Leaf litter decomposition in the pure and mixed plantations of <Emphasis Type="Italic">Cunninghamia lanceolata</Emphasis> and <Emphasis Type="Italic">Michelia macclurei</Emphasis> in subtropical China

Leaf litter decomposition of Cunninghamia lanceolata, Michelia macclurei, and their mixture in the corresponding stands in subtropical China was studied using the litterbag method. The objective was to assess the influence of native evergreen broadleaved species on leaf litter decomposition. The hypotheses were: (1) M. macclurei leaf litter with lower C/N ratio and higher initial N concentration decomposed faster than C. lanceolata litter, (2) decomposition rates in litter mixtures could be predicted from single-species decay rates, and (3) litters decomposed more rapidly at the site that contained the same species as in the litterbag. The mass loss of leaf litter was positively correlated with initial N concentration and negatively correlated with C/N ratio. The decomposition rate of M. macclurei leaf litter was significantly higher than that of C. lanceolata needle litter in the pure C. lanceolata stand. Contrary to what would be predicted, the litter mixture decomposed more slowly than expected based on the results from component species decomposing alone. There was no significant difference in litter decomposition rate between different habitats.     

Effects of tree species and topography on soil chemistry, litter quality, and decomposition in Northeast Turkey

Leaf litters from beech (Fagus orientalis Lipsky.) and oak (Quercus robur L.), and needle litters from fir (Abies nordmanniana Spach.) and pine (Pinus sylvestris L.) trees were collected from north-facing site and south-facing site and at three slope positions (top, middle and bottom) on each aspect that varied in soil chemical characteristics (soil pH, cation exchange capacity and base saturation). The litters were analysed for initial total carbon, nitrogen, acid detergent fibre, lignin and cellulose concentrations. Nitrogen, acid detergent fibre and lignin concentrations and carbon:nitrogen and lignin:nitrogen ratios varied significantly within and between species according to soil chemical characteristics on aspects and slope positions. Litter decomposition was studied in the field using the litterbag technique. The litters were placed on two aspects and at three slopes on each aspect in October 2001, and were sampled every 6-month for 2 years. The main effects of aspect, species and slope position on decomposition rates were all statistically significant. Oak leaf litter showed highest decomposition rates, followed by pine, fir and beech litter, and the litters placed on north-facing site decomposed faster than those on the south-facing site. The litters placed at the top slope position decomposed slower than at those at either the bottom or middle positions. Initial lignin concentrations explained most of the variation in decomposition rates between species, and within species for the aspects and the slope positions, but the explained variance showed differences between aspects and slope positions. This result illustrates the important point that litter quality may define the potential rates of microbial decomposition but these are significantly influenced by the biotic and abiotic environment in which decomposition takes place.     

Nonadditive effects of litter mixtures on decomposition and correlation with initial litter N and P concentrations in grassland plant species of northern China

We studied the occurrence of nonadditive effects of litter mixtures on the decomposition (the deviation of decomposition rate of litter mixtures from the expected values based on the arithmetic means of individual litter types) of litters from three plant species (i.e., Stipa krylovii Roshev., Artemisia frigida Willd., and Allium bidentatum Fisch. ex Prokh. & Ikonn.-Gal.) endemic to the grassland ecosystems of Inner Mongolia, northern China and the possible role of initial litter N and P on such effects. We mixed litters of the same plant species that differed in N and P concentrations (four gradients for each species) in litterbags and measured mass losses of these paired mixtures after 30 and 80 days under field conditions. We found the occurrence of positive, nonadditive effects of litter mixtures and showed that the magnitude of the nonadditive effects were related to the relative difference in the initial litter N and P concentrations of the paired litters.     

Use of a modified litterbag technique for the study of litter mixtures

The applicability of a modified, easily constructed litterbag is evaluated in this study. In this litterbag, designed for use in decomposition studies involving litter mixtures, litter species are decomposed in separated by mesh compartments so that they can interact with adjacent species and at the same time can be retrieved “non-contaminated” by them until late decomposition stages. The validity of the technique was examined by comparing mass loss rates of two litter species (Ailanthus altissima and Medicago sativa) incubated in single and mixed new litterbags, with rates in conventional litterbags. The results, referring to the disappearance of up to 65% of the material at constant moisture and temperature conditions, were not significantly affected by the use of the modified litterbag. Although differences between expected and measured decomposition rates of the mixture of the two species were not statistically significant, mixing resulted in a modification of decomposition rates of the individual components compared with their respective rates in single species litterbags.     

Litter decomposition and faunal activity in Mediterranean forest soils: effects of N content and the moss layer

Accumulation of soil carbon is mainly controlled by the balance between litter production and litter decomposition. Usually In Mediterranean forests there are contrasting conditions in the distribution of faunal activity and the moss layer that may have different effects on litter decomposition. Decomposition and faunal activity were studied by exposing litter of contrasting quality (Pinus halepensis Mill. and Quercus ilex L.) for 3.5 yr in three Mediterranean pine forests of the eastern Iberian Peninsula. The effects of mosses on decomposition and on faunal activity were studied by exposing P. halepensis litter either on moss patches or directly on the forest floor. Faecal pellet production was used as an indication of faunal activity. Water availability or soil characteristics seem to limit faunal activities in the drier sites. Faecal pellets were not found during the first stages of decomposition and in all sites they appeared when about a 30% of the initial litter had decomposed. Under wet conditions faecal pellet production was very high and a mass balance suggested that soil faunal activity may result in a net flow of organic matter from the lower organic horizons to the surface Oi horizon. Mosses slightly increased mass loss of pine litter probably as a consequence of high potentially mineralizable nitrogen in the Oa horizon of moss patches and also, perhaps, as a consequence of the higher moisture content measured in the Oi horizon needles sampled among the mosses. In contrast, moss patches reduced faunal activity. The effect of litter quality on mass loss was not always significant, suggesting an interaction between litter quality and site conditions. During the first stages of decomposition there was N immobilisation in P. halepensis litter (poorer in N) and N release from Q. ilex litter (richer in N). In conclusion, in these forests soil microclimate and/or N availability appear to be more important controlling litter decomposition than the distribution of faunal activity.     

Do invertebrate decomposers affect the disappearance rate of litter mixtures?

We designed an experiment using litter bags with fine and coarse mesh size to analyse interacting effects between invertebrate decomposers and the number of litter species on litter disappearance rates. We used litter of nine broad-leaved tree species to compare disappearance rates of litter from single species with mixtures of two to six species. Species composition of litter and invertebrates interacted strongly in their effects on litter disappearance rate. Contribution of invertebrates to litter disappearance increased with time mainly for litter which disappeared slower in the absence of invertebrates. Disappearance rates were positively correlated with initial N content and negatively correlated to initial C content of litter. These relationships were stronger in the presence of invertebrates, suggesting that their activity is positively related to initial litter chemistry. Number of component litter species, however, had no effect on disappearance rate irrespective of the activity of invertebrates. Using individual rates of disappearance for single species, we calculated the expected rates of disappearance for each of the experimental mixtures of leaf litters. We found that mixtures of several species of leaf litter resulted in significant deviations from the expected values. These deviations showed a significant effect of the number of component litter species. However, this result was caused by a strong negative deviation of one single mixture of six species. The presence of invertebrates resulted in even greater deviations from the expected values, suggesting an important contribution of invertebrates to the effects of litter mixing on litter disappearance rates. Hence, our results underline the importance of idiosyncratic effects of species traits in mixtures. Our results suggest that the influence of invertebrate decomposers interacts with litter chemistry during decomposition, but is not affected by litter species richness per se.     

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.     

茂兰喀斯特森林自然保护区凋落叶分解动态

[目的]研究喀斯特森林生态系统凋落叶分解特征,为喀斯特森林区石漠化防治及水土保持提供科学依据。[方法]采用1 mm网孔孔径分解袋,对茂兰喀斯特森林自然保护区不同树种凋落叶(落叶和常绿叶)在不同坡位的分解状况进行为期18个月的观测研究。研究茂兰喀斯特森林自然保护区凋落叶失重率和干重残留率动态变化、分解速率及养分释放特征。[结果]凋落叶分解过程呈现"快—慢—快"的周期变化,春夏季分解速度快于秋冬季,落叶树种凋落叶分解速度快于常绿树种凋落叶,不同坡位凋落叶的分解速度表现为:下坡中坡上坡。利用Olson模型对凋落叶分解50%和95%所需时间进行估测,发现落叶树种凋落叶分解50%和95%所需时间分别为0.95～1.66 a和4.13～7.19 a,常绿树种凋落叶分解50%和95%所需时间分别为1.14～1.69 a和4.92～7.30 a,二者无显著性差异。凋落叶分解速率低于中亚热带东部常绿阔叶林和常绿落叶阔叶混交林,但比同区域喀斯特次生林与人工林高。落叶树种凋落叶和常绿树种凋落叶的N元素释放模式为富集—释放模式,C含量随分解时间的波动差异显著,总体在不断减少,而C/N比呈逐步下降的趋势。[结论]由于不同树种凋落叶初始养分含量和叶片理化结构的差异,落叶比常绿叶具有更快的分解速率和养分释放速率,对促进喀斯特森林生态系统物质循环起着积极作用。     

Succession of collembolan communities during decomposition of leaf and root litter: Effects of litter type and position

Little is known about the collembolan community involved in the decomposition of fine root (≤2.0 mm in diameter) litter, which is largely different from leaves in both litter quality and position. The collembolan communities involved in root and leaf litter decomposition were compared in a litterbag experiment in a coniferous forest of Chamaecyparis obtusa. A two-factor experiment (litter type × litter position) was conducted to evaluate the relative effects of litter quality and position. Litterbags of roots and leaves were each placed at two positions (on the soil surface and in the soil), and were collected at seven different times over three years. Abundance and biomass of Collembola involved in root decomposition in the soil were higher than those involved in leaf decomposition on the soil surface, and the collembolan community composition largely differed between these two types of litterbag. Differences between root and leaf decomposition were mainly caused by litter position, but effects of litter type were also detected at species-level. Species that preferred roots were abundant at an early stage of litter decomposition in the soil. Because the early stage of decomposition in the soil is naturally achieved only by root litter initially deposited in the soil, root litter may function as an essential resource for certain species. The results of this study indicate that root litter contributes to collembolan community organization as a spatially and qualitatively different resource than leaf litter. This also suggests that root litter is decomposed via different soil faunal processes than leaf litter.     

Litter mixture effects on decomposition in tropical montane rainforests vary strongly with time and turn negative at later stages of decay

In a litterbag study in a tropical montane rainforest in Ecuador we assessed the impact of leaf litter species identity and richness on decomposition. We incubated leaf litter of six native tree species in monocultures and all possible two and four species combinations and analysed mass loss over a period of 24 months. Mass loss in monocultures averaged 30.7% after 6 month and differed significantly between species with variations being closely related to initial concentrations of lignin, Mg and P. At later harvests mass loss in monocultures averaged 54.5% but did not vary among leaf litter species and, unexpectedly, did not increase between 12 and 24 months suggesting that litter converged towards an extremely poor common quality retarding decomposition. After 6 months mass loss of leaf litter species was significantly faster in mixtures than in monocultures, resulting in synergistic non-additive mixture effects on decomposition, whereas at later harvests mass loss of component litter species was more variable and leaf litter mixture effects differed with species richness. Mass loss in the two species mixtures did not deviate from those predicted from monocultures, while we found antagonistic non-additive mixture effects in the four species mixtures. This suggests that litter species shared a poor common quality but different chemistry resulting in negative interactions in chemically diverse litter mixtures at later stages of decomposition. Overall, the results suggest that interspecific variations in diversity and composition of structural and secondary litter compounds rather than concentrations of individual litter compounds per se, control long term leaf litter decomposition in tropical montane rainforests. Plant species diversity thus appears to act as a major driver for decomposition processes in tropical montane rainforest ecosystems, highlighting the need for increasing plant conservation efforts to protect ecosystem functioning of this threatened biodiversity hotspot.     

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 Mediterranean ecosystems: Modelling the controlling role of climatic conditions and litter quality

A new process-based model of litter decomposition, characterized by detailed climatic data input and simple litter quality parameters, is proposed. Compared to existing litter carbon models, specific implementations for temperature and moisture limiting effects have been adopted. The model is capable to represent decomposition processes in Mediterranean ecosystems, with summer drought slowing down, even at optimal temperatures, the litter decay rates of sclerophyll plants whose leaf masses are rich in structural compounds and low in N content. The model was calibrated by a best fitting procedure of two different datasets. First, unpublished results of litterbag experiments on leaf litter of 9 Mediterranean species, decomposing under controlled and not limiting temperature and water conditions, have been used to estimate the decay rate dependency from litter quality that was defined by only three initial C pools (labile, stable and recalcitrant compounds) instead of traditional N-based indices. Second, a set of published data from three medium-term field experiments on a single species, Phillyrea angustifolia, decomposing under different climatic conditions, have been used to estimate the limiting effects of temperature and moisture. The model was then validated against published data on seven other species and showed a correct reproduction of the major patterns of litter mass loss during decomposition processes of other seven different Mediterranean species. The model simulations, satisfactory for different litter types under a wide range of climatic conditions, suggest that factors which were not taken into account, such as initial litter N contents, microclimatic variations related to stand structure, soil chemistry and texture, and microbial communities, are not very significant for assessing decomposition dynamics in Mediterranean ecosystems. The minimal requirements of input data, the simple structure, and the easiness of parameterisation make our model, among the many other available litter carbon models, an attractive alternative for different research purposes, at least for Mediterranean ecosystems.     

Early-stage decomposition of maize litter at different positions in a semi-arid cropland

ABSTRACT

Litter decomposition plays a crucial role in controlling carbon (C) cycling and nutrient turnover in agroecosystems. In this study, the litterbag method was used to investigate the mass loss and nitrogen (N) dynamics of maize litters (culms, leaves and sheaths) at aerial, surficial and belowground positions in the initial 191 d of decomposition. For any tissue, the decomposition rates in the air and on the soil surface were similar, but both were less than the decomposition rates below the ground. The sheaths always decomposed at a lower rate than the other two tissues at any position. During decomposition, the N concentrations for all tissues decreased at both the aerial and the surficial positions but increased for belowground leaves and sheaths in the last months. For the N amount, these three tissues generally exhibited a net N release during the experiment irrespective of the position. Overall, position plays a crucial role in controlling early-stage litter decomposition in croplands, and this role will be modified by litter quality. Therefore, further studies on litter decomposition should fully consider the litter position to comprehensively evaluate the biogeochemical cycles in agroecosystems.     

Estimating N transfers between N<Subscript>2</Subscript>-fixing actinorhizal species and the non-N<Subscript>2</Subscript>-fixing<Emphasis Type="Italic"> Prunus avium</Emphasis> under partially controlled conditions

Two methods of N transfer between plants—by litter decomposition and root-to-root exchange—were examined in mixed plantations of N-fixing and non-fixing trees. Nitrogen transfers from decaying litters were measured by placing ¹⁵N-labelled litters from four actinorhizal tree species around shoots of containerized Prunus avium. Nitrogen transfers by root-to-root exchanges were measured after foliar NO₃-¹⁵N fertilization of Alnus subcordata and Elaeagnus angustifolia growing in containers in association with P. avium. During the first 2 years of litter decomposition, from 5–20% of the N, depending on the litter identity, was released and taken up by P. avium. N availability in the different litters was strongly correlated with the amount of water-soluble N, which was highest in leaves of E. angustifolia. In the association between fixing and non-fixing plants, 7.5% of the A. subcordata N and 25% of E. angustifolia N was transferred to P. avium by root exchange. These results showed that the magnitude of N transfers by root exchange depended on the associated N₂-fixing species. Among the species investigated, E. angustifolia displayed the highest capacity for exudating N from roots as well as for releasing N from litters. These qualities make this tree a promising species for enhancing wood yields in mixed stands.     

Tree species effects on microbial respiration from decomposing leaf and fine root litter

Tree species have an impact on decomposition processes of woody litter, but the effects of different tree species on microbial heterotrophic respiration derived from decomposing litter are still unclear. Here we used leaf and fine root litter of six tree species differing in chemical and morphological traits in a temperate forest and elucidated the effects of tree species on the relationships between litter-derived microbial respiration rates and decomposition rates and morphological traits, including specific leaf area (cm² g⁻¹) and specific root length (m g⁻¹) of litter at the same site. Litterbags set in forest soil were sequentially collected five times over the course of 18 months. During litter decomposition, microbial respiration from leaf and fine root litter differed among the six tree species. Temporal changes in the remaining mass and morphology (specific leaf area and specific root length) were observed, and the magnitude of these changes differed among species. Positive correlations were observed between respiration and mass loss or morphology across species. These results revealed that litter mass loss and morphological dynamics during decomposition jointly enhanced microbial respiration, and these carbon-based litter traits explained species differences in decomposition of leaves and fine roots. In conclusion, tree species influenced the magnitude and direction of microbial respiration during leaf/fine root litter decomposition. Tree species also affected the relationship between microbial respiration and litter decomposition through direct effects of litter traits and indirect effects mediated by regulation of heterotroph requirements.     

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.     

Particle size alters litter diversity effects on decomposition

Nutrient transfer between decomposing leaves may explain non-additive species diversity effects on decomposition. The influence of the diversity of litter species on decomposition was compared in mixtures composed of large (>200 mm²) or small (<9 mm²) litter fragments. The increase in the number of species (aspen, oak, alder and pine, from monocultures to four species in all possible combinations) initially (at day 43) suppressed respiration, but eventually (after 142 days) did not affect the mass loss of the mixtures of small litter fragments. In contrast, the decomposition of litter in large fragments increased with increased diversity, and 93% of all mixtures decomposed faster than would be predicted from monocultures. The results suggest that the active transport of nutrients by fungal hyphae, rather than passive diffusion, drives positive effect of the litter species diversity on decomposition.     

Leaf litter decomposition in a chaparral ecosystem, Southern California

Decomposition losses from leaves of three evergreen chaparral species, scrub oak (Quercus dumosa), ceanothus (Ceanothus crassifolius), and manzanita (Arctostaphylos glauca), were quantified over a 2-y field exposure using litterbags. Changes in ash-free dry mass, C, and N were monitored at 2- to 6-month intervals at four replicate sites composed of patches of these three chaparral species. Three proximate C fractions were extracted from fresh and decomposing litter samples: polar and non-polar extractives (EXT), acid-solubles (ACID), and acid-insolubles (KLIG). The chemical structure of fresh and decomposed litter was additionally characterized using high-resolution solid-state ¹³C NMR spectroscopy, while morphological properties were examined by scanning electron microscopy (SEM). After 2 y, the litters had lost between 20.7%±1.2 (Ceanothus) and 35.2%±6.8 (Quercus) of their original ash-free dry mass. The manzanita decomposed at a significantly faster rate than the other two litter types during the first few months of field exposure. Yet, after 2 y, mass loss was greater for the oak. Differences in decomposition rates could not be accounted for based on a single litter quality index. Fresh manzanita exhibited a significantly higher N content, which could explain its initially faster decay rate. Fresh oak litter, on the other hand, had a relatively high ACID and O-alkyl C (O-ALK) content, which may have been responsible for its decay pattern. Fresh ceanothus contained a relatively low KLIG content, yet it decomposed more slowly than the two other species. The solid-state ¹³C NMR spectra of the ceanothus litter had two peaks characteristic of proanthocyanidins, which likely contributed to the recalcitrance of this litter type. SEM revealed that ceanothus leaf surfaces were left nearly unchanged after field exposure. In comparison, the oak and manzanita leaf surfaces were pitted and covered by microbial growth to the point of being unrecognizable. Taken together, our results indicate that a combination of biological, physical and chemical factors need to be examined to clarify the different decomposition rates and patterns of these three chaparral species.     

Secondary compounds can reduce the soil micro-arthropod effect on lichen decomposition

Phenolic compounds have been shown in several studies to have important ‘carryover effects’ on litter decomposition, microbial nutrient immobilization and nutrient availability. These effects arise in part because of the adverse effect they have on the feeding activities of litter-feeding invertebrates such as micro-arthropods that drive decomposition processes. However, the interactive effects of phenolic compounds and soil micro-arthropods on litter decomposition are poorly understood. Phenolic compounds can easily be removed by acetone rinsing from living lichens, allowing us to specifically test the role that phenolic compounds (and their removal) have in controlling the effects of micro-arthropods on the decomposition of their litter. We performed a litter-bag experiment aimed at exploring how lichen litter mass loss and nutrient release during decomposition was affected by phenolics (by using acetone rinsed and non-rinsed lichen material) and micro-arthropod activity (by using different mesh sizes to allow or exclude entry by micro-arthropods) for each of six contrasting lichen species (Cladonia rangiferina, Cladonia stellaris, Evernia prunastri, Hypogymnia physodes, Pseudevernia furfuracea and Usnea dasypoga). Both the removal of phenolic compounds and the presence of micro-arthropods accelerated mass and nutrient release overall, but not for either of the two Cladonia species. Removal of phenolics also had an overall positive effect on the effects of arthropods on the loss of P, but not mass and N, from the decomposing lichens. Further, for U. dasypoga, but not the other species, natural levels of phenolic compounds deterred micro-arthropods from accelerating mass loss, and the removal of these compounds enabled micro-arthropods to enhance its decomposition. Our findings that lichen phenolic compounds can sometimes interact with micro-arthropods to influence lichen litter mass loss and nutrient release during decomposition assists our understanding of how lichens and their consumers may impact on organic matter dynamics, biochemical nutrient cycling and other related ecosystem processes.     

Nutrient dynamics in litter mixtures of four Mediterranean maquis species decomposing in situ

In natural conditions, litters shed from different species become mixed with each other, and decompose together. Most studies deal with decomposition of individual species; few studies investigate the influence of litter mixing on decomposition and nutrient dynamics; the results are contradictory as positive, negative, or no effect, of litter mixing have been observed. In this study we test the hypothesis: i) that litter mixing in the Mediterranean maquis, a nutrient poor, high diversity ecosystem, produces non-additive effects on nutrient dynamics; ii) that the effects vary with the composition in species of the mixture and with the relative amount of the species component the mixture. Two types of 3-species mixtures were set up; one contained three sclerophylls, Phillyrea angustifolia, Pistacia lentiscus and Quercus ilex; the other contained the first two species with the mesophyll Cistus. Litterbags, containing monospecific litters and even and uneven mixtures, were incubated under natural condition in situ; even mixtures had the 3 species in equal proportion, whereas uneven mixtures had one of the species as dominant (50%) and the other two species in equal proportion (25%:25%). Litterbags were retrieved after 92, 188 and 403 days; litters from the mixtures were separately analyzed for mass loss and content of nitrogen (N), potassium (K), sodium (Na), calcium (Ca), magnesium (Mg), manganese (Mn), iron (Fe), copper (Cu), and zinc (Zn). Results indicate that mixing influences the dynamics of N, Mn, Ca, Mg, Fe, Cu and Zn, but scarcely affects the dynamics of K and Na. The comparison of observed to expected values for changes of nutrients in litterbags indicates the occurrence of non-additive effects of litter mixing on movements of N, Fe, Cu, and Zn to or from the litterbags containing the mixtures. The effects depend on the composition in species of the mixture, whereas the relative amount of the species component the mixture is not relevant.