance of climatic, soil, and historical factors. Based on nematode distribution data in 16 natural sites at a large scale (ranging from 22 to 40°N) in mainland China, we conducted elastic net regression model to test the effects of climatic (e.g., mean and seasonality of temperature/precipitation), soil (e.g., soil carbon, nitrogen, and pH), and historical (e.g., temperature/precipitation anomaly and the velocity of the change since the Last Glacial Maximum) variables on nematode genus richness and Shannon's diversity. Additionally, variation partitioning was used to determine the contribution of the three predictor sets to the explanation of both Jaccard and Bray–Curtis community dissimilarity. We found that climate generally explained more variations in both diversity and composition than soil and historical predictors in our samples. We also showed that although historical factors (e.g., temperature change velocity) were correlated with nematode diversity and composition, the pure effects of these historical factors were negligible. In other words, the historical effects were commonly represented by their interactions with current climatic and soil factors within our selected sites. Our results indicated that contemporary factors, especially climate, may outperform historical factors in regulating soil nematode diversity patterns at large scales.     

Plant diversity does not buffer drought effects on early‐stage litter mass loss rates and microbial properties

Human activities are decreasing biodiversity and changing the climate worldwide. Both global change drivers have been shown to affect ecosystem functioning, but they may also act in concert in a non‐additive way. We studied early‐stage litter mass loss rates and soil microbial properties (basal respiration and microbial biomass) during the summer season in response to plant species richness and summer drought in a large grassland biodiversity experiment, the Jena Experiment, Germany. In line with our expectations, decreasing plant diversity and summer drought decreased litter mass loss rates and soil microbial properties. In contrast to our hypotheses, however, this was only true for mass loss of standard litter (wheat straw) used in all plots, and not for plant community‐specific litter mass loss. We found no interactive effects between global change drivers, that is, drought reduced litter mass loss rates and soil microbial properties irrespective of plant diversity. High mass loss rates of plant community‐specific litter and low responsiveness to drought relative to the standard litter indicate that soil microbial communities were adapted to decomposing community‐specific plant litter material including lower susceptibility to dry conditions during summer months. Moreover, higher microbial enzymatic diversity at high plant diversity may have caused elevated mass loss of standard litter. Our results indicate that plant diversity loss and summer drought independently impede soil processes. However, soil decomposer communities may be highly adapted to decomposing plant community‐specific litter material, even in situations of environmental stress. Results of standard litter mass loss moreover suggest that decomposer communities under diverse plant communities are able to cope with a greater variety of plant inputs possibly making them less responsive to biotic changes.     

Global climate change and litter decomposition: more frequent fire slows decomposition and increases the functional importance of invertebrates

Of the many mechanisms by which global climate change may alter ecosystem processes perhaps the least known and insidious is altered disturbance regimes. We used a field-based experiment to examine the climate change scenario of more frequent fires with altered invertebrate assemblages on the decomposition of Eucalyptus leaves. Our design comprised three fire regimes [long-term fire exclusion (FE), long-term frequent burning (FB) and FE altered to FB (FEFB)] and two litter bag mesh sizes (8.0 and 0.2 mm) that either permitted or denied access to the leaf litter by most invertebrates. We found a significant interaction effect between fire regime and mesh size in losses of litter mass and net carbon (C). Compared with the regime of FE, with more frequent burning (FB and FEFB) the pace of decomposition was slowed by 41% (when access to litter by most invertebrates is not impeded). For the regime of FE, denying access to leaf litter by most invertebrates did not alter the pace of decomposition. Conversely, under regimes of frequently burning, restricting access to the litter by most invertebrates altered the pace of decomposition by 46%. Similar results were found for net C. For net losses of nitrogen (N), no interaction effects between fire regime and mesh size were detected, although both main effects were significant. Our results show that by modifying disturbance regimes such as fire frequency, global climate change has the potential to modify the mechanism by which ecosystems function. With more FB, decomposition is driven not only by fire regime induced changes in substrate quality and/or physiochemical conditions but through the interaction of disturbance regime with animal assemblages mediating ecosystem processes.     

Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes

Climate and litter quality have been identified as major drivers of litter decomposition at large spatial scales. However, the role played by soil fauna remains largely unknown, despite its importance for litter fragmentation and microbial activity. We synthesised litterbag studies to quantify the effect sizes of soil fauna on litter decomposition rates at the global and biome scales, and to assess how climate, litter quality and soil fauna interact to determine such rates. Soil fauna consistently enhanced litter decomposition at both global and biome scales (average increment ~ 27%). However, climate and litter quality differently modulated the effects of soil fauna on decomposition rates between biomes, from climate‐driven biomes to those where climate effects were mediated by changes in litter quality. Our results advocate for the inclusion of biome‐specific soil fauna effects on litter decomposition as a mean to reduce the unexplained variation in large‐scale decomposition models.     

森林土壤微生物对干旱和氮沉降的响应

干旱和氮沉降深刻影响着人类世森林生态系统的生命活动与物质循环,进而影响全球碳平衡、并反馈作用于气候变化。土壤微生物驱动元素的生物地球化学循环和关键土壤生态过程,在气候变化生物学研究方面具有核心地位和全球重要性。本文综述了干旱和氮沉降对森林土壤细菌和菌根真菌的影响。提出未来应加强全球变化多因子交互作用对土壤微生物多样性、活性与生态功能的研究;建立野外长期定位站,强化亚热带森林生态系统与全球变化研究;注重土壤生物之间互作及网络研究;利用微生物大数据建立相关的机理模型等。从认识微生物多样性和群落组成对全球变化的响应与适应,逐步发展为调控利用微生物群落服务于森林的优化管理、生态资源的合理保护与可持续利用,为充分发挥微生物减缓全球气候变化的作用提供理论基础。     

Timing and magnitude of drought impacts on carbon uptake across a grassland biome

Although drought is known to negatively impact grassland functioning, the timing and magnitude of these impacts within a growing season remain unresolved. Previous small-scale assessments indicate grasslands may only respond to drought during narrow periods within a year; however, large-scale assessments are now needed to uncover the general patterns and determinants of this timing. We combined remote sensing datasets of gross primary productivity and weather to assess the timing and magnitude of grassland responses to drought at 5 km² temporal resolution across two expansive ecoregions of the western US Great Plains biome: the C₄-dominated shortgrass steppe and the C₃-dominated northern mixed prairies. Across over 700,000 pixel-year combinations covering more than 600,000 km², we studied how the driest years between 2003–2020 altered the daily and bi-weekly dynamics of grassland carbon (C) uptake. Reductions to C uptake intensified into the early summer during drought and peaked in mid- and late June in both ecoregions. Stimulation of spring C uptake during drought was small and insufficient to compensate for losses during summer. Thus, total grassland C uptake was consistently reduced by drought across both ecoregions; however, reductions were twice as large across the more southern and warmer shortgrass steppe. Across the biome, increased summer vapor pressure deficit (VPD) was strongly linked to peak reductions in vegetation greenness during drought. Rising VPD will likely exacerbate reductions in C uptake during drought across the western US Great Plains, with these reductions greatest during the warmest months and in the warmest locations. High spatiotemporal resolution analyses of grassland response to drought over large areas provide both generalizable insights and new opportunities for basic and applied ecosystem science in these water-limited ecoregions amid climate change.     

Highly consistent effects of plant litter identity and functional traits on decomposition across a latitudinal gradient

Plant litter decomposition is a key process in terrestrial carbon cycling, yet the relative importance of various control factors remains ambiguous at a global scale. A full reciprocal litter transplant study with 16 litter species that varied widely in traits and originated from four forest sites covering a large latitudinal gradient (subarctic to tropics) showed a consistent interspecific ranking of decomposition rates. At a global scale, variation in decomposition was driven by a small subset of litter traits (water saturation capacity and concentrations of magnesium and condensed tannins). These consistent findings, that were largely independent of the varying local decomposer communities, suggest that decomposer communities show little specialisation and high metabolic flexibility in processing plant litter, irrespective of litter origin. Our results provide strong support for using trait-based approaches in modelling the global decomposition component of biosphere-atmosphere carbon fluxes.     

Litter mass loss rates in pine forests of Europe and Eastern United States: some relationships with climate and litter quality

The purpose of this study was to relate regional variation in litter mass-loss rates (first year) in pine forests to climate across a large, continental-scale area. The variation in mass-loss rate was analyzed using 39 experimental sites spanning climatic regions from the subarctic to subtropical and Mediterranean: the latitudinal gradient ranged from 31 °N to 70 °N and may represent the the largest geographical area that has ever been sampled and observed for the purpose of studying biogeochemical processes. Because of unified site design and uniform laboratory procedures, data from all sites were directly comparable and permitted a determination of the relative influence of climateversus substrate quality viewed from the perspective of broad regional scales. Simple correlation applied to the entire data set indicated that annual actual evapotranspiration (AET) should be the leading climatic constraint on mass-loss rates (R_adj ² = 0.496). The combination of AET, average July temp. and average annual temp. could explain about 70% of the sites' variability on litter mass-loss. In an analysis of 23 Scots pine sites north of the Alps and Carpatians AET alone could account for about 65% of the variation and the addition of a substrate-quality variable was sufficiently significant to be used in a model. The influence of litter quality was introduced into a model, using data from 11 sites at which litter of different quality had been incubated. These sites are found in Germany, the Netherlands, Sweden and Finland. At any one site most ( ≫ 90%) of the variation in mass-loss rates could be explained by one of the litter-quality variables giving concentration of nitrogen, phosphorus or water solubles. However, even when these models included nitrogen or phosphorus even small changes in potential evapotranspiration resulted in large changes in early-phase decay rates. Further regional subdivision of the data set, resulted in a range of strength in the relationship between loss rate and climatic variables, from very weak in Central Europe to strong for the Scandinavian and Atlantic coast sites (R_adj ² = 0.912; AETversus litter mass loss). Much of the variation in observed loss rates could be related to continentalversus marine/Atlantic influences. Inland locations had mass-loss rates lower than should be expected on the basis of for example AET alone. Attempts to include seasonality variables were not successful. It is clear that either unknown errors and biases, or, unknown variables are causing these regional differences in response to climatic variables. Nevertheless these results show the powerful influence of climate as a control of the broad-scale geography of mass-loss rates and substrate quality at the stand level. Some of these relationships between mass-loss rate and climatic variables are among the highest ever reported, probably because of the care taken to select uniform sites and experimental methods. This suggest that superior, base line maps of predicted mass-loss rates could be produced using climatic data. These models should be useful to predict the changing equilibrium litter dynamics resulting from climatic change.     

Long‐term amelioration of acidity accelerates decomposition in headwater streams

The secondary production of culturally acidified streams is low, with a few species of generalist detritivores dominating invertebrate assemblages, while decomposition processes are impaired. In a series of lowland headwater streams in southern England, we measured the rate of cellulolytic decomposition and compared it with values measured three decades ago, when anthropogenic acidification was at its peak. We hypothesized that, if acidity has indeed ameliorated, the rate of decomposition will have accelerated, thus potentially supporting greater secondary production and the longer food chains that have been observed in some well‐studied recovering freshwater systems. We used cellulose Shirley test cloth as a standardized bioassay to measure the rate of cellulolytic decomposition, via loss in tensile strength, for 31 streams in the Ashdown Forest over 7 days in summer 2011 and 49 days in winter 2012. We compared this with data from an otherwise identical study conducted in 1978 and 1979. In a secondary study, we determined whether decomposition followed a linear or logarithmic decay and, as Shirley cloth is no longer available, we tested an alternative in the form of readily available calico. Overall mean pH had increased markedly over the 32 years between the studies (from 6.0 to 6.7). In both the previous and contemporary studies, the relationship between decomposition and pH was strongest in winter, when pH reaches a seasonal minimum. As in the late 1970s, there was no relationship in 2011/2012 between pH and decay rate in summer. As postulated, decomposition in winter was significantly faster in 2011/2012 than in 1978/1979, with an average increase in decay rate of 18.1%. Recovery from acidification, due to decreased acidifying emissions and deposition, has led to an increase in the rate of cellulolytic decomposition. This response in a critical ecosystem process offers a potential explanation of one aspect of the limited biological recovery that has been observed so far, an increase in larger bodied predators including fish, which in turn leads to an increase in the length of food chains.     

Decoupling the direct and indirect effects of climate on plant litter decomposition: Accounting for stress‐induced modifications in plant chemistry

Decomposition of plant litter is a fundamental ecosystem process that can act as a feedback to climate change by simultaneously influencing both the productivity of ecosystems and the flux of carbon dioxide from the soil. The influence of climate on decomposition from a postsenescence perspective is relatively well known; in particular, climate is known to regulate the rate of litter decomposition via its direct influence on the reaction kinetics and microbial physiology on processes downstream of tissue senescence. Climate can alter plant metabolism during the formative stage of tissues and could shape the final chemical composition of plant litter that is available for decomposition, and thus indirectly influence decomposition; however, these indirect effects are relatively poorly understood. Climatic stress disrupts cellular homeostasis in plants and results in the reprogramming of primary and secondary metabolic pathways, which leads to changes in the quantity, composition, and organization of small molecules and recalcitrant heteropolymers, including lignins, tannins, suberins, and cuticle within the plant tissue matrix. Furthermore, by regulating metabolism during tissue senescence, climate influences the resorption of nutrients from senescing tissues. Thus, the final chemical composition of plant litter that forms the substrate of decomposition is a combined product of presenescence physiological processes through the production and resorption of metabolites. The changes in quantity, composition, and localization of the molecular construct of the litter could enhance or hinder tissue decomposition and soil nutrient cycling by altering the recalcitrance of the lignocellulose matrix, the composition of microbial communities, and the activity of microbial exo‐enzymes via various complexation reactions. Also, the climate‐induced changes in the molecular composition of litter could differentially influence litter decomposition and soil nutrient cycling. Compared with temperate ecosystems, the indirect effects of climate on litter decomposition in the tropics are not well understood, which underscores the need to conduct additional studies in tropical biomes. We also emphasize the need to focus on how climatic stress affects the root chemistry as roots contribute significantly to biogeochemical cycling, and on utilizing more robust analytical approaches to capture the molecular composition of tissue matrix that fuel microbial metabolism.     

Effects of diversity loss on ecosystem function across trophic levels and ecosystems: A test in a detritus‐based tropical food web

Abstract We investigated the effects of biodiversity loss across trophic levels and across ecosystems (terrestrial to aquatic) on ecosystem function, in a detritus‐based tropical food web. Diversities of consumers (stream shredders) and resources (leaf litter) were experimentally manipulated by varying the number of species from 3 to 1, using different species combinations, and the effects on leaf breakdown rates were examined. In single‐species shredder treatments, leaf diversity loss affected breakdown rates, but the effect depended on the identity of the leaves remaining in the system: they increased when the most preferred leaf species remained, but decreased when this species was lost (leaf preferences were the same for all shredders). In multi‐species shredder assemblages, breakdown rates were lower than expected from single‐species treatments, suggesting an important role of interspecific competition. This pattern was also evident when oneleaf species was available but not with higher leaf diversity, suggesting that lowered leaf diversity promotes competitive interactions among shredders. The influence of diversity and identity of species across trophic levels and ecosystems on stream functioning points to complex interactions that may well be reflected in other types of ecosystem.     

Outcomes of fungal interactions are determined by soil invertebrate grazers

Saprotrophic fungal community composition, determined by the outcome of competitive mycelial interactions, is one of the many key factors affecting soil nutrient mineralisation and decomposition rates. Fungal communities are not generally predicted to be regulated by top-down factors, such as predation, but rather by bottom-up factors, including resource availability. We show that invertebrate grazers can exert selective pressures on fungal decomposer communities in soil, reversing the outcomes of competitive interactions. By feeding selectively on the cord-forming fungus Resinicium bicolor, isopods prevented the competitive exclusion of Hypholoma fasciculare and Phanerochaete velutina in soil and wood. Nematode populations also reversed the outcomes of competitive interactions by stimulating growth of less competitive fungi. These represent two opposing mechanisms by which soil fauna may influence fungal community composition and diversity. Factors affecting soil invertebrate communities will have direct consequences for fungal-mediated nutrient cycling in woodland soils.     

Investigating the long‐term legacy of drought and warming on the soil microbial community across five European shrubland ecosystems

We investigated how the legacy of warming and summer drought affected microbial communities in five different replicated long‐term (>10 years) field experiments across Europe (EU‐FP7 INCREASE infrastructure). To focus explicitly on legacy effects (i.e., indirect rather than direct effects of the environmental factors), we measured microbial variables under the same moisture and temperature in a brief screening, and following a pre‐incubation at stable conditions. Specifically, we investigated the size and composition of the soil microbial community (PLFA) alongside measurements of bacterial (leucine incorporation) and fungal (acetate in ergosterol incorporation) growth rates, previously shown to be highly responsive to changes in environmental factors, and microbial respiration. We found no legacy effects on the microbial community size, composition, growth rates, or basal respiration rates at the effect sizes used in our experimental setup (0.6 °C, about 30% precipitation reduction). Our findings support previous reports from single short‐term ecosystem studies thereby providing a clear evidence base to allow long‐term, broad‐scale generalizations to be made. The implication of our study is that warming and summer drought will not result in legacy effects on the microbial community and their processes within the effect sizes here studied. While legacy effects on microbial processes during perturbation cycles, such as drying–rewetting, and on tolerance to drought and warming remain to be studied, our results suggest that any effects on overall ecosystem processes will be rather limited. Thus, the legacies of warming and drought should not be prioritized factors to consider when modeling contemporary rates of biogeochemical processes in soil.     

Synergistic effects of water temperature and dissolved nutrients on litter decomposition and associated fungi

In woodland streams, the decomposition of allochthonous organic matter constitutes a fundamental ecosystem process, where aquatic hyphomycetes play a pivotal role. It is therefore greatly affected by water temperature and nutrient concentrations. The individual effects of these factors on the decomposition of litter have been studied previously. However, in the climate warming scenario predicted for this century, water temperature and nutrient concentrations are expected to increase simultaneously, and their combined effects on litter decomposition and associated biological activity remains unevaluated. In this study, we addressed the individual and combined effects of water temperature (three levels) and nutrient concentrations (two levels) on the decomposition of alder leaves and associated aquatic hyphomycetes in microcosms. Decomposition rates across treatments varied between 0.0041 day^?1 at 5 °C and low nutrient level and 0.0100 day^?1 at 15 °C and high nutrient level. The stimulation of biological variables at high nutrients and temperatures indicates that nutrient enrichment of streams might have a higher stimulatory effect on fungal performance and decomposition rates under a warming scenario than at present. The stimulation of fungal biomass and sporulation with increasing temperature at both nutrient levels shows that increases in water temperature might enhance fungal growth and reproduction in both oligotrophic and eutrophic streams. The stimulation of fungal respiration and litter decomposition with increasing temperature at high nutrients indicates that stimulation of carbon mineralization will probably occur at eutrophied streams, while oligotrophic conditions seem to be ‘protected’ from warming. All biological variables were stimulated when both factors increased, as a result of synergistic interactions between factors. Increased water temperature and nutrient level also affected the structure of aquatic hyphomycete assemblages. It is plausible that if water quality of presently eutrophied streams is improved, the potential stimulatory effects of future increases in water temperature on aquatic biota and processes might be mitigated.     

Drought‐resistant fungi control soil organic matter decomposition and its response to temperature

Microbial‐mediated decomposition of soil organic matter (SOM) ultimately makes a considerable contribution to soil respiration, which is typically the main source of CO₂ arising from terrestrial ecosystems. Despite this central role in the decomposition of SOM, few studies have been conducted on how climate change may affect the soil microbial community and, furthermore, on how possible climate‐change induced alterations in the ecology of microbial communities may affect soil CO₂ emissions. Here we present the results of a seasonal study on soil microbial community structure, SOM decomposition and its temperature sensitivity in two representative Mediterranean ecosystems where precipitation/throughfall exclusion has taken place during the last 10 years. Bacterial and fungal diversity was estimated using the terminal restriction fragment length polymorphism technique. Our results show that fungal diversity was less sensitive to seasonal changes in moisture, temperature and plant activity than bacterial diversity. On the other hand, fungal communities showed the ability to dynamically adapt throughout the seasons. Fungi also coped better with the 10 years of precipitation/throughfall exclusion compared with bacteria. The high resistance of fungal diversity to changes with respect to bacteria may open the controversy as to whether future ‘drier conditions’ for Mediterranean regions might favor fungal dominated microbial communities. Finally, our results indicate that the fungal community exerted a strong influence over the temporal and spatial variability of SOM decomposition and its sensitivity to temperature. The results, therefore, highlight the important role of fungi in the decomposition of terrestrial SOM, especially under the harsh environmental conditions of Mediterranean ecosystems, for which models predict even drier conditions in the future.