Local distribution of native and invasive earthworms and effects on a native salamander

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.     

Effects of non-native Asian earthworm invasion on temperate forest and prairie soils in the Midwestern US

Effects of invasive European earthworms in North America have been well documented, but less is known about ecological consequences of exotic Asian earthworm invasion, in particular Asian jumping worms (Amynthas) that are increasingly reported. Most earthworm invasion research has focused on forests; some Amynthas spp. are native to Asian grasslands and may thrive in prairies with unknown effects. We conducted an earthworm-addition mesocosm experiment with before–after control-impact (BACI) design and a complementary field study in southern Wisconsin, USA, in 2014 to investigate effects of a newly discovered invasion of two Asian jumping worms (Amynthas agrestis and Amynthas tokioensis) on forest and prairie litter and soil nutrient pools. In both studies, A. agrestis and A. tokioensis substantially reduced surface litter (84–95 % decline in foliage litter mass) and increased total carbon, total nitrogen, and available phosphorus in the upper 0–5 cm of soils over the 4-month period from July through October. Soil inorganic nitrogen (ammonium– and nitrate–N) concentration increased across soil depths of 0–25 cm, with greater effects on nitrate–N. Dissolved organic carbon concentration also increased, e.g., 71–108 % increase in the mesocosm experiment. Effects were observed in both forest and prairie soils, with stronger effects in forests. Effects were most pronounced late in the growing season when earthworm biomass likely peaked. Depletion of the litter layer and rapid mineralization of nutrients by non-native Asian jumping worms may make ecosystems more susceptible to nutrient losses, and effects may cascade to understory herbs and other soil biota.     

Culturable microorganisms from the earthworm digestive tract

The cultured aerobic copiotrophic bacteria and fungi from food-free digestive tracts of Aporrectodea caliginosa, Lumbricus terrestris, and Eisenia fetida earthworms, soil (compost), and fresh earthworm excrements were investigated. The microorganisms were isolated on nutrient media and identified by sequencing the fragments of bacterial 16S rRNA and fungal 28S rRNA (D1/D2 domain) gene sequences with subsequent phylogenetic analysis. Bacteria isolated from the digestive tracts of earthworms belonged to the families Aeromonadaceae, Comamonadaceae, Enterobacteriaceae, Flavobacteriaceae, Moraxellaceae, Pseudomonadaceae, and Sphingobacteriaceae (Bacteroidetes), as well as Actinobacteria. For five strains, namely Ochrobactrum sp. 341-2 (α-Proteobacteria), Massilia sp. 557-1 (β-Proteobacteria), Sphingobacterium sp. 611-2 (Bacteroidetes), Leifsonia sp. 555-1, and a bacterium from the family Microbacteriaceae, isolate 521-1 (Actinobacteria), the similarity to known 16S rRNA sequences was 93–97%; they therefore, probably belong to new species and genera. Bacterial groups isolated from the digestive tracts of earthworms were significantly different from those isolated from soil and excrements. Some bacterial taxa occurred in different sections of A. caliginosa intestine and in intestines of different earthworm species; however, the overall composition of bacterial communities in these objects is different. Existence of bacterial groupings symbiotically associated with intestines is proposed. Among the fungi, Bjerkandera adusta and Syspastospora parasitica were isolated from the cleaned digestive tracts as light-colored, sterile mycelium, as well as Geotrichum candidum, Acremonium murorum (A. murorum var. felina), Alternaria alternata, Aspergillus candidus, A. versicolor, Cladosporium cladosporioides, Rhizomucor racemosus, Mucor hiemalis, Fusarium (F. oxysporum, Fusarium sp.), and Penicillium spp. These fungi survive for a long time in the earthworm’s digestive environment. Investigation of the functional characteristics and role in the host organism is required to confirm the symbiotic status of the microorganisms associated with the earthworm digestive tract.     

Earthworm Communities in the Bamboo Plantations of West Tripura (India)

Present study revealed the presence of 16 earthworm species belonging to 11 genera and four families viz. Megascolecidae (Amynthus alexandri, Metaphire houlleti, Lampito mauritii, Kanchuria sp1, Perionyx excavatus), Octochaetidae (Eutyphoeus gigas, Eutyphoeus comillahnus, Eutyphoeus orientalis, Octochaetona beatrix, Dichogaster bolaui, Lennogaster chittagongensis, Lennogaster yeicus), Moniligastridae (Drawida papillifer papillifer, Drawida assamensis, Drawida nepalensis) and Glossoscolecidae (Pontoscolex corethrurus) in the soils of five bamboo species [Bambusa balcooa (Sil Barak), Melocanna baccifera (Muli), Bambusa polumorpha (Bari), Bambus cacharensis (Bom) and Bambus bambus (Katabarak)] of West-Tripura. While four earthworm species viz. Metaphire houlleti, Drawida assamensis, Drawida papillifer papillifer and Pontoscolex corethrurus were common to all species of bamboo plantations, the rest showed restricted distribution. Among the earthworm species 4 were exotic (Amynthus alexandri, Metaphire houlleti, Dichogaster bolaui and Pontoscolex corethrurus) and the others were native to the Indian sub-continent. In general, earthworms under the bamboo plantations occurred within temperature range of 21.6 °C–28.0 °C, pH 4.0–7.0, organic matter 0.56–5.99 %, moisture 9.6–31.7 %, water holding capacity 14.6–43.9 % and bulk density 0.7–1.8 g cm^?3. The average density and biomass of the earthworms in the studied places were 108 ind m^?2 and 44 g m^?2 respectively. Earthworm diversity, dominance and evenness indices showed the values 1.00, 0.47 and 0.70 respectively. Earthworm density and biomass showed a negative correlation with temperature whereas those had a strong positive correlation with pH, moisture and organic matter of the soils.     

Tracking an invasion: community changes in hardwood forests following the arrival of <Emphasis Type="Italic">Amynthas agrestis</Emphasis> and <Emphasis Type="Italic">Amynthas tokioensis</Emphasis> in Wisconsin

Because Upper Midwest temperate forests lack native earthworms, the invasions of European and Asian earthworms can significantly alter soils and understory vegetation. Earthworms’ ability to increase leaf litter decay, alter nutrient cycling by mixing the organic layer with mineral soil, and decrease plant species richness leads to concern about the Asian ‘jumping earthworm’ (Amynthas agrestis and A. tokioensis) species that were recorded in the University of Wisconsin—Madison Arboretum in 2013. In 2015, we found A. agrestis and A. tokioensis in a distinct 8-ha region of a 23-ha hardwood forest surveyed in the Arboretum; by 2016 A. agrestis and A. tokioensis had spread over an additional 7 ha. Plots also contained the European earthworm species Lumbricus terrestris, L. rubellus, and Apporectodea spp., whose distributions decreased from 2015 to 2016. While leaf litter, plant species richness, and tree and shrub seedling abundance were generally reduced in areas with European earthworms, they were typically slightly increased in areas with A. agrestis and A. tokioensis versus those without. Although our results do not show substantial impacts of A. agrestis and A. tokioensis on vegetation in the initial years of invasion, the rapid replacement of European earthworms by A. agrestis and A. tokioensis suggests continued monitoring of these new invasive species is important to better understand their potential to change the Upper Midwest’s forests.     

Decrease in heathland soil labile organic carbon under future atmospheric and climatic conditions

Characterization of the impacts of climate change on terrestrial carbon (C) cycling is important due to possible feedback mechanisms to atmospheric CO₂ concentrations. We investigated soil organic matter (SOM) dynamics in the A1 and A2 horizons (~0–5.1 and ~5.1–12.3 cm depth, respectively) of a shrubland grass (Deschampsia flexuosa) after 8 years of exposure to: elevated CO₂ (CO₂), summer drought (D), warming (T) and all combinations hereof, with TDCO₂ simulating environmental conditions for Denmark in 2075. The mean C residence time was highest in the heavy fraction (HF), followed by the occluded light fraction and the free light fraction (fLF), and it increased with soil depth, suggesting that C was stabilized on minerals at depth. A2 horizon SOM was susceptible to climate change whereas A1 horizon SOM was largely unaffected. The A2 horizon fLF and HF organic C stocks decreased by 43 and 23% in response to warming, respectively. Organic nitrogen (N) stocks of the A2 horizon fLF and HF decreased by 50 and 17%, respectively. Drought decreased the A2 horizon fLF N stock by 38%. Elevated CO₂ decreased the A2 horizon fLF C stock by 39% and the fLF N stock by 50%. Under TDCO₂, A2 horizon fLF C and N stocks decreased by 22 and 40%, respectively. Overall, our results indicate that shrubland SOM will be susceptible to increased turnover and associated net C and N losses in the future.     

A comparative study of gut microbiota profiles of earthworms fed in three different substrates

The gut microbiome of earthworms has a complex interdependence with the host. When the soil minerals pass through earthworm’s gut, they may affect the gut microbiota. To gain insight into the response of gut microbiota to the passed minerals, we fed earthworm (Eisenia fetida) on nutrient-poor soil and ore powder, and used high throughput sequencing to characterize the earthworm intestinal microbial community to find evidence for a core bacterial community of the E. fetida. The results showed that earthworms’ gut maintained a core microbiome that appeared in all samples. These core microbiota may play a significant role in a species’ environmental interactions. The composition of intestinal microbiomes varied with substrates. The earthworm guts from two nutrient-poor substrates had similar microbial communities and they were different from nutrient-rich substrate. Proteobacteria and Bacteroidetes were more abundant in the gut of earthworms kept on a nutrient-poor substrate such as ore powder or mineral soil than in the gut of earthworms kept in organic-rich compost soil; some of these microorganisms may help earthworms survive in nutrient-poor substrates.     

Reaction of microorganisms to the digestive fluid of earthworms

The reaction of soil bacteria and fungi to the digestive fluid of the earthworm Aporrectodea caliginosa was studied. The fluid was obtained by centrifugation of the native enzymes of the digestive tract. The inhibition of growth of certain bacteria, spores, and fungal hyphae under the effect of extracts from the anterior and middle sections of the digestive tract of A. caliginosa was discovered for the first time. In bacteria, microcolony formation was inhibited as early as 20–30 s after the application of the gut extracts, which may indicate the nonenzymatic nature of the effect. The digestive fluid exhibited the same microbicidal activity whether the earthworms were feeding on soil or sterile sand. This indicates that the microbicidal agents are formed within the earthworm’s body, rather than by soil microorganisms. The effect of the digestive fluid from the anterior and middle divisions is selective in relation to different microorganisms. Of 42 strains of soil bacteria, seven were susceptible to the microbicidal action of the fluid (Alcaligenes faecalis 345-1, Microbacterium sp. 423-1, Arthrobacter sp. 430-1, Bacillus megaterium 401-1, B. megaterium 413-1, Kluyvera ascorbata 301-1, Pseudomonas reactans 387-2). The remaining bacteria did not die in the digestive fluid. Of 13 micromycetes, the digestive fluid inhibited spore germination in Aspergillus terreus and Paecilomyces lilacinus and the growth of hyphae in Trichoderma harzianum and Penicillium decumbens. The digestive fluid stimulated spore germination in Alternaria alternata and the growth of hyphae in Penicillium chrysogenum. The reaction of the remaining micromycetes was neutral. The gut fluid from the posterior division of the abdominal tract did not possess microbicidal activity. No relation was found between the reaction of microorganisms to the effects of the digestive fluid and the taxonomic position of the microorganisms. The effects revealed are similar to those shown earlier for millipedes and wood lice in the following parameters: quick action of the digestive fluid on microorganisms, and the selectivity of the action on microorganisms revealed at the strain level. The selective effect of the digestive gut fluid of the earthworms on soil microorganisms is important for animal feeding, maintaining the homeostasis of the gut microbial community, and the formation of microbial communities in soils.     

Soil microarthropods and macrofauna in monsoon tropical forests of Cat Tien and Bi Dup-Nui Ba National Parks,southern Vietnam

The abundance, biomass, vertical distribution, and taxonomic composition of soil invertebrates (springtails, macrofauna, and termites) were studied in forest formations differing in edaphic and climatic conditions: lowland forests dominated by Lagerstroemia spp. or Dipterocarpus spp. in the Cat Tien National Park and in a mountain pine (Pinus kesiya) forest on the Da Lat Plateau, southern Vietnam. In the lowland forests, springtails had a relatively low density (10000–12000 ind./m²), but their diversity was high (41–43 species in each forest). The density of large soil invertebrates (without ants and termites) reached 500–700 ind./m² at a biomass of approximately 30 g/m² (with earthworms accounting for up to 230 ind./m² and 19–28 g/m²). Among termites, species of the genera Macrotermes and Odontotermes were dominant. Their total biomass in some areas exceeded 15–20 g/m². In the mountain pine forest, the total biomass of soil macrofauna was approximately 11 g/m², the abundance and diversity of springtails were low (7500 ind./m², 28 species), and wood-destroying species of the genera Schedorhinotermes sp. and Coptotermes sp. dominated among termites.     

Crop rotation associating a legume and the nickel hyperaccumulator <Emphasis Type="Italic">Alyssum murale</Emphasis> improves the structure and biofunctioning of an ultramafic soil

Nickel (Ni) agromining aims to phytoextract heavy metals using hyperaccumulators whilst at the same time rehabilitating ultramafic soils. After removing the bioavailable metal, ultramafic soils are improved in terms of their agronomic properties with the aim of future agricultural uses. The low fertility of ultramafic soils can be compensated by integrating legumes already used in traditional agro-systems because of their importance in soil nitrogen enrichment. However, few studies have evaluated the potential profits of legumes on Ni agromining and their potential benefits on soil biological fertility. Here, we characterized the effect of a crop rotation with two plants, a legume (Vicia sativa) and a hyperaccumulator (Alyssum murale), on the phytoextraction efficiency and on soil structure and biofunctioning. A pot experiment was set up in controlled conditions to grow A. murale and four treatments were tested: rotation with V. sativa (Ro), fertilized mono-culture (FMo), non-fertilized mono-culture (NFMo) and bare soil without plants (BS). No significant difference was found between the Ro and NFMo treatments for the dry biomass yield. However, the Ro treatment showed the highest Ni concentrations ([Ni]) in A. murale shoots compared to FMo and NFMo treatments. The Ro treatment plants had more than twice as many leaves [Ni] compared to FMo. Soil physico-chemical analyses showed that the Ro treatment was better structured and showed the highest presence of bacterial micro-aggregates, as well as less non-aggregated particles. Legumes integration in Ni-agromining systems could be a pioneering strategy to reduce chemical inputs and to improve soil biofunctioning and thus fertility.     

High abundance of non-mycorrhizal plant species in severely phosphorus-impoverished Brazilian <Emphasis Type="Italic">campos rupestres</Emphasis>

Background and aims

We sought to describe the species and functional composition of Brazilian campos rupestres plant communities on severely nutrient-impoverished white sands, to test hypotheses relating plant communities and physiological adaptations to infertile soils. Based on recently-published information on a south-western Australian dune chronosequence, we hypothesised that campos rupestres plant communities would similarly contain a relatively large proportion of non-mycorrhizal species, because of the phosphorus-(P) impoverished nature of the soils. We also sought to test the hypothesis that many of these non-mycorrhizal species have high leaf manganese (Mn) concentrations as a consequence of carboxylate exudation to mobilise soil P.

Methods

We conducted flora surveys and quantified mycorrhizal status and foliar Mn concentrations in field sites with strongly-weathered sandy soils. Rhizosphere carboxylates were collected from glasshouse-grown plants to assess a potential correlation of carboxylates and leaf Mn concentrations.

Results

Soils were depleted of all major plant nutrients. Non-mycorrhizal plants were abundant in most field sites (mean relative cover = 48%). Vellozia species were dominant aboveground; belowground, roots were colonised more by dark septate endophytic fungi than by mycorrhizal fungi. From the field sites, foliar Mn concentrations in non-mycorrhizal species increased with decreasing soil P concentrations, but only when soil Mn concentrations were above a minimum threshold (exchangeable [Mn] above detection limit). Across all species, however, there was no relationship of foliar Mn concentrations with soil P concentrations.

Conclusions

Our hypothesis that white-sand campos rupestres communities contain a relatively large proportion of non-mycorrhizal plants was supported. Comparison with similar ecosystems in south-western Australia suggests that plant communities on severely P-impoverished sandy soils, despite differing evolutionary histories and little overlap in plant families, follow convergent evolutionary paths towards increasing abundance of non-mycorrhizal species.

     

A qPCR assay that specifically quantifies <Emphasis Type="Italic">Tricholoma matsutake</Emphasis> biomass in natural soil

Tricholoma matsutake is an ectomycorrhizal basidiomycete that produces prized, yet uncultivable, “matsutake” mushrooms along densely developed mycelia, called “shiro,” in the rhizosphere of coniferous forests. Pinus densiflora is a major host of this fungus in Japan. Measuring T. matsutake biomass in soil allows us to determine the kinetics of fungal growth before and after fruiting, which is useful for analyzing the conditions of the shiro and its surrounding mycorrhizosphere, predicting fruiting timing, and managing forests to obtain better crop yields. Here, we document a novel method to quantify T. matsutake mycelia in soil by quantifying a single-copy DNA element that is uniquely conserved within T. matsutake but is absent from other fungal species, including close relatives and a wide range of ectomycorrhizal associates of P. densiflora. The targeted DNA region was amplified quantitatively in cultured mycelia that were mixed with other fungal species and soil, as well as in an in vitro co-culture system with P. densiflora seedlings. Using this method, we quantified T. matsutake mycelia not only from shiro in natural environments but also from the surrounding soil in which T. matsutake mycelia could not be observed by visual examination or distinguished by other means. It was demonstrated that the core of the shiro and its underlying area in the B horizon are predominantly composed of fungal mycelia. The fungal mass in the A or A₀ horizon was much lower, although many white mycelia were observed at the A horizon. Additionally, the rhizospheric fungal biomass peaked during the fruiting season.     

Exotic earthworms dispersion through protected forest areas and their potential impacts on nitrous oxide production

It is generally accepted that human activities are responsible for the dispersal of exotic earthworms in northeastern North America. We know little, however, about the relative effects of concurrent human activities on the structure of these earthworm communities in protected forest areas, nor on their impacts on soil biological activities. Our first objective was to infer the relative importance of recreational fishing and road traffic on the structure of Lumbricidae communities in Mont-Tremblant National Park, the oldest conservation area in the province of Quebec, Canada. Our second objective was to test the relationship between earthworm species abundances and soil properties related to microbial and nitrogen dynamics. We sampled earthworm communities around 61 lakes, which included 23 heavily-fished lakes and 20 non-fished lakes located near roads, as well as 18 non-fished lakes located in remote areas of the park. Our results revealed that fishing and proximity to roads both have a positive effect on the abundance of earthworms, as does the soil pH. Fishing activities had a greater effect than road proximity on the abundance and diversity of earthworm communities, notably on the abundance of the anecic species Lumbricus terrestris. To assess at a finer scale the effects of earthworm community structure on soil microbial and nitrogen dynamics, we collected and analyzed soils from 47 sampling points around two lakes with high earthworm densities. Exploratory redundancy analysis found a negative correlation between epigeic and anecic earthworm species, with the former correlating positively to microbial biomass and the latter correlating positively to nitrification and denitrification. Confirmatory path analysis established a positive indirect effect of Lumbricus terrestris, the preferred fishing bait, on potential soil nitrous oxide emissions. We conclude that the human-mediated dispersion of earthworms in the most pristine ecosystems of Quebec affects ecosystem functioning and thus requires a review of current policies regarding the use of live-bait by fishermen.     

Human-mediated introduction of geoengineering earthworms in the Fennoscandian arctic

It is now well established that European earthworms are re-shaping formerly glaciated forests in North America with dramatic ecological consequences. However, few have considered the potential invasiveness of this species assemblage in the European arctic. Here we argue that some earthworm species (Lumbricus rubellus, Lumbricus terrestris and Aporrectodea sp.) with great geomorphological impact (geoengineering species) are non-native and invasive in the Fennoscandian arctic birch forests, where they have been introduced by agrarian settlers and most recently through recreational fishing and gardening. Our exploratory surveys indicate no obvious historical dispersal mechanism that can explain early arrival of these earthworms into the Fennoscandian arctic: that is, these species do not appear to establish naturally along coastlines mimicking conditions following deglaciation in Fennoscandia, nor were they spread by early native (Sami) cultures. The importance of anthropogenic sources and the invasive characteristics of L. rubellus and Aporrectodea sp. in the arctic is evident from their radiation outwards from abandoned farms and modern cabin lawns into adjacent arctic birch forests. They appear to outcompete previously established litter-dwelling earthworm species (i.e. Dendrobaena octaedra) that likely colonized the Fennoscandian landscape rapidly following deglaciation via hydrochory and/or dispersal by early Sami settlements. The high geoengineering earthworm biomasses, their recognized ecological impact in other formerly glaciated environments, and their persistence once established leads us to suggest that geoengineering earthworms may pose a potent threat to some of the most remote and protected arctic environments in northern Europe.     

The population of earthworms (Lumbricidae) in the main types of dark coniferous forests in Pechora-Ilych Nature Reserve

The population of earthworms has been studied in the main types of old-growth dark coniferous forests of Pechora-Ilych Nature Reserve (Komi Republic) that have not been subject to anthropogeniс impact for a long time. Ten species of earthworms have been identified. The greatest diversity (7 species), abundance, and biomass of earthworms has been revealed in the tall-grass fir–spruce forests. P. diplotetratheca had the greatest abundance. E. nordenskioldi nordenskioldi and E. atlavinyteae had the greatest biomass. The lowest species diversity (3 species), abundance, and biomass of earthworms have been found in the largefern, blueberry–green moss, and sphagnum–horsetail fir–spruce forests. The role of deadwood in maintaining the species diversity of Lumbricidae in dark coniferous forests has been demonstrated. The complexes of Lumbricidae have been considered in anthropogenically disturbed territories, where the following species with a habitat range to the south of the northern and middle taiga have been identified: L. rubellus, A. rosea, A. caliginosa caliginosa, and E. fetida.     

Biocontrol of the toxigenic plant pathogen <Emphasis Type="Italic">Fusarium culmorum</Emphasis> by soil fauna in an agroecosystem

In 2011 and 2013, a field experiment was conducted in a winter wheat field at Adenstedt (northern Germany) to investigate biocontrol and interaction effects of important members of the soil food web (Lumbricus terrestris, Annelida; Folsomia candida, Collembola and Aphelenchoides saprophilus, Nematoda) on the phytopathogenic fungus Fusarium culmorum in wheat straw. Therefore, soil fauna was introduced in mesocosms in defined numbers and combinations and exposed to either Fusarium-infected or non-infected wheat straw. L. terrestris was introduced in all faunal treatments and combined either with F. candida or A. saprophilus or both. Mesocosms filled with a Luvisol soil, a cover of different types of wheat straw and respective combinations of faunal species were established outdoors in the topsoil of a winter wheat field after harvest of the crop. After a time span of 4 and 8 weeks, the degree of wheat straw coverage of mesocosms was quantified to assess its attractiveness for the soil fauna. The content of Fusarium biomass in residual wheat straw and soil was determined using a double-antibody sandwich (DAS)-ELISA method. In both experimental years, the infected wheat straw was incorporated more efficiently into the soil than the non-infected control straw due to the presence of L. terrestris in all faunal treatments than the non-infected control straw. In addition, Fusarium biomass was reduced significantly in all treatments after 4 weeks (2011: 95–99%; 2013:15–54%), whereupon the decline of fungal biomass was higher in faunal treatments than in non-faunal treatments and differed significantly from them. In 2011, Fusarium biomass of the faunal treatments was below the quantification limit after 8 weeks. In 2013, a decline of Fusarium biomass was observed, but the highest content of Fusarium biomass was still found in the non-faunal treatments after 8 weeks. In the soil of all treatments, Fusarium biomass was below the quantification limit. The earthworm species L. terrestris revealed a considerable potential as an effective biocontrol agent contributing to a sustainable control of a Fusarium plant pathogen in wheat straw, thus reducing the infection risk for specific plant diseases in arable fields.     

Interactions of the manganese hyperaccumulator <Emphasis Type="Italic">Phytolacca americana</Emphasis> L. with soil pH and phosphate

Hyperaccumulators are plants that store exceptionally high concentrations of heavy metals or metalloids in their leaves. Phytolacca americana is one of the few species known to hyperaccumulate manganese (Mn); however, it is a common weedy species and has no specific association with high-Mn soils. Neither the mechanism by which P. americana hyperaccumulates Mn nor the ecological significance of this trait are well understood. It has recently been suggested that P. americana secretes acids into the rhizosphere as a means of acquiring phosphate, which might coincidentally increase Mn uptake. To determine whether P. americana acidifies the surrounding soil, plants were grown in rhizoboxes providing access to living roots. A thin layer of agar containing bromocresol green pH indicator dye was placed on the roots to observe color changes indicating acidification. Comparative studies showed that P. americana acidifies the rhizosphere significantly more than the non-accumulating plant Acalypha rhomboidea. A second experiment studied whether adjustment of soil pH and phosphate affect foliar Mn concentrations of P. americana. Concentrations of Mn in leaves were highest when plants were grown in acidified soils but were significantly lower in soils that were alkaline and/or enriched with phosphate. These results suggest that Mn hyperaccumulation may be a side effect of rhizosphere acidification as a phosphorus-acquisition mechanism, rather than an adaptation in its own right. The findings provide fundamental information about hyperaccumulator physiology and evolution, and may be relevant to attempts to utilize P. americana for phytoremediation.     

Concentration of some metals in soil and plant organs and their biochemical profiles in <Emphasis Type="Italic">Tulipa luanica</Emphasis>, <Emphasis Type="Italic">T. kosovarica</Emphasis> and <Emphasis Type="Italic">T. albanica</Emphasis> native plant species

The purpose of this study was to determine the concentration of some metals (Al, Cd, Co, Cr, Cu, Fe, Mn, Ni, Pb, Zn, Ca and Mg) in soil of serpentine and limestone sites, their bioaccumulation and impact on some biochemical parameters in T. luanica, T. kosovarica and T. albanica plants. T. kosovarica and T. albanica grows in serpentine soil, while T. luanica grow in limestone soil. The research showed that concentrations of Cd, Co, Cr, Fe, Mn and Ni were significantly higher at serpentine soil sites in comparison with limestone sites, while concentrations of Pb, Cd, Co and Cr in bulbs, leaves and seeds were under the limit of detection. The concentration of Ni in plant samples of T. kosovarica was significantly higher in comparison with its concentration in T. albanica, but it was under the limit of detection in T. luanica. Moreover, concentrations of Al and Fe in leaves of T. kosovarica and T. albanica were higher in comparison with T. luanica. The concentration of Mg was significantly higher in T. kosovarica and T. albanica than in T. luanica. The δ-aminolevulinic acid dehydratase activity, malondialdehyde and glutathione contents in leaves of T. luanica were higher in comparison with T. kosovarica and T. albanica. In addition, the amounts of total chlorophyll and δ-aminolevulinic acid (ALA) in leaves of T. albanica were higher in comparison with T. kosovarica and T. luanica. Our findings show that target organs of metal accumulation in three Tulip species appears to be leaves?>?seeds?>?bulbs, while the biochemical parameters show that limestone sites represent a less stressful habitat for growing these plant species in comparison with serpentine sites.     

Strong increase of durum wheat iron and zinc content by field-inoculation with arbuscular mycorrhizal fungi at different soil nitrogen availabilities

Background and aims

The changes in the characteristics of Panicum virgatum, an exotic invasive species, after invading various plant communities on the Loess Plateau in China and the main soil nutrient factors in these communities closely associated with invasion remain unclear.

Methods

A pot culture experiment was carried out to simulate the changes in photosynthesis, biomass, and biomass allocation in P. virgatum and to identify the main soil nutrient factors in various soils collected from local plant communities. P. virgatum was grown in soils collected from communities of P. virgatum (PS treatment), Setaria viridis (SS treatment), Bothriochloa ischaemum (BS treatment), and Artemisia sacrorum (AS treatment) and in a mixed soil from the communities of S. viridis, B. ischaemum, and A. sacrorum (MS treatment).

Results

Photosynthesis in P. virgatum differed significantly among the soil treatments. Net photosynthetic rate, stomatal conductance, and photochemical efficiency (Fv/Fm) were highest in PS, whereas single-photon avalanche diode values were highest in PS and SS. The variation of biomass differed significantly in different tissues of P. virgatum in the treatments. Leaf and stem biomasses were highest in PS and SS, and root biomass was highest in PS and MS. Total biomass differed significantly among the treatments, except between BS and MS. Both the leaf to total and stem to total biomass ratios were highest in AS and SS, but the root to total biomass ratio was lowest in these two treatments. A constrained redundancy analysis and a path analysis suggested that the water-soluble nitrate-nitrogen (W-NN) concentration of the soil could significantly affect photosynthesis, biomass, and biomass allocation in P. virgatum.

Conclusions

Photosynthesis, biomass, and biomass allocation in P. virgatum differed significantly when grown in soils from different local plant communities on the Loess Plateau. The soil W-NN concentration in these local plant communities likely has a large impact on the invasive success of P. virgatum.

     

Local-scale correlates of native and non-native earthworm distributions in juniper-encroached tallgrass prairie

European and Asian earthworms have invaded much of North America with profound impacts to soils, plant communities, and animal populations. However, few studies have assessed local-scale correlates of earthworm distributions, and most invasive earthworm research has occurred in northern forests. Additionally, despite several studies showing facilitative relationships between invasive earthworms and invasive plants, no research has assessed a potential facilitative interaction between earthworms and woody plants encroaching into prairies. We conducted the first assessment of factors influencing local-scale distributions of native and non-native earthworms for the U.S. Great Plains in a tallgrass prairie-woodland mosaic experiencing eastern redcedar (Juniperus virginiana) encroachment. We documented both native and non-native earthworms, including non-native species from Eurasia (Aporrectodea spp.) and South America (Family Ocnerodrilidae). Native and non-native earthworm distributions were strongly correlated, yet local-scale predictors of distribution also differed between the groups. Native earthworms were more likely to occur near roads and in areas with moist soils. Contrary to expectation, we found no evidence that non-native earthworms occurred more frequently in areas with eastern redcedar-encroachment; instead, non-native earthworms were most likely to occur in tallgrass prairie. Our results suggest that, within prairies and woodlands of the Great Plains, native and non-native earthworms occur most frequently near roadways and in locations with moist soil. Because the few approaches for controlling invasive earthworms are only likely to be feasible on a small scale, findings from such local-scale studies are important for directing management to reduce earthworm impacts on biodiversity and ecosystem services.