Anatomy and ultrastructure of ventral pharyngeal organs and their phylogenetic importance in Polychaeta (Annelida)

Summary A comparative anatomical and ultrastructural study of ventral pharyngeal organs (pharyngeal bulbs) was carried out in two species of the Dinophilidae: Dinophilus gyrociliatus and Trilobodrilus axi. Special attention was paid to the fine structure of the stomodeal epithelium, cuticle, glands, muscles, and myoepithelial junctions. The differences between the species are very slight. The pharyngeal organ of the Dinophilidae is characterized by the following features: solid muscle bulbus made up of muscle cells only, bulbus muscle cells with two myofilament systems crossing at an angle of about 90°, gap junctions between these muscle cells, bulbus projects into a pharyngeal sac and bears rostrally a specific epithelium and cuticle, no bulbus glands, no investing (= sagittal) muscles, specific cuticle ultrastructure, cilia of ascending oesophagus with asymmetric tips, specific structure and position of salivary gland openings. The phylogenetic importance of these structures is discussed. Some of these characters are clearly autapomorphic features of the Dinophilidae and no common derived structures to other families with a ventral pharyngeal organ are present. Therefore, it is most likely that the dinophilid pharyngeal organ evolved independently. These findings do not agree with the hypothesis of the unity of the archiannelid families (Polygordiidae, Protodrilidae, Saccocirridae, Nerillidae, Dinophilidae, and Diurodrilidae) established on the basis of an assumed structural similarity of their ventral pharyngeal organs.Abbreviations bb basal body - bep bulbus epithelium - bl basal lamina - bm bulbus muscle - c cilium - cc coelenchyme cell - cm circular muscle - cr caudal rootlet - cu cuticle - dblm dorsal bulbus longitudinal muscle - dlm dorsal longitudinal muscle - dsn dorsal stomatogastric nerve - dy dyad - el electron-dense layer - fl fibrous layer - fi filaments - g Golgi apparatus - gl gland cell - hv homogeneous vesicle - l lipid droplet - la external lamina - lal lamellar layer - ll lower lip - lm longitudinal muscle - ly lysosome - m mitochondrion - mo mouth opening - mt microtubule - mv microvillus - mvp microvillar process - n nucleus - nu nucleolus - oes oesophagus - pcom preoral commissure - phf pharyngeal fold - phl pharyngeal lumen - phs pharyngeal sac - pms peripheral myofilament system - r rootletlike structure - rer rough endoplasmic reticulum - rr rostral rootlet - s sarcoplasmic reticulum - sc salivary canal - scom suboesophageal commissure - sd septate desmosome - ser smooth endoplasmic reticulum - sg secretory granule - sgl salivary gland - sn stomatogastric nerve - st stomach - step stomodeal epithelium - tep transitional epithelium - tf tonofilaments - va vacuole - vlm ventral longitudinal muscle - vsn ventral stomatogastric nerve - z z-element - za zonula adherens     

Anatomy and ultrastructure of ventral pharyngeal organs and their phylogenetic importance in Polychaeta (Annelida)

Summary Transmission electron microscopic studies were carried out on the ventral pharyngeal organs in Ctenodrilus serratus and Scoloplos armiger. The pharyngeal organs are composed of a muscle bulbus and a tongue-like organ. In both species the muscle bulbus consists of transverse muscle fibres and interstitial cells with voluminous cell bodies and dorsoventral tonofilaments; the investing muscle runs into the tongue-like organ; the nuclei of the investing muscle fibres are located in caudal bulges; salivary glands are not present, but numerous gland cells occur in the bulbus epithelium. The tongue-like organ, however, is formed by lateral folds (C. serratus) or a bridge-like structure (S. armiger). The specific structure of the bulbus muscle is probably a homologous characteristic also occurring in several other polychaete families. The phylogenetic importance of this ventral pharynx is discussed and a hypothesis is suggested to explain the differentiation of certain other ventral pharyngeal organs from this probably primitive type.     

The Baikalian genus Rhyacodriloides in Europe: phylogenetic assessment of Rhyacodriloidinae subfam. n. within the Naididae (Annelida)

Martin, P., Martínez‐Ansemil, E. & Sambugar, B. (2010). The Baikalian genus Rhyacodriloides in Europe: phylogenetic assessment of Rhyacodriloidinae subfam. n. within the Naididae (Annelida). —Zoologica Scripta, 39, 462–482. Two new species of the oligochaete genus Rhyacodriloides Chekanovskaya, Rhyacodriloides aeternorum sp. n. and Rhyacodriloides latinus sp. n., are described from subterranean water bodies of Italy and Slovenia. A comparison with the known species of this genus, Rhyacodriloides abyssalis Chekanovskaya, 1975 and Rhyacodriloides gladiiseta Martin & Brinkhurst, 1998, both from Lake Baikal, shows that the enigmatic ‘cellular masses’ of the latter two species must be interpreted as different, not homologous structures. As a result, R. gladiiseta is to be ascribed to the Phallodrilinae, a primarily marine naidid subfamily, mentioned for the first time in Lake Baikal, and placed in its own genus, Phallobaikalus gen. n. The two new species are morphologically very similar, but their penial setae differ slightly. The phylogenetic relationships of R. latinus sp. n. and R. abyssalis within the Naididae (formerly the Tubificidae) were investigated using a combination of three genes, one nuclear (18S rDNA) and two mitochondrial (12S rDNA and 16S rDNA). A fragment of the mitochondrial COI gene, used as a barcode, also genetically characterized all Rhyacodriloides species. Sequences of 34 Naididae were obtained from EMBL, representative of five naidid subfamilies, and including five oligochaete outgroups. The data were analysed by parsimony, maximum likelihood and Bayesian inference. Taken in combination, the three genes investigated confirm that the two Rhyacodriloides species analysed are closer to each other than to any other naidid species. However, they are separated by 16S and COI distances that amount to 18.5% and 27.2%, respectively, suggesting an ancient separation between species, in good accordance with their present biogeographic distribution. Rhyacodriloides cannot be considered as a rhyacodriline, as assumed so far, as they never appeared related to this subfamily in any analysis considered. In contrast, they appear at the base of a naidid group, including the Tubificinae, the Phallodrilinae, the Limnodrilinae, as well as Branchiura sowerbyi, a species whose phylogenetic association with the rhyacodrilines has been questioned for a long time. Despite a lack of phylogenetic support, this position is congruent with a morphological reassessment of the Rhyacodrilinae, and strongly supports the erection of a new naidid subfamily to accommodate Rhyacodriloides.     

Inference of phylogenetic relationships within Fabriciidae (Sabellida,Annelida) using molecular and morphological data

Cladistic relationships among fabriciids have to date been explored in the context of adult morphology, but resolution has been declining as more species are described. In this study, we incorporated data on the reproductive system, including features related to the male sperm and sperm storage by females, to supplement existing data on adult morphology (for a total of 50 characters). Three nuclear DNA markers (18S rDNA approximately 1800 bp, the D1 region of 28S rDNA approximately 320 bp, and histone H3 approximately 330 bp) were sequenced from 21 species of fabriciids. We assessed the phylogeny of Fabriciidae based on an integrative analysis of these morphological and molecular characters. Our results show that, in addition to three previously recovered apomorphies for Fabriciidae (absence of ventral lips, modification of abdominal uncini to an elongate manubrium, and presence of branchial hearts), six more apomorphies associated with the reproductive system can be used to support this clade—spermiogenesis only in the thorax, spermiogenesis in large clusters with a central cytophore, single dorsal sperm duct, sperm nuclear projection, thickening of the sperm nuclear membrane and the sperm extra‐axonemal sheath. The results require the erection of two new genera and two new species, which are described. © The Willi Hennig Society 2010.     

Arhynchobdellida (Annelida: Oligochaeta: Hirudinida): phylogenetic relationships and evolution

A remarkable diversity of life history strategies, geographic distributions, and morphological characters provide a rich substrate for investigating the evolutionary relationships of arhynchobdellid leeches. The phylogenetic relationships, using parsimony analysis, of the order Arhynchobdellida were investigated using nuclear 18S and 28S rDNA, mitochondrial 12S rDNA, and cytochrome c oxidase subunit I sequence data, as well as 24 morphological characters. Thirty-nine arhynchobdellid species were selected to represent the seven currently recognized families. Sixteen rhynchobdellid leeches from the families Glossiphoniidae and Piscicolidae were included as outgroup taxa. Analysis of all available data resolved a single most-parsimonious tree. The cladogram conflicted with most of the traditional classification schemes of the Arhynchobdellida. Monophyly of the Erpobdelliformes and Hirudiniformes was supported, whereas the families Haemadipsidae, Haemopidae, and Hirudinidae, as well as the genera Hirudo or Aliolimnatis, were found not to be monophyletic. The results provide insight on the phylogenetic positions for the taxonomically problematic families Americobdellidae and Cylicobdellidae, the genera Semiscolex, Patagoniobdella, and Mesobdella, as well as genera traditionally classified under Hirudinidae. The evolution of dietary and habitat preferences is examined.     

Ultrastructure and functional morphology of the female reproductive organs inProtodrilus (Polychaeta,Annelida)

The morphology and function of the female reproductive organs in 6Protodrilus species are investigated by light- and transmission electron microscopy. Possible ways in which spermatozoa may enter the female coelom after leaving the spermatophore are discussed for species with and without special female reception organs. Only femaleP. rubropharyngeus andP. flavocapitatus have “dorsal organs” for spermatophore reception. The structure and function of these organs are described, as well as those of the oviduct found in 3 of the species investigated. The possible phylogenetic origin of gonoducts and different modes of oviposition within the genus are discussed. Finally, the high taxonomic significance of female traits such as dorsal organs, oviducts, cocoon glands and lateral ciliary rows in this genus is stressed.     

Further revisions of the Sabellidae subfamilies and cladistic relationships among the Fabriciinae (Annelida: Polychaeta)

Cladistic relationships of the sabellid subfamilies Fabriciinae and Sabellinae are examined in light of recent revisions of fabriciin taxa. The potential placement of Caobangia in the Sabellinae is suggested from an initial analysis of selected fabriciin species and genera. Subsequent cladistic analyses at the family level produced over 1800 cladograms, among which Caobangia is the most plesiomorphic taxon of either the Fabriciinae or Sabellinae. Successive approximations weighting reduced this ambiguity, consistently placing Caobangia in the Sabellinae. Subfamilies are emended on the basis of these results. Cladistic relationships of fabriciin taxa, exclusive of Caobangia , display topological instability among some genera and species. Genera are, however, monophyletic in all topologies. Incorporating ontogenetic data for branching ventral filamentous appendages in Augeneriella reduces ambiguity among genera and suggests alternative transformation series for ventral filamentous appendages.     

Phylogenetic relationships and evolution of Orbiniidae (Annelida, Polychaeta) based on molecular data

The phylogenetic relationships of orbiniid taxa were reconstructed based on sequence data of the mitochondrial 16S rRNA and nuclear 18S rRNA genes. Both genes were analysed separately and in combination using maximum likelihood, Bayesian inference and maximum parsimony. Regardless of the method used, a clade consisting of the investigated Orbiniidae, Methanoaricia dendrobranchiata and Questa was strongly supported by the 18S dataset. The analysis of the combined dataset suggests inclusion of M. dendrobranchiata within the Orbiniidae with close relationships to species of Orbinia and Phylo, rather than as a sister taxon to all other orbiniids. Evidence is given for the paraphyletic status of Leitoscoloplos , Naineris , Orbinia , Phylo and Scoloplos , which represent the most species-rich genera of the Orbiniidae. It is thus reasoned that the morphological characters presently used for genus diagnosis are not informative for cladistic analysis. No support is found for the hypothesis that taxa of the Protoariciinae represent juveniles of Orbiniinae. Instead, in the case of Protoaricia oerstedi , strong support for a progenetic origin is found.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 59−73.     

Molecular and morphological evidence of Alvinellidae relationships (Terebelliformia, Polychaeta, Annelida)

In this study we assessed the phylogenetic relationships of the hydrothermal vent polychaete group Alvinellidae, based on parsimony analyses of combined morphological and molecular data. Morphological data were obtained from newly examined specimens and literature information of 16 terminal taxa belonging to Alvinellidae, Ampharetidae, Pectinariidae, Terebellidae, Trichobranchidae, and the outgroups Oweniidae and Sabellidae. Molecular data were based on 28S rRNA from 13 of the 16 morphological terminals (10 previously published sequences plus three new ones). The combined analysis indicated the clades ((Alvinellidae, Trichobranchidae) Pectinariidae) and (Ampharetidae, Terebellidae). Alvinellidae, Ampharetidae and Terebellidae, as currently delineated, are monophyletic. The positions of Trichobranchidae and Pectinariidae contradicted traditional views, and they also had low Bremer support and merit further studies. Well-supported clades included Alvinellidae and Terebellinae. Previous statements that Alvinellidae are either nested within Ampharetidae or the sister to this taxon were not supported. The traditional but here contradicted view that Terebellidae and Trichobranchidae are closely related may be based on plesiomorphic similarities between these two taxa.     

Spermatozoon ultrastructure and phylogenetic relationships in the monogeneans (platyhelminthes)

Justine J.-L., Lambert A. and Mattei X. 1985. Spermatozoon ultrastructure and phylogenetic relationships in the monogeneans (Platyhelminthes). International Journal for Parasitology15: 601–608. New observations reported in this study together with bibliographical data allow comparisons of spermatozoon ultrastructure in 28 genera of monogeneans, belonging to 19 families. The authors propose to compare and classify monogenean spermatozoa using two simple ultrastructural characteristics: (a) the number of axonemes, 1 or 2, (b) the presence or absence of cortical microtubules. These traits make it possible to group monogenean spermatozoa in four patterns. Pattern 1 (2 axonemes plus microtubules) is characteristic of the polyopisthocotyleans (9 families). The three other patterns are found in the monopisthocotyleans. Pattern 2 (2 axonemes without microtubules) is found in the Capsalidae and Dionchidae, which seem closely related, and also in the Udonellidae, Gyrodactylidae and Euzetrema. Pattern 3 (1 axoneme plus 1 altered axoneme plus microtubules) is found in the Monocotylidae and Loimoidae. Pattern 4 (1 axoneme without microtubules) is found in the Amphibdellatidae, Ancyrocephalidae, Calceostomatidae and Diplectanidae. A phylogeny of the monogeneans is drawn from the data of comparative spermatology; this scheme coincides in many points with the phylogeny of Lambert (1980) which was based on the study of chaetotaxy and ciliated cells of the oncomiracidium.     

Fossil onychophorans from Dominican and Baltic amber: Tertiapatus dominicanus n.g., n.sp. (Tertiapatidae n.fam.) and Succinipatopsis balticus n.g., n.sp. (Succinipatopsidae n.fam.) with a proposed classification of the subphylum Onychophora

Abstract. Tertiapatus dominicanus n.g., n.sp. (Tertiapatidae n.fam.) and Succinipatopsis balticus n.gen., n.sp. (Succinipatopsidae n.fam.) (Lobopodia: Onychophora), the first Tertiary fossils of the Lobopodia, are described from Dominican and Baltic amber, respectively. Both families are characterized by the presence of simple legs lacking foot portions with claws and pads. Tertiapatidae is further characterized by soluble body pigments and oral papillae shorter than the legs. Succinipatopsidae is characterized by non-soluble body pigments and oral papillae longer than the legs. Nomenclatural changes include the erection of the class Udeonychophora n. nom. for terrestrial onychophorans with a ventral mouth, the order Ontonychophora n.nom. for extant onychophorans possessing legs with a differentiated "foot" portion, and the family Helenodoridae n.nom. for the genus Helenodora from the Carboniferous. The biogeographical significance of these fossils and their phylogenetic relationship with previously described onychophorans are discussed.     

Comparative ultrastructure of juvenile and adult nuchal organs of an annelid (Polychaeta: Opheliidae)

Opheliid nuchal organs are composed of ciliated cells, retractor muscles, and sensory cells. The perikarya of sensory cells are located in the posterior portion of the brain, and their distal processes extend along the body wall, as the nuchal nerve, and terminate just anterior to the ciliated region. The nuchal nerve of the juvenile is composed of 30–35 dendrites; the adult nuchal nerve has 35–40 dendrites. The ends of the sensory dendrites form sensory bulbs which are clustered around the olfactory chamber, and each bulb bears a modified cilium. Sensory cilia lose their axonemes and extend as microvillous-like structures into the olfactory chamber. Supportive cells delineate approximately the posterior and dorsal portions of the chamber with sensory bulbs forming the remaining ventral and anterior portions. On the lateral aspect of the chamber, cuticular matrix extends into it, and in this area supportive cells bear microvilli which extend into the matrix. The adult nuchal organ is larger than that of the juvenile, and the sensory portion of the olfactory chamber wall is expanded. Expansion of the sensory area is apparently the result of size increase in sensory bulbs and by intrusion of supportive cells between sensory bulbs.     

Ultrastructure and functional morphology of male genital organs and spermatophore formation inProtodrilus (Polychaeta,Annelida)

Summary The structure and functional morphology of lateral organs and sperm ducts, as well as the mechanisms of spermatophore formation and transfer, are investigated by means of light and electron microscopy in the genusProtodrilus. The sperm ducts are simple, ciliated, intercellular gonoducts with a funnel section surrounded by a thin muscle layer and a tube section opening externally in the anterior region of the lateral organs. No glands are present in the sperm ducts. The lateral organs are formed by long epidermal invaginations enclosing an elongate lumen into which numerous cilia project and a large number of glands open. Five to ten different gland types with strikingly distinctive secretory granules are found in the different species. In addition, special supporting cells, the so-called sponge cells, sensory cells and an underlying nervous tissue are developed in the lateral organs. It is stated that apart from some similarities to the ventral atrium ofNerilla antennata no corresponding organs are known within the Annelida. It is argued that inProtodrilus the spermatophores are formed by the lateral organs as there are a high number of glands opening into the lumen of the organ. The possible origin and genesis of the male gonoducts as well as the mode of spermatophore transfer inProtodrilus is discussed.Abbreviations used in the figures bl basal lamina - cc coelomic cell - ci ciliated cell - cir ciliary root - cr ciliary ring - cu cuticle - cv bs contractile ventral blood sinus - d dissepiment/septum - dbs dorsal blood sinus - es euspermatozoa - f funnel - fi filament - g gut - glo gland openings - lgl lateral organ gland - lm longitudinal muscle - lo lateral organ - lu lumen - mi mitochondrion - mt microtubules - mu muscle - mv microvilli - mvc microvillar crown - n nucleus - ne nervous tissue - o opening - ps paraspermatozoa - rer rough endoplasmatic reticulum - s spermatozoa - sc sponge cell - sg salivary gland - spd sperm duct - spdo sperm duct opening - t tube - tm transverse muscle - vc ventral ciliary band     

Sabellaria tottoriensis n. sp. (Annelida: Polychaeta: Sabellariidae) from shallow water off Tottori, the Sea of Japan

A new species of the genus Sabellaria Lamarck, 1812 (Annelida: Polychaeta: Sabellariidae), is described from shallow water off Tottori, the Sea of Japan. Sabellaria tottoriensis n. sp., is gregarious with tubes constructed of sand and shell debris. The new species is distinguished by the character combination of 1 or 2 pairs of nuchal spines, two forms (long and short) of opercular paleae in the middle row, with the slender blades of long ones recurved outward. Detailed morphological features of the species are described and compared with other Japanese and worldwide congeners.     

Ovalopodium desertum n. sp. and the phylogenetic relationships of Cochliopodiidae (Amoebozoa)

An amoeba isolated from a weakly saline semi-desert pond in Kazakhstan (Central Asia) resembles a small Cochliopodium in the light microscope, but has a dorsal fibrous cell coat without scales. Thus it can be identified morphologically as a new species of Ovalopodium Sawyer, 1980, and it is herein named O. desertum. Phylogenetic analysis of the SSU rRNA gene sequences of the new species and four Cochliopodium spp. sequenced additionally shows that Ovalopodium desertum is a sister clade to a robustly monophyletic Cochliopodium. The close relationship between Ovalopodium and Cochliopodium is also confirmed by the analysis of SSU rRNA secondary structure showing the specific helices in the region V5 in all species of both genera. Analysis of actin gene sequences fails to resolve the position of Ovalopodium but demonstrates that Parvamoeba Rogerson, 1993 is probably related to Cochliopodium. The position of Cochliopodiidae within Amoebozoa remains unresolved, despite our efforts to resolve it using broader taxonomic sampling of Amoebozoa, testing alternative tree topologies and removing the fast-evolving sites. Among sequenced genera, Parvamoeba and Endostelium Olive et al., 1984 are probable relatives to Cochliopodiidae. Molecular trees weakly support an inclusion of the family in Flabellinia (Discosea), but more phylogenomic data are necessary to test this hypothesis.     

Morphology and ultrastructure of the nephridial system of Hypania invalida (Grube, 1860) (Annelida,Polychaeta, Ampharetidae)

The excretory organs of the freshwater polychaete Hypania invalida have been examined using scanning and transmission electron microscopy. Three pairs of macroscopically and ultrastructurally different nephridia are present in the thorax. Intersegmental septa in the thorax are absent, with the exception of a single diaphragm between second and third chaetiger. The first pair of nephridia is anterior to this septum, the second pair crosses the septum, with the nephrostomes anterior and the ducts and the nephridiopori posterior to it, and the third pair of nephridia is entirely posterior to the diaphragm. The first two pairs of nephridia have ciliated nephrostomes of moderate size and long nephridial ducts that extend the length of the thorax. In contrast, the third pair is characterized by short ducts and very prominent nephrostomes. Macroscopically, seven different sections of nephridial duct cells can be distinguished along the length of the first two pairs of nephridia, whereas, on an ultrastructural basis, only six different regions can be identified. Only two regions of different duct cells can be recognized in the third pair of nephridia. Cells of the two anterior pairs of nephridia show typical characteristics of transport epithelia and most likely function as excretory organs. In contrast, the duct cells of the third pair are not that much differentiated and might primarily be responsible for the release of sexual products, as sperm was observed passing through these ducts. Podocyte‐like cells were observed to accompany nephridial ducts. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.