Effect of food type and placement on earthworm (Aporrectodea tuberculata) burrowing and soil turnover

Subsurface-dwelling Aporrectodea tuberculata, a common earthworm in Upper-Midwest (USA) agricultural fields, may be a significant component of agroeco-systems with regard to soil mixing and preferential transport of water and chemicals. In this study we looked at effects of food residue placement and food type on A. tuberculata burrowing and soil turnover in two-dimensional Evans box microcosms. Four food residue placements mimicked patterns induced by primary tillage and two food types, readily available and natural food sources, with no food as a control. An average earthworm population of 100 earthworms m^-2 was calculated to generate 1058 km ha^-1 of new burrows and turnover 7.9 Mg ha^-1 of soil in 1 week of activity at 20°C. Burrowing was random until food sources were encountered, at which time burrowing appeared to center around the food source.     

Growth rates of Aporrectodea caliginosa (Oligochaetae: Lumbricidae) as influenced by soil temperature and moisture in disturbed and undisturbed soil columns

Earthworm growth is affected by fluctuations in soil temperature and moisture and hence, may be used as an indicator of earthworm activity under field conditions. There is no standard methodology for measuring earthworm growth and results obtained in the laboratory with a variety of food sources, soil quantities and container shapes cannot easily be compared or used to estimate earthworm growth in the field. The objective of this experiment was to determine growth rates of the endogeic earthworm Aporrectodea caliginosa (Savigny) over a range of temperatures (5–20 °C) and soil water potentials (−5 to−54 kPa) in disturbed and undisturbed soil columns in the laboratory. We used PVC cores (6 cm diameter, 15 cm height) containing undisturbed and disturbed soil, and 1 l cylindrical pots (11 cm diameter, 14 cm height) with disturbed soil. All containers contained about 500 g of moist soil. The growth rates of juvenile A. caliginosa were determined after 14–28 days. The instantaneous growth rate (IGR) was affected significantly by soil moisture, temperature, and the temperature×moisture interaction, ranging from −0.092 to 0.037 d⁻¹. Optimum growth conditions for A. caliginosa were at 20 °C and −5 kPa water potential, and they lost weight when the soil water potential was −54 kPa for all temperatures and also when the temperature was 5 °C for all water potentials. Growth rates were significantly greater in pots than in cores, but the growth rates of earthworms in cores with undisturbed or disturbed soil did not differ significantly. The feeding and burrowing habits of earthworms should be considered when choosing the container for growth experiments in order to improve our ability to extrapolate earthworm growth rates from the laboratory to the field.     

Effects of endogeic earthworms on soil processes and plant growth in coniferous forest soil

Summary The effects of the endogeic earthworm, Aporrectodea caliginosa tuberculata (Eisen) on decomposition processes in moist coniferous forest soil were studied in the laboratory. The pH preference of this species and its effects on microbial activity, N and P mineralization, and the growth of birch seedlings were determined in separate pot experiments. Homogenized humus from a spruce stand was shown to be too acid for A. c. tuberculata. After liming, the earthworms thrived in the humus and their biomass increased (at pH above 4.8). In later experiments in which the humus was limed, the earthworms positively influenced the biological activity in humus and also increased the rate of N mineralization. A. c. tuberculata increased the growth of birch seedlings, with increases observed in stems, leaves, and roots. Neither NH ₄ ⁺ -N fertilizer nor mechanical mixing with artificial worms affected seedling growth. No plant-growth-affecting compounds (e.g., hormone-like compounds) due to the earthworms were present in the humus. The shoot: root ratio in the birch seedlings was not affected by either the earthworms or the fertilizer. The experiments revealed the impact of earthworm activity on soil processes and plant growth.     

The response of Aporrectodea rosea and Aporrectodea trapezoides (Oligochaeta: Lubricidae) to moisture gradients in three soil types in the laboratory

The question of whether the response of earthworms to soil moisture is governed by their reaction to soil wetness (moisture content) or to soil water energy (matric suction) was examined in two species of earthworm using moisture gradients in three contrasting soil types with clay contents varying from 4 to 39%. Gravimetric moisture gradients ranging over 5–30% were established in horizontal cores comprising 12 or 14 sections containing loosely packed soil. Earthworms were introduced to each section at the beginning of each experiment. The earthworms moved from sections containing dry soil into adjacent sections containing moister soil. Clear effects were evident after 6 h but these became more obvious after 96 h. For the earthworm Aporrectodea rosea, the threshold soil mositure level at which earthworms were induced to move away from dry soil was a matric suction of about 300 kPa (pF 3.4) and was independent of soil type. In contrast, for A. trapezoides, the threshold soil moisture varied with soil type (sandy loam 15 kPa, loam 25 kPa, clay 300 kPa). We conclude that, for the earthworm A. rosea, matric suction and not water content of soil provided the cue by which the earthworm recognized dry soil. For A. trapezoides, there was an interaction between matric suction and soil type in which the response of A. trapezoides to soil moisture varied with soil texture and the threshold for avoidance of dry soil ranged from a matric suction of 300 kPa (20% w/w) in clay to 15 kPa (10% w/w) in sandy loam.     

Population dynamics of the earthworm Aporrectodea trapezoides (Annelida: Lumbricidae) in a Western Australian pasture soil

Summary The changes in size and age-composition of an earthworm population were studied in a Western Australian pasture developed since 1912. The population size in the surface 10 cm was estimated by handsorting during the cool wet season (19 weeks from May to September). Two species, both exotic, were found, Aporrectodea tranpezoides (Lumbricidae) and Microscolex dubius (Acanthodrilidae). Of the 615 individual speciemens collected, 99.7% were A. trapezoides. The abundance of A. trapezoides increased from 58 m^-2 at the time of the opening rains to 170 m^-2 (88.6 g live weight m^-2) after 10.5 weeks. Near the end of the wet season (in October) the density was 37 m^-2. At the time of the opening rains the population consisted of juvenile and semimature individuals. Clitellate earthworms were found 1 month later and predominated from August (10.5 weeks) to the end of the season. Egg capsules were found from August through October. Egg capsules incubated in the laboratory at 16°C hatched on average after 42 days and produced two juveniles. Juvenile and immature earthworms collected from a quiescent state at the end of summer matured within 1 month when reared in moist soil in the laboratory.     

Responses of co-occurring populations of Dendrobaena octaedra (Lumbricidae) and Cognettia sphagnetorum (Enchytraeidae) to soil pH, moisture and resource addition

Dendrobaena octaedra (Lumbricidae) and Cognettia sphagnetorum (Enchytraeidae) are the two most dominating soil invertebrates in terms of biomass in boreal coniferous forest soils. A microcosm experiment was set up in order to study the influence of pH, moisture and resource addition on D. octaedra and C. sphagnetorum when both species are simultaneously present. Two kinds of coniferous forest humus were used as substrate, pine stand humus (pH 4.2), and spruce stand humus (pH 4.6); in the third treatment the pine stand humus was adjusted with slaked lime (CaOH₂) to the same initial pH as the spruce stand humus. Each substrate was adjusted to water contents of 25%, 42.5% and 60% of WHC (referred to as ‘dry’, ‘moist’ and ‘wet’). In the second part of the experiment, spruce needle litter and birch leaf litter were separately added into the pine stand humus (‘moist’, unlimed) and compared with a control without litter. The microcosms were plastic jars with 75 g (d.m.) of humus, into which 4 specimens of D. octaedra and 70 specimens of C. sphagnetorum were added. D. octaedra showed the highest biomass and C. sphagnetorum the lowest biomass in the spruce stand humus with higher pH. Moisture did not affect earthworms, while C. sphagnetorum thrived best at the highest moisture. Addition of both kinds of litter increased the numbers and biomass of D. octaedra, while on C. sphagnetorum resource addition had little effect. The results help to explain the abundance of these two species in coniferous forests differing in soil acidity, moisture and fertility.     

稻草还田对油菜生长、土壤温度及湿度的影响

采用盆栽和田间试验相结合的方法,研究了不同稻草还田方式及还田量对油菜出苗及苗情和不同稻草覆盖量对油菜生长、养分吸收、产量及土壤水热状况的影响。盆栽试验结果表明,稻草覆盖和翻埋还田对油菜出苗均有抑制作用,出苗率及出苗速率随稻草还田量的增加均有降低趋势,但覆盖条件下油菜出苗所受抑制更加明显。稻草覆盖还田对油菜苗情影响不大,而稻草翻埋还田则对苗期油菜生长有明显的抑制作用。大田试验结果表明,稻草覆盖可以显著提高油菜苗期干物质积累及氮、磷、钾养分吸收的速率,但进入薹期之后油菜生长基本不受稻草覆盖的影响。稻草覆盖对油菜有显著的增产效果,平均增产幅度为18.1%,但不同覆盖量之间差异不显著。另外,稻草覆盖后,土壤温度日变化趋于缓和,土壤含水率明显提高,且随覆盖量增加稻草的保温保水效果更为明显。     

Reproduction and growth of three deep-burrowing earthworms (Lumbricidae) in laboratory culture in order to assess production for soil restoration

Laboratory experiments were conducted to examine the growth and reproduction of three deep-burrowing lumbricids, Aporrectodea longa, Lumbricus terrestris, and Octolasion cyaneum. The reproductive output was recorded as 18.8, 38.0, and 32.3 cocoons per worm per year for A. longa, L. terrestris, and O. cyaneum, respectively. For the same species, maturity was reached at a mean mass of 3.9, 5.0 and 2.4 g, within 3 months from the hatchling stage by L. terrestris and within 4 months by the other two species. The hatching success of cocoons at 15 and 20°C was within the range of 70–80% for each species, except A. longa at the higher temperature, where a viability of 47% was recorded. Twenty percent of viable O. cyaneum cocoons produced twin hatchlings, compared with only one percent for A. longa and L. terrestris. A combination of these results suggests that a complete life-cycle for each species could be achieved within 6 months (L. terrestris and A. longa) or 7–8 months (O. cyaneum). Each species has particular life-cycle strategies that would aid survival and colonisation, under field conditions, if inoculated into restored soils.     

Ability of the earthworms Aporrectodea rosea and Aporrectodea trapezoides to increase plant growth and the foliar concentration of elements in wheat (Triticum aestivum cv. spear) in a sandy loam soil

In a greenhouse study, the ability of the earthworms Aporrectodea rosea and A. trapezoides to influence the foliar concentration of elements and the growth of wheat plants was assessed 27 days after sowing in a sandy loam soil. The presence of A. rosea and A. trapezoides (at densities equivalent to 314 and 471 m^-2, respectively) caused a significant increase in the shoot dry weight of wheat. The presence of A. rosea and A. trapezoides (at densities equivalent to 314 and 157 m^-2, respectively) was also associated with a significant increase in the root dry weight of wheat. The presence of A. rosea caused a significant increase in the foliar concentration of Ca, Cu, K, Mn, N, Na, and P, but did not influence the foliar concentration of Al, B, Fe, Mo, Mg, S, and Zn. The presence of A. trapezoides was associated with a significant increase in the foliar concentration of Al, Ca, Fe, K, Mn, N, and Na, but did not influence the foliar concentration of B, Cu, Mo, Mg, P, S, and Zn. These results demonstrate the potential of A. rosea and A. trapezoides to increase the growth of wheat in a sandy loam soil and suggest that the mechanism by which they increased plant growth was, in part, through increasing the availability and uptake of nutrients from this soil.     

Microcosm investigations into the influence of sheep manure on the behaviour of the geophagous earthworms Aporrectodea trapezoides and Microscolex dubuis

A series of experiments was conducted over 96 h in 240-mm-deep soil microcosms, to assess the effect of the presence and distribution of sheep manure over the soil surface on the vertical and horizontal distribution of burrows and numbers of the earthworms Aporrectodea trapezoides and Microscolex dubius. Within some microcosms the dung was placed on half of the soil surface and this caused aggregation, with over two-thirds of the earthworms being found in the soil directly under the manure. The presence of surface-applied sheep manure caused both species to aggregate in the surface soil. In contrast, without manure, A. trapezoides was evenly distributed throughout the soil profile while M. dubius aggregated in the deeper soil. The pattern of burrow construction was also influenced by the presence of surface manure. In the absence of manure, burrows of both species were evenly distributed through the soil, but in the presence of surface manure M. dubius constructed proportionally more burrows close to the surface. Both species constructed approximately twice the burrow area in the absence than in the presence of surface manure. For both species the daily rate of burrow construction decreased over the experimental period. From these data we inferred that there was more widespread and active foraging behaviour in both species when organic food material was scarce. M. dubius differed from A. trapezoides in that it more strongly concentrated foraging activity in the vicinity of the manure food source.     

The effect of earthworms (Lumbricidae) on nitrogenase activity in soil

Summary Nitrogenase activity associated with earthworms, their faeces and activity in soil was measured by the acetylene reduction technique. A clear increase in nitrogenase activity was found in field-deposited casts of Aporrectodea caliginosa in comparison with surrounding soil, although potential nitrogenase activity was significantly higher in soil than in casts. Nitrogenase activity associated directly with earthworms (Lumbricus rubellus ) was detected, indicating the presence of active N₂-fixing bacteria on the body surface and/or in the gut. Laboratory experiments showed that nitrogenase activity in the casts of L. rubellus was higher than in unmodified soil, and that nitrogenase activity in soil was significantly increased by the burrowing and feeding activity of these worms. This paper discusses the possible causes of these earthworm effects on soil nitrogenase activity and some methodological problems of determining the nitrogenase activity.     

可降解地膜对土壤、温度水分及玉米生长发育的影响

针对普通地膜覆盖导致的农田土壤污染现象,进行了可降解地膜、普通地膜覆盖及露地栽培玉米对比试验,探讨可降解地膜对土壤温度、土壤水分、玉米生长发育、产量及相关农艺性状的影响。结果表明,与露地栽培相比,可降解地膜覆盖能明显提高玉米播种后2个月的地表和地下10 cm的土壤温度,增加玉米播种至大喇叭口期0～20 cm、>20～40 cm的土壤水分含量,使玉米生育进程加快,出苗率和拔节期节根层数及条数增加,玉米不同生育时期株高、叶面积及地上部干物质积累量增加,玉米穗粒数增加9.6%,千粒质量增加20.9%,产量增加35.1%。可降解地膜和普通地膜间差异不显著。研究认为,以可降解地膜替代普通地膜应用于农业生产是可行的。     

定植孔密封方式对土壤水热盐及番茄苗存活率的影响

为了提高河套灌区盐碱地番茄移栽成活率,以番茄定植孔采用土封孔为对照,监测了沙封孔、不封孔株间和根际的土壤水分、土壤盐分、土壤温度及番茄幼苗的生长状况。结果表明,不同处理株间土壤水分、土壤盐分、土壤温度无显著差异。沙封孔、不封孔与土封孔相比,根际0～10cm土层土壤水分降低了11.73%和14.80%,≥10～20cm土层降低了9.60%和13.64%;根际5cm土层最高温度降低了1.2和3.6℃,日均温降低了1.6和2.2℃;根际0～10cm土层土壤盐分降低了19.11%和24.84%,≥10～20cm土层降低了11.48%和19.67%。沙封孔、不封孔番茄幼苗移栽成活率较土封孔提高了20.57%和19.40%。河套灌区盐碱地地膜覆盖栽培番茄,定植时采用沙封孔或不封孔,可降低根际土壤水分及土壤温度,阻碍盐分表聚于根际土壤,提高了番茄幼苗的移栽成活率,该研究为河套灌区盐碱地种植番茄提供指导。     

Effects of temperature regime on the respiratory activity of developmental stages of Lumbricus rubellus (Lumbricidae)

Rates of carbon dioxide production of cocoons, juveniles and matures of Lumbricus rubellus were measured at five constant (2°C, 5°C, 10°C, 15°C, 20°C) and three diurnally fluctuating temperature regimes (0–10°C, 5–15°C, 10–20°C) covering the whole range of temperature conditions experienced by this species in forests in Central Germany. Respiration rates of developmental stages significantly increased with temperature both under constant and fluctuating regimes. Overall, at constant temperature regimes Q₁₀ values for cocoons, juvenile and mature earthworms were 1.6, 2.7 and 2.0, respectively. At fluctuating temperature regimes Q₁₀ values were generally higher with 2.4, 3.6 and 3.5 for cocoons, juvenile and mature earthworms, respectively. At the same mean temperature respiration rates at fluctuating regimes exceeded those at constant regimes in all developmental stages.     

Influence of the earthworm Aporrectodea trapezoides (Lumbricidae) on the colonization of alfalfa (Medicago sativa L.) roots by Rhizobium meliloti L5-30R and the survival of R. meliloti L5-30R in soil

A greenhouse study was performed to examine the ability of the earthworms Microscolex dubius and Aporrectodea trapezoides to transport Rhizobium meliloti L5-30R through soil. When R. meliloti L5-30R was inoculated into either ezi-mulch (a pelleted formulation of cereal-pea straw), oat hay, pea hay, or sheep dung and placed on the soil surface together with either A. trapezoides or M. dubius, >10⁴ colony-forming units (CFU) of R. meliloti L5-30R g^-1 soil were detected at 90 mm soil depth after 18 days. In the absence of earthworms, R. meliloti L5-30R was not detected at 90 mm soil depth after this time. In a second experiment using ezi-mulch as the inoculant material and in the presence of A. trapezoides (in a number equivalent to 471 or 785 m^-2), ca. 10³ CFU of R. meliloti L5-30R per 10 mm of alfalfa root were detected at 0–30, 30–60, and 60–90 mm soil depth after 18 days, while <3 CFU were detected per 10 mm of root in the absence of A. trapezoides. In a third experiment in which R. meliloti L5-30R was distributed evenly through soil at the start of the experiment, A. trapezoides (in a number equivalent to 157, 471, or 785 m^-2) significantly decreased the survival of L5-30R in soil after 40 days of incubation at 15°C, but not after 20 days. The decrease in survival of R. meliloti L5-30R was correlated with the density of A. trapezoides. These results demonstrate that A. trapezoides can increase root colonization of alfalfa by R. meliloti L5-30R, but may also reduce the ability of R. meliloti L5-30R to survive in soil.     

可降解地膜的降解性能及对土壤温度、水分和玉米生长的影响

针对生产中使用普通农用塑料地膜导致农田土壤污染的现状,进行了不同厚度可降解(光、生物降解)地膜、普通地膜和露地栽培玉米对比试验,探讨可降解地膜的降解性能及对土壤水分、温度和玉米生长的影响。结果表明,0.005mm厚可降解地膜的降解速度及强度均优于0.008mm厚膜,二者在覆膜后90d分别达降解5级、4级水平,地膜质量损失率达55.48%、39.99%。两种可降解地膜对土壤水分、温度和玉米生长的影响与普通地膜相当,均使0～20、>20～40cm土壤水分含量、地表及地下10cm土壤温度明显高于露地对照,使玉米出苗率提高,生育进程加快,株高、叶面积和地上部干物质积累量增加;其中0.008mm厚膜覆盖玉米效果优于0.005mm厚膜。研究认为,以可降解地膜替代普通地膜应用于农业生产具有可行性。     

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.     

Growth and survival of Eisenia fetida (Sav.) (Oligochaeta: Lumbricidae) in relation to temperature,moisture and presence of Enchytraeus albidus (Henle) (Enchytraeidae)

Summary The aim of this study was to investigate the environmental requirements of Eisenia fetida (Lumbricidae) and its relation to Enchytraeus albidus (Enchytraeidae). Specimens of Eisenia fetida were cultured at two different temperatures (15°C and 25°C) and moisture conditions (50% and 80%) in presence or absence of Enchytraeus albidus. Eisenia fetida grew best at the higher temperature and amount of moisture. Enchytraeus albidus impaired the growth of Eisenia fetida at the higher temperature. When Enchytraeus albidus was present, the mortality of Eisenia fetida increased.Interaction between Eisenia fetida and Enchytraeus albidus was shown, although the mechanisms remain unclear. These species, coexisting in the same habitat (compost), appear to have slightly different niches: Eisenia fetida favours warmer, moister conditions than Enchytraeus albidus.Dedicated to the late Prof.Dr. M.S. Ghilarov     

Density, biomass and vertical distribution ofAporrectodea caliginosa (Savigny 1826) (Oligochaeta, Lumbricidae) in Benghazi, Libya

Studies were conducted during the period May 1993 to April 1994 on the density, biomass and vertical distribution ofAporrectodea caliginosa, a common species of earthworm found in the Benena farms in Benghazi, Libya. The soil of the farm was a clay loam, pH 7.3, and the organic matter averaged 4.2%. Maximum density and biomass ofA. caliginosa were recorded from October to December 1993 and again during March and April 1994, and minimum values were recorded from May to July 1993. A juvenile dominant population was observed during the summer. This was followed by an increase in subadults and adults during the autumn, winter and spring seasons. A positive correlation of the moisture and an inverse correlation of soil temperature on the density and biomass of these animals were discernible. A. caliginosa mainly confined their activities to the surface organic zone (0–10 cm depth) of the soil for most of the months studied.