Leaf litter selection by detritivore and geophagous earthworms

Summary Litterbag experiments with 10 different kinds of leaf litter showed that detritivore (Lumbricus species) and geophagous (Aporrectodea species) earthworms prefer certain litter types over others, since different numbers of worms were found below the litter after 50–52 days of exposure in a pasture. The detritivores preferred Fraxinus, Tilia, and predecomposed Ulmus and Fagus litter to Fagus litter and paper, while geophages preferred Tilia litter to Alnus and Ulmus litter, so that the two groups of earthworms showed different preferences. The detritivores seemed to be more selective than the geophages. The palatability of the litter was examined in relation to the C: N ratio, the lignin concentration and the initial and final polyphenol concentration. The numbers of detritivores were significantly correlated with the C: N ratio and the final polyphenol concentration, so that selection of litter seems to be related to palatability. The numbers of geophages were not significantly correlated with any of the parameters for palatability. The disappearance of litter after 50–52 days appeared to be due to detritivore activity, since the numbers found below the litter were positively and significantly correlated with the litter disappearance. There was no significant correlation with geophage activity. This indicates that detritivores use litter as food, and therefore influence the composition of the litter layer.     

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).     

Stabilisation of soil organic matter in invertebrate faecal pellets through leaf litter grazing

Soils were amended with either leaf litter or faeces from pill millipedes fed on the leaf litter, then incubated at 20 °C for 130 days whilst monitoring the respiration rates. Significantly more CO₂ was respired from soil containing leaf litter than that amended with an equivalent weight of faecal matter, whilst the unamended soil exhibited a respiration rate similar to soil amended with faecal material. Consideration of these findings with recently observed differences in biochemical compositions of litter and faeces suggests that processing of plant litter by detritivores leads to more stabilised forms of organic matter by removal of biochemical components essential to the nutrient requirements of the invertebrate and the soil microbial biomass.     

Decomposer animals (Lumbricidae, Collembola) and organic matter distribution affect the performance of Lolium perenne (Poaceae) and Trifolium repens (Fabaceae)

Decomposer animals stimulate plant growth by indirect effects such as increasing nutrient availability or by modifying microbial communities in the rhizosphere. In grasslands, the spatial distribution of organic matter (OM) rich in nutrients depends on agricultural practice and the bioturbation activities of large detritivores, such as earthworms. We hypothesized that plants of different functional groups with contrasting nutrient uptake and resource allocation strategies differentially benefit from sites in soil with OM accumulation and the presence of decomposer animals. In a greenhouse experiment we investigated effects of spatial distribution of ¹⁵N-labelled grass litter, earthworms and collembola on a simple grassland community consisting of Lolium perenne (grass) and Trifolium repens (legume). Litter aggregates (compared to homogeneous litter distribution) increased total shoot biomass, root biomass and ¹⁵N uptake by the plants. Earthworms and collembola did not affect total N uptake of T. repens; however, the presence of both increased ¹⁵N uptake by T. repens and L. perenne. Earthworms increased shoot biomass of T. repens 1.11-fold and that of L. perenne 2.50 fold. Biomass of L. perenne was at a maximum in the presence of earthworms, collembola and with litter concentrated in a single aggregate. Shoot biomass of T. repens increased in the presence of collembola, with L. perenne generally responding opposingly. The results indicate that the composition of the decomposer community and the distribution of OM in soil affect plant competition and therefore plant community composition.     

Influence of local illumination and plant composition on the spatial and seasonal distribution of litter-dwelling arthropods in a tropical rainforest

Using pitfall traps, we evaluated the spatial and seasonal variance in arthropod abundance, species richness, higher taxonomic and species composition, and guild structure within the ground litter of seven sites in a relatively undisturbed rainforest in Panama. We examined each of these five arthropod-dependent variables at two spatial scales (a few meters and a few hundred meters) and one temporal scale (a few months encompassing dry and wet periods), against environmental variables including local illumination and plant composition. Trap catches (9458 arthropods collected during 630 trap-days) were high compared to similar studies in temperate forests. We observed spatial and seasonal differences in abundance, species richness and composition of litter-dwelling arthropods. Often these differences appeared weakly related to geographical coordinates. They reflected forest structure (basal area) and local plant composition, and less so illumination patterns or seasonal changes in radiation. Seasonal variance was high and may relate to surrogate variables accounting for seasonal changes in litter moisture. The composition of higher taxa and species was often predicted by different independent variables at the three scales studied. Guild structure was difficult to predict. Our study lead us to expect that litter-dwelling arthropods may be more seasonal than soil microarthropods in tropical rainforests; and that tropical litter-dwelling arthropods may also be more spatially variable and seasonal than their temperate counterparts. We also recommend that conservation studies using pitfall traps in tropical rainforests should focus on: (1) taxonomic resolution to understand the functional complexity of soil organisms; (2) spatial replication to address subtle changes in plant composition throughout the study area; and (3) seasonal replicates to be commensurate with seasonal changes in litter moisture.     

Soil macroinvertebrate fauna of a Mediterranean arid system: Composition and temporal changes in the assemblage

Temporal variability is a key factor to understand the structure of belowground communities. Seasonal and annual variations are especially relevant in unpredictable desert ecosystems, where macroinvertebrates are poorly known, despite constituting an important group of soil organisms. In the present study, we analyse the composition and temporal (seasonal and annual) variations of soil macroinvertebrates in an arid area of southern Spain. During two years, macroinvertebrates were sampled in litter and belowground levels by means of soil cores. Results show that the assemblage was dominated by arthropods, especially Formicidae and Coleoptera. The assemblage differed between litter and belowground levels. In litter, detritivores dominated the community, while belowground fauna showed a similar proportion of detritivores and herbivores and a low percentage of predators. Litter and belowground assemblages showed seasonal variations in richness, abundance, biomass and composition, although variations were more marked in litter than belowground. Patterns of seasonal variation also differed between the two study years for both litter and belowground invertebrates. The seasonal and annual variability of the assemblage has potentially important implications for community dynamics in the study system, since the changes in species composition and trophic structure of soil invertebrate assemblages may affect species interactions and food web dynamics over time. Therefore, integrating temporal variability is likely to be crucial to understand soil community dynamics and food webs, especially in heterogeneous, variable systems as deserts.     

Soil microbial activity and litter turnover in native grazed and ungrazed rangelands in a semiarid ecosystem

Long-term overgrazing is known to influence soil microbiological properties and C sequestration in soil organic matter. However, much remains to be known concerning overgrazing impacts on soil microbial activity and litter turnover in heavily grazed rangelands of Central Iran. Aboveground litter decomposition of three dominant species (Agropyron intermedium, Hordeum bulbosum, and Juncus sp.) were studied using a litter bag experiment under field conditions in three range sites of Central Iran, a site with continuous grazing, a site ungrazed for 17 years with dominant woody species (80% cover), and a site ungrazed for 17 years with dominant pasture species (70% cover). Soil samples were taken from 0 to 30 cm depth and analyzed for their chemical and microbiological properties. Results demonstrate that soil organic C and total N contents and C/N ratios were similar for both ungrazed and grazed sites, while available P and K concentrations significantly decreased under grazed conditions. It was also evident that range grazing decreases soil respiration and microbial biomass C, suggesting a lower recent annual input of decomposable organic C. Nevertheless, grazing conditions had no significant effect on litter decomposition indicating soil microclimate is not affected by grazing animals in this ecosystem. It is concluded that overgrazing may presumably depress microbial activity through either reduced input of fresh plant residue into the surface soil or lack of living roots and exudates for stimulating microbial activity. This study also suggests that 17 years of livestock exclusion might be insufficient time for expected C accumulation in soil.     

Effects of seasonal grazing and precipitation regime on the soil macroinvertebrates of a Mediterranean old-field

Soil macroinvertebrate communities (SMC) are well known to influence major ecosystem processes, but relatively few investigations have examined the mechanisms and factors involved in SMC regulation. We conducted a factorial experiment with combinations of seasonal grazing by sheep and irrigation (simulating different precipitation regimes) to assess their effects on the SMC of a semiarid Mediterranean old-field. We also analyzed effects on plant species richness, total aboveground biomass, and litter. The data were collected in autumn and spring, the two favorable seasons for SMC and primary production in the region, and season was included as an additional random factor. Main results were: 1) Ungrazed plots accumulated more aboveground plant biomass and litter during spring, providing extra food for soil biota. However, grazing during autumn or spring did not affect SMC characteristics. 2) Reduction of inter-annual precipitation variability in autumn and spring increased the abundance of two decomposer taxa: Oligochaeta and Diplopoda. Additionally, if summer drought was reduced, plant species richness, litter and the abundance of Isopoda were increased. 3) Oligochaeta and Diplopoda increase their abundance in spring, particularly, the most abundant taxon (Oligochaeta). We conclude that inter- and intra-annual variability in precipitation is a key environmental factor for the decomposer soil fauna in Mediterranean ecosystems, modifying the physical characteristics of the soils (humidity, hardness, etc.), as well as affecting the amount or characteristics of plant biomass or litter. The respiration system of the macroinvertebrates (cutaneous, tracheal or branquial) and the capacity to migrate vertically into the soil may determine the decomposers' responses to precipitation.     

Microbial function in adjacent subtropical forest and agricultural soil

Soil microbial communities and their activities are altered by land use change; however impacts and extent of these alterations are often unclear. We investigated the functional responses of soil microbes in agricultural soil under sugarcane and corresponding native soil under Eucalyptus forest to additions of contrasting plant litter derived from soybean, sugarcane and Eucalyptus in a microcosm system, using a suite of complimentary techniques including enzyme assays and community level physiological profiles (CLPP). Initially agricultural soil had 50% less microbial biomass and lower enzyme activities than forest soil, but significantly higher nitrification rates. In response to litter addition, microbial biomass increased up to 11-fold in agricultural soil, but only 1.8-fold in forest soil, suggesting a prevalence of rapidly proliferating ‘r’ and slower growing ‘K’ strategists in the respective soils. Litter-driven change in microbial biomass and activities were short lived, largely returning to pre-litter addition levels by day 150. Decomposition rates of sugarcane and soybean litter as estimated via CO₂ production were lower in agricultural than in forest soil, but decomposition of more recalcitrant Eucalyptus litter was similar in both soils, contradicting the notion that microbial communities specialise in decomposing litter of the dominant local plant species. Enzyme activities and community level physiological profiles (CLPP) were closely correlated to microbial biomass and overall CO₂ production in the agricultural soil but not the forest soil, suggesting contrasting relationships between microbial population dynamics and activity in the two soils. Activities of enzymes that break down complex biopolymers, such as protease, cellulase and phenol oxidase were similar or higher in the agricultural soil, which suggests that the production of extracellular biopolymer-degrading enzymes was not a factor limiting litter decomposition. Enzyme and CLPP analyses produced contrasting profiles of microbial activity in the two soils; however the combination of both analyses offers additional insights into the changes in microbial function and community dynamics that occur after conversion of forest to agricultural land.     

Intermediate tidal stress promotes the detritivore-mediated decomposition of Spartina litter

Saltmarshes, functionally important habitats in the marine–terrestrial ecotone that are regularly affected by tidal inundation, are mainly detritus-based in terms of fluxes of nutrients and energy. With respect to the mediating influence of saltmarsh detritivores on microbial colonisation of detritus and on decomposition processes, we tested whether the “intermediate disturbance hypothesis” (IDH) is also applicable to the effects of stress in this stressful environment. Decomposition experiments with litter of the cordgrass, Spartina anglica, and with terrestrial [Porcellio scaber (Isopoda)] and marine/semi-terrestrial [Orchestia gammarellus (Amphipoda)] detritivores as well as animal-free controls were carried out in an artificial saltmarsh system. Different daily flooding regimes served as experimental levels of stress. Both litter mass loss and microbial respiration were mostly higher under aquatic than under terrestrial conditions, no matter whether detritivores were present or not. Considering the intertidal zone, low to intermediate daily inundation rates resulted in increased microbial respiration and an increased influence of detritivores on litter mass loss in early stages of cordgrass decomposition with high rates of detritus mass loss, and intermediate tidal stress led to higher microbial cell counts throughout the entire experiment. Summarised over 3 months, regression analyses suggested that microbial activity and detritus mass loss show a trend towards highest values at low inundation rates and under permanent inundation, although microbial density was higher under longer daily inundation. Access to detritus by detritivores enhanced both litter mass loss and microbial respiration, especially in later decomposition stages, whereas microbial density was reduced by detritivores. In conclusion, we predict that overall the decomposition of cordgrass detritus in saltmarshes is promoted in the intermediate to high area of the intertidal zone with daily inundation of ca. 4–10 hours where both marine and terrestrial detritivores have access to promote decomposition processes through feeding and mediating microbial activity.     

Soil arthropods beneficially rather than detrimentally impact plant performance in experimental grassland systems of different diversity

Impacts of belowground insecticide application on plant performance and changes in plant community structure almost uniformly have been ascribed to reduced belowground herbivory, although recent studies reported distinct side effects on detritivore soil animals, particularly on Collembola. Consequently, it remains controversial if the resulting soil feedbacks on plants are due to alterations in arthropod herbivory or to changes in the activity of detritivores. We investigated the impacts of the application of a commonly used belowground insecticide (chlorpyrifos) on soil animals and soil feedbacks on model plant species representing two main plant functional groups of grassland communities, the grass Lolium perenne and the forb Centaurea jacea.Insecticide application decreased soil insect herbivore densities considerably. However, also Collembola densities and diversity decreased markedly due to insecticide application and this was most pronounced in Entomobryidae, Isotomidae, Hypogastruridae, and Sminthuridae. While densities of other detritivore taxa were not affected or even increased (Oribatida) in insecticide subplots, that of predators mostly decreased.Both model plant species built considerably more biomass in control subplots than in insecticide subplots irrespective of characteristics of the resident plant community. This suggests that soil feedbacks on plants were not due to belowground herbivory and highlights the significance of alternative mechanisms responsible for insecticide-mediated soil feedbacks on plants. The deterioration of model plant species’ performances in insecticide subplots most likely was due to decreased densities of Collembola resulting in the deceleration of nutrient cycling and plant nutrition. The results suggest that it is oversimplistic to only ascribe insecticide-mediated soil feedbacks on plants to belowground herbivores. The results further indicate that in the present study the impact of arthropod detritivores on plant productivity was more important than that of belowground herbivores. This emphasizes that plant-soil arthropod interactions in grassland might be based on both facilitative and antagonistic interrelationships.     

The fate of condensed tannins during litter consumption by soil animals

Condensed tannins (CT) can strongly affect litter decomposition, but their fate during the decomposition process, in particular as influenced by detritivore consumption, is not well understood. We tested the hypothesis that litter CT are reduced by the gut passage of two functionally distinct detritivores of Mediterranean forests, the millipede Glomeris marginata, and the land snail Pomatias elegans, as a fixed proportion of initial litter CT, but more so in Pomatias since snails are known to have a more efficient enzymatic capacity. Contrary to our hypothesis, both detritivore species reduced litter CT to near zero in their faecal pellets irrespective of the wide range in initial leaf litter CT concentrations of 9-188 mg g⁻¹ d m among three Mediterranean tree species (Pistacia terebinthus, Quercus ilex, Alnus glutinosa) and different decomposition stages of their litter. The almost complete disappearance of CT even from some litter types highly concentrated in CT, due to either degradation by gut microorganism or complexation of CT into insoluble high molecular weight structures, suggests a high “de-tanning” efficiency across functionally distinct detritivore species. The transformation of CT-rich litter into virtually CT-free faecal pellets by detritivores might be highly relevant for the subsequent decomposition process in ecosystems with a high macrofauna abundance and CT-rich plant species such as Mediterranean forests.     

Leaf litter decomposition and soil microarthropods affected by sulphur dioxide fallout

A field experiment was set up in northeastern Italy to investigate the effects of sulphur dioxide fallout on leaf litter decomposition rates and soil microarthropods. The pollution fallout, which affected part of the Po River delta, originated from the activity of an oil-fired power plant located at Isola Camerini (Porto Tolle, Rovigo). Four sampling sites, exposed to different amounts of pollutant fallout, were selected along the river bank. Two sites, located 1·5 km from the power-plant stack, received minimum sulphur dioxide fallout and were used as controls; two high-deposition sites were about 13 km away from the power-plant settlement. Soil cores were taken to compare sulphur concentrations in the upper layers of the sampling site soil profiles. Litter bags filled with plant material of various types, laid down over two consecutive sampling periods, were used to study leaf litter decomposition and sulphur accumulation on plant tissues. Microarthropods were extracted from the litter bags by means of a modified Tullgren apparatus. Soil chemical analysis showed the highest sulphur concentrations at the high pollutant deposition sites along the plume path. Litter bag dry weight loss over time was reduced by sulphur accumulation in plant tissues. Sulphur accumulation in litter bags gave an indirect measure of the differential pollutant deposition over the land. High-deposition sampling sites showed a significant reduction in the total number of some decomposers. Collembola, in particular, appeared to be a robust bio-indicator of pollutant fallout. Conclusions were drawn about the possible detrimental effects of sulphur compounds on soil and leaf chemistry, litter decomposition and microarthropod decomposer populations.© 1997 John Wiley & Sons, Ltd.     

Organic By-Product Effects on Soil Chemical Properties And Microbial Communities

Soil management practices that contribute to increased soil productivity and longterm sustainable agricultural production have been neglected over the last four decades. The need to increase soil productivity led to the evaluation of a system of disposing of large quantities of organic by-products and poultry litter on agricultural land. Our objectives were to evaluate the effects of applying noncomposted municipal solid waste (MSW), amended with either poultry litter (PL) or NH₄NO₃ to adjust C:N ratios in the soil surface in either the spring or fall. Changes in soil chemical properties, bacteria population shifts, changes in species richness and evenness of indigenous soil bacteria, and response by cotton (Gossypium hirsutum L.) were evaluated. Soil P, K, Ca, and Mg were increased in the surface 0–15 cm by a factor of three or four times by application of organic by-products. After two annual applications, soil Cu increased slightly, Zn doubled, Co and Cr decreased, while Pb increased by a factor of two. Soil organic matter content increased on average by 89 percent for treatments containing newsprint, yard trimmings, and cotton gin trash. Newsprint plus NH₄NO₃ resulted in a shift to more Gram positive bacteria, while newsprint plus poultry litter resulted in a shift to more Gram negative bacteria. Both N sources resulted in a reduction in Bacillus sp. Shifts in the bacterial populations and changes in species richness (number of species detected) and evenness (relative abundance of each species) were induced by organic by-product additions. These shifts appear to be the result of increased substrate for C mineralization rather than any properties of biological control. Shifts in the microbial community structure towards Gram negative organisms may benefit plant growth and may be useful as an indicator of soil quality.     

Changes in the community structure and diversity of soil invertebrates across the Franz Josef Glacier chronosequence

Following the creation of new land surfaces, there is an initial build-up phase of ecosystem development, but after a prolonged absence of major disturbance a retrogressive (decline) phase often follows due to reduced nutrient availability over time. Although many studies have considered how the soil community changes during the build-up phase, the response of this community to the retrogressive phase is poorly known. We measured litter and soil communities of microfauna and macrofauna along the Franz Josef Glacier chronosequence in New Zealand that spans ca. 120,000 years, and includes well-established build-up and retrogressive stages. We aimed to assess whether the abundances, community structure and diversity of these groups show the same pattern across the sequence as that for vegetation. With regard to microfaunal abundances, litter-dwelling microbe-feeding nematodes were most abundant in the first stage of the chronosequence, but several other groups of microfauna in both the soil and litter increased sharply during the first few stages and declined sharply during the last (retrogressive) stages. The ratios of bacterial- to fungal-feeding nematodes in both soil and litter were lowest for the final stages of the chronosequence, and (in the case of soil) for some of the early stages, pointing to domination by the fungal-based energy channel at those stages for which soil organic matter content or quality were lowest. This is consistent with the fungal-based energy channel being better adapted than the bacterial-based channel for resource-poor conditions. The main groups of macroinvertebrates typically had their lowest abundances at the very early and late stages of the chronosequence, although the relative abundances of different taxa differed during the intermediate stages. Taxonomic diversity of nematodes and macroinvertebrates in both litter and soil varied strongly with chronosequence stage but differed among taxa; diversity of only one group (macroinvertebrates in litter) declined significantly during retrogression. Diversity of nematodes and macroinvertebrates along the sequence did not closely match tree diversity or soil chemical properties, but community composition of these groups was often related to tree community composition and ratios of soil C to N, C to P and N to P. Different groups of soil invertebrates show contrasting responses to chronosequence stage, probably because they differ in their relative response to bottom-up and top-down controls. However, the abundance of most groups increased during the build-up phase and declined during retrogression. As such, the build-up and decline phases observed for plant communities and ecosystem processes across long-term chronosequences also apply to soil communities, pointing to the importance of resource availability as a major driver of soil biota during long-term ecosystem change.     

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.     

Effects of Soil Development Time and Litter Quality on Soil Carbon Sequestration: Assessing Soil Carbon Saturation with a Field Transplant Experiment along a Post‐mining Chronosequence

In previous studies, the rate of soil carbon (C) sequestration decreased with increasing age of post‐mining soils. It was also shown to depend on plant biomass and earthworm bioturbation. Here, a soil transplant experiment was used to determine whether this decrease is caused by soil C saturation or other factors (such as bioturbation and litter input). Soils collected on 15‐, 25‐ and 50‐year‐old successional sites, dominated by willow (Salix caprea L.) and birch (Betula pendula Roth), and on a 50‐year‐old site reclaimed by the planting of alder (Alnus glutinosa L.) were placed in plastic boxes that were accessible to soil macrofauna. The boxes were buried in the 50‐year‐old reclaimed site and supplemented with either alder litter or successional (willow and birch) litter. Soil C content and soil C fractions (hot water C, particulate organic C, particulate organic C bound in aggregates and C bound to mineral soil) were studied. After 1 year, the C content increased by 2–5%, but there was no effect of soil source or litter type. For all C fractions, the relationships between change in C content and initial C content were described by bell‐shaped curves. Easily available C fractions were saturated earlier than more recalcitrant fractions. Despite these saturation tendencies in individual soil organic matter pools, the soil was evidently far from saturation after 50 years of soil development. The decrease in C sequestration with soil age previously observed for this soil was probably caused by a decrease in litter input rather than by C saturation. Copyright © 2016 John Wiley & Sons, Ltd.     

Composition of lipophilic compounds and carbohydrates in the accumulated plant litter and soil organic matter in boreal forests

Carbohydrates and lipophilic compounds constitute an important component of litter and soil organic matter in boreal forests, but are still poorly identified. We characterized needle litter and coarse tree litter (cones, seeds, bark and twigs) from coniferous trees (Pinus sylvestris L. and Picea abies Karst.), and moss litter (Pleurozium schreberi, Hylocomnium speldens), fermentation (F) and humus (H) layers in four boreal forest sites in Finland using a combination of sequential fractionation (non‐polar extractions, NPE; water‐soluble extractions, WSE; acid‐soluble fraction, AS) and detailed analyses on the soluble fractions using GC‐MS. Comparisons among the different layers of the soil organic horizon were used to assess which lipophilic compounds and carbohydrates increased in the F and H layers in proportion to their relative abundance in the litter layer and thus might have a large potential to accumulate in soil organic matter. Both concentrations and relative proportions of different compounds varied among the soil layers. Several of the fatty acids (FAs) found in the litter samples were absent in the F and the H layers. Needle and coarse tree litter contained a wide range of di‐ and triterpenes, but in the F and H layers oxidized forms of dehydroabietic acid and sterols were abundant. The large proportion of dehydroabietic acid in the lipophilic fraction in the H layer suggests that it may be poorly degradable by soil microorganisms, probably because of its anti‐microbial function in trees. The composition of the acid‐soluble fraction indicated that the proportion of cellulose in relation to hemicelluloses increased from the litter layer to the F and H layers. Put together, changes in the relative proportions of organic compounds in soluble fractions indicate that selective preservation of compounds, litter input by plant roots and microbial synthesis of compounds all contribute to the accumulation of aliphatic compounds in the H layer of boreal forests.     

Plant‐available N:P alters root litter N recycling in a Mediterranean tree–grass ecosystem

Background: Nitrogen deposition can cause an ecosystem‐level shift in available N (nitrogen) to P (phosphorus) availability. However, most plant N nutrition is from edaphic sources rather than deposition and in seasonally dry grassland systems, root litter is the predominant nutrient source. Aims: We were interested how litter turnover and altered nutrient recycling from dead biomass can compensate for these shifts in ecosystem stoichiometry. Methods: We studied a Mediterranean savanna amended with N or NP treatments three years prior. We measured root and plant‐available soil N:P stoichiometry in two micro‐habitats: open pasture and beneath oak canopies. ¹⁵N‐labelled root litter incubated in topsoils without litterbags was used to trace uptake of litter N by herbaceous strata roots. Results: Since fertilization, NP added sites have become relatively P enriched, resulting in lower N:P ratios in living roots than either when N was added alone or control sites. Total litter‐derived ¹⁵N uptake by roots was proportional to root ingrowth response but higher in the NP than N treatment, indicating a higher N demand when N and P were added together. We observed more ¹⁵N uptake by plants under tree canopies, indicating a tighter nutrient recycling loop in these micro‐habitats in contrast to treatment level ‘fertility' trends. Conclusions: Root stoichiometry responded to manipulated soil nutrient availability and N uptake was altered as plants attempted to compensate for nutrient availability imbalances, indicating that these ecosystem perturbations have long term effects on nutrient cycling which can propagate to whole system function. This was also related to functional community‐level adaptions between micro‐habitats with under canopy communities more able to take advantage of the litter nutrient source.     

Plant growth and microbial activity in a tropical soil amended with faecal pellets from millipedes and woodlice

The effect of soil fauna-mediated leaf litter (faecal pellets) versus mechanically fragmented (finely ground) leaf litter on biomass production of rice (Oryza sativa, var. Primavera) was assessed in pot tests. Rice seedlings were either grown in soil samples amended with faecal pellets of diplopods and isopods fed on leaf litter of a legume cover crop (Pueraria phaseoloides (Roxb.) Benth) and a peach palm (Bactris gasipaes) or in soil amended with finely ground leaf litter. The addition of faecal pellets caused a significant and dose-related increase in plant biomass compared to pure soil. Ground leaf litter induced a significantly smaller positive effect on plant biomass development with Pueraria litter > Bactris litter > mixed primary forest litter. In contrast, soil microbial biomass development during the 4 weeks plant test was higher in the soil amended with ground litter as compared to soil amended with feacal pellets. The results show a clear positive effect of the soil fauna on soil fertility and indicate differences in the availability of nutrients from the organic substrates to higher plants and soil microorganisms.