Changes in potassium availability and other soil properties due to soil ingestion by earthworms

An incubation experiment was conducted to study the changes that occur in potassium availability and other soil properties with ingestion of soil by earthworms. Two soils were used. Raumai soil with high non-exchangeable K and Milson soil with low non-exchangeable K were incubated with two species of earthworm, Aporrectodea caliginosa and Lumbricus rubellus, for 8 weeks. The casts and soil samples were analysed for exchangeable K, Ca, Mg, Na, and H, pH, organic C, and texture. The results indicated that in Raumai soil, the exchangeable K levels of the casts of both earthworm species were significantly higher than for the control soil, the effect being more marked for L. rubellus than for A. caliginosa. In Milson soil, the exchangeable K levels were significantly lower in the casts of both types of earthworm than in the control soil. The nitric acid-extractable K of the soil and casts was not markedly different for either soil type, but available non-exchangeable K values were significantly higher for the casts of L. rubellus from Milson soil than for the noningested Milson soil. In Raumai soil, the exchangeable Ca was higher in the casts of L. rubellus, exchangeable Mg and H were reduced, and exchangeable Na did not change markedly in the cast compared to the control soil. For Milson soil, the casts contained lower exchangeable Ca and H but higher Na and Mg than the control. The casts of both species of earthworm had significantly higher pH values for both soil types. There was no marked difference in the organic C content of the control soil and cast samples for Milson but a reduction in the casts of A. caliginosa for the Raumai soil. Finer fractions increased in the casts of both earthworm species in both soil types.     

Earthworm channels in cultivated clayey and loamy Norwegian soils

Studies of earthworm species and their activity, expressed as channels, on cultivated loamy (Humic Cryaquept) and clayey soils (Typic Endoaqualf and Typic Cryaquept) were conducted in southern and central Norway before conversion from conventional to organic cropping systems. At all the three study areas: Landvik (Grimstad), Voll (Ås) and Kvithamar (Stjørdal), the earthworm species Aporrectodea caliginosa, Aporrectodea rosea, Allolobophora chlorotica and Lumbricus rubellus were found. At Landvik, where the deep-burrowing species Aporrectodea longa, A. caliginosa and Lumbricus terrestris dominated, medium (4–6 mm) and coarse (>6 mm) earthworm channels were most numerous below the plough layer. Almost no coarse pores were found at Voll and Kvithamar. The volume of earthworm channels below the plough layer was 0.6–0.8% of total soil volume at Landvik and Voll and 0.3–0.4% at Kvithamar. Earthworm channels >6 mm below the plough layer were useful to identify present and previous activity of deep-burrowing earthworms such as L. terrestris. Below the plough layer, roots were almost entirely restricted to earthworm channels and interaggregate spaces.     

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.     

Earthworm burrowing behaviour in 2D terraria with single- and multi-species assemblages

Experiments were performed in 2D terraria to investigate the burrowing behaviour of different earthworm species from various ecological categories in single- and multi-species assemblages. The burrowing behaviour was quantified using image analysis software during a 2-week period. Terraria were found to reveal realistic impressions of the burrowing behaviour of various species according to the ecological classification of Bouché into epigeic, endogeic and anecic species. Results of the study also permit the recommending of classifying various earthworms as intermediate species, e.g. Aporrectodea longa as endo-anecic and Lumbricus rubellus as epi-endogeic. Burrowing activity of endogeic species was significantly reduced in multi-species compositions compared to single-species treatments. Moreover, burrowing activity of Octolasion tyrtaeum was significantly reduced in the presence of Lumbricus terrestris compared to the specific single treatment. This endogeic species profited from the burrowing behaviour, e.g. bioturbation, of L. terrestris, and as such this is circumstantial evidence for commensalism (species interaction, in which one partner benefits while the other is unaffected) between anecic and endogeic earthworm species. Simultaneous burrowing activity of a combined assemblage of both endogeic species, Aporrectodea caliginosa and O. tyrtaeum, was also significantly reduced compared with the particular single treatments. Thus, this seems likely to be a response of interspecific competition and trophic niche separation between endogeic species.     

Interactive effects of functionally different earthworm species on aggregation and incorporation and decomposition of newly added residue carbon

The interactive effects of two functionally different earthworm species (Aporrectodea caliginosa (endogeic species) and Lumbricus rubellus (epigeic species)) on the incorporation of fresh residue into large macroaggregates and formation of microaggregates within these large macroaggregates were investigated during a short-term laboratory experiment using ¹³C-labelled sorghum (Sorghum bicolor (L.) Moench) residues. Soil was collected from a long-term no-tillage agricultural field, crushed through a 250-μm sieve and incubated under laboratory conditions. The following earthworm treatments were applied: (i) soil+¹³C-labelled residue+A. caliginosa; (ii) soil+¹³C-labelled residue+L. rubellus; (iii) soil+¹³C-labelled residue+A. caliginosa+L. rubellus and; (iv) soil+¹³C-labelled residue. Two residue placement treatments (i.e. surface and incorporated) were superimposed on the earthworm treatments. Earthworms were added after 8 days of incubation. Aggregate size distribution and total C and ¹³C were measured after 22 days. Microaggregates, fine inter-microaggregate particulate organic matter (inter-POM) and intra-microaggregate POM (intra-POM) were isolated from macroaggregates. Earthworms had a greater stimulating effect on the formation of large macroaggregates (>2000 μm) and microaggregates within large macroaggregates when residue was incorporated in the soil, especially in the presence of A. caliginosa. When residue was placed on the surface, residue-derived intra-POM C was highest when L. rubellus was present and significantly lower in the presence of A. caliginosa. Residue-derived inter-POM C was highest when a mix of both species was present. These results indicate that earthworm species differentially affect incorporation of fresh organic matter into stable microaggregates within macroaggregates, and that interactive effects of earthworm species might have important consequences for the incorporation and protection of C inside of microaggregates within macroaggregates especially when residues are placed on the soil surface.     

A simple and effective method to keep earthworms confined to open-top mesocosms

Earthworms can have a profound effect on a myriad of soil physical, chemical and microbial parameters. To better understand their role in the soil, they are often studied under controlled conditions. However, a persistent problem in such controlled experiments is the ability of earthworms to escape from experimental units with open tops (e.g. for plant growth). Here, we tested whether adhesive hook tape applied to the inside of mesocosms is effective in confining them to their experimental units. A mesocosm study was set up with hook tape treatments (control, one layer, two layers), mesocosm material (polyvinylchloride – PVC, polypropylene – PP) and earthworm species (Lumbricus rubellus (Hoffmeister), Aporrectodea caliginosa (Savigny), Lumbricus terrestris (L.) + Aporrectodea longa (Ude)) as different factors to study the escape of earthworms during 24 h. In the treatments without hook tape, individuals of L. rubellus and A. caliginosa escaped, with highest escape rates (80%) for L. rubellus from the PP mesocosms, and lowest escape rates (20%) for A. caliginosa from the PVC mesocosms. When hook tape was applied, in either one or two layers, no individuals of those species escaped. The two anecic earthworm species, L. terrestris and A. longa did not escape from any mesocosms, irrespective of the presence of hook tape. As not a single earthworm escaped from the hook tape treatments, we conclude that applying hook tape is a simple, inexpensive and effective method to keep earthworms confined to experimental units.     

Distribution of earthworms and influence of soil properties across a successional sand dune ecosystem in NW England

An investigation of earthworms across a sand dune system in NW England examined species distribution and abundance with respect to soil physico-chemical conditions and management. Replicated 0.1 m² quadrats were examined every 50 m along 700 m transects across areas of vegetation succession and samples were then taken every 10 m where earthworms were first encountered. A translocation of Aporrectodea longa assessed the ability of this species to live in soils with a high sand content and laboratory work examined growth, maturation and survival of this species in soils with increasing proportions of sand (0–100%). Nine earthworm species were found on the dunes, but none in yellow dunes where organic matter (OM) content was <1%. Dendrobaena octaedra and Lumbricus rubellus were located 300 m from the strand line in grey dunes with an OM content of 3.9%. Allolobophora chlorotica and Lumbricus castaneus occurred within a wet dune slack at 340 m (OM content 11%). In areas of human disturbance (dune car park), A. longa, Aporrectodea caliginosa and Lumbricus terrestris were present, and in soil below pine trees L. castaneus and L. rubellus were present in litter. Laboratory growth of A. longa demonstrated significant (p < 0.05) increases in growth (2–2.75 g) with 25 and 50% sand compared with 0, 75 and 100% over 24 weeks and a greater rate of maturation. Initial results show earthworm colonisation to be influenced by dune successional stage. Laboratory findings suggested A. longa could exist in higher sand content areas but experimental design needs development. The translocation was unsuccessful. Future investigations could examine soil properties more closely and undertake monitoring on site throughout the year.     

Earthworm effects on selected physical and chemical properties of soil aggregates

Summary Some physical and chemical properties of 1-to 2-mm aggregates obtained from casts and the burrow-wall material of the earthworm species Lumbricus terrestris, Aporrectodea longa, and Aporrectodea caliginosa were determined in order to show the effects of earthworms on the stabilization of soil aggregates. The results were compared with those of the natural soil from the Ap horizon of a Parabraunerde (Luvisol, FAO). Both the tensile strength and the water stability of aggregates from casts and burrow-wall material were reduced compared with those of the natural aggregates but were increased compared with those of remoulded aggregates. These results indicate that to a great extent existing bonds are destroyed by earthworm ingestion. Nevertheless, earthworm activities are advantageous for the stabilization of reformed aggregates. The coarse sand fraction is reduced by selective ingestion by earthworms. The organic C content is increased by 4.1–21.0% for burrow-wall material and by 21.2–43.0% for casts. The carbonate content of aggregates from casts and burrow-wall material of L. terrestris was reduced by more than 50%, while that of A. longa showed no noticeable changes and that of A. caliginosa was increased by more than 60%. The total content of polysaccharides was increased by 35–87% for casts and by 33–46% for the burrow-wall material of all earthworm species. The most frequently detected monosaccharides were glucose, galactose, and glucosamine. L. terrestris appeared to have the strongest effect on the interparticle bonding of the reformed aggregates, measured both as tensile strength and water stability, followed by A. longa and A. caliginosa.     

Earthworm feeding activity and development of the humus profile

Earthworm casts and digestive tract contents were simultaneously examined, using the same methods, in a recently formed humus profile in a mountain spruce forest. Earthworm species had distinct diets and an earthworm foodweb could be distinguished. Lumbricus terrestris and Aporrectodea icterica were distinct from the other species examined: the former to some extent as a litter consumer, and both species because they excavated mineral material which was deposited within new top layers of the mull humus. Aporrectodea nocturna and Aporrectodea caliginosa both had a non-specific soil feeding mode. Most of the species enriched the humus profile with amorphous organic matter finely incorporated within a mineral matrix. Besides different food selection, a network of burrows was produced as a consequence of the different burrowing behaviour of each earthworm species. Received: 2 January 1997     

Phosphatase activity in earthworm faeces

Wormcasts of Allolobophora caliginosa have been shown to have higher phosphatase activity than occurs in uningested soil, resulting in an increase in inorganic P released by mineralization of organic P. This paper describes similar observations on organic wastes which may be used for vermiculture. Besides A. caliginosa, three species which will feed on dung or other organic substrates were investigated, Eisenia foetida, Dendrobaena veneta and Lumbricus rubellus. When cow dung was used as a substrate, background phosphatase activity was too high for the effects of earthworm activity to be detected and a culture medium was therefore developed from sterilized paper waste sludge with added phytin. Phosphatase activity, assayed by a modified Hoffman method, was higher in the presence of all four earthworm species than in controls. It showed two peaks in relation to pH, at 3–5 and 9–10. The former is attributed to microbial activity and the latter to microbial activity or earthworm alkaline phosphatase. After 1 month, cultures of paper waste sludge with phytin contained about twice as much water soluble P when worms were present than in their absence.