Stable isotopes revisited: Their use and limits for oribatid mite trophic ecology

In this review we summarize our knowledge of using stable isotopes (¹⁵N/¹⁴N, ¹³C/¹²C) to better understand the trophic ecology of oribatid mites. Our aims are (a) to recapitulate the history of stable isotope research in soil animals with a focus on oribatid mites, (b) to present new stable isotope data for oribatid mites and overview the current state of knowledge of oribatid mite trophic niche differentiation, (c) to compile problems and limitations of stable isotope based analyses of trophic relationships and (d) to suggest future challenges, questions and problems that may be solved using stable isotope analyses and other novel techniques for improving our understanding on the trophic ecology of soil invertebrates. We conclude that (1) in addition to ¹⁵N/¹⁴N ratios, ¹³C/¹²C ratios contribute to our understanding of the trophic ecology of oribatid mites, allowing, e.g. separation of lichen- and moss-feeding species, (2) there likely are many lichen but few moss feeding oribatid mite species, (3) oribatid mite species that are endophagous as juveniles are separated by their stable isotope signatures from all other oribatid mite species, (4) fungivorous oribatid mite species cannot be separated further, e.g. the fungal taxa they feed on cannot be delineated. A particular problem in using stable isotope data is the difficulty in determining signatures for basal food resources, since decomposing material, fungi and lichens comprise various components differing in stable isotope signatures; ¹³C/¹²C ratios and potentially other isotopes may help in identifying the role of these resources for decomposer animal nutrition.     

Compartmentalization of the soil animal food web as indicated by dual analysis of stable isotope ratios (N/N and C/C)

The soil animal food web has become a focus of recent ecological research but trophic relationships still remain enigmatic for many taxa. Analysis of stable isotope ratios of N and C provides a powerful tool for disentangling food web structure. In this study, animals, roots, soil and litter material from a temperate deciduous forest were analysed. The combined measurement of δ¹⁵N and δ¹³C provided insights into the compartmentalization of the soil animal food web. Leaf litter feeders were separated from animals relying mainly on recent belowground carbon resources and from animals feeding on older carbon. The trophic pathway of leaf litter-feeding species appears to be a dead end, presumably because leaf litter feeders (mainly diplopods and oribatid mites) are unavailable to predators due to large size and/or strong sclerotization. Endogeic earthworms that rely on older carbon also appear to exist in predator-free space. The data suggest that the largest trophic compartment constitutes of ectomycorrhizal feeders and their predators. Additionally, there is a smaller trophic compartment consisting of predators likely feeding on enchytraeids and potentially nematodes.     

Changes in the community composition and trophic structure of microarthropods in sporocarps of the wood decaying fungus Fomitopsis pinicola along an altitudinal gradient

Soil microarthropods colonize a wide range of habitats including microhabitats such as earthworm burrows, ant nests, tree trunks, moss mats and wood decaying fungi. While many of these microhabitats have been investigated intensively, the role of wood decaying fungi as a habitat and food resource for microarthropods found little attention. We investigated the density, community structure, reproductive mode and trophic structure of microarthropods, in particular oribatid mites, in the wood decaying fungus Fomitopsis pinicola (Schwarts: Fr) Karst. along an altitudinal gradient in Germany spanning from 350 m to 1160 m. Microarthropods were extracted from sporocarps, and stable isotope ratios (¹⁵N/¹⁴N; ¹³C/¹²C) of the fungus and the microarthropods were measured. Densities of most microarthropod taxa were highest at lower altitudes and decreased with increasing altitude. Oribatid mites were the dominant animal taxon. Their community structure gradually changed with altitude. Stable isotope ratios indicated that oribatid mite and other arthropod species occupy distinct trophic niches but most do not feed on F. pinicola. Notably, species of the same genus, e.g. Carabodes, occupied different trophic niches. Most oribatid mite species in F. pinicola reproduced sexually which is similar to the bark of trees but in contrast to the soil where most species reproduce via parthenogenesis. The findings indicate that (1) at high altitudes microarthropod density in fungal fruiting bodies is limited by low temperatures reducing animal metabolism and reproduction, and this also affects oribatid mite community structure, (2) despite the uniform habitat trophic niches of oribatid mite species differ and this also applies to morphologically similar species of the same genus, and (3) feeding on F. pinicola or associated resources facilitates the dominance of sexual reproducing species.     

Trophic structure of oribatid mite communities from six different oak forests (Quercus robur)

The degree of trophic plasticity in soil animals is intensely debated. We used stable isotope ratios (¹⁵N/¹⁴N, ¹³C/¹²C) of oribatid mite species from six oak (Quercus robur) forests to investigate (1) if trophic niches vary between forests and (2) the range of trophic levels spanned by oribatid mites. Using litter as baseline stable isotope signatures of most oribatid mite species differed between forests. Therefore, the stable isotope signatures were re-calibrated using stable isotope values of Platynothrus peltifer as primary decomposer species occurring in each of the six forests. Re-calibrated values of nine species (Cerachipteria jugata, Damaeus clavipes, Neotrichoppia variabilis, Oppia denticulata, Hermaniella dolosa, Steganacarus magnus, Ceratozetes peritus, Nanhermannia nana, Xenillus tegeocranus, Eremaeus cordiformis) differed significantly between forests indicating trophic plasticity in most of the studied oribatid mite species. Overall, calibrated stable isotope ratios spanned over 8.7 δ units for ¹⁵N and 5.9 δ units for ¹³C indicating that in forest ecosystems oribatid mite species span about three trophic levels.     

Feeding guilds in Collembola based on nitrogen stable isotope ratios

In soil a high number of species co-exist without extensive niche differentiation, which was assigned as ‘the enigma of soil animal species diversity’. In particular, the detritivores are regarded as food generalists. We have investigated nitrogen stable isotope ratios (¹⁵N/¹⁴N) of a major decomposer group, the Collembola, to evaluate trophic relationship and determine feeding guilds. Additionally, the δ¹⁵N values of potential food sources such as mosses, lichens and other plant derived material (bark, nuts, leaves) were analysed. The natural variation in nitrogen isotopes was assessed in 20 Collembola taxa from three deciduous forest stands. The δ¹⁵N signature formed a continuum from phycophages/herbivores to primary and secondary decomposers, reflecting a gradual shift from more detrital to more microbial diets. The δ¹⁵N gradient spanned over 9 δ units, which implies a wide range in food sources used. Assuming a shift in ¹⁵N of about 3 ‰ per trophic level, the results indicate a range of three trophic levels. These variations in ¹⁵N/¹⁴N ratios suggest that trophic niches of Collembola species differ and this likely contributes to Collembola species diversity.     

Molecular detection of nematode predation and scavenging in oribatid mites: Laboratory and field experiments

Recent stable isotope analyses indicate that a number of putative detritivorous soil microarthropods is not typical detritivores but rather live as predators or scavengers. Using molecular gut content analyses the present study investigates if nematodes indeed form part of the diet of oribatid mites. First, in a no-choice laboratory feeding experiment two nematode species (Phasmarhabditis hermaphrodita and Steinernema feltiae) were offered to eight species of oribatid mites and one gamasid mite. Second, after feeding for 4 and 48 h on each nematode species the detection time of prey DNA in the oribatid mite species Steganacarus magnus was investigated. Third, in a field experiment nematode prey (P. hermaphrodita and S. feltiae) in the diet of microarthropods was investigated distinguishing between scavenging and predation. In the no-choice laboratory experiment not only the gamasid mite but also several of the studied oribatid mite species consumed nematodes. After feeding on nematodes for 4 h prey DNA was detectable in S. magnus for only 4 h, but after feeding for 48 h prey DNA was detectable for 128 h, indicating that the duration of feeding on prey is an important determinant for prey DNA detection. The field experiment confirmed that oribatid mite species including Liacarus subterraneus, Platynothrus peltifer and S. magnus intensively prey on nematodes. Interestingly, DNA of dead P. hermaphrodita was detectable to a similar degree as that of living individuals indicating that scavenging is of significant importance in decomposer food webs. Results of our study indicate that predation and scavenging on nematodes by “detritivorous” microarthropods in soil food webs need to be reconsidered.     

Trophic stability of soil oribatid mites in the face of environmental change

A key issue in ecology is the degree to which trophic structure within communities responds to environmental change. Organisms with generalist diets are more flexible in their feeding habits than are specialists, and may be affected less in a changing environment. Soil fauna fulfill crucial ecosystem functions in terrestrial ecosystems and many are thought to have generalized diets. They may therefore be buffered from negative effects of environmental change. Here, we used ¹⁵N isotope analysis to study trophic differentiation among 91 species of oribatid mites and their responses to chronic atmospheric N deposition. Combining our own measurements with published data, we established that the trophic positions of mite species were remarkably stable within and among forests, as well as between ambient and experimental N deposition. Trophic stability indicates a higher than expected level of feeding specialization, which may foster diversity, but limit the ability to switch food resources in a changing environment.     

Oribatid mite and collembolan diversity, density and community structure in a moder beech forest (Fagus sylvatica): effects of mechanical perturbations

The effects of mechanical perturbations on two soil microarthropod communities (oribatid mites and collembolans) were investigated in a moder beech forest on sandstone. We disturbed the soil matrix by sieving and mixing the litter and soil of the moder profile. The top litter layer (L material) and the deep mineral soil (Bv) remained intact. Three amounts of disturbance were established: a single perturbation, perturbations once every 2 months (60 d) and once every 2 weeks (14 d). Densities of most groups of oribatid mites and all groups of collembolans declined in the disturbance treatments. In most cases, densities were lowest in the strong perturbation treatment (14 d). Desmonomata were the only group of oribatid mites that benefited from intermediate amounts of disturbance but not from the strongest disturbance. Also, disturbances reduced diversity of oribatid mites and collembolans. According to their sensitivity to disturbances oribatid mites ranked Poronota=Enarthronota=Suctobelbidae (the most sensitive)>Oppiidae>Tectocepheus>Desmonomata. The ranking of collembolans was Folsomia (the most sensitive)>Hypogastruridae/Neanuridae>Onychiuridae=Isotomidae>Entomobryidae. Generally, tolerance of disturbance was wider for oribatid mites than for collembolans. The results indicate that disturbances such as mixing of litter and soil and comminution of litter material strongly affect the density and diversity of soil microarthropods. However, they also indicate that the soil microarthropod community is resistant to weaker disturbances. In the field, mechanical disturbances are often caused by burrowing of earthworms. Our results suggest that the high density of microarthropods in moder soils may be due to the low intensities of mechanical disturbances by earthworms.     

Dual stable isotope analysis (δC and δN) of soil invertebrates and their food sources

More research is required to validate and refine natural abundance stable isotope ratio techniques as a tool for the investigation of the feeding ecology of soil animals and trophic relations in soil food webs. Isotope ratios of C (δ¹³C) and N (δ¹⁵N) were measured in herbivorous and detritivorous invertebrate groups, namely lumbricid earthworms (7 species), enchytraeid worms (3 species), slugs (3 taxa), and their potential food sources in an arable system. Intrapopulation δ¹⁵N variation in the slug Deroceras reticulatum (n=52) was large (range 4.2‰), possibly reflecting spatial variability in the food sources. Significant correlations between C:N ratios and isotope ratios in earthworms suggest that factors other than feeding may influence isotopic patterns. One enchytraeid species, Enchytraeus buchholzi, was enriched in ¹³C and strongly depleted in ¹⁵N compared to all other groups. Invertebrates formed a continuum when considered in relation to C and N separately, but fell into two distinct groups on the basis of combined C and N isotope ratios. The less enriched group represents herbivorous and litter-feeding species, while the more enriched group represents soil feeders. It is concluded that δ¹³C measurements could provide a means of assigning separate baseline δ¹⁵N values to primary and secondary decomposers, which in turn could improve the inference of higher trophic levels, omnivory and intraguild predation.     

The fate of glucose,a low molecular weight compound of root exudates,in the belowground foodweb of forests and pastures

Increasing evidence suggests that much of belowground, heterotrophic activity in terrestrial ecosystems is fueled by inputs of low molecular weight carbon compounds (LMWCCs). Root exudation (rhizodeposition) is a primary source of these inputs and will likely increase with rising atmospheric CO₂. Yet the fates of these compounds belowground, as well as the environmental factors that influence them, are relatively unexplored. Using stable isotopes we track the fate of one dominant LMWCC, glucose, in three pasture and three forest sites located in South Carolina, USA. We resolve glucose-derived C in CO₂, dissolved and soil organic C (DOC, SOC), microbial biomass, and microarthropods (Collembola, oribatid and mesostigmatid mites). After 72 h, the greatest proportions of glucose-C are in microbial biomass and SOC, followed by CO₂, DOC, and microarthropods. Within this short time frame, glucose-C propagates through the foodweb to the highest trophic level, predatory mesostigmatid mites. The biomass of these predators is the only variable that explains the relative partitioning across sites of glucose-C, with higher biomass associated with reduced partitioning of glucose-C to respiration and hence greater retention belowground. Our results suggest that LMWCCs entering belowground systems may propagate through soil foodwebs rapidly, and that their partitioning belowground may potentially be determined by higher trophic levels.     

Origin of the nitrogen assimilated by soil fauna living in decomposing beech litter

We investigated the nitrogen source for main taxa of soil fauna in two beech forests of contrasted humus type using ¹⁵N-labelled beech litter and ¹⁵N analysis of soil fauna. ¹⁵N-labelled beech litter was deposited on the topsoil in December 2000 in four stands of different ages at Leinefelde (Germany) with mull humus and in one mature stand at Sorø (Denmark) with moder humus. The fate of the tracer isotope was measured in litter and soil, as well as in the soil fauna, and for each taxa, we calculated the proportion of N in the animal derived from the labelled substrate. Of the original N contained in the litter, 20-41% was lost after 9 months at Leinefelde, and only 10% at Sorø. This loss was counterbalanced by the incorporation of 24-31% external N at Leinefelde, and 31% at Sorø, partly originating from fungal colonisation of the added litter. The proportion of N assimilated from the labelled litter by the different soil animals varied in relation to their mobility and feeding preferences. Large and mobile soil animals, especially predators, derived on average less ¹⁵N because they were also able to feed outside the labelled litter boxes. Detritivores assimilated at most 15% of their nitrogen content at Leinefelde and 11% at Sorø from the decomposing labelled litter. The most labelled taxa at Leinefelde were small fungivorous and coprophagous species, mainly isotomid Collembola such as Isotomiella and Folsomia. At Sorø, best labelled taxa were saprophagous species such as Enchytraeidae, Glomeridae and Phthiracaroidea. These low rates of ¹⁵N assimilation indicate that fresh litter is not directly the main N source for soil animals. The results obtained suggest that soil fauna fed preferentially upon microorganisms colonising the litter at Leinefelde (mull) and from litter itself at Sorø (moder).     

Estimating N2 fixation by Sesbania rostrata and S. cannabina (syn. S. aculeata) in lowland rice soil by the 15N dilution method

Summary A field experiment in concrete-based plots was conducted to estimate the contribution of N derived from air (Ndfa) or biological N₂ fixation in Sesbania rostrata and S. cannabina (syn. S. aculeata), using various references, by the ¹⁵N dilution method. The two Sesbania species as N₂-fixing reference plants and four aquatic weed species as non-N₂-fixing references were grown for 65 days after sowing in two consecutive crops, in the dry and the wet seasons, under flooded conditions. Soil previously labeled with ¹⁵N at 0.26 atom % ¹⁵N excess in mineralizable N was further labeled by ammonium sulfate with 3 and 6 atom % ¹⁵N excess. The results showed that ¹⁵N enrichment of soil NH ₄ ⁺ -N dropped exponentially in the first crop to half the original level in 50 days while in the second crop, it declined gradually to half the level in 130 days. The decline in ¹⁵N enrichment, in both N₂-fixing and non-fixing species, was also steeper in the first crop than in the second crop. Variations in ¹⁵N enrichment among non-fixing species were smaller in the second crop. The ratio of the uptake of soil N to that of fertilizer N in N₂-fixing and non-fixing species was estimated by the technique of varying the ¹⁵N level. In the second crop, this ratio in non-fixing species was higher than that in N₂-fixing species. Comparable estimates of % Ndfa were obtained by using ¹⁵N enrichment of various non-fixing species. There was also good agreement between the estimates obtained by using ¹⁵N enrichment of non-fixing species and those by using soil NH ₄ ⁺ -N, particularly in the second crop. By 25 days after sowing, the first crop of both Sesbania spp. had obtained 50% of total N from the atmosphere and the second crop had obtained 75%. The contribution from air increased with the age of the plant and ranged from 70% to 95% in 45–55 days. S. rostrata fixed substantially higher amounts of N₂ due to its higher biomass production compared with S. cannabina. Mathematical considerations in applying the ¹⁵N dilution method are discussed with reference to these results.     

Evaluating N/N and C/C isotope ratio analysis to investigate trophic relationships of elaterid larvae (Coleoptera: Elateridae)

Carbon and nitrogen isotope ratios in consumer tissues can be used to analyse the diet and trophic level of soil animals. However, life history traits may significantly influence stable isotope patterns. We evaluated in a series of experiments how stable isotope ratios of carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) at natural abundance can be used to study the diet and trophic position of long-lived macro-invertebrates, elaterid larvae, which are major below-ground herbivores. Small, but significant differences in δ¹³C signatures were found between the larvaes’ anterior and posterior body segments, whereas exuvia reflected the body's overall isotopic composition. The species-specific trophic shift (±SE) in δ¹⁵N for Agriotes obscurus and Agriotes sputator (1.62±0.24‰ and 1.08±0.27‰, respectively) was significantly lower than “mean enrichment estimates” reported in the literature, showing the limited applicability of such generalised estimates in studies of invertebrate trophic ecology. To avoid false-positive assignments to two trophic levels due to variation in δ¹⁵N values, a minimum sample size of three and five individuals for A. obscurus and A. sputator, respectively, was needed to reduce this risk to below α=5%. Keeping elaterid larvae for up to 128 days without food did not affect their isotopic signatures, in contrast to previous studies on starving animals. Switching wireworms to isotopically different diets induced changes in their isotopic signatures within 2 weeks. Changes, however, were significant only when the isotopic difference between diets was large. We conclude that experimental studies evaluating how specific life history traits affect stable isotope signatures in consumers have to precede any interpretation of stable isotope data gathered in the field.     

Estimates of Gross Transformation Rates of Dairy Manure N Using 15N Pool Dilution

Abstract

Most measurements of dairy manure nitrogen (N) availability depend on net changes in soil inorganic N concentration over time, which overlooks the cycling of manure N in the soil. Gross transformations of manure N, including mineralization (m), immobilization (i), and nitrification (n), can be quantified using ¹⁵N pool dilution methods. This research measures gross m, n, and i resulting from application of four freeze‐dried dairy manures that had distinctly different patterns of N availability. A sandy loam soil (coarse‐loamy, mixed, frigid Typic Haplorthod) was amended with four different freeze‐dried dairy manures and incubated at 25°C with optimal soil water content. The dilution of ¹⁵ammonium (NH4⁺) during a 48‐h interval (7–9 d and 56–58 d after manure application) was used to estimate m, whereas the dilution of ¹⁵nitrate (NO₃ ^?) was used to estimate n. Gross immobilization was calculated as gross minus net mineralization. Gross mineralization in the unamended soil was similar at 7‐ to 9‐d and 56‐ to 58‐d intervals and was significantly increased by the application of manures. For both amended and unamended soil, m was much greater (i.e., three‐ to nine‐fold) than estimated net mineralization, illustrating the degree to which manure N can be cycled in soil. At the early interval, both m and i were directly related to the manure C input, demonstrating the linkage between substrate C availability and N utilization by soil microbes. This research clearly shows that the application of dairy manures stimulates gross N transformation rates in the soil, improving our understanding of the impact of manure application on soil N cycling.     

Niche differentiation among neotropical soldierless soil-feeding termites revealed by stable isotope ratios

Termites represent one of the most abundant belowground animal taxa in tropical rainforests, where their species richness is much higher than in any other ecosystem. This high diversity in soil ecosystems is however difficult to explain by classical Hutchinsonian niche theory, as there is little evidence for spatial or temporal separation between species. Using δ¹³C and δ¹⁵N isotopic ratios, we tested if resource partitioning along the humification gradient occurs in neotropical soldierless termites of the Anoplotermes-group. Two distinct sites were investigated to check if interspecific differences are transposable between sites. Significant differences in δ¹⁵N were found between species of the Anoplotermes-group. Although some species displayed higher intersite δ¹⁵N variation than others, species-average δ¹⁵N values for both sites were highly correlated, showing that sympatric soldierless soil-feeding termites feed on distinct components of the soil. Our data also suggest that some species are more likely to shift along this gradient than others, in response to overall habitat conditions or to the presence of competitors. Feeding niche differentiation can therefore account for the high species richness and diversity of soldierless soil-feeding termites in neotropical rainforests.     

Effects of food quality, starvation and life stage on stable isotope fractionation in Collembola

Naturally occurring stable isotopes of carbon and nitrogen are powerful tools to investigate food webs, where the ratio of ¹⁵N/¹⁴N is used to assign trophic levels and of ¹³C/¹²C to determine the food source. A shift in δ¹⁵N value of 3‰ is generally suggested as mean difference between two trophic levels, whereas the carbon isotope composition of a consumer is assumed to reflect the signal of its diet. This study investigates the effects of food quality, starvation and life stage on the stable isotope fractionation in fungal feeding Collembola. The fractionation of nitrogen was strongly affected by food quality, i.e. the C/N ratio of the fungal diet. Collembola showed enrichment in the heavier isotope with increasing N concentration of the food source. Δ¹⁵N varied between 2.4‰, which assigns a shift in one trophic level, and 6.3‰, suggesting a shift in two trophic levels. Starvation up to 4 weeks resulted in an increase in the total δ¹⁵N value from 2.8‰ to 4.0‰. Different life stages significantly affected the isotope discrimination by Collembola with juveniles showing a stronger enrichment (Δ¹⁵N=4.9‰) compared to adults (Δ¹⁵N=3.5‰). Δ¹³C varied between −2.1‰ and −3.3‰ depending on the food quality, mainly due to compensational feeding on low quality diet. During starvation δ¹³C value decreased by 1.1‰, whereas the life stage of Collembola had no significant effect on isotopic ratios. The results indicate that the food resource and the physiological status of the consumer have important impact on stable isotope discrimination. They may cause differences in fractionation rate comparable to trophic level shifts, a fact to consider when analysing food web structure.     

Tree identity surpasses tree diversity in affecting the community structure of oribatid mites (Oribatida) of deciduous temperate forests

The role of tree diversity and identity as determinants of soil animal community structure is little understood. In a mature deciduous forest dominated by beech we identified clusters of one, two and three tree species of beech, ash and lime allowing to investigate the role of tree species diversity and identity on the density and community structure of oribatid mites. To relate oribatid mite community structure to environmental factors we measured leaf litter input, fine root biomass, mass of organic layers, topsoil pH and C and N content. We expected oribatid mite density to increase with increasing tree diversity, but we expected the effects of tree species identity to override effects of tree diversity. Further, we hypothesized the density of oribatid mites to be reduced by the presence of beech but increased by the presence of lime and ash. As expected tree diversity little affected oribatid mite communities, whereas tree species identity strongly altered density and community structure of oribatid mites. However, in contrast to our expectations the density of oribatid mites was highest in presence of beech indicating that many oribatid mite species benefit from the presence of recalcitrant litter forming thick organic layers. Especially Oppioidea benefited from the presence of beech presumably due to an increased availability of food resources such as fungi and nematodes. Lower density of oribatid mites in monospecific clusters of lime and ash suggests that oribatid mites did not benefit from high quality litter of these species. Notably, large and strongly sclerotized oribatid mite species, such as Steganacarus magnus and Chamobates voigtsi, benefited from the presence of ash and lime. Presumably, these large species better resist harsh microclimatic conditions in shallow organic layers.     

Effects of forest stand type on oribatid mite (Acari: Oribatida) assemblages in a southwestern Quebec forest

Despite the ubiquity of oribatid mites in soil and litter systems, and their importance in decomposition and nutrient cycling processes, little is known of the factors underlying the composition of their assemblages. Our objective was to address this by determining how oribatid assemblage composition changes by forest stand type. This work was done in and near a hardwood forest in southwestern Quebec, Canada. We sampled mites by collecting 1 L of litter and 170 cm³ of soil from four sites in each of four distinct habitat types: American beech stands, sugar maple stands, mixed deciduous stands and mixed conifer plantations. Samples were collected in July and September 2005, and June 2006, and over 6500 oribatid mites were collected and identified to species. Abundance and species richness differed between forest types: for abundance conifer>beech>maple>mixed deciduous while for species richness beech and conifer>maple>mixed deciduous. Ordination analyses revealed that conifer plantations and beech stands supported distinct assemblages, while there were some overlap in the assemblages found in maple stands and mixed deciduous stands. These data support the importance of aboveground plant communities in affecting the composition of oribatid assemblages even at local scales and provide insight into additional impacts that may be caused by shifts in plant species ranges due to global changes.     

Rhizosphere activity, grass species and N availability effects on the soil C and N cycles

We examined whether grass species and soil nitrogen (N) availability could enhance Carbon (C) and N turnover during root litter decay in grassland. Three species with increasing competitiveness (Festuca ovina, Dactylis glomerata and Lolium perenne) were grown at two N fertiliser levels in an undisturbed grassland soil, in which soil organic fractions derived for the last 9 years from Lolium root litter which was ¹³C-depleted. During the subsequent experimental year, the C turnover was calculated using the respective δ¹³C values of the old and new C in the root phytomass, in two Particulate Organic Matter (POM) fractions above 200 μm and in the lightest part of the aggregated soil fraction between 50 and 200 μm. Soil N availability was monitored during the regrowth periods with ion exchange resins (IER). The C decay rates of each particle size fraction were calculated with a simple mechanistic model of C dynamics. The N mineralisation immobilisation turnover (MIT) was characterised by dilution of ¹⁵N-labelled fertiliser in the N harvestThe C:N ratio and the residence time of C in the fractions decreased with particle size. The presence of a grass rhizosphere increased the decay rate of old C. Accumulation of new C in particle size fractions increased with species competitiveness and with N supply. Species competitiveness increased C turnover in the aggregated fraction, as a result of greater accumulation of new C and faster decay of old C. Fertiliser N increased N turnover and C mineralisation in the SOM. Species competitiveness decreased soil -N exchanged with the IER and increased dissolved organic C (DOC) content. The nature of the current rhizosphere is thus an important factor driving C and N transformations of the old root litter, in relation with grass species strategy. Plant competitiveness may stimulate the C and N turnover in the more evolved SOM fractions in a similar way to the mineral N supply.     

A 15N tracing technique-based analysis of the fate of fertilizer N: a 4-year case study in eastern China

The fate of fertilizer N applied with different irrigation amounts in tobacco fields was quantitatively studied by applying ¹⁵N double-labelled NH₄NO₃ in lysimeters. The ¹⁵N (fertilizer N originating from the fertilizer applied in 2011) in tobacco plants, ¹⁵N in soils and ¹⁵N loss were observed continuously from 2011 to 2014. The results showed that 21.6% of ¹⁵N was utilized by tobacco plants, 72.1% remained in the 0–60 cm soil layer and 6.3% was lost from the soil–plant system after the first season’s harvest (2011) of flue-cured tobacco. During the four seasons from 2011 to 2014, cumulative utilization of ¹⁵N by tobacco plants was 34.3%, while 54.2% remained in the 0–60 cm soil layer, and 11.5% was lost via mechanisms such as leaching and volatilization. The fate of ¹⁵N in terms of accumulation in plants and soils or losses from the soil–plant system from 2012 to 2014 was greatly affected by the fertilizer and irrigation management strategies in 2011. The results of this investigation suggest that the major amount of fertilizer N applied during the first season remains available in the soil for utilization by tobacco plants after 4 years.