Progenesis in Eunicida ("Polychaeta," Annelida)--separate evolutionary events? Evidence from molecular data

The Eunicida are a well-defined annelid taxon currently comprising Dorvilleidae, Lumbrineridae, Eunicidae, Onuphidae, and Hartmanniellidae. Especially the Dorvilleidae include several species of small body size and apparently simple organisation, resembling larvae and juveniles of larger species. One hypothesis to explain the evolution of such species is progenesis. By this criterion the Dinophilidae were included in this group as well. Although their overall similarity suggests monophyly, parallel evolution of such progenetic taxa cannot be excluded. To infer the phylogeny of Eunicida and their presumed relationship to Dinophilidae and progenetic dorvilleids, 18S rDNA sequences from various Dinophilidae, Dorvilleidae, and members of other Eunicida were determined. The data matrix were extended by adding sequences from several annelid and molluscan species and analysed with maximum-parsimony, distance, and maximum-likelihood algorithms. Highly significant bootstrap-values support monophyly of the Dinophilidae, of a clade enclosing the Eunicidae and the Onuphidae, and of a dorvilleid group comprising six species including the presumed progenetic Parapodrilus psammophilus. The position of the Dinophilidae remains unresolved, so that their inclusion within the Eunicida can neither be corroborated nor rejected. However, a constraint monophyly of Dinophilidae and P. psammophilus is rejected by several tests, substantiating the hypothesis of separate progenetic evolutionary events. Furthermore, the molecular data indicate polyphyly of the Dorvilleidae.     

Interstitial Dorvilleidae (Annelida, Polychaeta) from Europe, Australia and New Zealand

Four dorvilleid species new to science are described from intertidal and subtidal sandy sediments. For three of them new genera are erected: Coralliotrocha natans gen. et sp.n., Microdorvillea otagensis gen. et sp.n. and Pusillotrocha Åkessoni gen. et sp.n. One species represents the first record of the genus Pettiboneia from Australia, P. australiensis sp.n. Ophryotrocha minuta Levi, 1954 redescribed from Helgoland is transferred to a different genus, Arenotrocha gen.n. The present taxa belong to a morphological series of species with an increasing degree of neoteny in the family Dorvilleidae.     

A generic revision of the Dorvilleidae (Polychaeta), with six new species from the deep North Pacific

PETER A. JUMARS
The Dorvilleidae are divided into eight genera: Dorvillea, Schistomeringos (new name), Protodorvillea, Meiodorvillea (new genus), Exallopus (new genus), Ophryotrocha, Parophryotrocha , and Apophryotrocha (new genus). Periodic maxillary replacement is found to occur in a manner which makes many often cited maxillary characters taxonomically unreliable. Therefore, the pharyngeal characters of all the type-species are redescribed or newly described. The discovery of new forms and the present interpretation of hard jaw parts permit construction of a phylogenetic scheme with Dorvillea as the most primitive genus containing living species. Suggestions are given for the advantageous use of the phenomenon of maxillary replacement in autecoiogical investigation.
Evidence is presented against the cosmopolitan distribution of Schistomeringos rudolphi , a species often noted in nearshore pollution studies. Newly described species are Dorvillea batia, Schistomeringos mediofurca, Meiodorvillea apalpata, Exallopus cropion, Ophryotrocha profunda , and Apophryotrocha mutabiliseta , all from the deep North Pacific. A key to the genera is provided, as are listings of all the known species within each genus.     

Petrocha notogaea gen. et sp.n., a neotenic interstitial polychaete (Dorvilleidae) from the Hauraki Gulf, New Zealand

Petrocha notogaea represents a new interstitial dorvilleid genus from subtidal medium coarse sands. With its distinctly neotenic characters, such as one pair of short palps, ciliary rings on all parts of the body, setigers with uniramous parapodia, simple capillaries, simple heterogomph falcigers and comparatively small size (1.2–1.4 mm), it shows close phylogenetic affinity to several other taxa of the Dorvilleidae, such as Ikosipodus, Microdorvillea, Arenotrocha and Pusillotrocha , which have only been described in the last four years.     

Phylogeny of Nerillidae (Polychaeta, Annelida) as inferred from combined 18S rDNA and morphological data

A phylogeny of the meiofaunal polychaete family Nerillidae based on morphological, molecular and combined data is presented here. The data sets comprise nearly complete sequences of 18S rDNA and 40 morphological characters of 17 taxa. Sequences were analyzed simultaneously with the morphological data by direct optimization in the program POY, with a variety of parameter sets (costs of gaps: transversions: transitions). Three outgroups were selected from the major polychaete group Aciculata and one from Scolecida. The 13 nerillid species from 11 genera were monophyletic in all analyses with very high support, and three new apomorphies for Nerillidae are identified. The topology of the ingroup varied according to the various parameter settings. Reducing the number of outgroups to one decreased the variance among the phylogenetic hypotheses. The congruence among these was tested and a parameter set, with equal weights (222) and extension gap weighted 1, yielded minimum incongruence (ILD). Several terminal clades of the combined analysis were highly supported, as well as the position of Leptonerilla prospera as sister terminal to the other nerillids. The evolution of morphological characters such as segment numbers, chaetae, appendages and ciliation are traced and discussed. A regressive pathway within Nerillidae is indicated for several characters, however, generally implying several convergent losses. Numerous genera are shown to require revision. © The Willi Hennig Society 2005.     

Progenetic species in polychaetes (Annelida) and problems assessing their phylogenetic affiliation

Progenesis is defined as the retention of ancestral juvenilecharacters by adult stages of descendants due to an accelerationof the sexual maturation and thus is often regarded as a fastevolutionary process. Several small, meiofaunal polychaetes,such as Dinophilidae, some "Dorvilleidae" (for example, Parapodrilus),and Protodrilida, exhibit morphological simplicity in that theylack features typical of larger polychaetes, for example, parapodiaand/or head appendages. Due to the general resemblance of adultmeiofaunal polychaetes to juveniles of larger forms, progenesishas been invoked to explain evolutionary origins of many smallertaxa with increasing frequency over the past 4 decades. In thisreview, I summarize the interstitial species of polychaetesfor which progenetic origin has been suggested on the basisof morphology. However, critical examination of morphologicaldata that includes larval features reveals that autapomorphiccharacters uniting supposed progenetic taxa to specific annelidlineages are often missing. Typically larval and juvenile characters,which are argued to support hypotheses of progenetic origin,are often widely dispersed, homoplastic features. Because ofthis situation, molecular data seem to be the most reliablesource for phylogenetic inference. However, other biologicaldata, for example, from life history and morphology, are necessitiesto substantiate the progenetic evolution of these species.     

Parapodrilus psammophilus nov. gen. nov. spec., eine neue Polychaeten-Gattung aus dem Mesopsammal der Nordsee

Zusammenfassung 1. Ein neuer Polychaet,Parapodrilus psammophilus nov. gen. nov. spec., wurde im Eulitoral der Nordsseinsel Sylt entdeckt.2. Körperform, Größe und Gliederung dieses Interstitialbewohners zeigen gewisse Beziehungen zur Familie Dinophilidae der Archiannelida.3. Die Existenz einästiger, mit Borsten und Aciculae versehener Parapodien unterscheidet die neue Art jedoch scharf von allen Genera der Dinophilidae. Der Mangel eines für die Archianneliden typischen muskulösen Schlundsackes macht darüber hinaus die Einreihung in diese Gruppe problematisch.4. Der Verfasser beschränkt sich daher vorläufig auf eine Beschreibung des neuen Polychaeten und auf eine kurze Diagnose der neuen Gattung. Die Einordnung in das System der Polychaeta bleibt einer eingehenderen anatomischen Untersuchung vorbehalten.

---

Parapodrilus psammophilus nov. gen. nov. spec., a new polychaete genus from the mesopsammal of the North Sea

A new polychaete genus,Parapodrilus psammophilus, is described on the basis of individuals caught in the sandy interstitial zone on the island Sylt (North Sea, German Bight). The main features of the new polychaete are: length up to 650µm; body divided into a prostomium (without any appendages), 7 segments and pygidium; segments 3 to 6 carry simple parapods as well as setae and aciculae.

     

Ikosipodus carolensis gen. et sp.n., an Interstitial Neotenic Polychaete from North Carolina, U.S.A., and Its Phylogenetic Relationships within Dorvilleidae

Ikosipodus carolensis gen. et sp.n. from shallow subtidal sands off the coast of North Carolina is described. The species shows conspicuous neotenic features. The sexes are separate and dimorphic. The juveniles resemble the adults greatly. A seminal receptacle in the male, of which the fine structure was investigated, is the first proof of direct sperm transmission in the Dorvilleidae. The new form has been classified into a tentative phylogenetic dendrogram of the dorvilleid genera     

Anatomy and Ultrastructure of Ventral Pharyngeal Organs and their Phylogenetic Importance in Polychaeta (Annelida). IV. The Pharynx and Jaws of the Dorvilleidae

An anatomical and ultrastructural investigation of the ventral pharyngeal organ, jaws and replacement of jaws was carried out in Ophryotrocha gracilis and Protodorvillea kefersteini (Dorvilleidae). The pharynx exhibits the following features: jaw apparatus present, consisting of paired mandibles and rows of maxillary plates, the latter are fused to form a single piece; cuticular jaws electron-dense, in P. kefersteini with collagen fibres; muscle bulbus solid, composed of muscle cells only; parallel running myofilaments, centrally located mitochondria and nuclei, bulbus epithelium containing the mandibles and gland cells, maxillary plates lying on folds corresponding to a tongue-like organ, connected with mandibles by longitudinal investing muscles; numerous gland cells not united to distinct salivary glands. Development of jaw replacements occurs in epithelial cavities beside the functional maxillae. Shape of maxillary plates is preformed by microvilli carrying cell processes. Maxilloblasts change their shape during the development. Synapomorphic structures occurring in ventral pharyngeal organs of other species outside the Eunicea are not present and even the closely related Dinophilidae exhibit a completely different pharyngeal organ. Therefore, convergent evolution of these organs is the most probable explanation. These findings do not agree with the hypothesis of the homology of the ventral pharyngeal organs in the Polychaeta.     

La famille des Cicerinidae (Turbellaria, Kalyptorhynchia)

The present study deals with 12 species of Cicerinidae. Further details are given for 6 species previously described: Toia ycia Marcus, Cicerina eucentrota Ax, Nan-norhynchides vividus Brunet, JV. harparius Brunet, N. corneus Brunet, Didiadema picardi Brunet. Two new species are described, Toia calceformis sp.n., different from T. ycia by half-boot shape of the cirrus, and Paracicerina globulosa sp.n., characterized by very short (4 μm) cuticular spermatic ducts and a spherical copulatory bulb provided with a spiny cirrus (35–40 μm long). Histological studies of Ptyalorhynchus coecus Ax confirm the generic separation of Ptyalorhynchus and Paracicerina; the main difference is the glandular girdle structure of the proboscis in Ptyalorhynchus which is split into two areas (inner and outer one) by basal circular muscles; the intensively stained secretion of the inner area is localized in 15 membranous canals. Pocillorhynchus gen.n., with two species, P. agilis sp.n. and P. minutus sp.n., is closely related to the genera Toia and Nannorhynchides , but differs from them by the following characters: proboscis with broad glandular girdle, devoid of circular muscle bulb, cuticular spermatic ducts long and thin (95–125 μm). From the comparison of their morphological and anatomical features, two groups of genera are distinguished in Cicerinidae; first group including Toia - Nannorhynchides - Pocillorhynchus , second group including Cicerina - Paracicerina - Etmorhynchus - Ptyalorhynchus - Zonorhynchus - Xenocicerina - Didiadema. Special attention is paid to proboscis structure. The study ends with an identification key for all known Cicerinidae.     

Revision of some comasterid genera from Australia (Echinodermata: Crinoidea), with descriptions of two new genera and nine new species

Morphological characters used in comasterid taxonomy are reappraised. As a result, form and distribution of pinnule combs are newly promoted as primary characters of generic recognition. Form and arrangement of cirri and division series, as used by A. H. Clark (1931), are rejected as characters important in recognizing taxa higher than species. Five of 12 comasterid genera reported as occurring in Australian waters (H. L. Clark, 1946) are critically examined. Of these genera, Comantheria A. H. Clark, 1909b is referred to the synonymy of Comanthus A. H. Clark, 1908a, Comanlhoides A. H. Clark, 1931 is referred to the synonymy of Cenolia A. H. Clark, 1916b, and Comanthina A. H. Clark, 1909b is maintained. In addition, two new genera are described. These genera are related by the occurence of the first brachial syzygy at 3 + 4 on all arms. Twenty-six species within the five genera are now recognized in Australian waters. Nine new species are described, and the 17 existing species are rediagnosed and critically reappraised. The revision involves taxa distributed throughout the Indo-west Pacific region, as far west as S Africa.     

Ultrastructure of Tricornia muhezae N. G., N. Sp. (Microspora, Thelohaniidae), a Parasite of Mansonia africana (Diptera: Culicidae) from Tanzania

ABSTRACT. Collections of Mansonia africana mosquito larvae were made at one site in N.E. Tanzania in 1985 and 1987 and from two additional sites, both within about 2 mi of the original one in 1987. An octosporous microsporidian, present at all three sites, was found in both years infecting from 7 to 22% of larvae. Spores (stained in Giemsa) measured 3.0 μ m × 0.25 μ m × 2.25 μ m × 0.26 μ m. Ultrastructurally, spores were seen to have an anterior rim surrounding a depressed area where the endospore was at its thinnest. In transmission electron microscopy section, the rim appeared as two processes into which all layers of the wall extended. At the posterior end all layers of the wall extended into a simple knob-like structure which could be interpreted as a section through a crest running longitudinally around the spore. The polar filament was anisofilar, with two anterior coils of greater diameter than the three posterior coils. Although most closely resembling the genera Amblyspora and Parathelohania in the family Thelohaniidae, the species in M. africana differs from the former, which has oval spores, broadly rounded at the ends, and from the latter, which has a prominent, ridged posterior extension to the spores. The new species has been placed in a new genus and the name Tricornia muhezae proposed.     

A reappraisal of the family Comasteridae A. H. Clark, 1908 (Echinodermata: Grinoidea), with the description of a new subfamily and a new genus

The family Comasteridae is divided into four subfamilies on the basis of syzygy positions in the division series and proximal arm. Use of this character results in major differences in perceived relationships between genera compared with that obtained when traditional subfamilial characters are used. The primary character previously used to determine comasterid subfamilies, cirrus ornamentation, is rejected as unimportant in recognizing taxa any higher than species. Seventeen genera, including a new genus from the tropical western Atlantic, and 99 species are now recognized in the Comasteridae. Four of these genera and five species cannot be placed within the newly diagnosed subfamilies. Characters of generic importance are considered to be pinnule comb form, mouth position and, to a lesser extent, the characters traditionally used to distinguish genera.     

Phylogenetic relationships within Nephtyidae (Polychaeta,Annelida)

Ravara, A., Wiklund, H., Cunha, M. R. & Pleijel, F. (2010). Phylogenetic relationships within Nephtyidae (Polychaeta, Annelida). —Zoologica Scripta, 39, 394–405. We present the first phylogeny of nephtyids, a common, soft‐bottom living polychaete family comprising five genera and over 100 species. Characters used to distinguish nephtyid genera are a matter of controversy and considerable confusion remains as to the generic delineations. The phylogeny is estimated with molecular data from the mitochondrial genes cytochrome oxidase I and 16S rDNA, the nuclear genes 18S rDNA and 28S rDNA and morphological data. The results reveal two well‐supported major clades, corresponding in part to the two main genera of the family, Aglaophamus and Nephtys. The species Nephtys pulchra and Nephtys australiensis are transferred to Aglaophamus, and new diagnoses for the genera are provided. Dentinephtys is synonymized with Nephtys, and Nephtys cornuta is sister to the remaining nephtyids and is referred to the new genus Bipalponephtys, together with Nephtys danida and Micronephthys neotena. Micronephthys is sister to Nephtys and Inermonephtys is of uncertain position.     

A systematic account of the Questidae (Annelida, Polychaeta), with description of new taxa

Questidae is a poorly known polychaete family with many unusual characters, particularly with regard to its reproductive organs. Its taxonomic position remains uncertain, which can be partly referred to the limited material described to date. New material, mostly from shallow sublittoral sediments, shows that questids have a circummundane distribution. Questa Hartman, 1966 is revised to include Novaquesta Hobson, 1970 as a junior synonym. Eight species of questids are assigned to Questa , four of them are new to science: Questa bicirrata sp. n., Q. paucibranchiata sp. n., Q. mediterranea sp. n., and Q. riseri sp. n. Character patterns of the questid polychaetes described so far are cladistically analysed. The limitation of informative characters causes ambiguity in the most parsimonious reconstruction of the phylogeny of the family, and therefore, the Questidae is proposed to be classified as comprising only one genus.     

Taxonomic Review of the Orders Mysida and Stygiomysida (Crustacea,Peracarida)

The order Mysida (2 families, 178 genera, 1132 species) contains species across a broad range of habitats, such as subterranean, fresh, brackish, coastal, and surface to deep-sea habitats. The Stygiomysida (2 families, 2 genera, 16 species), however, are found primarily in subterranean waters, but always in waters with a marine influence. The Mysida and Stygiomysida body is divided into three main regions: cephalon, thorax, and abdomen. They are shrimp-like in appearance, containing morphological features earlier referred to as defining a "caridoid facies". The shrimp-like morphology was to some extent diagnostic for the historic Decapod taxon Schizopoda, containing the Nebalia, Mysida, Lophogastrida, and Euphausiacea. In 1904 the concept of Schizopoda was abandoned, and the Mysidacea (Mysida and Lophogastrida) along with Cumacea, Amphipoda, Isopoda, and Tanaidacea were placed in a new taxon, the Peracarida. Later discoveries of groundwater mysids led to the establishment of Stygiomysida, but placement to either Lophogastrida or Mysida remained unclear. The presence of oostegites and absence of podobranchiae, coupled with non-statocyst bearing uropods have been used to classify the Stygiomysida as a primitive Mysida family, comparable to Petalophthalmidae. On the other hand, equally suggestive characters, but for a Lophogastrida affiliation, was suggested for the archaic foregut characters and again, non-statocyst bearing uropods. With the inclusion of DNA sequence data of ribosomal genes, sister group relationships between Stygiomysida, Lophogastrida, and Mictacea within the Peracarida are observed, which supports a classification of the Stygiomysida as a separate order removed from the Mysida.