Search

A new genus of ectoparasitic marine annelids living on ctenophores, Ctenophoricola gen. nov., is described and its feeding behaviour, reproduction and developmental stages are discussed. Its unusual morphology challenged its placement within the known marine families. However, analyses of mitochondrial and nuclear sequence data showed the new genus as member of the Alciopini, a group of holopelagic annelids included within the Phyllodocidae. Ctenophoricola masanorii sp. nov. from Japan and Ctenophoricola rousei sp. nov. from the Canary Islands (Spain) are described. A third species from the Gulf of California is not formally described because the specimens are in poor condition. The new genus is characterized by having: 1) two distinctive body regions, the anterior with reduced parapodia lacking chaetae, and the posterior with long parapodia and chaetae and 2) a pair of large, elongate lensed eyes. These eyes are here described using histology and 3D reconstruction based on a Californian specimen. The two new species mainly differ in colour pattern, shape of parapodia, number of chaetae and body ciliation.

We assess the phylogenetic position of Siboglinidae (previously known as the phyla Pogonophora and Vestimentifera, but now referred to Annelida) in parsimony analyses of 1100 bp from 18S rRNA, 320 bp from the D1 region of 28S rRNA, and 107 morphological characters, totaling 667 parsimony informative characters. The 34 terminal taxa, apart from six siboglinids, include polychaete members of Sabellida, Terbelliformia, Cirratuliformia and Spionida, plus two Aciculata polychaetes as outgroups. Our results contradict most recent hypotheses in showing a sistergroup relationship between Siboglinidae and Oweniidae, and in that the latter taxon is not a member of Sabellida. Furthermore, our results indicate that Sabellariidae is not closely related to Sabellida, that Serpulidae may be nested within Sabellidae, and that Alvinellidae is nested within Ampharetidae. � The Willi Hennig Society 2004.

Phylogenetic analyses of molecular data (COI, 16SrDNA, 18SrDNA, and 28SrDNA) show that the hesionid genus Gyptis Marion & Bobretzky in Marion, 1874, and the tribe Gyptini Pleijel, 1998, are nonmonophyletic as currently delineated. We introduce the new tribe Amphidurini and the new genus Neogyptis to accommodate these new findings. Amphidurini is sister to Gyptini and Ophiodromini and includes Amphiduros Hartman, 1959, Amphiduropsis, Pleijel, 2001, Neogyptis gen. nov., and, possibly, Parahesione Pettibone, 1956. Morphologically, Amphidurini can be separated from Gyptini by the distally, rather than subdistally, inserted ventral cirri, and Neogyptis by the combination of this feature, the presence of a median antenna, and a distal ring with papillae on the proboscis. We redescribe and transfer the four species Gyptis crypta Pleijel, 1993, Gyptis mediterranea Pleijel, 1993, Gyptis plurisetis Hilbig, 1992, and Gyptis rosea (Malm, 1874) from Gyptis to Neogyptis gen. nov., and describe five new species from shallow waters in Belize, Hong Kong, off Vladivostok, and deep-sea hydrothermal vents in the Lau Basin off Fiji.

Nereimyra Blainville, 1828 (Psamathini, Hesionidae, Aciculata, Annelida) is revised based on examination of all available types and newly collected specimens. We assessed the phylogeny of Nereimyra in an analysis based on cytochrome oxidase c subunit I (COI), 16S rDNA, 18S rDNA, and 28S rDNA. The genus is delineated to include the three species Nereimyra aphroditoides (Fabricius, 1780), Nereimyra punctata (O.F. Muller, 1776), and Nereimyra woodsholea (Hartman, 1965), which are redescribed. Nereimyra punctata has a characteristic pigmentation, but otherwise there are no clear morphological characters for separating the species. Based on the molecular data we obtained strong support both for the monophyly of Nereimyra and for each of the three included species. Nereimyra punctata and N. woodsholea are sister species, and the Kimura two-parameter (K2P)-corrected COI distances between the three species are 16–23%. Syllidia Quatrefages, 1866, is sister group to Nereimyra. Previous uncertainties regarding the type species of the genus are settled to Nereis rosea Fabricius, 1780, junior synonym of N. aphroditoides. A neotype is designated for N. aphroditoides. The distribution of Nereimyra is at present restricted to the Arctic and the boreal parts of the North Atlantic, possibly extending to the Gulf of Mexico on the United States east coast. Records outside this area require verification. Castalia multipapillata Theel, 1879, and Nereimyra alvinae Blake, 1985, are of uncertain affinity and are treated as nomina dubia. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 36–51.

We investigate the genetic structure of European lineages of the polychaete Eumida sanguinea with mitochondrial COI and nuclear ITS, and demonstrate the presence of ten cryptic species. Within the E. sanguinea species complex there are six different white pigmentation patterns but only three of these are unique for a single species. No other consistent morphological differences were observed. We give new species names to seven of the lineages from our study, providing examples of combined morphological and molecular diagnoses, apply the available name E. sanguinea to one species, and leave two lineages with single representatives unnamed. We also include new data and a diagnosis for the poorly known E. notata, which also belongs to the E. sanguinea species complex.

We investigated whether the different colour morphs in Harmothoe imbricata constitute a single polymorphic species or if there are several species present. We sequenced the mitochondrial COI and the nuclear ITS1–5.8SrDNA–ITS2 region from 57 specimens representing 10 distinct colour morphs collected from Svalbard to the Swedish west coast. The resulting minimum spanning haplotype network based on mitochondrial COI unequivocally shows H. imbricata to be a single and colour polymorphic species, whereas variation in the ITS region was very limited.

We redescribeImajimaea draculaibased on newly collected specimens from the Swedish west coast and describe for the first time its association with the sea penFuniculina quadrangularis. We assess the phylogenetic position ofI. draculaiwithin Autolytinae using nuclear 18SrDNA, together with mitochondrial COI and 16SrDNA, and we estimate its intraspecific variation using nuclear ITS and mitochondrial COI. Our molecular data unequivocally show thatI. draculaibelongs to the tribe Procerini in a clade consisting ofProcerastea, VirchowiaandImajimaeathat is separated from all other Procerini taxa for which molecular data are available. No intraspecific variation is found inI. draculaifor the sequenced molecular markers.

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.

We report five different colour morphs of the hesionid polychaete genus Gyptis co-occurring in a small area in shallow water under Edithburgh jetty, South Australia. The five morphs cannot be separated using standard morphological features, but phylogenetic analyses of sequence data (COI and ITS1) unequivocally show that three species are present, introduced as Gyptis simpsonorum, new species, G. paucilineata, new species and G. polymorpha, new species. Gyptis simpsonorum has a speckled pigmentation pattern and G. paucilineata a few transverse lines on specific segments. Gyptis polymorpha is polymorphic with three different, distinct pigmentation patterns, either as dense transverse lines, as a thin longitudinal, mid-dorsal line, or as an uniformly dark dorsum. The speckled pigmentation likely represents the plesiomorphic condition, and G. propinqua is the closest known relative to these new species.

Vrijenhoekia balaenophila gen. nov., sp. nov. (Polychaeta, Hesionidae) is described from a whale carcass at near 3000 m depth in Monterey Canyon off the coast of California. The phylogenetic relationships of V. balaenophila are assessed in a parsimony analysis of morphological data together with nucleotide data from 28S rDNA, 16S rDNA and cytochrome oxidase I genes. Within the hesionids V. balaenophila belongs to Psamathini, where it is the sister group to Sirsoe. Among psamathins it is morphologically distinguished by having six glandular lip pads around the mouth opening, papilla-shaped neuropodial lobes on segment 3, extreme length of the dorsal cirri, and by a characteristic growth pattern in which the maximum number of segments is already formed in subadults, and further growth takes place through size increase of the segments. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 625–634.

A combined molecular (18S rDNA, 28S rDNA, 16S rDNA and COI) and morphological analysis of the benthic phyllodocids is presented for the first time. Nineteen phyllodocids and two outgroup taxa are assessed using parsimony, maximum likelihood and Bayesian analyses. We demonstrate high degrees in homoplasy in the traditionally used morphological phyllodocid characters, and show that all the three current subfamilies Phyllodocinae, Eteoninae and Notophyllinae are non-monophyletic. The genera Eulalia, Eumida, Protomystides, Pseudomystides, Pterocirrus and Sige form a well-supported group, as does Mystides and Nereiphylla. Another clade with strong support includes Eteone and Paranaitis, although with Eteone nested within a paraphyletic Paranaitis. The relationship between these two taxa indicate that the unusual arrangement of modified cirri on the first segments in Eteone is due to a fusion of segment 1 and 2 where the cirri of segment 1 have been reduced. Eulalia is non-monophyletic and should be split, minimally into two groups. Our results are ambiguous regarding the ancestral phyllodocid condition of absence-presence of median antenna or nuchal papilla and uniramous or biramous parapodia, but shows that the absence of cirri on segment 3 (previously an apomorphy, for e.g., Mystides, Pseudomystides and Hesionura) is maximally homoplastic.

We present parsimony analyses of annelids based on the largest taxon sample and most extensive molecular data set yet assembled, with two nuclear ribosomal genes (18S rDNA and the D1 region of 28S rDNA), one nuclear protein coding-gene (Histone H3) and one mitochondrial ribosomal gene (16S rDNA) from 217 terminal taxa. Of these, 267 sequences are newly sequenced, and the remaining were obtained from GenBank. The included taxa are based on the criteria that the taxon must have 18S rDNA or at least two other loci. Our analyses show that 68% of annelid family ranked taxa represented by more than one taxon in our study are supported by a jackknife value > 50%. In spite of the size of our data set, the phylogenetic signal in the deepest part of the tree remains weak and the majority of the currently recognized major polychaete clades (except Amphinomida and Aphroditiformia) could not be recovered. Terbelliformia is monophyletic (with the exclusion of Pectinariidae, for which only 18S data were available), whereas members of taxa such as Phyllodocida, Cirratuliformia, Sabellida and Scolecida are scattered over the trees. Clitellata is monophyletic, although Dinophilidae should possibly be included, and Clitellata has a sister group within the polychaetes. One major problem is the current lack of knowledge on the closest relatives to annelids and the position of the annelid root. We suggest that the poor resolution in the basal parts of the trees presented here may be due to lack of signal connected to incomplete data sets both in terms of terminal and gene sampling, rapid radiation events and/or uneven evolutionary rates and long-branch attraction. © The Willi Hennig Society 2006.

We assess phylogenetic relationships within the polychaete family Hesionidae from morphological data combined with nucleotide data from 18S rDNA, 28S rDNA, 16S rDNA and COI. Parsimony and Bayesian analyses were performed on two data sets; the first was based on a more restricted set of terminals with both morphological and molecular data (17 ingroup terminals), while the second included additional taxa with morphological data only (25 ingroup terminals). The different sets of terminals yielded fully congruent results, as did the parsimony and the Bayesian analyses. Our results indicate high levels of homoplasy in traditionally used morphological characters in the group, and that Hesioninae, Gyptini and Gyptis are nonmonophyletic. Hesionini (mainly Hesione and Leocrates), Psamathini (mainly Hesiospina, Micropodarke, Nereimyra, Psamathe and Syllidia), Ophiodrominae (Gyptini and Ophiodromini) and Ophiodromini (mainly Heteropodarke, Ophiodromus and Podarkeopsis) are monophyletic and agree with previous classifications, and Hesionini is probably the sister to all other hesionids. The placements of the small hesionids capricornia and Lizardia, the hydrothermal vent taxa Hesiodeira and Hesiolyra, and the newly described Hesiobranchia, remain uncertain.

Syllidia Quatrefages, 1866 (Psamathini, Hesionidae, Polychaeta) is revised based on examination of all extant types, other museum specimens, and a large number of newly collected specimens. Syllidia hongkongensis, new species, is described from Hong Kong and Hawaii, and Syllidia armata Quatrefages, 1866 from western France is redescribed. Syllidia hongkongensis is unique within the genus in having neuropodial lobes and neurochaetae from segment 3, rather than from segment 4. The generic names Magalia Marion and Bobretzky in Marion 1874 and Pseudosyllidia Czerniavsky, 1882 are treated as junior synonyms of Syllidia, and the specific names Magalia assimilis Pryde, 1914, M. capensis McIntosh, 1924, and Psamathe britannica Chamberlin, 1919 as junior synonyms of S. armata; the latter two represent new synonymies. A lectotype is designated for P. britannica. Syllidia inermis (Ehlers, 1912) from the Antarctic is treated as incertae sedis; it belongs within Psamathini but is not a member of Syllidia. Syllidia lin...

We present a COI-based parsimony analysis of the relationships between the shallow water, pigmented hesionid polychaete Nereimyra punctata, and a deep-water, unpigmented form with sympatric distribution in Norway and Sweden. Apart from the pigmentation differences, the two forms exhibit no observed morphological differences. The terminals are represented by four specimens each of the two forms from the Trondheimsfjord in Norway, and four each of the two forms from northern Bohuslan in Sweden, plus members of the two hesionids Heteropodarke and Ophiodromus as outgroups. In addition, the analysis includes a topotype of the morphologically similar and unpigmented Nereimyra woodsholea from the Middle Atlantic Bight off the US east coast. The equally weighted matrix includes 132 informative characters. All most-parsimonious trees unequivocally indicate that specimens belonging to the same form (pigmented or unpigmented) from different areas are cladistically closer related to each other than different forms from the same areas. Nereimyra woodsholea is nested within the unpigmented deeper group of the Norwegian and Swedish specimens, thus indicating that this name should be applied to the deep-water form in Norway and Sweden.

Micropodarke Okuda, 1938 (Psamathini, Hesionidae, Polychaeta) is revised based on examination of all available types, other museum specimens, and a large number of newly collected specimens from Japan, Hong Kong, Papua New Guinea, Australia, New Caledonia, California, the west coast of Canada, and the Red Sea. The previous synonymy of Micropodarke amemiyai Okuda, 1938 with M. dubia (Hessle, 1925) is substantiated, and M. trilobata Hartmann‐Schroder, 1983 is newly synonymized with M. dubia, leaving Micropodarke monotypic. A lectotype is selected for M. dubia. Micropodarke dubia is identified by the apomorphies of a proboscis diaphragm, segmental ventral adhesive papillae, distally nobbed and slightly curved neuroaciculae, and median neurochaetae with few, basally situated prolonged teeth (“spurs”), with abrupt transition to the following, much shorter teeth. Micropodarke dubia has a main distribution in the tropical and warm temperate Indo‐Pacific region, and appears to be absent from the Atlantic Ocean.

Two new species of Aberranta Hartman, 1965 are described from southeast France and Hong Kong, China. The two previously known species, A. enigmatica Hartman, 1965 and A. palpata Wolf, 1987 are redescribed from type and other material. Aberranta banyulensis n. sp. resembles A. palpata, and its internal and external morphology are examined further using histological, transmission and scanning electron microscopic techniques. Aberranta sulcata n. sp. is morphologically closest to A. enigmatica. The relationships between Aberranta and other polychaetes are discussed in light of a recent morphological and molecular analysis involving Nerillidae and Aciculata. The relationships among the Aberranta species are examined in a morphology-based cladistic analysis, using an amphinomid and three nerillids as outgroups.

An introduction is provided to the literature and to issues relating to phylogenetic nomenclature and the PhyloCode, together with a critique of the current Linnaean system of nomenclature. The Linnaean nomenclature fixes taxon names with types, and associates the names with ranks (genus, family, etc.). In phylogenetic nomenclature, names are instead defined with reference to cladistic relationships, and the names are not associated with ranks. We argue that taxon names under the Linnaean system are unclear in meaning and provide unstable group–name associations, notwithstanding whether or not there are agreements on relationships. Furthermore, the Linnaean rank assignments lack justification and invite unwarranted comparisons across taxa. On the contrary, the intention of taxon names in phylogenetic nomenclature is clear and stable, and the application of the names will be unambiguous under any given cladistic hypothesis. The extension of the names reflects current knowledge of relationships, and will shift as new hypotheses are forwarded. The extension of phylogenetic names is, therefore, clear but is associated to (and thus dependent upon) cladistic hypotheses. Stability in content can be maximized with carefully formulated name definitions. A phylogenetic nomenclature will shift the focus from discussions of taxon names towards the understanding of relationships. Also, we contend that species should not be recognized as taxonomic units. The term ‘species’ is ambiguous, it mixes several distinct classes of entities, and there is a large gap between most of the actual concepts and the evidence available to identify the entities. Instead, we argue that only clades should be recognized. Among these, it is useful to tag the smallest named clades, which all represent non-overlapping groups. Such taxa – LITUs (Least Inclusive Taxonomic Units) – are distinguished from more inclusive clades by being spelled with lower-case initial letter. In contrast to species, LITUs are conceptually straightforward and are, like other clades, identified by apomorphies. Zusammenfassung Die Arbeit ist eine Einfuhrung in die Kernpunkte der phylogenetischen Nomenklatur und des PhyloCodes mit einer Zusammenstellung der dazu vorliegenden Literatur sowie einer Kritik am gultigen System der Linneschen Nomenklatur. Die Linnesche Nomenklatur macht Taxonnamen an Typen fest und verbindet diese Namen mit einem Rank (Gattung, Familie etc.). In der Phylogenetischen Nomenklatur werden die Namen dagegen auf der Basis von Cladogrammen festgelegt und erhalten keinen Rank. Wir vertreten den Standpunkt, dass Bedeutung und Umfang der Linneschen Namen nicht wirklich klar sind und zu vagen Verbindungen zwischen Gruppen Veranlassung geben, ungeachtet, ob verwandtschaftliche Beziehungen zwischen ihnen akzeptiert werden oder nicht. Daruberhinaus fehlt den Linneschen Rankzuordnungen die Begrundung, was zu ungerechtfertigten Vergleichen uber die Taxa hinweg verleitet. Im Gegensatz dazu ist die Intention eines Taxonnamens in der Phylogenetischen Systematik klar und gleichbleibend; sein Gebrauch ist eindeutig, welche cladistische Hypothese auch gerade angenommen wird. Der Umfang eines derartigen Namens spiegelt das aktuelle Wissen uber die Verwandtschaftsbeziehungen wider; er wird sich verandern, wenn neue Hypothesen dazu vorliegen. Der Inhalt phylogenetisch definierter Namen ist daher klar, allerdings auch abhangig von der jeweiligen unterliegenden cladistischen Hypothese. Inhaltliche Stabilitat kann am besten durch sorgfaltig formulierte Namen erreicht werden. Eine phylogenetische Nomenklatur lenkt so den Blick von Diskussionen uber Taxonnamen hin auf das Verstehen verwandtschaftlicher Beziehungen. Wir setzen uns auch stark dafur ein, dass Arten nicht als taxonomische Einheiten angesehen werden. Der Begriff ,,Art“ ist unscharf und vieldeutig; er vermischt verschiedene Wesensklassen. So existiert zwischen den meisten aktuellen Artkonzepten und dem Erkennen dieser Einheiten eine grose Kluft. Dagegen pladieren wir dafur, dass nur Kladen anerkannt werden sollten, wobei es sinnvoll erscheint, die jeweils kleinsten bekannten Kladen, die fur sich nicht uberschneidende Gruppen stehen, zu bezeichnen. Derartige Taxa – LITUs (Last Inclusive Taxonomic Units) – unterscheiden sich von ubergeordneten Kladen durch kleine Anfangsbuchstaben. Im Gegensatz zu den Spezies sind LITUs begrifflich eindeutig und werden wie andere Kladen durch Autapomorphien charakterisiert.

Paranaitis Southern, 1914 (Phyllodocidae, Polychaeta) is revised based on an examination of all available types and newly collected specimens. Redescriptions are provided of the 11 previously described species considered valid: P. wahlbergi (Malmgren, 1865), P. abyssalis (Hartmann-Schroder, 1975), P. benthicola (Knox, 1960), P. bowersi (Benham, 1927), R caeca (Moore, 1903), R gardineri Perkins, 1984, P. inflata (Hutchings & Murray, 1984), R kosteriensis (Malmgren, 1867), P. polynoides (Moore, 1909), P. speciosa (Webster, 1879) and P. uschakovi Eibye-Jacobsen, 1991. Paranaitis misakiensis sp. nov., P. moritai sp. nov. and P. pumila sp. nov. are described from Japan. Anaitis peremptoria Claparede, 1870; Anaitis zeylanica Willey, 1905; Phyllooloce (Anaitis) papillosa Ehlers, 1887; and Phyllodoce (Anaitis) rubens Grube, 1880 are referred to as Phyllodocidae incertae sedis, and P. capensis (Day, 1960), P. formosa (Verrill, 1885) and R picta (Verrill, 1885) to as Paranaitis incertae sedis. Phyllodoce truncata (Hartmann-Schroder 1965) comb. nov. is removed from Paranaitis. Some previously unreported charaefers are introduced, including a series of proboscis characters, morphology of dorsal cirrophores, and symmetry of rostrum of chaetal shaft. Distinguishing characters for all recognized species of Paranaitis are provided in a table. In order to assess the position and delineation of Paranaitis and the relationships within this taxon, we present a morphology-based parsimony analysis of relationships within the Phyllodocidae. Paranaitis is shown to be paraphyletic at the exclusion of Chaetoparia, although current support does not allow for any formal synonymy. Phyllodoce and the Eteone-group appear as consecutive sister taxa to the Paranaitis-Chaetoparia clade. The monophyly of Notophyllinae is well supported, but low consensus resolution is obtained for the positions of major taxa such as Eulalia, Eumida, and the Mystides-group. (C) 2003 The Linnean Society of London.

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.

Hesionidae Grube, 1850 currently comprises over 175 species in 28 genera, placed in several subfamilies. Discoveries in recent years have largely been of deep-sea taxa. Here we describe a further four new hesionid species, mainly from methane ‘cold’ seeps at around 1000–1800 m depths off the Pacific coast of Costa Rica and new record of another species. Several of these taxa also occur at methane seeps in the Guaymas Basis (Mexico) and off the USA west coast (California and Oregon). The phylogenetic relationships within Hesionidae are reassessed via maximum parsimony and maximum likelihood analyses of DNA sequences from nuclear (18S rRNA and 28SrRNA) and mitochondrial (16SrRNA and Cytochrome c oxidase I) loci for the new samples. On the basis of these results, we refer one of the new species to Gyptis Marion & Bobretzky in Marion, 1874, one to Neogyptis Pleijel, Rouse, Sundkvist & Nygren, 2012, and two to Sirsoe Pleijel, 1998. The new species Gyptis robertscrippsi n. sp., Neogyptis jeffruoccoi n. sp., Sirsoe dalailamai n. sp. and Sirsoe munki n. sp. We refer to a collection of individuals from seeps ranging from Oregon to Costa Rica as Amphiduropsis cf. axialensis (Blake & Hilbig, 1990), even though this species was described from hydrothermal vents off Oregon. Neogyptis jeffruoccoi n. sp. was generally found living inside the solemyid clam Acharax johnsoni (Dall, 1891). The position of Hesiolyra bergi Blake, 1985 is resolved on the basis of newly-collected specimens from near the type locality and, as a result, Hesiolyrinae Pleijel, 1998 is synonymized with Gyptini Pleijel, 1998 (and Gyptinae Pleijel, 1998). http://zoobank.org/urn:lsid:zoobank.org:pub:9C0E88EE-34F8-4F25-9EC8-91797618AC86

Notophyllum Orsted, 1843 (Phyllodocidae, Polychaeta) is revised based on all available types and a large number of newly collected specimens. Redescriptions are provided of the seven species considered valid: N. americanum Verrill, 1885, N. foliosum (Sars, 1835), N. imbricatum Moore, 1906, N. japonicum Marenzeller, 1879, N. multicirris (Grube, 1878), N. sagamianum Izuka, 1912 and N. splendens (Schmarda, 1861). Notophyllum imajimai sp. n. is described from Japan. Some previously unreported characters are introduced, including a series of proboscis characters, morphology of dorsal cirri and cirrophores, and first appearance of notoaciculae in non-cephalized segments. A parsimony analysis of these taxa is presented based on 20 morphological characters and indicates two well-supported clades: Notophyllum as traditionally delineated and (N. foliosum, N. americanum (N. imajimai sp. n., N. imbricatum)). Characters for distinguishing all species of Notophyllum are provided in a table. Previous identifications of ...

A revision is presented of the taxonomy of Amphiduros Hartman, 1959 based on re-examined types, newly collected topotypes, and other specimens. Amphiduros axialensis Blake & Hilbig, 1990 is referred to Amphiduropsis, new genus. Amphiduros fuscescens (Marenzeller, 1875), A. izukai Hessle, 1925, A. pacificus Hartman, 1961 and A. setosus Hessle, 1925 cannot be conclusively diagnosed, wherefore the three latter are synonymised with A. fuscescens. Amphiduros fuscescens is newly recorded from Papua New Guinea and the Great Barrier Reef; the taxon appears to have a circumtropical and warm-water temperate distribution. Following a parsimony analysis based on morphological characters, Amphiduros and Amphiduropsis constitute a monophyletic group, either with Gyptis Marion & Bobretzky in Marion, 1874, or part of Gyptis, as sister taxon. In parallel to the traditional Linnean nomenclature, I present an alternative phylogenetic nomenclature of the group, with apomorphy-based definitions of taxon names and wit...

Nygren, A. & Fredrik, P. (2010). Chimaeras and the origins of the holopelagic annelids Typhloscolecidae and Lopadorhynchidae: a reply to Struck & Halanych. —Zoologica Scripta, 40, 112–114.

The phylogeny of Nereidiformia is assessed in a parsimony analysis of combined morphological and DNA data, with special focus on previously questioned relationships between Chrysopetalidae and Hesionidae, between Pilargidae and Hesionidae, and the affinities of Hesionides and Microphthalmus. A 660 bp segment of the mtDNA cytochrome c oxidase subunit I gene was sequenced for two chrysopetalids, one nereidid, one pilargid, one pisionid, two hesionids, plus the two questionable hesionids Hesionides arenaria and Microphthalmus sp. Phylogenetic resolution was poor for the cytochrome c oxidase subunit I gene data alone, but the combined analysis yielded partially robust topologies, suggesting that nereids are the sister group to chrysopetalids, and that pilargids, Hesionides and Microphthalmus do not belong within the hesionids

Phylogenetic taxonomy is applied for the systematization of a new Hesionidae, rather than the traditional Linnean system, with an apomorphy-based definition of the name and without reference to rank. It is argued that biological diversity is better represented without species concepts, but that it is useful to specify when a name refers to a smallest known clade which currently cannot be further subdivided; for this we apply the newly introduced concept LITU (Least-Inclusive Taxonomic Unit) for the new taxon. LITUs are made identifiable by being italicised with lower-case initial letter; all other taxon names are italicised with capital initial letter. The Hesionidae is accordingly named capricornia, new taxon, and was found in shallow water at One Tree Island, Capricorn Group, southernmost part of the Great Barrier Reef. capricornia is small (length

Cladistic relationships between 7 parts of the hesionid polychaete group Heteropodarke are assessed in a parsimony analysis based on 34 morphological characters. Taxon names are de- fined by apomorphy-based phylogenetic definitions, without reference to Linnean ranks or types. Species entities are omitted and denied any role in taxonomy; taxon names refer to monophyletic groups only. Linnean binomial species names are not employed, and all taxa are assigned uninomi- als. Previously known parts of Heteropodarke (Africana Hartmann-Schroder, 1974; Formalis, Perkins, 1984; Heteromorpha Hartmann-Schroder, 1962; Lyonsi Perkins, 1984; Xiamenensis Ding, Wu, and Wes- theide, 1997) are reexamined, and Bidentata, new taxon, and "Zmyrina" (informal name) are de- scribed from Papua New Guinea and Belize, respectively. The new taxon Crassichaetae is named for a subgroup of Heteropodarke, which is diagnosed by enlarged, anteriorly situated falcigers, and in- cludes Africana, Heteromorpha, Lyonsi, and Xiamenensis . Within this group Africana and Heteromorpha are treated as taxa inquirendae. {Apomorphy-based definitions; Linnean nomenclature; phylogenetic taxonomy; polychaetes; ranks; species concepts; species names; types.}

Previous studies suggest that the polychaete taxa Hesionidae and Chrysopetalidae may not represent separate groups, that Pilargidae constitute a subgroup within Hesionidae, and that Hesionides and Microphthalmus are highly derived hesionids. Phylogenetic systematic analyses of Phyllodocida and the subgroup Nereidiformia are presented in order to clarify the position and delineation of these taxa. The phyllodocida analysis includes 18 families representing the majority of the taxa in the group, is rooted with Onuphidae, and is based on 42 absent/present coded morphological characters, obtained mainly from literature. All 69 resulting shortest trees include the clade (Chrysopetalidae, Nereididae, Hesionidae), but with either Syllidae, Nautiliniellidae, Pilargidae or (Aphroditiformia, Pisionidae) as sister. In- and outgroup taxon selection for the Nereidiformia study is dictated by the outcome of Phyllodocida analysis, with scores based on examined species of two chrysopetalids, four hesionids, one nereid, one pilargid, one pisionid, one syllid, plus the putative hesionids Hesionides arenaria and Microphthalmus sp. It is based on 46 absent/present coded morphological characters. Two equally parsimonious trees indicate that chrysopetalids and hesionids are well delineated, that pilargids and hesionids are non-overlapping, and that Microphthalmus and Hesionides are not hesionids.

Hesionid interrelationships are assessed in a parsimony analysis of 97 (86 informative) absent/present coded morphological characters of 37 terminal taxa within the group. Character information is based on examination of available types, newly collected topotypes and additional non-types, with introduction of many not previously recorded characters. Polarity is assessed differently in two separate analyses, one by outgroup comparison with nereids and chrysopetalids, yielding two equally parsimonious trees, and one by observations from juvenile hesionids, yielding 10 equally parsimonious trees. The outgroup rooted trees are selected for a reclassification, and three main groups are recognised: Hesiolyrinae, new subfamily, Hesioninae Grube, 1850, and Ophiodrominae, new subfamily. Hesiolyrinae is monotypic for Hesiolyra, Hesioninae includes Hesionini Grube, 1850 and Psamathini, new tribe, and Ophiodrominae includes Gyptini and Ophiodromini, both new tribes. Sirsoe, new genus and member of Psamathini, is established for Orseis grasslei Blake, 1985 and the yet undescribed Sirsoe A. High degrees of homoplasy are demonstrated for a number of characters traditionally used in hesionid taxonomy, e.g., distribution of anterior, enlarged cirri, and first segment provided with neurochaetae and neuropodial lobes. Diagnoses and descriptions are provided for all supraspecific hesionid taxa and a checklist of hesionids is included. For improved precision in polychaete descriptions it is suggested that the term ‘tentacular cirri’ should be replaced by ‘anteriorly situated cirri’ with specified position and morphology. The following new generic synonyms are introduced: Lamproderma Grube, 1877, objective junior synonym of Leocrates Kinberg, 1866; Neopodarke Hartman, 1965 junior synonym of Nereimyra Blainville, 1828; Orseis Ehlers, 1864, junior synonym of Ophiodromus Sars, 1862; Paragyptis Pocklington, 1984 junior synonym of Gyptis Marion & Bobretzky in Marion, 1874; and Podarke Ehlers, 1864, objective junior synonym of Ophiodromus. Additionally a number of new species group synonyms and new combinations are proposed. Parallel to the Linnaean, type-based definitions of taxon names, a hesionid classification is introduced with phylogenetic taxon name definitions.

The phylogenetic relationships of spionid genera are estimated from parsimony analyses of morphological characters, with Trochochaetidae, Poecilochaetidae and Uncispionidae as outgroups. A first analysis of currently recognised genera proved inconclusive and even exclusion of six of the most polymorphic genera resulted in 13 305 equally parsimonious trees and a fully collapsed consensus tree. A second analysis using only the type species of each genus, yielded four equally parsimonious trees; reduced to two after successive weighting. The topologies of these two trees indicated division of the family into four main groups: (1) Aonidella and Xandaros ; (2) Prionospio ( sensu lato )-complex, Laonice , Spiophanes and Aonides ; (3) a large assemblage of genera, including Polydora -( sensu lato ), Scolelepis , Malacoceros and Spio ; (4) Atherospio , Pseudatherospio and Pygospiopsis . Earlier literature classifications of the group are evaluated and compared with the new results.

Taxon names should be founded on phylogenetic relationships. and the names defined on the basis of common ancestry. Definitions based on evolutionary relationships relate the names to a phylogeny, and while the inclusiveness of the name may change with changing hypotheses of monophyly, the actual name remains unaltered. The limits of the name arc fixed by pointing to a monophyletic clade, where group membership is determined by the relationship to this clade. and not to subjective decisions of taxon delineations. Since phylogenetic definitions unambiguously connect the name to a specified clade, and not to a type, the conventional type concept becomes superfluous. We furthermore consider the Linnean categories poorly suited to convey the information in evolutionary trees, and suggest that these categories are abandoned.

The phylogeny of scale-worms, benthic polychaetes carrying dorsal scales (elytra), including taxa from Acoetidae, Aphroditidae, Eulepethidae, Pholoidae, Pholoididae, Polynoidae and Sigalionidae (Aphroditiformia), is assessed from the nuclear markers 18SrRNA and 28SrRNA, and mitochondrial 16SrRNA and cytochrome c oxidase subunit I (COI), and 24 morphological characters. The data sets are analyzed both separately and combined, with Bayesian analyses, maximum likelihood and parsimony. In total, 56 terminal taxa are examined, including 48 taxa from all scale-worm families, and eight out-group species. The results indicate that Aphroditidae and Eulepethidae are the most basally placed families among the scale-worms. The Pholoididae and Pisionidae are positioned within and synonymized with the Sigalionidae, and Pholoidae may be part of the same group. The subfamily Iphioninae falls out as sister group to a clade consisting of Polynoidae and Acoetidae and is elevated to Iphionidae. The families now included in the Aphroditiformia are Acoetidae, Aphroditidae, Eulepethidae, Pholoidae, Polynoidae, Iphionidae and Sigalionidae, and the subfamily name Harmothoinae and Acholoinae are treated as a junior synonyms of Polynoinae.

The phylogeny of 13 species of the polychaete genus Phyllodoce Lamarck is estimated from 23 morphological characters, totalling 54 character states. The genus is large, with over 100 nominal species, but, since both available material and descriptions for many species are inadequate, a smaller number of species was included in the analysis. Following recent, higher-level phylogenetic studies, Chaetoparia nilssoni Malmgren and a group of Paranaitis Southern species were chosen as outgroups. Six equally parsimonious trees with a consistency index (CI) of 0.78 were obtained. The original character matrix is compared to randomized character states in a permutation tail probability test, the result of which indicates the presence of non-random, hierarchical pattern in the original data. Phyllodoce is reclassified and divided into three subgenera: Phyllodoce, Zverlinum Averincev, and Anaitides Czerniavsky. Synapomorphies are also used for subgeneric allocation of those species not included in the cladistic analysis. Several new synonymies are introduced: Austrophyllum monroi Hartman, junior synonym to Phyllodoce bulbosa Wesenberg-Lund; Anaitides compsa , Chamberlin and Anaitides carloensis Kudenov, junior synonyms to Phyllodoce panamensis Treadwell.

We describe Proceraea albocephala, new species, from Madeira, and Erseia oligochaeta, new genus and new species, from Istria, Croatia, and we provide redescriptions of Proceraea madeirensis (Nygren, 2004) from topotype material, and Myrianida longoprimicirrata (Lopez, San Martin & Jimenez, 1997) from material collected at Istria, Croatia, and Banyuls-sur-Mer, France. Proceraea albocephala, new species is morphologically separated from similar species by a prostomial white spot, and E. oligochaeta, new genus, new species is unique in having only a few (1–2) simple unidentate chaetae in all chaetigers, and a trepan with a single large and 25–28 smaller teeth. We assess the phylogenetic positions of the four species using nuclear 18SrDNA, together with mitochondrial COI and 16SrDNA. Our molecular data show that among the sequenced autolytines 1) P. albocephala, new species is most closely related to P. nigropunctata Nygren & Gidholm, 2001, P. okadai (Imajima, 1966), and P. cornuta (Agassiz, 1862), 2) E. oligochaeta, new genus, new species belongs within a clade together with Procerastea nematodes Langerhans, 1884, Virchowia clavata Langerhans, 1879, and Imajimaea draculai (San Martin & Lopez, 2002), 3) M. longoprimicirrata is sister species to M. pentadentata (Imajima, 1966), and 4) P. madeirensis has a basal position within Procerini. The molecular data suggests that Proceraea Ehlers, 1864 as currently delineated is paraphyletic.

The phylogenetic relationships of the benthic phyllodocid genera are estimated by a parsimony analysis based on 26 morphological characters. It is concluded that the cephalization within Eteone has led to a reduction of segment 1 and loss of the first pair of tentacular cirri. Biramous parapodia (like in Austrophyllum and Notophyllum) constitute an ancestral state, which has evolved independently also within Phyllodoce. A new subfamily, Notophyllinae, is erected, and the two additional subfamilies, Phyllodocinae and Eteoninae, are redefined. Pareteoninae is treated as a junior synonym to Phyllodocinae, and Lugiinae as a nomen dubium. A number of genera are redefined and new combinations introduced. Based on type material Zverlinum and Phyllouschakovius are synonymized with Phyllodoce, Nipponophyllum with Notophyllym, Vitiaziphyllum with Eumida, Pareteone, Eulalia (Euphylla) and Compsanaitis with Paranaitis. More tentatively proposed synonymizations include Bergstroemia with Austrophyllum, Prophyllodoce with Phyllodoce, and Pseudeulalia with Protomystides. Eulalia lapsus is proposed as a replacement name for E. anoculata Fauchald, 1972, junior homonym to E. anoculata Hartman & Fauchald, 1971, and a number of species are synonymized and new combinations introduced. A checklist of phyllodocids together with information on the location of their type specimens is included.

We review the taxonomic history of pogonophores (frenulates and vestimentiferans), from the species in first described 1914 to the recently described bone-eating worm Osedax. Previous systematists have referred both groups to the rank of phylum, and the animals have been treated as deuterostomes with a dorsal nerve cord. Further knowledge on their embryology, the discovery of the previously overlooked posterior, segmented part provided with chaetae, and access to molecular data, have completely changed earlier views on their affinities. They are now referred to as a single family of polychaete annelids, Siboglinidae.

The genus Sige Malmgren, 1865 is revised and the type species S.fusigera Malmgren, 1865, redescribed from syntypes and newly collected topotype material. Based on examination of type specimens, six species described from other genera are transferred to Sige: Pirakia brunnea Fauchald, 1972; Vitiazia dogieli Ushakov, 1953; Eulalia longocirrata Stop-Bowitz, 1948; Eumida (Eumidaj parvicirrus Perkins, 1984; Eulalia sandwichensis Ushakov, 1975; Eulalia sigeformis Annekova, 1937. The genus Vitiazia Ushakov, 1953, being monotypic, becomes a junior synonym to Sige. Pirakia lanceolata Hartman & Fauchald, 1971 is synonymized with Sige longocirrata (Stop-Bowitz, 1948) and Sige oliveri sp. nov. is described from western Norway and Sweden. Ten species are recognized as belonging to Sige. The relationships of Sige to other genera are discussed.

We describe Myrianida gidholmi sp. n. from shallow waters in Belize. It is characterized by a unique colour pattern consisting of red transverse and longitudinal bands. We determine its phylogenetic position within Myrianida using a combined approach with morphological and molecular data. The new species is compared to relevant Myrianida taxa and important features for morphological identification are listed in a table. The new combinations Myrianida tyrrhenica (Cognetti, 1953) and M. cognetti (Cinar & Gambi, 2005) are introduced, and Autolytus antondohrni Cinar & Gambi, 2005, is synonymized with M. tyrrhenica. We also provide a checklist of all taxa belonging to Myrianida.

We provide a description of newly collected specimens of Axiokebuita Pocklington & Fournier from Norway, previously known only from east Canada and the Antarctic. Due to delineation problems between the only two described species, A. minuta (Hartman) and A. millsi Pocklington & Fournier, these new specimens cannot unambiguously be referred to either species. Previously unnoticed adhesive papillae on the pygidium are present in both species and may constitute an apomorphy for Axiokebuita. The taxon lacks many morphological features otherwise characteristic for scalibregmatids, and to assess its affinities we present 18S rDNA and 28S rDNA-based analyses together with six other scalibregmatids and twenty other polychaetes. A nemertean is used as outgroup. All analyses support that Axiokebuita is a scalibregmatid. Furthermore, Travisia Johnston, traditionally referred to the Opheliidae, is nested within the scalibregmatids, as sister to Neolipobranchius Hartman & Fauchald. Arenicolidae and Maldanidae may constitute the sister group of scalibregmatids.

The phylogeny of Aphroditiformia, benthic polychaetes carrying dorsal elytra, is assessed from nuclear 18S rDNA, mitochondrial cytochrome c oxidase subunit I (COI), and 31 morphological characters. Two non-elytrabearing taxa, Palmyra and Pisione, are included to assess their relationship to the elytrabearers. The data are analysed both separately and combined, with parsimony, maximum likelihood and Bayesian analyses. In total, 19 terminal taxa are examined, including 12 elytrabearing taxa from all scale-worm groups, Palmyra, Pisione, and five outgroup taxa. The results show that Palmyra and Pisione are nested within Aphroditiformia. Palmyra is sister to Aphrodita, and both Pisione and Pholoe are positioned within Sigalionidae, suggesting that both family names Pisionidae and Pholoidae should be treated as junior synonyms of Sigalionidae.

Some of the intriguing issues in current polychaete systematics are reviewed. (1) The root of the ‘polychaete’ tree. Currently there are two major hypotheses concerning the root position among polychaetes. One is based on rooting cladograms with outgroups such as Mollusca and result in simple-bodied taxa such as Opheliidae and Questidae forming a basal annelid grade along with Clitellata. Other hypotheses do not use outgroup rooting but involve scenarios on the evolution of the group and would place taxa in Aciculata as basal annelids, thus making Aciculata and Phyllodocida paraphyletic. Molecular sequence data has been of little help in resolving this issue thus far, largely due to limited taxon sampling. (2) Paraphyly. Owing, in part, to a tradition involving the emphasis on differences among taxa, and the application of Linnean ranks (e.g., family), paraphyly is undoubtedly a widespread phenomenon in polychaete systematics. An example of this has been proposed already for Spionidae. If the tree topology and rooting used by Blake & Arnofsky (1999) is correct, Spionidae is made paraphyletic by the recognition of the following four family-ranked taxa; Trochochaetidae, Poecilochaetidae, Longosomatidae and Uncispionidae. Another possible example is seen with Cirratulidae. A preliminary cladistic analysis shows that it is entirely possible that seven other taxa recognised as families may be nested within Cirratulidae. These include Acrocirridae, Ctenodrilidae, Fauveliopsidae, Flabelligeridae, Flotidae, Poeobiidae and Sternaspidae. (3) Problematic taxa. Apart from the problems exposed by the analysis of Cirratuliformia, the position of some of these groups, such as Aberranta, Alciopidae, Hesionides, Lopadorhynchidae, Microphthalmus, Nerillidae, Spinther, Tomopteridae and Sabellariidae, is discussed.

Nereididae Johnston, 1865, part of the large clade Phyllodocida, is one of the best-known annelid groups. Presently, there are some 500 nominal species grouped into 42 genera, although there is little consensus among different authors as to how they should be classified. The relationships of nereidids were assessed in a morphology-based parsimony analysis of 41 terminal taxa, with members of Chrysopetalidae and Hesionidae used as outgroups. Type species for the majority of currently recognised nereidid genera were used as terminal taxa, and character information was based on examination of type and non-type specimens, together with literature descriptions. High degrees of homoplasy were found for several features that are traditionally applied to delineate subgroups of Nereididae, including the presence of paragnaths and the distribution of different kinds of chaetae. Six major groups were recovered: Namanereidinae, including Namalycastis and Namanereis, characterised by spherical shape of palpostyles and ventrally displaced notoaciculae; one clade corresponding in part to previous authors concepts of Nereidinae, including Nereis, Eunereis, Hediste and Platynereis (the relationships of several well known nereidids, such as Neanthes and Perinereis, commonly referred to Nereidinae, could not be unambiguously resolved); one unnamed and not previously recognised clade (A), including Australonereis, Laeonereis, Dendronereides and Olganereis, characterised by the presence of papillae on the maxillary ring; a second unnamed clade (B), including Leptonereis, Sinonereis, Tylonereis and Tylorrhynchus, characterised by enlarged notopodial ligulae; a well supported Gymnonereidinae, restricted to Ceratocephale, Gymnonereis, Tambalagamia and Micronereides; and a third unnamed clade (C), including Ceratonereis, Solomononereis, Unanereis, Cheilonereis and Websterinereis, characterised by unilobated neuropodial postchaetal lobes. Among these groups we found good support for the Namanereidinae, the Gymnonereidinae and for the whole of Nereididae. The subfamilies Dendronereidinae and Notophycinae (based on Micronereis, senior synonym of Notophycus) are regarded here as monotypic.

Descriptions of species of Phyllodoce Savigny, 1818 from Northern Europe [P. laminosa Savigny, 1818, P. citrna Malmgren, 1865, P. groenlandicaOrsted, 1843, P. lineata (Claparede, 1870), P. longipes Kinberg, 1866, P. maculata (Linnaeus, 1767), P. mucosaOrsted, 1843 and P. rosea (Melntosh, 1877) comb.n.] are given. A key to the species is included. The examined material consists of newly collected material as well as collections in museums. Available type material has been checked. A number of junior synonyms are proposed. Anaitides Czerniavsky, 1882 and Aponaitides MeCammon & Montagne, 1979 are considered not valid.

The phylogenetic position of the most speciose meiofaunal polychaete family, Nerillidae, has remained contentious. Recent hypotheses have generally focused on the fact that Nerillidae shares with Aciculata (a major polychaete subgroup) features such as compound chaetae, ventral buccal organ and short ventrolateral palps. Here we present the first phylogenetic analysis of Aciculata, together with Nerillidae, combining morphological and molecular data. We also include Aberrantidae, previously referred to or placed near to spiomorph polychaetes, but recently referred to Aciculata, possibly close to Nerillidae. The data sets of 24 terminals contain 53 morphological characters and nearly complete sequences of 18S rRNA. The sequences were analysed simultaneously with the morphological data by direct optimization in the program POY with a variety of parameter settings (costs of gaps: transversions: transitions). The various settings resulted in markedly different phylogenetic hypotheses, but on the basis of congruence (ILD) the results of two parameter settings were chosen. In all analyses, the three included nerillid species constituted a monophyletic group. Only two analyses provided fully resolved cladograms. The morphological analysis gave poor resolution and the position of the nerillids was equivocal. The two molecular-based cladograms (minimizing ILD) were also poorly resolved, but one provided a position for nerillids next to Eunice pennata and Nothria conchilega, from the subgroup Eunicida within Aciculata. The two cladograms of the combined analyses (minimizing ILD) were fully resolved and placed nerillids in a terminal position next to Aberranta sp., within a clade of eunicidan species. The study showed that the analytical conditions for the homology assignment of 18S rRNA strongly influenced the phylogenetic results. The various previous proposals on the phylogenetic position of the Nerillidae are reviewed, some of which are in accordance with the results of the present study.