Plant community impacts on the structure of earthworm communities depend on season and change with time

Declining plant diversity potentially threatens essential ecosystem functions driven by the decomposer community, such as litter decomposition and nutrient cycling. Currently, there is no consensus on the interrelationships between plant diversity and decomposer performance and previous studies highlighted the urgent need for long-term experiments.In the Jena Experiment we investigated the long-term impacts of plant community characteristics on the structure of earthworm communities representing key decomposers in temperate grassland. We repeatedly sampled plots varying in plant species richness (1-16 species), plant functional group richness (1-4 groups), and presence of certain plant functional groups (grasses and legumes) three, four, and six years after establishment of the experiment in spring and autumn.The results show that earthworm performance is essentially driven by the presence of certain plant functional groups via a variety of mechanisms. Plant productivity (root biomass) explained most of the detrimental grass impacts (decrease in earthworm performance), while beneficial legume effects likely were linked to high quality inputs of plant residues (increase in earthworm performance). These impacts depended on the functional group of earthworms with the strongest effects on surface feeding anecic earthworms and minor effects on soil feeding endogeic species. Remarkably, effects of plant community characteristics on the composition and age structure of earthworm communities varied between seasons. Moreover, plant diversity effects reported by a former study decreased and detrimental effects of grasses increased with time.The results indicate that plant community characteristics, such as declining diversity, indeed affect the structure of earthworm communities; however, loss of key plant functional groups is likely to be more important than plant species number per se. However, in frequently disturbed ecosystems plant species richness might be important for the recovery and resilience of belowground functions. Moreover, the results accentuate the importance of long-term repeated measurements to fully appreciate the impacts of plant community composition and diversity on ecosystem properties. Single point observations may be misleading and potentially mask the complexity of above-belowground interrelationships.     

Plant diversity enhances the reliability of belowground processes

Theory predicts that the probability that an ecosystem will provide a consistent level of functioning over a given unit of time, i.e. the reliability of ecosystem processes, should increase with species richness. There is growing evidence that plant diversity increases the temporal stability of productivity, but only a few studies have investigated its impact on the reliability of ecosystem processes, and information on whether this propagates to the belowground system is virtually lacking. Using a microcosm experiment with plant communities varying in species and functional group diversity and two decomposer groups (earthworms and Collembola) we investigated the effects of plant diversity on the reliability of the belowground system and vice versa, the effect of decomposers on the reliability of plant productivity. Generally, plant diversity increased the reliability of above- and belowground processes by significantly increasing the reliability of eight out of eleven measured ecosystem parameters (six out of nine belowground responses). Plant functional group diversity had a stronger stabilising effect than species richness on above- and belowground processes including plant shoot and total biomass, microbial basal respiration and Collembola densities. By contrast, in the presence of both decomposer groups the reliability of decomposer populations was reduced. The results indicate that plant diversity effects propagate into the belowground system and increase the reliability of belowground processes via more consistent plant derived belowground inputs.     

Interactive effects of warming, soil humidity and plant diversity on litter decomposition and microbial activity

Human activity has induced a multitude of global changes that are likely to affect the functioning of ecosystems. Although these changes act in concert, studies on interactive effects are scarce. Here, we conducted a laboratory microcosm experiment to explore the impacts of temperature (9, 12 and 15 °C), changes in soil humidity (moist, dry) and plant diversity (1, 4, 16 species) on soil microbial activity and litter decomposition.We found that changes in litter decomposition did not mirror impacts on microbial measures indicating that the duration of the experiment (22 weeks) may not have been sufficient to determine the full magnitude of global change effects. However and notably, changes in temperature, humidity and plant litter diversity/composition affected in a non-additive way the microbial parameters investigated. For instance, microbial metabolic efficiency increased with plant diversity in the high moisture treatment but remained unaffected in low moisture treatment suggesting that climate changes may mask beneficial effects of biodiversity on ecosystem functioning. Moreover, litter decomposition was unaffected by plant litter diversity/composition but increased with increasing temperature in the high moisture treatment, and decreased with increasing temperature in the low moisture treatment.We conclude that it is inevitable to perform complex experiments considering multiple global change agents in order to realistically predict future changes in ecosystem functioning. Non-additive interactions highlight the context-dependency of impacts of single global change agents.     

Plant species richness leaves a legacy of enhanced root litter-induced decomposition in soil

Increasing plant species richness generally enhances plant biomass production, which may enhance accumulation of carbon (C) in soil. However, the net change in soil C also depends on the effect of plant diversity on C loss through decomposition of organic matter. Plant diversity can affect organic matter decomposition via changes in litter species diversity and composition, and via alteration of abiotic and/or biotic attributes of the soil (soil legacy effect). Previous studies examined the two effects on decomposition rates separately, and do therefore not elucidate the relative importance of the two effects, and their potential interaction. Here we separated the effects of litter mixing and litter identity from the soil legacy effect by conducting a factorial laboratory experiment where two fresh single root litters and their mixture were mixed with soils previously cultivated with single plant species or mixtures of two or four species. We found no evidence for litter-mixing effects. In contrast, root litter-induced CO₂ production was greater in soils from high diversity plots than in soils from monocultures, regardless of the type of root litter added. Soil microbial PLFA biomass and composition at the onset of the experiment was unaffected by plant species richness, whereas soil potential nitrogen (N) mineralization rate increased with plant species richness. Our results indicate that the soil legacy effect may be explained by changes in soil N availability. There was no effect of plant species richness on decomposition of a recalcitrant substrate (compost). This suggests that the soil legacy effect predominantly acted on the decomposition of labile organic matter. We thus demonstrated that plant species richness enhances root litter-induced soil respiration via a soil legacy effect but not via a litter-mixing effect. This implies that the positive impacts of species richness on soil C sequestration may be weakened by accelerated organic matter decomposition.     

Above- and below-ground plant inputs both fuel soil food webs

Soil food webs depend almost exclusively on plant derived resources; however, it is still subject to debate how plants affect soil biota. We tested the effects on soil decomposers of three components of soil inputs of plant species identity: presence of live plants (representing rhizodeposits), identity of shoot litter input and identity of root litter input; using all combinations of these for Trifolium pratense and Plantago lanceolata. We assessed impacts on soil microorganisms, Collembola, Oribatida and earthworms in a full-factorial greenhouse experiment. Species identity of shoot litter input had greatest effect on decomposers, following by species identity of live plant. Microbial carbon use efficiency and Oribatida density were significantly higher in the presence of T. pratense shoot litter input than in that of P. lanceolata shoot litter input, while earthworm body mass ratio was significantly higher in the presence of P. lanceolata plants than in that of T. pratense plants. Oribatida density was at minimum in the presence of P. lanceolata plants, shoot and root litter input, resulting in a significant three-way interaction and pointing to the relevance of all investigated plant input pathways. Live plant identity effects were not due to differences in living root biomass among species and treatments. Detrimental P. lanceolata effects may have been due to significantly lower N concentrations than in T. pratense tissue. Our results indicate that both above- and below-ground plant inputs into soil determine the performance of decomposers, and thus suggest due consideration of both types of inputs fueling soil food webs in future studies.     

Collembola switch diet in presence of plant roots thereby functioning as herbivores

Collembola are abundant and ubiquitous soil decomposers, being particularly active in the rhizosphere of plants where they are assumed to be attracted by high microbial activity and biomass. While feeding on root associated microorganisms or organic matter they may also ingest plant roots, e.g. particularly root hairs and fine roots. Employing stable isotope analysis we investigated Collembola (Protaphorura fimata Gisin) feeding preferences and types of ingested resources. We offered Collembola two resources with distinct isotope signatures: a C4 plant (Zea mays L.) planted in soil mixed with ¹⁵N labelled litter of Lolium perenne L. (C3 plant). We hypothesised that Collembola obtain their nutrients (C and N) from different resources, with their carbon being mainly derived from resources that are closely associated to the plant root, e.g. root exudates, causing enrichment in ¹³C in Collembola tissue, while the incorporated nitrogen originating from litter resources. In contrast to our hypothesis, stable isotope analysis suggests that in absence of plant roots Collembola derived both the incorporated C and N predominantly from litter whereas in presence of plant roots they switched diet and obtained both C and N almost exclusively from plant roots.The results indicate that Collembola in the rhizosphere of plants, being assumed to be mainly decomposers, in fact predominately live on plant resources, presumably fine roots or root hairs, i.e. are herbivorous rather than detritivorous or fungivorous. These findings have major implications on the view how plants respond to decomposers in the rhizosphere.     

Management intensity affects traits of soil microarthropod community in montane spruce forest

This study examined the influence of forest management intensity (3 unmanaged, 3 mild managed, 5 intensively managed stands) on soil microarthropods in montane spruce forest. We particularly focused on Oribatida and Collembola which play important roles in organic matter decomposition and nutrient cycling. Our results showed a significant shift from fungivory and carnivory to detritivory in the Oribatida community accompanying management intensification. Similarly, parthenogenetic oribatid mite species contributed more to the community in intensively managed forests and the presence of Collembola species with developed furca increased with management intensification. Although there was no remarkable influence of management intensity on total densities or diversity indices, important and significant shifts in species composition and functional groups showed that soil functions and processes were affected by forest management. Trait assessment indicates a shift in roles Oribatida play in decomposition; fragmentation and comminuting of undecomposed litter seems to gain importance in the intensively managed forest, whereas fungivorous species affect primary decomposers through feeding on fungi in the unmanaged forest.     

Effects of the annual invasive plant Impatiens glandulifera on the Collembola and Acari communities in a deciduous forest

Invasive plants can disturb interactions between soil organisms and native plants and thereby alter ecosystem functions and/or reduce local biodiversity. Collembola and Acari are the most abundant microarthropods in the leaf litter and soil playing a key role in the decomposition of organic material and nutrient cycling. We designed a field experiment to examine the potential effects of the annual invasive plant Impatiens glandulifera on species diversity, abundance and community composition of Collembola and Acari in leaf litter and soil in a deciduous forest in Switzerland. Leaf litter and soil samples were obtained from plots invaded by I. glandulifera since 6 years, from plots in which the invasive plant had been removed for 4 years and from plots which were not yet colonized by the invasive plant. The 45 leaf litter and soil samples were equally distributed over three forest areas, which were differently affected by a wind throw 12 years prior to sampling representing a natural gradient of disturbance. Collembola species richness and abundance in the leaf litter and soil samples were not affected by the presence of the invasive plant. However, the species composition of Collembola was altered in plots with I. glandulifera. The abundance of leaf-litter dwelling Acari was increased in invaded plots compared to the two other plot types. Furthermore, the presence of the invasive plant shifted the composition of Acari individuals belonging to different groups. Our field experiment demonstrates that an annual invasive plant can affect microarthropods which are important for nutrient cycling in various ecosystems.     

Legumes Functional Group Promotes Soil Organic Carbon and Nitrogen Storage by Increasing Plant Diversity

Community composition strongly affected the soil C and N storages. However, the influences of community composition on native grassland remain poorly understood. The purpose of this study is to investigate the ability of plant communities including how legumes affect the soil C and N storages in the semi‐arid grassland. Experimental grassland communities were separated by whether or not containing legumes. We measured soil C and N storages and determined above‐ground and below‐ground biomass, litter biomass, plant species richness, and species diversity to understand the mechanisms underlying the changes of soil C and N storages and to determine the relationship of species diversity and productivity. The results showed that legumes increased above‐ground and below‐ground biomass and C and N storages. Soil C and N storages were significantly and positively related to above‐ground and below‐ground biomass, litter biomass, plant species richness, and diversity in the presence of legumes. The presence of legumes increased soil C and N simultaneously but not synchronously, which resulting in a higher C:N ratio. This study indicated that legumes increased soil C and N storages possibly through increasing biomass and soil C and N inputs. The increases are mediated by plant diversity and plant functional complementarity. We suggest that the combination of legumes‐grass species may greatly enhance ecosystem services such as soil C and N storages, productivity, and diversity in semi‐arid grassland. Copyright © 2016 John Wiley & Sons, Ltd.     

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.     

Lack of home-field advantage in the decomposition of leaf litter in the Atlantic Rainforest of Brazil

Experiments using litter monocultures have indicated that litter decomposes faster on its home site owing to specialised decomposers leading to a home-field advantage (HFA). However, most natural forests, in particular tropical rainforests, harbour more than one species of trees, all of which contribute to the local litter layer. Since interactions among different litter types that cause non-additive decomposition dynamics may prevent HFA, the occurrence of HFA in such multispecies ecosystems is still a matter of debate. Here we studied whether there is an HFA in a highly diverse forest ecosystem in the Atlantic Rainforest of Brazil. We used a litter decomposition experiment using natural litter mixtures with reciprocal transfers among three forest successional stages that differed in their tree species composition and general litter quality. We also investigated the role of soil macro- and meso-invertebrates for HFA and their relative importance along a successional gradient. Results of various statistical procedures failed to demonstrate HFA. A reason for this lack of a HFA may be rapid shifts in the composition of local microbial communities in response to local litter quality. Our experiments indicate a rapid resilience of the microbial decomposition during forest regeneration.     

The effects of earthworm functional group diversity on earthworm community structure

A comprehensive understanding of how species/functional group interactions determine population dynamics, community composition and their effect on ecosystem functioning is necessary if we are to understand the consequences of species loss. This paper presents data from a mesocosm experiment, based on the simplex design, to examine the effect of interactions between earthworm functional groups, food supply and initial overall biomass on community structure. All communities containing anecic species moved towards domination by anecics. The survival of anecics remained constant irrespective of initial conditions, with no effect of initial community structure, food supply or initial biomass. The proportional biomass of epigeics increased when they were placed in communities dominated by anecics. Initial overall biomass had a significant effect on the survival of endogeics, with increased survival at low biomass. Juvenile production was significantly increased in communities that contained a higher initial abundance of epigeics. The anecics had significantly increased production of juveniles at lower levels of initial biomass. Overall, earthworm functional group diversity had an idiosyncratic effect on earthworm assemblage structure but the presence of synergistic interactions suggests that the persistence and optimal contribution of the earthworm assemblage to ecosystem service is dependent on the presence of a number of functional groups.     

Additive and interactive effects of functionally dissimilar soil organisms on a grassland plant community

The productivity and diversity of plant communities are affected by soil organisms such as arbuscular mycorrhizal fungi (AMF), root herbivores and decomposers. However, it is unknown how interactions between such functionally dissimilar soil organisms affect plant communities and whether the combined effects are additive or interactive. In a greenhouse experiment we investigated the individual and combined effects of AMF (five Glomus species), root herbivores (wireworms and nematodes) and decomposers (collembolans and enchytraeids) on the productivity and nutrient content of a model grassland plant community as well as on soil microbial biomass and community structure. The effects of the soil organisms on productivity (total plant biomass), total root biomass, grass and forb biomass, and nutrient uptake of the plant community were additive. AMF decreased, decomposers increased and root herbivores had no effect on productivity, but in combination the additive effects canceled each other out. AMF reduced total root biomass by 18%, but decomposers increased it by 25%, leading to no net effect on total root biomass in the combined treatments. Total shoot biomass was reduced by 14% by root herbivores and affected by an interaction between AMF and decomposers where decomposers had a positive impact on shoot growth only in presence of AMF. AMF increased the shoot biomass of forbs, but reduced the shoot biomass of grasses, while root herbivores only reduced the shoot biomass of grasses. Interactive effects of the soil organisms were detected on the shoot biomasses of Lotus corniculatus, Plantago lanceolata, and Agrostis capillaris. The C/N ratio of the plant community was affected by AMF.In soil, AMF promoted abundances of bacterial, actinomycete, saprophytic and AMF fatty acid markers. Decomposers alone decreased bacterial and actinomycete fatty acids abundances but when decomposers were interacting with herbivores those abundances were increased. Our results suggests that at higher resolutions, i.e. on the levels of individual plant species and the microbial community, interactive effects are common but do not affect the overall productivity and nutrient uptake of a grassland plant community, which is mainly affected by additive effects of functionally dissimilar soil organisms.     

Consequence of grazing pattern and vegetation structure on the spatial variations of net N mineralisation in a wet grassland

The aim of this study was to determine whether the spatial heterogeneity of grassland vegetation structure would lead to spatial heterogeneity in the net nitrogen mineralisation process in the soil and therefore in the quantity of mineral nitrogen available for the plants. The net nitrogen mineralisation in the soil was compared between different vegetation patches generated by grazing, on two different types of plant communities: mesophilous and meso-hygrophilous.In ungrazed conditions, the net soil nitrogen mineralisation rates did not vary significantly between the two plant communities and remained relatively constant with time. Grazing by cattle or horses appeared to have two effects on the process of net soil nitrogen mineralisation. Firstly, it significantly stimulated net nitrogen mineralisation compared to ungrazed conditions and secondly, it led to spatial heterogeneity in mineralisation rates in the grazed enclosures. This spatial heterogeneity of nitrogen available for plants occurred both between and within plant communities.In the meso-hygrophilous plant community, net nitrogen mineralisation increased with grazing pressure. We suggest that a decrease of C inputs to the soil, concomitant with increasing grazing pressure, could decrease microbial nitrogen immobilisation.By contrast, in the mesophilous plant community net nitrogen mineralisation did not vary with grazing pressure. These differences in the functional responsiveness to grazing and biomass between the two plant communities could be related to the differences in the functional traits characterizing their dominant species along the grazing gradient. In the meso-hygrophilous community, the species composition switch with grazing intensity gradient led to the replacement of the perennial plant species by annual plant species which could lead to an improvement in the litter nitrogen content and an acceleration in the litter decomposition rate. By contrast, in the mesophilous plant community, the perennial species remained dominant along the grazing intensity gradient and could explain the absence of effect on the net nitrogen mineralisation rates.We suggest that at the scale of the vegetation patch, the decrease in plant biomass linked to grazing could regulate soil microorganism activity, in relation with shift in plant functional traits which improve litter decomposability.     

黄顶菊(Flaveria bidentis)凋落物对土壤无脊椎动物群落的影响

以黄顶菊(Flaveria bidentis)入侵的林地、荒地和沟渠作为调查样地,探讨黄顶菊凋落物对土壤无脊椎动物群落的影响。利用环刀进行取样,3种生境共捕获土壤无脊椎动物54 315头,隶属2门10纲17目。3种生境的优势类群皆为蜱螨目和弹尾目,其余类群的相对多度较小。黄顶菊凋落物能够为土壤无脊椎动物提供栖息地与食物来源,进而影响其群落结构及多样性,该影响与黄顶菊自身群体的生长状况有关,在长势较弱的林地生境影响较小,而在长势较强的荒地和沟渠生境影响较大。综上所述,黄顶菊入侵3种生境后,植株及其凋落物能为优势类群提供更好的栖息地和隐蔽所,并通过改变表层腐殖土的主要养分含量,引起土壤无脊椎动物多样性的升高,且土壤无脊椎动物在凋落物层中多样性呈自上而下的升高趋势。     

Resource dynamics in an early-successional plant community are influenced by insect exclusion

The exclusion of insects from terrestrial ecosystems may change productivity, diversity and composition of plant communities and thereby nutrient dynamics. In an early-successional plant community we reduced densities of above- and below-ground insects in a factorial design using insecticides. Beside measuring vegetation dynamics we investigated the effects of insect exclusion on above- and below-ground plant biomass, below-ground C and N storage by plants, litter quality, decomposition rate, soil water content, soil C:N ratio, nutrient availability and soil microbial activity and biomass.The application of soil insecticide had only minor effects on above- and below-ground biomass of the plant community but increased carbon content in root biomass and total carbon and nitrogen storage in roots. In one of the three investigated plant species (Cirsium arvense), application of soil insecticide decreased nitrogen concentration of leaves (−12%). Since C. arvense responded positively to soil insecticide application, this effect may be due to drought stress caused by root herbivory. Decomposition rate was slightly increased by the application of above-ground insecticide, possibly due to an impact on epigeic predators. The application of soil insecticide caused a slightly increased availability of soil water and an increased availability of mineralised nitrogen (+30%) in the second season. We explain these effects by phenological differences between the plant communities, which developed on the experimental plots. Microbial biomass and activity were not influenced by insecticide application, but were correlated to above-ground plant biomass of the previous year. Overall, we conclude that the particular traits of the involved plant species, e.g. their phenology, are the key to understand the resource dynamics in the soil.     

Benefactor and allelopathic shrub species have different effects on the soil microbial community along an environmental severity gradient

Patches where shrubs have either positive or negative effects on their understory plant community are common in arid ecosystems. The intensity and balance of these effects change along environmental severity gradients but, despite the major role of soil microbes in plant interactions, little is known about the differences among soil microbial communities under these species and their possible influence on such contrasting shrub effects. We hypothesized that microbial communities associated to benefactor and allelopathic shrubs would differ among them and that differences would increase with environmental severity. To test these hypotheses we characterized soil microbial biomass, activity and community composition under a benefactor shrub species, Retama sphaerocarpa, an allelopathic shrub species, Thymus hyemalis, and in bare soil among plants (gaps) at three sites along an environmental severity gradient. Shrubs promoted an increase in soil bacterial diversity, being bacterial communities associated to benefactor shrubs, allelopathic shrubs and gaps different in composition. Microbial enzymatic activity and biomass increased under shrubs and under more mesic conditions; nonetheless, they were highest under benefactor shrubs at the most arid site and under allelopathic shrubs at the less severe site. Compared to gaps, the presence of shrubs induced changes in microbial activity and community composition that were larger at the most severe site than at the less severe site. Along the gradient, benefactor shrubs enhanced the abundance of bacterial groups involved in organic matter decomposition and N fixation as well as plant pathogens, which could contribute to Retama's outstanding positive effects on understory plant biomass and diversity. Plant patches mitigate the effects of extreme conditions on associated plant and soil microbial communities and promote soil biodiversity and ecosystem functioning in arid ecosystems, with shrubs actively selecting for specific microbial groups in their understory.     

Soil fauna feeding activity in temperate grassland soils increases with legume and grass species richness

Edaphic fauna contributes to important ecosystem functions in grassland soils such as decomposition and nutrient mineralization. Since this functional role is likely to be altered by global change and associated shifts in plant communities, a thorough understanding of large scale drivers on below-ground processes independent of regional differences in soil type or climate is essential. We investigated the relationship between abiotic (soil properties, management practices) and biotic (plant functional group composition, vegetation characteristics, soil fauna abundance) predictors and feeding activity of soil fauna after accounting for sample year and study region. Our study was carried out over a period of two consecutive years in 92 agricultural grasslands in three regions of Germany, spanning a latitudinal gradient of more than 500 km. A structural equation model suggests that feeding activity of soil fauna as measured by the bait-lamina test was positively related to legume and grass species richness in both years. Most probably, a diverse vegetation promotes feeding activity of soil fauna via alterations of both microclimate and resource availability. Feeding activity of soil fauna also increased with earthworm biomass via a pathway over Collembola abundance. The effect of earthworms on the feeding activity in soil may be attributed to their important role as ecosystem engineers. As no additional effects of agricultural management such as fertilization, livestock density or number of cuts on bait consumption were observed, our results suggest that the positive effect of legume and grass species richness on the feeding activity in soil fauna is a general one that will not be overruled by regional differences in management or environmental conditions. We thus suggest that agri-environment schemes aiming at the protection of belowground activity and associated ecosystem functions in temperate grasslands may generally focus on maintaining plant diversity, especially with regard to the potential effects of climate change on future vegetation structure.     

Chemical composition and diversity influence non-additive effects of litter mixtures on soil carbon and nitrogen cycling: Implications for plant species loss

Diverse plant litter mixtures frequently decompose differently than expected compared to the average of the component species decomposing alone, and it remains unclear why decomposition may respond non-additively to diversity. Here, we hypothesized that litter chemical composition and chemical diversity would be important determinants of the strength and direction (synergistic versus antagonistic) of non-additive soil carbon (C) and nitrogen (N) cycling responses to litter mixtures. To test this, we performed a soil incubation experiment using litter mixtures comprised of up to four plant species, and we measured three components of decomposition: respiration, net N mineralization, and microbial biomass N accumulation. We used nine chemical traits to calculate the chemical composition and diversity of the litter mixtures. First, we found that respiration responded as the average of the individual species in the mixture (i.e. additively), rather than non-additively as initially predicted. Second, litter mixtures stimulated significantly more net N immobilization than expected in 64% of cases, and non-additive responses were highly dependent on mixture chemical composition, and were influenced to a lesser degree by chemical diversity. Specifically, concentrations of tannins and certain low molecular weight phenolics in the mixtures were positively correlated with greater N immobilization than expected. Non-additive N mineralization responses were poorly correlated with traditional measures of litter chemistry like N concentration, C:N, lignin:N, and phenolic:N. Our results also show that non-additive N mineralization responses were affected by loss of some species significantly more than others, and the effects of species loss could depend on 1) whether a species contains compounds with strong effects on non-additive responses; and 2) whether those compounds are also found in other species. Finally, litter mixtures stimulated more microbial biomass N than expected in 45% of cases, but non-additive responses were only weakly dependent on the litter chemistry variables that we measured.     

真菌群落沿气候梯度与植物凋落物分解之间的关系研究

The decomposition of plant litter is a major process of equivalent status to primary production in ecosystem functioning. The spatiotemporal changes in the composition and dynamics of litter fungal community along a climate gradient ranging from arid desert to humid-Mediterranean regions in Israel was examined using wheat straw litter bags placed at four selected sites along the climate gradient, arid, semi-arid, Mediterranean, and humid-Mediterranean sites. Litter samples were collected over a two-year decomposition period to evaluate litter weight loss, moisture, C：N ratio, fungal composition, and isolate density. The litter decomposition rate was found to be the highest during the first year of the study at the Mediterranean and arid sites. Although the Shannon-Wiener index values of the fungal communities in the litter samples were the highest at the humid-Mediterranean site, the number of fungal species was not significantly different between the four study sites. Different fungal groups were found to be related to different study sites： Basidiomycota, Mucoromycotina, and teleomorphic Ascomycota were associated with the humid-Mediterranean site, while Coelomycetes were mostly affected by the arid site. Our results indicate that climate factors play an important role in determining the structure of saprotrophic fungal communities in the decomposing litter and in mediating plant litter decomposition processes.