Seasonal population dynamics of Eisenia fetida (Savigny, 1826) (Oligochaeta, Lumbricidae) in the field

In order to assess the response of epigeic earthworms to seasonal changes we monitored the population dynamics of Eisenia fetida (Oligochaeta, Lumbricidae) in a manure heap in the field during a year. Earthworms were hand-sorted from five 0.25 x 0.25 x 0.20 m blocks around the heap in November (autumn) 1999 and in January (winter), April (spring) and August (summer) 2000 to determine earthworm population dynamics. Earthworms of each block were classified into different age classes: mature, preclitellate, juvenile, hatchling and cocoon, and afterwards counted and weighed. Seasonality had a strong effect on the density, biomass and reproductive activity of the population. The population of E. fetida was characterized by a high density of individuals and the predominance of mature individuals throughout the year. Maximum density, mating activity and size of cocoons were achieved in spring, but there were not changes in the number of cocoons per mature earthworm throughout the year. Unexpectedly, the smallest cocoons were produced in winter by the largest individuals. These results suggest that E. fetida is able to allocate resources to growth and/or reproduction in response to environmental fluctuations.     

Distribution of aquatic Oligochaeta in the Finnish Lake District

SUMMARY. The distribution of oligochaetes was examined at sampling stations grouped into three types: (1), in eutrophic and dystrophic areas or in areas polluted by pulp-mill wastewaters; (2), slightly eutrophic stations; (3), 'clean' stations unaffected by sewage or other inputs. Limnodrilus hoffmeisteri and Potamothrix hammoniensis were practically the only species in group 1, and the same species accounted for two thirds of the total numbers in group 2, although Peloscolex ferox and some other species also occurred. At the cleanest stations the dominating species were P. ferox, Psammoryctides barbatus, Stylodrilus heringianus and Tubifex tubifex . A factor analysis showed that L. hoffmeisteri and P. hammoniensis were associated with increased nutrients and water colour in the lower part of the epilimnion. In the metalimnion the relationship of S. heringianus and P. ferox to water colour and total nitrogen differed from that of L. hoffmeisteri and P. hammoniensis . Winter oxygen content was important for S. heringianus .     

Ultrastructural features of gametogenesis during the life cycle in Branchiura sowerbyi Beddard (Oligochaeta,Tubificidae)

An ultrastructural analysis of the gametogenetic phases in Branchiura sowerbyi, a tubificid oligochaete, has been accomplished. These phases mostly conform to the usual pattern for the family, however, some interesting peculiarities are pointed out. The regression of sexual apparatus after reproductive period and its regeneration up to a new period of sexual maturity, has been followed throughout the year.     

Effects of lime and artificial acid rain on the enchytraeid (Oligochaeta) fauna in coniferous forest

Effects of lime and artificial rain of varying acidity on the enchytraeid fauna were studied in 4 field experiments in coniferous forest. Artificial rain was applied 5 months per year in quantities of 25 or 50 mm per month and with pH values from about 6 to 2. The rain was produced by mixing groundwater and sulphuric acid. The greatest total abundance of enchytraeids and the greatest population densities of Cognettia sphagnetorum and Mesenchytraeus pelicensis were found in plots supplied rain of pH 6, pH 4 and pH 3. Their abundances were lowered by additional acidification (pH 2.5 and 2) and also by liming. Species like Enchytronia parva and especially Enchytraeus buchholzi and Fredericia parnoniana were mostly stimulated by liming.     

Terrestrial Enchytraeidae (Oligochaeta) and Parergodrilidae (Polychaeta) from Sweden, with description of a new enchytraeid species

Forty-four cnchytraeid species and one parergodrilid species are reportcd from terrestrial habitats in Sweden. Eighteen of the Enchytraeidae are ncw records, including Achueta bifollicula sp.n. Thc new species has 4046 segments; dorsal sctal follicles; follicle-like bodics vcntrally at IV and V; clitellum dorsally interrupted, with fields of hyaline cells; three pairs of primary septa1 glands, and one pair of secondary ones behind 5/6; dorsal blood vessel originating in VII; anteclitellar nephridia in three pairs at 6/7-8/9; long, large spermathecae reaching to X-XII; sperm funnels with a 1ength:width ratio G7:l; seminal vesicles extending through three scgments. The usefulness of the clitellar structure of A chaeta species as a taxonomic critcrion is discussed.     

The histochemistry and ultrastructure of the clitellum of the enchytraeid Lumbricittus rivalis (Oligochaeta: Annelida)

The monostratified clitellar epidermis of Lumbricillus rivalis consists of supporting cells, granular secretory cells, and globular secretory cells, together with the acid mucous cells typical of normal skin in the anterior and posterior transition zones. The secretion of the granular cells is a neutral glycoprotein with low levels of bound lipid, and that of the globular cells is a sulphated acid mucopolysaccharide lacking detectable protein or lipid.
Ultrastructurally the granular cells possess 1 μm, membrane-bound granules of variable electron density suggestive of maturation changes. The formative granules arise from Golgi vesicles and are moderately electron translucent. They contain parallel-aligned, tubule-like inclusions of 14 nm diameter, observed also in the mature regions of the Golgi. Granules in the mid or apical part of the cell show increased electron density, the formative pattern being wholly or partially obscured and the subunit alignment, when discernable, no longer uniform. Moiré fringe patterns are evident in some granules. The patterns and electron density are lost after pronase digestion. The globular cells contain electron translucent, membrane-bound globules of 2–3 μm diameter with finely fibrous contents. They arise from mature Golgi vesicles and are unaffected by pronase treatment.
The results suggest that the granular cells secrete the cocoon wall and the globular cells the material that surrounds the developing embryos, and are compared with published accounts of other microdrile clitella and with those of the clitellum of lumbricid earthworms.     

Freeze tolerance and accumulation of cryoprotectants in the enchytraeid Enchytraeus albidus (Oligochaeta) from Greenland and Europe

The freeze tolerance and accumulation of cryoprotectants was investigated in three geographically different populations of the enchytraeid Enchytraeus albidus (Oligochaeta). E. albidus is widely distributed from the high Arctic to temperate Western Europe. Our results show that E. albidus is freeze tolerant, with freeze tolerance varying extensively between Greenlandic and European populations. Two populations from sub Arctic (Nuuk) and high Arctic Greenland (Zackenberg) survived freezing at −15 °C, whereas only 30% of a German population survived this temperature. When frozen, E. albidus responded by catabolising glycogen to glucose, which likely acted as a cryoprotectant. The average glucose concentrations were similar in the three populations when worms were frozen at −2 °C, approximately 50 μg glucose mg⁻¹ tissue dry weight (DW). At −14 °C the glucose concentrations increased to between 110 and 170 μg mg⁻¹ DW in worms from Greenland. The average glycogen content of worms from Zackenberg and Nuuk were about 300 μg mg⁻¹ DW, but only 230 μg mg⁻¹ DW in worms from Germany showing that not all glycogen was catabolised during the experiment. Nuclear magnetic resonance spectrometry (NMR) was used to screen for other putative cryoprotectants. Proline, glutamine and alanine were up regulated in frozen worms at −2 °C but only in relatively small concentrations suggesting that they were of little significance for freeze survival. The present study confirms earlier reports that freeze tolerant enchytraeids, like other freeze tolerant oligochaete earthworms, accumulate high concentrations of glucose as a primary cryoprotectant.     

Laboratory studies on the life-histories of four enchytraeid worms (Oligochaeta) which inhabit sewage percolating filters

The life-histories of four enchytraeid worms, Lumbricillus rivalis, Enchy-traeus coronatus, E. buchholzi, and E. albidus which occur in sewage percolating filters, were studied under laboratory conditions at 8 , 15 and 20°C. The number of ova per cocoon varied from 0 to 50 (L. rivalis), 0 to 33 (E. coronatus), 1 to 9 (E. buchholzi) and 0 to 22 (E. albidus). The mean number of ova per cocoon was highest at 15°C for all species except E. coronatus which had a highest mean value at 8°C. The number of ova in cocoons was correlated with cocoon length (P < 0.001) for all species. Cocoon production usually increased with temperature ranging from 0.8 cocoons per adult per week at 8°C to 2.0 at 20°C for L. rivalis, and from 1–4 to about 2.6 for E. coronatus and E. buchholzi. The total number of ova produced by each E. coronatus (350 at 8°C to 550 at 20°C) was similar to that produced by each L. rivalis (600 at 8°C to 350 at 20°C) and was about five times greater than the total numbers produced by the other two species. Cocoon and ova production and the number of ova per cocoon varied with the age of the adult, usually reaching a peak soon after maturity. Hatching success was low and generally 40–50 % of ova failed to develop; subsequent mortality among immature worms was about 10–20%. Growth was more rapid at the higher temperatures; L. rivalis matured in about 26 days at 20°C, the clitellum forming when the worm was 13–14 mm long; data for the other species are 13 days and 5–6 mm (E. coronatus); 16 days and 3–4 mm (E. buchholzi); 28 days and 13–14 mm (E. albidus). The maturation period at 8°C was at least twice that at 20°C. The generation period (cocoon to cocoon) was about a month at 20°C for all species except E. albidus (2 months), but as some species had longer reproductive periods than others the actual number of generations per year was highest in E. buchholzi, 7.0 per year, and lowest in E. albidus, about 3.3 per year, At 8°C all four species had between 1.4 and 2.8 generations a year. A comparison of expected and observed population densities of L. rivalis and E. coronatus in a sewage percolating filter showed that neither achieved values approaching their potential summer densities although ample food was apparently available. Of the four species studied only E. buchholzi produced viable ova without pairing.     

Observations on the population dynamics of Potamothrix hammoniensis (Michaelsen, 1901) (Tubificidae,Oligochaeta) in Lake Isacova in the Danube Delta

The Danube Delta is one of the widest wetland systems in Europe and Lake Isacova is one of the biggest lakes in the Delta. The oligochaete community in the Lake comprised 52% of the total benthic fauna and Potamothrix hammoniensis (Michaelsen, 1901) (Tubificidae, Oligochaeta) represented more than 90% of the oligochaetes sampled. During September 1991 – August 1994, the proportion of ovigerous individuals of P. hammoniensis represented less than 20% of the total population, with localised differences in the Lake occurring over time. Breeding and cocoon production in the study period started in March of the year, when the water temperature was about 7 °C, and lasted until May 1992 and June 1993, respectively. The newly hatched individuals reached the highest proportion in May. Although large individuals were found again by the end of September (1991) and the beginning of October (1993), no spermatozeugmata in their spermathecae or cocoons were observed during November and October, respectively. Member of this cohort bred and started cocoon deposition during the following spring, after about 11 months of maturation (from April to March). A second, less intensive period of breeding was recorded during August 1992 and this lasted until the beginning of September 1993. However, cocoons from this latter period did not show embryonic growth and hatching until the autumn floods bring oxygenated water to the bottom level. This resulted in a period of rapid development followed by the mass hatching of cocoons. These specimens needed a maximum of 9 month (from October to July) to reach sexual maturity.     

Life cycle of the earthworm Octodrilus complanatus (Oligochaeta, Lumbricidae)

The earthworm Octodrilus complanatus (Dugès, 1828) is a large-sized species often found in grasslands and pastures, with a wide distribution area in Europe and North Africa. In order to determine the characteristics of its life cycle, earthworms of this species were cultured from hatching until they reached maturity. Development rates, cocoon production, and incubation period were recorded under controlled conditions. On average, individuals of O. complanatus reared in isolation from hatching reached maturity at day 150 and body weight at maturity ranged between 6 and 8 g. The cocoon incubation period was 66 days, and only one individual hatched from each cocoon, with a mean hatchability of 55%. Reproductive rate showed by O. complanatus after mating was of 52 cocoons per earthworm per year. The results showed that the life cycle of O. complanatus was characterized by both low growth rates and low fecundity. Specific features in life history traits of O. complanatus were compared with those of species of the genus Octodrilus and other related earthworms.     

The effect of temperature on the life cycle, growth and fecundity of Branchiura sowerbyi (Oligochaeta: Tubificidae)

Branchiura sowerbyi Beddard was probably introduced to Britain with exotic plants consigned to botanic gardens. The British populations occur mainly in artificially warmed habitats such as lily ponds in botanic gardens and power station effluents but a few are found at natural temperatures.
Branchiura completed its life cycle in about a year both in the river Avon, Wiltshire, at natural temperatures and in the warm effluent from a power station discharging into the river Thames. In a worm culture maintained at natural temperatures the rate of cocoon production reached a peak in summer and was closely related to temperature.
In the laboratory it was found that the optimum temperature for cocoon laying in mature worms, also growth in sexually immature worms, was near 25°C. The optimum temperature for growth in sexually mature worms, however, was lower (10°C in the Avon population and 15°C in the Thames population). It followed that at 20°–25°C the growth of sexually mature worms was probably depressed by the high rate of cocoon production.     

Propappidae and aquatic Enchytraeidae (Oligochaeta) from the farthest southeast of Russia

Two mountain streams north and northeast of Vladivostok were studied in 1983–1984. In the Komarovka Stream, a great variety of Mesenchytraeus occurred in the spring zone, among them M. crenobius sp. n. with giant setae in the ventral bundles of VI–VIII and M. vshivkovae sp. n. with numerous spermathecal diverticula. In the stream zone of the Frolovka Stream Cernosvitoviella pensau sp. n., characterized by a complicated penial bulb and a very long spermathecal ampulla, was the dominant oligochaete. Propappus arhynchotus Sokolskaya was the second most abundant oligochaete in this stream. It proved to be a true member of this genus and not synonymous with any other species. These are the first records of aquatic enchytraeids for the Primorski Region of Russia.     

Spatial patterns and population dynamics of plant-associated microcrustacea (Cladocera) in an English shallow lake (Little Mere, Cheshire)

Little Mere (Cheshire) is a small (2.7 ha) and shallow (average depth 0.7 m) fertile lake in Cheshire, UK. Nymphaeids cover almost 40 % of its entire surface during the growing season (April to October) and practically all the rest is covered by a mixed community of submerged plants. The lake was intensively sampled for plant-associated Cladocera and zooplankters from April 1998-April 2000. Samples were collected at five sites across the lake, three of them located within lily beds, the other two over submerged plant beds of mixed composition. Specific sampling techniques were developed for floating lily leaves, petioles, submerged plants and water. Significant horizontal differences were identified for most cladoceran species, both open-water and plant-associated, for chydorid periphyton scrapers and for filter-feeders. Daphnia hyalina (L.) and Ceriodaphnia sp were significantly more abundant in lily beds than in more open water in both growing seasons, suggesting lily beds are an effective refuge against fish predation. Size-structure and egg-ratio data support this contention. Egg-ratio models were examined for Daphnia hyalina and Simocephalus vetulus (O.F. Müller), a plant-associated cladoceran. The fertility of S. vetulus in lily beds was generally high throughout growing seasons. The construction of egg-ratio models for this species was hampered by their generally very patchy distributions.     

The life cycle, population dynamics, growth and secondary production of the snail Xeropicta arenosa Ziegler (Gastropoda: Pulmonata) in northern Greece

The life cycle, population dynamics and secondary production of the xerothermophilic land snail Xeropicta arenosa were studied in northern Greece. The study of biology and the demographic analysis of the populations of X. arenosa revealed that it is a semelparous annual species. Reproduction takes place in autumn and the majority of the adults die after egglaying. Growth is continuous throughout the year and no seasonal fluctuations occur. Estimation of annual secondary production with the Hynes' size frequency method revealed a mean standing crop ( aa ) of 0.075 g m^-2 year^-1 and a productivity ( P ) of 0.267+ 0.043 gm^-2 year^-1. The annual turnover ratio ( P/ ) was equal to 3.56.     

Preliminary study of the aquatic Oligochaeta of the Changjiang (Yangtze) River

The aquatic oligochaetes in mainstream of the Changjiang River have not been studied previously. Since 1983, benthic samples have been collected mainly from mid-upper reach of the river. Altogether 19 species belonging to 4 families of the Oligochaeta were found. Among them, Nais inflata is the commonest species throughout the river, whereas Nais sp., Arcteonais lomondi, Limnodrilus silvani, Rhyacodrilus riabuschinskii and Telmatodrilus sp. ae recorded from China for the first time.Quantitatively, the aquatic oligochaetes constitute the majority of zoobenthos in the mainstream. In 1984, the standing crop of the Tubificidae was 231–343 ind·m^-2 in density and 0.73–0.97 g·m^-2 (wet weight) in biomass, and that of the Naididae was 21–4982 ind·m^-2 and 0.003–1.75 g·m^-2 (wet weight) respectively.     

Growth,reproductive biology and life cycle of the vermicomposting earthworm, Perionyx ceylanensis Mich. (Oligochaeta: Megascolecidae)

In the present study, an attempt has been made to study the growth, reproduction and life cycle of the earthworm, Perionyx ceylanensis Mich. in cowdung for the period of 340 days. Results showed that the overall mean growth rate was 1.79, 1.57 and 1.34 mg/worm/day respectively for the worms cultured singly, in batches of four and eight. Cocoon production rate was found between 0.85 and 0.94 cocoons/worm/day and the hatching success between 74.67% and 82.67%. The majority of the cocoons (95.16–96.77%) hatched only one hatchling. Worms raised singly also produced viable cocoons indicating that P. ceylanensis reproduce parthenogenetically. The life cycle of the worms cultured singly was ±57 days and it was ±50 days for the worms cultured in batches of four and eight. There is a vast scope to utilize P. ceylanensis for vermiculture practices due to short period of life cycle.     

Observations on the population dynamics, life cycle and ecology of the plant parasitic nematode Rotylenchus robustus

In a forest nursery growing Sitka spruce Rotylenchus robustus was most numerous at a soil depth of 10–19 cm, the vertical distribution being associated with that of the tree roots. Although no consistent seasonal fluctuations in nematode numbers were obvious there was a seasonal pattern in the relative frequency with which adult and larval nematodes were recovered.
Experimentally, the life cycle of R. robustus was completed in 14–18 months depending upon the time of year when nematodes were added to the trees. Soil moisture content of less than 8.7% (pF 3.8) restricted the movement of 50% of the population. The overall mean basal temperature for the completion of the life cycle of R. robustus was calculated to be 6.5°C.
Soil temperature was considered to be the dominant factor controlling the rate of development under field conditions in Scotland; only occasionally did soil moisture become limiting.     

A parthenogenetic life cycle in a population of Canthocamptus Staphylinus (Copepoda,Harpacticoida)

The harpacticoid copepod Canthocamptus staphylinus (Jurine) was shown to reproduce parthenogenetically in an oligotrophic Finnish lake. The population was univoltine with peaks of egg production in winter and early spring. Young from both peaks became adults in spring and aestivated as cysts. Laboratory experiments suggested that either high temperature (12 °C or more) or long days will inhibit egg production, but that both factors together are needed to induce encystment. Embryonic survival was low, on the average only 60%. The population was limited by low food levels in winter, and it is suggested that food shortage prevents the species from colonizing the profundal zone of oligotrophic lakes.     

Distribution and population dynamics of Porphyra (Bangiales, Rhodophyta) in the southern Western Cape, South Africa

Although Porphyra is commercially farmed in many countries, in South Africa only small harvests of wild populations for sale as nori have been carried out. The discovery that Porphyra improves growth of South African abalone (Haliotis midae) farmed inland-based tanks has led to increased pressure to harvest wild populations. This paper reports on a survey of the distribution and seasonality of Porphyra in the southern Western Cape. Porphyrawas present at all sites surveyed, and showed considerable temporal variation. A significant amount of the Porphyra present is in reserves and therefore protected from harvesting. Close rexamination of one site revealed seasonal populations of Porphyra that occupied different niches dependent on season. Recruitment peaked in spring and autumn, leading to dense summer and winter populations. Summer populations generally grew lower in the eulittoral than winter populations. No pattern in the mortality of larger thalli wasde tected, though sporeling mortality was high following recruitment peaks. Although it seems that most sites in the southern Western Cape are suitable for harvesting, the taxonomy of the genus in the region urgently needs revision if populations are to be appropriately managed. This revised version was published online in August 2006 with corrections to the Cover Date.