A phylogenetic analysis of the genus Eunice (Eunicidae, polychaete, Annelida)

Species of Eunice are distributed worldwide, inhabiting soft and hard marine bottoms. Some of these species play significant roles in coral reef communities and others are commercially important. Eunice is the largest and most poorly defined genus in Eunicidae. It has traditionally been subdivided in taxonomically informal groups based on the colour and dentition of subacicular hooks, and branchial distribution. The monophyly of Eunice and of its informal subgroups is tested here using cladistic analyses of 24 ingroup species based on morphological data. In the phylogenetic hypothesis resulting from the present analyses Eunice and its subgroups are paraphyletic; the genus may be divided in at least two monophyletic groups, Eunice s.s. and Leodice , but several species do not fall inside these two groups. Most of the traditional characters used in the taxonomy of Eunice are homoplasies; however, characters used for the first time in this study, such as certain jaw characters and characters derived from a close examination of chaetal variation along the body, are promising sources of phylogenetic signal.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 413–434.     

Jaw growth and replacement in<Emphasis Type="Italic"> Ophryotrocha labronica</Emphasis> (Polychaeta,Dorvilleidae)

Ophryotrocha labronica, as typical for Eunicida, has a complex jaw apparatus consisting of ventral mandibles and dorsal maxillae. Mandibles are not replaced but are retained throughout life. Larval mandibles have adult-sized cutting plates but their proximal shafts lengthen and enlarge as the worm grows. The maxillary apparatus of O. labronica undergoes three moults or replacements. The initial, or larval maxillae, consisting of two paired basal plates and two paired free denticles, develop in the unreleased larvae. They are replaced in the 5-setiger juvenile by the P1-maxillae consisting of falcate forceps and six denticles. The second moult occurs in the 8- to 9-setiger juveniles and results in the P2-maxillae with bidentate forceps and seven denticles, and the third and final moult results in the K-maxillae and seven denticles. The K-maxillae develop in 9- to 12-setiger males and 13- to 15-setiger females and are not replaced but enlarge proximally. Thus the K-forceps can be traced back through the P2-forceps, P1-forceps, to the larval basal plates, indicating the apomorphic state of the K-forceps. Three pulp cavities, separated by darker fusion lines are visible in weakly sclerotised young K-forceps suggesting the fusion of three separate elements. It is concluded that the Ophryotrocha forceps are homologous to the superior and probably inferior basal plates of other dorvilleids. The internal structure of the Ophryotrocha forceps demonstrates that they are not homologous to the labidognath maxilla I as has been suggested.     

Homology of prostomial and pharyngeal structures in eunicida (Annelida) based on innervation and morphological similarities

Eunicidan bristle worm families are commonly identified by the shape of their prostomia and pharyngeal structures. However, current hypotheses of homology among these structures are conflicting, making it difficult to assess morphological evolution, reconstruct phylogeny, and produce a stable classification. To generate more consistent hypotheses of homology among eunicidan anterior structures, the author examined the anterior morphology and the nervous system stained with anti‐α‐tubulin and serotonin antibodies in representative species of Eunicidae, Onuphidae, Oenonidae, Dorvilleidae and Lumbrineridae. The shape of the brain varied conspicuously among families; however, it has mostly the same commissures (usually two of the dorsal and five of the ventral roots of the circumoesophageal connective). The stomatogastric system is also conservative in composition, having two main pairs of stomatogastric nerves which vary in their relative position among the different families. Innervation similarities combined with correspondence and topological morphological similarities made it possible to present explicit hypotheses of primary homology of features, such as buccal lips, pharyngeal fold, and dorsolateral fold anterior extension. Buccal lips are present in all families; however, ventral pads on the prostomium of the Dorvillea line of Dorvilleidae are anterior prolongations of the pharyngeal fold and not buccal lips. All examined taxa, except dorvilleid species, have conspicuous dorsolateral fold anterior extension. In Eunicidae, this anterior extension is a transverse band, while in other families it is a pair of folds. Observed similarities also gave insights on the homology of maxillary elements of Dorvilleidae, providing background knowledge for future studies. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Ultrastructure of pigmented adult eyes in errant polychaetes (Annelida): implications for annelid evolution

The evolution of photoreceptor cells and eyes in Metazoa is far from being resolved, although recent developmental and structural studies have provided strong evidence for a common origin of photoreceptor cells and existence of sister cell types already in early metazoans. These sister cell types are ciliary and rhabdomeric photoreceptor cells, depending on which part of each cell is involved in photoreception proper. However, a crucial point in eye evolution is how the enormous structural diversity of photoreceptor cells and visual systems developed, given the general molecular conservation of the photoreceptor cells. One example of this diversity can be observed in Annelida. Within the polychaetes the errant forms, taxon Aciculata, constitute the only group possessing true multicellular eyes in the adult stage. Thus far these organs have been investigated only in taxa of Phyllodocida, a subgroup of Aciculata. Data on Eunicida and Amphinomida as well as certain phyllodocidan taxa had been lacking. The ultrastructure of these adult eyes was investigated in various species of errant polychaetes, belonging to Amphinomidae, Eunicidae and Hesionidae, to elucidate whether they provide any phylogenetic clues regarding either the evolution of visual systems in Annelida or lophotrochozoan phylogeny in general. These eyes are composed of numerous supportive pigment cells and rhabdomeric photoreceptor cells and sometimes additional cell types. As a rule the pigment and rhabdomeric cell types form a continuous epithelium in which the two types intermingle. Presence of granules with shading pigment in sensory cells is a common feature but is apparently restricted to a taxon comprising Phyllodocida and Eunicida s. str. Very likely a lens-like structure does not belong to the ground pattern of annelid eyes, despite its presence in Phyllodocida. These lens-like structures are formed by secretions or cellular processes of the pigment cells. In many species the eye cup communicates with the exterior via a small cuticularized canal. This canal is interpreted as a rudiment due to the mode of formation in the epidermis. With respect to current phylogenetic hypotheses, these multicellular eyes have either been developed in the stem species of a taxon Aciculata nested within the polychaetes or have been evolved in the stem lineage of Annelida. Similarities to gastropod eyes are interpreted as convergent and not as indication of common origin. Except for the photoreceptor cells proper, the structure of the adult eyes in polychaetes most likely does not help to resolve lophotrochozoan phylogeny.     

Ultrastructure of pigmented eyes in Dorvilleidae (Annelida,Errantia, Eunicida) and their importance for understanding the evolution of eyes in polychaetes

Among polychaetes, the errant forms are the only group known so far possessing true multicellular eyes in adults which are preceded by bicellular larval eyes in many species. Most likely, two pairs of such eyes showing a specific structure belong to the ground pattern of Errantia = Aciculata. However, these eyes have primarily been investigated in only two subgroups of Errantia, but data on the third main taxon, Eunicida, are available for only two taxa. In the present investigation, the eyes in two additional species of Eunicida, the dorvilleids Protodorvillea kefersteini and Schistomeringos neglecta, were studied. In P. kefersteini, usually described as possessing one pair of small eyes, two pairs could be detected, whereas in S. neglecta only one pair was found. Each eye is made up of rhabdomeric photoreceptor cells, pigment cells and unpigmented supportive cells. Lenses or vitreous bodies are absent. From their structure most likely all eyes represent adult eyes and even the small anterior eyes in P. kefersteini structurally resemble miniaturized adult eyes. Neither persisting larval eyes nor unpigmented rhabdomeric ocelli were found in the two species. The observations in Dorvilleidae confirm the hypothesis of a common origin of adult eyes in Errantia.     

Comparative analysis of the nervous systems in presumptive progenetic dinophilid and dorvilleid polychaetes (Annelida) by immunohistochemistry and cLSM

Entire nervous systems of the dinophilids Dinophilus (two species) and Trilobodrilus (three species) and the dorvilleids Parapodrilus psammophilus and Ophryotrocha gracilis (larva) were stained with antisera directed against serotonin, Phe‐Met‐Arg‐Phe‐NH₂ (FMRFamide) and acetylated α‐tubulin and analysed by confocal laser scanning microscopy (cLSM). Adult dinophilids and the dorvilleid larva exhibit the same structure of the ventral nerve cord, with two main nerves spaced far apart, one median and two paramedian nerves. A serotonergic plexus is situated between the paramedian nerve pair, above the ventral locomotory ciliary band. These similarities between adults and larva corroborate the presumed progenetic origin of dinophilids. However, since larval nervous systems of other polychaete taxa also seem to be organized in this way, this result cannot support the view that dinophilids originate from dorvilleids. In P.psammophilus the main nerve cords are widely separated only in the last segment, indicating that this pattern may be correlated with the absence of parapodia. The unpaired median nerve of dinophilids, P. psammophilus and many other polychaetes, is considered to be part of the basic annelid body plan. The ground pattern of the ventral paired dinophilid ganglia is represented by three (anterior, main, posterior) commissures, conserved in most of the ganglia in the Dinophilus species and mostly reduced to a main commissure in the Trilobodrilus species. The dinophilid species and P. psammophilus possess six pairs of ganglia indicating six trunk segments – in contrast to former views. The two rings behind the prostomium in both the dinophilid and the dorvilleid species contain one pair of ganglia only, corroborating the presumed homology of this peristomial region in the two taxa. The Dinophilus nervous system with 12 longitudinal nerves and three perpendicular nerve rings per segment resembles orthogonal nervous structures characteristic of platyhelminths.     

Early Middle Ordovician scolecodonts from north‐western Argentina and the emergence of labidognath polychaete jaw apparatuses

Scolecodonts provide fossil evidence of the evolution and diversification of jaw‐bearing polychaetes from the latest Cambrian onwards. However, their record before the Darriwilian (Middle Ordovician) is scarce worldwide, which limits our understanding of key evolutionary events. One such event is the emergence of taxa possessing the asymmetrical labidognath‐type jaw apparatus architecture, which became common in the Middle Ordovician and is often dominant throughout the Palaeozoic. Here, we document a small collection of Dapingian scolecodonts from the Capillas section, Sierras Subandinas, north‐western Argentina. The isolated elements recovered allowed us to reconstruct the distinctive jaw apparatus, and to introduce a new taxon, Andiprion paxtonae gen. et sp. nov. The maxillary apparatus of Andiprion is intermediate between the symmetrognath type of the Early Ordovician Kadriorgaspis and the labidognath type that is present in polychaetaspids and related taxa. The apparatus architecture of Andiprion corresponds best to the labidognath type, but the morphology of the individual jaws suggests that it may be the most primitive representative of this lineage currently known. We propose that Andiprion‐like forms were ancestral to polychaetaspids, polychaeturids and ramphoprionids. The Capillas collection provides supporting evidence for the evolutionary homology of the ‘basal plate’ and the left first maxilla. Thus the labidognath‐type asymmetry, with an unpaired left maxilla III, developed as a result of gradual reduction in size of the first right jaw (‘basal plate’) in front of the carriers, instead of loss or fusion of anterior maxillae.