Search

Copyright of Conservation Biology is the property of Wiley-Blackwell and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Plant nutrient uptake and productivity are driven by a multitude of factors that have been modified by human activities, like climate change and the activity of decomposers. However, interactive effects of climate change and key decomposer groups like earthworms have rarely been studied. In a field microcosm experiment, we investigated the effects of a mean future climate scenario with warming (+ 0.50 °C to + 0.62 °C) and altered precipitation (+ 10% in spring and autumn, − 20% in summer) and earthworms (anecic—two Lumbricus terrestris, endogeic—four Allolobophora chlorotica and both together within 10 cm diameter tubes) on plant biomass and stoichiometry in two land-use types (intensively used meadow and conventional farming). We found little evidence for earthworm effects on aboveground biomass. However, future climate increased above- (+40.9%) and belowground biomass (+44.7%) of grass communities, which was mainly driven by production of the dominant Festulolium species during non-summer drought periods, but decreased the aboveground biomass (− 36.9%) of winter wheat. Projected climate change and earthworms interactively affected the N content and C:N ratio of grasses. Earthworms enhanced the N content (+1.2%) thereby decreasing the C:N ratio (− 4.1%) in grasses, but only under ambient climate conditions. The future climate treatment generally decreased the N content of grasses (aboveground: − 1.1%, belowground: − 0.15%) and winter wheat (− 0.14%), resulting in an increase in C:N ratio of grasses (aboveground: + 4.2%, belowground: +6.3%) and wheat (+5.9%). Our results suggest that climate change diminishes the positive effects of earthworms on plant nutrient uptakes due to soil water deficit, especially during summer drought. [ABSTRACT FROM AUTHOR]

Biological invasions have major impacts on a variety of ecosystems and threaten native biodiversity. Earthworms have been absent from northern parts of North America since the last ice age, but non‐native earthworms were recently introduced there and are now being spread by human activities. While past work has shown that plant communities in earthworm‐invaded areas change towards a lower diversity mainly dominated by grasses, the underlying mechanisms related to changes in the biotic interactions of the plants are not well understood. Here, we used a trait‐based approach to study the effect of earthworms on interspecific plant competition and aboveground herbivory. We conducted a microcosm experiment in a growth chamber with a full‐factorial design using three plant species native to northern North American deciduous forests, Poa palustris (grass), Symphyotrichum laeve (herb) and Vicia americana (legume), either growing in monoculture or in a mixture of three. These plant community treatments were crossed with earthworm (presence or absence) and herbivore (presence or absence) treatments. Eight out of the fourteen above‐ and belowground plant functional traits studied were significantly affected by earthworms, either by a general effect or in interaction with plant species identity, plant diversity level and/or herbivore presence. Earthworms increased the aboveground productivity and the number of inflorescences of the grass P. palustris. Further, earthworms and herbivores together affected root tissue density of P. palustris and the specific leaf area of V. americana. In this study, earthworm presence gave a competitive advantage to the grass species P. palustris by inducing changes in plant functional traits. Our results suggest that invasive earthworms can alter competitive and multitrophic interactions of plants, shedding light on some of the mechanisms behind invasive earthworm‐induced plant community changes in northern North America forests. [ABSTRACT FROM AUTHOR]

Microbial necromass is a central component of soil organic matter (SOM), whose management may be essential in mitigating atmospheric CO2 concentrations and climate change. Current consensus regards the magnitude of microbial necromass production to be heavily dependent on the carbon use efficiency of microorganisms, which is strongly influenced by the quality of the organic matter inputs these organisms feed on. However, recent concepts neglect agents relevant in many soils: earthworms. We argue that the activity of earthworms accelerates the formation of microbial necromass stabilized in aggregates and organo‐mineral associations and reduces the relevance of the quality of pre‐existing organic matter in this process. Earthworms achieve this through the creation of transient hotspots (casts) characterized by elevated contents of bioavailable substrate and the efficient build‐up and quick turnover of microbial biomass, thus converting SOM not mineralized in this process into a state more resistant against external disturbances, such as climate change. Promoting the abundance of earthworms may, therefore, be considered a central component of management strategies that aim to accelerate the formation of stabilized microbial necromass in wide locations of the soil commonly not considered hotspots of microbial SOM formation. [ABSTRACT FROM AUTHOR]

Diversity and community composition of earthworms, key drivers of ecosystem functions, are increasingly threatened by global change, including climate and land-use change. However, empirical evidence for interactions of these concurrent drivers in affecting earthworm communities is scarce. Here, we investigated the effects of an experimentally imposed climate change scenario, including warming and altered precipitation patterns, and land use with two croplands (both conventional farming and organic farming characterize a three-year crop rotation) and two grasslands (intensively-used meadow and extensively-used meadow) on earthworm communities across different seasons and years in a field experiment. Compared with grasslands, earthworms in croplands have lower species richness (-26 %), abundance (-80 %), and biomass (-73 %), particularly the abundance of juveniles (-83 %) and Aporrectodea rosea (-76 %) as well as the biomass of juveniles (-84 %), A. rosea (-72 %), Octolasion cyaneum (-47 %), and Lumbricus terrestris (-83 %). Due to extreme droughts in Central Europe from 2018 to 2020, earthworm abundance and biomass were low across land-use types, but in grassland they increased (abundance: +80 %; biomass: +85 %) in 2021 presumably due to increased moisture conditions. Main effects of experimental climate change and intensified management practices as well as interaction of experimental climate change and land use on abundance and biomass of earthworms were non-significant. Notably, experimental climate change and land use interactively altered earthworm community composition, with the most pronounced difference between ambient and future climate in croplands than in grasslands. This indicates that earthworm community composition more sensitively reflects changes in environmental conditions than earthworm abundance and biomass, but the latter two negatively responded to prolonged drought conditions. Our results indicate that grasslands have a higher resilience of earthworm populations to buffer adverse environmental conditions than croplands. Overall, this study provides a comprehensive overview of the response of earthworms to inter-annual climatic variability and experimental climate change under different land-use types. • Earthworm communities in grasslands have a higher diversity than in croplands. • Earthworm communities in grasslands are more resilient to climate extremes. • Climate extremes have a stronger impact on earthworms than mean climate changes. • Climate change–land use interactions impact earthworm community composition. [ABSTRACT FROM AUTHOR]

Anthropogenic pressures alter the biodiversity, structure and organization of biological communities with severe consequences for ecosystem processes. Species invasion is such a human‐induced ecosystem change with pronounced impacts on recipient ecosystems. Around the globe, earthworms invade habitats and impact abiotic soil conditions and a wide range of above‐ and belowground organisms. In northern North America, where earthworms have been largely absent since the last glaciation period and most earthworm species present today have only been (re‐)introduced a few hundred years ago, invasion impacts have been intensively studied. However, despite several studies assessing impacts of invasive earthworms on soil fauna, studies have rarely investigated the simultaneous responses of different soil‐fauna size groups and biodiversity facets which might respond differently to earthworm invasion and independently affect ecosystem processes. Our study goes beyond previously‐established knowledge on earthworm‐invasion effects by simultaneously assessing differences in four biodiversity facets, namely the abundance, biomass, richness and Shannon index of soil invertebrate macro‐, meso‐ and microfauna communities between high‐ and low‐invasion status plots (n = 80) and in relation to invasion intensity measured as earthworm biomass across four northern North American forests sampled between 2016 and 2017. Across forests and soil‐fauna groups, we found reduced abundance (−33 to −45%) and richness (−18 to −25%) in high compared to low‐invasion status areas. Additionally, meso‐ (−14%) and microfauna biomass (−38%) and macro‐ (−7%) and microfauna Shannon index (−8%) were reduced. Higher invasion intensity (earthworm biomass) was additionally related to reduced soil‐fauna biodiversity. While the studied biodiversity facet was important for the soil fauna response, soil‐fauna size group was comparably unimportant. Given the global ubiquity of earthworm invasion and the importance of soil fauna for key ecosystem processes, our observational results help to assess future impacts of this invasion and the consequences for anthropogenically‐altered ecosystem functioning. [ABSTRACT FROM AUTHOR]

As ecosystem engineers, invasive earthworms are one of the main drivers of plant community changes in North American forests previously devoid of earthworms. One explanation for these community changes is the effects of earthworms on the reproduction, recruitment, and development of plant species. However, few studies have investigated functional trait responses of native plants to earthworm invasion to explain the mechanisms underlying community changes. In a mesocosm (Ecotron) experiment, we set up a plant community composed of two herb and two grass species commonly found in northern North American forests under two earthworm treatments (presence vs. absence). We measured earthworm effects on above- and belowground plant biomass and functional traits after 3 months of experiment. Our results showed that earthworm presence did not significantly affect plant community biomass and cover. Furthermore, only four out of the fifteen above- and belowground traits measured were affected by earthworm presence. While some traits, such as the production of ramets, the carbon and nitrogen content of leaves, responded similarly between and within functional groups in the presence or absence of earthworms, we observed opposite responses for other traits, such as height, specific leaf area, and root length within some functional groups in the presence of earthworms. Plant trait responses were thus species-specific, although the two grass species showed a more pronounced response to earthworm presence with changes in their leaf traits than herb species. Overall, earthworms affected some functional traits related to resource uptake abilities of plants and thus could change plant competition outcomes over time, which could be an explanation of plant community changes observed in invaded ecosystems. [ABSTRACT FROM AUTHOR]

Recent research shows that earthworms can alter defense traits of plants against herbivores and pathogens by affecting soil biochemistry. Yet, the effects of invasive earthworms on defense traits of native plants from previously earthworm‐free ecosystems as well as the consequences for multitrophic interactions are virtually unknown.Here we use a combination of an observational study and a complementary experimental study to investigate the effects of invasive earthworms on leaf defense traits, herbivore damage and pathogen infection in two poplar tree species (Populus balsamifera and Populus tremuloides) native to North American boreal forests.Our observational study showed that earthworm invasion was associated with enhanced leaf herbivory (by leaf‐chewing insects) in saplings of both tree species. However, we only detected significant shifts in the concentration of chemical defense compounds in response to earthworm invasion for P. balsamifera. Specifically, leaf phenolic concentrations, including salicinoids and catechin, were lower in P. balsamifera from earthworm‐invaded sites.Our experimental study confirmed an earthworm‐induced reduction in leaf defense levels in P. balsamifera for one of the defense compounds, tremulacin. The experimental study additionally showed that invasive earthworms reduced leaf dry matter content, potentially increasing leaf palatability, and enhanced susceptibility of trees to infection by a fungal pathogen, but not to aphid infestation, in the same tree species.Synthesis. Our results show that invasive earthworms can decrease the concentrations of some chemical defense compounds in P. balsamifera, which could make them susceptible to leaf‐chewing insects. Such potential impacts of invasive earthworms are likely to have implications for tree survival and competition, native tree biodiversity and ecosystem functioning. [ABSTRACT FROM AUTHOR]

The impacts of climate change on biodiversity can be modulated by other changing environmental conditions (e.g. induced by land‐use change). The potential interactive effects of climate change and land use have rarely been studied for soil organisms. To test the effects of changing climatic conditions and land use on soil invertebrates, we examined earthworm communities across different seasons in different grassland‐use types (intensively managed grassland, extensively managed meadow and extensively managed sheep pasture). We predicted that the strength of climate change effects would vary with season and land use. Overall, extracted earthworm populations showed the strongest variations in response to the season, indicating major differences in activity patterns and extraction efficiency, whereas climate change and different grassland‐use types had fewer and weaker effects. Future climate, characterized by slightly higher precipitation in spring and autumn but a strong reduction during the summer, had positive effects on the abundance of extracted adult earthworms in spring but then reduced the abundance of active earthworms across the remaining seasons. In contrast, the total biomass of juveniles tended to be consistently lower under future climate conditions. Earthworm species responded differently to the climate change and different grassland management types, and these species‐specific responses further varied strongly across seasons. Intensive grassland management had negative effects, due to plant community composition, whereas sheep grazing favoured earthworm populations, due to dung deposition. There were only limited interactive effects between climate and land use, which thus did not support our main hypothesis. Nevertheless, these results highlight the complex and context‐dependent responses of earthworm communities and activity patterns to climate change, with potential consequences for long‐term population dynamics and crucial ecosystem functions. Highlights: We explored earthworm communities in response to climate change, different grassland‐use types and seasonsClimate had species‐specific effects on active earthworms, but few interactions with land‐use typeIntensive grassland management decreased, but sheep grazing favoured, active earthworm populationsStrong seasonal variations in earthworm activity periods will be modulated by climate change [ABSTRACT FROM AUTHOR]

Aim: Lumbricid earthworms are invasive across northern North America, causing notable changes in forest ecosystems. During their range expansion, they encountered harsher climatic conditions compared to their native ranges in short time (~400 years). This study investigated if (a) dispersal barriers, (b) climatic selection or (c) anthropogenic activities, that is fishing bait disposal, structure the dispersal of free‐living earthworm populations. Location: North America, forest habitats along former Wisconsinan glaciation line. Taxon: Lumbricus terrestris, L. rubellus. Methods: Lumbricus terrestris and L. rubellus co‐occur in the same habitats but differ in ecology and use as fishing bait. Both species were sampled in five transects ranging from the east to the west coast of northern North America, including major dispersal barriers, three different climate zones, and bait shops near sampling locations. Genetic diversity and structure were compared between the two species, and the presence of free‐living bait shop genotypes was assessed using four markers (COI, 16S rDNA, 12S rDNA, and H3). Results: Populations of both species were genetically diverse with some geographic structure, which was more pronounced in L. terrestris than in L. rubellus. Common haplotypes were present in all regions, but locally restricted haplotypes also occurred. Furthermore, two distinct genetic clades of L. terrestris co‐occurred only in the two most distant transects (Alberta and Minnesota). Genotypes identical to bait individuals were omnipresent in field populations of L. terrestris. Main Conclusions: Genetic diversity was high in both species, and invasive populations represented a genetic subset of European earthworms. Geographic and climatic dispersal barriers affected the less mobile species, L. terrestris, resulting in differences in genetic structure between the two species. Our results indicate common long‐distance dispersal vectors and vectors affecting only L. terrestris. The roles of climate and anthropogenic activities are discussed, providing additional explanations of dispersal and new insights into establishment of invasive earthworm populations. [ABSTRACT FROM AUTHOR]

Recent studies have shown that invasive earthworms can dramatically reduce native biodiversity, both above and below the ground. However, we still lack a synthetic understanding of the underlying mechanisms behind these changes, such as whether earthworm effects on soil chemical properties drive such relationships. Here, we investigated the effects of invasive earthworms on soil chemical properties (pH, water content, and the stocks and fluxes of carbon, nitrogen, and phosphorus) by conducting a meta‐analysis. Invasive earthworms generally increased soil pH, indicating that the removal of organic layers and the upward transport of more base‐rich mineral soil caused a shift in soil pH. Moreover, earthworms significantly decreased soil water content, suggesting that the burrowing activities of earthworms may have increased water infiltration of and/or increased evapotranspiration from soil. Notably, invasive earthworms had opposing effects on organic and mineral soil for carbon and nitrogen stocks, with decreases in organic, and increases in mineral soil. Nitrogen fluxes were higher in mineral soil, whereas fluxes in organic soil were not significantly affected by the presence of invasive earthworms, indicating that earthworms mobilize and redistribute nutrients among soil layers and increase overall nitrogen loss from the soil. Invasive earthworm effects on element stocks increased with ecological group richness only in organic soil. Earthworms further decreased ammonium stocks with negligible effects on nitrate stocks in organic soil, whereas they increased nitrate stocks but not ammonium stocks in mineral soil. Notably, all of these results were consistent across forest and grassland ecosystems underlining the generality of our findings. However, we found some significant differences between studies that were conducted in the field (observational and experimental settings) and in the lab, such as that the effects on soil pH decreased from field to lab settings, calling for a careful interpretation of lab findings. Our meta‐analysis provides strong empirical evidence that earthworm invasion may lead to substantial changes in soil chemical properties and element cycling in soil. Furthermore, our results can help explain the dramatic effects of invasive earthworms on native biodiversity, for example, shifts towards the dominance of grass species over herbaceous ones, as shown by recent meta‐analyses. [ABSTRACT FROM AUTHOR]

Non‐native, invasive earthworms are altering soils throughout the world. Ecological cascades emanating from these invasions stem from rapid consumption of leaf litter by earthworms. This occurs at a midpoint in the trophic pyramid, unlike the more familiar bottom‐up or top‐down cascades. These cascades cause fundamental changes ("microcascade effects") in soil morphology, bulk density, and nutrient leaching, and a shift to warmer, drier soil surfaces with a loss of leaf litter. In North American temperate and boreal forests, microcascade effects can affect carbon sequestration, disturbance regimes, soil and water quality, forest productivity, plant communities, and wildlife habitat, and can facilitate other invasive species. These broader‐scale changes ("macrocascade effects") are of greater concern to society. Interactions among these fundamental changes and broader‐scale effects create "cascade complexes" that interact with climate change and other environmental processes. The diversity of cascade effects, combined with the vast area invaded by earthworms, leads to regionally important changes in ecological functioning. [ABSTRACT FROM AUTHOR]

• Warming and earthworms interactively affect soil biota. • Earthworms buffer detrimental warming effects on soil biodiversity. • Declining earthworm densities alter the structure of soil food webs. Anthropogenic climate change is altering the functioning of terrestrial ecosystems. Agricultural systems are particularly vulnerable to climate change as they are frequently disturbed by intensified management practices. This also threatens belowground organisms that are responsible for providing crucial ecosystem functions and services, such as nutrient cycling and plant disease suppression. Amongst these organisms, earthworms are of particular importance as they can modulate the effects of climate change on soil organisms by modifying the biotic and abiotic soil conditions. However, they are also known to decline under intensified management, justifying their use as key biotic indicators of intensified agriculture. Yet, our knowledge of the responses of belowground species to the interacting effects of warming and land-use intensification (simulated by earthworm reduction in the experimental setup) remains limited. Here, we tested the interactive effects of soil warming and reduced earthworm densities on soil protists, nematodes, meso- and macrofauna, and their diversity in a common barley system in the Hohenheim Climate Change Experiment. We found that belowground species richness was lowest at elevated temperature and reduced earthworm densities, indicating that earthworms can buffer warming effects on belowground biodiversity. Furthermore, warming increased the densities of plant-feeding nematodes, and herbivorous macrofauna benefitted from reduced earthworm densities. Our results indicate that warming and reduced earthworm densities may simultaneously modify the functioning and service provisioning of soils via shifts in diversity and density of soil biota that would likely lead to simplified belowground food webs. These findings thus highlight the importance of maintaining greater densities of ecosystem engineers like earthworms that may help buffering the detrimental effects of climate warming in agricultural systems. [ABSTRACT FROM AUTHOR]

Earth is experiencing a substantial loss of biodiversity at the global scale, while both species gains and losses are occurring at local and regional scales. The influence of these nonrandom changes in species distributions could profoundly affect the functioning of ecosystems and the essential services that they provide. However, few experimental tests have been conducted examining the influence of species invasions on ecosystem functioning. Even fewer have been conducted using invasive ecosystem engineers, which can have disproportionately strong influence on native ecosystems relative to their own biomass. The invasion of exotic earthworms is a prime example of an ecosystem engineer that is influencing many ecosystems around the world. In particular, European earthworm invasions of northern North American forests cause simultaneous species gains and losses with significant consequences for essential ecosystem processes like nutrient cycling and crucial services to humanity like soil erosion control and carbon sequestration. Exotic earthworms are expected to select for specific traits in communities of soil microorganisms (fast-growing bacteria species), soil fauna (promoting the bacterial energy channel), and plants (graminoids) through direct and indirect effects. This will accelerate some ecosystem processes and decelerate others, fundamentally altering how invaded forests function. This project aims to investigate ecosystem responses of northern North American forests to earthworm invasion. Using a novel, synthetic combination of field observations, field experiments, lab experiments, and meta-analyses, the proposed work will be the first systematic examination of earthworm effects on (1) plant communities and (2) soil food webs and processes. Further, (3) effects of a changing climate (warming and reduced summer precipitation) on earthworm performance will be investigated in a unique field experiment designed to predict the future spread and consequences of earthworm invasion in North America. By assessing the soil chemical and physical properties as well as the taxonomic (e.g., by the latest next-generation sequencing techniques) and functional composition of plant, soil microbial and animal communities and the processes they drive in four forests, work packages I-III take complementary approaches to derive a comprehensive and generalizable picture of how ecosystems change in response to earthworm invasion. Finally, in work package IV, meta-analyses will be used to integrate the information from work packages I-III and existing literature to investigate if earthworms cause invasion waves, invasion meltdowns, habitat homogenization, and ecosystem state shifts. Global data will be synthesized to test if the relative magnitude of effects differs from place to place depending on the functional dissimilarity between native soil fauna and exotic earthworms. Moving from local to global scale, the present proposal examines the influence of earthworm invasions on biodiversity-ecosystem functioning relationships from an aboveground-belowground perspective in natural settings. This approach is highly innovative as it utilizes the invasion by exotic earthworms as an exciting model system that links invasion biology with trait-based community ecology, global change research, and ecosystem ecology, pioneering a new generation of biodiversity-ecosystem functioning research. [ABSTRACT FROM AUTHOR]

Bottom-up effects of plants on soil communities can be modified by the activity of exotic earthworms, by altering resource availability for soil food webs through feeding, burrowing, and casting activities. The present study explored effects of plants (planting of shrubs) on soil micro-food webs (composition of soil microbial and nematode communities), and whether these effects were altered by the activity of exotic earthworms (exotic earthworms addition). Planted shrubs resulted in a non-significant increase of bacterial biomass and significantly increased the abundance of different nematode trophic groups and total nematode biomass, indicating that planted shrubs had significant bottom-up effects on soil bacteria and nematodes. Planted shrubs decreased nematode diversity, evenness, and richness, but increased nematode dominance in the plots where the abundance of exotic earthworms was not amended. By contrast, these effects of shrub presence on soil biodiversity were not found in the plots that received exotic earthworms. In addition, planted shrubs increased the total energy flux to the nematode community. By contrast, the elevated activity of exotic earthworms mitigated the increase in total energy flux to nematodes in the presence of shrubs, and increased the ratio of fungal to bacterial PLFAs. Both of these changes indicate reduced energy flux in the plots with added exotic earthworms. Nematode diversity decreased, while nematode dominance increased with increasing total energy flux to nematodes, probably because few species benefited from high energy flux. Our study indicates that exotic earthworms can maintain soil biodiversity by reducing the energy flux through soil food webs. [ABSTRACT FROM AUTHOR]

Biological invasions pose a serious threat to biodiversity and ecosystem functioning across ecosystems. Invasions by ecosystem engineers, in particular, have been shown to have dramatic effects in recipient ecosystems. For instance, invasion by earthworms, a below-ground invertebrate ecosystem engineer, in previously earthworm-free ecosystems alters the physico-chemical characteristics of the soil. Studies have shown that such alterations in the soil can have far-reaching impacts on soil organisms, which form a major portion of terrestrial biodiversity., Here, we present the first quantitative synthesis of earthworm invasion effects on soil micro-organisms and soil invertebrates based on 430 observations from 30 independent studies., Our meta-analysis shows a significant decline of the diversity and density of soil invertebrates in response to earthworm invasion with anecic and endogeic earthworms causing the strongest effects. Earthworm invasion effects on soil micro-organisms were context-dependent, such as depending on functional group richness of invasive earthworms and soil depth. Microbial biomass and diversity increased in mineral soil layers, with a weak negative effect in organic soil layers, indicating that the mixing of soil layers by earthworms (bioturbation) may homogenize microbial communities across soil layers., Our meta-analysis provides a compelling evidence for negative effects of a common invasive below-ground ecosystem engineer on below-ground biodiversity of recipient ecosystems, which could potentially alter the ecosystem functions and services linked to soil biota. [ABSTRACT FROM AUTHOR]

Human activities have been a significant driver of environmental changes with tremendous consequences for carbon (C) dynamics. Peatlands are critical ecosystems because they store ~30% of the global soil organic C pool and are particularly vulnerable to anthropogenic changes. The Zoige peatland on the eastern Tibet Plateau, as the largest alpine peatland in the world, accounts for 1‰ of global peat soil organic C storage. However, this peatland has experienced dramatic climate change including increased temperature and reduced precipitation in the past decades, which likely is responsible for a decline of the water-table and facilitated earthworm invasion, two major factors reducing soil organic carbon ( SOC) storage of peatlands., Because earthworms often are more active in low- than in high-moisture peatlands, we hypothesized that the simultaneous occurrence of water-table decline and earthworm invasion would synergistically accelerate the release of SOC from peatland soil. We conducted a field experiment with a paired split-plot design, i.e. presence vs. absence of the invasive earthworms ( Pheretima aspergillum) nested in drained vs. undrained plots, respectively, for 3 years within the homogenous Zoige peatland., Water-table decline significantly decreased soil water content and bulk density, resulting in a marked reduction of SOC storage. Moreover, consistent with our hypothesis, earthworm presence dramatically reduced SOC in the drained but not in the undrained peatland through the formation of deep burrows and decreasing bulk density of the lower soil layer over 3 years. The variation in SOC likely was due to changes in above-ground plant biomass, root growth and earthworm behaviour induced by the experimental treatments., Synthesis and applications. We suggest that incentive measures should be taken to prevent further water-table decline and earthworm invasion for maintaining the soil carbon pool in Zoige peatland. Artificial filling of drainage canals should be implemented to increase the water-table level, facilitating the recovery of drained peatlands. Moreover, the dispersal of earthworms and their cocoons attached to the roots of crop plants and tree saplings from low-lying areas to the Zoige region should be prevented. [ABSTRACT FROM AUTHOR]

The activity and spread of exotic earthworms often are spatially correlated with N deposition because both arise from human activities. Exotic earthworms, in turn, can also greatly affect soil abiotic and biotic properties, as well as related ecological processes. Previous studies showed, for example, that earthworms can counteract the detrimental effects of plant-feeding nematodes on plant growth. However, potential interactive effects of N deposition and exotic earthworms on ecosystems are poorly understood., We explored the changes in density of plant-feeding nematodes in response to the presence of exotic earthworms, and whether these changes are altered by elevated N deposition in a two-factorial field mesocosm experiment at the Heshan National Field Research Station of Forest Ecosystem, in southern China., Our results show that earthworm addition marginally significantly increased the density of exotic earthworms and significantly increased the mass of earthworm casts. The total density of plant-feeding nematodes was not significantly affected by exotic earthworms or N deposition. However, exotic earthworms tended to increase the density of plant-feeding nematode taxa that are less detrimental to plant growth (r-strategists), while they significantly reduced the density of more harmful plant-feeding nematodes (K-strategists). Importantly, these earthworm effects were restricted to the ambient N deposition treatment, and elevated N deposition cancelled out the earthworm effect. Although exotic earthworms and N deposition interactively altered foliar N : P ratio in the target tree species, this did not result in significant changes in shoot and root biomass in the short term., Overall, our study indicates that N deposition can cancel out exotic earthworm-induced reductions in the density of harmful plant-feeding nematodes. These results suggest that anthropogenic N deposition can alter biotic interactions between exotic and native soil organisms with potential implications for ecosystem functioning. [ABSTRACT FROM AUTHOR]

Aims: The passage of seeds through the earthworm gut potentially damages seeds, altering seed and seedling performances depending on seed traits. This work was conducted to study to what extent chemical and morphological seed traits determine the seed attractiveness for earthworms. Methods: We tested seed selection via the ingestion and digestion of 23 grassland plant species spanning a range of 14 morphological and chemical traits by two common earthworm species: the anecic Lumbricus terrestris and the endogeic Allolobophora chlorotica. Results: Both earthworm species ingested seeds from all plant species. A. chlorotica digested almost all ingested seeds (out of the 15 % ingested), whereas L. terrestris excreted them in varying quantities (out of the 86 % ingested), depending on plant species identity. Seed ingestion rate by L. terrestris was driven by seed oil content and earthworm initial weight. The apparent effect of seed length was explained via seed oil content. Seed digestion rate by L. terrestris was negatively impacted by seed size. Seed ingestion rate by A. chlorotica tended to be impacted by seed protein content and seed length. Conclusion: Earthworms-seed interactions depend on a variety of seed traits and earthworm identity. Thus, earthworms, via their specific feeding behavior, might facilitate or impede the regeneration of certain plant species and drive plant communities. [ABSTRACT FROM AUTHOR]

Management of biodiversity and ecosystem services requires a better understanding of the factors that influence soil biodiversity. We characterized the species (or genera) richness of 10 taxonomic groups of invertebrate soil animals in replicated monocultures of 14 temperate tree species. The focal invertebrate groups ranged from microfauna to macrofauna: Lumbricidae, Nematoda, Oribatida, Gamasida, Opilionida, Araneida, Collembola, Formicidae, Carabidae, and Staphylinidae. Measurement of invertebrate richness and ancillary variables occurred ~34 years after the monocultures were planted. The richness within each taxonomic group was largely independent of richness of other groups; therefore a broad understanding of soil invertebrate diversity requires analyses that are integrated across many taxa. Using a regression-based approach and ~125 factors related to the abundance and diversity of resources, we identified a subset of predictors that were correlated with the richness of each invertebrate group and richness integrated across 9 of the groups (excluding earthworms). At least 50% of the variability in integrated richness and richness of each invertebrate group was explained by six or fewer predictors. The key predictors of soil invertebrate richness were light availability in the understory, the abundance of an epigeic earthworm species, the amount of phosphorus, nitrogen, and calcium in soil, soil acidity, and the diversity or mass of fungi, plant litter, and roots. The results are consistent with the hypothesis that resource abundance and diversity strongly regulate soil biodiversity, with increases in resources (up to a point) likely to increase the total diversity of soil invertebrates. However, the relationships between various resources and soil invertebrate diversity were taxon-specific. Similarly, diversity of all 10 invertebrate taxa was not high beneath any of the 14 tree species. Thus, changes to tree species composition and resource availability in temperate forests will likely increase the richness of some soil invertebrates while decreasing the richness of others. [ABSTRACT FROM AUTHOR]

Common buckthorn ( Rhamnus cathartica L.) is one of the most abundant and ecologically harmful non-native plants in forests of the Upper Midwest United States. At the same time, European earthworms are invading previously glaciated areas in this region, with largely anecdotal evidence suggesting they compound the negative effects of buckthorn and influence the invasibility of these forests. Germination and seedling establishment are important control points for colonization by any species, and manipulation of the conditions influencing these life history stages may provide insight into why invasive species are successful in some environments and not others. Using a greenhouse microcosm experiment, we examined the effects of important biotic and abiotic factors on the germination and seedling establishment of common buckthorn. We manipulated light levels, leaf litter depth and earthworm presence to investigate the independent and interactive effects of these treatments on buckthorn establishment. We found that light and leaf litter depth were significant predictors of buckthorn germination but that the presence of earthworms was the most important factor; earthworms interacted with light and leaf litter to increase the number and biomass of buckthorn across all treatments. Path analysis suggested both direct and moisture-mediated indirect mechanisms controlled these processes. The results suggest that the action of earthworms may provide a pathway through which buckthorn invades forests of the Upper Midwest United States. Hence, researchers and managers should consider co-invasion of plants and earthworms when investigating invasibility and creating preemptive or post-invasion management plans. [ABSTRACT FROM AUTHOR]

Background: Recent studies suggested that the earthworm Lumbricus terrestris might act as a seedling predator by ingesting emerging seedlings, and individuals were observed damaging fresh leaves of various plant species in the field. To evaluate the significance of herbivore behavior of L. terrestris for plant and earthworm performance we exposed 23- to 33-days-old seedlings of six plant species to earthworms in two microcosm experiments. Plants belonged to the three functional groups grasses, non-leguminous herbs, and legumes. Leaf damage, leaf mortality, the number of leaves as well as mortality and growth of seedlings were followed over a period of up to 26 days. In a subset of replicates 0.1 g of soil surface litter of each of the six plant species was provided and consumption was estimated regularly to determine potential feeding preferences of earthworms. Results: There was no difference in seedling growth, the number of live seedlings and dead leaves between treatments with or without worms. Fresh leaves were damaged eight times during the experiment, most likely by L. terrestris, with two direct observations of earthworms tearing off leaf parts. Another nine leaves were partly pulled into earthworm burrows. Lumbricus terrestris preferred to consume legume litter over litter of the other plant functional groups. Earthworms that consumed litter lost less weight than individuals that were provided with soil and live plants only, indicating that live plants are not a suitable substitute for litter in earthworm nutrition. Conclusion: Our results demonstrate that L. terrestris damages live plants; however, this behavior occurs only rarely. Pulling live plants into earthworm burrows might induce microbial decomposition of leaves to make them suitable for later consumption. Herbivory on plants beyond the initial seedling stage may only play a minor role in earthworm nutrition and has limited potential to influence plant growth. [ABSTRACT FROM AUTHOR]

Climate change causes species range shifts and potentially alters biological invasions. The invasion of European earthworm species across northern North America has severe impacts on native ecosystems. Given the long and cold winters in that region that to date supposedly have slowed earthworm invasion, future warming is hypothesized to accelerate earthworm invasions into yet non-invaded regions. Alternatively, warming-induced reductions in soil water content (SWC) can also decrease earthworm performance. We tested these hypotheses in a field warming experiment at two sites in Minnesota, USA by sampling earthworms in closed and open canopy in three temperature treatments in 2010 and 2012. Structural equation modeling revealed that detrimental warming effects on earthworm densities and biomass could indeed be partly explained by warming-induced reductions in SWC. The direction of warming effects depended on the current average SWC: warming had neutral to positive effects at high SWC, whereas the opposite was true at low SWC. Our results suggest that warming limits the invasion of earthworms in northern North America by causing less favorable soil abiotic conditions, unless warming is accompanied by increased and temporally even distributions of rainfall sufficient to offset greater water losses from higher evapotranspiration. [ABSTRACT FROM AUTHOR]

European earthworms are invading many ecosystems worldwide and fundamentally transform habitats by acting as dominant ecosystem engineers. However, there is little knowledge of the consequences of earthworm invasion on the composition and diversity of native soil organisms. Particularly functionally similar groups, such as enchytraeids (Annelida: Enchytraeidae), may be affected through changes in the chemical and physical properties of the soil, but also due to competition for resources. In 2010–2011, we studied the impact of earthworm invasion on enchytraeids at two sites in the northern hardwood forests of North America: one site within the Chippewa National Forest in northern Minnesota and one site in the Chequamegon-Nicolet National Forest in northern Wisconsin, USA. At each site, three plots were sampled along a transect, representing (1) a non-invaded or very slightly invaded area, (2) the leading edge of earthworm invasion and (3) a heavily invaded area with an established population of the anecic earthworm Lumbricus terrestris (among other species). In total, 29 enchytraeid (morpho)species were identified (some yet to be formally described, several first or second records for the continent); of those 24 occurred at the Minnesota site and 17 at the Wisconsin site. The structure of enchytraeid assemblages differed significantly among the three invasion stages, although this was not equally pronounced at the two sites. Each stage was characterized by one or several indicator species. Mean enchytraeid densities (10,700–30,400 individuals/m 2 ) did not differ significantly among the invasion stages, but were lowest at the leading edge of earthworm invasion at both sites. In the heavily invaded plot at the Minnesota site, the mean enchytraeid density and biomass in L. terrestris middens were significantly higher than in soil in-between the middens. This was due to a pronounced effect of L. terrestris middens in the uppermost 3 cm of soil. Differences in biomass among earthworm invasion stages were most apparent for mean individual biomass. This was significantly higher in the heavily invaded area than at the leading edge or in the non-invaded area at the Minnesota site. Compositional changes of the enchytraeid assemblage are likely to result in changes in the functioning of soil foods webs. Our results suggest that earthworm invasions can cause a loss of native species in soil, including heretofore unknown ones, that might go unnoticed. [ABSTRACT FROM AUTHOR]

Background: Infiltration is a key process in determining the water balance, but so far effects of earthworms, soil texture, plant species diversity and their interaction on infiltration capacity have not been studied. Methodology/Principal Findings: We measured infiltration capacity in subplots with ambient and reduced earthworm density nested in plots of different plant species (1, 4, and 16 species) and plant functional group richness and composition (1 to 4 groups; legumes, grasses, small herbs, tall herbs). In summer, earthworm presence significantly increased infiltration, whereas in fall effects of grasses and legumes on infiltration were due to plant-mediated changes in earthworm biomass. Effects of grasses and legumes on infiltration even reversed effects of texture. We propose two pathways: (i) direct, probably by modifying the pore spectrum and (ii) indirect, by enhancing or suppressing earthworm biomass, which in turn influenced infiltration capacity due to change in burrowing activity of earthworms. Conclusions/Significance: Overall, the results suggest that spatial and temporal variations in soil hydraulic properties can be explained by biotic processes, especially the presence of certain plant functional groups affecting earthworm biomass, while soil texture had no significant effect. Therefore biotic parameters should be taken into account in hydrological applications. [ABSTRACT FROM AUTHOR]

Agri-environmental schemes involving organic farming or set-aside management aim at promoting biodiversity and restoring ecosystem functioning in agrarian landscapes. Application of pesticides in these crop fields is strongly regulated facilitating the spread of weeds but also allowing for the establishment of endangered herbs and a variety of animals. Recent studies found gastropods and earthworms to be legitimate dispersers of seeds of wild plants. We assumed that both groups also play a significant role in the spread and establishment of wild plants within crop fields. Therefore, we are conducting a series of experiments in three different study systems on (1) the role of earthworms and gastropods as dispersers of rare herbs and weeds in an organic rye field in Germany, (2) the seed feeding behavior of gastropods of plants sown in fallow ground in Switzerland, and (3) weed dispersal in irrigated rice fields by golden apple snails in the Philippines. [ABSTRACT FROM AUTHOR]

Abstract: 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. [Copyright &y& Elsevier]

Abstract: Non-trophic interactions are shaping soil food web structure and functions. Particularly, the action of ecosystem engineers, such as earthworms, are likely to fundamentally impact the abiotic and biotic properties of their environment. The present study aimed to identify the main mechanisms through which earthworms belonging to varying ecological groups – epigeic, endogeic and anecic species – affect soil microarthropods by reviewing the literature on this topic and by performing meta-analyses. Earthworm ecological groups differed considerably in their impacts on microarthropods, whereas effects did not vary significantly between microarthropod taxa at the habitat scale. Inconsistent impacts of epigeic species on soil microarthropods are most likely due to differences in earthworm densities. Effects can thus be positive in the case of moderate densities or negative in the case of high densities and associated distinct changes in the physical structure of the upper soil organic layers. By contrast, impacts of endogeic earthworms appeared to be mainly negative and were primarily due to competition with microarthropods for food resources. Consequently, negative impacts on soil microarthropods intensified with increasing earthworm density and biomass. This interaction between endogeic earthworms and microarthropods is better referred to as amensalism due to the competitive predominance of earthworms. Impacts of anecic earthworm species differed significantly from that of endogeic ones; they were neutral at the habitat scale and positive on the microhabitat scale. Moreover, impacts were independent of earthworm densities due to the quasi-territorial behaviour of anecic earthworms. Positive effects were mainly attributed to the formation of stable microhabitats by anecic species; namely burrows/middens, rich in nutrients and microorganisms. The present study points to the relevance of the non-trophic biotic interactions that drive the composition of belowground food webs by identifying the most essential mechanisms underlying the impacts of animal ecosystem engineers on soil microarthropods. Moreover, as earthworms emerge as important biological invaders, the results of the present study may help to fully appreciate, estimate and model the consequences of this momentous global change phenomenon. Particularly, the spread of exotic epigeic and endogeic earthworm species likely threatens soil microarthropod density, diversity and functions. [Copyright &y& Elsevier]

Abstract: Anecic earthworms have been shown to collect, concentrate and bury seeds in their burrows. Moreover, recent studies suggest that earthworms function as granivores and seedling herbivores thereby directly impacting plant community assembly. However, this has not been proven unequivocally. Further, it remains unclear if earthworms benefit from seed ingestion, i.e., if they assimilate seed carbon. We set up a series of three laboratory experiments in order to test the following hypotheses: (1) anecic earthworms (Lumbricus terrestris L.) not only ingest seeds but also seedlings, (2) ingestion of seedlings is lower than that of seeds due to a ‘size refuge’ of seedlings (i.e., they are too big to be swallowed), and (3) seeds and seedlings contribute to earthworm nutrition. L. terrestris readily consumed legume seedlings in the radicle stage, whereas legume seeds and seedlings in the cotyledon stage, and grass seeds and seedlings in the radicle and cotyledon stage were ingested in similar but lower amounts. Importantly, ingestion of seedlings, in contrast to seeds, was lethal for all plant species. Moreover, earthworm weight change varied with the functional identity and vitality of seeds and natural 15N signatures in earthworm body tissue underlined the importance of seedlings for earthworm nutrition. The results indicate that the anecic earthworm L. terrestris indeed functions as a granivore and seedling herbivore. The selectivity in seedling ingestion points at the potential of direct earthworm effects on plant community assembly. Further, seeds and seedlings most likely contribute significantly to earthworm nutrition potentially explaining the collection and concentration of seeds by L. terrestris in its middens and burrows; however, the present results call for experiments under more natural conditions. [Copyright &y& Elsevier]

Copyright of Basic & Applied Ecology is the property of Urban & Fischer Verlag and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Abstract: 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. [Copyright &y& Elsevier]

Abstract: Knowledge of the role of decomposers in the plant diversity–productivity relationship is scarce. In the framework of the Jena Experiment, we observed regrowth of grassland plant communities varying in plant species and functional group richness three weeks after mowing. We investigated earthworm subplots and subplots with reduced earthworm density in order to explore if earthworms enhance plant regrowth and if earthworm effects depend on plant diversity. Earthworms significantly enhanced each of the plant regrowth parameters (plant coverage and maximum and average height of the vegetation) suggesting that particularly fast growing species, such as grasses, benefit from earthworm activity. However, the average height of the vegetation was not affected in 16-species mixtures suggesting compensation of the impact of earthworms on plant regrowth in complex plant communities. [Copyright &y& Elsevier]

Summary: The soil seed bank functions to escape unfavourable environmental conditions and seed predation. Anecic earthworms are increasingly recognised as important dispersers and predators of plant seeds. In contrast, the role of the usually more abundant endogeic earthworms which live and feed in the soil on plant seeds is largely unknown. We tested whether (A) endogeic earthworms (Aporrectodea rosea, Allolobophora chlorotica, Octolasion tyrtaeum, Aporrectodea caliginosa) ingest and digest grassland plant seeds (Phleum pratense, Bellis perennis, Trifolium repens, Poa trivialis, Plantago lanceolata, Medicago varia), (B) the passage of seeds through the gut of endogeic earthworms modifies seed germination, and (C) excreta (mucus and casts) of endogeic earthworms modify seed germination. As a reference effects of the well-studied anecic species Lumbricus terrestris were determined. Endogeic earthworms ingested and digested all of the studied plant seeds; however, both ingestion and digestion were earthworm and plant species specific. Moreover, passage through the gut of endogeic earthworms and their excreta modified plant seed germination (gut passage: B. perennis; excreta: Ph. pratense and Pl. lanceolata). The results indicate that endogeic earthworms may strongly impact the composition of the soil seed bank and, consequently, plant community assembly via direct and indirect effects on plant seeds. Since post-dispersal seed predation is a key factor for the structure of plant communities, with the effect on seed survival potentially exceeding that of pre-dispersal predation, seed predation and changes in germination of seeds by endogeic earthworm species is likely to drive plant community composition. [Copyright &y& Elsevier]

Abstract: Earthworms and arbuscular mycorrhizal fungi (AMF) might interactively impact plant productivity; however, previous studies reported inconsistent results. We set up a three-factorial greenhouse experiment to study the effects of earthworms (Aporrectodea caliginosa Savigny and Lumbricus terrestris L.) and AMF (Glomus intraradices N.C. Schenck & G.S. Sm.) on the performance (productivity and shoot nutrient content) of plant species (Lolium perenne L., Trifolium pratense L. and Plantago lanceolata L.) belonging to the three functional groups grasses, legumes and herbs, respectively. Further, we investigated earthworm performance and plant root mycorrhization as affected by the treatments. Our results accentuate the importance of root derived resources for earthworm performance since earthworm weight (A. caliginosa and L. terrestris) and survival (L. terrestris) were significantly lower in microcosms containing P. lanceolata than in those containing T. pratense. However, earthworm performance was not affected by AMF, and plant root mycorrhization was not modified by earthworms. Although AMF effectively competed with T. pratense for soil N (as indicated by δ 15N analysis), AMF enhanced the productivity of T. pratense considerably by improving P availability. Remarkably, we found no evidence for interactive effects of earthworms and AMF on the performance of the plant species studied. This suggests that interactions between earthworms and AMF likely are of minor importance. [Copyright &y& Elsevier]

Abstract: Grasses and legumes are grown together worldwide to improve total herbage yield and the quality of forage, however, the causes of population oscillations of grasses and legumes are poorly understood. Especially in grasslands, earthworms are among the most important detritivore animals functioning as ecosystem engineers, playing a key role in nutrient cycling and affecting plant nutrition and growth. The objectives of the present greenhouse experiment were to quantify the effects of earthworms on grass–legume competition in model grassland systems at two harvesting dates – simulating the widespread biannual mowing regime in Central European grasslands. The presence of earthworms increased the productivity of grasses and legumes after 6 weeks but only that of grasses after another 10 weeks. In mixed treatments, the presence of grasses and earthworms decreased legume shoot biomass, the amount of nitrogen (N) in shoot tissue and the number of legume flowerheads while the presence of legumes and earthworms increased the amount of N in grass shoots and the infestation of grasses with aphids. Analyses of 15N/14N ratios indicate that, compared to legumes, grasses more efficiently exploit soil mineral N and benefit from legume presence through reduced “intra-functional group” competition. In contrast to previous experiments, we found no evidence for N transfer from legumes to grasses. However, legume presence improved total herbage and N yield. Earthworms likely modulate the competition between grasses and legumes by increasing soil N uptake by plants and thereby increasing the competitive strength of grasses. Earthworms function as essential driving agents of grass–legume associations by (I) increasing grass yield, (II) increasing the amount of N in grass hay, (III) increasing the infestation rate of grasses with aphids, and (IV) potentially reducing the attractiveness of grass–legume associations to pollinators. [Copyright &y& Elsevier]

Background: Invasions of natural communities by non-indigenous species are currently rated as one of the most important global-scale threats to biodiversity. Biodiversity itself is known to reduce invasions and increase stability. Disturbances by ecosystem engineers affect the distribution, establishment, and abundance of species but this has been ignored in studies on diversity-invasibility relationships. Methodology/Principal Findings: We determined natural plant invasion into 46 plots varying in the number of plant species (1, 4, and 16) and plant functional groups (1, 2, 3, and 4) for three years beginning two years after the establishment of the Jena Experiment. We sampled subplots where earthworms were artificially added and others where earthworm abundance was reduced. We also performed a seed-dummy experiment to investigate the role of earthworms as secondary seed dispersers along a plant diversity gradient. Horizontal dispersal and burial of seed dummies were significantly reduced in subplots where earthworms were reduced in abundance. Seed dispersal by earthworms decreased with increasing plant species richness and presence of grasses but increased in presence of small herbs. These results suggest that dense vegetation inhibits the surface activity of earthworms. Further, there was a positive relationship between the number of earthworms and the number and diversity of invasive plants. Hence, earthworms decreased the stability of grassland communities against plant invasion. Conclusions/Significance: Invasibility decreased and stability increased with increasing plant diversity and, most remarkably, earthworms modulated the diversity-invasibility relationship. While the impacts of earthworms were unimportant in low diverse (low earthworm densities) and high diverse (high floral structural complexity) plant communities, earthworms decreased the stability of intermediate diverse plant communities against plant invasion. Overall, the results document that fundamental processes in plant communities like plant seed burial and invader establishment are modulated by soil fauna calling for closer cooperation between soil animal and plant ecologists. [ABSTRACT FROM AUTHOR]

Abstract: Reliable non-destructive extraction methods are required for the assessment of the size and composition of earthworm communities where physical disturbances are not acceptable. The aim of the present study was to investigate the efficiency of the electrical octet method and the mustard extraction method for sampling of different ecological groups of earthworms (anecics, endogeics and epigeics) under dry soil conditions. We hypothesized that: (1) the extraction efficiency of the mustard method and the octet method will vary with ecological earthworm group; and (2) beforehand water addition to dry soil will increase the extraction efficiency of the octet method but not that of the mustard method. Endogeic earthworm species were extracted in low numbers irrespective of the extraction method indicating their inactivity during dry periods. The mustard method was more efficient for the extraction of anecic earthworms even under dry soil conditions, whereas the octet method was inappropriate in reflecting the actual earthworm community structure. Surprisingly, the efficiency of both methods was not improved by beforehand water addition. These findings are essential to be considered when working under dry soil conditions e.g. in the context of environmental monitoring. [Copyright &y& Elsevier]

Abstract: Anecic earthworm species function as ecosystem engineers by structuring the soil environment, incorporating large amounts of litter and seeds into soil and, thereby influence the composition of plant communities. The aim of the present greenhouse experiment was to investigate the effects of three apparently anecic earthworm species on wheat seed burial, seedling establishment, wheat growth and litter incorporation. The three species differed substantially in their behavior and effect on plant establishment. Aporrectodea longa did not incorporate litter into the soil while Lumbricus terrestris (−69%) and Lumbricus rubellus friendoides (−75%) reduced the litter layer considerably during 9 weeks of incubation. Moreover, L. terrestris and L. rubellus friendoides buried more wheat seeds than A. longa. Fewer seeds germinated when buried by A. longa compared to L. terrestris. The behavior of L. terrestris and L. rubellus friendoides was characteristic for anecic earthworm species whereas that of A. longa rather resembled that of endogeic species. The present study is the first experimental evidence for anecic behavior in L. rubellus friendoides. [Copyright &y& Elsevier]

Abstract: Ecosystems of northern North America existed without earthworm fauna until European settlers arrived and introduced European species. The current extent of invasion by some of these species, Lumbricus terrestris L., Octolasion tyrtaeum Savigny and Dendrobaena octaedra Savigny, into an aspen forest in the Canadian Rocky Mountains and the effects of the invasion on soil chemistry, microflora, soil microarthropods and vegetation were investigated. Densities of earthworm species, soil structure, plant coverage and abundance were determined along three transects starting at the edge of the forest. At locations with L. terrestris, litter was incorporated into the soil, and where O. tyrtaeum was present, organic layers were mixed with mineral soil layers. Organic layers disappeared almost entirely when both species occurred together. Carbon and nitrogen concentrations were reduced in organic layers in the presence of L. terrestris and O. tyrtaeum. Microbial biomass and basal respiration were reduced when L. terrestris and O. tyrtaeum were present, presumably due to resource competition and habitat destruction. Microarthropod densities and the number of microarthropod species were strongly reduced in the presence of O. tyrtaeum (−75% and −22%, respectively), probably through mechanical disturbances, increasing compactness of the soil and resource competition. The coverage of some plant species was correlated with earthworm abundance, but the coverage of others was not. Despite harsh climatic conditions, the invasion of boreal forest ecosystems by mineral soil dwelling earthworm species is proceeding and strongly impacts soil structure, soil chemistry, microorganisms, soil microarthropods and vegetation. [Copyright &y& Elsevier]

The interactions between earthworms and soil microbial communities have attracted a lot of scientific attention, but if and how important soil invertebrates may modulate climate effects on soil microorganisms in different land-use types is less clear. Here we collected earthworm casts as an indicator of earthworm effects on soil microbial communities and bulk soil from the Global Change Experiment Facility (GCEF) situated at Bad Lauchstädt, Germany. We tested the interactive effect of two climate treatments (ambient and future), land-use type (conventional farming, organic farming, and extensively used meadows), and sample type (cast and bulk soil) on the soil microbial community. The biomass and community composition of the bacterial and fungal communities were assessed by fatty acid analysis. Climate, alone and in interaction with land-use type, had no significant effects on the biomass and composition of soil microorganisms in bulk soil and casts produced by earthworms. The biomasses of almost all microbial markers (except for 18:2ω6t and 18:1ω9t) were significantly higher in the cast than in bulk soil in all land-use types (+185%), with the most substantial impact on all other fungal makers in soil. However, the land-use effect was significantly modulated by soil type, i.e., earthworms. Mainly extensively managed meadows had more than double (+237%) the microbial biomass in casts than in bulk soil, whereas in the other land-use types, the positive effect of cast soil was much smaller (conventional farming: +166%; organic farming: +123%). The microbial community composition showed no difference between the two cropping systems (conventional farming and organic farming). Here, bulk soil was more associated with fungal biomarkers, whereas casts were more dominated by bacterial markers; however, this effect was rather weak. Extensively managed meadows showed strong differences to both cropping systems, mainly due to higher microbial biomass and a more heterogeneous association of either more fungal or bacterial communities. We suggest that the main factors that led to differences between land-use types were a more diverse plant community in meadows and the less intensive management regime with undisturbed soils. The conditions in extensively managed meadows allowed earthworms to strongly amplify those positive effects in casts, suggesting that earthworms can act as important biotic filters modulating land-use effects on soil microbes across climate treatments. • We explored climate and land-use effects on soil microbial communities in bulk soil and earthworm casts. • Extensive land use enhanced soil microbial biomass, while climate had no significant effects. • Microbial biomass was significantly higher in earthworm casts than in bulk soil. • Extensively managed meadows had more than double the microbial biomass in casts than in bulk soil • Earthworms intensified beneficial effects of extensive land use on soil microbial communities. [ABSTRACT FROM AUTHOR]