Seasonal variations in growth and reproduction of the earthworm Perionyx excavatus (Oligochaeta: Megascolecidae)

 One-week-old juveniles of Perionyx excavatus were cultured in urine-free cow manure for 16 weeks in three different seasons to find out the influence of environmental factors on growth and reproduction. The biomass of worms increased gradually until the 16th week in all seasons. There was no significant (F=2.51) variation in biomass between the three seasons; however, the growth rate varied significantly (F=50.49, P<0.01). During the pre-clitellar period, the growth rate (mg/g) increased steeply (1013.17±41.33 in the monsoon season, 728.64±15.5 in winter and 463.16±22.96 in summer) until the worms attained maturity (5–8 weeks) and from then onwards gradually decreased after the commencement of cocoon production. Worms attained maturity when the mean weight of each worm ranged from 167.46 mg to 197.13 mg. The cumulative number of cocoons varied significantly (F=26.07, P<0.01) with season. Similarly, the rate of cocoon production also varied significantly (F=62.06, P<0.01). The average rate of cocoon production was 7.23, 0.99 and 0.53/worm/week in the monsoon season, winter and summer, respectively. By week 16, the ratio of weight gain to the initial weight of 1-week-old juveniles was 1 : 43.13 in the monsoon season, 1 : 30.49 in the winter and 1 : 23.69 in the summer. The maximum mass gain, growth rate, and cocoon production and earliest attainment of maturity were recorded during the monsoon season, and were attributed to the lowest range of fluctuating temperatures and high humidity compared with during winter and summer. The growth rate of worms in all seasons was inversely proportional to the number of cocoons produced. Further, the higher the rate of cocoon production, the lower the weight of individual cocoons. Received: 1 July 1997     

Assessment of recovery of population, biomass and reproduction of the earthworm Drawida willsi following the application of malathion under field conditions

 The recovery of population, biomass and reproduction of a dominant crop field earthworm (Drawida willsi, Michaelsen) was assessed following the application of two recommended agricultural doses of malathion (2.2 mg kg^–1– single dose and 4.4 mg kg^–1– double dose) in a rice field agroecosystem for a period of 105 days. The average worm population and biomass showed 12% and 21% decline, respectively, in plots treated with single and double doses of malathion, compared with control. The peak of worm population was observed after 60 days in control (plot I) and this peak value was shifted by 15 days (i.e. observed after 75 days) in malathion-treated plots (plots II and III) indicating delay in growth and maturation of the worms following the application of malathion. Inhibition in the total number of cocoons produced, and thus in the rate of reproduction, was also observed in malathion-treated plots. The peak rate of reproduction (1.71 cocoons adult^–1) was observed after 60 days in control plots and after 75 days (1.57 and 1.40 cocoons adult^–1) in single and double dose malathion-treated plots, respectively. One-way analysis of variance showed significant differences in the population size and biomass of D. willsi up to 75 days, in rate of reproduction up to 90 days, and no difference thereafter. This indicates that, under field conditions, D. willsi worms took about 75–90 days after application of malathion to resume normal population, biomass and reproduction, and hence we suggest that a second application of malathion in single and double agricultural doses should be avoided before this time. Received: 7 September 1999     

The effects of low lead levels on the growth and reproduction of the African earthworm Eudrilus eugeniae (Oligochaeta)

 The effects of sublethal concentrations of lead nitrate on the growth and reproduction of the African composting earthworm species, Eudrilus eugeniae, was studied by exposing worms in an organic substrate to lead-nitrate-contaminated food over a period of 76 days. The results revealed that growth was initially affected negatively by the presence of lead, while the maturation rate and cocoon production were not affected. In agreement with other studies on Eisenia fetida, cocoon viability was affected negatively by lead, making this a sensitive toxicity endpoint. Received: 14 October 1998     

The influence of feeding patterns on growth and reproduction of the vermicomposting earthworm Eisenia fetida (Oligochaeta)

Summary Observations in outdoor culture beds indicated that food availability and frequency of feeding influenced worm density. In order to study the role of feeding status on the growth and reproduction of Eisenia fetida, pre-clitellate specimens were kept for 180 days under controlled conditions. A control group was regularly provided with freshly produced cattle manure while the experimental groups were fed only sporadically and at some stage even starved by removing some of the substrate. Worm growth, maturation, and cocoon production were monitored. The role of food availability was highlighted by this study. Both worm growth and cocoon production were correlated closely with the feeding pattern that was followed. This study also showed that freshly produced, urine-free, cattle manure proved an excellent food source provided it was added in such a way as to prevent the development of anaerobic conditions. Regular feeding gave high growth and reproduction rates.     

A defined medium for the study of growth and reproduction of the earthworm Eisenia fetida (Oligochaeta)

Summary The feeding requirements of earthworms have not been fully established. A vermiculite matrix was used to determine the dietary effects of various organic substances. Vitamins, DNA, and humic acid as additives improved growth and allowed reproduction of Eisenia fetida in a cellulose-based medium. Mature worms were not able to adapt to the medium and 20-day-old worms were used. Rates of growth and cocoon production were slightly less in a defined medium (7% organic content) than in a cow-manure control medium (70% organic content). The medium is likely to prove useful as a reference medium for obtaining biological information.     

Growth and cocoon production by the earthworm Metaphire houletti (Oligochaeta) in different food sources

 Quantitative observations on the biology of Metaphire houletti were made in cow and horse manure and oak litter in laboratory conditions over a period of 240 days. The study revealed that copulation is not a prerequisite for production of viable cocoons, indicating that M. houletti may be parthenogenetic. The mean growth rate was 2.86 mg worm^–1 day^–1 reaching sexual maturity at 45 days and producing 0.015 cocoons worm^–1 day^–1 (kept singly) and 2.82 mg worm^–1 day^–1, reaching sexual maturity at 45 days and producing 0.03 cocoons worm^–1 day^–1 (kept in batches) in cow manure; 4.08 mg worm^–1 day^–1, reaching sexual maturity at 46 days and producing 0.02 cocoons worm^–1 day^–1 (kept singly) and 2.97 mg worm^–1 day^–1, reaching sexual maturity at 45 days and producing 0.016 cocoons worm^–1 day^–1 (kept in batches) in horse manure; 3.73 mg worm^–1 day^–1 reaching sexual maturity at 45 day and producing 0.023 cocoons worm^–1 day^–1 (kept singly) and 2.73 mg worm^–1 day^–1, reaching sexual maturity at 47 days and producing 0.028 cocoons worm^–1 day^–1 (kept in batches) in oak litter. After an incubation period of 31.9±1.2 days 82% of the cocoons hatched with a mean of 1.12±0.06 hatchlings per cocoon. The earthworms reared in batches did not demonstrate any advantage over those reared singly in all substrates. Higher growth rates were observed in earthworms raised singly than those raised in batches in all substrates. Received: 30 April 1998     

Uptake and toxicity of copper and zinc for the African earthworm,Eudrilus eugeniae (Oligochaeta)

Growth rate change in earthworms is considered to be a suitable endpoint when determining sublethal effects. In this study we evaluated growth and maturation in the vermicomposting earthworm speciesEudrilus eugeniae as marker of sublethal toxicity of copper and zinc. We also compared routes of uptake. Apart from exposing worms experimentally for 73 days to contaminated food, a series of contact filter paper tests was also performed to determine LD₅₀ for copper and zinc. Both copper and zinc at sublethal concentrations affected growth and maturation in worms exposed to contaminated food. These worms had a copper content of 34.5 g g^–1 after 73 days and a zinc content of 184.9 g g^–1, showing a differential uptake. Copper was more toxic than zinc. Also in the contact test worms did take up more zinc than copper and the LD₅₀ (48 h) for copper was 0.011 mg cm^–2 and for zinc 0.066 mg cm^–2, which translated to body burdens of 6 g g^–1 for copper and 131 g g^–1 for zinc. Indications were that a regulatory mechanism existed for both metals. Both metals were taken up through the body wall at a relatively fast rate. This study indicated that the skin was the major route of metal uptake. This study also showed a poor relation between the two types of tests for purposes of evaluating lethality of zinc and copper.     

Rainfall effects on erosion of earthworm casts and phosphorus transfers by water runoff

 We investigated whether, under a temperate climate and in a maize crop, earthworm casts could contribute to soil erosion and further favour the exportation of phosphorus by runoff waters. Recording of casts was made in compacted (wheel-tracks) and non-compacted inter-rows, for a 2-month period in spring. To assess the rainfall impact on cast evolution, half of the observation sites were protected against rain splash by a nylon mesh placed above the soil surface. The water runoff was collected and analysed for sediment contents and phosphorus concentration. The mean annual production of surface casts was calculated to be 34 kg (dry weight) year^–1 kg^–1 earthworm (fresh weight). Synchronization between cast erosion and rainfall events was shown under natural conditions (unprotected sites). The erosion rate was 4 times greater over rainy periods than dry ones, reaching 80% of cast numbers. It appeared that not the runoff effect but the splash effect, due to the kinetics of the drops, disrupted casts. Newly formed casts disappeared first, with the erosion rate decreasing twofold for casts more than 10 days old. Cast erosion and runoff, as well as worm casting activity, were greater under compacted sites than under non-compacted sites, indicating an influence of earthworms on soil erosion from compacted soils. The total phosphorus content was similar in casts and uningested soil (0.80 mg phosphorus g^–1). Potential phosphorus losses from cast erosion was calculated to reach 25–49 mg phosphorus m^–2 per rainfall event depending on soil compaction. The amounts of particulate phosphorus recovered in water runoff after each rainfall event varied from 1 mg to 11 mg phosphorus. These results are compared and discussed. Received: 20 October 1998     

Effects of trefoil cover crop and earthworm inoculation on maize crop and soil organisms in Reunion Island

 Traditional tree fallows have been abandoned on the western coast of the Reunion Island because of the increasing need for cultivated land. Soil fertility is no longer restored and crop yields have decreased drastically. The leguminous plant, Lotus uliginosus (trefoil), used as a cover crop, has made possible the control of erosion, the restoration of soil macrofauna, especially earthworms, and the increase in crop yields. When trefoil was associated with earthworms (Amynthas corticis), the densities of maize, the yields of maize stalk and dry matter, the yield of trefoil fodder dry matter, and the biomass and respiratory activity of soil microflora were considerably increased. The combined effects of their association led to a significant decrease in populations of the plant-parasitic nematode, Pratylenchus vulnus, in maize roots, and in the population of borers. Some soil chemical features were modified. Received: 10 September 1997     

Effect of coffee residues on growth and reproduction of Hyperiodrilus africanus (Oligochaeta, Eudrilidae) in Ivory Coast

Hyperiodrilus africanus (Beddard) is a 12-cm to 16-cm-long earthworm, which is widely distributed in West and Central Africa. It lives in the upper 10–20 cm of the soil, and feeds on a mixture of soil and above-ground litter. Cocoons obtained in the laboratory hatched on average 17 days after deposition and produced two juveniles on average. Paired individuals fed soil amended with 2% coffee residues grew significantly (P<0.05) faster than those in the control soil. Daily individual weight increments were respectively 6.1 mg worm^–1 day^–1 and 1.0 mg worm^–1 day^–1 in supplemented and control soil. The generation time was short, and cocoon production reached 9.6 month^–1 (i.e. 115 cocoons adult^–1 year^–1). When H. africanus collected from the field were raised in the laboratory, they grew slowly, laid fewer cocoons and mortality was high. Demographic parameters indicated an improvement when H. africanus were raised in batches rather than individually. Mating enhanced cocoon production although parthenogenesis was possible. Received: 4 April 1997     

Moisture preferences,growth and reproduction of the compost worm Eisenia fetida (Oligochaeta)

Summary Although moisture requirements of Eisenia fetida are considered to be generally known, very few in-depth studies have been performed. The moisture preferences of juveniles as well as adults of this species were determined in moisture towers containing cow manure with a particle size bigger than 500 and smaller than 1000 m and an organic matter content of 54%. The clitellate worms showed a wider preference range than the juveniles but deposited most of their cocoons within the narrow range of 65%–70% moisture — a moisture range preferred by 80% of the juveniles. Growth studies of newly hatched worms at various moisture levels over a period of 90 days showed that growth rate and final biomass were influenced by moisture availability with a very low rate of growth at low as well as at high moisture levels. The change in biomass of clitellate specimens as well as their consecutively produced cocoons were monitored at different moisture levels. A clear relationship between mass of cocoons and biomass of the producing worms was established at presumably favourable moisture conditions. Furthermore, the relation between cocoon mass and the production rate of cocoons was also determined, which showed that cocoon mass increased concomitantly with increase in cocoon production. The study also showed that clitellum development was increased at moisture levels above 64%. The onset of cocoon production was, however, earliest at slightly lower moisture levels, from which it is concluded that the optimal moisture level for growth and development of clitella are not necessarily optimal for the production of cocoons.Dedicated to the late Prof. Dr. M.S. Ghilarov     

Effect of climate, soil type and earthworm activity on nitrogen transfer from a nitrogen-15-labelled decomposing material under field conditions

 N transfer from ¹⁵N-labelled decomposing material into the microbial biomass and inorganic N forms was studied for more than 2 years at three experimental sites differing in climatic conditions and earthworm abundance. The ¹⁵N-labelled decomposing material was mixed with low-elevation soil (LES), mid-elevation soil (MES) and high-elevation soil (HES). The amended soils were put into two kinds of plastic cylinders closed on both sides with nets preventing (0.1 cm mesh) and allowing (0.5 cm mesh) access by earthworms, and were buried in soil (20 cm depth) to monitor the transfer of N from the ¹⁵N-labelled decomposing material. Climate and soil type play an important role in the release of N from decomposing material. LES transplanted to more humid sites (mid- and high-elevation sites) showed an increase in most of its biological activities (N atom % excess, and microbial biomass C and N). Furthermore, LES was a sandy soil in which the ¹⁵N-labelled decomposing material was less bound than in MES and HES, which contained more silt and clay. This resulted in faster organic matter turnover when climatic conditions were favourable. The presence of earthworms greatly increased the quantity of inorganic N (mainly NH₄ ⁺) in the soils and enhanced the release of N from the ¹⁵N-labelled decomposing material and the native organic matter, compared to soil without earthworms. Received: 21 January 1999     

幼虫密度对小菜蛾生长发育和繁殖的影响

为明确小菜蛾幼虫密度对其种群增长的影响,在室内条件下[温度(22±1)℃,RH70%~80%,光周期12L∶12D]对不同幼虫密度[1头.皿1、10头.皿1、20头.皿1、30头.皿1、40头.皿1(直径为9 cm)]饲养的小菜蛾生长发育及繁殖进行了研究。结果表明,幼虫密度对小菜蛾幼虫、蛹的发育历期和存活率、蛹重及成虫生殖等有显著影响。幼虫和蛹历期均以10头.皿1处理最短,且随幼虫密度增加而显著延长。幼虫至蛹的存活率以10头.皿1、20头.皿1处理较高,显著高于1头.皿1和40头.皿1处理。蛹重以1头.皿1处理最重,并随幼虫密度的增加而显著变轻。不同幼虫密度下羽化的成虫畸形率差异显著,以10头.皿1处理最低,其余随幼虫密度的增加而升高。成虫产卵量以1头.皿1处理最高,随着幼虫密度增加,产卵量逐渐下降。雌、雄蛾寿命分别以40头.皿1、10头.皿1处理最长,密度过高时或过低雌雄蛾寿命显著缩短。生命表分析表明,幼虫密度对小菜蛾种群增长指数影响显著,以10头.皿1处理种群增长指数最高,幼虫密度过高或过低,种群增长指数下降。因此,幼虫密度是影响小菜蛾种群增长的重要因子之一。     

Effects of earthworm density on growth, development, and reproduction in Lumbricus rubellus (Hoffm.) and possible consequences for the intrinsic rate of population increase

In this paper, the influence of earthworm density is assessed on the life-history parameters: growth, development, reproduction, and survival of Lumbricus rubellus (Hoffm.). Density ranges from two to nine earthworms in 1-l containers, corresponding to field densities of 300-1350 earthworms m⁻². Earthworms were kept under optimal laboratory conditions, with a surplus of food. The results show that at high earthworm density, individual growth is retarded, maturation delayed and cocoon production decreased, even when food is optimal. The integration of these results into a Dynamic Energy Budget model suggests that the food intake of individual earthworms is lower at the higher tested densities. This lower food intake is most probably driven by competition for space at the higher densities. The lower food intake results in a lower intrinsic rate of population increase which is negative at the highest tested density.     

Beneficial effects of blue-green algae and Azolla, excluding supplying nitrogen, on wetland rice fields: a review

 The role of blue-green algae (BGA) and Azolla in supplying N to rice fields is well documented. In addition, they also bring about, directly or indirectly, a number of changes in the physical, chemical and biological properties of the soil and soil-water interface in rice fields. For example, BGA liberate extracellular organic compounds and photosynthetic O₂ during their growth, while Azolla prevent a rise in the pH, reduce water temperature, curb NH₃ volatilisation and suppress weeds; and both of them contribute biomass. On decomposing, they influence the redox activity and result in the formation of different organic acids in soil. All such changes brought about by BGA and Azolla in soil may ultimately influence plant-available nutrients and also soil characteristics. An attempt has been made in this review to highlight these effects of BGA and Azolla in rice fields and discuss their possible implications relating to management and productivity of rice-field systems. Received: 4 February 1998     

Effects of yeast on the growth and reproduction of the saprophagous millipedePolydesmus angustus (Diplopoda, Polydesmidae)

The millipedePolydesmus angustus was reared in the laboratory from hatching to maturity, reproduction and death. Two food types were used: dead leaves alone or dead leaves supplemented monthly with dry food yeast (Saccharomyces cerevisiae) at a rate not exceeding 5% of leaf dry weight. Growth, survival, adult live weight and fertility were compared between females reared on the two diets. Although the species was able to complete its life cycle on dead leaves alone, several parameters were strongly affected by the addition of yeast: growth was significantly faster, adult females became significantly larger and there was a 4.3-fold increase in fertility. Only survival was unaltered by the addition of yeast. The comparison between these laboratory results and field data on female fertility and live weight suggests that the natural diet of millipedes includes foods of higher quality than dead leaves. Possible sources of high-quality food in natural conditions are discussed.     

The effect of soil liming on shoot development, root growth, and cluster root activity of white lupin

 The effects of a limed soil upon root and shoot growth of white lupin (Lupinus albus L.) were investigated using soil tubes and pots. After 75 days in the soil tubes, the combined taproot and lateral root dry weight in limed soil (2.5% CaO w/w) was significantly less than in neutral pH soil (by 57%). However, the dry weight and numbers of cluster roots remained comparable between the treatments, demonstrating for the first time that the cluster roots respond differently to the rest of the root system. Cluster roots accounted for 17% of the total root biomass in neutral soil, increasing significantly to over 30% in limed soil. When grown for 43 days in pots containing soil with different additions of lime (0.5–2.5% CaO w/w), soil citrate concentrations were higher than in the neutral pH soil treatment in all except the 2.5% lime treatment, in which they were lower. In both experiments, shoot dry weights were lower in the presence of the limed soil compared with those in the neutral pH soil. Although a reduction in shoot dry weight was not apparent at 21 days in the limed-soil tubes, the initiation of fewer mainstem leaf primordia indicated a slower shoot development than occurred in the neutral soil. Plants grown in the limed-soil tubes showed leaf yellowing and some chlorosis within 9 days. At the final harvest, the shoot phosphorus and manganese concentrations were significantly lower in plants grown in limed soil compared with those in the neutral pH soil, whereas the concentration of calcium was higher. Received: 11 October 1999     

Soil organic carbon stocks, storage profile and microbial biomass under different crop management systems in a tropical agricultural ecosystem

 We investigated the soil organic C and N stocks, storage profiles and microbial biomass as influenced by different crop management systems in a tropical agricultural ecosystem. The different crop management systems significantly affected the C and N stocks and microbial biomass C and N at different soil depths. Amongst the systems evaluated, the rice-wheat system maintained a higher soil organic C content. Inclusion of legumes in the system improved the soil organic matter level and also soil microbial biomass activity, vital for the nutrient turnover and long-term productivity of the soil. Irrespective of the cropping system, approximately 58.4%, 25.7% and 15.9% of the C was distributed in 0–15, 15–30 and 30–60 cm depths, respectively. Received: 10 October 1999     

Life cycle and reproductive traits of the earthworm Aporrectodea trapezoides (Dugès, 1828) in laboratory cultures

The parthenogenetic earthworm Aporrectodea trapezoides (Dugès, 1828) is widely distributed all over the world due to European agricultural practices. In order to provide baseline life cycle data, cocoons were obtained from field-collected individuals and their features and viability, incubation period, number of hatchlings and mortality rate were recorded. Singleton and twin earthworms from this first experiment were cultured from hatching during a 490-day period under controlled conditions with biomass, survival, reproductive condition and cocoon production recorded at intervals of 15 days. On average, individuals of isolated-reared A. trapezoides reached maturity at day 153 and body weight at maturity was approximately 1 g. In order to record reproductive traits and differences between field-collected and laboratory-reared individuals, 40 microcosms with an isolated earthworm (20 with field-collected individuals and 20 with laboratory-reared ones) and 40 containing groups of three (20 with each type of individual) were maintained during a complete year under controlled conditions. The amount of soil per individual was the same in both types of microcosm. Both the individuals kept in isolation and those cultured in groups produced cocoons, hence completely proving the obligatory parthenogenetic reproduction in this species without copulation or need of any physical-chemical stimulus. In general, isolated earthworms produced a significantly higher number of cocoons than those in groups of three, and the same was recorded for laboratory-reared earthworms when compared with field-collected ones. This study highlights the importance of knowing the life cycle and reproductive traits of one possible key species in soil management due to its vast distribution and high density in soils, and the species’ highly recommended use in applied studies because of its ease of culture.