Soil characteristics determine soil carbon and nitrogen availability during leaf litter decomposition regardless of litter quality

Climate and litter quality have been identified as major drivers of litter decomposition, but our knowledge of how soil characteristics (e.g. microbial community and chemical properties) determine carbon (C) and nitrogen (N) availability derived from the decomposition of litter of different qualities is still scarce. We conducted a microcosm experiment to evaluate how soils with contrasting microbial communities and soil properties (denoted Soils A and B hereafter, where Soil B has higher bacterial and fungal abundance, fungal:bacterial ratio, and organic C than Soil A) determine the availability of soil C (carbohydrates, proteins, amino acids and phenols) and N (dissolved organic and inorganic N, microbial biomass N and available N) during the decomposition of litter of contrasting quality (C:N ratios ranging from 20 to 102). We also evaluated the relative importance of soil characteristics and litter quality as drivers of C and N inputs to the soil during this process. Overall, higher soil C and N availability after litter decomposition was found in Soil B than in Soil A. Soil characteristics had a higher positive effect on soil C and N contents than litter quality during litter decomposition. We also found that changes in N availability and organic matter quality registered after litter decomposition, linked to different soil characteristics, were able to promote dissimilarities in the potential mineralization rates. In conclusion, our study provides evidence that soil characteristics (e.g. microbial communities and chemical properties) can be more important than litter quality in determining soil C and equally important for N availability during the decomposition of leaf litter.     

Leaf decomposition in two semi-evergreen tropical forests: influence of litter quality

Decomposition processes in tropical semi-evergreen forests are still poorly understood. The influence of soil properties and litter quality on decomposition rate was studied in two semi-evergreen forests of Guadeloupe, a forest plantation and a secondary forest, located on different soils. Leaf litter of four tree species was enclosed in litterbags for a 14-month period. Non-linear correlations were calculated between mass loss and the concentration of major leaf components (soluble C, N, lignin, cellulose, tannins, total soluble phenols) in order to determine the best predictor of leaf litter decomposition. Soil physico-chemical properties and ratios between some of the above-mentioned litter quality parameters were also examined as mass loss predictors. In addition, non-linear correlations were calculated between mass loss and litter quality parameters, at successive periods. Litter quality was the main determinant of litter decomposition in the studied forests. Several litter quality parameters were correlated with leaf disappearance, varying according to stages of decomposition. Between 1 month and 2.5 months, the mass loss was correlated negatively with the initial phenol content and with initial lignin:N and (lignin+phenol):N ratios. From 2.5 to 5.5 months, the mass loss was correlated negatively with the initial phenol content and positively with the initial cellulose content. At later stages of decomposition (9-14 months), the mass loss was correlated negatively with the initial tannin content. Differences in soil characteristics and fauna did not seem to be enough to affect decomposition.     

川西3种亚高山针叶林的养分和凋落物格局分析

Investigations were conducted to quantify litterfall, and litter and nutrient accumulation in forest floor, and to acquire information on litter decomposition and nitrogen and phosphorus release patterns in three different subalpine coniferous forests, a plantation （P1）, a secondary forest （SF）, and a primitive forest （PF）, in western Sichuan, China. The litter trap method was used to evaluate litterfall with the litterbag method being utilized for litter decomposition. Seasonal patterns of litterfall were similar in the three forests, with two peaks occurring in September-November and March-May. The plantation revealed an annual litterfall of 4.38 x 103 kg ha-1, which was similar to those of SF and PF, but P1 had a lower mass loss rate and a higher C/N ratio. The C/N ratio may be a sound predictor for the decomposition differences. N concentrations of leaf litter in both the secondary forest and primitive forest increased first and then decreased, and the percentages of their final/initial values were 108.9% and 99.9%, respectively. P concentration in the three forests increased by the end of the study. The results of litterfall and decomposition indicated that in the plantation the potential to provide nutrients for soil organic matter was similar to those of SF and PF; however, its slower decomposition rate could result in a somewhat transient accumulation of litter in the forest floor.     

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.     

Effects of sulfuric,nitric, and mixed acid rain on litter decomposition,soil microbial biomass,and enzyme activities in subtropical forests of China

Acid rain pollution is changing gradually from sulfuric acid rain (SAR) to mixed acid rain (MAR) and then to nitric acid rain (NAR) with the rapidly growing number of motor vehicles. The influences of changed acid rain types on ecosystem functions, particularly on litter decomposition, remain unclear. Two dominant litter types from a coniferous forest and a broad-leaved forest were incubated in microcosms with original forest soils and treated by five types of acid rain with different SO₄²⁻ to NO₃⁻ ratios (1:0, 5:1, 1:1, 1:5, and 0:1). During a six-month incubation period, litter mass losses, soil microbial biomass, and enzyme activities were investigated. Results showed that various acid treatments inhibited litter decomposition, soil microbial biomass, and most enzyme activities, and the inhibitory effects of NAR were more significant than those of SAR and MAR. The resistance to external acid of microbial communities in broad-leaved forest was higher than that in coniferous forest. NAR and MAR treatments slowed down soil carbon (C), nitrogen (N), and phosphorus (P) mineralization by attenuating the correlations between litter mass losses and the enzymes involved in C, N, and P cycling. Results reveal that the ratio of SO₄²⁻ to NO₃⁻ in acid rain is an important factor which profoundly influences litter decomposition process. In the future, a decreasing ratio of SO₄²⁻ to NO₃⁻ in acid rain will be observed in subtropical forests. Thus, soil C would accumulate as a consequence of future acid precipitation, and this may seriously affect the balance of ecosystem C, N flux.     

Effect of earthworm addition on soil nitrogen availability,microbial biomass and litter decomposition in mesocosms

The aim of the study was to determine the effect of adding two tropical earthworm species, Rhinodrilus contortus and Pontoscolex corethrurus, to mesocosms on the availability of mineral N (NH₄ ⁺ and NO₃ ^– concentrations), soil microbial biomass (bio-N), and the decomposition rates of three contrasting leaf litter species, in a glasshouse experiment. The mesocosms were filled with forest soil and covered with a layer of leaf litter differing in nutritional quality: (1) Hevea brasiliensis (C/N=27); (2) Carapa guianensis (C/N=32); (3) Vismia sp., the dominant tree species in the second growth forest (control, C/N= 42); and, (4) a mixture of the former three leaf species, in equal proportions (C/N=34). At the end of the 97-day experiment, the soil mineral N concentrations, bio-N, and leaf litter weight loss were determined. Both earthworm species showed significant effects on the concentrations of soil NO₃ ^– (p<0.01) and NH₄ ⁺ (p<0.05). Bio-N was always greater in the mesocosms with earthworms (especially with R. contortus) and in the mesocosms with leaf litter of H. brasiliensis (6 µg N g^–1 soil), the faster decomposing species, than in the other treatments (0.1–1.6 µg N g^–1). Thus, earthworm activity increased soil mineral-N concentrations, possibly due to the consumption of soil microbial biomass, which can speed turnover and mineralization of microbial tissues. No significant differences in decomposition rate were found between the mesocosms with and without earthworms, suggesting that experiments lasting longer are needed to determine the effect of earthworms on litter decomposition rates.     

Effects of soil microarthropods on plant litter decomposition across an elevation gradient in the Wuyi Mountains

Soil microarthropods are considered to be among the strongest determinants of plant litter decomposition in warm, humid sites. Little is known, however, about the regulation of plant litter decomposition dynamics along an elevation gradient in such sites. Our study examined the contributions of soil microarthropods to leaf litter decomposition of a single substrate (Castanopsis carlesii) along an elevation gradient across four types of zonal vegetations in southeastern China: evergreen broadleaf forest (EVB): coniferous forest (COF): dwarf forest (DWF): and alpine meadow (ALM) during April 2007 to March 2008. Leaf litter decomposition of C. carlesii was significantly accelerated by the presence of fauna in all four sites. After 360 days in the field, mass loss rates with the full decomposer assemblage and in the reduction of microarthropods were 62.9% and 41.2% in EVB, 48.1% and 30.6% in COF, 36.4% and 27.8% in DWF, 30.3% and 23.7% in ALM, respectively. The percentage of total decomposition due to the presence of soil fauna was 37% in EVB, 25% in COF, 12% in DWF, and 8% in ALM, thus showing strong systematic variation along the elevational gradient. The mass losses in control plots at the four sites were significantly correlated with the abundances of total Acari, Collembolans, and Mesostigmata mites. Although the proportion of Oribatid mites at EVB was not the highest among the four sites, there were elevated proportions of Mesostigmatid and Prostigmata mites, many of which were microbe-consuming species and induce an indirect influence on litter decomposition. Moreover, Shannon Index (F = 2.455, p = 0.093) and Group Number (F = 5.830, p = 0.005) both decreased along the elevation gradients. Mass losses were also found to be distinctively related to H′ (r² = 0.984, p = 0.016), and GN (r² = 0.952, p = 0.048) across the four sites. Our results suggest that the faunal contribution to plant litter decomposition varies markedly across environmental gradients that differ in litter faunal diversity.     

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.     

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.     

Organic matter and nutrient dynamics of the litter layer on a forest Rendzina under beech

Summary The decomposition of beech (Fagus sylvatica L.) leaf litter was investigated in a calcareous beech forest using mesh cages containing two layers, fresh leaf litter (O layer), and partly decomposed leaf litter (F layer). C loss was monitored, together with the changes in the contents of total N, hexosamines, ash, Na, K, Mg, Ca, Fe, Mn, Al, Cl, Sulphate, and Phosphate.In 1-mm mesh cages, which excluded access to the macrofauna, the mean annual loss rates for C were 28% in the O leaf litter and 17% in the F leaf litter, totalling approximately 23% for the two layers. The mean loss rates from the 12-mm mesh cages were 54% in the O leaf litter and 58% in the F leaf litter. Degradation processes and feeding activities caused increased contents of ash, total N, and hexosamines in the O layer of both treatments. This increase was greater for the ash and smaller for N, glucosamine, and galactosamine in the 12-mm mesh cages. The sum of ions (Na+K+Mg+Ca+Fe+Mn+Al+Cl+SO₄+PO₄) and also the contents of most single ions were not markedly affected, despite the much higher ash content in the O leaf litter of the 12-mm mesh cages. The ash content increased mainly as a consequence of contamination by soil, which increased the contents of Fe and Al in the ash.     

Soil communities and plant litter decomposition as influenced by forest debris: Variation across tropical riparian and upland sites

Forest debris on ground surface can interact with soil biota and consequently change ecosystem processes across heterogeneous landscape. We examined the interactions between forest debris and litter decomposition in riparian and upland sites within a tropical wet forest. Our experiment included control and debris-removal treatments. Debris-removal reduced leaf litter decomposition rates in both the riparian and upland sites. Debris-removal also reduced soil microbial biomass C in the upland site, but had no effect on microbial biomass C in the riparian site. In contrast, debris-removal altered the density of selected arthropod groups in the riparian site. Litter decomposition rates correlated with both soil microbial biomass and the density of millipedes in a multiple stepwise regression model. Removal of forest debris can substantially reduce rates of leaf litter decomposition through suppressing soil activities. This influence can be further modified by landscape position. Forest debris plays an essential role in maintaining soil activities and ecosystem functioning in this tropical wet forest.     

Interactions between litter quality, decomposition and soil fertility: a laboratory study

Leaf litters from beech (Fagus sylvatica L.) and oak (Quercus robur L.) trees were collected from mixed, deciduous woodlands growing on three soil types that varied in mineral nutrient concentrations and N mineralisation potential. Litter quality, including %N, %Mn, %P, acid detergent fibre, cellulose, Klason lignin, phenylpropanoid constituents of lignin, hexose and pentose sugar (mainly from hemicelluloses) varied within species according to soil type. However, oak and beech showed the opposite responses to soil nutrient status for most of these variables. The litters were incubated in the laboratory for 12 months (at 18 °C and constant moisture) on beds of forest floor material from two soils of contrasting high nutrient material (HNM) or low nutrient material (LNM) nutrient status to investigate litter quality and substrate interactions. At 4, 8 and 12 months there were significant differences in mass losses from oak and beech litters from all sites, and for each litter type exposed to the HNM and LMN soils. At 12 months mean mass losses were higher for HNM treatment (38.7% oak, 27.8% beech) than for the LNM treatment (30.6% oak, 25.5% beech). However, the beech and oak litters from the different sites consistently responded in opposite ways on the same soil treatment reflecting site-related effects on litter quality. Initial concentration of Klason lignin was the best predictor for mass losses from litter species and litter types. Intra-specific variation in rates of litter decomposition of beech and oak litters from different sites, and differences in their interactions with the two forest floor materials, illustrate the complexities of proximate controls on decomposition that are often masked in system-level studies.     

Litter decomposition and nitrogen mineralization in newly formed gaps in a Danish beech (Fagus sylvatica) forest

Gap formation is suggested as an alternative forest management approach to avoid extreme changes in the N cycle of forest ecosystems caused by traditional management practises. The present study aimed to investigate the effect of gap formation on N availability in beech litter and mineral soil on sites, which experienced only little soil disturbance during tree harvest. N pools, litter decomposition, and N mineralization rates in mineral soil were studied in two gaps (17 and 30 m in diameter) in a 75-year-old managed European beech (Fagus sylvatica L.) forest in Denmark and related to soil temperature (5 cm depth) and soil moisture (15 cm depth). Investigations were carried out during the first 2 years after gap formation in measurement plots located along the north-south transect running through the centre of each gap and into the surrounding forest.An effect of gap size was found only for soil temperatures and litter mass loss: soil temperatures were significantly increased in the northern part of the large gap during the first year after gap formation, and litter mass loss was significantly higher in the smaller gap. All other parameters investigated revealed no effect of gap size. Nitrification, net mineralization, and soil N concentrations tended to be increased in the gaps. Cumulative rates of net mineralization were two fold higher in the gaps during the growing season (June-October), but a statistically significant increase was found only for soil NH₄-N concentrations during this period. Forest floor parameters (C:N ratios, mass loss, N release) were not significantly modified during the first year after gap formation, neither were the total C content nor the C:N ratio in mineral soil at 0-10 cm depth.     

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.     

Leaf litterfall and decomposition of different above- and belowground parts of birch (Betula ermanii) trees and dwarf bamboo (Sasa kurilensis) shrubs in a young secondary forest in Northern Japan

In many Japanese forests, the forest understory is largely dominated by dwarf bamboo (Sasa) species, which compete with overstory vegetation for soil nutrients. We studied the rate of leaf litterfall, and decomposition and mineralization of carbon (C) and nitrogen (N) from various components (leaf, root, wood, and rhizome) of overstory and understory vegetation in a young Betula ermanii forest from 2002 to 2004. Total litterfall was 377 g m⁻² year⁻¹, of which the overstory vegetation contributed about two thirds. A litter decomposition experiment conducted for 770 days indicated that mass loss of different litter components varied significantly, except for Sasa kurilensis wood and rhizome. Relative decomposition rates were significantly greater in the first growth period (June to October) than the dormant period (November to May) in most cases. Rainfall was the most important abiotic variable, explaining 75–80% of the variability in mass loss rates. Concentrations of ethanol soluble substances and N were significantly positively correlated (r=0.77 to 0.97, P<0.05) with mass loss at an early stage (41 days). The ratios of lignin/N and C/N were found to be negatively correlated with mass loss rates at all stages of litter decomposition. C stock loss was similar to that of mass loss, whereas N stock loss was slower, except for S. kurilensis fine root litter. The evergreen understory species S. kurilensis exhibited greater N use efficiency than B. ermanii, suggesting better competitive ability that might favor the production of a high biomass and invasion under tree species like B. ermanii.     

杨树刺槐混交林枯落物分解速率的研究

试验研究杨树纯林、刺槐纯林以及杨树刺槐混交林枯落物分解速率结果表明 ,枯落物分解速率随土壤水分的增大而增加 ,土壤含水量达最大持水量时其分解速率达最大值。起始分解速率以 70 %田间持水量为最大 ,其达最大分解速率的时间为 4d ,其他土壤含水量起始分解存在滞后现象 ,达最大分解速率的时间为 7d左右。不同枯落物分解速率为刺槐纯林 >杨树刺槐混交林 >杨树纯林 ,其原因是由刺槐纯林枯落物C N值较高 ,而杨树纯林枯落物C N值较小缘故所致。刺槐与杨树混交可降低枯落物C N值 ,加快枯落物分解速率     

Fine litter chemistry, early-stage decay, and nitrogen dynamics under plantations and primary forest in Lowland Amazonia

One year field exposures of leaf litter from replicated plots of Pinus caribaea var. hondurensis Barrett and Golfari, Carapa guianensis Aubl., Euxylophora paraensis Hub., a Leguminosae combination (Dalbergia nigra Fr. All., Dinizia excelsa Ducke, Parkia multijuga Benth.), and adjacent upland (terra firme) forest at the Curuá-Una Forest Reserve, Pará, Brazil were used to examine the factors controlling leaf litter decay and N dynamics in a lowland tropical environment. Initial leaf litter N concentrations ranged from 4.4 (P. caribaea) to 16.3 mg g⁻¹ dry matter (Leguminosae), and initial lignin concentrations from 190.8 (Leguminosae) to 459.3 mg g⁻¹ dry matter (forest). Pinus caribaea leaf litter lost the least mass (28%), and the Leguminosae leaf litter the most (61%), during the year long incubations. Initial and 1-y proximate C fractions, N concentrations and polyphenol concentrations were not related to mass loss. Annual N accumulation or depletion from leaf litter under the plantations and forest was related to C loss (R²=0.93, P=0.007) and holocellulose loss (R²=0.84, P=0.02). When leaf litter was placed outside its stand of origin, there was a significant location effect on decay rates, indicating that differences in the physical and biological microenvironments under the monospecific plots affected litter decomposition.     

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.     

How browsing by red deer impacts on litter decomposition in a native regenerating woodland in the Highlands of Scotland

Herbivores can indirectly affect ecosystem productivity and processes such as nutrient cycling and decomposition by altering the quantity and quality of resource inputs into the decomposer subsystem. Here, we tested how browsing by red deer impacts on the decomposition of, and nutrient loss from, birch leaf litter (Betula pubescens), and tested whether effects of browsing on these measures were direct, via alteration of the quality of leaf litter, or indirect through long term impacts of deer browsing on soil biological properties. This was tested in a microcosm experiment using soil and litter taken from inside and outside three individual fenced exclosures located at Creag Meagaidh National Nature Reserve, Scotland. We found that litter of un-browsed trees decomposed faster than that from browsed trees, irrespective of whether soil was sourced from inside or outside exclosures. These findings suggest that effects of browsing on litter quality, rather than on soil biological properties, are the key determinant of enhanced decomposition in un-browsed areas of this ecosystem. Despite this, we found no consistent impact of browsing on litter C:N, a key indicator of litter quality; however, the rate of litter decomposition was linearly and negatively related to litter C:N when analysed across all the sites, indicating that this measure, in part, contributed to variation in rates of decomposition in this ecosystem. Our findings indicate that herbivores impact negatively on rates of decomposition in this ecosystem, ultimately retarding nutrient cycling rates, and that these effects are, in part, related to changes in litter quality.