Differential effects of lichens and mosses on soil enzyme activity and litter decomposition

In the New Jersey Pinelands, canopy gaps in the pine-dominated forest support patches of lichens, mosses, and caespitose grasses. We tested the hypotheses that non-vascular plants and lichens can affect nutrient cycling processes and that mosses and lichens would differ from each other. We predicted that (1) lichen tissues would decompose more slowly than pine or moss tissues, (2) all plant materials would decompose more slowly beneath lichens than beneath mosses, and (3) soil enzyme activities would be higher under lichens than under mosses or grasses, reflecting greater nutrient limitation. We compared rates of decomposition of the litter of Pinus rigida and moss and lichen tissues, and measured soil enzyme activities responsible for nutrient mineralization from litter (acid and alkaline phosphatases, chitinase, β-glucosidase, aminopeptidase, and phenol oxidase) under three types of groundcover (lichens, mosses, and grasses) and unvegetated soil at two sites. While groundcover affected enzyme activities, the patterns of enzyme activities differed markedly between the two sites. In general, the enzyme activities were uniformly low. Decomposition rates were more strongly affected by the groundcover than by litter materials. While all litters tended to decompose more slowly under lichens than under mosses, supporting one of our initial hypotheses, the rates of decomposition were markedly different between the two sites. These results suggest that while mosses and lichens create patches of different soil function in both sites, the differences between the sites in unknown factors cause the enzyme activities and decomposition rates to differ.     

Moss and lichen decomposition in old-growth and harvested high-boreal forests estimated using the litterbag and minicontainer methods

Bryophytes and lichen are important components of many boreal forest ecosystems, making the quantification of moss and lichen decomposition rates critical to understanding the C cycle of these forests. Cryptogam decomposition has been predominantly studied in wetlands, while few studies exist for forest-habitat mosses and even fewer for foliose ground lichen. We used a) the litterbag and b) the minicontainer (MC) method to quantify the decomposition rates of i) feathermoss, ii) forest peatmoss, iii) foliose ground lichen, and iv) alder leaves (reference litter) in cool, wet high-boreal Labrador black spruce forests. A total of 1560 litter samples (360 litterbags, 1200 MCs) were incubated for two years in six forest stands of different disturbance origin: three old-growth stands of wildfire origin, and three recently clearcut stands. Litter samples were retrieved after 6.5, 13, 47, 57 and 109 weeks of field incubation, and analysed for mass loss, C, N, nutrient, and fibre content.While clearcut harvesting had no significant effect on litter decomposition at all, decomposition rates significantly differed between litter types, with residual mass increasing in the order alder ≤ lichen < feathermoss ≤ peatmoss. Compared to wetlands, forest moss litter was more labile in the studied forest types, with lichen producing especially fast-decomposing litter. Litter type was a better predictor of decomposition rates than individual chemical parameters, indicating that, even in extreme climates like in Labrador, substrate quality is more important in determining decomposition rates than environmental factors. For all litter types, decomposition models accounting for the seasonality of decay dynamics performed better than models assuming constant or continuously decreasing decay rates. Compared to the litter bag method, which yielded decomposition rates comparable to previous studies, the MC method overestimated decomposition rates for alder and lichen due to fragmentation artefacts. The small sample size of the MC method therefore outweighs its statistical advantages arising from ease of replication. In order to derive reliable estimates of litter decomposition, both the field incubation method and the applied decomposition model must therefore be selected to suit the studied litter types.     

Determinants of litter mixing effects in a Swedish boreal forest

When the litter of a given species decomposes, it will often break down in the proximity of litters from other species. We investigated the effects of litters of 10 different species in a boreal forest of northern Sweden on each others' decomposition and N release rates; this was done through the use of litterbags containing two compartments separated by single mesh partition. Different litters could be placed on opposite sides of this mesh so that they were in contact with each other. Treatments consisted of all the possible pairwise combinations of the 10 species, with members of each pair placed in different compartments of the same litterbag. Litterbags were harvested after 1, 2 and 4 years in the field. Species differed significantly in their effects on decomposition and N loss rates of associated litters. Generally, litters from feather mosses and lichens showed the greatest promotion of decomposition on associated litters, while some vascular plant species, notably Empetrum hermaphroditum, showed the least. At year four, feather mosses also had the greatest positive effects on N loss from the litters of associated species. There were several instances in which litter of a given species decomposed at different rate when litter from its own species, rather than that of a different species, was placed in the adjacent litterbag compartment. This was particularly apparent in the second year, when across the entire data set, litters decomposed fastest when associated with their own litters. Generally, slowly decomposing litters had the greatest positive effects on decomposition of associated litters. It is proposed that in boreal forests slow decomposing litters (particularly those of feather mosses) may contribute to enhancing moisture attention in the litter layer, which in turn promotes the decomposition and N release of associated litters. Further, while litter mixing effects were clearly demonstrated in our study, they were also shown to be of secondary importance to the effects of species identity on decomposition.     

Tree influence on soil biological activity: What can be inferred from the optical examination of humus profiles?

Humus forms may vary in different forest stands, but the local influence of trees upon soil microbial and faunal activities is still imperfectly known. Optical methods could help to discern processes of litter transformation and formation of organo-mineral assemblages, allowing a better diagnostic of tree influences upon humus-soil development. The microstratification of humus was studied under a beech (Fagus crenata), a mixed oak forest (Quercus crispula and Quercus serrata), and a cedar (Cryptomeria japonica) plantation. The three sites are located in Kyoto (Japan), and share similar environmental conditions. Litter decomposition rates and soil fauna were also investigated. At the beech site, which had the thickest O horizon, the main process was the gradual fragmentation of litter. This process, together with shallow root and weak fungal development, gave rise to a stable sandwich-like structure in the O horizon. In contrast, the oak site showed a two-step transformation of litter. Initially, litter decomposition was triggered by the activity of white rot fungi, and the discarded litter decayed much more slowly thereafter. The cedar site exhibited a sharp vertical delineation between upper thick Oe horizon developed since plantation time and a relict A horizon. The optical method thus demonstrated differences in soil biological activities and litter transformation patterns under the three sites.     

The bacterial community inhabiting temperate deciduous forests is vertically stratified and undergoes seasonal dynamics

Bacterial communities living in forest soils contribute to the decomposition of organic matter and the recycling of nutrients in these ecosystems and form one of the most diverse habitats on Earth. Unfortunately, due to difficulty in culturing soil bacteria, the understanding of their ecology is still limited. In the case of temperate deciduous forests, soil microbial communities face large seasonal variations in environmental conditions, such as temperature or moisture. Moreover, the supply of nutrients also differs due to seasonal processes, such as the allocation of photosynthates into soil by the roots of primary producers or the seasonal input of fresh litter. The aim of this study was to reveal how the bacterial community responds to these seasonal processes in the litter and soil of a Quercus petraea forest. Bacterial communities from litter and from the organic and mineral horizons of soil were analyzed during the four seasons of the year by 16S rRNA gene pyrosequencing. The results revealed that the composition of the bacterial community is horizon specific. The litter horizon had a higher relative abundance of Proteobacteria and Bacteroidetes than soil, while the organic and mineral horizons had a higher abundance of Acidobacteria, Firmicutes and Actinobacteria than litter. Moreover, the bacterial community was significantly affected by seasonality in all horizons. Bacterial communities in the litter showed significant differences between the vegetation season (May and July) and the autumn and winter seasons (October, February). In mineral soil, bacterial community composition was specific in the summer, when it was significantly different from all other seasons, with a larger number of taxa described as rhizosphere and mycorrhizosphere inhabitants. The results indicate that litter decomposition is the main driver of bacterial community composition in litter horizon. In contrast to reports on fungal communities, bacterial community composition in mineral soil responds to the seasonal peaks of rhizodeposition in the summer.     

Does moder development along a pure beech (Fagus sylvatica L.) chronosequence result from changes in litter production or in decomposition rates?

The development of temperate deciduous and conifers forests stands usually results in accumulation of forest floor organic matter and a shift from mull to moder humus forms. It has been suggested that an increase in nutrient uptake by trees during their rapid growth phase leads to topsoil acidification, decrease in earthworm density and thereby a decrease in litter turnover. The focus of this paper was to examine if the mull-moder shift with forest ageing results from higher leaf litter production and/or lower litter decay rates. The objectives of this research were to determine (1) changes in macro-morphological properties of humus forms, leaf litter production, litter decay rates, soil nutrients content and pH along a 130-year pure beech (Fagus sylvatica L.) chronosequence in Normandy (Northwest France), (2) if humus form varied from mull to moder with increasing stand age, and (3) if a shift from mull to moder resulted from increased litter production, decreased litter decay rates, or both. Annual litter production did not change significantly along the chronosequence (mean 2.41 t ha⁻¹). In contrast, litter decay rates decreased significantly during the rapid growth phase of trees. In consequence, the litter turnover time (1/k) was lower in the youngest stands (20 months) compared to the oldest ones (31 months). Even in the absence of a significant pattern of variation, litter production was positively correlated with the thickness of the OF (Oi) horizon. In contrast, litter decay was strongly negatively correlated with maximum thickness of the OH (Oa) horizon, suggesting that the appearance of the humification layer was mainly due to a decrease in litter decay rate. We did not find significant changes in the main properties of the organo-mineral horizon, suggesting that soil nutrient availability may not directly affect litter dynamics. We concluded that moder development along the chronosequence resulted in decreasing litter decay rates during the aggradation phase while litter production was stable. Further studies are required to identify the ecological factors responsible for moder development along forest ageing.     

Low levels of nitrogen addition stimulate decomposition by boreal forest fungi

Climate warming and associated increases in nutrient mineralization may increase the availability of soil nitrogen (N) in high latitude ecosystems, such as boreal forests. These changes in N availability could feed back to affect the decomposition of litter and organic matter by soil microbes. Since fungi are important decomposers in boreal forest ecosystems, we conducted a 69-day incubation study to examine N constraints on fungal decomposition of organic substrates common in boreal ecosystems, including cellulose, lignin, spruce wood, spruce needle litter, and moss litter. We added 0, 20, or 200 μg N to vials containing 200 mg substrate in factorial combination with five fungal species isolated from boreal soil, including an Ascomycete, a Zygomycete, and three Basidiomycetes. We hypothesized that N addition would increase CO₂ mineralization from the substrates, particularly those with low N concentrations. In addition we predicted that Basidiomycetes would be more effective decomposers than the other fungi, but would respond weakly or negatively to N additions. In support of the first hypothesis, cumulative CO₂ mineralization increased from 635 ± 117 to 806 + 108 μg C across all fungal species and substrates in response to 20 μg added N; however, there was no significant increase at the highest level of N addition. The positive effect of N addition was only significant on cellulose and wood substrates which contained very little N. We also observed clear differences in the substrate preferences of the fungal species. The Zygomycete mineralized little CO₂ from any of the substrates, while the Basidiomycetes mineralized all of the substrates except spruce needles. However, the Ascomycete (Penicillium) was surprisingly efficient at mineralizing spruce wood and was the only species that substantially mineralized spruce litter. The activities of β-glucosidase and N-acetyl-glucosaminidase were strongly correlated with cumulative respiration (r = 0.78 and 0.74, respectively), and Penicillium was particularly effective at producing these enzymes. On moss litter, the different fungal species produced enzymes that targeted different chemical components. Overall, our results suggest that fungal species specialize on different organic substrates, and only respond to N addition on low N substrates, such as wood. Furthermore, the response to N addition is non-linear, with the greatest substrate mineralization at intermediate N levels.     

Effects of nitrogen addition and litter properties on litter decomposition and enzyme activities of individual fungi

Litter decomposition is an important process of C and N cycling in the soil. Variation in the response of litter decomposition to nitrogen (N) addition (positive, negative or neutral) has been observed in many field studies. However, mechanism about variability in individual fungal species response to N addition has not yet been well demonstrated in the literature. Therefore, the objective of this study was to investigate the effects of N addition and litter chemistry properties on litter decomposition and enzyme activities of individual fungi. Three fungal species (Penicillium, Aspergillus, and Trichoderma) were isolated from a subtropical mixed forest soil. An incubation experiment was conducted using the individual fungi with two types of litter (leaf of Pinus massoniana and needle of Cryptocarya chinensis) and different N addition levels (0, 50 and 100 for N-deficient treatments, and 500 and 1000 μg N for N-excessive treatments). Cumulative CO₂-C, enzyme activities, and lignin and cellulose loss were measured during the incubation period of 60 days. Litter decomposition and enzyme activities significantly varied with the fungal species, while the N addition and litter types greatly affected fungal enzyme activities. The N treatments significantly increased lignin-rich needle decomposition by lignocellulose decomposers (Penicillium and Aspergillus) but did not affect their leaf decomposition. On the contrary, The N treatments stimulated leaf decomposition by cellulolytic species (Trichoderma) but did not affect its needle decomposition. Correlation analysis showed that lignin in the litter was the key component to affect litter decomposition. Activities of N-acetyl-β-glucosaminidase and phenol oxidase were both positively correlated to litter decomposition. The fungi (Penicillium and Aspergillus) with higher production of N-acetyl-β-glucosaminidase showed higher litter decomposition ability. The low N addition levels stimulated Penicillium and Aspergillus litter decomposition, but they still required more N source (e.g., litter N source) to support decomposition. Depressed fungal litter N uptake (lower N-acetyl-β-glucosaminidase activities) only occurred at the highest N addition level. Litter decomposition of Trichoderma depended more on external N and its litter decomposition capability was the lowest among the three species.     

Microbial activity and quality changes during decomposition of Quercus ilex leaf litter in three Mediterranean woods

Changes in enzyme activities during litter decomposition provide diagnostic information on the dynamics of decay and functional microbial succession. Here we report a comparative study of enzyme activities involved in the breakdown of major plant components and of other key parameters (microbial respiration, fungal biomass, N, lignin and cellulose contents) in homogeneous leaf litter of Quercus ilex L. incubated in three evergreen oak woods in Southern Italy (Campania), differing for chemical and physical soil characteristics and microclimatic conditions. The results showed that the litter mass loss rates were similar in the three wood sites. Independently of the incubation sites, cellulase, xylanase and peroxydase activities showed seasonal variations with maximum and minimum levels in wet and dry periods, respectively, and this pattern closely matched microbial respiration. Activities of α- and β-amylase, instead, were high at the beginning of incubation and quickly decreased with decomposition progress because their substrate was rapidly depleted. Laccase activity, in contrast, was low at the beginning of incubation but after 6 months it increased significantly. The increase of laccase activity was correlated to an increase in fungal biomass, probably reflecting a major shift in the litter microbial community. As concerns quality changes, N and lignin content did not significantly change during decay. The cellulosic component started being degraded after about 6 months in the litter incubated in two of the three wood sites and from the start of decomposition in the third site. Apart from minor differences in the levels of certain enzyme activities, the data showed that the functional microbial succession involved in the decomposition of Q. ilex leaf litter did not change appreciably in response to differences in soil and microclimatic conditions in the incubation sites.     

Plant litter quality influences the contribution of soil fauna to litter decomposition in humid tropical forests, southwestern China

The aim of this field experiment was to quantify the contribution of soil fauna to plant litter decomposition in three forest sites differing in C/N ratio under natural conditions in Xishuangbanna, southwestern China. We conducted a survey of soil fauna communities, the forest floor litter and investigated mass loss of mixed tree species leaf litter for two years in a tropical secondary forest, an evergreen broad-leaf forest and a tropical rain forest. Exclusion treatments of different sized soil fauna from the leaf litter by using varying mesh size litter bags (2 mm and 0.15 mm) were also performed. Mass loss and C and N concentrations in litter bag leaf materials were determined at monthly intervals. We found that: (1) the three forests differed in floor litter biomass and nutrient contents but not in soil fauna richness and abundance; (2) litter mass loss and decomposition rate were slower when soil macrofauna and most of mesofauna were excluded; and (3) greatest soil fauna contribution to plant litter decomposition occurred in the rain forest, where leaf litter C/N ratio was also highest (41.5% contribution: 54.8 C/N ratio), in comparison to 8.69% in the broad-leaf forest and 19.52% in the secondary forest, both with low leaf litter C/N ratios (<32). Our results suggested that, soil fauna played a more pronounced role in the decomposition of mixed leaf litter in tropical rain forest, and significantly bigger effects from fauna were ascribed to the enhancement of N concentration and decrease of C concentration of the initially high C/N ratio litter in this forest site.     

Litter decomposition in primary forest and adjacent fire-disturbed forests in the Gran Sabana,southern Venezuela

Litter decomposition and nutrient dynamic were studied in tall primary forest (TF) and in adjacent slightly fire-affected (MF) and strongly fire-affected (LF) forests of the Gran Sabana, southern Venezuela. The aim of the study was to compare the mass and nutrient loss of litter in undisturbed forest and adjacent fire-disturbed forests growing under the same soil conditions. The results showed no significant differences in the dry-mass reduction among TF, MF, and LF after 1-year of decomposition. At the end of the decomposition period, the mass loss was 31% in TF, 24% in MF, and 25% in LF. With few exceptions, the initial nutrient content of the litter did not show significant differences among the studied forests. The initial litter was poor in nutrients, especially in P, with C/N ratio extremely high, particularly in LF. Both residual P content and C/N ratio were the most effective predictors of dry-mass loss. The general trend in the three studied forests was net N and to less extend P immobilization and release of K, Mg, and Ca. We concluded that during a 1-year period, the decomposition process was similar in undisturbed and adjacent fire-disturbed forests in the Gran Sabana and that the low litter decomposition seems to be mainly controlled by the low chemical quality of the decomposing litter.     

Mass loss and nutrient release during litter decay in peatland: The role of microbial adaptability to litter chemistry

In peatlands the reduced decomposition rate of plant litter is the fundamental mechanism making these peat-accumulating ecosystems effective carbon sinks. A better knowledge of litter decomposition and nutrient cycling is thus crucial to improve our predictions of the effects of anthropogenic perturbation on the capacity of peatlands to continue to behave as carbon sinks. We investigated patterns of plant litter decomposition and nutrient release along a minerotrophic-ombrotrophic gradient in a bog on the south-eastern Alps of Italy. We determined mass loss as well as P, N, K, and C release of seven vascular plant species and four moss species after 1 year in both native and transplanted habitats. Hence, differences in litter decay were supposed to reflect the degree of adaptability of microbial communities to litter quality. Polyphenols/nutrient and C/nutrient quotients appeared as the main parameters accounting for decomposition rates of Sphagnum litter. In particular, litter of minerotrophic Sphagnum species decomposed always faster than litter of ombrotrophic Sphagnum species, both in native and transplanted habitats. Decomposition rates of vascular plant litter in native habitats were always higher than the corresponding mass loss rates of Sphagnum litter. Minerotrophic forbs showed the fastest decomposition both in native and transplanted habitats in accordance with low C/P and C/N litter quotients. On the other hand, C/P quotient seems to play a primary role also in controlling decomposition of graminoids. Decomposition of deciduous and evergreen shrubs was negatively related to their high lignin content. Nitrogen release from Sphagnum litter was primarily controlled by C/N quotient, so that minerotrophic Sphagnum litter released more N than ombrotrophic Sphagnum litter. Overall, we observed slower N release from litter of ombrotrophic vascular plant species compared to minerotrophic vascular plant species. No single chemical parameter could predict the variability associated with different functional groups. The release of K was very high compared to all the other nutrients and rather similar between ombrotrophic and minerotrophic litter types. In Sphagnum litter, a higher C/P quotient was associated with a slower P mineralisation, whereas a faster P release from vascular plant litter seems primarily associated with lower C/P and polyphenols/P quotients.     

Impacts of anthropogenic N additions on nitrogen mineralization from plant litter in exotic annual grasslands

Urban regions of southern California receive up to 45 kg N ha^-1 y^-1 from nitrogen (N) deposition. A field decomposition study was done using ¹⁵N-labelled litter of the widespread exotic annual grass Bromus diandrus to determine whether elevated soil N is strictly from N deposition or whether N mineralization rates from litter are also increased under N deposition. Tissue N and lignin concentrations, which are inversely related in field sites with high and low N deposition, determine the rate at which N moves from plant litter to soil and becomes available to plants. The effect of soil N on N movement from litter to soil was tested by placing litter on high and low N soil in a factorial experiment with two levels of litter N and two levels of soil N. The litter quality changes associated with N deposition resulted in faster rates of N cycling from litter to soil. Concentrations of litter-derived N in total N, NH₄⁺, NO₃⁻, microbial N and organic N were all higher from high N/low lignin litter than from low N/high lignin litter. Litter contributed more N to soil NH₄⁺ and microbial N in high N than low N soil. At the end of the study, N mineralized from high N litter on high N soil accounted for 46% of soil NH₄⁺ and 11% of soil NO₃⁻, compared to 35% of soil NH₄⁺ and 6% of soil NO₃⁻ from low N litter on low N soil. The study showed that in high N deposition areas, elevated inorganic soil N concentrations at the end of the summer N deposition season are a result of N mineralized from plant litter as well as from N deposition.     

Moss-nitrogen input to boreal forest soils: Tracking 15N in a field experiment

Cyanobacteria living epiphytically on mosses in pristine, unpolluted areas fix substantial amounts of atmospheric nitrogen (N) and therefore represent a primary source of N in N-limited boreal forests. However, the fate of this N is unclear, in particular, how the fixed N₂ enters the soil and becomes available to the ecosystem. In this study, we applied ¹⁵N-ammonium chloride (¹⁵N-NH₄Cl) onto carpets of the feather moss Pleurozium schreberi and traced the ¹⁵N label into green (living) and brown (senescent) moss and into the upper soil layer over time. Further, we placed filters between moss and soil to assess the role of moss-associated fungi for N-transfer to the soil. The experiment was conducted at endpoints of a N₂ fixation gradient in Northern Sweden. Feather moss retained the applied N in the green moss parts for up to 1 year and no increase of excess ¹⁵N was found in the brown moss parts or in the soil within that same time frame. The filter treatment did not alter the ¹⁵N-distribution in moss or soil. Nitrogen retention in the moss was similar regardless of position along the N₂ fixation gradient. Our results suggest that mosses represent a short-term inorganic N sink and that transfer of N to the soil is not facilitated by fungal hyphae.     

Exotic plant influences soil nematode communities through litter input

The impact of exotic plant invasions on soil communities and nutrient cycling processes has received an increasing attention in recent years. To test whether the exotic plant invasions affect nematode communities through altering litter quality, we compared mass loss and nematode colonization during the stem litter decomposition of invasive Spartina alterniflora and native Phragmites australis in salt marshes of the Yangtze River estuary, China. Plastic drinking straws were synchronously used as controls. The addition of plant residues was found stimulating the growth of nematodes, particularly bacterial feeders on day 16 after burial. A top-down control of bacterivous nematodes by carnivores existed in nematode succession during the litter decomposition. With higher nitrogen content and lower C:N ratio, stem litter of the invasive S. alterniflora decayed faster and supported more abundant nematodes than the native P. australis. The greater nematode abundance in S. alterniflora was mainly due to two dominant genera of bacterial nematodes, namely Diplolaimelloides and Diplolaimella. Lower values of maturity index and structure index in S. alterniflora than in P. australis litter indicate that a more degraded food web condition resulted from the faster litter decay. A considerable difference in nematode community structures between two litter types only occurred in a certain period of the decomposition (from 8 to 32 days after burial), suggesting that the changes in faunal community structure are time dependent. In summary, this study confirmed the hypothesis that the invasion of S. alterniflora stimulates the growth of bacterial nematodes by producing higher quality of litter than the native P. australis. The results obtained here suggest that the invasion of exotic plant is likely to alter ecosystem functions indirectly through exerting its effect on soil decomposer communities such as nematodes.     

Limitations of faunal effects on soil carbon flow: density dependence, biotic regulation and mutual inhibition

Soil animals are known to stimulate soil microbial activity and thereby to accelerate decomposition of soil organic matter. In this paper, we investigate potential limitations of soil animal effects on soil carbon flow by analysing how animal effects relate to the density of four major faunal groups. Specifically, we analyse the extent to which faunal effects are subject to biotic regulation or to mutual inhibition between groups under different levels of resource supply.In an extensive laboratory experiment, 96 microcosms established in three consecutive blocks were inoculated with nematodes, enchytraeids, microarthropods, and lumbricids. Each faunal group was inoculated in three densities, including combinations of groups. Introduced animal densities were within the natural range of densities in fallow soil. Bare agricultural soil and soil covered with maize litter were used as substrates. The microcosms were kept under constant conditions at 12 °C and 50% water holding capacity for 8 weeks. Soil CO₂ evolution was measured daily by means of gas chromatography.Animal effects were on an average relatively stronger in bare soil (+95% CO₂; R²=0.76) than in soil with litter (+14% CO₂; R²=0.40), where organic matter decomposition was seven times more intense. Higher animal densities generally led to accelerated decomposition up to three times that of the controls. However, beyond a specific density, decomposition rates stopped increasing or even declined, depending on the faunal group. In addition, animal effects were limited by mutual inhibition between groups in bare soil where effects were strong, while stimulatory interactions were prominent in the litter treatments where effects were generally weak.We interpret the limitation of soil faunal effects on soil carbon flow in terms of incomplete habitat exploitation and biotic regulation. Under conditions of substrate homogeneity, such as in the bare soil treatments, animal effects were stronger, but they were limited by overexploitation. Under conditions of substrate heterogeneity, such as in the litter treatments, animal effects were limited by incomplete habitat utilisation. We assume that complementary habitat colonisation by different faunal groups in the litter treatments gave rise to positive diversity effects, but that these effects did not compensate for reduced overall habitat utilisation. We infer that a knowledge of faunal resource utilisation and of mutual inhibition of faunal groups can be exploited for ecological soil management towards stabilisation of soil organic matter.     

Changes in the fauna and its contribution to mass loss and N release during leaf litter decomposition in two deciduous forests

Investigations on the mass loss of leaf litter were carried out between 1992 and 1994 using litter bags of 0.02 mm and 5 mm mesh sizes in a beech and a mixed forest in northern Germany. The two forests on moder humus differed in soil faunal composition, vegetation type, and nutrient supply. Mass loss and N and C concentrations were determined from the litter at bimonthly intervals. From subsamples macrofauna were sorted by hand and mesofauna was extracted by heat. The biomass and N content of the litter bag fauna was estimated. Mass loss, particularly that attributed to the fauna, was different between the two sites with highest rates in the mixed forest and lowest at the beech site. A significantly higher rate of N release was found for the litter extracted from 5 mm mesh size litter bags in the mixed forest but not in the beech forest. Collembola and Cryptostigmata changed in numbers during litter breakdown. Collembola reached high numbers in the beginning, whereas Cryptostigmata dominated later. The diversity of Cryptostigmata increased at both sites during litter breakdown, whereas collembolan diversity only increased in the beech forest and remained at the same level in the mixed forest. Several species of Collembola and Cryptostigmata occurred earlier in the mixed forest than in the beech forest. Mass loss rate attributable to the fauna did not correspond to total faunal biomass. Only Isopoda, Diplopoda and Cryptostigmata appeared to affect the mass loss positively, whereas the biomass of Lumbricidae was negatively correlated with mass loss, particularly in the beech forest. On the other hand, the release of N attributable to the fauna was positively correlated with the total faunal biomass in the beech forest and Lumbricidae in particular were positively correlated with N-release at both sites.     

Major mechanisms contributing to the macrofauna-mediated slow down of litter decomposition

To understand why excrements of soil macrofauna often decompose more slowly than leaf litter, we fed Bibio marci larvae the litter of tree species differing in litter quality (Alnus glutinosa, Salix caprea, and Quercus robur) and then measured respiration induced by litter and excrements. We also measured respiration induced by the same litter artificially modified to mimic faunal effects; the litter was modified by grinding, grinding with alkalinization to pH = 11, grinding with coating by kaolinite, and grinding with both alkalinization and coating. Decomposition of excrements tended to be slower for willow and was significantly slower for oak and alder than for the corresponding litter. With oak, decomposition was slower for all artificially modified litter than for non-modified litter. The reduction in the decomposition was similar for excrements and for alder and willow litter that was ground, coated, and alkalinized. In alder, a similar reduction was found in ground and alkalinized litter. ¹³C NMR indicated that gut passage increases aliphatic components and decreases polysaccharides. Pyrolysis indicated that gut passage increases the ratio of guaiacyl to hydroxymethyl derivatives in lignin. Our findings indicate that the decreased decomposition rate of excrements might result from the removal of easily available polysaccharides, the increase in aliphatic components, an increase in the resistant components of lignin, the accumulation of microbial cell walls, and the binding of nitrogen into complexes with aromatic components. Several of these mechanisms are supported or determined by litter alkalinization during gut passage.     

The oribatid mite Scheloribates moestus (Acari: Oribatida) alters litter chemistry and nutrient cycling during decomposition

It is widely accepted that microarthropods influence decomposition dynamics but we know relatively little about their effects on litter chemistry, extracellular enzyme activities, and other finer-scale decomposition processes. Further, few studies have investigated the role of individual microarthropod species in litter decomposition. The oribatid mite Scheloribates moestus Banks (Acari: Oribatida) is abundant in many U.S. ecosystems. We examined the potential effects of S. moestus on litter decomposition dynamics and chemical transformations, and whether these effects are influenced by variation in initial litter quality. We collected corn and oak litter from habitats with large populations of S. moestus and in microcosms with and without mites measured respiration rates, nitrogen availability, enzyme activities, and molecular-scale changes in litter chemistry. Mites stimulated extracellular enzyme activities, enhanced microbial respiration rates by 19% in corn litter and 17% in oak litter over 62 days, and increased water-extractable organic C and N. Mites decreased the relative abundance of polysaccharides in decomposing corn litter but had no effect on oak litter chemistry, suggesting that the effects of S. moestus on litter chemistry are constrained by initial litter quality. We also compared the chemistry of mite feces to unprocessed corn litter and found that feces had a higher relative abundance of polysaccharides and phenols and a lower relative abundance of lignin. Our study establishes that S. moestus substantially changes litter chemistry during decomposition, but specific effects vary with initial litter quality. These chemical transformations, coupled with other observed changes in decomposition rates and nutrient cycling, indicate that S. moestus could play a key role in soil C cycling dynamics.     

Effects of addition of maize litter and earthworms on C mineralization and aggregate formation in single and mixed soils differing in soil organic carbon and clay content

C mineralization and aggregate stability directly depend upon organic matter and clay content, and both processes are influenced by the activity of microorganisms and soil fauna. However, quantitative data are scarce. To achieve a gradient in C and clay content, a topsoil was mixed with a subsoil. Single soils and the soil mixture were amended with 1.0 mg maize litter C g soil⁻¹ with and without endogeic earthworms (Aporrectodea caliginosa). The differently treated soils were incubated for 49 days at 15 °C and 40% water holding capacity. Cumulative C mineralization, microbial biomass, ergosterol content and aggregate fractions were investigated and litter derived C in bulk soil and aggregates were determined using isotope analyses. Results from the soil mixture were compared with the calculated mean values of the two single soils. Mixing of soil horizons differing in carbon and clay content stimulated C mineralization of added maize residues as well as of soil organic matter. Mixing also increased contents of macro-aggregate C and decreased contents of micro-aggregate C. Although A. caliginosa had a stimulating effect on C mineralization in all soils, decomposition of added litter by A. caliginosa was higher in the subsoil, whereas A. caliginosa decreased litter decomposition in the soil mixture and the topsoil. Litter derived C in macro-aggregates was higher with A. caliginosa than with litter only. In the C poor subsoil amended with litter, A. caliginosa stimulated the microbial community as indicated by the increase in microbial biomass. Furthermore, the decrease of ergosterol in the earthworm treated soils showed the influence of A. caliginosa on the microbial community, by reducing saprotrophic fungi. Overall, our data suggest both a decrease of saprotrophic fungi by selective grazing, burrowing and casting activity as well as a stimulation of the microbial community by A. caliginosa.