Isoenzyme differences between three closely related species of Lineus (Heteronemertea)

Starch-gel electrophoresis was employed to compare six enzymes in three closely related species of nemertean worms, Lineus ruber (Müller, 1774), Lineus sanguineus (Rathke, 1799), and Linens viridis (Müller, 1774). Differences in mobility recorded for most of the enzyme loci examined support the hypothesis that these nemerteans are distinct taxa.     

Impact of the nemertean Lineus viridis on its polychaete prey on an intertidal sandflat

Interactions of the predator Lineus viridis (O. F. Muller 1774) with infaunal polychaetes were investigated in the German Wadden Sea. L. viridis is the dominant heteronemertean in this intertidal area with a mean abundance of 14 individuals per square meter. Predator enclosure experiments in the field revealed that L. viridis preyed on predatory polychaetes. The preferred prey species was Nereis diversicolor; others were Anaitides maculata. Nephtys hombergii and Harmothoe sarsi. These results were supported by a series of feeding experiments in the laboratory. The rate of consumption was estimated by laboratory and field experiments. The feeding rate of Lineus viridis was estimated by laboratory enclosure experiments to 5.6 g ashfree dry weight (AFDW) per square meter annually.     

The fate of the larval epidermis in the Desor-larva of <Emphasis Type="Italic">Lineus viridis</Emphasis> (Pilidiophora,Nemertea) displays a historically constrained functional shift from planktotrophy to lecithotrophy

Pilidiophora constitutes a clade of nemerteans characterized by a peculiar larval type, the pilidium. A characteristic of this larva is the transitory epidermis in which the juvenile develops from imaginal discs. The primary function of this larval envelope is assumed to be feeding and dispersal. When juvenile development is complete, the larval epidermis is ruptured and swallowed by the juvenile. According to recent cladistic and molecular analyses of the Nemertea, the intracapsular Desor-larva of the sibling species Lineus viridis and L. ruber is thought to have evolved from a pelagic pilidium. The general course of development has been demonstrated to be similar to that of the pilidium, in which the juvenile forms from imaginal discs under the larval epidermis. The two Lineus species, however, differ in their mode of larval feeding: L. ruber being ootrophic and L. viridis being lecithotrophic. In order to elucidate the transition from the planktotrophic pilidum to lecithotrophic development, I studied the early cleavage and metamorphosis from intracapsular Desor-larva to juvenile stages in L. viridis from the island of Sylt, using light microscopical, electron microscopical, and fluorescent staining methods. Due to the specific cleavage pattern with equally sized 1st quartet animal blastomeres and vegetal blastomeres in L. viridis, the larval epidermis later contains a considerable amount of the yolk reserve. During metamorphosis, the larval epidermis is ingested by the juvenile thus displaying behavior similar to that of the pilidium larva. In contrast to the pilidium, the function of the larval epidermis of the Desor-larva has shifted from feeding and dispersal to direct food supply. Thus, the development of L. viridis is a perfect example for strong historical constraints that prevent ancestral larval structures from being lost.     

Molecular Phylogeny of Some European Heteronemertean (Nemertea) Species and the Monophyletic Status ofRiseriellus, Lineus,andMicrura

The 16S rRNA mitochondrial gene was used to reconstruct the relationships among 10 heteronemertean species (subclass Heteronemertea, phylum Nemertea);Lineus ruberandL. viridisare represented by more than one specimen to assess intraspecific variation in these enigmatic species, and the analysis includes in total 14 terminal taxa incorporating one palaeonemertean species (Tubulanus annulatus) for outgroup rooting. The aligned sequences were subjected to maximum parsimony, maximum-likelihood, and neighbor-joining analyses to estimate the phylogenetic relationship of the species. The results were concordant from all analyses and indicate that neitherLineusnorMicruraare monophyletic taxa, and that there is no support from a phylogenetic point of view to establish the monotypic genusRiseriellus.     

Fortgesetzte Studien Über die Regeneration der Nemertinen

Ohne Zusammenfassung Wir bezeichnen diesen Abschnitt unsrer fortgesetzten Studien über die Regeneration der Nemertinen betreffendLineus lacteus Rathke mit II, da der erste Abschnitt sich aufLineus ruber Müll, bezog (vgl. das Literaturverzeichnis Nr. 8–10, 13 u. 14).     

Enzyme electrophoresis,genetic identity and description of a new genus and species of heteronemertean (Nemertea,Anopla) from northwestern Spain and North Wales

The anoplan order Heteronemertea, particularly the genera Cerebratulus, Lineus and Micrura, contains a very large number of nominate species, many of which are inadequately described. As a consequence, systematic difficulties are encountered with the identification of many taxa in this group, especially those originally established primarily on the basis of their external features. The present paper concerns heteronemerteans collected from two locations, the Foz Estuary (north-western Spain) and Llandudno (North Wales). The Spanish collection included specimens identified as Lineus longissimus (Gunnerus), whilst samples from Llandudno contained large numbers of Lineus viridis (Müller); samples of a third similar but apparently undescribed species were found at both locations. Starch gel electrophoresis showed that samples of the apparent third species were genetically almost identical from each of the two locations, but were clearly different from the two described Lineus species. Histological studies of the unknown specimens revealed anatomical characters, including the unique feature of a proboscis epithelium ciliated throughout its length, which exclude it from any known heteronemertean taxon; it is accordingly placed in a new genus and species, for which the name Riseriellus occultus is proposed.     

Monoamine- and peptide-containing elements in the body wall and nervous trunks of nemerteans

Localization and morphological peculiarities of cateholamine-containing (CA-C), serotonin-(5HTIR), neurotensin-(NT-IR) and FMRFamide-immunoreactive (FMRFa-Ir) cells and processes in the body wall and lateral nervous trunks of nemerteans were investigated. The following nemertean species from three orders occurring in the White Sea were studied by fluorescent histochemistry and by immunohistochemistry: Lineus viridis, Lineus ruber and Cerebratulus sp. (Heteronemertini), Cephalothrix linearis (Palaeonemertini), and Amphiporus lactifloreus (Hoplonemertini). The similarity and characteristics of morphological types of monoamine-and peptide-containing elements, their distribution and possible participation in regulation of some functions in different nemertean species and in other invertebrates are discussed. We suggest that 5HT-IR and CA-C bi-and multipolar intra-and subepidermal open-type cells prevailing in the body wall perform a receptive function, while the FMRFa-Ir and NT-IR neurons of the nerve trunks and body wall are responsible for efferent innervation of musculature and connections within the nervous system.     

A reassessment of the systematics and a proposal for the phylogeny of some cosmopolitan Lineus species (Nemertea)

Morphological, behavioural and biochemical characters of 9 cosmopolitan, benthic species of the nemertean genus Lineus are used both to clarify their taxonomy and to discuss their phylogenetic relationships. Data analysis shows that all fissiparous species of Lineus collected in the world seas up to date can be placed in a single species, L. sanguineus, with three recognisable subspecies, L. s. sanguineus, L. s. nigricans and L. s. pseudolacteus. These taxa were originally described as separate species mainly according to their geographic origin under the names L. sanguineus (shores of European seas), L. socialis (Atlantic North American shores), L. vegetus (Pacific North American shores), L. pseudolacteus (shores of the English Channel) and L. nigricans (Mediterranean coasts of Italy and France). Examination of the character value matrix suggests a phylogenetic tree developing from two ancestral branches. The first branch includes the two related species L. ruber and L. viridis, the second branch the three species L. longissimus, L. lacteus and L. sanguineus. This approach to the taxonomic and phylogenetic relationships of Lineus nemertean species is in agreement with the intra- and interspecies histocompatibility data in nemerteans: (i) grafts transplanted from donors to recipients of the same species succeed and grafts transplanted from donors to recipients of different species fail; (ii) grafts are rejected more rapidly when the donor and recipient species diverged earlier in evolution and, a contrario, grafts are accepted better when the donor and recipient species have been more recently isolated.     

Ultrastructure of sperm and male reproductive system in Lineus viridis (Heteronemertea, Nemertea)

Nemerteans possess serially arranged gonads that lie between the midgut pouches. In both sexes the gonads are lined with an epithelium. During maturity, they gain contact to the exterior by a ciliated duct, which is generally assumed to be a derivative of the gonad. Gonad lining and sperm ultrastructure are little known in heteronemerteans, a group of nemerteans belonging to the Anopla, one of the two large nemertean subgroups. Reproduction biology in heteronemertean Lineus viridis allows predicting a modified sperm type, so-called introsperm for this taxon. Nothing is known on the fate of the testes at the end of the reproductive period of this perennial species. In order to test the predictions and to broaden the data base, males of L. viridis were collected at different times of the year. Histological and ultrastructural data show that the gonad wall is lined with different aciliated endothelial cells and germ cells, while the gonoduct is formed by densely ciliated cells. The testes are completely filled with sperm cells during maturity; there is no hint at ongoing spermiogenesis at this time. The sperm consists of head, midpiece and tail. Externally, head and midpiece cannot be discriminated. The acrosome is cup-shaped and lies anterior to the nucleus which contains 6–8 lateral ridges. Three long mitochondria mark the midpiece. They line the posterior section of the nucleus and extend up to the level of the ciliary basal structures. The sperm morphology corroborates the predictions derived from the mode of reproduction. At the end of the reproductive period the male gonads change cellular composition, while the gonoduct degenerates. Provided that both sexes show the same growth rate, male offspring acquire sexual maturity earlier than female offspring, since L. viridis males are always smaller than the females. In contrast to the males, females keep their gonads and gonoducts during most time of the year. Since large males were never found within the studied population, these data indicate that L. viridis might be a consecutive hermaphrodite.     

PHYLOGENY AND SYSTEMATICS OF EUGLENA (EUGLENACEAE) SPECIES WITH AXIAL,STELLATE CHLOROPLASTS BASED ON MORPHOLOGICAL AND MOLECULAR DATA—NEW TAXA,EMENDED DIAGNOSES,AND EPITYPIFICATIONS1

Morphological and molecular studies, as well as original literature reexamination, necessitate establishment of five Euglena species with a single axial, stellate chloroplast [Euglena viridis (O. F. Müller) Ehrenberg 1830 , Euglena pseudoviridis  Chadefaud 1937 , Euglena stellata  Mainx 1926 , Euglena pseudostellata sp. nov., and Euglena cantabrica  Pringsheim 1956 ], three species with two chloroplasts (Euglena geniculata Dujardin ex Schmitz 1884 , Euglena chadefaudii  Bourrelly 1951 , and Euglena pseudochadefaudii sp. nov.), and one species with three chloroplasts (Euglena tristella  Chu 1946 ). The primary morphological features, allowing distinction of the considered species are the presence and the shape of mucocysts, as well as the number of chloroplasts. Spherical mucocysts occur in E. cantabrica and E. geniculata, while spindle‐shaped mucocysts are present in E. stellata, E. pseudostellata, E. chadefaudii, E. pseudochadefaudii, and E. tristella. No mucocysts are observed in E. viridis and E. pseudoviridis. Two new species (E. pseudochadefaudii sp. nov. and E. pseudostellata sp. nov.) differ from the respective species, E. chadefaudii and E. stellata, only at the molecular level. Molecular signatures and characteristic sequences are designated for nine distinguished species. Emended diagnoses for all and delimitation of epitypes for seven species (except E. viridis and E. tristella) are proposed.     

Life history of Lineus viridis (Müller, 1774) (Heteronemertea, Nemertea)

Lineus viridis is a common nemertean species of North-Atlantic intertidal sand flats. Its mating behaviour is peculiar insofar as this species is reported to be polyandric. However, detailed information on this topic is lacking. In order to get more data on the reproduction, oogenesis and life history of this species, a population in the Wadden Sea on the Isle of Sylt (North Sea) was studied between 2005 and 2011. We conducted regular surveys, during which we sampled, measured and recorded the sex status of 25–100 individuals at each sampling event; at least three individuals were fixed for histological studies at each sampling date. In addition, animals were kept in the laboratory for 3?years to complement field data on sexual identity. Lineus viridis was found to reproduce annually in several successive year; the females are significantly larger than the males. Oogenesis starts in spring, shortly after the preceding reproductive period, and continues until the end of December. Spermiogenesis starts in late autumn and also ends late in December. During mating, several males are generally found crawling on a single female, which forms a cocoon that encloses both the female and the associated males. Fertilization is internal. While females discharge all of their eggs during a single mating event and lose more than 40% of their wet weight, males only empty a few of their gonads, and are thus able to fertilize more than one female. Our study clearly shows that Lineus viridis is a perennial, iteroparous species with a pronounced sexual size dimorphism. During this long-term study, no evidence for sequential hermaphroditism has been found. The observed polyandric mating system in this species raises further questions regarding mate and sperm competition that deserve additional research.     

Zur Fortpflanzungsbiologie vonLineus viridis (Nemertini)

The fertilization of the heteronemerteanLineus viridis Müller takes place in a cocoon built up by the female surrounding itself and the male. After the male has left the cocoon, the female builds a mucous layer within the cocoon, in which the fertilized eggs enclosed in egg capsules are embedded. Observations on mating behavior and spawning are described and discussed.     

Ultrastructure des photorécepteurs de Lineus ruber (O. F. Müller) (Hétéronémertes Lineidae)

Résumé Lors de la régénération traumatique des yeux de Lineus ruber la différenciation biochimique des nouvelles cellules pigmentaires est séquentielle. On assiste à l'apparition successive des chaînes enzymatiques nécéssaires à la biosynthèse de porphyrine et de mélanine.Les études ultrastructurale et autoradiographique à haute résolution de ces phénomènes — montrent que la mélanisation s'opère au niveau d'organites cellulaires spécialisés (prémélanosomes, mélanosomes). La porphyrinogénèse se développe dans des vacuoles et organites d'origine golgienne qui participent également à la mélanogénèse.

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Ultrastructure of photoreceptors in Lineus ruber (O. F. Müller)

Summary In Lineus ruber the biochemical differentiation of the new pigmentary cells is sequential during the reparative regeneration of the eyes. The enzymatic pathways for porphyrin and melanin biosynthesis appear successively.Ultrastructural and high resolution radioautography studies, show that melanization occurs in specialized organelles called premelanosomes, melanosomes and melanin granules. Prophyrogenesis occurs in Golgi vesicles which are also involved in melanogenesis.

     

Survey tools and demographic parameters of Slovakian Agrilus associated with beech and poplar

Woodborers in the Agrilus genus (Coleoptera: Buprestidae) pose high invasiveness risk as indicated by the recent invasion and continental spread of emerald ash borer, and the associated threat to ash resources in North America. In that context, development of detection tools for potentially invasive Agrilus spp. is a research priority. Experiments carried out in 2013 in Slovakian beech and poplar forests evaluated the attraction of multiple Agrilus species to green and purple sticky prism traps baited with various lures [blank, cubeb oil, (Z)‐3‐hexenol]. The two most abundant species were Agrilus viridis L. in beech (Fagus spp.) forest (146 adults, >95% of which were females) and Agrilus convexicollis Redtenbacher in poplar (Populus spp.) forest (158 adults, two‐thirds of them males). The two species exhibited opposite responses to color: purple traps attracted 2–3× more adult A. viridis than green traps, whereas most (>95%) specimens of A. convexicollis were captured on green traps. Volatile baits did not influence captures of adults for either species. The introduction and establishment of A. viridis in North America is of particular concern owing to its feeding niche (primary pest that can attack healthy trees), large body size, and high level of polyphagy (>10 genera of host trees). Additional experiments conducted in beech forests in 2014 found purple prism traps more attractive to female A. viridis than green prism traps, especially those baited with cubeb oil. No analysis was conducted for males because of their low abundance. Female A. viridis flew earlier in 2013 than in 2014, but neither their body size nor fecundity varied between years. In both years, large females had more eggs in their abdomen than small females, and the number of eggs steadily declined over time, which suggests that female A. viridis are reproductively mature at emergence.     

The coralline genera Sporolithon and Heydrichia (Sporolithales,Rhodophyta) clarified by sequencing type material of their generitypes and other species

Interspecific systematics in the red algal order Sporolithales remains problematic. To re‐evaluate its species, DNA analyses were performed on historical type material and recently collected specimens assigned to the two genera Sporolithon and Heydrichia. Partial rbcL sequences from the lectotype specimens of Sporolithon ptychoides (the generitype species) and Sporolithon molle, both from El Tor, Egypt, are exact matches to field‐collected topotype specimens. Sporolithon crassum and Sporolithon erythraeum also have the same type locality; material of the former appears to no longer exist, and we were unable to PCR amplify DNA from the latter. A new species, Sporolithon eltorensis, is described from the same type locality. We have not found any morpho‐anatomical characters that distinguish these three species. No sequenced specimens reported as S. ptychoides from other parts of the world represent this species, and likely reports of S. ptychoides and S. molle based on morpho‐anatomy are incorrect. A partial rbcL sequence from the holotype of Sporolithon dimotum indicates it is not a synonym of S. ptychoides, and data from the holotype of S. episporum confirm its specific recognition. DNA sequences from topotype material of Heydrichia woelkerlingii, the generitype species, and isotype material of Heydrichia cerasina confirm that these are distinct species; the taxon reported to be H. woelkerlingii from New Zealand is likely an undescribed species. Type specimens of all other Sporolithon and Heydrichia species need to be sequenced to confirm that they are distinct species; morpho‐anatomical studies have proved inadequate for this task.     

Phylogeography of the Lacerta viridis complex: mitochondrial and nuclear markers provide taxonomic insights

Based on broad, nearly rangewide sampling, we reanalysed the phylogeography of the Lacerta viridis complex using the mitochondrial cytochrome b gene and the intron 7 of the nuclear β‐fibrinogen gene. Using the mitochondrial marker, we identified in phylogenetic analyses 10 terminal clades clustering in four deeply divergent main lineages whose relationships are weakly resolved. These lineages correspond to Lacerta bilineata, L. viridis, the previously identified Adriatic or West Balkan lineage and a newly discovered fourth lineage from the Anatolian Black Sea coast and the south‐eastern Balkan Peninsula. Except for the latter lineage, there is considerable phylogeographic structuring in each lineage, with higher diversity in the south of the distribution ranges. This pattern indicates the existence of two distinct microrefugia in the Italian Peninsula and Sicily and of up to seven microrefugia in the Balkan Peninsula, but of only one refugium along the Black Sea coast of Anatolia. We identified secondary contact zones of the main lineages and of terminal clades within these lineages. However, most of the formerly described putative contact zone of L. bilineata and L. viridis turned out to be a contact zone between the Adriatic lineage and L. viridis, but L. bilineata seems to be involved only marginally. Our nuclear marker could not unambiguously resolve whether there is gene flow in contact zones. Thus, further research is necessary to decide whether the four main lineages are conspecific or whether they represent distinct biological species. We restrict the name L. v. meridionalis to the newly identified genetic lineage from Turkey and south‐eastern Europe, synonymize some previously recognized taxa and suggest a tentative nomenclature for the L. viridis complex.     

A taxonomic catalogue of Japanese nemerteans (phylum Nemertea)

A literature-based taxonomic catalogue of the nemertean species (Phylum Nemertea) reported from Japanese waters is provided, listing 19 families, 45 genera, and 120 species as valid. Applications of the following species names to forms previously recorded from Japanese waters are regarded as uncertain: Amphiporus cervicalis, Amphiporus depressus, Amphiporus lactifloreus, Cephalothrix filiformis, Cephalothrix linearis, Cerebratulus fuscus, Lineus vegetus, Lineus bilineatus, Lineus gesserensis, Lineus grubei, Lineus longifissus, Lineus mcintoshii, Nipponnemertes pulchra, Oerstedia venusta, Prostoma graecense, and Prostoma grande. The identities of the taxa referred to by the following four nominal species require clarification through future investigations: Cosmocephala japonica, Dicelis rubra, Dichilus obscurus, and Nareda serpentina. The nominal species established from Japanese waters are tabulated. In addition, a brief history of taxonomic research on Japanese nemerteans is reviewed.     

Weitere Studien über die Regeneration der Nemertinen

Ohne ZusammenfassungDie drei ersten Teile dieser Studien, welche I. Technisch-methodische Vorbemerkungen, Beobachtungen am Lebenden usw., II. Manche Strukturunterschiede zwischen der breiten und dünnen Form desLineus ruber Müll. usw., und III. Die inneren Vorgänge bei der Regeneration der Vorderteile (Kopfabschnitte) desLineus ruber — enthielten, wurden in diesem Archiv, Bd. XXX (Festband I), 1910 veröffentlicht.     

The turbellaria fauna of a small water reservoir in Central Russia

The Turbellaria fauna has been studied in the Uvod’ Reservoir. A total of 18 species have been found representing three orders: Catenulida, Macrostoimida, and Neorhabdocoela. The patterns of Turbellaria distribution in the reservoir have been analyzed. The most abundant species have been determined: Stenostomum leucops Duges, 1828, Microstomum lineare (O.F. Müller, 1774), Bothromesostoma essenii M. Braun, 1885, and Mesostoma lingua (Abildgaard, 1789).