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.     

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.     

A taxonomic and faunistic survey of the marine Tubificidae and Enchytraeidae (Oligochaeta) of Italy. Introduction and preliminary results

On the basis of material collected along the Ligurian, Tyrrhenian, Ionic and Adriatic coasts of Italy, a tentative list of 26 species (18 tubificids, 8 enchytraeids) of marine Oligochaeta is presented. Most of the species are new to science, and it can therefore be concluded that there is a very high diversity of oligochaetes in the Mediterranean Sea.     

Histophysiology of digestion and observations on the structure of the alimentary canal in the ectosymbiont Chaetogaster limnaei limnaei Baer, 1827 (Annelida: Oligochaeta)

The naidid oligochaete Chaetogaster limnaei limnaei has an alimentary canal consisting of a mouth, pharynx with a dorsal pharyngeal pad, esophagus, stomach, anterior and posterior intestine, and anus. The diet is omnivorous but limited by particle size. Unattached food organisms are sucked into the pharynx while sessile organisms are plucked from the substratum. Granules of acid mucosubstances that stain purple with neutral red are secreted into the stomach lumen after food enters, rapidly increasing the acidity from pH 3 to 1.5. Acid induced lysis of the organisms initiates autolysis before the food is passed into the alkaline, pH 7 to 8, anterior intestine. Ciliated intestinal cells showed arylamidase, acid phosphatase and C-esterase active granules indicating primary lysosomes with secondary lysosomes being recognized in electron micrographs suggesting intracellular digestion. Arylamidase and alkaline phosphatase activity appears in the intestinal margins during the alkaline phase of digestion. Scattered, pyramidal cells found only in the anterior intestine contain yellow refractile spheres. The spheres stain alcian blue pH 2.5 and bromophenol blue positive and exhibit a strong acid phosphatase activity all the time with A-esterase active granules surrounding them. Glycogen and lipids are stored mainly in the chlorogague cells. Many of the yellow refractile granules in the stomach and intestinal cells are bacteria.     

Preliminary report on the Enchytraeidae (Oligochaeta) of West Florida

A wide range of habitats, both terrestrial and marine littoral, were sampled in an area between Pensacola and Tallahassee. Enchytraeids were present in all moist or wet soils and in marine habitats enriched by organic matter but were absent or rare in very dry soils, saturated or flooded substrates and clean intertidal sand. Between 70 and 75 species were distinguished of which 50–55 are new to North America and about 25 are probably new to science. The genera represented are the same as those occurring in Western Europe but their relative importance differs and there are some tropical elements. The number of species occurring in terrestrial sites was greater than recorded in similar surveys in southwestern France and Ireland (approximately the same-sized area, number of sites and size of sample), but the number of species per sample was lower than in either of the other countries. It is concluded that species distribution in western Florida is very patchy. Possible reasons for this patchiness are discussed.     

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.     

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.     

Ecology and importance of Oligochaeta in the biocenosis of zoobenthos in lakes of the National Park of the Lithuanian SSR

This paper aims to describe the ecology and determine the distribution of Oligochaeta according to sediment type, depth and other environmental factors in 21 lakes of the National Park of the Lithuanian SSR. Diagrams are included showing the distribution of the lakes according to the index of density and biomass of Oligochaeta in the trophic structure of biocenosis of the zoobenthos. Light is thrown on the problem of sediment composition as a significant factor in the distribution and development of aquatic Oligochaeta.     

Diversity and zoogeography of marine Tubificidae (Annelida,Oligochaeta) with notes on variation in widespread species

The specific and generic diversities of the marine Tubificidae (Annelida, Oligochaeta) of the NE Pacific are compared to those of the NE and NW Atlantic as well as to those of Heron Island, Australia. Diversity in the NE Pacific is relatively high when compared to that of the NE Atlantic. The Tubificidae of the NW Atlantic (limited to the eastern coast of the USA) show two distinct zoogeographic regions: Florida to Cape Hatteras; Cape Hatteras to Massachusetts. Diversity, both in terms of the number of species and number of genera, is approximately the same in these two regions, and is similar to that of both the NE Pacific and Heron Island. Evidence suggests that the widespread marine species, in particular Tubificoides pseudogaster, have a range of morphotypes across their distributions. The apparent wide distributions of these species may be due to a taxonomy unable to resolve the differences between the morphotypes. The tubificid oligochaete fauna of the NE Atlantic appears impoverished compared to the other regions examined. The NE Pacific, NW Atlantic, and Heron Island regions are not dominated by one group of species while the NE Atlantic fauna is dominated by Tubificoides benedeni and Clitellio arenarius.     

Molecular phylogeny of Enchytraeidae (Oligochaeta) indicates separate invasions of the terrestrial environment

Enchytraeidae is a family of soil inhabiting small‐ to medium‐sized oligochaete worms using degradable plant material as a food source and primarily adapted to terrestrial or semi‐terrestrial environments. The molecular phylogeny based upon both mitochondrial and nuclear genes indicates early segregations of the two genera Enchytraeus and Lumbricillus leaving the remaining genera included in this study as a later segregated major monophyletic branch. Extant members of the two former genera dominate in decaying seaweed in the littoral zone along the sea although members of in particular the genus Enchytraeus have also invaded other habitats. Historically the littoral zone of the sea is undoubtedly the first terrestrial or semi‐terrestrial habitat where dead plant material accumulates to any greater extent and Enchytraeus and Lumbricillus may represent early successful attempts to exploit this resource. Inland soils probably had to await the emergence of land plants in order to provide a similar food resource and here the major branch of enchytraeid genera diversified into a high number of species in the numerous decomposer networks of this varied environment. A subdivision into the genera Enchytraeus and Lumbricillus on the one hand and a branch of mainly inland genera on the other is supported by differences in two somewhat neglected morphological features. Firstly, in Enchytraeus and Lumbricillus the testes are enclosed in a testis sac within which the male cells mature, by one possible exception a unique feature among Oligochaeta, The other enchytraeid genera studied and Oligochaeta in general lack this sac and the male cells mature directly in the cavity of the testicular segment. Secondly, species of Enchytraeus and Lumbricillus generally have a higher reproductive output than species of the inland terrestrial branch and this may represent an adaptation to the unpredictable littoral zone compared to the more stable nature of inland habitats. In the older literature the genus Mesenchytraeus is considered to have a basic position within the entire family but our molecular data do not support this expectation. In Enchytraeidae the nephridia are elaborate organs of a characteristic and constant shape covering species from different genera in a pattern following the molecular phylogeny. Other much used morphological features such as shape of setae, anteclitellar origin of the dorsal vessel and various modifications of the intestine have arisen more than once.     

Population dynamics of Tubificoides amplivasatus (Oligochaeta,Tubificidae) in the Oresund,Denmark

A two year fortnightly collection of data from the 27 m station in the Oresund on the tubificid species Tubificoides amplivasatus was analyzed for population trends and life stages. Recruitment patterns from year to year were consistent in timing but very irratic in magnitude, being similar to patterns seen in other estuarine and marine annelids. Sexually mature T. amplivasatus were found throughout the study period, but mature mated specimens were more abundant during late summer and autumn. It is estimated that from 120 to 150 days is spent, by this species, in the cocoon and individuals may reach maturity in less than one year. There was some indication of individuals with regressed genital organs indicating that they may live longer than one year. At the 27 m station T. amplivasatus is part of a complex and well developed benthic community. Temporal variation in this species was highly correlated to that of other major taxa (bivalves, ophiuroids).     

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.     

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.     

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.     

A comparative cytological investigation of the reproductive cycle of an amphimictic diploid and a parthenogenetic triploid form of Lumbricillus lineatus (O.F.M.) (Oligochaeta,Enchytraeidae)

Summary The cytology of a diploid amphimictic (2x=26) and a triploid parthenogenetic (3x=39) cytotype of Lumbricillus lineatus has been described. The triploid type never produces mature sperm, but it must be fertilized by sperm from the diploid cytotype in order to produce viable eggs. However, the sperm does not enter the egg, but eggs which have not been activated by sperm die at an early stage. Oogenesis in the triploid cytotype, which results ultimately in the restoration of the somatic chromosome number, follows a pattern which has not been described in other parthenogenetic organisms. The first meiotic division is asynaptic and no typical metaphase plate is formed; the univalents are distributed to opposite poles without undergoing an equational division. The first division is therefore numerically approximately reductional. The nuclei are at anaphase when the eggs leave the animal, whereas the nuclei of the diploid amphimictic cytotype are in MI when the eggs are laid. Following activation by spermatozoa, the first anaphase spindle is much elongated, and becomes V-shaped. The first anaphase spindle persists and no second anaphase spindle is formed. At the second division the chromosomes divide mitotically. The resulting four chromosome groups fuse two by two in such a way that two nuclei are formed each with the full somatic (triploid) chromosome set (cf. Figs. 14–16).The different steps involved in the evolution of this mechanism are discussed in relation to the evolution of the triploid cytotype.     

Oligochaeta from profundal zones of Spanish reservoirs

The profundal benthic fauna of 63 Spanish reservoirs was studied during 1972–75. Samples were taken in the deepest part of each reservoir with a modified Van Veen grab. The fauna was dominated by tubificid worms and chironomids. Sixteen species of Oligochaeta were found, two of them new for the Spanish fauna (Potamothrix heuscheri and Haber pyrenaicus). This was the first time H. pyrenaicus was found since the species was described in 1974 from the Pyrenees. Four species were frequent in the reservoirs (Limnodrilus hoffmeisteri, Tubifex tubifex, L. claparedeianus and Dero digitata) but only the first two were abundant as mature individuals. Another interesting fact is the high abundance of immature animals in all samples, accounting for more than 80% of the total individuals examined. The geographical distribution of all species in Spain and their relationship with the typology of Spanish reservoirs are discussed. There seems to be a correlation between the throphic status of the reservoirs and the relative proportion of L. hoffmeisteri as compared to T. tubifex, the number of individuals of the first species increasing with water eutrophication.     

A phenetic and cladistic study of spermatozoal ultrastructure in the Oligochaeta (Annelids)

Spermatozoal ultrastructure of nine oligochaete families has been examined for congruence with phylogenetic and taxonomic systems for the Oligochaeta based on general morphology, particularly the holomorphological hennigian analysis of Jamieson (1978a, 1980, 1983). Estimation of congruence has been made following phenetic and cladistic (phylogenetic) analysis. Correspondence, in phenograms and phylograms, of sperm types with taxonomic and phylogenetic groupings previously recognized is generally good. Departure from this rule in the similarity of the phreodrilid sperm to that of the Lumbricina suggests a corresponding alteration of fertilization biology in the phreodrilids. The results indicate that the Haplotaxidae lie at the base of the opisthopores though they do not unequivocally contraindicate acceptance of a Haplotaxis like form as a stem form of the Haplotaxida (opisthopores and Haplotaxidae) and Tubificida. An even more basal position for prosopores, now represented by the Lumbriculida, cannot yet be dismissed.     

Two atavistic characters of some Lumbriculidae and their importance for the classification of Oligochaeta

The rudimentary atria, and the posterior sperm funnels and sperm ducts, which occur in some species of the Lumbriculidae, are discussed. It is shown that the posterior location of funnels and sperm ducts is the result of a forward shift of the atria, which refutes Stephenson's supposition that the Lumbriculidae is the most archaic family of the present-day Oligochaeta.     

Phylogeny and species delimitation of North European Lumbricillus (Clitellata,Enchytraeidae)

The enchytraeid genus Lumbricillus comprises about 80 described species of clitellate worms, which are up to a few centimetres long, and they mostly inhabit the littoral zone of non‐tropical marine and brackish waters world‐wide. The phylogeny of this genus is poorly studied, but previous work has suggested that Lumbricillus is a non‐monophyletic group. In this study, species boundaries and the phylogeny of this genus is re‐assessed using more than 300 DNA‐barcoded specimens (using COI mtDNA), part of which was also sequenced for two additional mitochondrial and four nuclear molecular markers. Statistical and coalescent based applications were used for the delimitation of a total of 24 species, of which 20 were identified as belonging to 17 described morphospecies; one morphospecies was found to be a complex of four delimited species, and another four delimited species could not be matched with any described species. Furthermore, gene trees, concatenation and multispecies coalescent based species trees were estimated using Bayesian inference. The estimated phylogenies confirm a non‐monophyletic Lumbricillus as L. semifuscus is clearly excluded from the genus. Furthermore, the placement of a monophyletic clade consisting of L. arenarius, L. dubius, and an unidentified species varies between analyses; they are either found as the sister‐group to the genus Grania or as sister‐group to the remaining Lumbricillus, where the latter relationship is supported by the multispecies coalescent, which we consider as the most reliable method.