Molecular Biomonitoring during Rhizoremediation of Oil-Contaminated Soil

Rhizoremediation is the use of microbial populations present in the rhizosphere of plants for environmental cleanup. The idea of this work was that bacteria living in the rhizosphere of a nitrogen-fixing leguminous plant, goat's rue (Galega orientalis), could take part in the degradation of harmful monoaromatic hydrocarbons, such as benzene, toluene and xylene (BTEX), from oil-contaminated soils. In addition to chemical (e.g. pollutant concentration) and physical (e.g. soil structure) information, the knowledge of biological aspects (e.g. bacteria and their catabolic genes) is essential when developing the rhizoremediation into controlled and effective bioremediation practice. Therefore, the need for reliable biomonitoring methods is obvious. The main aims of this thesis were to evaluate the symbiotic G. orientalis - Rhizobium galegae system for rhizoremediation of oil-contaminated soils, to develop molecular methods for biomonitoring, and to apply these methods for studying the microbiology of rhizoremediation. In vitro, Galega plants and rhizobia remained viable in m-toluate concentrations up to 3000 mg/l. Plant growth and nodulation were inhibited in 500 mg/l m-toluate, but were restored when plants were transferred to clean medium. In the greenhouse, Galega showed good growth, nodulation and nitrogen fixation, and developed a strong rhizosphere in soils contaminated with oil or spiked with 2000 mg/l m-toluate. The high aromatic tolerance of R. galegae and the viability of Galega plants in oil-polluted soils proved this legume system to be a promising method for the rhizoremediation of oil-contaminated soils. Molecular biomonitoring methods were designed and/or developed further for bacteria and their degradation genes. A combination of genomic fingerprinting ((GTG)5-PCR), taxonomic ribotyping of 16S rRNA genes and partial 16S rRNA gene sequencing were chosen for molecular grouping of culturable, heterogeneous rhizosphere bacteria. PCR primers specific for the xylE gene were designed for TOL plasmid detection. Amplified enzyme-coding DNA restriction analysis (AEDRA) with AluI was used to profile both TOL plasmids (xylE primers) and, in general, aromatics-degrading plasmids (C230 primers). The sensitivity of the direct monitoring of TOL plasmids in soil was enhanced by nested C23O-xylE-PCR. Rhizosphere bacteria were isolated from the greenhouse and field lysimeter experiments. High genetic diversity was observed among the 50 isolated, m-toluate tolerating rhizosphere bacteria in the form of five major lineages of the Bacteria domain. Gram-positive Rhodococcus, Bacillus and Arthrobacter and gram-negative Pseudomonas were the most abundant genera. The inoculum Pseudomonas putida PaW85/pWW0 was not found in the rhizosphere samples. Even if there were no ecological niches available for the bioaugmentation bacterium itself, its conjugative catabolic plasmid might have had some additional value for other bacterial species and thus, for rhizoremediation. Only 10 to 20% of the isolated, m-toluate tolerating bacterial strains were also able to degrade m-toluate. TOL plasmids were a major group of catabolic plasmids among these bacteria. The ability to degrade m-toluate by using enzymes encoded by a TOL plasmid was detected only in species of the genus Pseudomonas, and the best m-toluate degraders were these Pseudomonas species. Strain-specific differences in degradation abilities were found for P.oryzihabitans and P. migulae: some of these strains harbored a TOL plasmid - a new finding observed in this work, indicating putative horizontal plasmid transfer in the rhizosphere. One P. oryzihabitans strain harbored the pWW0 plasmid that had probably conjugated from the bioaugmentation Pseudomonas. Some P. migulae and P. oryzihabitans strains seemed to harbor both the pWW0- and the pDK1-type TOL plasmid. Alternatively, they might have harbored a TOL plasmid with both the pWW0- and the pDK1-type xylE gene. The breakdown of m-toluate by gram-negative bacteria was not restricted to the TOL pathway. Also some gram-positive Rhodococcus erythropolis and Arthrobacter aurescens strains were able to degrade m-toluate in the absence of a TOL plasmid. Three aspects of the rhizosphere effect of G. orientalis were manifested in oil-contaminated soil in the field: 1) G. orientalis and Pseudomonas bioaugmentation increased the amount of rhizosphere bacteria. G. orientalis especially together with Pseudomonas bioaugmentation increased the numbers of m-toluate utilizing and catechol positive bacteria indicating an increase in degradation potential. 2) Also the bacterial diversity, when measured as the amount of ribotypes, was increased in the Galega rhizosphere with or without Pseudomonas bioaugmentation. However, the diversity of m-toluate utilizing bacteria did not significantly increase. At the community level, by using the 16S rRNA gene PCR-DGGE method, the highest diversity of species was also observed in vegetated soils compared with non-vegetated soils. Diversified communities may best guarantee the overall success in rhizoremediation by offering various genetic machineries for catabolic processes. 3) At the end of the experiment, no TOL plasmid could be detected by direct DNA analysis in soil treated with both G. orientalis and Pseudomonas. The detection limit for TOL plasmids was encountered indicating decreased amount of degradation plasmids and thus, the success of rhizoremediation. The use of G. orientalis for rhizoremediation is unique. In this thesis new information was obtained about the rhizosphere effect of Galega orientalis in BTEX contaminated soils. The molecular biomonitoring methods can be applied for several purposes within environmental biotechnology, such as for evaluating the intrinsic biodegradation potential, monitoring the enhanced bioremediation, and estimating the success of bioremediation. Environmental protection by using nature's own resources and thus, acting according to the principle of sustainable development, would be both economically and environmentally beneficial for society. Keywords: molecular biomonitoring, genetic fingerprinting, soil bacteria, bacterial diversity, TOL plasmid, catabolic genes, horizontal gene transfer, rhizoremediation, rhizosphere effect, Galega orientalis, aerobic biodegradation, petroleum hydrocarbons, BTEX

Physical properties of a boreal clay soil under differently managed perennial vegetation

The physical properties of surface soil horizons, essentially pore size, shape, continuity and affinity for water, regulate water entry into the soil. These properties are prone to changes caused by natural forces and human activity. The hydraulic properties of the surface soil greatly impact the generation of surface runoff and accompanied erosion, the major concern of agricultural water protection. The general target of this thesis was to improve our understanding of the structural and hydraulic properties of boreal clay soils. Physical properties of a clayey surface soil (0 - 10 cm, clay content 51%), with a micaceous/illitic mineralogy subjected to three different management practices of perennial vegetation, were studied. The study sites were vegetated buffer zones located side by side in SW Finland: 1) natural vegetation with no management, 2) harvested once a year, and 3) grazed by cattle. The soil structure, hydraulic properties, shrinkage properties and soil water repellency were determined at all sites. Two distinct flow domains were evident. The surface soil was characterized by subangular blocky, angular blocky and platy aggregates. Hence, large, partially accommodated, irregular elongated pores dominated the macropore domain at all sites. The intra-aggregate pore system was mostly comprised of pores smaller than 30 μm, which are responsible for water storage. Macropores at the grazed site, compacted by hoof pressure, were horizontally oriented and pore connectivity was poorest, which decreased water and air flux compared with other sites. Drying of the soil greatly altered its structure. The decrease in soil volume between wet and dry soil was 7 - 10%, most of which occurred in the moisture range of field conditions. Structural changes, including irreversible collapse of interaggregate pores, began at matric potentials around -6 kPa indicating, instability of soil structure against increasing hydraulic stress. Water saturation and several freezethaw cycles between autumn and spring likely weakened the soil structure. Soil water repellency was observed at all sites at the time of sampling and when soil was dryer than about 40 vol.%. (matric potential < -6 kPa). Therefore, water repellency contributes to water flow over a wide moisture range. Water repellency was also observed in soils with low organic carbon content (< 2%), which suggests that this phenomenon is common in agricultural soils of Finland due to their relatively high organic carbon content. Aggregate-related pedofeatures of dense infillings described as clay intrusions were found at all sites. The formation of these intrusions was attributed to clay dispersion and/or translocation during spring thaw and drying of the suspension in situ. These processes generate very new aggregates whose physical properties are most probably different from those of the bulk soil aggregates. Formation of the clay infillings suggested that prolonged wetness in autumn and spring impairs soil structure due to clay dispersion, while on the other hand it contributes to the pedogenesis of the soil. The results emphasize the dynamic nature of the physical properties of clay soils, essentially driven by their moisture state. In a dry soil, fast preferential flow is favoured by abundant macropores including shrinkage cracks and is further enhanced by water repellency. Increase in soil moisture reduces water repellency, and swelling of accommodated pores lowers the saturated hydraulic conductivity. Moisture- and temperature-related processes significantly alter soil structure over a time span of 1 yr. Thus, the pore characteristics as well as the hydraulic properties of soil are time-dependent.

Collapse behaviour of a residual soil

Residually derived red soils occur in Bangalore District of Karnataka State, India. The porous and unsaturated nature of the red soils makes them susceptible to collapse on wetting under load. The present study analyses the collapse behaviour of an unsaturated bonded (undisturbed) red soil from Bangalore referenced to tests on samples in an unbonded (remoulded) state. A filter paper method was used to determine the matric suction of the bonded and unbonded specimens, and mercury intrusion porosimetry (MIP) was used to determine their soil structure. Analysis of the experimental results shows that bonding plays an important role in the collapse behaviour of the unsaturated residual soil. The results of the study also provide insight into the volume change behaviour of unsaturated bonded soils on wetting within and beyond the yield locus.

MODELING AND SIMULATION OF HYDRODYNAMIC INTERACTION OF DNA IN A MICRO-FLUIDIC CHANNEL

The motion of DNA (in the bulk solution) and the non-Newtonian effective fluid behavior are considered separately and self-consistently with the fluid motion satisfying the no-slip boundary condition on the surface of the confining geometry in the presence of channel pressure gradients. A different approach has been developed to model DNA in the micro-channel. In this study the DNA is assumed as an elastic chain with its characteristic Young's modulus, Poisson's ratio and density. The force which results from the fluid dynamic pressure, viscous forces and electromotive forces is applied to the elastic chain in a coupled manner. The velocity fields in the micro-channel are influenced by the transport properties. Simulations are carried out for the DNAs attached to the micro-fluidic wall. Numerical solutions based on a coupled multiphysics finite element scheme are presented. The modeling scheme is derived based on mass conservation including biomolecular mass, momentum balance including stress due to Coulomb force field and DNA-fluid interaction, and charge transport associated to DNA and other ionic complexes in the fluid. Variation in the velocity field for the non-Newtonian flow and the deformation of the DNA strand which results from the fluid-structure interaction are first studied considering a single DNA strand. Motion of the effective center of mass is analyzed considering various straight and coil geometries. Effects of DNA statistical parameters (geometry and spatial distribution of DNAs along the channel) on the effective flow behavior are analyzed. In particular, the dynamics of different DNA physical properties such as radius of gyration, end-to-end length etc. which are obtained from various different models (Kratky-Porod, Gaussian bead-spring etc.) are correlated to the nature of interaction and physical properties under the same background fluid environment.

Nonlinear analysis of pile foundation jacket platform

In this paper, the nonlinear collapse of the BOHAI-8 pile foundation jacket platform has been analyzed. The ultimate load and collapse process of two computational models of the structure are given. One model is of fixed support whose length is eight times the pile leg diameter and the other considers the nonlinearity of the soil-pile interaction.

Fluid-structure Interactions of Inverted Leaves and Flags

Interactions between fluid flows and elastic bodies are ubiquitous in nature. One such phenomena that is encountered on a daily basis is the flapping and fluttering of leaves in the wind. The fluid-structure interaction that governs the physics of a leaf in the wind is poorly understood at best and has potential applications in biomechanics, vehicle design, and energy conversion. We build upon previous work on the flapping dynamics of inverted flags, which are cantilevered elastic sheets with free leading edge and fixed trailing edge that display unique large amplitude oscillatory behaviors. We model a leaf in the laboratory using modified inverted flags, experimentally probing the governing parameters behind leaf fluttering as well as shedding light on the physics behind the inverted flag phenomena. The behavior of these "inverted leaves" studied here display sensitive dependence on two biomechanically relevant parameters, stem-to-leaf rigidity and stem-to-leaf length. In addition, leaves on a tree are not often found alone. We seek to understand the complex interactions of multiple fluttering and flapping leaves by way of examining the interactions between pairs of inverted flags. Coupling through their flow fields, pairs of inverted flags exhibit striking emergent phenomena. We report these observed dynamical behaviors and the conditions upon which they arise.

Deformation and fracture of impulsively loaded sandwich panels

Light metal sandwich panel structures with cellular cores have attracted interest for multifunctional applications which exploit their high bend strength and impact energy absorption. This concept has been explored here using a model 6061-T6 aluminum alloy system fabricated by friction stir weld joining extruded sandwich panels with a triangular corrugated core. Micro-hardness and miniature tensile coupon testing revealed that friction stir welding reduced the strength and ductility in the welds and a narrow heat affected zone on either side of the weld by approximately 30%. Square, edge clamped sandwich panels and solid plates of equal mass per unit area were subjected to localized impulsive loading by the impact of explosively accelerated, water saturated, sand shells. The hydrodynamic load and impulse applied by the sand were gradually increased by reducing the stand-off distance between the test charge and panel surfaces. The sandwich panels suffered global bending and stretching, and localized core crushing. As the pressure applied by the sand increased, face sheet fracture by a combination of tensile stretching and shear-off occurred first at the two clamped edges of the panels that were parallel with the corrugation and weld direction. The plane of these fractures always lay within the heat affected zone of the longitudinal welds. For the most intensively loaded panels additional cracks occurred at the other clamped boundaries and in the center of the panel. To investigate the dynamic deformation and fracture processes, a particle-based method has been used to simulate the impulsive loading of the panels. This has been combined with a finite element analysis utilizing a modified Johnson-Cook constitutive relation and a Cockcroft-Latham fracture criterion that accounted for local variation in material properties. The fully coupled simulation approach enabled the relationships between the soil-explosive test charge design, panel geometry, spatially varying material properties and the panel's deformation and dynamic failure responses to be explored. This comprehensive study reveals the existence of a strong instability in the loading that results from changes in sand particle reflection during dynamic evolution of the panel's surface topology. Significant fluid-structure interaction effects are also discovered at the sample sides and corners due to changes of the sand reflection angle by the edge clamping system. © 2012 Elsevier Ltd. All rights reserved.

Effect of desert soil algae on the stabilization of fine sands

Four filamentous cyanobacteria, Microcoleus vaginatus, Phormidium tenue, Scytonema javanicum (Kutz.) and Nostoc sp., and a single-celled green alga, Desmococcus olivaceus, all isolated from Shapotou (Ningxia Hui Autonomous Region of China), were batch cultured and inoculated onto unconsolidated sand in greenhouse and field experiments. Their ability to reduce wind erosion in sands was quantified by using a wind tunnel laboratory. The major factors related to cohesion of algal crusts, such as biomass, species, species combinations, bioactivity, niche, growth phase of algae, moisture, thickness of the crusts, dust accretion (including dust content and manner of dust added) and other cryptogams (lichens, fungi and mosses) were studied. The best of the five species were M. vaginatus and P. tenue, while the best mix was a blend of 80% M. vaginatus and 5% each of P. tenue, S. javanicum, Nostoc sp. and D. olivaceus. The threshold friction velocity was significantly increased by the presence of all of the cyanobacterial species, while the threshold impact velocity was notably increased only by the filamentous species. Thick crusts were less easily eroded than thin crusts, while biomass was more effective than thickness. Dust was incorporated best into Microcoleus crust when added in small amounts over time, and appeared to increase growth of the cyanobacterium as well as strengthen the cohesion of the crust. Microbial crust cohesion was mainly attributed to algal aggregation, while lichens, fungi and mosses affected more the soil structure and physico-chemical properties.

Methods for physical separation and characterization of soil organic matter fractions

Turnover of soil organic matter (SOM) is coupled to the cycling of nutrients in soil through the activity of soil microorganisms. Biological availability of organic substrate in soil is related to the chemical quality of the organic material and to its degree of physical protection. SOM fractions can provide information on the turnover of organic matter (OM), provided the fractions can be related to functional or structural components in soil. Ultrasonication is commonly used to disrupt the soil structure prior to physical fractionation according to particle size, but may cause redistribution of OM among size fractions. The presence of mineral particles in size fractions can complicate estimations of OM turnover time within the fractions. Densiometric separation allows one to physically separate OM found within a specific size class from the heavier-density mineral particles. Nutrient contents and mineralization potential were determined for discrete size/density OM fractions isolated from within the macroaggregate structure of cultivated grassland soils. Eighteen percent of the total soil C and 25% of the total soil N in no-till soil was associated with fine-silt size particles having a density of 2.07-2.21 g/cm3 isolated from inside macroaggregates (enriched labile fraction or ELF). The amount of C and N sequestered in the ELF fraction decreased as the intensity of tillage increased. The specific rate of mineralization (mug net mineral N/mug total N in the fraction) for macroaggregate-derived ELF was not different for the three tillage treatments but was greater than for intact macroaggregates. The methods described here have improved our ability to quantitatively estimate SOM fractions, which in turn has increased our understanding of SOM dynamics in cultivated grassland systems.

Characterization, structure and function of linker polypeptides in phycobilisomes of cyanobacteria and red algae: An overview

Cyanobacteria and red algae have intricate light-harvesting systems comprised of phycobilisomes that are attached to the outer side of the thylakoid membrane. The phycobilisomes absorb light in the wavelength range of 500-650 nm and transfer energy to the chlorophyll for photosynthesis. Phycobilisomes, which biochemically consist of phycobiliproteins and linker polypeptides, are particularly wonderful subjects for the detailed analysis of structure and function due to their spectral properties and their various components affected by growth conditions. The linker potypeptides are believed to mediate both the assembly of phycobiliproteins into the highly ordered arrays in the phycobilisomes and the interactions between the phycobilisomes and the thylakoid membrane. Functionally, they have been reported to improve energy migration by regulating the spectral characteristics of colored phycobiliproteins. In this review, the progress regarding linker polypeptides research, including separation approaches, structures and interactions with phycobiliproteins, as well as their functions in the phycobilisomes, is presented. In addition, some problems with previous work on linkers are also discussed. (c) 2005 Elsevier B.V. All rights reserved.

Dynamic fluid–structure interaction using finite volume unstructured mesh procedures

A three-dimensional finite volume, unstructured mesh (FV-UM) method for dynamic fluid–structure interaction (DFSI) is described. Fluid structure interaction, as applied to flexible structures, has wide application in diverse areas such as flutter in aircraft, wind response of buildings, flows in elastic pipes and blood vessels. It involves the coupling of fluid flow and structural mechanics, two fields that are conventionally modelled using two dissimilar methods, thus a single comprehensive computational model of both phenomena is a considerable challenge. Until recently work in this area focused on one phenomenon and represented the behaviour of the other more simply. More recently, strategies for solving the full coupling between the fluid and solid mechanics behaviour have been developed. A key contribution has been made by Farhat et al. [Int. J. Numer. Meth. Fluids 21 (1995) 807] employing FV-UM methods for solving the Euler flow equations and a conventional finite element method for the elastic solid mechanics and the spring based mesh procedure of Batina [AIAA paper 0115, 1989] for mesh movement. In this paper, we describe an approach which broadly exploits the three field strategy described by Farhat for fluid flow, structural dynamics and mesh movement but, in the context of DFSI, contains a number of novel features: • a single mesh covering the entire domain, • a Navier–Stokes flow, • a single FV-UM discretisation approach for both the flow and solid mechanics procedures, • an implicit predictor–corrector version of the Newmark algorithm, • a single code embedding the whole strategy.

Dynamic fluid-structure interactions using finite volume unstructured mesh procedures

A three-dimensional finite volume, unstructured mesh (FV-UM) method for dynamic fluid–structure interaction (DFSI) is described. Fluid structure interaction, as applied to flexible structures, has wide application in diverse areas such as flutter in aircraft, wind response of buildings, flows in elastic pipes and blood vessels. It involves the coupling of fluid flow and structural mechanics, two fields that are conventionally modelled using two dissimilar methods, thus a single comprehensive computational model of both phenomena is a considerable challenge. Until recently work in this area focused on one phenomenon and represented the behaviour of the other more simply. More recently, strategies for solving the full coupling between the fluid and solid mechanics behaviour have been developed. A key contribution has been made by Farhat et al. [Int. J. Numer. Meth. Fluids 21 (1995) 807] employing FV-UM methods for solving the Euler flow equations and a conventional finite element method for the elastic solid mechanics and the spring based mesh procedure of Batina [AIAA paper 0115, 1989] for mesh movement. In this paper, we describe an approach which broadly exploits the three field strategy described by Farhat for fluid flow, structural dynamics and mesh movement but, in the context of DFSI, contains a number of novel features: a single mesh covering the entire domain, a Navier–Stokes flow, a single FV-UM discretisation approach for both the flow and solid mechanics procedures, an implicit predictor–corrector version of the Newmark algorithm, a single code embedding the whole strategy.

Numerical simulations of fluid-structure interactions in single-reed mouthpieces

Most single-reed woodwind instrument models rely on a quasistationary approximation to describe the relationship between the volume flow and. the pressure difference across the reed channel. Semiempirical models based on the quasistationary approximation are very useful in explaining the fundamental characteristics of this family of instruments such as self-sustained oscillations and threshold of blowing pressure. However, they fail at explaining more complex phenomena associated with the fluid-structure interaction during dynamic flow regimes, such as the transient and steady-state behavior of the system as a function. of the mouthpiece geometry. Previous studies have discussed the accuracy of the quasistationary approximation but the amount of literature on the subject is sparse, mainly due to the difficulties involved in the measurement of dynamic flows in channels with an oscillating reed. In this paper, a numerical technique based on the lattice Boltzmann method and a finite difference scheme is proposed in order to investigate the characteristics of fully coupled fluid-structure interaction in single-reed mouthpieces with different channel configurations. Results obtained for a stationary simulation with a static reed agree very well with those predicted by the literature based on the quasistationary approximation. However, simulations carried out for a dynamic regime with dn oscillating reed show that the phenomenon associated with flow detachment and reattachment diverges considerably frorn the theoretical assumptions. Furthermore, in the case of long reed channels, the results obtained for the vena contracta factor are in significant disagreement with those predicted by theory. For short channels, the assumption of constant vena contracta was found to be valid for only 40% of the duty cycle. (c) 2007 Acoustical Society of America.

Just a matter of time: Fungi and roots significantly and rapidly aggregate soil over four decades along the Tagliamento River, NE Italy

Fluvial islands are emergent landforms which form at the interface between the permanently inundated areas of the river channel and the more stable areas of the floodplain as a result of interactions between physical river processes, wood and riparian vegetation. These highly dynamical systems are ideal to study soil structure development in the short to medium term, a process in which soil biota and plants play a substantial role. We investigated soil structure development on islands along a 40 year chronosequence within a 3 km island-braided reach of the Tagliamento River, Northeastern Italy. We used several parameters to capture different aspects of the soil structure, and measured biotic (e.g., fungal and plant root parameters) and abiotic (e.g. organic carbon) factors expected to determine the structure. We estimated models relating soil structure to its determinants, and, in order to confer statistical robustness to our results, we explicitly took into account spatial autocorrelation, which is present due to the space for time substitution inherent in the study of chronosequences and may have confounded results of previous studies. We found that, despite the eroding forces from the hydrological and geomorphological dynamics to which the system is subject, all soil structure variables significantly, and in some case greatly increased with site age. We interpret this as a macroscopic proxy for the major direct and indirect binding effects exerted by root variables and extraradical hyphae of arbuscular mycorrhizal fungi (AMF). Key soil structure parameters such as percentage of water stable aggregates (WSA) can double from the time the island landform is initiated (mean WSA = 30%) to the full 40 years (mean WSA = 64%) covered by our chronosequence. The study demonstrates the fundamental role of soil biota and plant roots in aggregating soils even in a system in which intense short to medium term physical disturbances are common.

Environmental filtering vs. resource-based niche partitioning in diverse soil animal assemblages

Terrestrial invertebrates constitute most of described animal biodiversity and soil is a major reservoir of this diversity. In the classical attempt to understand the processes supporting biodiversity, ecologists are currently seeking to unravel the differential roles of environmental filtering and competition for resources in niche partitioning processes: these processes are in principle distinct although they may act simultaneously, interact at multiple spatial and temporal scales, and are often confounded in studies of soil communities. We used a novel combination of methods based on stable isotopes and trait analysis to resolve these processes in diverse oribatid mite assemblages at spatial
scales at which competition for resources could in principle be a major driver. We also used a null model approach based on a general neutral model of beta diversity. A large and significant fraction of community variation was explainable in terms of linear and periodic spatial structures in the distribution of organic C, N and soil structure: species were clearly arranged along an environmental, spatially structured gradient. However, competition related trait differences did not map onto the distances separating species along the environmental gradient and neutral models provided a satisfying approximation of beta diversity patterns. The results represent the first robust evidence
that in very diverse soil arthropod assemblages resource-based niche partitioning plays a minor role while environmental filtering remains a fundamental driver of species distribution.