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International audience; AimSpecies shift their ranges as a consequence of climate change, hence modifying the structure of local assemblages. This may have important consequences for ecosystem functioning in the case of ecosystem engineers such as earthworms, especially when community restructuring leads to an alteration of their functional diversity. Here, we aimed to model the potential modification of the functional diversity of French earthworm assemblages in a context of climate change.LocationMetropolitan France.MethodsWe fitted boosted regression trees to earthworm data collected using a standardized protocol across France in the 1960s. We used model projections constrained by a macroecological model of species richness to predict the composition of earthworm assemblages in the present and in two scenarios of climate change and two future time periods. We coupled these results with a large set of species traits to calculate predicted changes in functional diversity, which we summarized by ecoregion.ResultsModels predicted a clear decline in functional richness between the period of sampling and nowadays which are expected to continue in the future, with substantial differences depending on ecoregions and on whether species will be able to disperse or not. However, predicted changes in functional evenness and divergence are much weaker, suggesting that climate change will not affect all facets of functional diversity in the same way.Main ConclusionsOur results mostly pointed to a potential reduction of the functional richness of earthworm communities in the future, but this predicted loss of diversity could be weaker if species are able to colonize new suitable sites or to persist in microclimate refugia. There are concerns, though, that these changes lead to an alteration of soil processes and of the ecosystem services they provide.

The present study investigates the bioaccumulation of the insensitive munition compounds 2,4-dinitroanisole (DNAN) and 3-nitro-1,2,4-triazol-5-one (NTO), developed for future weapons systems to replace current munitions containing sensitive explosives. The earthworm Eisenia andrei was exposed to sublethal concentrations of DNAN or NTO amended in Sassafras sandy loam. Chemical analysis indicated that 2- and 4-amino-nitroanisole (2-ANAN and 4-ANAN, respectively) were formed in DNAN-amended soils. The SumDNAN (sum of DNAN, 2-ANAN, and 4-ANAN concentrations) in soil decreased by 40% during the 14-d exposure period. The SumDNAN in the earthworm body residue increased until day 3 and decreased thereafter. Between days 3 and 14, there was a 73% decrease in tissue uptake that was greater than the 23% decrease in the soil concentration, suggesting that the bioavailable fraction may have decreased over time. By day 14, the DNAN concentration accounted for only 45% of the SumDNAN soil concentration, indicating substantial DNAN transformation in the presence of earthworms. The highest bioaccumulation factor (BAF; the tissue-to-soil concentration ratio) was 6.2 ± 1.0 kg/kg (dry wt) on day 3 and decreased to 3.8 ± 0.8 kg/kg by day 14. Kinetic studies indicated a BAF of 2.3 kg/kg, based on the earthworm DNAN uptake rate of 2.0 ± 0.24 kg/kg/d, compared with the SumDNAN elimination rate of 0.87 d-1 (half-life = 0.79 d). The compound DNAN has a similar potential to bioaccumulate from soil compared with trinitrotoluene. The NTO concentration in amended soil decreased by 57% from the initial concentration (837 mg NTO/kg dry soil) during 14 d, likely due to the formation of unknown transformation products. The bioaccumulation of NTO was negligible (BAF ≤ 0.018 kg/kg dry wt). Environ Toxicol Chem 2021;40:1713-1725. © 2021 SETAC. This article has been contributed to by US Government employees and their work is in the public domain in the USA.

The present study is aimed to evaluate the effects of different levels of earthworm (EW) powder and vermihumus (VH) on the growth performance, humoral immunity response, plasma constituents and reproductive performance of Japanese breeder quails. A total of 336 birds were assigned to seven treatments in a completely randomized design from day 30 to 90 of age. The dietary treatments were carried out, one without any addition of EW and VH, the second had a standard rate of VH at 0.8%, while a range of EW additions was supplied from 0%, 0.5%, 1%, 1.5%, 2% and 2.5%. The effects of treatments on body weight and mortality were not significant (p > .05). The highest feed intake was observed in birds fed the diets supplemented with 1% and 1.5% EW (p

Soil structure is closely linked to biological activities. However, identifying, describing and quantifying soil aggregates remain challenging. X‐ray micro computed tomography (X‐ray μCT) provides a detailed view of the physical structure at a spatial resolution of a few microns. It could be a useful tool to discriminate soil aggregates, their origin and their formation processes for a better comprehension of soil structure properties and genesis. Our study aims to (a) determine different X‐ray μCT‐based aggregate parameters for differentiating earthworm casts belowground (earthworm aggregates) from aggregates that are not formed by earthworms (non‐earthworm aggregates), and (b) to evaluate if these parameters can also serve as specific “tomographic signatures” for the studied earthworm species. For this purpose, we set up a microcosm experiment under controlled conditions during 8 weeks, including three species of earthworms tested separately: the epigeic Lumbricus rubellus, the anecic Lumbricus terrestris and the endogeic Allolobophora chlorotica. Our results show that X‐ray μCT analysis helps distinguish earthworm aggregates from non‐earthworm ones using (a) the relative volume of the components within aggregates and (b) the volumetric mass of aggregates and their global volume. In particular, the volume ratio of mineral grains within the aggregates is significantly different according to earthworm species. So, X‐ray μCT is a powerful and promising tool for studying the composition of earthworm casts and their formation. However, future research is needed to take into account the shapes and spatial distribution of the aggregates' components, in particular the different states of organic matter decomposition. HIGHLIGHTS: Can earthworm belowground casts be differentiated from other soil aggregates using X‐ray μCT? The use of X‐ray μCT for characterizing soil aggregates at a spatial resolution of a few microns. A combination of X‐ray μCT variables discriminates earthworm casts from non‐earthworm aggregates. X‐ray μCT, used alone, is relevant for defining species‐specific signatures of earthworm casts.

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 seasons Climate had species‐specific effects on active earthworms, but few interactions with land‐use type Intensive grassland management decreased, but sheep grazing favoured, active earthworm populations Strong seasonal variations in earthworm activity periods will be modulated by climate change

Microplastic pollution is a topic of increasing concern for the world's oceans, freshwaters and, most recently, soils. Microplastics have been found in soils across the globe. Like other anthropogenic pollutants, they can negatively affect a range of soil organisms through several mechanisms, though often dependent on particle size, shape and polymer type. However, microplastics are unique among pollutants due to the diversity of ways in which soil organisms may themselves be able to affect their occurrence and distribution and mediate their effects on the rest of the soil food web. In this review, we argue for a more explicitly ecological framing of this novel issue for the soil environment and discuss their potential interactions with soil communities, including microplastic formation via microbial and faunal fragmentation of large plastic debris and organisms such as earthworms placing microplastic particles in unique pedological contexts they could not otherwise reach. Ecological interactions may be crucial for dictating microplastics’ ultimate fate and effect on terrestrial ecosystems. A free Plain Language Summary can be found within the Supporting Information of this article.

Four common earthworm species, the anecic Lumbricus terrestris, the endogeic Octolasion tyrteum as well as the epigeic Eisenia fetida and Dendrobaena veneta, were examined for the presence of the microbial gut symbiont Plagiotoma lumbrici. The evolutionary origin of this endobiotic microbe was reconstructed, using the 18S rRNA gene, the ITS1-5.8S-ITS2 region, and the first two domains of the 28S rRNA gene. Plagiotoma lumbrici was exclusively detected in the anecic Lumbricus terrestris. Multigene analyses and the ITS2 secondary structure robustly determined the phylogenetic home of Plagiotoma lumbrici populations within the oxytrichid Dorsomarginalia (Spirotrichea: Hypotrichia) as a sister taxon of the free-living Hemiurosomoida longa. This indicates that earthworms obtained their gut endosymbiont by ingesting soil/leaf litter containing oxytrichine ciliates that became adapted to the intestinal tract of earthworms. Interestingly, according to the literature data, Plagiotoma lumbrici was detected in multiple anecic and some epigeic but never in endogeic earthworms. These observations suggest that Plagiotoma lumbrici might be adapted to certain gut conditions and the lifestyle of anecic Lumbricidae, such as Lumbricus, Aporrectodea, and Scherotheca, as well as of some co-occurring epigeic Lumbricus species.

The effects of five conservation tillage drills with crop residue levels covering between 17% and 79% of the soil, and tillage depths ranging from 25 mm to 200 mm, were examined over three years. The tillage systems ranged from a relatively disruptive Farm System to a Low Disruption system, with three intermediate treatments labelled Sumo DTS, Claydon, and Mzuri. The study involved field sites on a clay or clay loam soil, where winter wheat and oilseed rape were grown in rotation. In the clay field, the Mzuri and Low Disruption treatments, which produced the highest residue coverage, showed the greatest increase in surface total soil organic carbon (1.1 and 0.48 Mg C ha−1 respectively) between year 1 and 3. The least disruptive tillage system also resulted in the highest density of earthworms (181‐228 m−2), and the most disruptive system produced the lowest densities (75‐98 m−2). In the third year, the least disruptive system also showed a higher proportion of water stable aggregates (29.8%) than the other treatments (22.7‐25.3%). Linear regressions showed positive relationships of both soil organic carbon and earthworm density with surface residue cover, and of the proportion of water stable aggregates with soil organic carbon.

Earthworm activity is beneficial in most natural and agricultural systems, but excessive earthworm casting is a problem on sports fields worldwide. Expulsion of soil-rich earthworm fecal matter, or casts, as mounds of soil on the turf canopy can muddy the surface, reduce photosynthesis, and lead to thinning, weed invasion and surface softening. Casts affect ball roll on sports fields, cause fouling of maintenance equipment, and dull mower blades. Build-up of cast material on reel mower units can affect height and quality of cut. Casting activity is dependent on environmental conditions such as soil texture, temperature, moisture, pH, and organic matter. Response to environmental conditions varies by species. Management options are limited, because no pesticides are specifically labeled for earthworm control at this time, and cultural control methods such as soil modification, turfgrass clipping removal, and sand topdressing have limited and inconsistent efficacy. Products containing plant-derived saponins and irritants show promise for earthworm management. Pest management practices to mitigate excessive earthworm casting will likely need to be species-specific, but limited knowledge of earthworm identification by end-users further inhibits the efficacy of control measures. © 2019 Society of Chemical Industry.

The alimentary canal of the earthworm is representative of primitive gut ecosystems, and gut fermenters capable of degrading ingested biomass-derived polysaccharides might contribute to the environmental impact and survival of this terrestrial invertebrate. Thus, this study evaluated the postulation that gut microbiota of the model earthworm Lumbricus terrestris ferment diverse biomass-derived polysaccharides. Structural polysaccharides (e.g. cellulose, chitin) had marginal impact on fermentation in anoxic gut content treatments. In contrast, nonstructural polysaccharides (e.g. starch, glycogen) greatly stimulated (a) the formation of diverse fermentation products (e.g. H2 , ethanol, fatty acids) and (b) the facultatively fermentative families Aeromonadaceae and Enterobacteriaceae. Despite these contrasting results with different polysaccharides, most saccharides derived from these biopolymers (e.g. glucose, N-acetylglucosamine) greatly stimulated fermentation, yielding 16S rRNA gene-based signatures of Aeromonadaceae-, Enterobacteriaceae- and Fusobacteriaceae-affiliated phylotypes. Roots and litter are dietary substrates of the earthworm, and as proof-of-principle, gut-associated fermenters responded rapidly to root- and litter-derived nutrients including saccharides. These findings suggest that (a) hydrolysis of certain ingested structural polysaccharides may be a limiting factor in the ability of gut fermenters to utilize them and (b) nonstructural polysaccharides of disrupted biomass are subject to rapid fermentation by gut microbes and yield fatty acids that can be utilized by the earthworm.

Improper reuse of manure could not only lead to low nitrogen-use efficiency but also result in environmental pollution. Therefore, we studied the effect of manure amendment with or without organisms (earthworm and bacteria) on soil nitrogen cycling, aiming to develop a more scientific way to reuse manure. In this study, six treatments, including control (CK), manure (M), manure + slurry (MS), manure + slurry + earthworm (MSE), manure + slurry + bacteria (MSB), and manure + slurry + earthworm + bacteria (MSEB) were divided into two groups to study the effect of manure and the effect of organism on the soil nitrogen pools and nitrogen cycling, respectively. The results showed that manure amendment coupled with organisms increased total nitrogen from 0.538 ± 0.06 to 1.380 ± 0.22 g kg ¹, and available nitrogen from 67.28 ± 5.22 to 113.11 ± 12.21 mg kg⁻¹, and improved soil organic nitrogen pool by increasing acid-hydrolyzable nitrogen (AHN) from 404 ± 39 to 974 ± 94 mg kg⁻¹ and non-acid-hydrolyzable nitrogen (NAHN) from 133 ± 26 to 406 ± 155 mg kg⁻¹. Based on the results of soil inorganic nitrogen and soil enzyme activities, we further found that earthworms and bacteria accelerated soil nitrogen cycling by stimulating soil mineralization (ammonification and nitrification) and nitrifier-denitrification, providing more available nitrogen to plants. Thus, we conclude that manure coupled with organisms would be a more efficient way to reuse manure via increasing soil nitrogen storage and availability simultaneously.

Water-treatment residuals (WTRs), by-products of drinking water clarification, are increasingly recycled to land to promote circular economy and reduce disposal costs, yet there is a lack of published literature on their effects on soil ecology. In the present study, the effects of WTRs on earthworm growth, soil respiration, and soil porewater chemistry were investigated throughout a 7-wk outdoor mesocosm trial. We derived WTRs from both aluminum and iron coagulants and applied them to a loam soil at 0 to 20% (w/w). In addition, soil from a field that had received long-term WTR applications and that of an adjacent nontreated reference field were included in the study. Earthworm mass increase was significantly higher in all but one laboratory-treated soil when compared to the control. Furthermore, a linear regression model was used to predict increases in weekly soil respiration based on the application rates of both Al and Fe WTRs. In addition, a significant increase in soil respiration was observed from the treated farm soils during the first 4 wk of the trial. Measured sodium, magnesium, potassium, and iron porewater concentrations were higher in the treated farm soils than the reference site soil in a majority of samples, although these differences may be related to land management. Laboratory-treated soils had elevated porewater arsenic concentrations (e.g., ~17 µg L-1 in controls vs ~62 µg L-1 in the 20% w/w Al WTR treatment in week 1), whereas porewater nickel concentrations were, respectively, elevated and lowered in Al WTR- and Fe WTR-amended samples. Overall, observed disturbances to soil ecology were determined to be minimal. Environ Toxicol Chem 2021;40:1964-1972. © 2021 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

Biological invasions by cryptic ecosystem engineers can alter the ecological and socio‐economic values of ecosystems in ways that may take decades to detect. The invasion of North American glacial refuges by non‐native earthworms offers a prominent but understudied example of a cryptic invasion. Non‐native earthworms are known to alter carbon sequestration, disrupt mycorrhizal networks and homogenize plant communities, but natural resource managers still lack robust protocols to identify and safeguard high conservation value communities (HCVCs) from such invasions. In the absence of such protocols, or reliable methods to eradicate non‐native earthworm populations once established, there is an urgent need for methods to identify HCVCs at risk of or potentially shielded from earthworm invasion by the existence of abiotic barriers to their dispersal or persistence in such sites. We developed species distribution models (SDMs) using in situ field surveys and remotely sensed data to (1) identify factors limiting the occurrence of non‐native Lumbricus earthworms in imperilled coastal Douglas‐fir (Pseudotsuga menziesii) forest and Garry oak (Quercus garryana) and maritime meadow ecosystems endemic to glacial refugia of the Pacific Northwest of North America and to (2) evaluate their influence on plant species diversity and identify abiotic factors capable of preventing their dispersal or persistence at a site. As predicted, shallow, drier soils and steeper terrain limited earthworm occurrence and abundance in our highly heterogeneous study area, and earthworm presence appeared to reduce plant species richness. Our results indicated that HCVCs endemic to shallow‐soil (

Mounded, tallgrass prairie ecosystems are a unique, endangered ecosystem due to severe disturbance for agricultural production and/or urban expansion. Limited research exists on earthworm (Lumbricus terrestris) dynamics in mounded, tallgrass prairies, particularly in the Ozark Highlands region of northwestern Arkansas. The objectives of this study were to evaluate the effects of soil moisture regime (SMR), landscape position (LP), and sample date on earthworm densities in the top 20 cm of soil in two mounded, native prairies in the Ozark Highlands region of northwestern Arkansas. Juvenile and total earthworm densities were five times greater (P

To mitigate the loss of tallgrass prairies through fragmentation and conversion to other land uses, prairie restoration efforts have been undertaken to reclaim areas that have been previously used for agricultural production. Earthworms are a key component of the soil ecology of natural, undisturbed ecosystems. However, limited research exists on earthworm populations in prairie restorations and how earthworm densities may change over time. The objectives of this study were (a) to evaluate earthworm densities in the upper 20 cm of soil over time (i.e., May 2007 and May 2019) in a native prairie and chronosequence of three prairie restorations (i.e., established in 1979, 2000, and 2002), and (b) to identify relationships between earthworm (i.e., juvenile, adult, and total) density and soil physical and chemical properties. Juvenile (P = .03) earthworm density decreased 25 and 35% and total (P = .03) earthworm density decreased by 33 and 42% in the 2000 and 2002 restorations, respectively, from May 2007 to May 2019. Adult earthworm density did not change over time (P > .05) in any prairie ecosystem. Combined across prairie sites, adult earthworm density was positively correlated (P

In many terrestrial ecosystems, earthworms operate at the interface between plants and soil. As ecosystem engineers, they affect key ecosystem functions such as decomposition, nutrient cycling and bioturbation. Their incidence and abundance depends on several soil properties, yet simultaneously they also impact soil properties themselves. The existence of a positive feedback loop in which earthworm activity maintains their own niche—by promoting turnover rate in the forest floor, thereby increasing topsoil pH and creating suitable living conditions for themselves—has been suggested before, yet lacks supporting evidence. Here we assessed how tree species litter traits relate to such below‐ground interactions in forests across Europe. Specifically, we hypothesized a below‐ground feedback loop between burrowing earthworm biomass, humus form and pH, affected by litter quality. We tested this hypothesis by means of structural equation modelling. Our results demonstrate that litter nutrient concentrations affect both burrowing and litter dwelling earthworm biomass, which in turn directly impacts humus form and indirectly soil pH. At a continental scale, that is, including all edaphic conditions, soil pH did not feed into earthworm biomass nor could we link leaf structural recalcitrance (e.g. lignin) or functional diversity to below‐ground interactions. However, in forests where moisture is not limiting, soil acidity proved an important factor determining the context of below‐ground interactions. Therefore, we were able to confirm the hypothesized feedback loop for forest ecosystems with soil pH ≤ 5. In calcareous and/or periodically dry forests, other factors than soil chemistry and litter quality became determinant for earthworm biomass. The activity of burrowing earthworms is pivotal in below‐ground ecosystem functioning of mesic forest soils, impacting litter accumulation and forest floor conditions above‐ground, the pH and nutrient status below‐ground and ultimately their own living conditions. This highlights earthworm bioturbation as a key mechanism for understanding plant–soil interactions in forests. A free Plain Language Summary can be found within the Supporting Information of this article. ispartof: FUNCTIONAL ECOLOGY vol:34 issue:12 pages:2598-2610 status: published

Aim\ud Soils and their biological communities face increasing pressure from multiple global drivers, including land management and climate change. In soils, earthworms play key roles in ecosystem functioning, but the environmental controls on their global communities are not fully understood. Here, an earthworm dataset was compiled to investigate the effects of environmental variables and land management on global earthworm communities.\ud \ud Location\ud 40° S–65° N.\ud \ud Time period\ud 1962 to 2016.\ud \ud Major taxa studied\ud Earthworms.\ud \ud Methods\ud A dataset of 899 earthworm community observations, together with environmental variables, was compiled across 169 globally distributed sites. Sites included natural forest and grassland or managed arable, pasture and plantation ecosystems. Total, anecic, endogeic and epigeic abundances and total species richness were compared in natural and managed ecosystems to quantify the effects of land management across climates. A hierarchical model was used to test the importance of environmental controls in predicting the relationship between total earthworm species richness and abundance at a global scale.\ud \ud Results\ud Land management prompted little change in total earthworm abundance at the global scale, but reduced species richness and shifted community composition. Endogeic earthworms were more abundant in managed ecosystems, while anecic and epigeic earthworms showed variable responses across ecosystem types. Global relationships between total earthworm species richness and abundance were explained by climate, soil pH and land management.\ud \ud Main conclusions\ud Land management modulates the effects of environmental controls on global earthworm communities, through direct disturbance and indirect changes in edaphic conditions.

Municipal biosolids are increasingly used as a low-cost fertilizer in agricultural soil. Biosolids are contaminated by low concentrations (nanograms per gram dry wt range) of a large variety of organic contaminants, such as triclosan. The effect of exposure to low concentrations of organic contaminants on soil biota remains largely undocumented. We evaluated the sublethal effects of triclosan on the earthworm Eisenia andrei using an artificial soil amended with a nominal concentration of triclosan of 50 ng g-1 dry weight soil. Using a 56-d reproduction test, we monitored the effect of triclosan exposure on adult earthworm survival, growth, and reproduction. The bioaccumulation of triclosan in earthworm tissue (adults and juveniles) and degradation of triclosan were monitored. The genotoxicity of triclosan was evaluated using a comet assay (DNA damage) on adult earthworm coelomocytes. Exposure to a low concentration of triclosan had no significant effects on adult earthworm survival and DNA damage but significantly stimulated growth (p

Nanopesticides are novel plant protection products offering numerous benefits. Because nanoparticles behave differently from dissolved chemicals, the environmental risks of these materials could differ from conventional pesticides. We used soil-earthworm systems to compare the fate and uptake of analytical-grade bifenthrin to that of bifenthrin in traditional and nanoencapsulated formulations. Apparent sorption coefficients for bifenthrin were up to 3.8 times lower in the nano treatments than in the non-nano treatments, whereas dissipation half-lives of the nano treatments were up to 2 times longer. Earthworms in the nano treatments accumulated approximately 50% more bifenthrin than those in the non-nano treatments. In the non-nano treatments, most of the accumulated material was found in the earthworm tissue, whereas in the nano treatments, the majority resided in the gut. Evaluation of toxicokinetic modeling approaches showed that models incorporating the release rate of bifenthrin from the nanocapsule and distribution within the earthworm provided the best estimations of uptake from the nano-formulations. Overall, our findings indicate that the risks of nanopesticides may be different from those of conventional formulations. The modeling presented provides a starting point for assessing risks of these materials but needs to be further developed to better consider the behavior of the nanoencapsulated pesticide within the gut system. Environ Toxicol Chem 2018;37:1420-1429. © 2018 SETAC.

Underground community assemblies have not been studied well compared with aboveground communities, despite their importance for our understanding of whole ecosystems. To investigate underground community assembly over evolutionary timescales, we examined terrestrial earthworm communities (Oligochaeta: Haplotaxida) in conserved mountainous primary forests in Japan as a model system. We collected 553 earthworms mostly from two dominant families, the Megascolecidae and the Lumbricidae, from 12 sites. We constructed a molecular taxonomic unit tree based on the analysis of three genes to examine the effects of a biogeographic factor (dispersal ability) and an evolutionary factor (habitat adaptation) on the earthworm community assembly process. The phylogenetic distance of the earthworm communities among sites was positively correlated with geographic distance when intraspecific variation was included, indicating that the divergence within species was affected by biogeographic factors. The community assembly process in the Megascolecidae has also been affected by environmental conditions in relation to an evolutionary relationship between habitat environment and intestinal cecum type, a trait closely related to habitat depth and diet, whereas that in the Lumbricidae has not been affected as such. Intestinal cecum type showed a pattern of niche conservatism in the Megascolecidae lineage. Our results suggest that investigating the evolution of a key trait related to life history can lead to the clear description of community assembly process over a long timescale and that the community assembly process can differ greatly among related lineages even though they live sympatrically.

Management of both municipal and industrial organic wastes remains a major threat to biota and the environment due to the presence of pathogens in abundance. Vermicomposting employing earthworms is increasingly gaining attention as a sustainable and ecofriendly technique to transform and sanitize a variety of organic wastes into nutrient-rich biofertilizer. Although considerable research has been undertaken to show that vermicomposting can significantly reduce pathogenic contents, there is little effort to summarize the various mechanisms responsible for it. With the aim to assess the fate of pathogens during vermicomposting of various organic wastes, this article provides a comprehensive summary on the occurrence of pathogens in a variety of wastes vis-à-vis pathogens standards, the efficacy of the process for pathogen reduction, and current knowledge of the plausible mechanisms involved. It is evident from the present study that earthworms and endosymbiotic microbes during vermicomposting tend to eliminate pathogens by enhancing enzymatic activities in both gut- and cast-associated processes. Pathogen reduction during vermicomposting can be plausibly attributed to direct actions like microbial inhibition due to intestinal enzymatic action, and secretion of coelomic fluids with antibacterial properties, as well as indirect actions like stimulation of endemic microbes leading to competition and antagonism, and aeration by burrowing activity. Further, the pathogen reduction during vermicomposting is largely selective, and earthworms exert a differential effect according to the earthworm species and whether the pathogen considered is Gram-positive or -negative, owing to its cell wall composition. However, further research is necessary to understand the exact mechanisms involved for pathogen reduction during vermistabilization of municipal and industrial organic wastes.

Lead concentrations in soil organisms are usually well below those in the associated soil and tend to decrease with each higher trophic level in a food chain. Earthworms of the species Eisenoides lonnbergi provide an exception to this observation, accumulating very high concentrations of lead from acidic soils. Earthworms belonging to this species were collected from strongly to extremely acidic soils at 16 sites on a wildlife refuge in Maryland, USA. A lead concentration as high as 766 mg/kg, dry weight, was detected in depurated E. lonnbergi collected from soil containing only 17 mg/kg of lead. Concentration factors (ratio of lead concentration in earthworms to lead concentration in soil, dry weights) were highly variable at the sites, from 1.0 to 83. As suggested previously by M. Ireland and J. Morgan, lead absorption by earthworms is enhanced in low calcium soils. The anomalously high concentrations of lead found in E. lonnbergi are more closely correlated with the uptake of calcium from acidic soils than with bioaccessibility of soil lead. This article is protected by copyright. All rights reserved

Atrazine toxicity to earthworms is still not fully understood, particularly at sub-lethal concentrations. Because of the ubiquity of atrazine in the environment, it is imperative to understand the impacts of atrazine presence on soil-dwelling organisms. To examine this in detail, we used 1H nuclear magnetic resonance (NMR)-based metabolomics to elucidate earthworm (Eisenia fetida) responses after a 48 h of atrazine exposure in contact tests. Earthworms were exposed to four sub-lethal concentrations of 362.4 ng/cm2, 181.2 ng/cm2, 90.6 ng/cm2 and 45.3 ng/cm2, which correspond to 1/8th, 1/16th, 1/32nd and 1/64th of the LC50 value respectively. After exposure, polar metabolites were isolated from earthworm tissues and analyzed using 1H NMR spectroscopy. Sub-lethal atrazine exposure induced a non-monotonic response with respect to exposure concentration and caused an overall suppression in earthworm metabolism. Maltose, fumarate, malate, threonine/lactate, adenosine-5'-triphosphate (ATP), betaine, scyllo-inositol, glutamate, arginine and glutamine were the metabolites identified as most sensitive to atrazine exposure. These observed fluctuations in the metabolic profile suggest that atrazine reduced ATP synthesis and negatively impacted the health of earthworms with acute sub-lethal exposure. Our study also demonstrates the utility of NMR-based metabolomics for the basic assessment of sub-lethal toxicity which can then be used for more targeted approaches with other molecular techniques. This article is protected by copyright. All rights reserved

Summary As ecosystem engineers, earthworms greatly affect plant communities. They create persistent soil structures enriched by nutrients that improve the conditions for plant growth and modify competition between plant species. We therefore hypothesized that earthworm activity would be more important in early stages of the primary succession, when the soil is not modified by earthworms, than in the late stages of the succession, when the soil is already improved by earthworms. On the other hand, earthworms also affect plants via many other effects such as seed predation or excreting hormone-like compounds, which could make earthworm presence important in late successional soil. To explore earthworm effects on plant community succession, we performed a laboratory microcosm experiment without and with earthworms (Lumbricus rubellus and Aporrectodea caliginosa), with early successional plants (Poa compressa, Medicago lupulina, and Daucus carota) and late successional plants (Arrhenatherum elatius, Lotus corniculatus, and Plantago lanceolata), and with soil previously unaffected by earthworms (young soil) and soil substantially affected by earthworms (developed soil). These soils were taken from the early and late successional post-mining sites of the Sokolov coal mining district (northwest Czech Republic). When both early and late successional plants were grown separately, earthworms increased plant biomass proportionally more in the young soil than in the developed soil, indicating that earthworm activity is more important in undeveloped than in developed soil. When early and late successional plants were competing each other, the biomass of the early successional plants was reduced. In the young soil the reduction was independent of earthworm presence. In the developed soil the reduction was promoted by the earthworms. Late successional plants profited from the reduction of early successional plants and increased their biomass. This increase was promoted by the earthworm presence. Our results indicate that the direct effects of earthworm presence on plants decrease during succession because of the cumulative effects of earthworm activity on soil conditions. Such ecosystem engineering effects favor late successional competitors and therefore promote the replacement of species during succession. This article is protected by copyright. All rights reserved.

Uptake of polychlorinated biphenyls (PCBs) by fish is controlled by the bioavailability of ingested PCBs in the gut and the freely dissolved concentration in the water moving across the gills. The prediction of bioaccumulation in fish relies on models that account for these exposure routes; however, these models typically do not account for incidental ingestion of sediment by fish, which is not well studied. The literature values for the PCB assimilation efficiency in the gut have been reported for compounds in food matrices and not associated with sediment particles. It is also unclear how mitigation strategies that alter PCB bioavailability in sediments affect predictions made by the bioaccumulation models when sediment ingestion is involved. To test the bioavailability of PCBs from treated and untreated sediments, dietary assimilation efficiencies were measured for 16 PCB congeners in mummichogs (Fundulus heteroclitus) that were fed 4 experimental diets. Diets consisted of PCB-spiked earthworms, spiked untreated sediment mixed with earthworms, spiked activated carbon-treated sediment mixed with earthworms, and spiked activated carbon mixed with earthworms. Assimilation efficiencies were determined by calculating the ratio of PCB mass in the fish tissue to the PCB mass in the food after a pulse feeding experiment. Assimilation efficiencies of PCBs associated with earthworm diet were similar to the values reported in the literature. Fish that were fed the PCB-spiked untreated sediment and activated carbon particles exhibited the highest and lowest assimilation efficiencies, respectively, over a wide KOW range. Assimilation efficiencies of sediment-bound PCBs were significantly reduced (31-93% reduction for different congeners) after amendment with activated carbon. The present study indicates that assimilation of PCBs can be reduced by sorption to black carbon. Environ Toxicol Chem 2017;36:3480-3488. © 2017 SETAC.

Earthworms and plants greatly affect belowground properties; however, their combined effects are more attractive based on the ecosystem scale in the field condition. To address this point, we manipulated earthworms (exotic endogeic species Pontoscolex corethrurus) and plants (living plants [native tree species Evodia lepta] and artificial plants) to investigate their combined effects on soil microorganisms, soil nutrients, and soil respiration in a subtropical forest. The manipulation of artificial plants aimed to simulate the physical effects of plants (e.g., shading and interception of water) such that the biological effects of plants could be evaluated separately. We found that relative to the controls, living plants but not artificial plants significantly increased the ratio of fungal to bacterial phospholipid fatty acids (PLFAs) and fungal PLFAs. Furthermore, earthworms plus living plants significantly increased the soil respiration and decreased the soil NH4+-N, which indicates that the earthworm effects on the associated carbon, and nitrogen processes were greatly affected by living plants. The permutational multivariate analysis of variance results also indicated that living plants but not earthworms or artificial plants significantly changed the soil microbial community. Our results suggest that the effects of plants on soil microbes and associated soil properties in this study were largely explained by their biological rather than their physical effects.

The use of engineered silver nanoparticles (AgNPs) is widespread, with expected release to the terrestrial environment through the application of biosolids onto agricultural lands. The toxicity of AgNPs and silver nitrate (AgNO3 ; as ionic Ag+ ) to plant (Elymus lanceolatus and Trifolium pratense) and soil invertebrate (Eisenia andrei and Folsomia candida) species was assessed using Ag-amended biosolids applied to a natural sandy loam soil. Bioavailable Ag+ in soil samples was estimated using an ion-exchange technique applied to KNO3 soil extracts, whereas exposure to dispersible AgNPs was verified by single-particle inductively coupled plasma-mass spectrometry and transmission electron microscopy-energy dispersive X-ray spectroscopy analysis. Greater toxicity to plant growth and earthworm reproduction was observed in AgNP exposures relative to those of AgNO3 , whereas no difference in toxicity was observed for F. candida reproduction. Transformation products in the AgNP-biosolids exposures resulted in larger pools of extractable Ag+ than those from AgNO3 -biosolids exposures, at similar total Ag soil concentrations. The results of the present study reveal intrinsic differences in the behavior and bioavailability of the 2 different forms of Ag within the biosolids-soils pathway. The present study demonstrates how analytical methods that target biologically relevant fractions can be used to advance the understanding of AgNP behavior and toxicity in terrestrial environments. Environ Toxicol Chem 2017;36:2756-2765. © 2017 Crown in the Right of Canada. Published Wiley Periodicals Inc., on behalf of SETAC.

Earthworms enhance soil functioning and are, therefore, key species in the soil. Their presence is generally a positive sign for a terrestrial ecosystem, as these species serve as important biomarkers in soil quality evaluations. Here, we describe a novel bioassay, named “dispersal assay,” that is a simple and rapid technique for field-based soil quality evaluations. This bioassay was designed based on the premise that earthworms prefer optimal soils if given the choice. Thus, assay tubes containing a reference soil were inserted in target sites, and earthworms were placed into these tubes. According to their soil preference, the earthworms dispersed into the surrounding soil, remained in the initial soil within the tubes, avoided both by crawling up the tube, or died. Furthermore, sensitivity responses to metal concentrations, electrical conductivity, and soil pH were observed in field tests. Although the dispersal assay did not completely match traditional toxicity endpoints such as earthworm survival, we found that the dispersal assay can serve as an in situ screening test for assessing soil quality. Overall, our dispersal assay was relatively rapid (within 24 h), had low levels of variation, and showed high correlations between earthworm behavior and soil physicochemical properties. This article is protected by copyright. All rights reserved

The adoption of less intensive soil cultivation practices is expected to increase earthworm populations and their contributions to ecosystem functioning. However, conflicting results have been reported on the effects of tillage intensity on earthworm populations, attributed in narrative reviews to site-dependent differences in soil properties, climatic conditions and agronomic operations (e.g. fertilization, residue management and chemical crop protection). We present a quantitative review based on a global meta-analysis, using paired observations from 165 publications performed over 65 years (1950–2016) across 40 countries on five continents, to elucidate this long-standing unresolved issue. Results showed that disturbing the soil less (e.g. no-tillage and conservation agriculture [CA]) significantly increased earthworm abundance (mean increase of 137% and 127%, respectively) and biomass (196% and 101%, respectively) compared to when the soil is inverted by conventional ploughing. Earthworm population responses were more pronounced when the soil had been under reduced tillage (RT) for a long time (>10 years), in warm temperate zones with fine-textured soils, and in soils with higher clay contents (>35%) and low pH (

North America is home to both native and invasive earthworms acting as ecosystem engineers as they build burrows that can serve as habitat for other species or otherwise alter soil structure, affecting nutrient cycling and other ecosystem processes. Here I determine where and what earthworm species commonly occur in my study area, and compare effects of native and invasive earthworms on the common woodland salamander, Plethodon cinereus, in field surveys and laboratory experiments. The native earthworm Eisenoides carolinensis was the most common earthworm, followed by two invasive species Dendrobaena octaedra and Octolasion tyrtaeum. The presence of O. tyrtaeum was associated with a narrower O-horizon (i.e., organic layer in the soil). Using structural equation modeling to explore direct and indirect pathways of these three most common earthworm species on salamanders, I found O. tyrtaeum occurrence was negatively correlated with nighttime salamander counts, a proxy for total salamander numbers, mediated by negative effects on O-horizon depth and microinvertebrate numbers. In the laboratory, O. tyrtaeum and D. octaedra consumed more leaf litter per gram of earthworm per day than the native E. carolinensis. However, salamanders consumed earthworms and used burrows of all native and invasive species of earthworms similarly. The potential for negative indirect effects of the invasive earthworm O. tyrtaeum on P. cinereus was demonstrated both in the field and laboratory, highlighting that seemingly small differences between native and invasive ecosystem engineers have the potential to significantly alter the effects of these closely related native and invasive organisms.

Although the use of engineered nanomaterials continues to increase, how these materials interact with coexisting contaminants in the environment is largely unknown. The effect of silver (Ag) in bulk, ionic, and nanoparticle (NP; bare and polyvinyl pyrrolidone-coated) forms at 3 concentrations (0 mg/kg, 500 mg/kg, 1000 mg/kg, 2000 mg/kg; ion at 69 mg/kg, 138 mg/kg, 276 mg/kg) on the accumulation of field-weathered chlordane and dichlorodiphenyldichloroethylene + metabolites (DDX) by Eisenia fetida (earthworm) was investigated. Earthworm biomass and survival were unaffected by treatment. At the 500 mg/kg and 1000 mg/kg exposure levels, NP-exposed earthworms contained significantly greater Ag (194-245%) than did the bulk exposed organisms; NP size or coating had no impact on element content. Generally, exposure to Ag of any type or at any concentration significantly reduced pesticide accumulation, although reductions for DDX (35.1%; 8.9-47.0%) were more modest than those for chlordane (79.0%; 17.4-92.9%). For DDX, the reduction in pesticide accumulation was not significantly affected by Ag type or concentration. For chlordane, the 3 NP exposures suppressed chlordane accumulation significantly more than did bulk exposure; earthworms exposed to bulk Ag contained 1170 ng/g chlordane, but levels in the NP-exposed earthworms were 279 ng/g. At the 500 mg/kg exposure, the smallest coated NPs exerted the greatest suppression in chlordane accumulation; at the 2 higher concentrations, chlordane uptake was unaffected by NP size or coating. The findings show that in exposed earthworms Ag particle size does significantly impact accumulation of the element itself, as well as that of coexisting weathered pesticides. The implications of these findings with regard to NP exposure and risk are unknown but are the topic of current investigation. Environ Toxicol Chem 2017;36:1864-1871. © 2016 SETAC.

Biosolids can be a valuable fertilizer for agriculture and in ecological restoration, although there are concerns about contaminants. Earthworm activity, including vermicomposting of biosolids, may influence the efficacy of their use. We investigated how two New Zealand endemic anecic species of (cf. ) responded to biosolids amendment and affected the mobility of nutrients and trace elements as well as greenhouse gas emissions in biosolids-amended soil. Earthworms were incubated with mixtures of biosolids-amended soil (0, 6.25, 12.5, 25, and 50% biosolids by volume) for 21 d. All species survived in the soil-biosolids mixtures but not in 100% biosolids. The native earthworms, and sp.2, increased KCl-extractable NH and NO by up to 29%, substantially more than . All species significantly increased microbial biomass carbon and Ca(NO)-extractable Cu but significantly decreased dehydrogenase enzymes activity in biosolids-amended soil. Concentrations of Ca(NO)-extractable Mg, S, Fe, Mn, Cd, Co, and Zn varied between earthworm species and with biosolids addition rates. New Zealand native earthworms exacerbated NO emissions from soil, whereas did not. is clearly a preferred species for vermicomposting biosolids and is more tolerant of high concentrations of biosolids. However, New Zealand native earthworms may be more suitable for improving the fertility of soil amended with biosolids.

Changes in understory plant composition and biodiversity declines in northeastern North American forests are widespread. Preserving species and ecosystem function requires appropriate identification and management of important stressors. Coexistence of stressors, among them earthworm invasions and white-tailed deer, makes correct identification of mechanisms that cause diversity declines challenging. We used an established factorial experiment to assess survival and growth of native seedlings (Actaea pachypoda, Aquilegia canadensis, Cornus racemosa, Quercus rubra, and Prenanthes alba) in response to presence/absence of deer and earthworms. We expected deer and earthworms to reduce seedling survival and biomass, and we evaluated potential pathways to explain this impact (soil N and P concentrations and pools, root architecture, and arbuscular mycorrhizal fungi [AMF] colonization). We developed structural equation models (SEM) to identify specific pathways through which earthworms and deer were impacting plant species with different life histories. Seedling survival was not affected by our treatments nor the plant and soil variables we tested. Actaea biomass was smaller in earthworm-invaded plots, and with larger total N pools. In contrast, both deer and earthworm treatments were associated with lower soil nutrient concentrations, and earthworm-invaded plots had smaller N and extractable P pools. Actaea, Cornus, Prenanthes, and Quercus seedlings had a lower proportion of fine roots in earthworm-invaded plots, while fine roots in Aquilegia made up a higher proportion of the root system. AMF colonization in Quercus was reduced in sites colonized by earthworms, but AMF in other species were unaffected. Our SEMs showed high correlation among soil variables, but because we do not know which variables are drivers of this change and which are passengers, we can only conclude that they are changing together as deer and earthworms exert their respective influence. Different plant species responded in idiosyncratic ways to earthworm and deer effects on soil fertility, root architecture and limited effects on AMF colonization. While earthworm and deer-mediated changes to fine roots, soil nutrients, and AMF may lead to changes in plant performance over time, these changes rarely translated to lower plant performance in our seedlings.

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Above–belowground (AG–BG) studies typically focus on plant-mediated effects inflicted by living organisms. However, animal cadavers may also play an important role in AG–BG interactions. Here, we explore whether living and dead foliar-feeding and soil-dwelling invertebrates differentially affect plants and their associated AG and BG multitrophic communities. In a mesocosm study we separated effects of living and dead locusts (AG herbivores) and earthworms (BG detritivores) on experimental multitrophic communities consisting of eight plant species, an AG aphid and parasitoid community and a BG nematode community. We measured root and shoot biomass and determined plant community composition and densities of aphids, parasitoids and nematodes. Living locusts decreased total shoot and root biomass in the mesocosms, whereas living earthworms enhanced total root biomass. Cadavers of both invertebrates strongly increased total root and shoot biomass, and changed the plant community composition mainly via enhanced growth of grasses. Earthworm cadavers affected plant biomass and community composition more strongly than their living counterparts, while this was reversed for locusts. Structural equation models showed that aphids and parasitoids were influenced via changes in plant community composition. Nematode densities in the soil, especially those of bacterivorous and entomopathogenic nematodes, were strongly increased by dead invertebrates, but unaffected by living ones. We conclude that effects of invertebrates on plant growth and densities of AG and BG organisms strongly depend on whether the invertebrates are dead or alive. Remarkably, invertebrate cadavers may inflict even stronger effects than their living counterparts. Hence, our study reveals an important, but often neglected, role of animal cadavers in AG–BG studies.

The preferential movement of water and transport of substances play an important role in soil, but they are not yet fully understood, especially in degraded peat soil. In this study, we aimed to deduce changes in flow and transport patterns with titanium dioxide (TiO2) as a dye tracer during the course of soil degradation resulting from peat drainage. The dye tracer experiments were carried out on columns of eight types of differently degraded peat soil from three sites taken in both vertical and horizontal directions. The titanium dioxide suspension (average particle size of 0.3 µm; 10 g l−1) was applied in a pulse of 40 mm to each soil core. The cores were subsequently cut into six slices, photographed and the images were analysed for the extent of dye cover and number of pores. In addition, the saturated hydraulic conductivity (Ks) was determined. Preferential flow occurred in all of the peat types investigated. From the stained soil structural elements, we concluded that undecomposed plant remains are the major preferential flow pathways in less degraded peat. For more strongly degraded peat, bio-pores, such as root and earthworm channels, operated as the major transport network. Results show that Ks and the effective pore network in less degraded peat soil are anisotropic. With increasing peat degradation, Ks and the cross-section of effective pores decreased in the predominant direction only and the anisotropy of both properties decreased. The Ks was closely related to the number of macropores and pore continuity. Therefore, we conclude that changes in flow and transport pathways and Ks with increasing peat degradation result from the disintegration of the peat-forming plant material and the decrease in number and continuity of macropores. Highlights We aimed to deduce changes in flow and transport patterns during peat degradation. TiO2 (dye tracer) was used to visualize the flow and transport patterns in degraded peat soil. The preferential flow paths, Ks and pore structures changed with peat degradation. Changes in flow and transport paths and Ks relate to peat-forming plant materials and pore structure.