Nearctic earthworm fauna in the southern USA: biodiversity and effects on ecosystem processes

An important aspect of biodiversity is the relative importance of species in the functioning of ecosystems; this is particularly so for the soil biota which regulate organic matter and nutrient dynamics in soil. This paper explores some of the relationships between biodiversity and ecosystem processes, using the example of the nearctic earthworm fauna in the glacial refugium of the southern USA. Competitive exclusion of nearctic earthworm species by exotic species has been postulated but there is little direct evidence of it; habitat alteration is the likely cause of native species decline. Reduced earthworm diversity may or may not strongly affect certain ecosystem processes, but more diverse assemblages may more effectively exploit soil resources and influence a wider array of processes. Nearctic species may be better adapted than exotics to local conditions and thus more strongly influence ecosystem processes. Earthworm communities provide a clear case for the union of functional and taxonomic biodiversity studies, because of the recognized ecological strategies of many species. However, some nearctic taxa may deviate from these strategies. Earthworms utilize course woody debris in forests both as a refuge and a resource, while enhancing the decomposition of wood. Management strategies to maintain or increase biodiversity of soil biota should include residual wood on the forest floor. An important task for ecosystem management is to restore biodiversity in degraded ecosystems; introduction programmes and techniques such as periodic burning may increase the abundance and diversity of native earthworm species. Whole ecosystem conservation and management are probably the most practical ways to conserve biodiversity generally and may be the only ways to maintain soil biodiversity.     

农田土壤线虫多样性研究现状及展望

目前土壤生物多样性已成为土壤生态学研究的热点问题之一。土壤生物以不同的方式改变着土壤的物理、化学和生物学特性。在农田生态系统中, 土壤动物是分解作用和养分矿化作用等生态过程的主要调节者。线虫作为土壤中数量最丰富的后生动物, 其生活史和取食类型多样, 在生态系统中发挥着重要作用。本文介绍了农田生态系统中影响线虫多样性的主要因素; 回顾了土壤线虫的物种多样性、营养类群多样性、生活史多样性和功能多样性的研究现状; 并提出了今后农田生态系统线虫多样性研究的重点。建议通过综合土壤线虫的生活史策略和营养类群等信息, 深入了解其生物多样性和土壤生态系统功能, 从而更好地发挥土壤线虫对农田生态系统变化的生物指示作用。     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

The importance of the biodiversity of soil biota in arid ecosystems

The importance of soil biota in maintaining ecosytem integrity is examined by a review of studies of soil processes and soil biota in arid ecosystems. In decomposition and mineralization processes, there is a temporal succession of microarthropod and nematode species. Tydeid mites are keystone species in the early stages of decomposition. Soil pore neck size variation affects the spatial distribution of soil biota and requires that species differ in size to provide efficient processing of organic matter. In arid ecosystems, the sub-set of soil biota that is active at any point in time is determined by the soil water potential and soil temperature. The structure of soil microarthropod assemblages is similar in several arid regions of the world and abundance and diversity are directly related to quantity of litter accumulations and soil organic matter. The unique life histories and behavioural characteristics of desert soil macrofauna (termites and ants) determine the effects of these organisms on soil properties and soil formation. The soil biota, by affecting the spatial and temporal distribution of essential sesources (water and nutrients), are essential to the maintenance of the ecosystem integrity of arid ecosystems.     

Soil animal responses to moisture availability are largely scale,not ecosystem dependent: insight from a cross‐site study

Climate change will result in reduced soil water availability in much of the world either due to changes in precipitation or increased temperature and evapotranspiration. How communities of mites and nematodes may respond to changes in moisture availability is not well known, yet these organisms play important roles in decomposition and nutrient cycling processes. We determined how communities of these organisms respond to changes in moisture availability and whether common patterns occur along fine‐scale gradients of soil moisture within four individual ecosystem types (mesic, xeric and arid grasslands and a polar desert) located in the western United States and Antarctica, as well as across a cross‐ecosystem moisture gradient (CEMG) of all four ecosystems considered together. An elevation transect of three sampling plots was monitored within each ecosystem and soil samples were collected from these plots and from existing experimental precipitation manipulations within each ecosystem once in fall of 2009 and three times each in 2010 and 2011. Mites and nematodes were sorted to trophic groups and analyzed to determine community responses to changes in soil moisture availability. We found that while both mites and nematodes increased with available soil moisture across the CEMG, within individual ecosystems, increases in soil moisture resulted in decreases to nematode communities at all but the arid grassland ecosystem; mites showed no responses at any ecosystem. In addition, we found changes in proportional abundances of mite and nematode trophic groups as soil moisture increased within individual ecosystems, which may result in shifts within soil food webs with important consequences for ecosystem functioning. We suggest that communities of soil animals at local scales may respond predictably to changes in moisture availability regardless of ecosystem type but that additional factors, such as climate variability, vegetation composition, and soil properties may influence this relationship over larger scales.     

锦鸡儿属灌丛化对草原化荒漠区土壤线虫群落组成和代谢足迹的影响

调查了内蒙古草原化荒漠区3种锦鸡儿[垫状锦鸡儿(Caragana tibetica)、荒漠锦鸡儿(C.roborovskyi)和狭叶锦鸡儿(C.stenophylla)]灌丛内、外土壤线虫群落多样性、组成和代谢足迹,以及相关的土壤理化性质和植物群落特征,旨在探讨锦鸡儿属灌丛对土壤线虫群落的影响,并分析这种影响是否具有灌木种间差异,同时从线虫功能团水平上探究灌丛如何通过非生物因素和生物因素影响线虫群落组成。结果表明：锦鸡儿属灌丛对土壤线虫多度、丰富度和多样性无显著影响,但却显著影响土壤线虫群落组成。原因是灌丛对土壤线虫群落具有物种选择性;灌丛可能主要是通过根系分泌物、凋落物质量等因素,而不是通过土壤理化性质和林下植物影响线虫群落组成。土壤线虫组成的变化引起线虫代谢足迹发生明显变化。灌丛内土壤线虫功能代谢足迹大于灌丛外(除狭叶锦鸡儿外),说明灌丛内土壤线虫群落对碳的利用率更高。3种锦鸡儿属灌木中,狭叶锦鸡儿的结构代谢足迹最大,表明狭叶锦鸡儿对捕食杂食类线虫代谢活性的促进作用更强。土壤线虫组成的变化通过线虫代谢足迹导致土壤食物网结构发生显著性变化。狭叶锦鸡儿灌丛土壤干扰程度低,营养富集状况好...     

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。     

Increasing Levels of Physical Disturbance Affect Soil Nematode Community Composition in a Previously Undisturbed Ecosystem

Soil is essential for the sustenance of life. Diverse soil organisms support several biological processes such as organic matter decomposition, mineralization, nutrient cycling, and controlling pests and diseases. Among multicellular soil organisms, nematodes are ubiquitous, functionally diverse, and abundant. Notably, agricultural practices have diverse impacts on plants, soils, and soil organisms. Tillage affects nematodes directly by altering pore size and disrupting the continuity of water films and indirectly by affecting the lower trophic groups such as bacteria and fungi. The primary goal of this study was to examine the effect of increasing levels of physical disturbance on nematode communities in an undisturbed forest ecosystem. The experiment included four treatments: control with no disturbance, surface litter removed with no litter and no vegetation, tilling the soil with a rototiller every 2 mon, and every 2 wk. Tillage significantly reduced the overall abundance and overall richness of nematode communities over time. Among nematode trophic groups, tillage significantly reduced the abundance and richness of bacterial feeders, predators, and omnivores over time. The abundance and richness of c-p 2, c-p 4, and c-p 5 class nematodes were significantly decreased by tillage. Unlike tillage, minimal disturbance such as removal of surface litter resulted in a significant decrease in the abundance of only three genera: Acrobeles, Aporcelaimellus, and Boleodorus. Nonmetric multidimensional scaling analysis revealed that nematodes of higher c-p classes such as Dorylaimida, Aporcelaimellus, Alaimus, Clarkus, and Tripyla were sensitive to physical disturbances. Bacterial feeders belonging to the c-p 2 class such as Tylocephalus, Acrobeles, Ceratoplectus, Plectus, and Pseudacrobeles were significantly reduced by tillage. Moreover, tillage significantly reduced the functional metabolic footprint of nematodes, which indicates decreased metabolic activity, reduced C inflow, and poorly structured soil food webs. Previous studies conducted in agricultural ecosystems determined that Clarkus, Filenchus, and Plectus were tolerant to tillage; however, they were found sensitive to tillage in our study. Overall, our study suggests that increasing levels of physical disturbance are detrimental to nematode community abundance and diversity that could affect soil ecosystem stability and sustainability.     

Global patterns of stream detritivore distribution: implications for biodiversity loss in changing climates

Aim We tested the hypothesis that shredder detritivores, a key trophic guild in stream ecosystems, are more diverse at higher latitudes, which has important ecological implications in the face of potential biodiversity losses that are expected as a result of climate change. We also explored the dependence of local shredder diversity on the regional species pool across latitudes, and examined the influence of environmental factors on shredder diversity. Location World‐wide (156 sites from 17 regions located in all inhabited continents at latitudes ranging from 67° N to 41° S). Methods We used linear regression to examine the latitudinal variation in shredder diversity at different spatial scales: alpha (α), gamma (γ) and beta (β) diversity. We also explored the effect of γ‐diversity on α‐diversity across latitudes with regression analysis, and the possible influence of local environmental factors on shredder diversity with simple correlations. Results Alpha diversity increased with latitude, while γ‐ and β‐diversity showed no clear latitudinal pattern. Temperate sites showed a linear relationship between γ‐ and α‐diversity; in contrast, tropical sites showed evidence of local species saturation, which may explain why the latitudinal gradient in α‐diversity is not accompanied by a gradient in γ‐diversity. Alpha diversity was related to several local habitat characteristics, but γ‐ and β‐diversity were not related to any of the environmental factors measured. Main conclusions Our results indicate that global patterns of shredder diversity are complex and depend on spatial scale. However, we can draw several conclusions that have important ecological implications. Alpha diversity is limited at tropical sites by local factors, implying a higher risk of loss of key species or the whole shredder guild (the latter implying the loss of trophic diversity). Even if regional species pools are not particularly species poor in the tropics, colonization from adjacent sites may be limited. Moreover, many shredder species belong to cool‐adapted taxa that may be close to their thermal maxima in the tropics, which makes them more vulnerable to climate warming. Our results suggest that tropical streams require specific scientific attention and conservation efforts to prevent loss of shredder biodiversity and serious alteration of ecosystem processes.     

Landscape perspectives on agricultural intensification and biodiversity – ecosystem service management

Understanding the negative and positive effects of agricultural land use for the conservation of biodiversity, and its relation to ecosystem services, needs a landscape perspective. Agriculture can contribute to the conservation of high‐diversity systems, which may provide important ecosystem services such as pollination and biological control via complementarity and sampling effects. Land‐use management is often focused on few species and local processes, but in dynamic, agricultural landscapes, only a diversity of insurance species may guarantee resilience (the capacity to reorganize after disturbance). Interacting species experience their surrounding landscape at different spatial scales, which influences trophic interactions. Structurally complex landscapes enhance local diversity in agroecosystems, which may compensate for local high‐intensity management. Organisms with high‐dispersal abilities appear to drive these biodiversity patterns and ecosystem services, because of their recolonization ability and larger resources experienced. Agri‐environment schemes (incentives for farmers to benefit the environment) need to broaden their perspective and to take the different responses to schemes in simple (high impact) and complex (low impact) agricultural landscapes into account. In simple landscapes, local allocation of habitat is more important than in complex landscapes, which are in total at risk. However, little knowledge of the relative importance of local and landscape management for biodiversity and its relation to ecosystem services make reliable recommendations difficult.     

Plant species identity and diversity effects on different trophic levels of nematodes in the soil food web

Previous studies on biodiversity and soil food web composition have mentioned plant species identity, as well as plant species diversity as the main factors affecting the abundance and diversity of soil organisms. However, most studies have been carried out under limitations of time, space, or appropriate controls. In order to further examine the relation between plant species diversity and the soil food web, we conducted a three-year semi-field experiment in which eight plant species (4 forb and 4 grass species) were grown in monocultures and mixtures of two, four and eight plant species. In addition there were communities with 16 plant species. We analyzed the abundance and identity of the nematodes in soil and roots, including feeding groups from various trophic levels (primary and secondary consumers, carnivores, and omnivores) in the soil food web.
Plant species diversity and plant identity affected the diversity of nematodes. The effect of plant diversity was attributed to the complementarity in resource quality of the component plant species rather than to an increase in total resource quantity. The nematode diversity varied more between the different plant species than between different levels of plant species diversity, so that plant identity is more important than plant diversity. Nevertheless the nematode diversity in plant mixtures was higher than in any of the plant monocultures, due to the reduced dominance of the most abundant nematode taxa in the mixed plant communities. Plant species identity affected the abundances of the lower trophic consumer levels more than the higher trophic levels of nematodes. Plant species diversity and plant biomass did not affect nematode abundance. Our results, therefore, support the hypothesis that resource quality is more important than resource quantity for the diversity of soil food web components and that plant species identity is more important than plant diversity per se.     

Principal methods for isolation and identification of soil microbial communities

Soil microbial populations play crucial role in soil properties and influence below-ground ecosystem processes. Microbial composition and functioning changes the soil quality through decomposition of organic matter, recycling of nutrients, and biological control of parasites of plants. Moreover, the discovery that soil microbes may translate into benefits for biotechnology, management of agricultural, forest, and natural ecosystems, biodegradation of pollutants, and waste treatment systems maximized the need of scientists for the isolation and their characterization. Operations such as the production of antibiotics and enzymic activities from microorganisms of soil constitute objectives of industry in her effort to cope with the increase of population of earth and disturbance of environment and may ameliorate the effects of global climate change. In the past decades, new biochemical and molecular techniques have been developed in our effort to identify and classify soil bacteria. The goal of measuring the soil microbial diversity is difficult because of the limited knowledge about bacteria species and classification through families and orders. Molecular techniques extend our knowledge about microbial diversity and help the taxonomy of species. Measuring and monitoring soil microbial communities can lead us to better understanding of their composition and function in many ecosystem processes.     

Influence of ecological and edaphic factors on biodiversity of soil nematodes

Nematodes are the most diverse and highly significant group of soil-inhabiting microorganisms that play a vital role in organic material decomposition and nutrient recycling.Diverse geographical locations and environmental gradients show a significant impact on the diversity of nematodes. Present study aims to assess the effects of ecological (altitude, temperature, moisture) and edaphic (soil pH, nutrients, soil patches) factors on the soil nematode diversity and structure at five different landscape patches (forests, apple orchards, rice fields, pastures, and alpine zone) from ten different sites of Kashmir valley (India). Differences in the altitudinal gradients results in the shift of generic nematode population. Among the soil patches, highest nematode diversity was observed in forest soil and least in alpine soil; however, bacteriovorous nematodes dominated all the soil patches. The temperature and moisture have a significant effect on nematode diversity, the highest nematode trophic levels were observed above 21°C temperature, and 30% moisture. Nematode abundance decreased from alkaline to acidic pH of the soil. Soil nutrients such as, nitrogen (N) and phosphorus (P) have shown a detrimental effect in nematode richness at each site, where nematode diversity and richness of genera were higher at abundant soil N and P but decreased at low soil nutrients. Ecological indices like diversity index (DI), Shannon-Wiener Index (H'), enrichment index (EI), and maturity Index (MI) values demonstrated forest soil more favourable for nematodes and high soil health status than other soil patches. This study suggested that these indices may be helpful as soil monitoring tools and assessing ecosystem sustainability and biodiversity.     

Role of Nematodes in Soil Health and Their Use as Indicators

The composition of nematode communities (plant-parasitic and free-living) may be used as bioindicators of soil health or condition because composition correlates well with nitrogen cycling and decomposition, two critical ecological processes in soil. Maturity and trophic diversity indices withstand statistical rigor better than do abundances, proportions, or ratios of trophic groups. Maturity indices respond to a variety of land-management practices, based largely on inferred life history characteristics of families. Similarity indices may be more useful than diversity indices because they reflect taxon composition. Improving existing indices or developing alternative indices refined by a greater understanding of the biology of key taxa may enhance the utility of nematodes as bioindicators.     

The Function of Marine Critical Transition Zones and the Importance of Sediment Biodiversity

Estuaries and coastal wetlands are critical transition zones (CTZs) that link land, freshwater habitats, and the sea. CTZs provide essential ecological functions, including decomposition, nutrient cycling, and nutrient production, as well as regulation of fluxes of nutrients, water, particles, and organisms to and from land, rivers, and the ocean. Sediment-associated biota are integral to these functions. Functional groups considered essential to CTZ processes include heterotrophic bacteria and fungi, as well as many benthic invertebrates. Key invertebrate functions include shredding, which breaks down and recycles organic matter; suspension feeding, which collects and transports sediments across the sediment–water interface; and bioturbating, which moves sediment into or out of the seabed. In addition, macrophytes regulate many aspects of nutrient, particle, and organism dynamics above- and belowground. Animals moving within or through CTZs are vectors that transport nutrients and organic matter across terrestrial, freshwater, and marine interfaces. Significant threats to biodiversity within CTZs are posed by anthropogenic influences; eutrophication, nonnutrient pollutants, species invasions, overfishing, habitat alteration, and climate change affect species richness or composition in many coastal environments. Because biotic diversity in marine CTZ sediments is inherently low whereas their functional significance is great, shifts in diversity are likely to be particularly important. Species introductions (from invasion) or loss (from overfishing or habitat alteration) provide evidence that single-species changes can have overt, sweeping effects on CTZ structure and function. Certain species may be critically important to the maintenance of ecosystem functions in CTZs even though at present there is limited empirical evidence that the number of species in CTZ sediments is critical. We hypothesized that diversity is indeed important to ecosystem function in marine CTZs because high diversity maintains positive interactions among species (facilitation and mutualism), promoting stability and resistance to invasion or other forms of disturbance. The complexity of interactions among species and feedbacks with ecosystem functions suggests that comparative (mensurative) and manipulative approaches will be required to elucidate the role of diversity in sustaining CTZ functions. Received 25 February 2000; accepted 31 January 2001.     

The local introduction of strongly interacting species and the loss of geographic variation in species and species interactions

Species introductions into nearby communities may seem innocuous, however, these introductions, like long-distance introductions (e.g. trans- and intercontinental), can cause extinctions and alter the evolutionary trajectories of remaining community members. These 'local introductions' can also more cryptically homogenize formerly distinct populations within a species. We focus on several characteristics and the potential consequences of local introductions. First, local introductions are commonly successful because the species being introduced is compatible with existing abiotic and biotic conditions; many nearby communities differ because of historical factors and the absence of certain species is simply the result of barriers to dispersal. Moreover, the species with which they interact most strongly (e.g. prey) may have, for example, lost defences making the establishment even more likely. The loss or absence of defences is especially likely when the absent species is a strongly interacting species, which we argue often includes mammals in terrestrial communities. Second, the effects of the introduction may be difficult to detect because the community is likely to converge onto nearby communities that naturally have the introduced species (hence the perceived innocuousness). This homogenization of formerly distinct populations eliminates the geographic diversity of species interactions and the geographic potential for speciation, and reduces regional species diversity. We illustrate these ideas by focusing on the introduction of tree squirrels into formerly squirrel-less forest patches. Such introductions have eliminated incipient species of crossbills (Loxia spp.) co-evolving in arms races with conifers and will likely have considerable impacts on community structure and ecosystem processes.     

Microbe-mediated plant–soil feedback and its roles in a changing world

Plants affect soil conditions, which in turn alter plant growth and interspecific competition, forming plant?Csoil feedback (PSF) systems. PSF is a good example of bidirectional interactions between biomes and the non-living environments, acting as a major driving force of community structure and ecosystem function. Among the major types of PSF mediated by various soil components, there are many holes in our knowledge of the interactions between PSF mediated by plant species-specific litter and PSF mediated by soil microbes. Here I discuss the role of the functional diversity of microbial decomposers in litter-mediated PSF and also propose new hypotheses on the role of microbial diversity in PSF mediated by pathogenic and mutualistic soil microbes. I also review how PSF interacts with human-induced environmental change, i.e., direct drivers of change in the ecosystem (e.g. climate change and the invasion of alien species). Many authors have suggested that the impact of alien plant species on ecosystems is mediated by PSF, which also interacts with other direct drivers, such as climate change. Using a simple model of litter-mediated PSF with microbial decomposers, I confirm that the interactions between PSF and other direct drivers affect the invasion process of alien species. The model also demonstrates that the functional diversity of microbial decomposers accelerates or decelerates the speed of the invasion depending on the environmental change scenarios. Further theoretical and empirical studies are needed to derive general predictions on how exogenous environmental change induced by human activities alters communities and ecosystems through disturbance or modification of endogenous community?Cecosystem interactions, such as the functioning of PSF.     

Plant Diversity Impacts Decomposition and Herbivory via Changes in Aboveground Arthropods

Loss of plant diversity influences essential ecosystem processes as aboveground productivity, and can have cascading effects on the arthropod communities in adjacent trophic levels. However, few studies have examined how those changes in arthropod communities can have additional impacts on ecosystem processes caused by them (e.g. pollination, bioturbation, predation, decomposition, herbivory). Therefore, including arthropod effects in predictions of the impact of plant diversity loss on such ecosystem processes is an important but little studied piece of information. In a grassland biodiversity experiment, we addressed this gap by assessing aboveground decomposer and herbivore communities and linking their abundance and diversity to rates of decomposition and herbivory. Path analyses showed that increasing plant diversity led to higher abundance and diversity of decomposing arthropods through higher plant biomass. Higher species richness of decomposers, in turn, enhanced decomposition. Similarly, species-rich plant communities hosted a higher abundance and diversity of herbivores through elevated plant biomass and C:N ratio, leading to higher herbivory rates. Integrating trophic interactions into the study of biodiversity effects is required to understand the multiple pathways by which biodiversity affects ecosystem functioning.     

Trait convergence and trait divergence in herbaceous plant communities: Mechanisms and consequences

In landscapes subject to intensive agriculture, both soil fertility and vegetation disturbance are capable of impacting strongly, evenly and simultaneously on the herbaceous plant cover and each tends to impose uniformity on the traits of constituent species. In more natural and ancient grasslands greater spatial and temporal variation in both productivity and disturbance occurs and both factors have been implicated in the maintenance of species‐richness in herbaceous communities. However, empirical data suggest that disturbance is the more potent driver of trait differentiation and species co‐existence at a local scale. This may arise from the great diversity in opportunities for establishment, growth or reproduction that arise when the intensity of competition is reduced by damage to the vegetation. In contrast to the diversifying effects of local disturbances, productivity‐related plant traits (growth rate, leaf longevity, leaf chemistry, leaf toughness, decomposition rate) appear to be less variable on a local scale. This difference in the effects of the productivity and disturbance filters arises from the relative constancy of productivity within the community and the diversity in agency and in spatial and temporal scales exhibited by disturbance events. Also, evolutionary responses to disturbances involve minor adaptive shifts in phenological and regenerative traits and are more likely to occur as micro‐evolutionary steps than the shifts in linked traits in the core physiology associated with the capacity to exploit productive and unproductive habitats. During the assembly of a community and over its subsequent lifespan filters with diversifying and convergent effects may operate simultaneously on recruitment from the local species pool and impose contrasted effects on the similarity of the trait values exhibited by co‐existing species. Moreover, as a consequence of the frequent association of productivity with the convergence filter, an additional difference is predicted in terms of the effects of the two filters on ecosystem functioning. Convergence in traits selected by the productivity filter will exert effects on both the plant community and the ecosystem while divergent effects of the disturbance filter will be restricted to the plant community.     

Above- and belowground insect herbivores differentially affect soil nematode communities in species-rich plant communities

Interactions between above‐ and belowground invertebrate herbivores alter plant diversity, however, little is known on how these effects may influence higher trophic level organisms belowground. Here we explore whether above‐ and belowground invertebrate herbivores which alter plant community diversity and biomass, in turn affect soil nematode communities. We test the hypotheses that insect herbivores 1) alter soil nematode diversity, 2) stimulate bacterial‐feeding and 3) reduce plant‐feeding nematode abundances. In a full factorial outdoor mesocosm experiment we introduced grasshoppers (aboveground herbivores), wireworms (belowground herbivores) and a diverse soil nematode community to species‐rich model plant communities. After two years, insect herbivore effects on nematode diversity and on abundance of herbivorous, bacterivorous, fungivorous and omni‐carnivorous nematodes were evaluated in relation to plant community composition. Wireworms did not affect nematode diversity despite enhanced plant diversity, while grasshoppers, which did not affect plant diversity, reduced nematode diversity. Although grasshoppers and wireworms caused contrasting shifts in plant species dominance, they did not affect abundances of decomposer nematodes at any trophic level. Primary consumer nematodes were, however, strongly promoted by wireworms, while community root biomass was not altered by the insect herbivores. Overall, interaction effects of wireworms and grasshoppers on the soil nematodes were not observed, and we found no support for bottom‐up control of the nematodes. However, our results show that above‐ and belowground insect herbivores may facilitate root‐feeding rather than decomposer nematodes and that this facilitation appears to be driven by shifts in plant species composition. Moreover, the addition of nematodes strongly suppressed shoot biomass of several forb species and reduced grasshopper abundance. Thus, our results suggest that nematode feedback effects on plant community composition, due to plant and herbivore parasitism, may strongly depend on the presence of insect herbivores.