Positive relationship between herbaceous layer diversity and the performance of soil biota in a temperate forest

The current decline in biodiversity is particularly pronounced in the herbaceous layer of forest ecosystems. We explored the relationship between a naturally occurring plant diversity gradient in the understory vegetation of a deciduous forest and several above-and belowground ecosystem processes. We show that particularly soil microbial parameters and microarthropod densities are positively correlated with plant species richness. These results confirm recent findings in grassland ecosystems and highlight the intimate interconnectance between the diversity and functioning of above-and belowground compartments. We conclude that irrespective of a potential causal relationship between plant species richness and belowground processes, it is essential to consider the performance of soil biota in order to understand the relationship between herbaceous layer composition and ecosystem function.     

Soil engineering ants increase grass root arbuscular mycorrhizal colonization

The role of edaphic factors in driving the relationship between plant community structure and ecosystem processes is a key issue of the current debate on functional implications of biodiversity. In this study, we draw a direct link between aboveground/belowground relationships, vegetation structure, and aboveground management. We used ground nesting ants and arbuscular mycorrhizal fungi (AMF) as an example for quantifying the role of biotic interactions in soil. Although both groups are known to have a major impact on grasslands, the interactive effect of these taxa on vegetation structure and its sensitivity to grassland management is poorly understood. We show that the ant Lasius flavus increases the root arbuscular mycorrhizal colonization (AMC) of grasses by modifying biotic and abiotic soil properties. As a possible consequence, the shoot length of grass growing on ant mounds was shorter and shoot N and P concentrations were higher than in grass growing off of the mounds. In addition, management affected ant nest architecture and soil and, in turn, AMC. These results emphasize the need to consider the interactions between plants, soil microorganisms, soil fauna, and aboveground management to increase the understanding of the drivers of biodiversity and ecosystem functioning in grasslands both aboveground and belowground.     

Terrestrial ecosystem processes of Victoria Land, Antarctica

Terrestrial environments of Victoria Land, Antarctica are ideal systems to test hypotheses about the sensitivity of ecosystem processes to climate variability, and the relationships between soil biodiversity and ecosystem functioning because of their high sensitivity to climate change and their limited diversity. This region is also considered among the most pristine of ecosystems, and therefore may serve as an indicator for detecting the response of other ecosystems to global environmental change. Rates and controls over key ecosystem processes remain poorly documented over much of Victoria Land, but it is generally held that the distribution and functioning of soil communities are most limited by the availability of liquid water and organic carbon. Here we review examples of ecosystem processes from several sites in North and South Victoria Land and develop a regional synthesis accounting for variation in the availability of soil resources (i.e. liquid water, organic matter, inorganic nutrients). Variation in soil microclimate, organic matter, moisture and salinity encountered over gradients of coastal to interior sites, latitude, and soil chronosequences are the primary controls over the structure of soil communities and their functioning. Imbalanced stoichiometric nutrient ratios frequently encountered in Victoria Land ecosystems also contribute to limited distribution of soil biota, and where they occur these elemental imbalances indicate lower biological activity and little biotic control over bulk element ratios in soils. Priorities and future directions of Victoria Land soil and ecosystem research are also discussed.     

Biological crusts as a model system for examining the biodiversity-ecosystem function relationship in soils

Despite that soils may be the greatest repository of biodiversity on Earth, and that most terrestrial ecosystem functions occur in the soil, research on the role of soil biodiversity in ecosystem function has lagged behind corresponding research on aboveground organisms. Soil organisms pose special problems to biodiversity-function research, including the fact that we usually do not know their identity nor what they do in soil ecosystems, cannot easily estimate their biodiversity, and cannot culture the majority of the organisms for use in manipulative experiments. We propose here that biological soil crusts (BSCs) of deserts and many other ecosystems may serve as a useful model system for diversity-function research because the species concept is relatively well-defined within BSC organisms, their functional attributes are relatively well-known, and estimation and manipulation of biodiversity in experiments are feasible, at least within some groups of BSC biota. In spite of these features, there is a pronounced lack of research on biodiversity-function using these organisms. At least two complementary approaches are possible: experiments using artificially-constructed BSCs, and observational studies which statistically control for the effects of other factors which are likely to covary with biodiversity. We applied the latter to four observational datasets collected at multiple spatial scales in Spain and the United States using structural equation models or path analysis using ecosystem function indicators relating to hydrology, trapping and retention of soil resources, and nutrient cycling. We found that, even when total BSC abundance and key environmental gradients are controlled for, direct and approximately linear relationships between species richness and/or evenness and indicators of ecosystem functioning were common. Such relationships appear to vary independently of region or spatial scale, but their strength seems to differ in every dataset. Functional group richness did not seem to adequately capture biodiversity-function relationships, suggesting that bryophyte and lichen components of BSC may exhibit low redundancy. More research employing the multi-trophic, multi-functional, and manipulable BSC system may enable more rapid understanding of the consequences of biodiversity loss in soils, and help enable a biodiversity-function theory that is pertinent to the numerous ecosystem services provided by soil organisms.     

植物多样性对土壤动物影响的研究进展

生物多样性与生态系统功能一直是生态学研究的一个热点。近些年来的研究表明,植物多样性除了影响陆地生态系统地上部分的初级生产力等生态系统功能,还会间接影响地下生物多样性及土壤生态系统过程。本文概述了植物多样性对土壤动物的影响及其主要机制,归纳了植物多样性通过改变输入土壤中的资源数量与资源多样性、微生境结构、土壤环境因子等影响土壤动物的途径。目前关于土壤动物群落对植物多样性的响应仍存在很多问题和争议,本文总结了需要进一步深入研究的方向,特别指出了要加强研究影响植物多样性与土壤动物关系的生物与非生物因子、后续的生态效应和反馈、不同机制和途径的贡献定量化等。     

土壤生物结皮在干旱半干旱地区退化生态系统恢复中的作用(英文)

 土壤生物结皮是由细菌、真菌、藻类、地衣和苔藓等形成的一种混合体,作为干旱半干旱地区生态系统的组成部分,它对生态系统镶嵌格局和生态过程有不可忽视的影响;同时,土壤生物结皮通过影响局部水分条件,可以起到稳定土壤表层、减少土壤侵蚀、增加土壤氮养分的作用:因此,土壤生物结皮能够为土壤表层提供一种天然保护的作用,对干旱半干旱地区退化生态系统的恢复具有非常重要的作用。     

Grazing or Not Grazing: Implications for Ecosystem Services Provided by Biocrusts in Mediterranean Cork Oak Woodlands

Livestock grazing is one of the most common practices in managed woodlands affecting the abundance and diversity of plant and soil communities. While grazing effects have been studied thoroughly in vascular plants, little is known about grazing consequences for soil lichens and mosses (“biocrusts”) in managed woodlands. However, these complex communities have critical contributions to soil functioning in drylands. We evaluated grazing effects on biocrust abundance and functional composition in Mediterranean cork‐oak woodlands under long‐term grazing and after 7 and 17 years of grazing exclusion. We found four main groups in the region regarding the functional trait ‘growth form’: fruticose and foliose lichens, and short and tall mosses. Each group presented different water absorption and retention capacity, and showed group‐specific effects on soil water and temperature. Fruticose lichens were the most sensitive group to grazing, decreasing dramatically in cover with grazing (~7 times). Also, this group presented higher water retention capacity (~19 h), together with a consistent reducing effect on soil temperature along the grazing gradient (up to 0·9 °C). The shift in biocrust abundance and functional composition under grazing pressure has direct consequences in microclimate regulation, and is likely to influence other ecosystem processes such as CO₂ fixation, soil surface protection and habitat provision. In particular, microclimate regulation may affect cork‐oak regeneration processes, which is one of the main concerns in managed woodlands. Copyright © 2016 John Wiley & Sons, Ltd.     

Functional aspects of soil animal diversity in agricultural grasslands

There has been recent interest in the characterization of soil biodiversity and its function in agricultural grasslands. Much of the interest has come from the need to develop grassland management strategies directed at manipulating the soil biota to encourage a greater reliance on ecosystem self-regulation. This review summarises information on selected groups of soil animals in grasslands, the factors influencing their abundance, diversity and community structure and their relationships to the functioning and stability of grassland ecosystems. Observations on the impacts of agricultural managements on populations and communities of soil fauna and their interactions confirm that high input, intensively managed systems tend to promote low diversity while lower input systems conserve diversity. It is also evident that high input systems favour bacterial-pathways of decomposition, dominated by labile substrates and opportunistic, bacterial-feeding fauna. In contrast, low-input systems favour fungal-pathways with a more heterogeneous habitat and resource leading to domination by more persistent fungal-feeding fauna. In view of this, we suggest that low input grassland farming systems are optimal for increasing soil biotic diversity and hence self-regulation of ecosystem function. Research is needed to test the hypothesis that soil biodiversity is positively associated with stability, and to elucidate relationships between productivity, community integrity and functioning of soil biotic communities.     

生物多样性与生态系统功能的关系研究进展

在国内外研究生物多样性与生态系统功能的关系基础上 ,系统分析了物种多样性与生产力之间相互作用关系及其影响因子 ,探讨了生物多样性与稳定性的关系。     

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

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

Soil biodiversity and its implications for ecosystem functioning in a heterogeneous and variable environment

This paper evaluates theoretical arguments and empirical evidence on the interrelation of soil biodiversity and ecological functioning, in order to identify appropriate methodologies and promising fields of investigation. Predictions from a range of general theories are evaluated against the background of empirical findings and results from models on specific soil ecosystems with reference to the functional effects of species losses (1) within trophic levels, (2) in food-webs, and (3) those that depend on the size of the spatial scale. Difficulties in proving species richness effects empirically arise because (i) a range of mechanisms can compensate species richness effects, (ii) predictions on the effects of species richness are context-dependent, which renders valid generalisations less likely, and (iii) species richness effects are of a probabilistic, rather than a deterministic nature. Generally, the effects of changes in species richness are likely to be stronger in species-poor communities than in species-rich communities. The authors deduce the hypotheses that (a) the species richness within trophic levels is likely to reduce functional gaps in space and time, (b) the species richness across trophic levels of the decomposer food web can enhance synchronisation of subprocesses of nutrient cycling, and (c) differences in species richness are likely to be more pronounced on larger spatial scales. The authors infer that an explicit consideration of spatiotemporal variation is essential in experimentation, as well as in modelling, in order to analyse species richness–function relationships.     

放牧管理对青藏高原东缘高寒草甸土壤线虫的影响及作用机制

不同强度、季节的放牧是草地生态系统中主要的放牧管理措施,在生物多样性维持以及生态过程发挥中起着重要的作用,然而,关于青藏高原东缘高寒草甸地区放牧对土壤线虫群落的研究甚少。本文调查了放牧（轻度、中度和重度）对植物群落、土壤理化性质和线虫群落的影响。结果表明：植物群落、土壤理化性质和线虫群落受到放牧、时间以及放牧和时间共同作用的影响;食细菌线虫、植物凋落物生物量、根生物量、土壤含水量、土壤有机碳含量在轻度放牧草地内最高（p＜0.05）;杂类草生物量在中度放牧草地内最高;杂食线虫数量在重度放牧地内最低（p＜0.05）。植物群落和土壤特性与土壤线虫群落有明显的相关关系（p＜0.05）。结构方程模型显示食草动物主要通过植物群落影响植食和食细菌线虫,进一步的研究应针对植物群落多样性以及个体特征对线虫群落的影响。     

Cross-taxon surrogacy of biodiversity in the Indian Garhwal Himalaya

Biodiversity surveys were conducted in 13, 10×50 m² plots located between 1400 to 3700 m above mean sea level in a range of habitats in temperate mixed Oak and Coniferous forests through sub-alpine to the alpine grasslands in Chamoli district of Uttaranchal state in the Indian Garhwal Himalaya. Cross-taxon congruence in biodiversity (α-diversity and β-diversity) across macrolichens, mosses, liverworts, woody plants (shrubs and trees) and ants was investigated, so as to examine the extent to which these groups of organisms can function as surrogates for each other. Although woody plants provided a major substrate for macrolichens and mosses, there was no species-specific association between them. Woody plant species richness was highly positively correlated with mosses (r²=0.63, P<0.001), but the relationship was not particularly very strong with lichens and liverworts. While there was a significant correlation in the species turnover (β-diversity) of macrolichens with mosses (r²=0.21, P<0.005), the relationship was relatively poor with the woody plants. On the other hand, negative correlations emerged in the species richness of ants with those of macrolichens, mosses and woody plants (r²=−0.44, P<0.05), but most of the complementarity (turnover) relationships among them were positive. Since diversity between taxonomic hierarchies within the group was consistently significantly positively correlated in all these taxa, the higher taxonomic categories such as genus and family may be employed as surrogates for rapid assessment and monitoring of species diversity. Although no single group other than macrolichens has emerged as a good indicator of changes in species richness in all other groups, some concordant relationships between them conform to the hypothesis that species assemblages of certain taxonomic groups could still be used as surrogates for efficient monitoring of species diversity in other groups whose distribution may further predict the importance of conserving overall biodiversity in landscapes such as the Garhwal Himalaya.     

变化环境下土壤生物多样性潜在威胁与影响因素

作为地球上生物最为丰富的生境,土壤生态系统具有高度的生物多样性,并与地上生态系统有着密切联系。随着经济快速发展,人为干扰日益严重,土壤生物多样性遭受严重威胁,在物种大量灭绝及对土壤生物多样性总体认识相对局限的双重背景下,有效保护土壤生物多样性迫在眉睫。首先,提出了土壤生物多样性的威胁框架,并详述了土地利用变化、农业管理措施、土壤退化、转基因作物、外来植物入侵五大威胁因素在生态系统,物种,基因三个尺度下的不同作用过程。其次,阐明了各威胁因素对土壤生物多样性的具体影响及其驱动机制。最后指出,尽管人们对于上述威胁因素的研究已经有了部分成果,但对于研究各威胁因素在不同尺度下的影响,明确因素间相互作用方式及贡献程度,进行土壤生物多样性威胁的空间精细化制图等方面还面临着巨大挑战。分析土壤生物多样性潜在威胁能够为实现对其综合定量评估、保护土壤生物多样性以及城市规划等提供有力依据。     

Grassland management influences spatial patterns of soil microbial communities

Soil micro-organisms play a vital role in grassland ecosystem functioning but little is known about the effects of grassland management on spatial patterns of soil microbial communities. We compared plant species composition with terminal restriction fragment length polymorphism (T-RFLP) fingerprints of soil bacterial and fungal communities in unimproved, restored and improved wet grasslands. We assessed community composition of soil micro-organisms at distances ranging from 0.01 m to 100 m and determined taxa–area relationships from field- to landscape level. We show that land management type influenced bacterial but not fungal community composition. However, extensive grassland management to restore aboveground diversity affected spatial patterns of soil fungi. We found distinct distance–decay and small-scale aggregation of fungal populations in extensively managed grasslands restored from former arable use. There were no clear spatial patterns in bacterial communities at the field-scale. However, at the landscape level there was a moderate increase in bacterial taxa and a strong increase in fungal taxa with the number of sites sampled. Our results suggest that grassland management affects soil microbial communities at multiple scales; the observed small-scale variation may facilitate plant species coexistence and should be taken into account in field studies of soil microbial communities.     

Spatial distribution and physiology of biological soil crusts from semi-arid central Spain are related to soil chemistry and shrub cover

Despite the critical role of biological soil crusts (BSCs) in arid and semi-arid ecosystem function, few studies are found concerning the most important environmental variables affecting their distribution and physiology. This study seeks to determine soil and microenvironmental factors affecting the spatial distribution and pigment production of BSC-forming lichens and mosses in open patches of a semi-arid Mediterranean kermes oak thicket. We measured late-successional BSC cover, shrub cover, distance to nearest kermes oak (to test for effects of kermes oak thicket microenvironment on BSC), and pigment concentration of one lichen (Cladonia foliacea) and one moss (Pleurochaete squarrosa) species in the Nature Reserve El Regajal-Mar de Ontígola (Central Spain). At the macroscale (>0.5 m), results showed that BSC distribution and pigments were tightly coupled to a suite of soil properties, in particular soil pH, Fe, and Ca. Specifically, soil pH had a positive relationship with the cover of five individual BSC-forming lichen species and was negatively related to pigment production in C. foliacea. When pH was excluded from the analysis, Ca appeared as the main soil variable and was correlated with total BSC cover and total lichen cover. The micronutrient Fe had a significant positive relationship with the concentration of eight pigments in P. squarrosa and was also coupled with the cover of two BSC-forming lichens. Manganese, previously proposed as a key limiting micronutrient for BSCs, affected lichen diversity in a negative way. At the microscale (∼0.5 m), kermes oak microenvironment, shrub cover, and moss cover were determinants of BSC distribution, and total lichen and total BSC cover were overrepresented on N and E-facing shrub microsites. Our findings suggest that soil chemical variability and microsite diversity created by neighbouring vegetation affect BSC distribution in complex and essential ways and that studies aiming to explore BSC-environment relationships should be conducted at various spatial scales. Studies based on species- or group-specific responses are, thus, inadequate to unveil the main factors determining the distribution of the diverse organisms that constitute BSCs and/or to propose potential tools aiming to restore BSC in arid and semiarid ecosystems.     

农业景观生物多样性功能和保护对策

研究表明,农业集约化生产和农业景观均质化导致农业景观生物多样性降低,致使生物多样性相关野生资源保护、自然授粉、害虫调节、水土涵养等生态系统服务功能受损,进而影响了农业可持续发展。本文对国内外农业景观生物多样性及其生态系统服务功能研究和实践进行了综述。分别探讨了农业景观在生物多样性保护、授粉服务、调节服务、水土涵养等多方面生态服务功能中发挥的作用及其生物多样性保护的意义,总结了欧美在农业景观生物多样性保护所采取的实践措施,指出中国虽然在生物多样性保护方面做出很多努力,但却忽略了对农业景观的保护,近些年的城市化和集约化发展都进一步加剧了农业景观生物多样性的丧失,亟需借鉴欧美国家经验提出我国的农业景观生物多样性保护策略。我们认为保护和提高农业景观生物多样性的生态集约化对维护农作物产量,改善品种具有重要的作用。农业景观生物多样性保护需要从农田生态系统和农业景观两个尺度上开展农业景观综合管理,提高农田作物和景观植物多样性和异质性,恢复和提高生物多样性及其生态服务功能。最后,针对我国农业可持续发展面临的问题和需求,建议从政策和法规、监测和评估、工程技术研发、技术集成示范、生态补贴制度、培训和推广等方面,开展农业景观生物多样性保护。     

Rehabilitation of the nematode fauna in a phytostabilized,heavily zinc-contaminated,sandy soil

Background Aim and Scope  The Maatheide in Lommel, Belgium, is an extremely metal contaminated, sandy area where vegetation has disappeared over ca. 130 hectares due to the activities of a former pyrometallurgical zinc smelter. To reduce the environmental impact of this area a rehabilitation strategy had to be developed. Therefore, in the centre of this area, an experimental phytostabilization (grass) field of three hectares had been installed in 1990. After a grass cover had been established, the development of the nematode fauna in the phyto-stabilized soil was studied. Nematodes act at various levels in soil ecosystems: herbivorous species extract their food from plant roots, bacterivorous and fungivorous species feed on microbes, predatory species consume other nematodes, and omnivorous species have mixed diets. In a mature soil ecosystem that normally exercises its manifold functions, a diverse nematode fauna occurs, reflecting the intactness of the ecosystem. As such, this fauna is indicative of crop growth, vegetative diversity, organic matter decomposition, microbial activity and diversity, and the maturity of the soil ecosystem. Methods  A metal immobilizing soil amendment (beringite) and municipal waste compost (to improve the nutrient status and water-retaining capacity) were incorporated in the soil and metal tolerant ecotypes of grasses were sown. Soil samples for nematode analyses were taken four times. Results  As a result of the treatment, pH of the soil increased and the water extractable amount of Zn was strongly reduced. Grass growth revitalized the impoverished soil ecosystem, organisms as well as metabolic processes. The nematode fauna of the experimental field in the Maatheide has been studied since 1997 and the recovery of the various feeding guilds and taxa was checked again in 2002. Nematode densities and feeding guilds have normalized, with omnivores and predators as the last guilds to reappear. Up to 27 species with a current diversity were observed in the grass-covered experimental plot, but a number of ubiquitous species, present at a comparable site at some distance, remained absent. Conclusion  It can thus be concluded that rehabilitation of an impoverished soil ecosystem is possible in heavily contaminated soils by means of phytostabilization, but there are some limitations on rehabilitation, since a number of common nematode species remained absent. Further research should indicate if this absence is indicative of a loss of essential processes in the soil. Recommendation and Perspective. Phytostabilization of heavily zinc-contaminated, sandy soil also remediates impoverished soil ecosystems. In particular, the recovery of nematode feeding categories is indicative for the normalization of soil life. The absence/presence of a number of ubiquitous taxa should be checked again after some time to verify if recovery is completed.     

Functional diversity of nematodes

Nematodes are the most abundant metazoa. Their food specificity, the high number of species and high abundance in every habitat where decomposition takes place indicates that the structure of the nematode community has a high information content. Since nematodes respond rapidly to new resources, and the nematode fauna can be efficiently analyzed, the structure of the nematode community offers an instrument to assess (changes in) the conditions of soils. A functional grouping of nematodes is generally synonymous with allocation into feeding groups. However, soil quality assessment indices based on the presence of all feeding groups still provide insufficient information regarding the functioning of soil ecosystems and their threats. An alternative concept of functional groups is based on the life history of nematodes. In this paper we present the most recent colonizer–persister allocation and the application of this scaling in the Maturity Index, cp-triangles, MI(2–5) and PPI/MI-ratio. We propose to integrate the life strategy approach and trophic group classification to obtain a better understanding of nematode biodiversity and soil functioning. Attention is given to competitive exclusion and coexistence and we summarize present concepts regarding succession and degradation.     

Collembola species composition and diversity effects on ecosystem functioning vary with plant functional group identity

The relationship between decomposer diversity and ecosystem functioning is little understood although soils accommodate a significant proportion of worldwide biodiversity. Collembola are among the most abundant and diverse decomposers and are known to modify plant growth. We examined the effects of Collembola species diversity (one, two and three species belonging to different life history groups) and composition on litter decomposition and the performance of plant communities (above- and belowground productivity) of different functional groups (grasses, forbs and legumes). Collembola densities did not increase with diversity indicating niche overlap. Generally, Collembola species composition was a better predictor for ecosystem functioning than Collembola species number with the impacts of Collembola diversity and composition on ecosystem functioning strongly depending on plant functional group identity. Non-linear effects of Collembola diversity on litter decomposition and plant productivity suggest pronounced and context dependent species interactions and feeding habits. Net surface litter decomposition was decreased by Collembola, whereas root litter decomposition was at maximum in the highest Collembola diversity treatment. Forbs benefitted most from the presence of three Collembola species. Similarly, Collembola diversity influenced root depth distribution in a plant functional group specific way: while grass root biomass decreased with increasing Collembola diversity in the upper and lower soil layer, legume root biomass increased particularly in the lower soil layer. Idiosyncratic and context dependent effects of Collembola diversity and composition even in rather simple assemblages of one to three species suggest that changes in Collembola diversity may have unpredictable consequences for ecosystem functioning. The finding that changes in Collembola performance did not directly translate to alterations in ecosystem functioning indicates that response traits do not necessarily conform to effect traits. Distinct plant functional group specific impacts of Collembola diversity on root depth distribution are likely to modify plant competition in complex plant communities and add a novel mechanism how decomposers may affect plant community assembly.