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.     

Earthworm-mediated maternal effects on seed germination and seedling growth in three annual plants

Many ecological studies have pointed out maternal effects in plants and shown that plant maternal environment influences germination of their seed and subsequent seedling growth. However, few have tested for maternal effects induced by soil macroorganisms. We tested whether two earthworm species (Aporrectodea caliginosa and Lumbricus terrestris) trigger such maternal effects on seed germination and seedling growth of three plant species (Veronica persica, Poa annua and Cerastium glomeratum). Our results show that, through maternal effects, A. caliginosa enhanced seed germination (V. persica and P. annua) and seedling growth (C. glomeratum and P. annua) while L. terrestris reduced seed germination only in V. persica. In some cases, the increase in germination rates of seeds produced in the presence of earthworms was associated with a reduction of nitrogen content in seeds. These results show that earthworms induce maternal effects in plants and that the size and direction of these effects depend on the combination of plant and earthworm species.     

Earthworms as seedling predators: Importance of seeds and seedlings for earthworm nutrition

Anecic earthworms have been shown to collect, concentrate and bury seeds in their burrows. Moreover, recent studies suggest that earthworms function as granivores and seedling herbivores thereby directly impacting plant community assembly. However, this has not been proven unequivocally. Further, it remains unclear if earthworms benefit from seed ingestion, i.e., if they assimilate seed carbon. We set up a series of three laboratory experiments in order to test the following hypotheses: (1) anecic earthworms (Lumbricus terrestris L.) not only ingest seeds but also seedlings, (2) ingestion of seedlings is lower than that of seeds due to a ‘size refuge’ of seedlings (i.e., they are too big to be swallowed), and (3) seeds and seedlings contribute to earthworm nutrition. L. terrestris readily consumed legume seedlings in the radicle stage, whereas legume seeds and seedlings in the cotyledon stage, and grass seeds and seedlings in the radicle and cotyledon stage were ingested in similar but lower amounts. Importantly, ingestion of seedlings, in contrast to seeds, was lethal for all plant species. Moreover, earthworm weight change varied with the functional identity and vitality of seeds and natural ¹⁵N signatures in earthworm body tissue underlined the importance of seedlings for earthworm nutrition. The results indicate that the anecic earthworm L. terrestris indeed functions as a granivore and seedling herbivore. The selectivity in seedling ingestion points at the potential of direct earthworm effects on plant community assembly. Further, seeds and seedlings most likely contribute significantly to earthworm nutrition potentially explaining the collection and concentration of seeds by L. terrestris in its middens and burrows; however, the present results call for experiments under more natural conditions.     

Competitive interactions affect the growth of Aporrectodea caliginosa and Lumbricus terrestris (Oligochaeta: Lumbricidae) in single- and mixed-species laboratory cultures

Negative interactions between earthworms may arise from high earthworm population densities. Under high populations in the field, niche separation or migration away from competitive pressure may help to regulate a multi-species population to a given level. This may not be possible in laboratory experiments, leading to an increase in competitive interactions which may alter earthworm growth rates and affect decomposition and nutrient mineralization processes. The objective of this experiment was to determine how growth rates of the endogeic earthworm Aporrectodea caliginosa Sav. and the anecic earthworm Lumbricus terrestris L. are affected by increasing population density and container size in both single- and multi-species cultures. Earthworm growth responses were compared in 1-L cylindrical pots containing disturbed soil and in 2.3-L PVC cores containing undisturbed soil. The relationship describing intra- and inter-specific competition was not affected by container type for both species. Nonetheless, decreasing the container size restricted the growth of L. terrestris in both single- and multi-species cultures, but only restricted the growth of A. caliginosa in multi-species cultures. For both species, a population density greater than one individual per litre reduced earthworm growth rates significantly, while weight loss in monocultures occurred when there were more than 10 A. caliginosa, and more than three L. terrestris per litre. Growth rates of both species were restricted in all population density treatments including the lowest of 0.9 individuals per litre. Further work is needed to find the population density at which growth rates are not affected and which may be used as an appropriate population in laboratory pot experiments to measure the effects of earthworms on soil processes and plant growth.     

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.     

Influence of two exotic earthworm species with different foraging strategies on abundance and composition of boreal microarthropods

In North America, many species of European earthworms have been introduced to northern forests. Facilitative or competitive interactions between these earthworm species may result in non-additive effects on native plant and animal species. We investigated the combined versus individual effects of the litter-dwelling earthworm Dendrobaena octaedra Savigny, 1826 and the deep-burrowing species Lumbricus terrestris L., 1758 on microarthropod assemblages from boreal forest soil by conducting a mesocosm experiment. Soil cores from earthworm-free areas of northern Alberta, Canada, were inoculated with D. octaedra alone, L. terrestris alone, both worm species together, or no earthworms. After 4.5 months, microarthropods were extracted from the soil, counted, and identified to higher taxa. Oribatid mites were further identified to family and genus. Abundance of microarthropods was significantly lower in the treatment containing both species than in the no earthworm treatment and the L. terrestris treatment. Oribatida and Prostigmata/Astigmata differed significantly among treatments and were lowest in the treatment containing both earthworm species, followed by the D. octaedra treatment, although post-hoc pairwise comparisons were not significant. Within the Oribatida, composition differed between the control and L. terrestris treatments as compared to the D. octaedra and both-species treatments, with Suctobelbella and Tectocepheus in particular having higher abundances in the control treatment. Effects of the two earthworm species on microarthropods were neither synergistic nor antagonistic. Our results indicate that earthworms can have strong effects on microarthropod assemblages in boreal forest soils. Future research should examine whether these changes have cascading effects on nutrient cycling, microbial communities, or plant growth.     

Herbivory of an invasive slug in a model grassland community can be affected by earthworms and mycorrhizal fungi

Invasion of non-native species is among the top threats for the biodiversity and functioning of native and agricultural ecosystems worldwide. We investigated whether the herbivory of the slug Arion vulgaris (formerly Arion lusitanicus; Gastropoda), that is listed among the 100 worst alien species in Europe, is affected by soil organisms commonly present in terrestrial ecosystems (i.e. earthworms—Annelida: Lumbricidae and arbuscular mycorrhizal fungi—AMF, Glomerales). We hypothesized that slug herbivory would be affected by soil organisms via altered plant nutrient availability and plant quality. In a greenhouse experiment, we created a simple plant community consisting of a grass, a forb, and a legume species and inoculated these systems with either two earthworm species and/or four AMF taxa. Slugs were introduced after plants were established. Earthworms significantly reduced total slug herbivory in AMF-inoculated plant communities (P?=?0.013). Across plant species, earthworms increased leaf total N and secondary metabolites, AMF decreased leaf thickness. Mycorrhizae induced a shift in slug feeding preference from non-legumes to legumes; the grass was generally avoided by slugs. AMF effects on legume herbivory can partly be explained by the AMF-induced increase in total N and decrease in C/N ratio; earthworm effects are less clear as no worm-induced alterations of legume plant chemistry were observed. The presence of earthworms increased average AMF colonization of plant roots by 140 % (P?<?0.001). Total shoot mass was significantly increased by AMF (P?<?0.001). These data suggest that the feeding behavior of this invasive slug is altered by a belowground control of plant chemical quality and community structure.     

Plant species influence microbial diversity and carbon allocation in the rhizosphere

Plant species effects on microbial communities are attributed to changes in microbial community composition and biomass, and may depend on plant species specific differences in the quality of resources (carbon) inputs. We examined the idea that plant-soil feedbacks can be explained by a chance effect, which is the probability of a highly productive or keystone plant species is present in the community and will influence the functions more than the number of species per se. A ¹³C pulse labelling technique was applied to three plant species and a species mixture in a greenhouse experiment to examine the carbon flow from plants to soil microbial communities. The ¹³C label was given as CO₂ to shoots of a legume (Lotus corniculatus), a forb (Plantago lanceolata), a grass (Holcus lanatus) and a mixture of the three species. Microbial phospholipid fatty acids (PLFA) was analysed in order to determine the biomass and composition of the soil microbial community. The incorporation of the stable isotope into soil microorganisms was determined through GC-IRMS analyses of the microbial PLFAs. Plant species identity did not influence the microbial biomass when determined as total carbon of microbial phospholipid fatty acids. However, the labelled carbon showed that the grass monoculture (H. lanatus) and the plant mixture allocated more ¹³C into bacteria and actinomycete biomass than the other plant species. H. lanatus monocultures had also the highest amounts of ¹³C allocated to AM-fungi and saprophytic fungi. The carbon allocation from plants to soil microorganisms in a plant species mixture can thus be explained by the presence of a highly productive species that influence soil functions.     

Invasion of a deciduous forest by earthworms: Changes in soil chemistry, microflora, microarthropods and vegetation

Ecosystems of northern North America existed without earthworm fauna until European settlers arrived and introduced European species. The current extent of invasion by some of these species, Lumbricus terrestris L., Octolasion tyrtaeum Savigny and Dendrobaena octaedra Savigny, into an aspen forest in the Canadian Rocky Mountains and the effects of the invasion on soil chemistry, microflora, soil microarthropods and vegetation were investigated. Densities of earthworm species, soil structure, plant coverage and abundance were determined along three transects starting at the edge of the forest. At locations with L. terrestris, litter was incorporated into the soil, and where O. tyrtaeum was present, organic layers were mixed with mineral soil layers. Organic layers disappeared almost entirely when both species occurred together. Carbon and nitrogen concentrations were reduced in organic layers in the presence of L. terrestris and O. tyrtaeum. Microbial biomass and basal respiration were reduced when L. terrestris and O. tyrtaeum were present, presumably due to resource competition and habitat destruction. Microarthropod densities and the number of microarthropod species were strongly reduced in the presence of O. tyrtaeum (−75% and −22%, respectively), probably through mechanical disturbances, increasing compactness of the soil and resource competition. The coverage of some plant species was correlated with earthworm abundance, but the coverage of others was not. Despite harsh climatic conditions, the invasion of boreal forest ecosystems by mineral soil dwelling earthworm species is proceeding and strongly impacts soil structure, soil chemistry, microorganisms, soil microarthropods and vegetation.     

Zum Einfluß von obstbaulichen Bodenpflegemaßnahmen auf Lumbriciden,insbesondere Lumbricus terrestris L .

Influence of orchard soil management on lumbricids, especially Lumbricus terrestris L. In a long-term soil management experiment (apple orchard treated with cultivation methods such as grass mulch, grass harvest, straw mulch, clean cultivation) 6 plots were selected in order to study the abundance and biomass of earthworm populations. The following results were obtained: In all treatments with the exception of straw mulch L. terrestris represented the majority (60.5% on average) of the lumbricid population. Their share in the biomass amounted to 93.3%. The earthworm biomass was the highest in grass mulch; a close correlation was observed between the litter production (grass and leaves) and the biomass of L. terrestris. Added farm yard-manure on plots with grass mulch had no additional positive effect on earthworms. Although a higher earthworm biomass was found under grass mulch in comparison with straw mulch, the species diversity was higher in the latter. Clean cultivation had a highly detrimental effect on all earthworms, especially on the horizontally burrowing species. The annual nitrogen turnover by L. terrestris in mulched orchards was estimated to approx. 50 kg N/ha.     

Fat but slim: Criteria of seed attractiveness for earthworms

Earthworms were shown to significantly affect seeds and seedlings survival via their ingestion and digestion for nutritive purposes. Such selective feeding of earthworms on plant seeds is likely to favour certain plant species and to affect seed bank composition, plant recruitment and plant community structure. Relationships between earthworms and seeds, particularly seed traits that determine attractiveness of seeds for earthworms, are yet to be determined. In this study, the influence of six seed traits was tested on the ingestion, digestion and germination of seeds by two earthworm species (Lumbricus terrestris, anecic and Satchellius mammalis, epigeic). The seed traits tested were their length, width, weight, shape, oil content and the presence of trichomes on their surface. Each earthworm species was introduced into a microcosm with eleven seed species from a chalk grassland that represented those different traits. Ingested, digested and germinated seeds were counted after voiding the guts of the earthworms. Univariate and multivariate analyses showed that seed length, width, weight and seed oil content could significantly affect the ingestion of seeds for both earthworm species. Seed width and seed oil content were the two traits that influenced the digestion of seeds the most, but only for L. terrestris. We also found that seed ingestion was earthworm species-specific but we found no correlation between earthworm traits and number of ingested or digested seeds. Few seeds germinated from L. terrestris casts and no seeds germinated from S. mammalis casts. Implications in terms of plant evolution strategies are further discussed.     

Plant community impacts on the structure of earthworm communities depend on season and change with time

Declining plant diversity potentially threatens essential ecosystem functions driven by the decomposer community, such as litter decomposition and nutrient cycling. Currently, there is no consensus on the interrelationships between plant diversity and decomposer performance and previous studies highlighted the urgent need for long-term experiments.In the Jena Experiment we investigated the long-term impacts of plant community characteristics on the structure of earthworm communities representing key decomposers in temperate grassland. We repeatedly sampled plots varying in plant species richness (1-16 species), plant functional group richness (1-4 groups), and presence of certain plant functional groups (grasses and legumes) three, four, and six years after establishment of the experiment in spring and autumn.The results show that earthworm performance is essentially driven by the presence of certain plant functional groups via a variety of mechanisms. Plant productivity (root biomass) explained most of the detrimental grass impacts (decrease in earthworm performance), while beneficial legume effects likely were linked to high quality inputs of plant residues (increase in earthworm performance). These impacts depended on the functional group of earthworms with the strongest effects on surface feeding anecic earthworms and minor effects on soil feeding endogeic species. Remarkably, effects of plant community characteristics on the composition and age structure of earthworm communities varied between seasons. Moreover, plant diversity effects reported by a former study decreased and detrimental effects of grasses increased with time.The results indicate that plant community characteristics, such as declining diversity, indeed affect the structure of earthworm communities; however, loss of key plant functional groups is likely to be more important than plant species number per se. However, in frequently disturbed ecosystems plant species richness might be important for the recovery and resilience of belowground functions. Moreover, the results accentuate the importance of long-term repeated measurements to fully appreciate the impacts of plant community composition and diversity on ecosystem properties. Single point observations may be misleading and potentially mask the complexity of above-belowground interrelationships.     

Non-native anecic earthworms (Lumbricus terrestris L.) reduce seed germination and seedling survival of temperate and boreal trees species

Recent studies have shown that the introduction of non-native earthworms in previously earthworm-free soils may have negative impacts on the recruitment of certain understory plant species in northern temperate forests. There is a need, therefore, to understand the mechanisms that may underlie this phenomenon. A microcosm study was conducted to test the effects of the anecic earthworm, Lumbricus terrestris L., on the number of days for germination, % seed germination, seedling survival and seedling biomass of 14 tree species native to southern Quebec (Canada). Seeds of these species were germinated and grown in the presence or absence of L. terrestris. The presence of earthworms significantly reduced % seed germination of seven tree species, as well as seedling survival of three tree species. The germination date of three tree species was significantly affected, either positively or negatively, by the presence of earthworms. Earthworms had no effect on seedling biomass. Results suggest that the introduction of L. terrestris into forested ecosystems of southern Québec may potentially alter overstory composition through several mechanisms that differentially affect the recruitment of various tree species in the understory.     

The effect of feeding behavior on Hg accumulation in the ecophysiologically different earthworms Lumbricus terrestris and Octolaseon cyaneum: A microcosm experiment

 A soil microcosm experiment was performed to assess the uptake of Hg from various Hg-spiked food sources (soil, leaf litter and root litter of Trifolium alexandrinum) by two earthworm species, Lumbricus terrestris (anecic) and Octolaseon cyaneum (endogeic). Treatments were applied in which one of the three food sources was Hg spiked and the other two were not. Additional treatments in which all or none of the food sources were Hg spiked were used as controls. Uptake of Hg from soil into tissues of both earthworm species was significantly higher than uptake of Hg from leaf litter or root litter, indicating that soil may be the most important pool for the uptake of Hg into earthworms. In addition, the anecic L. terrestris significantly accumulated Hg from all Hg-spiked food sources (leaf litter, root litter and soil), whereas the endogeic O. cyaneum took up Hg mainly from soil particles. Interestingly, there was no further increase in Hg in L. terrestris when all food sources were Hg spiked compared to the single Hg-spiked sources. This may be attributed to the relatively high Hg content in the soil, which may have influenced the feeding behavior of the earthworms, although their biomass did not significantly decline. We suggest that, in addition to the physiological differences, feeding behavior may also play a role in the contrasting uptake of Hg by the two earthworm species.     

Short rotation forestry – Earthworm interactions: A field based mesocosm experiment

Short rotation forestry (SRF) which consists of planting rapidly growing native and non-native tree species has been introduced to the UK to increase woody biomass production. A largely unknown aspect of SRF species is their interaction with soil fauna, of which the earthworm community is a major component. Earthworms have a pronounced impact on litter decomposition, nutrient cycling and tree growth. Conversely, tree litter and root chemistry can impact on the associated earthworm community development. The aim of this study was to determine direct interactions between SRF species and earthworms. A field-based mesocosm experiment was conducted using Betula pendula (birch) and Eucalyptus nitens (eucalyptus) with two earthworm species Lumbricus terrestris and Allolobophora chlorotica. The one year experiment revealed that native birch and non-native eucalyptus had a similar influence on L. terrestris population development. However, birch had a positive impact on A. chlorotica population establishment compared with eucalyptus. In the presence of earthworms, total tree biomass and leaf nitrogen concentration of eucalyptus were increased respectively by 25% and 27% compared with an earthworm-free control. In the presence of earthworms, surface litter incorporation was greater for both tree species (almost 5 times for birch and 3 times for eucalyptus) compared with controls. This work showed direct SRF-earthworm interactions which differed for tree species.     

Evenness and plant species identity affect earthworm diversity and community structure in grassland soils

Diversity of plant species and the species composition (identity) are known to influence below-ground diversity. In this paper we examine the effects of plant species diversity (richness and evenness), rates of nitrogen application and planting density, on earthworm community structure in grassland. The study was carried out at three sites in Ireland using a Simplex experimental design to define the compositions of the experimental plant communities used.A negative relationship was detected between diversity (evenness) of plant species and diversity of earthworms in the soils. However, plant species identity also affected the structure of the earthworm assemblage. In particular, the legume, Trifolium repens had a strong effect but this was conditional on the rate of nitrogen application. No earthworm species favoured communities dominated by slow growing grasses (Phleum pratense and Dactylis glomerata) (P = 0.02).Higher N inputs reduced earthworm abundance and biomass under T. repens. Earthworm richness, was negatively influenced by elevated amounts of N inputs. No effect of planting density was detected but this factor also did not affect plant biomass production.     

Introducing deep burrowing earthworms (Lumbricus terrestris L.) into arable heavy clay under boreal conditions

An inoculation of the deep burrowing earthworm Lumbricus terrestris L. was carried out for soil amelioration purposes at an arable field on heavy clay in southern Finland. L. terrestris, whilst entirely absent from the field and its immediate surrounding, was present in many other local soils. Prior to inoculation, field management was changed in favour of L. terrestris by improving the subdrainage and by implementation of reduced tillage. In Autumn 1996, 82 earthworm inoculation units (EIUs) were introduced in three transects at one end of the field, one transect lying within a permanent grass strip outside the cultivated area. In Autumn 1998, L. terrestris had persisted outside the cultivated area with maximum density of 28 ind. m⁻² (median: 0 ind. m⁻²). No individuals were found inside the field. In Autumn 2003, maximum density was 43 ind. m⁻² (median: 9 ind. m⁻²) outside the cultivated area, with evidence for 8 m dispersal from inoculation points (1.1 m year⁻¹). In 2003 individuals were also found in low densities inside the field, close to subdrains, with a maximum density of 4 ind. m⁻². Results call for critical evaluation of this inoculation practice.     

Introduced earthworm species exhibited unique patterns of seasonal activity and vertical distribution,and <Emphasis Type="Italic">Lumbricus terrestris</Emphasis> burrows remained usable for at least 7 years in hardwood and pine stands

It is difficult to obtain non-destructive information on the seasonal dynamics of earthworms in northern forest soils. To overcome this, we used a Rhizotron facility to compile 7 years of data on the activity of anecic (Lumbricus terrestris) and endogeic (Aporrectodea caliginosa complex) earthworms in two contrasting soil/plant community types. We hypothesized that L. terrestris burrows would be used for longer than a typical L. terrestris lifetime, and that the distribution and activity pattern of the two earthworm species would respond differently to changes in soil moisture and temperature. For 7 years we recorded earthworm distribution and activity state bi-weekly to a depth of 1.5 m, tracked L. terrestris burrows using images captured annually, and measured soil temperature and moisture. Activity and vertical distribution of earthworms was closely linked to earthworm species and soil temperature in the fall, winter and spring. Lumbricus terrestris typically remained active through the winter, whereas the A. caliginosa complex was more likely to enter an aestivation period. Activity of all earthworms decreased substantially in July and August when soil temperature was at its highest and soil moisture at its lowest for the year. Most L. terrestris burrows were used continuously and moved very little during the 7-year study, likely creating spatiotemporally stable hotspots of soil resources. The different patterns of response of these species to soil temperature and moisture suggests that endogeic earthworms are more likely than anecic earthworms to adjust activity states in response to climate change mediated shifts in soil moisture and temperature.     

Effects of liming on survival and reproduction of two potentially invasive earthworm species in a northern forest Podzol

During the last several decades, colonization of soil by exotic earthworms and their effects on soil properties and biodiversity have been reported in forests of North America. In some northern hardwood stands, acid soils or harsh climate may have prevented earthworm colonization. However, climatic change and the increasing use of liming to restore the vigor of declining sugar maple (Acer saccharum Marsh.) stands, situated on base-poor soils in USA and Canada, could make many of these sites more suitable for earthworm colonization. We tested survival and reproduction of two exotic earthworm species (Lumbricus terrestris and Amynthas hawayanus) in unlimed and limed soils at the northern limit of the northern hardwood forest distribution in Canada. Improving soil parameters of base-poor, acidic soils by liming positively influenced activity, survivability and reproductive output of L. terrestris in this northern hardwood forest. In contrast, the high mortality and low vigor of L. terrestris observed in the unlimed plots show that soils in this area with a pH of 4.3 are not favorable to this species. Our results suggest that A. hawayanus was very active prior to winter at both soil pHs, but was not able to complete its life cycle during one year at this latitude. Both earthworm species significantly reduced organic C and total N, and increased the C/N ratio of the forest floor. Given that forest liming activities are increasing in proximity to human activities, there is high probability that some earthworm species, such as L. terrestris, will invade limed northern hardwood forests in the next decades, with possible consequences for soil organic matter turnover, nutrient cycling and forest biodiversity and dynamics.     

Nutrient content of earthworm casts influenced by different mulch types

The aim of this study was to investigate the effects of different mulch types on the quality and quantity of casts of different lumbricid earthworm species. Nutrient contents (organic carbon C_org, total nitrogen N_t and inorganic phosphate Pi), and acid phosphatase activity (SPA) of earthworm casts and soil aggregates were measured and then compared. This short time experiment was conducted in the laboratory, simulating field conditions of mulch management in temperate agroecosystems. In microcosms, the two common field species Lumbricus terrestris and Octolasion cyaneum were inoculated. Barley, lupin, maize, or sugarbeet were used as mulch according to amounts usually applied in the field. Nutrient contents and phosphatase activity in worm casts were generally enhanced compared to the soil. Nutrient values were higher in the casts of the detritivorous L. terrestris than those of the geophageous O. cyaneum. Conversely, the phosphatase activity was increased in the casts of O. cyaneum compared to L. terrestris in case of the barley and sugar-beet treatments. Cast production was related to the earthworms' change of biomass (ΔB) as well as, in the case of L. terrestris, to nitrogen content of the mulch. Due to their chemical compounds, the mulch types influence both the food selection of the worms and the acceptance by microorganisms.