Contrasted effect of biochar and earthworms on rice growth and resource allocation in different soils

Adding biochar to soils and maintaining high earthworm biomasses are potential ways to increase the fertility of tropical soils and the sustainability of crop production in the spirit of agroecology and ecological engineering. However, a thorough functional assessment of biochar effect on plant growth and resource allocations is so far missing. Moreover, earthworms and biochar increase mineral nutrient availability through an increase in mineralization and nutrient retention respectively and are likely to interact through various other mechanisms. They could thus increase plant growth synergistically. This hypothesis was tested for rice in a greenhouse experiment. Besides, the relative effects of biochar and earthworms were compared in three different soil treatments (a nutrient rich soil, a nutrient poor soil, a nutrient poor soil supplemented with fertilization). Biochar and earthworm effects on rice growth and resource allocation highly depended on soil type and were generally additive (no synergy). In the rich soil, there were both clear positive biochar and earthworm effects, while there were generally only positive earthworm effects in the poor soil, and neither earthworm nor biochar effect in the poor soil with fertilization. The analysis of earthworm and biochar effects on different plant traits and soil mineral nitrogen content, confirmed that they act through an increase in nutrient availability. However it also suggested that another mechanism, such as the release in the soil of molecules recognized as phytohormones by plants, is also involved in earthworm action. This mechanism could for example help explaining how earthworms increase rice resource allocation to roots and influence the allocation to grains.     

Earthworm impacts on soil organic matter and fertilizer dynamics in tropical hillside agroecosystems of Honduras

Earthworms are important processors of soil organic matter (SOM) and nutrient turnover in terrestrial ecosystems. In agroecosystems, they are often seen as beneficial organisms to crop growth and are actively promoted by farmers and extension agents, yet their contribution to agroecosystem services is uncertain and depends largely on management. The Quesungual slash-and-mulch agroforestry system (QSMAS) of western Honduras has been proposed as a viable alternative to traditional slash-and-burn (SB) practices and has been shown to increase earthworm populations, yet the effect of earthworms on soil fertility and SOM in QSMAS is poorly understood. This study examined the role of Pontoscolex corethrurus in QSMAS by comparing their influence on aggregate-associated SOM and fertilizer dynamics with their effects under SB and secondary forest in a replicated field trial. Both the fertilized QSMAS and SB treatments had plots receiving additions of inorganic ¹⁵N and P, as well as plots with no inorganic N additions. Earthworm populations were manipulated in field microcosms at the beginning of the rainy season within each management treatment via additions of P. corethrurus or complete removal of existing earthworm populations. Microcosms were destructively sampled at harvest of Zea mays and soils were wet-sieved (using 53, 250 and 2000 μm mesh sizes) to isolate different aggregate size fractions, which were analyzed for total C, N and ¹⁵N. The effects of management system were smaller than expected, likely due to disturbance associated with the microcosm installation. Contrary to our hypothesis that earthworms would stabilize organic matter in soil aggregates, P. corethrurus decreased total soil C by 3% in the surface layer (0-15 cm), predominantly through a decrease in the C concentration of macroaggregates (>250 μm) and a corresponding depletion of C in coarse particulate organic matter occluded within macroaggregates. Earthworms also decreased bulk density by over 4%, but had no effect on aggregate size distribution. Within the two fertilized treatments, the QSMAS appeared to retain slightly more fertilizer derived N in smaller aggregate fractions (<250 μm) than did SB, while earthworms greatly reduced the recovery of fertilizer N (34% decrease) in both systems. Although management system did not appear to influence the impact of P. corethrurus on SOM or nutrient dynamics, we suggest the lack of differences may be due to artificially low inputs of fresh residue C to microcosms within all management treatments. Our findings highlight the potential for P. corethrurus to have deleterious impacts on soil C and fertilizer N dynamics, and emphasize the need to fully consider the activities of soil fauna when evaluating agroecosystem management options.     

Species effects on earthworm density in tropical tree plantations in Hawaii

Summary Tree species differ in the quantity and quality of litter produced, and these differences may significantly affect ecosystem structure and function. I examined the importance of tree species in determining earthworm densities in replicated stands of Eucalyptus saligna Sm. and Albizia falcataria (L.) Fosberg, and in mixed stands (25% albizia and 75% eucalyptus). Mean earthworm densities ranged from 92 m^-2 in the pure eucalyptus, to 281 m^-2 in the mixture, and a maximum of 469 m^-2 in the pure albizia stands. Only two earthworm species were present, Pontoscolex corethrurus and Amynthas gracilis. Leaf biomass on the forest floor was highest in the pure eucalyptus and lowest in the pure albizia stands, whereas the annual fine litterfall production was lowest in the pure eucalyptus and highest in the albizia stands. The N content of fine litterfall was correlated positively with earthworm density, and the fine litterfall biomass: N ratio was correlated negatively with earthworm density. Greater leaf biomass on the forest floor under eucalyptus stands despite lower rates of litterfall suggests that litter quality, rather than litter quantity, was primarily responsible for the greater earthworm density in the albizia stands. Some biogeochemical effects of tree species in the tropics may be mediated through effects on earthworm populations.     

Earthworm and microbe response to litter and soils of tropical forest plantations with contrasting C:N:P stoichiometric ratios

1

The stoichiometry of resources is increasingly acknowledged as a major control of consumer activity and abundance. Chemical properties of litter, the main source of food for decomposers, are likely to be important drivers of decomposer activity.

2

Theory predicts a high control of resource stoichiometry on the abundance of consumer organisms that maintain strict homeostasis, due to costs associated with the regulation of nutrient balance in their body tissue. Decomposer efforts in nutrient acquisition should be related to imbalances in resource stoichiometry.

3

A 21 year old experimental plantation of monospecific plots of trees with leaves of contrasting chemistry was used to test four hypotheses: (i) soil and litter nutrient stoichiometry (C, N, P) are linked; (ii); soil enzyme activity ratios and stoichiometry are linked; (iii) earthworms’ tissue stoichiometry does not depend on soil and litter stoichiometry (homeostasis); (iv) earthworm density is dependent upon phosphorus availability, the most limiting nutrient in soils at this site, and, to a lesser extent, to nitrogen availability.

4

We found (i) no relationship between litter and soil stoichiometry, (ii) microbial activity was linked to soil stoichiometry, (iii) earthworms showed strict homeostasis in their tissue and (iv) earthworm abundance increased with P availability.

5

We discuss the mechanisms that might lead to these patterns.

     

Nitrogen mineralization and reorganization in casts of the geophagous tropical earthworm Pontoscolex corethrurus (Glossoscolecidae)

Summary Mineral N concentrations ranged from 133.1 to 167.8 g g^-1 dry soil in fresh casts of the endogeic earthworm Pontoscolex corethrurus fed on an Amazonian Ultisol; this was approximately five times the concentration in non-ingested soil. Most of this N was in the form of NH _inf4 ^sup+ . N also accumulated in microbial biomass, which increased from a control value of 10.5–11.3 to 67.5–74.1 g g^-1 in fresh casts. During a 16-day incubation, part of the NH _inf4 ^sup+ -N was nitrified and/or transferred to the microbial biomass. Total labile N (i.e., mineral+biomas N) decreased sharply at first (ca. 50% in the first 12 h), and then more slowly. The exact fate of this N (microbial metabolites, denitrification, or volatilization) is not known. After 16 days, the overall N content of the casts was still 28% higher than that of the control soil. Incubation of the soil before ingestion by the earthworms significantly increased the production of NH _inf4 ^sup+ in casts. We calculate that in a humid tropical pasture, 50–100 kg mineral N may be produced annually in earthworm casts. Part of this N may be conserved in the compact structure of the cast where the cast is not in close contact with plant roots.     

Stable C and N isotope values of selected components of a tropical australian sugarcane ecosystem

The ¹³C and ¹⁵N values of sugarcane plant tissues, decomposing harvest residues, soil and the casts and body tissues of the earthwormPontoscolex corethrurus were determined. Little variation in ¹³C values was found between plant parts. The ¹³C values of the decomposing harvest residues declined and became more variable after 148 days of exposure in the field. In the decomposing residues, ¹³C values of the neutral detergent fibre fraction were similar to those of the whole tissues while those of the proximate lignin were more negative. The ¹⁵N values of the residues also declined over time after a short initial delay.P. corethrurus populations are more intimately associated with the roots of sugarcane than with the bulk soil. Tissue ¹³C values suggest that the earthworm diet is similar to or more enriched in¹³C than sugarcane tissues and is substantially more enriched than the soil C. Earthworm tissues have similar levels of¹⁵N enrichment to both the soil and plant tissues. These data are consistent with the hypothesis that this earthworm derives much of its assimilated C relatively directly from organic matter associated with the roots and decomposing harvest residues.     

Field inoculation effectiveness of native and exotic arbuscular mycorrhizal fungi in a Mediterranean agricultural soil

In sustainable agriculture, arbuscular mycorrhizal (AM) fungal inoculation in agronomical management might be very important, especially when the efficiency of native inocula is poor. Here, we assessed the effect of native and exotic selected AM fungal inocula on plant growth and nutrient uptake in a low input Trifolium alexandrinum-Zea mays crop rotation. We evaluated the effects of four exotic AM fungal isolates on T. alexandrinum physiological traits in greenhouse. Then, the field performances of T. alexandrinum inoculated with the exotic AMF, both single and mixed, were compared to those obtained with a native inoculum, using a multivariate analysis approach. Finally, we tested the residual effect of AM fungal field inoculation on maize as following crop. Multivariate analysis showed that the field AM fungal inoculation increased T. alexandrinum and Z. mays productivity and quality and that the native inoculum was as effective as, or more effective than, exotic AM fungal isolates. Moreover, the beneficial effects of AMF were persistent until the second year after inoculation. The use of native AMF, produced on farm with mycotrophic plants species, may represent a convenient alternative to commercial AM fungal inocula, and may offer economically and ecologically important advantages in sustainable or organic cropping systems.     

Earthworm activity affects soil aggregation and organic matter dynamics according to the quality and localization of crop residues—An experimental study (Madagascar)

Soil organic matter (SOM) plays a central role in the functioning of ecosystems, and is beneficial from agronomic and from environmental point of view. Alternative cultural systems, like direct seeding mulch-based cropping (DMC) systems, enhance carbon (C) sequestration in agricultural soils and lead to an increase in soil macrofauna. This study aimed at evaluating in field mesocosms the effects of earthworms on SOM dynamics and aggregation, as influenced by residue quality and management.In the highlands of Madagascar, buckets were filled with 2 mm-sieved clayey Inceptisol. The effects of earthworm addition (Pontoscolex corethrurus), residue addition (rice, soybean, and no addition), and localization of the residues (mulched or buried) were studied. After 5 months, soil from mesocosms with earthworms had significantly lower C concentration and higher proportion of large water-stable macroaggregates (>2000 μm) than those without earthworms, because of the production of large macroaggregates by earthworms. Earthworm effect on soil aggregation was greater with rice than with soybean residues. Casts (extracted from mesocosms with earthworms) were slightly enriched in C and showed significantly higher mineralization than the non-ingested soil (NIS), showing that at the time scale of our study, the carbon contained in the casts was not protected against mineralization. No difference in microbial biomass was found between casts and NIS.Complementary investigations are necessary to assess long-term effects of earthworm addition on SOM dynamics, the conditions of occurrence of physical protection, and the impact of earthworms on the structure of the microbial community.     

Long-term soil organic carbon dynamics in a subhumid tropical climate: C data in mixed C3/C4 cropping and modeling with ROTHC

Scanty information on long-term soil organic carbon (SOC) dynamics hampers validation of SOC models in the tropics. We observed SOC content changes in a 16-year continuously cropped agroforestry experiment in Ibadan, south-western Nigeria. SOC levels declined in all treatments. The decline was most pronounced in the no-tree control treatments with continuous maize and cowpea cropping, where SOC levels dropped from the initial 15.4 to 7.3-8.0 Mg C ha⁻¹ in the 0-12 cm topsoil in 16 years. In the two continuously cropped alley cropping (AC) systems, one with Leucaena leucocephala and one with Senna siamea trees, SOC levels dropped to 10.7-13.2 Mg C ha⁻¹. Compared to the no-tree control treatments, an annual application of an additional 8.5 Mg ha⁻¹ (dry matter) of plant residues, mainly tree prunings, led to an extra 3.5 Mg C ha⁻¹ (∼0.2% C) in the 0-12 cm top soil after 11 years, and 4.1 Mg C ha⁻¹ after 16 years. The addition of NPK fertilizer had little effect on the quantities of above-ground plant residues returned to the soil, and there was no evidence that the fertilizer affected the rate of SOC decomposition. The fact that both C₃ and C₄ plants returned organic matter to the soil in all cropping systems, but in contrasting proportions, led to clear contrasts in the ¹³C abundance in the SOC. This ¹³C information, together with the measured SOC contents, was used to test the ROTHC model. Decomposition was very fast, illustrated by the fact that we had to double all decomposition rate constants in the model in order to simulate the measured contrasts in SOC contents and δ¹³C between the AC treatments and the no-tree controls. We hypothesized (1) that the pruning materials from the legume trees and/or the extra rhizodeposition from the tree roots in the AC treatments accelerated the decomposition of the SOC present at the start of the experiment (true C-priming), and/or (2) that the physical protection of microbial biomass and metabolites by the clay fraction on this site, having a sandy top soil in which clay minerals are mainly of the 1:1 type, is lower than assumed by the model.     

Effects of an exotic prey species on a native specialist: Example of the snail kite

Despite acknowledging that exotic species can exhibit tremendous influence over native populations, few case studies have clearly demonstrated the effects of exotic prey species on native predators. We examined the effects of the recently introduced island apple snail (Pomacea insularum) on the foraging behavior and energetics of the endangered snail kite (Rostrhamus sociabilis plumbeus) in Florida. We conducted time-activity budgets: (i) on kites foraging for native Florida apple snails (Pomacea paludosa) in major wetland units within the kites’ range that had not been invaded by the exotic island apple snail and (ii) on kites foraging for exotic apple snails in Lake Tohopekaliga, the only major wetland utilized by the snail kite that had suffered a serious invasion of P. insularum. When foraging for P. insularum, snail kites dropped a greater proportion of snails, and they experienced increased handling times and decreased consumption rates; however, kites foraging for P. insularum also spent a smaller proportion of the day in flight. Estimates of net daily energy balances between kites feeding on P. insularum versus P. paludosa were comparable for adults, but juveniles experienced energetic deficiencies when feeding on the exotic snail. Due to this discrepancy, we hypothesize that wetlands invaded by P. insularum, such as Lake Tohopekaliga, may function as ecological traps for the snail kite in Florida by attracting breeding adults but simultaneously depressing juvenile survival. This study highlights the conservation implications and importance of elucidating the effects that exotic species have on native specialists, especially those that are endangered, because subtle influences on behavior may have significant population consequences.     

Habitat and flower resource partitioning by an exotic and three native bumble bees in central Hokkaido, Japan

Studies of the interspecific relationships between alien and native pollinator species can help forecast the success of alien species as well as assess the extent of disturbance to native plant-pollinator interactions. We examined the habitat and flower resource occupancy by the invasive introduced bumble bee Bombus terrestris and three dominant native bumble bees in central Hokkaido, northern Japan, in relation to a landscape factor (forest cover ratio) and flower morphology with respect to the proboscis length of bees. Three years of monitoring the invasive boundary of B. terrestris indicated that this species, which dominates open agricultural areas, probably will not invade the forests in which the native species dominate. This habitat partitioning likely followed the displacement of the natives by the invader in open agricultural lands. In forested areas, the native species partitioned flower resources on the basis of the relationship between proboscis length and the lengths of the corolla tubes of flowers. However, in open agricultural areas, both the long- and short-tubed flowers were primarily visited by the exotic short-tongued B. terrestris, which foraged illegitimately (by nectar robbing) on long-tubed flowers (Trifolium pratense L.) and legitimately on short-tubed flowers (T. repens L. and Lavandula angustifolia Mill.). The invasion of B. terrestris into open natural vegetation, in addition to open agricultural areas, has recently been reported in Hokkaido. Even though the exotic and dominant native bumble bees partition their habitat according to landscape factors, the invasive bee has the potential to alter the overall interactions within plant-pollinator systems in the regions of open vegetation on Hokkaido Island.     

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.     

Contribution of anecic earthworms to biopore formation during cultivation of perennial ley crops

Large sized biopores (diameter >2 mm) in the subsoil can be created by tap roots, which leave voids after their decay, or by the burrowing activity of anecic earthworms which may benefit from the temporary lack in tillage in perennial cropping systems. However, the interactions between root growth and earthworm activity in the process of biopore formation during perennial ley cropping are not well understood. The aim of this field study was to quantify the development of the abundance of the anecic earthworm Lumbricus terrestris and the biopore density during the cultivation of lucerne (Medicago sativa L.), chicory (Cichorium intybus L.) and tall fescue (Festuca arundinacea Schreb.) grown for either one, two or three years. An increased abundance of L. terrestris was already recorded after two years of continuous ley when compared with one year cultivation. The ley crop species had only minor influence on the abundance of L. terrestris. Biopore densities of both diameter classes under study (2–5 mm and >5 mm) were not significantly affected by the duration of ley cropping. In contrast, biopore densities were influenced by ley crop species. More biopores of diameter class 2–5 mm were recorded after chicory than after fescue. Lucerne cropping resulted in intermediate biopore density. Additionally, in an incubation experiment under field conditions, we quantified whether L. terrestris preferentially created new biopores or colonized abandoned, previously existing ones. After three weeks of incubation, one third of the adult individuals incubated in the experimental area created new biopores at 0.4 m soil depth. A similar percentage of individuals colonized previously existing biopores, partially widening the lumen of smaller sized biopores. The remaining individuals remained in the topsoil. Sub-adult individuals rarely formed new pores. Half of the introduced sub-adults remained in the topsoil. We conclude that in crop rotations new biopores can be generated during perennial ley cropping by taproot systems of ley crops, but that a two to three- year period without tillage is not sufficient for populations of anecic earthworms to make a marked contribution to biopore density in the subsoil. The relevance of anecic earthworms for altering physical and chemical properties of biopores during ley cropping still needs further investigation.     

Seasonal variation in trehalose contents of roots and nodules of leguminous trees in a tropical deciduous forest in Mexico

Seasonal variations of trehalose contents in roots and root-nodules of five legumes in a tropical deciduous forest in Jalisco, México, were determined. The tree species were: Lonchocarpus eriocarinalis and Erythrina lanata (sub-family Papilionoideae) and Piptadenia constricta, Albizia occidentalis and Lysiloma microphylum (sub-family Mimosoideae). Trehalose accumulation in nodules and roots varied seasonally and among species. For example, the Papilionoid-species retained nodules longer than those in the Mimosoideae (5 and 4 months, respectively), and accumulated the highest amounts of trehalose (average values of 178 vs. 2.88 mg g⁻¹ nodule (d. wt), respectively). Generally, maximum trehalose contents in nodules and roots were observed in November, at the beginning of the dry season.     

Plant invasion of native grassland on serpentine soils has no major effects upon selected physical and biological properties

Plant invasions alter soil microbial community composition; this study examined whether invasion-induced changes in the soil microbial community were reflected in soil aggregation, an ecosystem property strongly influenced by microorganisms. Soil aggregation is regulated by many biological factors including roots, arbuscular mycorrhizal fungal hyphae, and microbially-derived carbon compounds. We measured root biomass, fungal-derived glomalin-related soil protein (GRSP), and aggregate mean weight diameter in serpentine soils dominated by an invasive plant (Aegilops triuncialis (goatgrass) or Centaurea solstitialis (yellow starthistle)), or by native plants (Lasthenia californica and Plantago erecta, or Hemizonia congesta). Root biomass tended to increase in invaded soils. GRSP concentrations were lower in goatgrass-dominated soils than native soils. In contrast, starthistle dominated soil contained a higher amount of one fraction of GRSP, easily extractable immunoreactive soil protein (EE-IRSP) and a lower amount of another GRSP fraction, easily extractible Bradford reactive soil protein (EE-BRSP). Soil aggregation increased with goatgrass invasion, but did not increase with starthistle invasion. In highly aggregated serpentine soils, small increases in soil aggregation accompanying plant invasion were not related to changes in GRSP and likely have limited ecological significance.     

Impacts of earthworms and arbuscular mycorrhizal fungi (Glomus intraradices) on plant performance are not interrelated

Earthworms and arbuscular mycorrhizal fungi (AMF) might interactively impact plant productivity; however, previous studies reported inconsistent results. We set up a three-factorial greenhouse experiment to study the effects of earthworms (Aporrectodea caliginosa Savigny and Lumbricus terrestris L.) and AMF (Glomus intraradices N.C. Schenck & G.S. Sm.) on the performance (productivity and shoot nutrient content) of plant species (Lolium perenne L., Trifolium pratense L. and Plantago lanceolata L.) belonging to the three functional groups grasses, legumes and herbs, respectively. Further, we investigated earthworm performance and plant root mycorrhization as affected by the treatments. Our results accentuate the importance of root derived resources for earthworm performance since earthworm weight (A. caliginosa and L. terrestris) and survival (L. terrestris) were significantly lower in microcosms containing P. lanceolata than in those containing T. pratense. However, earthworm performance was not affected by AMF, and plant root mycorrhization was not modified by earthworms. Although AMF effectively competed with T. pratense for soil N (as indicated by δ¹⁵N analysis), AMF enhanced the productivity of T. pratense considerably by improving P availability. Remarkably, we found no evidence for interactive effects of earthworms and AMF on the performance of the plant species studied. This suggests that interactions between earthworms and AMF likely are of minor importance.     

Isotopic fractionation during N2 fixation by four tropical legumes

To quantify the contribution of biological nitrogen fixation (BNF) to legume crops using the ¹⁵N natural abundance technique, it is necessary to determine the ¹⁵N abundance of the N derived from BNF—the B value. In this study, we used a technique to determine B whereby both legume and non-N₂-fixing reference plants were grown under the same conditions in two similar soils, one artificially labelled with ¹⁵N, and the other not. The proportion of N derived from BNF (%Ndfa) was determined from the plants grown in the ¹⁵N-labelled soil and it was assumed that the %Ndfa values of the legumes grown in the two soils were the same, hence the B value of the legumes could be calculated. The legumes used were velvet bean (Mucuna pruriens), sunnhemp (Crotalaria juncea), groundnut (Arachis hypogaea) and soybean (Glycine max) inoculated, or not, with different strains of rhizobium. The values of %Ndfa were all over 89%, and all the legumes grown in unlabelled soil showed negative δ¹⁵N values even though the plant-available N in this soil was found to be approximately +6.0‰. The B values for the shoot tissue (B_s) were calculated and ranged from approximately −1.4‰ for inoculated sunnhemp and groundnut to −2.4 and −4.5‰ for soybean inoculated with Bradyrhizobium japonicum strain CPAC 7 and Bradyrhizobium elkanii strain 29W, respectively. The B (B_wp) values for the whole plants including roots, nodules and the original seed N were still significantly different between the soybean plants inoculated with CPAC 7 (−1.33‰) and 29W (−2.25‰). In a parallel experiment conducted in monoxenic culture using the same soybean variety and Bradyrhizobium strains, the plants accumulated less N from BNF and the values were less negative, but still significantly different for soybean inoculated with the two different Bradyrhizobium strains. The results suggest that the technique utilized in this study to determine B with legume plants grown in soil in the open air, yields B values that are more appropriate for use under field conditions.     

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.     

The effects of earthworms on the demography of annual plant assemblages in a long-term mesocosm experiment

Earthworms have been shown to influence plant growth, survival and fecundity. They can therefore affect plant demography in plant communities changing their composition. A long term mesocosm experiment was set-up to test the effects of an endogeic (Aporrectodea caliginosa) and an anecic (Lumbricus terrestris) earthworm species on assemblages of four species of annuals: one grass (Poa annua), two forbs (Veronica persica and Cerastium glomeratum) and one legume (Trifolium dubium). The number of individuals and the biomass of each species were investigated. A. caliginosa and L. terrestris affected the density of T. dubium at each of the three monitored census dates. The other plant species responded to A. caliginosa and L. terrestris at the second and third generations. The presences of A. caliginosa and L. terrestris reduced the total number of plant individuals from the second to the third generation. At harvest (3rd generation), T. dubium and V. persica had more and larger individuals in the presence of A. caliginosa. When both earthworm species were present, T. dubium had few but larger individuals. Our study confirms that earthworms affect plant demography and plant community structure. Our results also show that accurate prediction of long-term effects of earthworms on plant communities cannot be achieved using results on their short-term effects on plant growth. This is due to the poor understanding of the effects of earthworms on plant resource allocation and demography, and also the possibility that earthworms may exert the opposite effect on the short and long-term availability of nutrients.