Tree Species Effect on Litter Decomposition and Nutrient Release in Mediterranean Oak Forests Changes Over Time

Tree species can affect the decomposition process through the quality of their leaf fall and through the species-specific conditions that they generate in their environment. We compared the relative importance of these effects in a 2-year experiment. Litterbags containing leaf litter of the winter-deciduous Quercus canariensis, the evergreen Q. suber and mixed litter were incubated beneath distinct plant covers. We measured litter carbon loss, 9 macro- and micronutrients and 18 soil chemical, physical and biological parameters of the incubation environment. Tree species affected decay dynamics through their litter quality and, to a lesser extent, through the induced environmental conditions. The deciduous litter showed a faster initial decomposition but left a larger fraction of slow decomposable biomass compared with the perennial litter; in contrast the deciduous environment impeded early decomposition while promoting further carbon loss in the latter decay stages. The interaction of these effects led to a negative litter–environment interaction contradicting the home-field advantage hypothesis. Leaf litter N, Ca and Mn as well as soil N, P and soil moisture were the best predictors for decomposition rates. Litter N and Ca exerted counteractive effects in early versus late decay stages; Mn was the best predictor for the decomposition limit value, that is, the fraction of slowly decomposable biomass at the later stage of decomposition; P and soil moisture showed a constant and positive relation with carbon loss. The deciduous oak litter had a higher initial nutrient content and released its nutrients faster and in a higher proportion than the perennial oak litter, significantly increasing soil fertility beneath its canopy. Our findings provide further insights into the factors that control the early and late stages of the decomposition process and reveal potential mechanisms underlying tree species influence on litter decay rate, carbon accumulation and nutrient cycling.     

Isopod effects on decomposition of litter produced under elevated CO2, N deposition and different soil types

The performance of Oniscus asellus (Isopoda) and its influence on litter mass loss and mineralization was assessed in a microcosm experiment, using beech (Fagus sylvatica) leaf litter that was produced on different soil types, contrasting atmospheric CO₂ concentrations, and different nitrogen deposition rates. Litter quality was significantly altered by these treatments, and many of the CO₂ and N effects differed between soil types. Litter quality affected subsequent litter mass loss rates, microbial respiration, and leaching of dissolved organic carbon (DOC) and nitrate. These effects were largely independent of the presence of isopods, even though isopods highly accelerated litter mass loss, stimulated microbial respiration by 37%, and increased nitrate leaching by 50%. Isopods did not change their relative rates of litter consumption and growth in response to litter quality. Isopod mortality, however, increased with increasing litter lignin/N ratios, and was significantly different between soil types, which may indicate long‐term effects on litter decomposition through altered isopod densities. Having the choice among the litter of three different species [maple (Acer pseudoplatanus), beech (Fagus sylvatica) and oak (Quercus robur)] grown at either ambient or elevated CO₂, isopods preferred maple to beech when all the litter was produced under elevated CO₂. This suggests that beyond changes in consumption, an altered food selection by isopods in a CO₂‐enriched atmosphere may affect the temporal and spatial composition of the litter layer in temperate forests. In contrast to previous findings, we conclude that isopods do not always increase their consumption rates, and hence do not differentially affect microbial decomposition in litter of poorer quality. Nevertheless changes in animal densities and/or shifts in their food preferences, could result in altered decomposition and carbon and nutrient turnover rates.     

Foliage litter turnover and earthworm populations in three beech forests of contrasting soil and vegetation types

Summary Leaf litter decomposition, levels of accumulated litter as well as the abundance and biomass of earthworms were measured in three mature beech forests in southern Sweden: one mor site, one poor mull site, and one rich mull site. The disappearance rate of beech litter, measured with litter bags, increased with increasing soil fertility. On the rich mull site, the disappearance rate was much higher than in the two other forests, due to the combined effects of higher earthworm activity, more favouable soil moisture conditions, and higher litter quality. Incubating the litter in finely meshed bags (1-mm mesh) to exclude macrofauna had a great effect on litter mass loss in the rich mull site, but it had only a minor effect in the other sites. Simultaneous incubations of local and transplanted leaf litter on the three study sites showed that the substrate quality of the litter increased in the order: mor site — poor mull site — rich mull site. Lignin, N, and P concentrations of the leaf litter failed to explain the observed differences in decomposition rates, and acid/base properties are suggested to be more important. Earthworm numbers per m² were 2.5 (1 species) in the mor, 40 (6 species) in the poor mull and 220 (9 species) in the rich mull forest. Soil chemical conditions, notably pH, were suggested as the main factors determining the inter-site differences in abundance and species composition of earthworms. The role of litter decomposition and earthworm activity in the accumulation of organic matter in the forest floor in different types of beech woodlands are discussed.     

Effects of aqueous extracts from leaves and leaf litter on the abundance and diversity of soil gymnamoebae in laboratory microcosm cultures

The distribution and abundance of microbiota in soil and litter may be significantly affected by the quality and quantity of localized patches of leaf organic matter. This study examined the relative effects of aqueous extracts of shed autumn leaves from American beech (Fagus grandifolia), sugar maple (Acer saccharum), red oak (Quercus rubra), and white oak (Quercus alba) on the density and diversity of gymnamoebae in laboratory cultures. Overall, the beech leaf extract produced the most growth of gymnamoebae followed by white oak with leaf extracts from maple and red oak producing least growth. Cultures using natural leaf litter from beneath beech trees had higher densities and diversity of gymnamoebae than leaf-litter cultures from a maple-oak stand. Soil microcosms confirmed that beech leaf extracts produced a higher density of gymnamoeba growth when added to soil cultures compared with maple and oak leaf extracts. Protein content, CHN (carbon and nitrogen content), and pH of the leaf extracts were assayed, but these alone were not sufficiently different to account for the effects. A dilution experiment indicated that some other concentration-dependent factor in the extract may produce the effects.     

地上枯落物的累积、分解及其在陆地生态系统中的作用

了解陆地生态系统地上枯落物的累积和分解过程对认识它的生态作用、通过管理地上枯落物调控陆地生态系统功能和服务有重要意义。综述了陆地生态系统地上枯落物的积累和分解过程及其影响因素,然后概括了通过这些过程地上枯落物所发挥的生态作用,最后,在全球变化背景下,基于当前研究进展提出陆地生态系统地上枯落物研究的前景。地上枯落物累积在时间尺度上一般遵循植物的生命周期,同时也受环境因子的调控。大的空间尺度上,枯落物累积主要受水热因子控制,伴随植被类型的变化,表现随纬度升高而减少的趋势。然而,在局域尺度内,枯落物累积除受水、热因子限制,还被群落结构、土壤条件、植食动物等因素影响,表现较大变异性。当前,人类干扰作为一个不可忽视的因素,正在强烈甚至不可逆转的改变地表植被覆盖和枯落物累积。地上枯落物的分解过程包括淋溶、光降解、土壤动物和微生物分解,这些过程同时进行并相互影响。尽管目前还不清楚,但区分这些分解过程和分解产物的去向对了解陆地生态系统物质循环有重要意义。枯落物分解首先被自身类型、化学组成、物种多样性决定,同时也受分解者群体、非生物环境影响。其中,枯落物分解与其化学特性、物种多样性及土壤养分状况的关系是研究的热点,也是广泛争议的焦点。通过累积和分解,地上枯落物对陆地生态系统有物理、化学、生物作用。目前,枯落物的物理和化学作用研究较为透彻,而由于受枯落物数量、环境条件、响应植物特征或一些有待挖掘的未知因素的共同限制,地上枯落物的生物作用,尤其对植物的作用在不同研究中仍没有达成普遍的共识。全球变化可能影响地上枯落物累积、分解和生态作用。在全球变化的背景,研究地上枯落物产量和性状变化、阐明枯落物分解的分室模型、继续分析枯落物性状和分解关系、深入揭示枯落物的生态作用及其制约因素,理解和预测地上枯落物数量和质量变化对陆地生态系统功能和服务的影响是必要的。     

Long-term Changes in Forest Carbon and Nitrogen Cycling Caused by an Introduced Pest/Pathogen Complex

Invasion of exotic forest pests and pathogens is a serious environmental problem for many forests throughout the world, and has been especially damaging to forests of eastern North America. We studied the impacts of an exotic pest/pathogen complex, the beech bark disease (BBD), in the Catskill Mountains of New York State, USA. In this region, BBD has caused a decline in the basal area of American beech (Fagus grandifolia Ehrh.) over the last 60 years and this decline has been accompanied by an increase in the basal area of sugar maple (Acer saccharum Marsh.). We studied the impacts of the BBD on carbon (C) and nitrogen (N) cycling using a series of stands that represented a sequence of disease impact and beech replacement by sugar maple. Our study showed that these long-term changes in tree species composition can lead to important changes in C and N cycling in the ecosystem, including an increase in litter decomposition, a decrease in soil C:N ratio, and an increase in extractable nitrate in the soil and nitrate in soil solution. Rates of potential net N mineralization and nitrification did not change across the BBD sequence, but the fraction of mineralized N that was nitrified increased significantly. Many of the observed changes in ecosystem function are larger in magnitude than those attributed to climate change or air pollution, suggesting that the impacts of invasive pests and pathogens on tree species composition could be one of the most important factors driving changes in C and N cycling in these forests in the coming decades.     

凋落物分解主场效应及其土壤生物驱动

凋落物分解主场效应是指凋落物具有在其生长的栖息地比在别的生境分解更快的特征,土壤生物的特化作用被认为是主场效应的产生机理.主场效应是除基质质量和物理化学环境外控制凋落物分解的重要因子,可影响模拟精度的8％.凋落物分解主场效应驱动机制的深入研究对促进分解模型中纳入生物因子,提高区域尺度模拟精度具有重要作用.虽然时间和基质质量可导致主场效应强度变化,但不能全面解释主场效应强度差异特别是负效应的产生.通过分析凋落物分解过程中土壤生物的作用机理,指出凋落物分解主场效应的土壤生物驱动可能包括土壤微生物的调节性适应,土壤动物的后期插入以及物理化学环境的间接影响.为深入了解主场效应土壤生物驱动机制,更好地模拟凋落物分解过程,提出延长凋落物分解交互移置实验时间,拓展实验空间,结合室内模拟分析和构建分解模型等方法与途径.     

Correlation of foliage and litter chemistry of sugar maple, Acer saccharum, as affected by elevated CO2 and varying N availability, and effects on decomposition

Rising atmospheric carbon dioxide has the potential to alter leaf litter chemistry, potentially affecting decomposition and rates of carbon and nitrogen cycling in forest ecosystems. This study was conducted to determine whether growth under elevated atmospheric CO₂ altered the quality and microbial decomposition of leaf litter of a widely distributed northern hardwood species at sites of low and high soil nitrogen availability. In addition, we assessed whether the carbon–nutrient balance (CNB) and growth differentiation balance (GDB) hypotheses could be extended to predict changes in litter quality in response to resource availability. Sugar maple (Acer saccharum) was grown in the field in open‐top chambers at 36 and 55 Pa partial pressure CO₂, and initial soil mineralization rates of 45 and 348 μg N g^?1 d^?1. Naturally senesced leaf litter was assessed for chemical composition and incubated in the laboratory for 111 d. Microbial respiration and the production of dissolved organic carbon (DOC) were quantified as estimates of decomposition. Elevated CO₂ and low soil nitrogen resulted in higher litter concentrations of nonstructural carbohydrates and condensed tannins, higher C/N ratios and lower N concentrations. Soil N availability appears to have had a greater effect on litter quality than did atmospheric CO₂, although the treatments were additive, with highest concentrations of nonstructural carbohydrates and condensed tannins occurring under elevated CO₂–low soil N. Rates of microbial respiration and the production of DOC were insensitive to differences in litter quality. In general, concentrations of litter constituents, except for starch, were highly correlated to those in live foliage, and the CNB/GDB hypotheses proved useful in predicting changes in litter quality. We conclude the chemical composition of sugar maple litter will change in the future in response to rising atmospheric CO₂, and that soil N availability will exert a major control. It appears that microbial metabolism will not be directly affected by changes in litter quality, although conclusions regarding decomposition as a whole must consider the entire soil food web.     

An integrated perspective to explain nitrogen mineralization in grazed ecosystems

Large herbivores are key drivers of nutrient cycling in ecosystems worldwide, and hence they have an important influence on the productivity and species composition in plant communities. Classical theories describe that large herbivores can accelerate or decelerate nitrogen (N) mineralization by altering the quality and quantity of resource input (e.g. dung, urine, plant litter) into the soil food web. However, in many situations the impact of herbivores on N mineralization cannot be explained by changes in resource quality and quantity.In this paper, we aim to reconcile observations of herbivores on N mineralization that were previously regarded as contradictory. We conceptually integrate alternative pathways via which herbivores can alter N mineralization. We illustrate our new integrated perspective by using herbivore-induced soil compaction and subsequent changes in soil moisture and soil aeration as an example.We show that the net effect of herbivores on mineralization depends on the balance between herbivore-induced changes in soil physical properties and changes in the quality and quantity of resource input into the soil food web. For example, soil compaction by herbivores can limit oxygen or water availability in wet and dry soils respectively, particularly those with a fine texture. This can result in a reduction in N mineralization regardless of changes in resource quality or quantity. In such systems the plant community will shift towards species that are adapted to waterlogging (anoxia) or drought, respectively. In contrast, soils with intermediate moisture levels are less sensitive to compaction. In these soils, N mineralization rates are primarily associated with changes in resource quality and quantity.We conclude that our integrated perspective will help us to better understand when herbivores accelerate or decelerate soil nutrient cycling and improve our understanding of the functioning of grazed ecosystems.     

Nitrogen supply differentially affects litter decomposition rates and nitrogen dynamics of sub-arctic bog species

High-latitude peatlands are important soil carbon sinks. In these ecosystems, the mineralization of carbon and nitrogen are constrained by low temperatures and low nutrient concentrations in plant litter and soil organic matter. Global warming is predicted to increase soil N availability for plants at high-latitude sites. We applied N fertilizer as an experimental analogue for this increase. In a three-year field experiment we studied N fertilization effects on leaf litter decomposition and N dynamics of the four dominant plant species (comprising >75% of total aboveground biomass) in a sub-arctic bog in northern Sweden. The species were Empetrum nigrum (evergreen shrub), Eriophorum vaginatum (graminoid), Betula nana (deciduous shrub) and Rubus chamaemorus (perennial forb). In the controls, litter mass loss rates increased in the order: Empetrum < Eriophorum < Betula < Rubus. Increased N availability had variable, species-specific effects: litter mass loss rates (expressed per unit litter mass) increased in Empetrum, did not change in Eriophorum and Betula and decreased in Rubus. In the leaf litter from the controls, we measured no or only slight net N mineralization even after three years. In the N-fertilized treatments we found strong net N immobilization, especially in Eriophorum and Betula. This suggests that, probably owing to substantial chemical and/or microbial immobilization, additional N supply does not increase the rate of N cycling for at least the first three years.     

Changes in Clonal Poplar Leaf Chemistry Caused by Stem Galls Alter Herbivory and Leaf Litter Decomposition

Gall-inducing insects are highly specialized herbivores that modify the phenotype of their host plants. Beyond the direct manipulation of plant morphology and physiology in the immediate environment of the gall, there is also evidence of plant-mediated effects of gall-inducing insects on other species of the assemblages and ecosystem processes associated with the host plant. We analysed the impact of gall infestation by the aphid Pemphigus spirothecae on chemical leaf traits of clonal Lombardy poplars (Populus nigra var. italica) and the subsequent effects on intensity of herbivory and decomposition of leaves across five sites. We measured the herbivory of two feeding guilds: leaf-chewing insects that feed on the blade (e.g. caterpillars and sawfly larvae) and skeletonising insects that feed on the mesophyll of the leaves (e.g. larvae of beetles). Galled leaves had higher phenol (35%) and lower nitrogen and cholorophyll contents (35% respectively 37%) than non-galled leaves, and these differences were stronger in August than in June. Total herbivory intensity was 27% higher on galled than on non-galled leaves; damage by leaf chewers was on average 61% higher on gall infested leaves, whereas damage by skeletonising insects was on average 39% higher on non-galled leaves. After nine months the decomposition rate of galled leaf litter was 15% lower than that of non-galled leaf litter presumably because of the lower nitrogen content of the galled leaf litter. This indicated after-life effects of gall infestation on the decomposers. We found no evidence for galling x environment interactions.     

Soil Moisture Alters the Response of Soil Organic Carbon Mineralization to Litter Addition

Increasing rainfall and longer drought conditions lead to frequent changes in soil moisture that affect soil organic carbon (SOC) mineralization. However, how soil moisture affects response of SOC mineralization to litter addition in forest ecosystems remains unexplored. We added ¹³C-labeled litter to subtropical forest soils with three mass water contents (L, 21%; M, 33%; H, 45%). Carbon dioxide production was monitored, and the composition of soil microbial communities was determined by phospholipid fatty acid (PLFA). When no litter was added, SOC mineralization was greater in the M-treated soil. Litter addition promoted SOC mineralization, but this promotion was altered by soil moisture and litter type. Priming effects induced by P. massoniana leaf litter in the M-moistened soil were significantly (P < 0.05) higher than those in other treatments. Litter-derived C was approximately 55% incorporated into 18:1ω9c and 16:0 PLFAs, and this proportion was not significantly affected by soil moisture. Soil moisture affected the distribution of litter-¹³C in i15:0, i17:0, and cy19:0 individual PLFAs. The primed C evolution was significantly related to the ratio of Gram-positive to Gram-negative bacteria. These results suggest that changes in soil moisture could affect SOC mineralization in forest ecosystems.     

Increasing Red Maple Leaf Litter Alters Decomposition Rates and Nitrogen Cycling in Historically Oak-Dominated Forests of the Eastern U.S.

Without canopy-opening fire disturbances, shade-tolerant, fire-sensitive species like red maple (Acer rubrum L.) proliferate in many historically oak-dominated forests of the eastern U.S. Here, we evaluate potential implications of increased red maple dominance in upland oak forests of Kentucky on rates of leaf litter decomposition and nitrogen (N) cycling. Over 5 years, we evaluated mass loss of leaf litter and changes in total N and carbon (C) within six leaf litter treatments comprised of scarlet oak, chestnut oak, and red maple, and three mixed treatments of increasing red maple contribution to the leaf litter pool (25, 50, and 75% red maple). Over a 1.5-year period, we conducted a plot-level leaf litter manipulation study using the same treatments plus a control and assessed changes in net nitrification, ammonification, and N mineralization within leaf litter and upper (0–5 cm depth) mineral soil horizons. Red maple leaf litter contained more “fast” decomposing material and initially lost mass faster than either oak species. All litter treatments immobilized N during initial stages of decomposition, but the degree of immobilization decreased with decreasing red maple contribution. The leaf litter plot-level experiment confirmed slower N mineralization rates for red maple only plots compared to chestnut oak plots. As red maple increases, initial leaf litter decomposition rates will increase, leading to lower fuel loads and more N immobilization from the surrounding environment. These changes may reduce forest flammability and resource availability and promote red maple expansion and thereby the “mesophication” of eastern forests of the U.S.     

Sapling herbivory,invertebrate herbivores and predators across a natural tree diversity gradient in Germany’s largest connected deciduous forest

Tree species-rich forests are hypothesised to be less susceptible to insect herbivores, but so far herbivory–diversity relationships have rarely been tested for tree saplings, and no such study has been published for deciduous forests in Central Europe. We expected that diverse tree communities reduce the probability of detection of host plants and increase abundance of predators, thereby reducing herbivory. We examined levels of herbivory suffered by beech (Fagus sylvatica L.) and maple saplings (Acer pseudoplatanus L. and Acer platanoides L.) across a tree species diversity gradient within Germany’s largest remaining deciduous forest area, and investigated whether simple beech or mixed stands were less prone to damage caused by herbivorous insects. Leaf area loss and the frequency of galls and mines were recorded for 1,040 saplings (>13,000 leaves) in June and August 2006. In addition, relative abundance of predators was assessed to test for potential top-down control. Leaf area loss was generally higher in the two species of maple compared to beech saplings, while only beech showed a decline in damage caused by leaf-chewing herbivores across the tree diversity gradient. No significant patterns were found for galls and mines. Relative abundance of predators on beech showed a seasonal response and increased on species-rich plots in June, suggesting higher biological control. We conclude that, in temperate deciduous forests, herbivory–tree diversity relationships are significant, but are tree species-dependent with bottom-up and top-down control as possible mechanisms. In contrast to maple, beech profits from growing in a neighbourhood of higher tree richness, which implies that species identity effects may be of greater importance than tree diversity effects per se. Hence, herbivory on beech appeared to be mediated bottom-up by resource concentration in the sampled forest stands, as well as regulated top-down through biocontrol by natural enemies. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

Leaf litter fall and soil acidity during half a century of secondary succession in a temperate deciduous forest

Vegetation, leaf litter fall and soil pH were sampled repeatedly within semipermanent plots in a South-Swedish deciduous forest, 1935–1983. Leaf litter fall was summarized in a litter quality index. Vegetation types were differentiated along similar gradients in soil pH and leaf litter quality. The greatest shifts in dominance among field layer species were found in those plots where the quality of the leaf litter had improved. These plots also showed a halt in the general tendency towards a decreasing pH in the top soil.     

Regional patterns and controls of leaf decomposition in Australian tropical rainforests

Future climates have the potential to alter decomposition rates in tropical forest with implications for carbon emissions, nutrient cycling and retention of standing litter. However, our ability to predict impacts, particularly for seasonally wet forests in the old world, is limited by a paucity of data, a limited understanding of the relative importance of different aspects of climate and the extent to which decomposition rates are constrained by factors other than climate (e.g. soil, vegetation composition). We used the litterbag method to determine leaf litter decay rates at 18 sites distributed throughout the Australian wet tropics bioregion over a 14‐month period. Specifically, we investigated regional controls on litter decay including climate, soil and litter chemical quality. We used both in situ litter collected from litterfall on site and a standardized control leaf litter substrate. The control litter removed the effect of litter chemical quality and the in situ study quantified decomposition specific to the site. Decomposition was generally slower than for other tropical rainforests globally except in our wet and nutrient‐richer sites. This is most likely attributable to the higher latitude, often highly seasonal rainfall and very poor soils in our system. Decomposition rates were best explained by a combination of climate, soil and litter quality. For in situ litter (native to the site) this included: average leaf wetness in the dry season (LWDS; i.e. moisture condensation) and the initial P content of the leaves, or LWDS and initial C. For control litter (no litter quality effect) this included: rainfall seasonality (% dry season days with 0‐mm rainfall), soil P and mean annual temperature. These results suggest that the impact of climate change on decomposition rates within Australian tropical rainforests will be critically dependent on the trajectory of dry season moisture inputs over the coming decades.     

Natural and human‐mediated selection in a landrace of Thai rice (Oryza sativa)

Landrace rice in Thailand consists of managed populations grown under traditional and long‐standing agricultural practices. These populations evolve both in response to environmental conditions within the local agro‐ecosystem and in response to human activities. Single landraces are grown across varying environments and recently have experienced temporal changes in local environments due to climate change. Here we assess the interplay between natural selection in a changing climate and human‐mediated selection on the population genetic structure of Muey Nawng, a local landrace of Thai rice. Genetic diversity and population structure of landrace rice were assessed by a STRUCTURE analysis of 20 microsatellite loci. The first exon–intron junction of the waxy gene was sequenced to determine genotypes for glutinous or non‐glutinous grain starch. Muey Nawng rice is genetically variable and is structured based on starch grain types and the level of resistance to gall midge pest. A strong positive correlation was found between genetic diversity and the percentage of gall midge infestation. Variation in the waxy locus is correlated with starch quality; selection for non‐glutinous rice appears to involve additional genes. The dynamics of genetic diversity within Muey Nawng rice depends on three factors: (a) a genetic bottleneck caused by strong selection associated with gall midge infestation, (b) selection by local farmers for starch quality and (c) variation introduced by farmer practices for cultivation and seed exchange. These results, when taken in total, document the ability of landrace rice to quickly evolve in response to both natural and human‐mediated selection.     

Foliage litter quality and annual net N mineralization: comparison across North American forest sites

The feedback between plant litterfall and nutrient cycling processes plays a major role in the regulation of nutrient availability and net primary production in terrestrial ecosystems. While several studies have examined site-specific feedbacks between litter chemistry and nitrogen (N) availability, little is known about the interaction between climate, litter chemistry, and N availability across different ecosystems. We assembled data from several studies spanning a wide range of vegetation, soils, and climatic regimes to examine the relationship between aboveground litter chemistry and annual net N mineralization. Net N mineralization declined strongly and non-linearly as the litter lignin:N ratio increased in forest ecosystems (r ² = 0.74, P < 0.01). Net N mineralization decreased linearly as litter lignin concentration increased, but the relationship was significant (r ² = 0.63, P < 0.01) only for tree species. Litterfall quantity, N concentration, and N content correlated poorly with net N mineralization across this range of sites (r ² < 0.03, P = 0.17–0.26). The relationship between the litter lignin:N ratio and net N mineralization from forest floor and mineral soil was similar. The litter lignin:N ratio explained more of the variation in net N mineralization than climatic factors over a wide range of forest age classes, suggesting that litter quality (lignin:N ratio) may exert more than a proximal control over net N mineralization by influencing soil organic matter quality throughout the soil profile independent of climate. Received: 16 December 1996 / Accepted: 8 February 1997     

Influence of Litter Diversity on Dissolved Organic Matter Release and Soil Carbon Formation in a Mixed Beech Forest

We investigated the effect of leaf litter on below ground carbon export and soil carbon formation in order to understand how litter diversity affects carbon cycling in forest ecosystems. ¹³C labeled and unlabeled leaf litter of beech (Fagus sylvatica) and ash (Fraxinus excelsior), characterized by low and high decomposability, were used in a litter exchange experiment in the Hainich National Park (Thuringia, Germany). Litter was added in pure and mixed treatments with either beech or ash labeled with ¹³C. We collected soil water in 5 cm mineral soil depth below each treatment biweekly and determined dissolved organic carbon (DOC), δ¹³C values and anion contents. In addition, we measured carbon concentrations and δ¹³C values in the organic and mineral soil (collected in 1 cm increments) up to 5 cm soil depth at the end of the experiment. Litter-derived C contributes less than 1% to dissolved organic matter (DOM) collected in 5 cm mineral soil depth. Better decomposable ash litter released significantly more (0.50±0.17%) litter carbon than beech litter (0.17±0.07%). All soil layers held in total around 30% of litter-derived carbon, indicating the large retention potential of litter-derived C in the top soil. Interestingly, in mixed (ash and beech litter) treatments we did not find a higher contribution of better decomposable ash-derived carbon in DOM, O horizon or mineral soil. This suggest that the known selective decomposition of better decomposable litter by soil fauna has no or only minor effects on the release and formation of litter-derived DOM and soil organic matter. Overall our experiment showed that 1) litter-derived carbon is of low importance for dissolved organic carbon release and 2) litter of higher decomposability is faster decomposed, but litter diversity does not influence the carbon flow.     

Earthworms,Collembola and residue management change wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis>) and herbivore pest performance (Aphidina: <Emphasis Type="Italic">Rhophalosiphum padi</Emphasis>)

Management practices of arable systems determine the distribution of soil organic matter thereby changing decomposer animal activity and their impact on nutrient mineralization, plant growth and plant-herbivore interactions. Decomposer-mediated changes in plant growth and insect pest performance were investigated in wheat-aphid model systems in the greenhouse. Three types of litter distribution were established: litter patch at the soil surface (simulating mulching), litter patch deeper in soil (simulating ploughing) and litter homogeneously mixed into soil (simulating disk cultivation). The litter was labelled with (15)N to follow the mineralization and uptake of nutrients by the plants. Earthworms (Aporrectodea caliginosa) and Collembola (Protaphorura armata) were included as representatives of major functional groups of decomposers. Wheat (Triticum aestivum) was planted and aphids (Rhophalosiphum padi) were introduced to leaves as one of the most important pests. Earthworms, Collembola and litter distribution affected plant growth, N acquisition and aphid development in an interactive way. Earthworms and Collembola increased biomass of seeds, shoots and roots of wheat. Increased plant growth by earthworms and Collembola was mainly due to increased transfer of N from soil (rather than litter) into plants. Despite increasing plant growth, earthworms reduced aphid reproduction. Aphid reproduction was not correlated closely with plant N concentrations, but rather with the concentration of litter N in wheat. Unexpectedly, both Collembola and earthworms predominantly affected the mobilization of N from soil organic matter, and by altering the distribution of litter earthworms reduced infestation of crops by aphids via reducing plant capture of litter N, in particular if the litter was concentrated deeper in soil. The results suggest that management practices stimulating a continuous moderate increase in nutrient mobilization from soil organic matter rather than nutrient flushes from decomposing fresh organic matter result in maximum plant growth with minimum plant pest infestation.