Spermiogenesis in Bythonomus lemani and the phylogenetic position of the Lumbriculidae (Oligochaeta,Annelida)

Spermiohistogenesis and spermatozoal morphology of the lumbriculid Bythonomus lemani have been investigated by means of electron microscopy. Though spermiohistogenetic events follow the general microdrile pattern, some features are peculiar: chromatin is clumped in the first stages and its condensation is very irregular, as in Eisenia. Manchette geometry is similar to that of hirudineans, as the microtubules are helically arranged from the very early stages. A number of mitochondria are always present in the collar region.The mature sperm also departs from the microdrile model and is more similar to the megadrile one in (1) the strong withdrawal of the base of the acrosome vesicle into the acrosome tube; (2) the apparent development of rudimentary connectives. On the other hand, some features seem to be unique among oligochaetes, including the inclined longitudinal axis of the axial rod and tilted anterior end of the nucleus.Spermatologically, lumbriculids may be interpreted as advanced microdriles which may be descended from a stock which also gave rise to Haplotaxidae, Moniligastridae, and the true Opisthopores.     

Seasonal dynamics and emergence pattern of a tropical earthworm,Drawida calebi (Oligochaeta)

Cocoon morphology (shape, size, colour and ornamentation), hatching, and seasonal dynamics of cocoon and juveniles of D. calebi have been studied in an Indian pasture site. The average live weight of cocoon is 15.17 mg, length/diameter 0.94, and moisture content 72.5 g%. Just-hatched juveniles weigh on an average 10.4 mg, and are 10.5–17.5 mm in length. The incubation period lasts 56.33 ± 2 days during November—December in laboratory cultures (25°C, and moisture 15 g%). A maximum of 64.0 live cocoons/m² with a live weight of 0.971 g/m² were deposited during October—November 1977, but very few during summer months. A surface-living species, D. calebi shows single peak emergence pattern in December—January. It is estimated that 78 cocoons (live and dead)/m² (780,000/hectare), were produced in the field by D. calebi population in a year (1976–77). Thus the total cocoon production is 11.83 kg/hectare/year. It is estimated that 3 cocoons/individual/year are produced by D. calebi in field conditions.     

Specific criteria in Grania (Oligochaeta,Enchytraeidae)

The structure of the penial bulb and male efferent duct system of Grania species may be used in addition to setal pattern and spermathecal shape to distinguish species. Six penial bulb types are distinguished: (1) a simple, small, glandular bulb surrounding the male pore; (2) a small, glandular bulb, with a large, associated, dorso-medial gland mass; (3) a small glandular bulb, medial to the male pore, with an elongate male bursa (the aglandular sac), the vas deferens exitting directly into the invaginated male pore; (4) a glandular bulb with an aglandular sac and a small, cuticular stylet embedded in the bulb, extending from the ectal end of the vas deferens; (5) a glandular bulb and an aglandular sac with a long stylet extending from the vas deferens, through the bulb into the sac; and (6) glandular bulb reduced or absent, with or without an aglandular sac; with a long stylet and other prominent modifications, usually muscular, of the vas deferens. The details of the male duct structure were consistent within specimens grouped on the basis of setal distribution and shape and detailed spermathecal structure. Diverse male duct patterns are found within the polytypic species G. macrochaeta and G. postclitellochaeta. The positions of the spermathecal and male pores in their respective segments are distinctive for some species.     

Seasonal dynamics of aufwuchs Naididae (Oligochaeta) on Phragmites australis in a eutrophic lake

The Oligochaeta and Aphanoneura in the aufwuchs on Phragmites australis in a eutrophic hardwater lake were studied at two sites over a period of one year, in order to elucidate the structure and dynamics of this assemblage. The naidids Chaetogaster diastrophus, Nais spp., and Stylaria lacustris dominated the assemblage at any season. At both sites these taxa showed the same distinct pattern of successive population maxima in spring and summer: Chaetogaster diastrophus reached its peak density first, followed by Nais species, and eventually by Stylaria lacustris. Differences in temporal dynamics between sites were small apart from a second Stylaria maximum which was only observed at one site. Total naidid densities reached peak values of 3.8 individuals per cm² reed stem surface area. With mean individual biomass of 2.2 µg dry mass for Chaetogaster diastrophus, 13.3 µg for Nais, and 86 µg for Stylaria lacustris, respectively, maximum total naidid biomass on reed stems was 44 µg dry mass per cm². The biomass peak occurred later than that of total naidid density because in summer larger naidids dominated the assemblage. The observed succession appears to be consistent with seasonal changes in periphytic algal communities on the reed stems.     

Immunocytochemical demonstration of vertebrate neuropeptides in the earthworm Lumbricus terrestris (Annelida,Oligochaeta)

Summary The localisation and distribution of 10 vertebrate-derived neuropeptides in the earthworm, Lumbricus terrestris, have been determined by an indirect immunofluorescence technique. The peptides are pancreatic polypeptide (PP), peptide tyrosine tyrosine (PYY), neuropeptide Y (NPY), glucagon (C-terminal), vasoactive intestinal polypeptide (VIP), peptide histidine isoleucine (PHI), gastrinreleasing peptide (GRP), calcitonin gene-related peptide (CGRP), neurotensin (NT), and met-enkephalin. For 6 of the peptides — PYY, NPY, PHI, glucagon, GRP and CGRP — this is the first demonstration of their presence in any annelid, and NT has not previously been described in an oligochaete. Cell bodies and nerve fibres immunoreactive to the 10 peptides occur throughout the CNS. In the PNS, epidermal sensory cells displayed immunoreactivities to PP and PYY, and PP-, PYY-, NPY-, PHI- and GRP-like immunoreactivities occurred in nerve fibres supplying the main body muscles. Nerve fibres immunoreactive to PP and PYY are also associated with the innervation of the gut (pharynx, oesophageal glands, and mid and posterior regions of the intestine). No endocrine cells immunoreactive for any of the antisera tested could be identified in the gut epithelium, suggesting that dual location of peptides in the brain and gut epithelium is a phenomenon that occurred at a later stage in evolution. No immunoreactive elements were detected in any of the organs and ducts of the reproductive and excretory systems.     

Method for distinguishingLimnodrilus hoffmeisteri andLimnodrilus claparedeianus (Oligochaeta,Tubificidae) and its applicability in Lake Suwa

It is still next to impossible to distinguish species using immature worms of theLimnodrilus genus. A method was developed to separate mixed immature populations ofLimnodrilus hoffmeisteri andLimnodrilus claparedeianus into each species. The ratio of setal upper tooth length to lower tooth length, inL. hoffmeisteri andL. claparedeianus, ranged from 0.9 to 1.6 and from 1.3 to 2.1, respectively. Even if the median value of the frequency overlapped, this indistinguishable portion did not exceed 6% of each population.     

Intrinsic and extrinsic variables controlling the productivity of asexual populations of Nais spp, (Naididae,Oligochaeta)

The productivity of Nais spp. from periphyton of fishponds of the Dombes area (Ain) was studied in semi-natural conditions by cultivation of zooids in experimental glass enclosures immersed in situ and filled with pond water receiving injections of fertilizers (P₂O₅) and natural filtered periphyton extracts (particles < 70 µm). The growth rate of the experimental populations was not significantly affected by the concentration of fertilizers added to culture media. On the contrary, the water management of the culture media (as renewal or non-renewal of the water in experimental enclosures), the closing procedure of the enclosures and the load and composition of the nutritive substrate controlled the produced biomass. Temperature and food supply were the principal extrinsic variables controlling the asexual growth rate of the Nais species. The stolonization rate was analyzed as a biological parameter implicated in the instantaneous birth rate of zooids and the growth of naidid populations.     

The species-area relationship of European Lumbricidae (Annelida,Oligochaeta)

Summary Studies throughout Europe reporting species lists of lumbricid earthworms and ranging from 100 m² to >500000 km² are analysed for the regression of species number S on size of area A [km²]. This species-area relation is described by: S=7.9*A^0.09 (r=0.76).Revised version of a poster presented at the Wilhelm Michaelsen Memorial Symposium (International Symposium on Terrestrial Oligochaeta), Hamburg Sept. 14–18th 1987     

Iodination activity in Eisenia foetida (Annelida,Oligochaeta)

Summary In the nervous system of the earthworm Eisenia foetida, we have previously found thyroglobulin-like immunoreactive neurons. In the present study iodination activity was investigated by injecting worms or incubating them in vivo with radioiodine; animals treated with methimazole (MMI), an inhibitor of peroxidase-catalyzed iodination, served as controls. Radioiodinated proteins were identified in soluble extracts from ¹²⁵I-incubated animals; the sedimentation pattern of soluble proteins from cephalic segments showed a peak of radioactivity in the 3–4 S region. In animals pretreated with 10^-3 M MMI for 48 h, ¹²⁵I-incorporation into soluble proteins from cephalic segments was drastically reduced. Light-microscopic autoradiographic studies showed silver-grains selectively concentrated in the brain and ventral nerve cord of ¹²⁵I-injected animals. The highest grain-density occurred in the cerebral ganglion beginning 5 min after tracer injection; the reaction product was mainly distributed between the neurosecretory cells and neuropile fibres in the zone of the presumptive plexiform neurohaemal complex. In MMI-pretreated animals the reaction product was not visible in either cerebral ganglion or ventral nerve cord. Of interest, the setae appeared consistently positive with or without MMI treatment. These observations indicate that protein iodination involving peroxidase occurs in the nervous system of earthworms and suggest that iodination mechanisms, other than peroxidase-catalyzed, may be operating in some scleroprotein structures such as setae.     

Life-cycle and karyology of Branchiura sowerbyi Beddard (Oligochaeta,Tubificidae)

Data on the life-cycle of a population of Branchiura sowerbyi Beddard in a water-lily tank at the Botanical Garden in Padua are reported. The breeding period is from April to July, after which the reproductive system is partially resorbed (August–September) and reformed later in the autumn. The karyology of the species was also studied, and revealed 38 mitotic chromosomes in the gonia, and 19 bivalents in the primary spermatocytes and in the primary oocytes.     

Growth of Stylaria lacustris (L.) (Oligochaeta,Naididae)

The peculiarities of the growth in weight and length of Stylaria lacustris (L.) on the basis of observations in experimental vessels are considered. The growth of this species fits a parabolic curve. The equations relating weight to absolute growth rate as well as weight to duration of life are given.     

New species of Phallodrilus (Oligochaeta,Tubificidae) from caves of northern Spain and southwestern France

A taxonomic account of a collection of Phallodrilus species inhabiting caves is given. The following three new species are described: P. subterraneus, P. crypticus and P. labouichensis. New material of P. aquaedulcis Hrabe, 1960 from northern Spain and southwestern France is reported. This species was previously known from West Germany. The relationship between Phallodrilus cave species and littoral and deep-sea species is discussed.     

Population dynamics of Stylaria lacustris (Linnaeus, 1767) (Oligochaeta,Naididae) in Los Talas,Argentina

Population dynamics of Stylaria lacustris were analyzed over 2 years in a pond located at Los Talas, Argentina. In both years, the peak in abundance, due to intense asexual reproduction, fell at the end of winter, and was followed by mature individuals. The species was not collected during summer. Temperature was the main factor regulating the population through both sexual and asexual reproduction. In the second year, density was lower and correlated with a lower water level and a reduced vegetation development. Individuals were randomly distributed in periods of high abundance and contagious when density was low. Generation time was 15 days.     

Oligochaeta of the middle Po River (Italy): principal component analysis of the benthic data

The application of principal component analysis to two types of habitat (the benthos of macrophytes and of central river bed) enabled us to single out some of the factors that affect the dynamics and the structure of the oligochaete population and its various reactions to environmental conditions. As regards macrophytes, the distribution of the variables on the basis of the first component is correlated, to a certain extent, with a seasonal factor without any significant differences among sites. The largest population is most closely correlated with the summer months. In fact, we found that the Naididae and Tubificidae species generally develop in larger numbers at higher temperatures. For the Tubificidae, we could detect a precise seasonal cycle. In the central river bed habitat, the first component was correlated with the river discharge, which determines the granulometric characteristics of the sediment; we noticed a correlation among the sites that have the same characteristics, regardless of sampling site or date. The species which correlate most closely among themselves are the Tubificidae Limnodrilus hoffmeisteri, Tubifex tubifex, L. udekemianus and L. profundicola, which are very characteristic of environments that contain abundant organic matter. The second component is correlated with temperature, and hence with the availability of oxygen, which determines the presence and the abundance of more sensitive species.     

The occurrence of species of semi-aquatic Enchytraeidae (Oligochaeta) in Ireland

The Enchytraeidae are essentially terrestrial oligochaetes but many species have marked aquatic tendencies. Over two thirds of recorded Irish species were found in soils which were submerged or frequently flooded and 35% showed a distinct preference for these conditions. Relatively few species were living in soils subject to drought. Red blood was present in 28 species, all but one from soils with more than 55% water. Cognettia sphagnetorum and C. glandulosa developed red blood in very wet conditions. In a survey of Irish wetlands, samples were taken from bog, heath, marsh, fen, margins of lakes and rivers, and saltmarsh. The influence of various environmental parameters was determined using ordination techniques. Magnesium and pH were found to be the most important factors. A high level of magnesium distinguished coastal sites and pH 5.2 separated two clusters representing acid peat and marsh-fen-aquatic sites. Groups of indicator species characterized each of the three clusters. The ecological distribution of the indicator species is described, and their usefulness in classifying enchytraeid communities is discussed.     

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.     

Seasonal dynamics and the structure of colonies of Proterospongia skujae Bourrelly

The seasonal dynamics, the structure of colonies and the abundance of cells in the colonies of the monad Proterospongia skujae have been simultaneously studied in the river Danube and in a backwater of the Danube. Significant differences have been found in both the dynamics and the abundance of P. skujae in selected localities. Colonies are formed of 2 up to 16 cells, most frequency of 4 and 8 cells.     

Phylogenetics of some Enchytraeidae (Annelida: Oligochaeta): A preliminary investigation of relationships to the Haplotaxidae

In phylogenetic considerations of the Oligochaeta and the family Enchytraeidae, Propappus has been considered the ancestral or even the archetypal enchytraeid genus. Three nominal species are presently included in Propappus Michaelsen, 1905, P. volki Michaelsen, 1916, P. arhyncotus Sokolskaja, 1972, and the type species of the genus, P. glandulosus Michaelsen, 1905. Examination of type material of P. glandulosus and P. volki, as well as other material of the latter, showed that these species have a single pair of testes in segment XI and a single pair of ovaries in segment XIII. According to current classifications of the Oligochaeta, these species of Propappus cannot be retained in the Enchytraeidae. A phylogenetic analysis including two species of Propappus and 29 species of Haplotaxidae suggest a sister group relationship between Propappus and some Enchytraeidae and Haplotaxidae.     

Species separation and identification in the Enchytraeidae (Oligochaeta, Annelida): combining morphology with general protein data

In the Enchytraeidae, species separation and identification is often problematic due to high morphological similarity of closely related species and considerable intraspecific variability of crucial characteristics. Immature specimens are almost undeterminable. To meet these difficulties, this paper recommends the consulting of general protein patterns as exhibited by non-specific silver-staining after isoelectric focusing. A method is presented which allows the successful inclusion of protein data in taxonomic studies and field surveys. Two examples from an investigation on Fridericia field populations show that, with the help of general protein pattern analysis, a clear taxonomic decision on the identity of morphologically aberrant forms and of juvenile specimens as well can be achieved with comparatively little expenditure of time. A combined use of morphological and protein data for taxonomic purposes is suggested.