Cropping history and peat amendment-induced changes in strawberry field earthworm abundance and microbial biomass

Common agricultural practices, e.g. soil tillage and organic amendment, may affect field earthworm communities considerably. However, there is little data to show how long the changes persist after a certain action. The effect of peat, commonly used in Finland to improve the horticultural soil structure, on key soil organisms is also largely unknown. Earthworm abundance and microbial biomass were studied in a strawberry field experiment (soil type silty clay) with a history of different crops (strawberry, timothy, caraway, rye, turnip rape, fiddleneck, onion and buckwheat) and peat treatments. Sampling was carried out after three years of perennial cropping of strawberry. Half of the area was peat-amended twice three years apart. The earthworm community consisted mainly of Aporrectodea caliginosa and Lumbricus terrestris. Soil peat amendment almost doubled the number of endogeic A. caliginosa, but had no effect on the anecic L. terrestris. The effect of cropping history on earthworms diminished after three years of strawberry cropping. Only the positive effect of caraway on juvenile Lumbricus spp. was detectable three years after its cropping had been finished. However, some crops had secondary effects on the earthworm distribution without significant influence on their numbers while they were grown, e.g. high numbers of A. caliginosa were recorded from soil with a history of timothy ley. The effect of strawberry cropping was contradictory: six years of continuous strawberry cropping decreased the number of the anecic L. terrestris, but during the last three years on strawberry, the proportion of L. terrestris increased from 6% to 40% in the experimental area with a concomitant great drop in the number of A. caliginosa. The role of different agricultural practices (no tillage, mulching, inter-row grass cover and pesticides) is discussed. The crop-induced changes persisted in the microbial biomass for three years (onion cropping reduced microbial biomass C), but soil amendment had no effect on microbes. The abundance of A. caliginosa was associated with soil organic C, but not with soil microbial biomass.     

Endogeic and anecic earthworm abundance in six Midwestern cropping systems

Endogeic and juvenile anecic earthworm abundance was measured in soil samples and anecic populations were studied by counting midden numbers at the sites of two long-term cropping systems trials in South-central Wisconsin. The three grain and three forage systems at each site were designed to reflect a range of Midwestern USA production strategies. The primary objectives of this work were to determine if the abundance of endogeic or anecic earthworms varied among cropping systems or crop phases within a cropping system and were there specific management practices that impacted endogeic or anecic earthworm numbers. The earthworms present in the surface soil were: Aporrectodea tuberculata (Eisen), A. caliginosa (Savigny), A. trapezoides (Dugés); and juvenile Lumbricus terrestris (L.). True endogeic abundance was greatest in rotationally grazed pasture [188 m^?2 at Arlington (ARL) and 299 m^?2 at Elkhorn (ELK)], and smallest in conventional continuous corn (27 m^?2 at ARL and 32 m^?2 at ELK). The only type of anecic earthworm found was L. terrestris L. There was an average of 1.2 middens per adult anecic earthworm and the population of anecics was greatest in the no-till cash grain system (28 middens m^?2 at ARL, 18 m^?2 at ELK) and smallest in the conventional continuous corn system (3 middens m^?2 at ARL, 1 m^?2 at ELK). Earthworm numbers in individual crop phases within a cropping system were too variable from year-to-year to recommend using a single phase to characterize a whole cropping system. Indices for five management factors (tillage, manure inputs, solid stand, pesticide use, and crop diversity) were examined, and manure use and tillage were the most important impacting earthworm numbers across the range of cropping systems. Manure use was the most important management factor affecting endogeic earthworm numbers; but no-tillage was the most important for the juvenile and adult anecic groups and had a significantly positive influence on endogeic earthworm counts as well. The pesticides used, which were among the most commonly applied pesticides in the Midwestern USA, and increasing crop diversity did not have a significant effect on either the endogeic or anecic earthworm groups in this study. Consequently, designing cropping systems that reduce tillage and include manure with less regard to omitting pesticides or increasing crop diversity should enhance earthworm populations and probably improve sustainability.     

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.     

Earthworms promote the reduction of Fusarium biomass and deoxynivalenol content in wheat straw under field conditions

A field experiment was conducted to elucidate ecosystem services provided by earthworms on the repression of phytopathogenic and toxinogenic fungi. The study focussed on decomposing Fusarium culmorum-infected and deoxynivalenol (DON)-contaminated wheat straw remaining on the soil surface as part in conservation tillage. Mesocosms were established in the topsoil of a winter wheat field located in Northern Germany, where conservation tillage has been practised for 20 years. Besides a non-earthworm treatment, two earthworm species were inoculated in the mesocosms either separately or combined: Lumbricus terrestris (anecic, detritivorous) and Aporrectodea caliginosa (endogeic, geophagous). The earthworms were exposed either to artificially Fusarium-infected wheat straw highly contaminated with DON or to non-infected straw serving as a control. The experiment was conducted during an eight week period after harvest from mid August to mid October. For both species, the artificially Fusarium-infected and DON-contaminated wheat straw was a more attractive food source than the non-infected control. In contrast to A. caliginosa, L. terrestris incorporated infected straw faster into the soil compared to control straw. Furthermore, the reduction of Fusarium biomass and DON concentration in wheat straw was significantly higher in the presence of L. terrestris than in treatments with A. caliginosa and without earthworms. Here, no significant differences could be measured between the Fusarium biomass and DON concentration in wheat straw. A. caliginosa seems not to be relevant for the reduction of Fusarium biomass and DON concentration. We concluded that amongst earthworms, anecic detritivorous species are the drivers to compensate possible negative consequences (like crop infection) of conservation tillage. They take an important role in the control of phytopathogenic and toxinogenic fungi surviving on plant residues and in the degradation of their mycotoxins.     

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.     

Worms from the cold: Lumbricid life stages in boreal clay during frost

The vertical distribution and activity of earthworm life stages were studied in an arable field during 0.5 m deep frost. The anecic Lumbricus terrestris L. were below the frost at the bottom of their home burrows (max. depth 1.0 m) and remained there apparently active. Their burrows were open, free of ice and water. The endogeic Aporrectodea caliginosa Sav., mainly small juveniles, were aestivating in the frost layer, which confirms freeze-tolerance in this species. Large A. caliginosa individuals were actively burrowing below the frost down to 1 m depth at soil temperatures close to +1 °C, frost evidently triggering much deeper burrowing than summer droughts. Demonstrating cold-hardiness, viable cocoons of both A. caliginosa and L. terrestris were obtained within a 0-0.25 m layer, frozen for ca. one month prior to sampling. These two common earthworms of boreal soils seem to over-winter in all life stages and remain active below the frost, potentially contributing to the maintenance of subsoil processes during the winter months.     

Phosphorus fractions in bulk subsoil and its biopore systems

Improving phosphorus (P) accessibility in subsoils could be a key factor for sustainable crop management. This study aims to explain the quantity of different P fractions in subsoil and its biopore systems, and to test the hypothesis that crops with either fibrous (fescue) or tap‐root systems (lucerne and chicory) leave behind a characteristic P pattern in bulk subsoil, biopore linings and the rhizosphere. The crops were cultivated for up to 3 years in a randomized field experiment on a Haplic Luvisol developed from loess. Aqua regia‐extractable P (referred to as total P) and calcium acetate lactate‐extractable P (P_CAL) were assessed at 0–30 (Ap horizon), 30–45 (E/B horizon), 45–75 and 75–105 cm subsoil depths. In addition, sequential P fractionation was performed on different soil compartments between 45 and 75 cm depths. The results showed that total P stocks below the Ap horizon (30–105 cm) amounted to 5.6 t ha^?1, which was twice as large as in the Ap, although the Ap contained larger portions of P_CAL. Both P_CAL and sequential P extractions showed that biopore linings and the rhizosphere at the 45–75 cm depth were enriched, rather than depleted, in P. The content of inorganic P (81–90% of total P) increased in the following order: bulk soil = biopores _{<2 mm} ≤ rhizosphere ≤ biopores _{>2 mm}. Biopores _{>2 mm} and rhizosphere soil were clearly enriched in resin‐ and NaHCO₃‐extractable P_i and P_o fractions. However, we failed to attribute these P distribution patterns to different crops, suggesting that major properties of biopore P originated from relict biopores, rather than being influenced by recent root systems. The stocks of the sum of these P fractions in the bulk subsoil (182 kg ha^?1 at 45–75 cm depth) far exceeded those in the biopores (3.7 kg ha^?1 in biopores _{>2 mm} and 0.2 kg ha^?1 in biopores _{<2 mm}). Hence, these biopores may form attractive locations for root growth into the subsoil but are unlikely to sustain overall plant nutrition.     

Influence of earthworm activity on aggregate-associated carbon and nitrogen dynamics differs with agroecosystem management

Earthworms are known to be important regulators of soil structure and soil organic matter (SOM) dynamics, however, quantifying their influence on carbon (C) and nitrogen (N) stabilization in agroecosystems remains a pertinent task. We manipulated population densities of the earthworm Aporrectodea rosea in three maize-tomato cropping systems [conventional (i.e., mineral fertilizer), organic (i.e., composted manure and legume cover crop), and an intermediate low-input system (i.e., alternating years of legume cover crop and mineral fertilizer)] to examine their influence on C and N incorporation into soil aggregates. Two treatments, no-earthworm versus the addition of five A. rosea adults, were established in paired microcosms using electro-shocking. A ¹³C and ¹⁵N labeled cover crop was incorporated into the soil of the organic and low-input systems, while ¹⁵N mineral fertilizer was applied in the conventional system. Soil samples were collected during the growing season and wet-sieved to obtain three aggregate size classes: macroaggregates (>250 μm), microaggregates (53-250 μm) and silt and clay fraction (<53 μm). Macroaggregates were further separated into coarse particulate organic matter (cPOM), microaggregates and the silt and clay fraction. Total C, ¹³C, total N and ¹⁵N were measured for all fractions and the bulk soil. Significant earthworm influences were restricted to the low-input and conventional systems on the final sampling date. In the low-input system, earthworms increased the incorporation of new C into microaggregates within macroaggregates by 35% (2.8 g m⁻² increase; P=0.03), compared to the no-earthworm treatment. Within this same cropping system, earthworms increased new N in the cPOM and the silt and clay fractions within macroaggregates, by 49% (0.21 g m⁻²; P<0.01) and 38% (0.19 g m⁻²; P=0.02), respectively. In the conventional system, earthworms appeared to decrease the incorporation of new N into free microaggregates and macroaggregates by 49% (1.38 g m⁻²; P=0.04) and 41% (0.51 g m⁻²; P=0.057), respectively. These results indicate that earthworms can play an important role in C and N dynamics and that agroecosystem management greatly influences the magnitude and direction of their effect.     

Homing ability widens the sphere of influence of the earthworm Lumbricus terrestris L.

We report that the deep-burrowing earthworm Lumbricus terrestris L. possesses short-range homing capability. For 2 months we followed the soil surface activity of 18 field-collected L. terrestris in an experimental laboratory population. We recorded 26 cases where an individual after completely leaving its burrow returned to the burrow by backing along its own outward trail. Maximally, homing occurred from a distance of 0.70 m and after more than 3 h on the soil surface. The evolution of homing ability in L. terrestris is most likely related to the importance of the burrow for survival and reproduction. This adaptation also guarantees individuals an expanded foraging area, which we believe can partly explain the remarkable efficiency of L. terrestris in the burial of surface litter.     

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.     

Biopore history determines the microbial community composition in subsoil hotspots

Biopores are hotspots of nutrient mobilisation and shortcuts for carbon (C) into subsoils. C processing relies on microbial community composition, which remains unexplored in subsoil biopores. Phospholipid fatty acids (PLFAs; markers for living microbial groups) and amino sugars (microbial necromass markers) were extracted from two subsoil depths (45–75 cm ; 75–105 cm) and three biopore types: (I) drilosphere of Lumbricus terrestris L., (II) 2-year-old root biopores and (III) 1.5-year-old root biopores plus six 6 months of L. terrestris activities. Biopore C contents were at least 2.5 times higher than in bulk soil, causing 26–35 times higher Σ PLFAs g^-1 soil. The highest Σ PLFAs were found in both earthworm biopore types; thus, the highest soil organic matter and nutrient turnover were assumed. Σ PLFAs was 33% lower in root pores than in earthworm pores. The treatment affected the microbial community composition more strongly than soil depth, hinting to similar C quality in biopores: Gram-positives including actinobacteria were more abundant in root pores than in earthworm pores, probably due to lower C bioavailability in the former. Both earthworm pore types featured fresh litter input, promoting growth of Gram-negatives and fungi. Earthworms in root pores shifted the composition of the microbial community heavily and turned root pores into earthworm pores within 6 months. Only recent communities were affected and they reflect a strong heterogeneity of microbial activity and functions in subsoil hotspots, whereas biopore-specific necromass accumulation from different microbial groups was absent.     

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.     

The drilosphere concept: Fine-scale incorporation of surface residue-derived N and C around natural Lumbricus terrestris burrows

Anecic (deep-burrowing) earthworms are important for soil biogeochemical functioning, but the fine-scale spatial range at which they incorporate C and N around their burrows (the drilosphere sensu stricto) needs to be investigated under realistic conditions. We conducted a field experiment to delimit spatially the extent to which soil around natural Lumbricus terrestris burrows is influenced biochemically. We placed plant litter dual-labelled with ¹³C and ¹⁵N stable isotope tracers on L. terrestris burrow openings and we measured residue-derived ¹³C and ¹⁵N in thin concentric layers (0–2, 2–4, 4–8 mm) around burrows with or without a resident earthworm. After 45 days, earthworms were significantly enriched in ¹³C and ¹⁵N as a result of feeding on the plant litter. At 0–5 cm soil depth, soil ¹⁵N concentrations were significantly higher around occupied than unoccupied burrows, and they were significantly higher in all burrow layers (including 4–8 mm) than in bulk soil (50–75 mm from burrow). This suggests that biochemical drilosphere effects of anecic earthworms, at least in the uppermost portion of the burrow, extend farther than the 2 mm layer assumed traditionally.     

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.     

Soil biopore estimation: effects of tillage, nitrogen, and photographic resolution

Networks of biopores created by plant and animal activity might accumulate in untilled cropping systems. These would be relatively well connected to the soil surface. The objective of this study was to count biopores after long-term no-till in comparison to recently tilled soil. Biopores were counted and measured to 80 cm depth at 10 cm increments in plots either under no-till wheat production for 1 year or for 17 years, and receiving zero or 130 kg ha⁻¹ N. The measurements were repeated using different photographic methods with increased resolution. The only significant difference between the long and short term no-till was in biopore sizes over 1 mm diameter, where long-term no-till produced from 30 to 100% more biopores, probably caused by increased earthworm activity. Over 99% of biopores measured were less than 1 mm diameter. There was no difference between tillage or N treatments in the number of these smaller biopores at any depth. This means small biopores did not accumulate either above or below the plow layer in an untilled cropping system. Improved resolution in the second set of measurements produced a 100-fold increase in detection of biopores in the 0.3–0.5 mm range. This provides evidence that a substantial portion of biopores are very small and were missed in the first year of this study and perhaps in other studies of this type. It is hypothesized that biopores of 0.05–0.5 mm diameter make up over half of total biopore volume and might have a significant role in movement of water and gases.     

Effects of perennial fodder crops on soil structure in agricultural headlands

In agricultural headlands, rooting and yield of crops may be limited because of soil‐structure changes as a consequence of multiple passes of turning machinery. We hypothesized that perennial forage crops can substantially alter soil structure in agricultural headlands. On one experimental field and two commercial farms on Haplic Luvisols from respectively loess and sandy loess in the Lower Rhine Bay (Germany), we investigated how 4 y of continuously grown grass/clover or alfalfa affected soil structure and the performance of subsequent spring wheat. Compared with a crop rotation with annual plowing to 30 cm soil depth, perennial forage crops led to increased soil C content (+1.3% to +22.8%) and N content (+4.2% to +15.1%), higher densities of medium and coarse biopores at a depth of 35 cm, more large water‐stable soil macroaggregates, higher biomass and abundance of anecic earthworms, and higher grain yield and grain protein content of spring wheat grown as the following crop. Root‐length density of spring wheat in the subsoil was not affected by the preceding perennial fodder crops in two of the three field trials. We concluded that besides increasing N input to the soil, perennial cropping of grass/clover or alfalfa has effects on soil structure that may substantially reduce yield losses in agricultural headlands.     

Earthworms and collembola relationships: effects of predatory centipedes and humus forms

Relationships between anecic earthworms (Lumbricus terrestris and Aporrectodea giardi) and the collembolan species Heteromurus nitidus (Templeton, 1835), which is known to be attracted to earthworms, were investigated in an 8-week laboratory experiment. Our aims were (1) to assess whether earthworms influence the population dynamics of H. nitidus, and (2) to study pathways of influence and how earthworm effects are modified by humus forms and predators. Using microcosms with three defaunated humus forms, then provided with earthworms and predators, we intended to demonstrate that, amongst possible favourable effects of earthworms on springtail populations, earthworm activity may provide greater access and more pathways for springtails to explore soil and avoid predation. We expected that the effects of predators (centipedes) on the abundance of H. nitidus would increase from less (calcic mull) to more (moder) compact soil, and we hypothesized that earthworms would reduce predation pressure on H. nitidus by providing escape routes through increased macroporosity. Humus forms and earthworms only affected the population size of H. nitidus under high predation pressure, when collembolan numbers were higher in calcic mull than in moder, and were increased by the presence of earthworms. These results corroborate the hypothesis that earthworms, by increasing soil macroporosity, improve the escape routes for Collembola and thus evade predation. In moder humus earthworms increased the density of H. nitidus whether predators were present or not, so we cannot exclude that earthworms were also directly beneficial to H. nitidus. However, the hypothesis of a functional relationship mediated by soil macroporosity seems relevant since it was supported by differences observed when considering body size. When two size classes were distinguished within populations of H. nitidus (1) the positive effect of earthworms in moder was observed only on larger Collembola (>1 mm), (2) the density of the larger Collembola was decreased by predation only in moder and not in mull, (3) the effects of predators on the smaller individuals were not influenced by the presence of earthworms whatever the humus form, and was not decreased by the presence of earthworms. Nevertheless, factors other than macroporosity may operate as the presence of earthworms in acidic mull led to an unexplained decrease in the abundance of small-sized H. nitidus.     

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.     

Deep-burrowing earthworm additions changed the distribution of soil organic carbon in a chisel-tilled soil

We investigated the influence of earthworms on the three-dimensional distribution of soil organic carbon (SOC) in a chisel-tilled soil. By burrowing, foraging, and casting at the surface and throughout the soil, anecic earthworms such as Lumbricus terrestris L. may play a major role in regulating the spatial distribution of organic matter resources both at the surface and within the soil. In the fall of 1994, we manipulated ambient earthworm communities, which were without deep burrowing species, by adding 100 earthworm individuals m⁻² in spring and fall for 3 years. Overall, the biomass of L. terrestris was increased with earthworm additions and total earthworm biomass declined compared with ambient control treatments. To investigate the spatial variability in soil organic carbon due to this shift in earthworm community structure, we sampled soil on a 28×24 cm grid from the surface to 40 cm in four layers, 10 cm deep. Samples were analyzed for total carbon. We found that additions of anecic earthworms significantly increased average soil organic carbon content from 16.1 to 17.9 g C kg⁻¹ for the 0–10 cm soil, and from 12.4 to 14.7 g kg⁻¹ at 10–20-cm depth, and also changed the spatial distribution of soil organic carbon from uniform to patchy, compared with the ambient treatment.     

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.