Development of soil microbial communities during tallgrass prairie restoration

Soil microbial communities were examined in a chronosequence of four different land-use treatments at the Konza Prairie Biological Station, Kansas. The time series comprised a conventionally tilled cropland (CTC) developed on former prairie soils, two restored grasslands that were initiated on former agricultural soils in 1998 (RG₉₈) and 1978 (RG₇₈), and an annually burned native tallgrass prairie (BNP), all on similar soil types. In addition, an unburned native tallgrass prairie (UNP) and another grassland restored in 2000 (RG₀₀) on a different soil type were studied to examine the effect of long-term fire exclusion vs. annual burning in native prairie and the influence of soil type on soil microbial communities in restored grasslands. Both 16S rRNA gene clone libraries and phospholipid fatty acid analyses indicated that the structure and composition of bacterial communities in the CTC soil were significantly different from those in prairie soils. Within the time series, soil physicochemical characteristics changed monotonically. However, changes in the microbial communities were not monotonic, and a transitional bacterial community formed during restoration that differed from communities in either the highly disturbed cropland or the undisturbed original prairie. The microbial communities of RG₉₈ and RG₀₀ grasslands were also significantly different even though they were restored at approximately the same time and were managed similarly; a result attributable to the differences in soil type and associated soil chemistry such as pH and Ca. Burning and seasonal effects on soil microbial communities were small. Similarly, changing plot size from 300 m² to 150 m² in area caused small differences in the estimates of microbial community structure. In conclusion, microbial community structure and biochemical properties of soil from the tallgrass prairie were strongly impacted by cultivation, and the microbial community was not fully restored even after 30 years.     

Effects of white-tailed deer on the native earthworm,Eisenoides carolinensis,in the southern Appalachian Mountains,USA

Research on earthworms in North America has focused on the effects of invasive earthworms, with few studies examining the ecology of native earthworm species. Deer have been shown to influence belowground processes through grazing, trampling, and fecal pellet deposition. We proposed that native earthworms in an oak-dominated forest in Virginia might benefit from increased organic matter provided by deer fecal material. We examined potential interactions between a common aboveground herbivore, the white-tailed deer (Odocoileus virginianus), and earthworms using laboratory and field experiments. In our laboratory experiment, we found that a native earthworm, Eisenoides carolinensis, and an invasive earthworm, Lumbricus terrestris both fared better in treatments with deer pellets compared with the treatment with leaf litter alone. In our field experiment, we used fences to exclude deer from six plots and left twelve plots unfenced to explore the effects of deer activity on earthworm biomass and density. We also examined the effects of deer on soil and vegetation characteristics. After three years, the amount of herbaceous cover was higher on fenced plots compared with unfenced plots. Although we found no other differences for vegetation and soil characteristics between fenced and unfenced plots, many of these variables were important as covariates in our models examining the effect of deer exclusion on earthworms, indicating plot-level (as opposed to treatment-level) variation in these variables. All identifiable earthworms were either E. carolinensis or Diplocardia spp. (both native species), with E. carolinensis making up 90% of the specimens. The total biomass of earthworms, as well as the biomass and density of adult and small juvenile earthworms, was greater on unfenced plots with deer activity compared with fenced plots. This study highlights the importance of above- and below-ground interactions in forest ecosystems by showing that E. carolinensis appears to benefit from the presence of deer and adds to our sparse knowledge of the ecology of this native earthworm.     

Different behavioral patterns of the earthworms Octolasion tyrtaeum and Diplocardia spp. in tallgrass prairie soils: potential influences on plant growth

This study addressed differences between Diplocardia spp. (a native North American earthworm) and Octolasion tyrtaeum (an introduced European species), with respect to behavior, influence on soil microbial biomass, and plant uptake of N in tallgrass prairie soils. We manipulated earthworms in PVC-encased soil cores (20 cm diameter) over a 45-day period under field conditions. Treatments included: (1) control with no earthworms, (2) Diplocardia spp. only, and (3) O. tyrtaeum only. Prior to addition of earthworms, seedlings of Andropogon gerardii (a dominant tallgrass) were established in each core, and a dilute solution of ¹³C-labeled glucose and ¹⁵N-labeled (NH₄)₂SO₄ was added to the soil to facilitate examination of earthworm/microbe/plant interactions. We found that Diplocardia spp. were significantly more active than O. tyrtaeum, and quickly assimilated ¹³C and ¹⁵N from the tracer. Individuals of Diplocardia spp. were present at shallower soil depths than O. tyrtaeum throughout the study. Contrary to expectation, this greater activity of Diplocardia spp. did not result in increased plant productivity. Rather, the activity of Diplocardia spp. was associated with less plant growth and smaller amounts of N acquired by A. gerardii seedlings compared to controls or O. tyrtaeum treatments. We observed few significant influences of earthworm treatments on microbial biomass C or N pool sizes, but the microbial C/N ratio was consistently greater in the presence of Diplocardia spp. relative to O. tyrtaeum. Results of this study indicate that activity of earthworms may enhance competition for N between microbes and plants during the growing season in tallgrass prairie.     

Cattle grazing increases microbial biomass and alters soil nematode communities in subtropical pastures

This study focused on examining the impacts of cattle grazing on belowground communities and soil processes in humid grasslands. Multiple components in the soil communities were examined in heavily grazed and ungrazed areas of unimproved and improved bahiagrass (Paspalum notatum Flugge) pastures in south-central Florida. By using small (1-m×1-m) sampling plots, we were able to detect critical differences in nematode communities, microbial biomass, and mineralized C and N, resulting from the patchy grazing pattern of cattle. Soil samples were collected on three occasions between June 2002 and June 2003. Microbial C and N were greater (P?0.01) in grazed than in ungrazed plots on all sampling dates. Effects of grazing varied among nematode genera. Most genera of colonizer bacterivores were decreased (P?0.10) by grazing, but more persistent bacterivores such as Euteratocephalus and Prismatolaimus were increased, as were omnivores and predators. Higher numbers of persisters indicated that grazing resulted in a more structured nematode community. Some herbivores, particularly Criconematidae, were decreased by grazing. Abundance of omnivores, predators, and especially fungivores were strongly associated with C mineralization potential. Strong correlation of microbial C and N with nematode canonical variables composed of five trophic groups illustrates important links between nematode community structure and soil microbial resources. Including the analysis of nematode trophic groups with soil microbial responses reveals detection of grazing impact deeper into the hierarchy of the decomposition process in soil, and illustrates the complexity of responses to grazing in the soil foodweb. Although highly sensitive to grazing impacts, small-scale sampling could not be used to generalize the overall impact of cattle grazing in large-scale pastures, which might be determined by the intensity and grazing patterns of various stocking densities at the whole pasture level.     

Can the invasive earthworm,Amynthas agrestis,be controlled with prescribed fire?

Biological invasions are one of the most significant global-scale problems caused by human activities. Earthworms function as ecosystem engineers in soil ecosystems because their feeding and burrowing activities fundamentally change the physical and biological characteristics of the soils they inhabit. As a result of this “engineering,” earthworm invasions can have significant effects on soil physical, chemical and biological properties. The species Amynthas agrestis (family Megascolecidae) was introduced to the United States from Asia, and has expanded its distribution range to include relatively undisturbed forests. Here, to clarify life history traits, we reared individuals under seven different conditions of food provision using litter, fragmented litter and soil, and also analyzed the stable isotope ratios of field-collected specimens to investigate their food resources in the field. Second, we examined whether prescribed fire can be used to manage invasive earthworms. We constructed eight experimental plots, each with 100 individuals of A. agrestis each, and burned half of the plots. The feeding experiment showed that the earthworms in units containing soil and some form of organic matter (litter and/or fragmented litter) produced many cocoons, indicating that litter and fragmented litter are important food resources for them. Stable isotope analyses also supported this result. During the experimental fires, average soil temperature at 5 cm depth increased by only 7.7 °C (average maximum of 32.2 °C). Litter mass was significantly reduced by the fires. Although numbers of A. agrestis and cocoons recovered from burned and unburned plots were not different, the viability of cocoons was significantly lower in burned plots. Fire may also reduce the survival rate of juveniles in the next year by depriving them of their preferred food resource. Most native earthworms in the United States live in the soil, while many invasive ones live in the litter layer and soil surface. Therefore, prescribed fire could be a viable tool for control of invasive earthworms without negatively impacting native earthworm populations.     

Nitrogen enrichment causes minimal changes in arbuscular mycorrhizal colonization but shifts community composition—evidence from rDNA data

Intracellular arbuscular mycorrhizal (AM) colonization was compared between nitrogen (NH₄NO₃) fertilized (10 g N m^–2) and nonfertilized tallgrass prairie plots. In the microscopic analyses of host roots, only intracellular coils showed an increasing trend as a result of N fertilization, whereas intracellular colonization by arbuscules, hyphae, or vesicles did not differ between the N treatments. Clone libraries established from pooled PCR products of AM fungi contained exclusively species of Glomus; no other genera were detected indicating that Glomus spp. dominated the host roots. Comparisons between observed and random topologies indicated that cloned sequence placement covaried with N treatment: unique clades within Glomus originated exclusively from N-fertilized or nonfertilized treatments. We conclude that the communities of dominant and most commonly occurring AM fungi changed in response to N amendment, although the root colonization showed minimal or no response.     

Heathland management in the New Forest,Hampshire, England

Heathland in the New Forest, Hampshire, England, is subject to systematic controlled burning, the main intention of which is to maintain and enhance the quality of the grazing for commoners' animals. Mature heathland vegetation is richer in bird, reptile and invertebrate life than younger age classes. However, a survey has confirmed that despite a recent reduction in the area burnt annually, burning had produced an imbalance in favour of young age classes of heather Calluna vulgaris and Erica spp., and that intensive browsing after fire was severely inhibiting the regeneration of gorse Ulex europaeus. The data obtained made it possible to recommend management policies which attempted to correct the imbalance in vegetation age structure and also satisfied the requirements of the grazier.     

14C Cycling in lignocellulose-amended soils: predicting long-term C fate from short-term indicators

We compared lignocellulose (the second most abundant component of plant material) degradation over 8 months in contrasting soils from each of five sites across the United States with the aim of assessing which soils are likely to store more C. The soils were collected from a tallgrass prairie restoration (farmland, and plots restored in 1993 and 1979), the semiarid shrub-steppe (cool, moist upper slope and warm, dry lower slope soils), long-term farmland (no-till and conventional-till), and from two forest soils (loblolly pine and Douglas fir; fertilized and nonfertilized). Soils that rapidly metabolized freshly added C exploited endogenous and newly transformed C to a lesser degree over the course of the incubation (lower slope shrub-steppe, nonfertilized Douglas fir, and tallgrass prairie farmed and 1993 restorations). We also pooled the data to find a strong relationship between sand content and lignocellulose C remaining in the soil after 8 months (R=0.68) and also between short-term storage of lignocellulose C (at 7 days) and lignocellulose C remaining after 8 months (R=0.94). To predict C storage, models of C and soil properties must be modified to reflect the structure and function of microbial communities. Communities in richer soils may be more competent to use native C following fresh C additions when compared with communities in poorer soils.     

Influence of reduced tillage on earthworm and microbial communities under organic arable farming

Although reduced tillage is an agricultural practice reported to decrease soil erosion and external inputs while enhancing soil fertility, it has still rarely been adopted by European organic farmers. The objective of this study was to assess the long-term interactive effects of tillage (conventional (CT) vs. reduced (RT)) and fertilization (slurry (S) vs. composted manure/slurry (MCS)) on earthworms and microbial communities in a clay soil under spelt in an organic 6-year crop rotation. Earthworm populations (species, density and biomass, cocoons) were investigated by handsorting the soil nine years after initial implementation of the treatments. Soil microbial carbon (C_mic) and nitrogen (N_mic) were measured by chloroform-fumigation extraction and a simplified phospholipid fatty acid (PLFA) analysis was used to separate for populations of bacteria, fungi and protozoa. Significantly increased total earthworm density in RT plots was mainly attributed to increased numbers of juveniles. Moreover, we found five times more cocoons with RT. Species richness was not affected by the treatments, but tillage treatments had differentially affected populations at the species-level. In addition, cluster analysis at the community level revealed two distinct groups of plots in relation to tillage treatments. In RT plots C_mic increased in the 0–10 cm and 10–20 cm soil layers, while PLFA concentrations indicative of Gram-negative bacteria, fungi and protozoa only increased in the topsoil. Lower bacteria-to-fungi ratios in the upper soil layer of RT plots indicated a shift to fungal-based decomposition of organic matter whereas a higher C_mic-to-C_org ratio pointed towards enhanced substrate availability. Slurry application decreased microbial biomass and enhanced density of juvenile anecic earthworms but overall fertilization effect was weak and no interactions with tillage were found. In conclusion, tillage is a major driver in altering communities of earthworms and microorganisms in arable soils. The use of reduced tillage provides an approach for eco-intensification by enhancing inherent soil biota functions under organic arable farming.     

Long-term land-use effects on soil invertebrate communities in Southern Piedmont soils,USA

Historically, a large percentage of land area in the Piedmont of the southeastern USA was under intensive agricultural management for the production of cotton. This intensive farming resulted in massive erosion, and general degradation of soil resources until insect pests and poor economic conditions forced large-scale abandonment of farmland around the 1930s. In subsequent decades, there have been four predominant land-uses in the region, and we sampled soil macroinvertebrates from three replicate sites of cultivated fields, grass-dominated fields, loblolly pine stands, and remnant hardwood stands for a period of 2 years, with the objective of examining soil invertebrate community composition in relation to these long-term land-uses. At each site we dug three or four soil pits that were 30 × 30 cm to a depth of 15 cm, and sorted the soil volume by hand for a time not more than 1 person h, collecting all invertebrates ~5 mm in length or larger. We recorded abundance data for all invertebrate taxa collected, and we calculated community indices including diversity, evenness, rank abundance and percent similarity in order to identify patterns of community assemblage within each land-use type. Results suggest that soils in hardwood stands support the most taxonomically diverse macroinvertebrate communities followed by pine stands, pastures, and cultivated fields in order of decreasing diversity. For earthworms, Diplocardia spp. (North American megascolecids) were most abundant in the hardwood stands, but sometimes made up a substantial fraction of the community in other land-uses; whereas lumbricid earthworms (primarily introduced Apporectodea spp.) were most abundant in the cultivated and pasture soils, or showed no consistent habitat preference (native Bimastos spp.). Scarab beetles (larvae and adults) were common in all four systems, but reached the highest densities in cultivated and grass sites. Carabid beetle larvae were collected most often from cultivated soils. Several taxa were collected either exclusively or predominantly from forested sites, including diplopods, chilopods, gastropods, and several taxa of Diptera. These results indicate that long-term soil disturbance and the attendant differences in vegetation structure have profoundly influenced the community composition of invertebrates in Southern Piedmont soils, and that more intense disturbance results in a less diverse invertebrate community composed of a few, frequently non-native, disturbance-tolerant taxa.     

季节性放牧和全年放牧对亚高山草地生态系统中土壤动物和微生物的影响

Grazing and over-grazing may drive changes in the diversity and functioning of below-ground meadow ecosystems.A field soil survey was conducted to compare microbial biomass carbon (Cmin) and soil fauna communities in the two main grassland management systems in subalpine regions of Yunnan Province,China:perennial grazing currently practiced due to increasing herd sizes and traditional seasonal grazing.A three-year exclosure experiment was then conducted to further compare the effects of different grazing practices,including treatments of no mowing,perennial grazing (NM + G),mowing followed by seasonal grazing (M + G),mowing and no grazing (M + NG),and no mowing or grazing (NM + NG).The comparative survey result revealed that Cmin and total density of soil fauna were significantly lower at a perennially grazed site than at a seasonally grazed site.The experiment results showed that in comparison to non-grazing treatments (M + NG and NM + NG),grazing (NM + G and M + G) reduced total fauna density (by 150 individuals m-2) and the number of taxonomic groups present (by 0.32 taxa m-2).Mowing decreased Cmin (by 0.31 mg g-1).Furthermore,the NM + G treatment (perennial grazing) had the lowest density of Collembola (16.24 individuals m-2),one of the two most common taxonomic groups,although other taxonomic groups responded differently to the treatments.Treatment effects on soil fauna were consistent with those on above-ground grasses,in which C:N ratios were greatly reduced by grazing,with this effect being the greatest for the NM + G treatment.In contrast,different grazing treatments had little effect on C:N ratio of soil.Furthermore,the traditional grazing method (mowing followed by seasonal grazing) may have less severe effects on some taxonomic groups than perennial grazing.Therefore,an appropriate management should aim to protect soil fauna and microbes in this area from over-grazing and against further degradation.     

Potential impacts of elevated CO2 and above- and belowground litter quality of a tallgrass prairie

Increased atmospheric CO₂ will likely impact the productivity of arid and semiarid ecosystems through increased C, N, and water use efficiencies at the individual plant level. Tallgrass prairie has had increased above- and belowground biomass production under elevated CO₂, primarily due to increased water use efficiency. There is an apparent decreased N requirement to sustain increased productivity in CO₂-enriched tallgrass prairie, and C∶N ratios of plant litter above and below ground have increased. The tallgrass prairie ecosystem level response to elevated CO₂ on the C cycle could potentially increase C storage. Reduced litter quality associated with elevated CO₂ in tallgrass prairie has the potential to reduce decomposition rates, and ruminant digestion rate of plant biomass apparently has been lowered. Reduced intake by ruminants would shunt more of the plant biomass directly into the detrital food chain, thereby slowing decomposition further. The potential impact is for increased C to be retained as soil organic matter in the tallgrass prairie.     

Earthworms influence metal transfer from soil to snails

Most of the studies focusing on metal transfer from soil to biota ignore the possible non-trophic influence of an organism on pollutant transfer to other species. We hypothesised that an earthworm (Aporrectodea tuberculata) might modify the bioavailability of metals in soil and thus, their transfer to the snail Helix aspersa. Snails were exposed for 2 weeks to a multicontaminated field soil with or without earthworms, under controlled conditions. When exposed with earthworms, snails had higher concentrations of Cd, Cu and Zn than when they were exposed alone, while no difference was detected for Pb. For Cd only, the difference in snail bioaccumulation corresponded to an increase in its water-soluble fraction. Internal concentrations of metals in earthworms remained similar in the presence or absence of snails. Two non-exclusive possible mechanisms, including variations in bioavailable fractions and/or total accessible pools of metals, are proposed to explain how earthworms could modulate the transfer of metals from soil to snails. This work demonstrated that metal transfer from soil to one invertebrate species was influenced by another invertebrate. We conclude that the concept of intermediary species, usually used to describe interactions among species, should be extended to the interactions between biota and pollutants in non-biotic compartments.     

Livestock grazing activities and wild boar rooting affect alpine earthworm communities in the Central Pyrenees (Spain)

In alpine areas, shifts in traditional grazing activities are globally affecting ecosystem properties and rural livelihoods. The ongoing decrease in extensive husbandry, with a decline in sheep numbers and a relative increase in cattle stocking rates, has resulted in the abandonment of large alpine grazing areas. This pastoral change has been recently associated with increased disturbances of wild boar (Sus scrofa), mainly within cattle-stocked ranges. In turn, cattle areas favor earthworm communities, a preferred trophic resource for wild boars in mountain environments. However, it is unknown whether wild boar disturbances, together with grazing activities, can affect earthworm communities. Our aim is to analyze the abundance, richness and ecological categories of earthworms and soil parameters (soil C and N concentrations, moisture, and C:N ratio) in relation to the occurrence of wild boar disturbances and grazing activities at different stocking pressures. We sampled two different grazing scenarios differing in the distribution of cattle along a grazing gradient, which was represented by three levels of stocking pressure (high, intermediate and low). Our results showed a complex effect of grazing activities and disturbances on the abundance and richness of earthworms, along with variations in C:N ratio and soil moisture, especially with increasing cattle presence. At high-stocking pressures differences in earthworm abundance and richness between disturbed and undisturbed areas were limited, whereas at intermediate-stocking pressures earthworms were favored by wild boar disturbances. Ecological categories of earthworms responded differently; endogeic species were the most affected by grazing pressures and wild boar rooting, with highest occurrence at high-stocking pressures and within boar disturbed areas. In sum, pastoral use and soil disturbances affected earthworm community structure and composition in complex ways. These results indicate an interaction of processes that is relevant to understand current changes in alpine ecosystems.     

Peat amendment and production of different crop plants affect earthworm populations in field soil

A field experiment was conducted to study the effects of peat amendment and crop production system on earthworms. The experiment was established on a field previously cultivated with oats and with silt as the main soil type. Perennial crops strawberry, timothy and caraway, and annual crops rye, turnip rape, buckwheat, onion and fiddleneck were cultivated with conventional methods. All the crops were grown with and without soil amendment with peat. Earthworms were sampled twice: 4 and 28 months after establishment of the experiment. In the former case part of the experimental plots were soil sampled and hand sorted for estimation of earthworms. In the latter case all experimental plots were sampled and both soil sampling and mustard extraction was carried out. Soil organic carbon and microbial biomass was measured at 14 and 28 months. Peat increased the abundance of juvenile Aporrectodea caliginosa by 74% in three growing seasons, but had no effect on adult numbers. Lumbricus terrestris numbers were not increased by peat treatment. Three season cultivation of caraway favoured both A. caliginosa and L. terrestris. An equal abundance of A. caliginosa was also found in plots cultivated with turnip rape and fiddleneck. Total earthworm and especially A. caliginosa numbers were very small in plastic-mulched strawberry beds. This was mainly attributed to repeated use of the insecticide endosulfan. With the strawberry plots omitted there was a significant correlation between soil microbial N measured at 14 months and juvenile Aporrectodea spp. and Lumbricus spp. numbers measured at 28 months. Adult earthworm numbers were not associated with either soil organic C or microbial biomass.     

Relationships between microbial community structure and soil environmental conditions in a recently burned system

Most wildfires, even the most severe, burn at mixed intensities across a landscape, depending on local fuel loads, fuel moistures, and wind strength and direction. This heterogeneous patchwork of fire effects can influence the patterns of above- and belowground biotic recovery through altered environmental conditions, nutrient availability, and biotic sources for microbial and vegetative re-colonization. We quantified the effects of low- and high-severity fire 14 months post-burn on key environmental variables typically limiting to microbial activity. We characterized the soil microbial community structure through ester-linked fatty acid analysis (EL-FAME) and identified the soil environmental factors that best explain the pattern of microbial community profiles through canonical correspondence analysis (CCA). Low-severity burning caused no change in soil moisture, pH or temperature while high-severity burning caused an increase in soil moisture, temperature, and a decrease in pH levels, relative to the unburned sites. Soil respiration rates were significantly lower in both the low- and high-severity burn sites, relative to unburned sites, likely due to initial root and microbial death. Overall microbial biomass did not change with either low- or high-severity burning, but the microbial community ordination biplots showed separation of communities by fire, and slight separation by fire severity along three axes. This community separation was driven primarily by a decrease in fungal biomarkers (18:2ω6c, 18:3ω6c) with both low- and high-severity fire. Only 23% of the variation in the microbial community distribution could be explained by three environmental variables: soil pH, temperature, and carbon. These results suggest that the microbial communities in both the low- and high-severity burn sites are structurally different from the populations in the unburned sites.     

Detritivorous earthworms modify microbial community structure and accelerate plant residue decomposition

The impact of detritivorous earthworms (Eisenia andrei) on microbial community structure and function in grape marc, a lignocellulosic enriched plant residue, was investigated in a mesocosm experiment. Analysis of carbon and nitrogen pools was also carried out in order to evaluate how changes in microbial communities affect plant residue decomposition. The grape marc was completely processed after fifteen days as a result of the high density of earthworms present and the rapid gut transit time. Eisenia andrei had a large impact on the structure of the microbial community, as revealed by phospholipid fatty acid analysis. Earthworm activity reduced the abundance of both bacterial (except for Gram-negative bacteria) and fungal PLFA biomarkers relative to the control values. Decreases in microbial activity and in protease and cellulase activities were also attributable to the presence of earthworms. Moreover, earthworms strongly modified the substrate utilization patterns of microbial communities, as revealed by BIOLOG analysis. The presence of earthworms led to an increase in the utilization of some amino acids and polymers, which reached a higher substrate diversity value than that in the control mesocosm. The differences in microbial communities were accompanied by a reduction in the total C content and the labile C pool, relative to the control, although there were no significant differences in either cellulose or hemicellulose contents. However, total N content increased in both mesocosms – with and without earthworms – and the concentration of NH₄⁺ was also enhanced by earthworm activity. The results indicate that detritivorous earthworms play a key role in decomposing fresh plant residues in the short term via their intensive interactions with microbial communities.     

Beneficial effects of earthworms and arbuscular mycorrhizal fungi on establishment of leguminous trees on Pb/Zn mine tailings

Planting trees to stabilize metalliferous mine tailings is a widely used form of land reclamation although substantial soil amendment is invariably required, both to improve the physico-chemical status of the tailings and to ameliorate toxicity prior to planting. Here, we report a glasshouse study of the combined effects of burrowing earthworms (Pheretima guillelmi) and arbuscular mycorrhizal fungi (Glomus spp., AMF) on establishment of a naturally invasive, woody, nitrogen-fixing legume, Leucaena leucocephala, on topsoil-amended Pb/Zn mine tailings. AMF provided the most effective preliminary inoculant, improving N, P and K uptake, but earthworms had more influence improving N nutrition. In most cases, the combined effects of AMF and earthworms were additive and proved to be beneficial to plant growth, plant nutrition and for protection against uptake of toxic metals. AMF influenced metal uptake more than earthworms, but together they reduced mobility of Pb and Zn in soil by as much as 25%. Some minor but significant negative interactions were also evident; for example, earthworms enhanced soil microbial activity but inhibited the beneficial effects of AMF on N₂-fixation. We argue that increased attention to ecological interactions in soil could reduce costs and improve the efficacy of restoring a vegetation cover to land impacted by contaminated spoils.     

Changes in net N mineralization rates and soil N and P pools in a pine forest wildfire chronosequence

The concern that climate change may increase fire frequency and intensity has recently heightened the interest in the effects of wildfires on ecosystem functioning. Although short-term fire effects on forest soils are well known, less information can be found on the long-term effects of wildfires on soil fertility. Our objective was to study the 17-year effect of wildfires on forest net mineralization rates and extractable inorganic nitrogen (N) and phosphorus (P) concentrations. We hypothesize that (1) burned forest stands should exhibit lower net mineralization rates than unburned ones; (2) these differences would be greatest during the growing season; (3) differences between soil variables might also be observed among plots from different years since the last fire; and (4) due to fire-resistant geochemical processes controlling P availability, this nutrient should recover faster than N. We used a wildfire chronosequence of natural and unmanaged Pinus canariensis forests in La Palma Island (Canary Islands). Soil samples were collected during winter and spring at 22 burned and unburned plots. We found significantly higher values for net N mineralization and extractable N pools in unburned plots. These differences were higher for the winter sampling date than for the spring sampling date. Unlike extractable N and N mineralization rates, extractable P levels of burned plots exhibited a gradual recovery over time after an initial decrease. These results demonstrate that P. canariensis forest soils showed low resilience after wildfires, especially for N, and that this disturbance might induce long-term changes in ecosystem functioning.     

Competition between invasive earthworms (Amynthas corticis, Megascolecidae) and native North American millipedes (Pseudopolydesmus erasus, Polydesmidae): Effects on carbon cycling and soil structure

Invasive earthworms can have significant impacts on C dynamics through their feeding, burrowing, and casting activities, including the protection of C in microaggregates and alteration of soil respiration. European earthworm invasion is known to affect soil micro- and mesofauna, but little is known about impacts of invasive earthworms on other soil macrofauna. Asian earthworms (Amynthas spp.) are increasingly being reported in the southern Appalachian Mountains in southeastern North America. This region is home to a diverse assemblage of native millipedes, many of which share niches with earthworm species. This situation indicates potential for earthworm-millipede competition in areas subject to Amynthas invasion.In a laboratory microcosm experiment, we used two ¹³C enriched food sources (red oak, Quercus rubra, and eastern hemlock, Tsuga canadensis) to assess food preferences of millipedes (Pseudopolydesmus erasus), to determine the effects of millipedes and earthworms (Amynthas corticis) on soil structure, and to ascertain the nature and extent of the interactions between earthworms and millipedes. Millipedes consumed both litter species and preferred red oak litter over eastern hemlock litter. Mortality and growth of millipedes were not affected by earthworm presence during the course of the experiment, but millipedes assimilated much less litter-derived C when earthworms were present.Fauna and litter treatments had significant effects on soil respiration. Millipedes alone reduced CO₂ efflux from microcosms relative to no fauna controls, whereas earthworms alone and together with millipedes increased respiration, relative to the no fauna treatment. CO₂ derived from fresh litter was repressed by the presence of macrofauna. The presence of red oak litter increased CO₂ efflux considerably, compared to hemlock litter treatments.Millipedes, earthworms, and both together reduced particulate organic matter. Additionally, earthworms created significant shifts in soil aggregates from the 2000-250 and 250-53 μm fractions to the >2000 μm size class. Earthworm-induced soil aggregation was lessened in the 0-2 cm layer in the presence of millipedes. Earthworms translocated litter-derived C to soil throughout the microcosm.Our results suggest that invasion of ecosystems by A. corticis in the southern Appalachian Mountains is unlikely to be limited by litter species and these earthworms are likely to compete directly for food resources with native millipedes. Widespread invasion could cause a net loss of C due to increased respiration rates, but this may be offset by C protected in water-stable soil aggregates.