Trickle-down effects of aboveground trophic cascades on the soil food web

Trophic cascades are increasingly being regarded as important features of aboveground and belowground food webs, but the effects of aboveground cascades on soil food webs, and vice versa, remains essentially unexplored. We conducted an experiment consisting of model synthesised communities containing grassland plant and invertebrate species, in which treatments included soil only, soil+plants, soil+plants+aphids, and soil+plants+aphids+predators; predator treatments consisted of the lacewing Micromus tasmaniae and ladybird beetle Coccinella undecimpunctata added either singly or in combination. Addition of Micromus largely reversed the negative effects of aphids on plant biomass, while both of the predator species caused large changes in the relative abundances of dominant plant species. Predators of aphids also affected several components of the belowground subsystem. Micromus had positive indirect effects on the primary consumer of the soil decomposer food web (microflora), probably through promoting greater input of basal resources to the decomposer subsystem. Predator treatments also influenced densities of the tertiary consumers of the soil food web (top predatory nematodes), most likely through inducing changes in plant community composition and therefore the quality of resource input to the soil. The secondary consumers of the soil food web (microbe‐feeding nematodes) were, however, unresponsive. The fact that some trophic levels of the soil food web but not others responded to aboveground manipulations is explicable in terms of top‐down and bottom‐up forces differentially regulating different belowground trophic levels. Addition of aphids also influenced microbial community structure, promoted soil bacteria at the expense of fungi, and enhanced the diversity of herbivorous nematodes; in all cases these effects were at least partially reversed by addition of Micromus. These results in tandem point to upper level consumers in aboveground food webs as a potential driver of the belowground subsystem, and provide evidence that predator‐induced trophic cascades aboveground can have effects that trickle through soil food webs.     

The response of a three trophic level soil food web to the identity and diversity of plant species and functional groups

Despite considerable recent interest in how biodiversity may influence ecosystem properties, the issue of how plant diversity and composition may affect multiple trophic levels in soil food webs remains essentially unexplored. We conducted a glasshouse experiment in which three plant species of each of three functional groups (grasses, N‐fixing legumes and forbs) were grown in monoculture and in mixtures of three species (with the three species being in the same or different functional groups) and all nine species. Plant species identity had important effects on the biomasses or population densities of belowground primary consumers (microbial biomass, herbivorous nematodes) and two groups of secondary consumers (microbe‐feeding nematodes and enchytraeids); the third consumer trophic level (predatory nematodes) was marginally not significantly affected at P=0.05. Plant species also influenced the relative importance of the bacterial‐based and fungal‐based energy channels for both the primary and secondary consumer trophic levels. Within‐group diversity of only the soil microflora and herbivorous nematodes (both representing the basal consumer trophic level) were affected by plant species identity. However, community composition within all trophic groupings considered (herbivorous nematodes, microbes, microbe‐feeding nematodes, predatory nematodes) was strongly influenced by what plant species were present. Despite the strong responses of the soil biota to plant species identity, there were few effects of plant species or functional group richness on any of the belowground response variables measured. Further, net primary productivity (NPP) was unaffected by plant diversity. Since some belowground response variables were correlated with NPP across treatments, it is suggested that belowground responses to plant diversity might become more apparent in situations when NPP itself responds to plant diversity. Our results point to plant species identity as having important multitrophic effects on soil food webs, both at the whole trophic group and within‐group levels of resolution, and suggest that differences in plant traits across species may be important in driving the decomposer subsystem.     

Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition

Despite the major importance of soil biota in nutrient and energy fluxes, interactions in soil food webs are poorly understood. Here we provide an overview of recent advances in uncovering the trophic structure of soil food webs using natural variations in stable isotope ratios. We discuss approaches of application, normalization and interpretation of stable isotope ratios along with methodological pitfalls. Analysis of published data from temperate forest ecosystems is used to outline emerging concepts and perspectives in soil food web research. In contrast to aboveground and aquatic food webs, trophic fractionation at the basal level of detrital food webs is large for carbon and small for nitrogen stable isotopes. Virtually all soil animals are enriched in ¹³C as compared to plant litter. This ‘detrital shift’ likely reflects preferential uptake of ¹³C‐enriched microbial biomass and underlines the importance of microorganisms, in contrast to dead plant material, as a major food resource for the soil animal community. Soil organic matter is enriched in ¹⁵N and ¹³C relative to leaf litter. Decomposers inhabiting mineral soil layers therefore might be enriched in ¹⁵N resulting in overlap in isotope ratios between soil‐dwelling detritivores and litter‐dwelling predators. By contrast, ¹³C content varies little between detritivores in upper litter and in mineral soil, suggesting that they rely on similar basal resources, i.e. little decomposed organic matter. Comparing vertical isotope gradients in animals and in basal resources can be a valuable tool to assess trophic interactions and dynamics of organic matter in soil. As indicated by stable isotope composition, direct feeding on living plant material as well as on mycorrhizal fungi is likely rare among soil invertebrates. Plant carbon is taken up predominantly by saprotrophic microorganisms and channelled to higher trophic levels of the soil food web. However, feeding on photoautotrophic microorganisms and non‐vascular plants may play an important role in fuelling soil food webs. The trophic niche of most high‐rank animal taxa spans at least two trophic levels, implying the use of a wide range of resources. Therefore, to identify trophic species and links in food webs, low‐rank taxonomic identification is required. Despite overlap in feeding strategies, stable isotope composition of the high‐rank taxonomic groups reflects differences in trophic level and in the use of basal resources. Different taxonomic groups of predators and decomposers are likely linked to different pools of organic matter in soil, suggesting different functional roles and indicating that trophic niches in soil animal communities are phylogenetically structured. During last two decades studies using stable isotope analysis have elucidated the trophic structure of soil communities, clarified basal food resources of the soil food web and revealed links between above‐ and belowground ecosystem compartments. Extending the use of stable isotope analysis to a wider range of soil‐dwelling organisms, including microfauna, and a larger array of ecosystems provides the perspective of a comprehensive understanding of the structure and functioning of soil food webs.     

Linking aboveground and belowground food webs through carbon and nitrogen stable isotope analyses

Carbon and nitrogen stable isotope ratios (δ¹³C and δ¹⁵N) have been used for more than two decades in analyses of food web structure. The utility of isotope ratio measurements is based on the observation that consumer δ¹³C values are similar (<1‰ difference) to those of their diet, while consumer δ¹⁵N values are about 3‰ higher than those of their diet. The technique has been applied most often to aquatic and aboveground terrestrial food webs. However, few isotope studies have examined terrestrial food web structure that includes both above- and belowground (detrital) components. Here, we review factors that may influence isotopic signatures of terrestrial consumers in above- and belowground systems. In particular, we emphasize variations in δ¹³C and δ¹⁵N in belowground systems, e.g., enrichment of ¹³C and ¹⁵N in soil organic matter (likely related to soil microbial metabolism). These enrichments should be associated with the high ¹³C (~3‰) enrichment in belowground consumers relative to litter and soil organic matter and with the large variation in δ¹⁵N (~6‰) of the consumers. Because such enrichment and variation are much greater than the trophic enrichment generally used to estimate consumer trophic positions, and because many general predators are considered dependent on energy and material flows from belowground, the isotopic variation in belowground systems should be taken into account in δ¹³C and δ¹⁵N analyses of terrestrial food webs. Meanwhile, by measuring the δ¹³C of key predators, the linkage between above- and belowground systems could be estimated based on observed differences in δ¹³C of primary producers, detritivores and predators. Furthermore, radiocarbon (¹⁴C) measurements will allow the direct estimation of the dependence of predators on the belowground systems.     

Grazing simplifies soil micro‐food webs and decouples their relationships with ecosystem functions in grasslands

Livestock grazing often alters aboveground and belowground communities of grasslands and their mediated carbon (C) and nitrogen (N) cycling processes at the local scale. Yet, few have examined whether grazing‐induced changes in soil food webs and their ecosystem functions can be extrapolated to a regional scale. We investigated how large herbivore grazing affects soil micro‐food webs (microbes and nematodes) and ecosystem functions (soil C and N mineralization), using paired grazed and ungrazed plots at 10 locations across the Mongolian Plateau. Our results showed that grazing not only affected plant variables (e.g., biomass and C and N concentrations), but also altered soil substrates (e.g., C and N contents) and soil environment (e.g., soil pH and bulk density). Grazing had strong bottom‐up effects on soil micro‐food webs, leading to more pronounced decreases at higher trophic levels (nematodes) than at lower trophic levels (microbes). Structural equation modeling showed that changes in plant biomass and soil environment dominated grazing effects on microbes, while nematodes were mainly influenced by changes in plant biomass and soil C and N contents; the grazing effects, however, differed greatly among functional groups in the soil micro‐food webs. Grazing reduced soil C and N mineralization rates via changes in plant biomass, soil C and N contents, and soil environment across grasslands on the Mongolian Plateau. Spearman's rank correlation analysis also showed that grazing reduced the correlations between functional groups in soil micro‐food webs and then weakened the correlation between soil micro‐food webs and soil C and N mineralization. These results suggest that changes in soil micro‐food webs resulting from livestock grazing are poor predictors of soil C and N processes at regional scale, and that the relationships between soil food webs and ecosystem functions depend on spatial scales and land‐use changes.     

Global change belowground: impacts of elevated CO2, nitrogen,and summer drought on soil food webs and biodiversity

The world's ecosystems are subjected to various anthropogenic global change agents, such as enrichment of atmospheric CO₂ concentrations, nitrogen (N) deposition, and changes in precipitation regimes. Despite the increasing appreciation that the consequences of impending global change can be better understood if varying agents are studied in concert, there is a paucity of multi‐factor long‐term studies, particularly on belowground processes. Herein, we address this gap by examining the responses of soil food webs and biodiversity to enrichment of CO₂, elevated N, and summer drought in a long‐term grassland study at Cedar Creek, Minnesota, USA (BioCON experiment). We use structural equation modeling (SEM), various abiotic and biotic explanatory variables, and data on soil microorganisms, protozoa, nematodes, and soil microarthropods to identify the impacts of multiple global change effects on drivers belowground. We found that long‐term (13‐year) changes in CO₂ and N availability resulted in modest alterations of soil biotic food webs and biodiversity via several mechanisms, encompassing soil water availability, plant productivity, and – most importantly – changes in rhizodeposition. Four years of manipulation of summer drought exerted surprisingly minor effects, only detrimentally affecting belowground herbivores and ciliate protists at elevated N. Elevated CO₂ increased microbial biomass and the density of ciliates, microarthropod detritivores, and gamasid mites, most likely by fueling soil food webs with labile C. Moreover, beneficial bottom‐up effects of elevated CO₂ compensated for detrimental elevated N effects on soil microarthropod taxa richness. In contrast, nematode taxa richness was lowest at elevated CO₂ and elevated N. Thus, enrichment of atmospheric CO₂ concentrations and N deposition may result in taxonomically and functionally altered, potentially simplified, soil communities. Detrimental effects of N deposition on soil biodiversity underscore recent reports on plant community simplification. This is of particular concern, as soils house a considerable fraction of global biodiversity and ecosystem functions.     

Conversion of rainforest to oil palm and rubber plantations alters energy channels in soil food webs

In the last decades, lowland tropical rainforest has been converted in large into plantation systems. Despite the evident changes above ground, the effect of rainforest conversion on the channeling of energy in soil food webs was not studied. Here, we investigated community‐level neutral lipid fatty acid profiles in dominant soil fauna to track energy channels in rainforest, rubber, and oil palm plantations in Sumatra, Indonesia. Abundant macrofauna including Araneae, Chilopoda, and Diplopoda contained high amounts of plant and fungal biomarker fatty acids (FAs). Lumbricina had the lowest amount of plant, but the highest amount of animal‐synthesized C20 polyunsaturated FAs as compared to other soil taxa. Mesofauna detritivores (Collembola and Oribatida) contained high amounts of algal biomarker FAs. The differences in FA profiles between taxa were evident if data were analyzed across land‐use systems, suggesting that soil fauna of different size (macro‐ and mesofauna) are associated with different energy channels. Despite that, rainforest conversion changed the biomarker FA composition of soil fauna at the community level. Conversion of rainforest into oil palm plantations enhanced the plant energy channel in soil food webs and reduced the bacterial energy channel; conversion into rubber plantations reduced the AMF‐based energy channel. The changes in energy distribution within soil food webs may have significant implications for the functioning of tropical ecosystems and their response to environmental changes. At present, these responses are hard to predict considering the poor knowledge on structure and functioning of tropical soil food webs.     

Coupled biochar amendment and limited irrigation strategies do not affect a degraded soil food web in a maize agroecosystem,compared to the native grassland

Climate change is predicted to increase climate variability and frequency of extreme events such as drought, straining water resources in agricultural systems. Thus, limited irrigation strategies and soil amendments are being explored to conserve water in crop production. Biochar is the recalcitrant, carbon‐based coproduct of biomass pyrolysis during bioenergy production. When used as a soil amendment, biochar can increase soil water retention while enhancing soil properties and stimulating food webs. We investigated the effects of coupled biochar amendment and limited irrigation on belowground food web structure and function in an irrigated maize agroecosystem. We hypothesized that soil biota biomass and activity would decrease with limited irrigation and increase with biochar amendment and that biochar amendment would mitigate the impact of limited irrigation on the soil food web. One year after biochar addition, we extracted, identified, and estimated the biomass of taxonomic groups of soil biota (e.g., bacteria, fungi, protozoa, nematodes, and arthropods) from wood‐derived biochar‐amended (30 Mg ha^?1) and nonamended soils under maize with limited (two‐thirds of full) and full irrigation. We modeled structural and functional properties of the soil food web. Neither biochar amendment nor limited irrigation had a significant effect on biomass of the soil biota groups. Modeled soil respiration and nitrogen mineralization fluxes were not different between treatments. A comparison of the structure and function of the agroecosystem soil food web and a nearby native grassland revealed that in this temperate system, the negative impact of long‐term conventional agricultural management outweighed the impact of limited irrigation. One year of biochar amendment did not mitigate nor further contribute to the negative effects of historical agricultural management.     

Trophic level and basal resource use of soil animals are hardly affected by local plant associations in abandoned arable land

Plants provide resources and shape the habitat of soil organisms thereby affecting the composition and functioning of soil communities. Effects of plants on soil communities are largely taxon‐dependent, but how different functional groups of herbaceous plants affect trophic niches of individual animal species in soil needs further investigation. Here, we studied the use of basal resources and trophic levels of dominating soil meso‐ and macrofauna using stable isotope ratios of carbon and nitrogen in arable fallow systems 3 and 14–16 years after abandonment. Animals were sampled from the rhizosphere of three plant species of different functional groups: a legume (Medicaco sativa), a nonlegume herb (Taraxacum officinale), and a grass (Bromus sterilis). We found virtually no consistent effects of plant identity on stable isotope composition of soil animals and on thirteen isotopic metrics that reflect general food‐web structure. However, in old fallows, the carbon isotope composition of some predatory macrofauna taxa had shifted closer to that of co‐occurring plants, which was particularly evident for Lasius, an aphid‐associated ant genus. Trophic levels and trophic‐chain lengths in food webs were similar across plant species and fallow ages. Overall, the results suggest that variations in local plant diversity of grassland communities may little affect the basal resources and the trophic level of prey consumed by individual species of meso‐ and macrofauna belowground. By contrast, successional changes in grassland communities are associated with shifts in the trophic niches of certain species, reflecting establishment of trophic interactions with time, which shapes the functioning and stability of soil food webs.     

Stable isotope analysis of aquatic invertebrate communities in irrigated rice fields cultivated under different management regimes

In this study we have used stable isotope analysis to identify major food resources driving food webs in commercial rice agroecosystems and to examine the effects of agricultural management practices on the trophic structure of these food webs. Potential carbon sources and aquatic macroinvertebrate consumers were collected from large-scale rice farms in south-eastern Australia cultivated under three different crop management regimes conventional-aerial (agrochemicals applied, aerially sown), conventional-sod (agrochemicals applied, directly sown) and organic-sod (agrochemical-free, directly sown). Evidence from stable isotope analysis demonstrated the importance of food sources, such as biofilm and detritus, as the principal energy sources driving aquatic food webs in rice agroecosystems. Despite the greater diversity of potential food sources collected from the organic-sod regime across all sampling occasions, the range of food resources directly assimilated by macroinvertebrate consumers did not differ substantially across management regimes. Trophic complexity of aquatic food webs, as evidenced by the number of trophic levels identified using δ¹⁵N data, differed across management regimes at the early season sampling. Sites with low or no agrochemical applications contained more than two trophic levels, but at the site with the highest pesticide application no primary or secondary consumers were found. Our data demonstrates that the choice of agricultural management regime has a season-long influence on aquatic food webs in rice crops, and highlights the importance of conserving non-rice food resources that drive these trophic networks.     

Leaf breakdown, detrital resources, and food webs in streams affected by mine drainage

Breakdown of leaf litter is essential for providing detrital resources for food webs but can be impaired by anthropogenic activities, which may disrupt energy flow to consumers. We investigated the relationship between leaf breakdown and food web structure in 12 streams with or without mining impacts on South Island, New Zealand. Six streams received inputs of acid mine drainage (pH 2.5–4.9), three were naturally acidic (pH ~5.0), and three were circumneutral (pH ~6.8). Streams affected by mining either had highly acidic water (pH <3) or iron precipitates present on substrata. Breakdown rates of leaves were significantly lower in mining-affected streams than circumneutral (by almost 50%) but not naturally acidic streams and were driven primarily by microbial activity, as shredding invertebrates were often absent. Mining-affected stream webs were simplified structures with fewer species and links than those in other streams. With few species to process leaf litter and transfer detrital resources, inputs of AMD disrupted both the mechanisms responsible for breakdown and links for energy flow. While faster breakdown rates were associated with larger food webs, limited function maintained in mining-affected streams was sufficient to support primary consumers and small food webs.     

From clear lakes to murky waters – tracing the functional response of high‐latitude lake communities to concurrent ‘greening’ and ‘browning’

Climate change and the intensification of land use practices are causing widespread eutrophication of subarctic lakes. The implications of this rapid change for lake ecosystem function remain poorly understood. To assess how freshwater communities respond to such profound changes in their habitat and resource availability, we conducted a space‐for‐time analysis of food‐web structure in 30 lakes situated across a temperature‐productivity gradient equivalent to the predicted future climate of subarctic Europe (temperature +3°C, precipitation +30% and nutrient +45 μg L^?1 total phosphorus). Along this gradient, we observed an increase in the assimilation of pelagic‐derived carbon from 25 to 75% throughout primary, secondary and tertiary consumers. This shift was overwhelmingly driven by the consumption of pelagic detritus by benthic primary consumers and was not accompanied by increased pelagic foraging by higher trophic level consumers. Our data also revealed a convergence of the carbon isotope ratios of pelagic and benthic food web endmembers in the warmest, most productive lakes indicating that the incorporation of terrestrial derived carbon into aquatic food webs increases as land use intensifies. These results, reflecting changes along a gradient characteristic of the predicted future environment throughout the subarctic, indicate that climate and land use driven eutrophication and browning are radically altering the function and fuelling of aquatic food webs in this biome.     

Long‐term effects of climate change on carbon flows through benthic secondary production in small lakes

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Food Web Structure and Basal Resource Utilization along a Tropical Island Stream Continuum, Puerto Rico

Tropical stream food webs are thought to be based primarily on terrestrial resources (leaf litter) in small forested headwater streams and algal resources in larger, wider streams. In tropical island streams, the dominant consumers are often omnivorous freshwater shrimps that consume algae, leaf litter, insects, and other shrimps. We used stable isotope analysis to examine (1) the relative importance of terrestrial and algal‐based food resources to shrimps and other consumers and determine (2) if the relative importance of these food resources changed along the stream continuum. We examined δ¹⁵N and δ¹³C signatures of leaves, algae, macrophytes, biofilm, insects, snails, fishes, and shrimps at three sites (300, 90, and 10 m elev.) along the Río Espíritu Santo, which drains the Caribbean National Forest, Puerto Rico. Isotope signatures of basal resources were distinct at all sites. Results of two‐source δ¹³C mixing models suggest that shrimps relied more on algal‐based carbon resources than terrestrially derived resources at all three sites along the continuum. This study supports other recent findings in tropical streams, demonstrating that algal‐based resources are very important to stream consumers, even in small forested headwater streams. This study also demonstrates the importance of doing assimilation‐based analysis (i.e., stable isotope or trophic basis of production) when studying food webs.     

Allometric constraints on,and trade‐offs in,belowground carbon allocation and their control of soil respiration across global forest ecosystems

To fully understand how soil respiration is partitioned among its component fluxes and responds to climate, it is essential to relate it to belowground carbon allocation, the ultimate carbon source for soil respiration. This remains one of the largest gaps in knowledge of terrestrial carbon cycling. Here, we synthesize data on gross and net primary production and their components, and soil respiration and its components, from a global forest database, to determine mechanisms governing belowground carbon allocation and their relationship with soil respiration partitioning and soil respiration responses to climatic factors across global forest ecosystems. Our results revealed that there are three independent mechanisms controlling belowground carbon allocation and which influence soil respiration and its partitioning: an allometric constraint; a fine‐root production vs. root respiration trade‐off; and an above‐ vs. belowground trade‐off in plant carbon. Global patterns in soil respiration and its partitioning are constrained primarily by the allometric allocation, which explains some of the previously ambiguous results reported in the literature. Responses of soil respiration and its components to mean annual temperature, precipitation, and nitrogen deposition can be mediated by changes in belowground carbon allocation. Soil respiration responds to mean annual temperature overwhelmingly through an increasing belowground carbon input as a result of extending total day length of growing season, but not by temperature‐driven acceleration of soil carbon decomposition, which argues against the possibility of a strong positive feedback between global warming and soil carbon loss. Different nitrogen loads can trigger distinct belowground carbon allocation mechanisms, which are responsible for different responses of soil respiration to nitrogen addition that have been observed. These results provide new insights into belowground carbon allocation, partitioning of soil respiration, and its responses to climate in forest ecosystems and are, therefore, valuable for terrestrial carbon simulations and projections.     

Correlation between interaction strengths drives stability in large ecological networks

Food webs have markedly non‐random network structure. Ecologists maintain that this non‐random structure is key for stability, since large random ecological networks would invariably be unstable and thus should not be observed empirically. Here we show that a simple yet overlooked feature of natural food webs, the correlation between the effects of consumers on resources and those of resources on consumers, substantially accounts for their stability. Remarkably, random food webs built by preserving just the distribution and correlation of interaction strengths have stability properties similar to those of the corresponding empirical systems. Surprisingly, we find that the effect of topological network structure on stability, which has been the focus of countless studies, is small compared to that of correlation. Hence, any study of the effects of network structure on stability must first take into account the distribution and correlation of interaction strengths.     

Above‐ and belowground linkages in Sphagnum peatland: climate warming affects plant‐microbial interactions

Peatlands contain approximately one third of all soil organic carbon (SOC). Warming can alter above‐ and belowground linkages that regulate soil organic carbon dynamics and C‐balance in peatlands. Here we examine the multiyear impact of in situ experimental warming on the microbial food web, vegetation, and their feedbacks with soil chemistry. We provide evidence of both positive and negative impacts of warming on specific microbial functional groups, leading to destabilization of the microbial food web. We observed a strong reduction (70%) in the biomass of top‐predators (testate amoebae) in warmed plots. Such a loss caused a shortening of microbial food chains, which in turn stimulated microbial activity, leading to slight increases in levels of nutrients and labile C in water. We further show that warming altered the regulatory role of Sphagnum‐polyphenols on microbial community structure with a potential inhibition of top predators. In addition, warming caused a decrease in Sphagnum cover and an increase in vascular plant cover. Using structural equation modelling, we show that changes in the microbial food web affected the relationships between plants, soil water chemistry, and microbial communities. These results suggest that warming will destabilize C and nutrient recycling of peatlands via changes in above‐ and belowground linkages, and therefore, the microbial food web associated with mosses will feedback positively to global warming by destabilizing the carbon cycle. This study confirms that microbial food webs thus constitute a key element in the functioning of peatland ecosystems. Their study can help understand how mosses, as ecosystem engineers, tightly regulate biogeochemical cycling and climate feedback in peatlands     

Soil acidification reduces the effects of short‐term nutrient enrichment on plant and soil biota and their interactions in grasslands

Soil nitrogen (N) and phosphorus (P) contents, and soil acidification have greatly increased in grassland ecosystems due to increased industrial and agricultural activities. As major environmental and economic concerns worldwide, nutrient enrichment and soil acidification can lead to substantial changes in the diversity and structure of plant and soil communities. Although the separate effects of N and P enrichment on soil food webs have been assessed across different ecosystems, the combined effects of N and P enrichment on multiple trophic levels in soil food webs have not been studied in semiarid grasslands experiencing soil acidification. Here we conducted a short‐term N and P enrichment experiment in non‐acidified and acidified soil in a semiarid grassland on the Mongolian Plateau. We found that net primary productivity was not affected by N or P enrichment alone in either non‐acidified or acidified soil, but was increased by combined N and P enrichment in both non‐acidified and acidified soil. Nutrient enrichment decreased the biomass of most microbial groups in non‐acidified soil (the decrease tended to be greatest with combined N and P enrichment) but not in acidified soil, and did not affect most soil nematode variables in non‐acidified or acidified soil. Nutrient enrichment also changed plant and microbial community structure in non‐acidified but not in acidified soil, and had no effect on nematode community structure in non‐acidified or acidified soil. These results indicate that the responses to short‐term nutrient enrichment were weaker for higher trophic groups (nematodes) than for lower trophic groups (microorganisms) and primary producers (plants). The findings increase our understanding of the effects of nutrient enrichment on multiple trophic levels of soil food webs, and highlight that soil acidification, as an anthropogenic stressor, reduced the responses of plants and soil food webs to nutrient enrichment and weakened plant–soil interactions.     

Free‐living nematodes as prey for higher trophic levels of forest soil food webs

Nematodes are the most abundant invertebrates in soils and are key prey in soil food webs. Uncovering their contribution to predator nutrition is essential for understanding the structure of soil food webs and the way energy channels through soil systems. Molecular gut content analysis of consumers of nematodes, such as soil microarthropods, using specific DNA markers is a novel approach for studying predator–prey interactions in soil. We designed new specific primer pairs (partial 18S rDNA) for individual soil‐living bacterial‐feeding nematode taxa (Acrobeloides buetschlii, Panagrellus redivivus, Plectus velox and Plectus minimus). Primer specificity was tested against more than 100 non‐target soil organisms. Further, we determined how long nematode DNA can be traced in the gut of predators. Potential predators were identified in laboratory experiments including nine soil mite (Oribatida, Gamasina and Uropodina) and ten springtail species (Collembola). Finally, the approach was tested under field conditions by analyzing five mite and three collembola species for feeding on the three target nematode species. The results proved the three primer sets to specifically amplify DNA of the respective nematode taxa. Detection time of nematode DNA in predators varied with time of prey exposure. Further, consumption of nematodes in the laboratory varied with microarthropod species. Our field study is the first definitive proof that free‐living nematodes are important prey for a wide range of soil microarthropods including those commonly regarded as detritivores. Overall, the results highlight the eminent role of nematodes as prey in soil food webs and for channelling bacterial carbon to higher trophic levels.     

A global study of relationships between leaf traits, climate and soil measures of nutrient fertility

Aim This first global quantification of the relationship between leaf traits and soil nutrient fertility reflects the trade‐off between growth and nutrient conservation. The power of soils versus climate in predicting leaf trait values is assessed in bivariate and multivariate analyses and is compared with the distribution of growth forms (as a discrete classification of vegetation) across gradients of soil fertility and climate. Location All continents except for Antarctica. Methods Data on specific leaf area (SLA), leaf N concentration (LNC), leaf P concentration (LPC) and leaf N:P were collected for 474 species distributed across 99 sites (809 records), together with abiotic information from each study site. Individual and combined effects of soils and climate on leaf traits were quantified using maximum likelihood methods. Differences in occurrence of growth form across soil fertility and climate were determined by one‐way ANOVA. Results There was a consistent increase in SLA, LNC and LPC with increasing soil fertility. SLA was related to proxies of N supply, LNC to both soil total N and P and LPC was only related to proxies of P supply. Soil nutrient measures explained more variance in leaf traits among sites than climate in bivariate analysis. Multivariate analysis showed that climate interacted with soil nutrients for SLA and area‐based LNC. Mass‐based LNC and LPC were determined mostly by soil fertility, but soil P was highly correlated to precipitation. Relationships of leaf traits to soil nutrients were stronger than those of growth form versus soil nutrients. In contrast, climate determined distribution of growth form more strongly than it did leaf traits. Main conclusions We provide the first global quantification of the trade‐off between traits associated with growth and resource conservation ‘strategies’ in relation to soil fertility. Precipitation but not temperature affected this trade‐off. Continuous leaf traits might be better predictors of plant responses to nutrient supply than growth form, but growth forms reflect important aspects of plant species distribution with climate.