Revision of Pseudonereis (Polychaeta, Nereididae)

A taxonomic revision of Pseudonereis (Polychaeta, Nereididae) shows that some of the described taxa are very similar in most morphological characteristics. The revision includes all ten taxa considered valid, and are redescribed from type material. Lectotypes are designated for Pseudonereis anomala Gravier, 1901, Pseudonereis noodti (Hartmann-Schröder, 1962) and Pseudonereis trimaculata Horst, 1924. The widely geographically distributed and well-known P. gallapagensis Kinberg, 1865 and P. variegata ( Grube, 1857 ) show striking morphological resemblance to less well-known taxa with similar distribution. Paragnath variation in populations of P. anomala is discussed relating to its geographical distribution. Pseudonereis trimaculata is recorded from Australia for the first time. Taxa belonging to Pseudonereis are predominantly tropical and subtropical. A cladistic analysis using parsimony is included to test for monophyly of Pseudonereis . A monophyletic clade including all Pseudonereis taxa is given low bootstrap support. This clade is supported by the synapomorphies: presence of paragnaths in closely spaced comb-like rows on the maxillary ring on the pharynx, and presence of p-bar paragnaths in Areas II–IV and VII–VIII. Several of the included taxa share the shield-shaped paragnath in Area VI, which serves to distinguish Pseudonereis spp. from Perinereis spp. Paragnaths of the type p-bars and shield-shaped bar is described for the first time; the latter character is different from the smooth bar-shaped paragnaths in Area VI as has previously been described in these taxa.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 145–176.     

Selectivity of Perinereis aibuhitensis (Polychaeta,Nereididae) feeding on sediment

Deposit feeding polychaetes play an important role in the acceleration of the biogeochemical processes of the sediment through bioturbation. Feeding is one of the important factors of bioturbation. However, knowledge of the feeding biology of polychaetes, especially the subsurface deposit feeder, is limited. The objective of this study is to characterize the feeding selection of Perinereis aibuhitensis with different body weights. The animals were fed with natural sediment from their original habitat in the lab. The feed intake and particle size of sediment were measured to find any evidence of feeding selection. A two-way ANOVA showed that the particle size class significantly affected the particle size frequency of the ingested and the remaining sediment (P?P?>?0.05). Bivariate correlation analysis showed that the particle size frequency of sediment, ingested and remaining sediment in different size classes were linearly related (P?P?P?P. aibuhitensis preferred smaller particles in the sediment, which was limited by the particle size distribution of the sediment in which they live. The nitrogen and organic carbon contents in the faeces might be the thresholds when P. aibuhitensis selects sediment particles as food. These results demonstrated the particle selectivity of P. aibuhitensis, and may be applicable to other subsurface deposit feeding polychaetes.     

Phylogeny and reproduction in Syllidae (Polychaeta)

The phylogeny of 12 representatives of the four syllid subfamilies is analysed using 44 morphological characters and 96 character states. The analysis is designed to answer two questions: are the four subfamilies monophyletic, and how are the different reproductive strategies related? Syllids show two modes of reproduction, epigamy and schizogamy. Epigamy is mainly constrained to two of the four subfamilies, Eusyllinae and Exogoninae but occurs also in some members of Autolytinae. Schizogamy can be of two types–scissiparity and gemmiparity–and both types are found in Syllinae and Autolytinae. Several modes of brood protection also occur in the family; in some epigamous species the eggs and even juveniles are attached to the main animal, whereas in brooding schizogamous species the stolons care for the young. Brood protection appears in Exogoninae, a few species of Eusyllinae, and some Autolytinae. The resulting tree indicates that members exhibiting brood protection are constrained to one clade, and that it is uncertain whether epigamy or schizogamy is the plesiomorphic reproductive mode. Three of the four subfamilies, Autolytinae, Exogoninae and Syllinae, are supported in the analysis whereas Eusyllinae is found to be polyphyletic.     

Behaviour, Setal Inversion and Phylogeny of Sabellida (Polychaeta)

The path taken by the faecal groove, following two right angle bends, appears inefficient in Sabellidae. It fits well with the thoracic folds of serpulids and follows a gentle spiral in caobangiids, but only in spirorbids is it of obvious adaptive value, being suited to life in a coiled tube. The associated setal inversion is not necessary for locomotion, which does not depend wholly on setal leverage, except in the smaller Fabriciinae where long-shafted uncini also thrust actively against the elastic tube wall. The collar setae of spirorbids and some serpulids thrust anteriorly, giving some backward movement, but such motion is mainly by body contraction. Short uncini are virtually passive, engaged by opposing setae acting as distance pieces. Some broad bladed setae have a hollow structure around a central core. Pick-axe setae of Sabellinae may have developed to counteract the thrust of respiratory peristalsis. This is not seen in spirorbids and serpulids, the thoracic folds of which are a sufficient respiratory supplement to the tentacles. — Numbers of thoracic segments in Sabellidae range from 1 to 16, and the abdomen becomes vestigial in some Fabriciinae. Abdominal inversion is important to spirorbids but not to sabellids, which are more varied. It seems that the former are archaic and the latter had a coiled ancestry. Sabellariids also show abdominal inversion suggesting a distant relationship with Sabellida.     

Phylogeny and classification of the Phyllodocidae (Polychaeta)

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.     

Phylogeny of Syllidae (Polychaeta) based on morphological data

The phylogeny of Syllidae is assessed in two parsimony analyses of 107 morphological characters. The first analysis included one species of each of the 71 genera of the Syllidae, as well as members of other close families. In the second analysis, 23 poorly known genera were excluded. Character information is based on the examination of available types, additional non-types and newly collected material. Syllidae, except Bollandia Glasby, 1994 is monophyletic. Both analyses supported three of the four traditional subfamilies (Exogoninae, Syllinae and Autolytinae) as monophyletic, whereas Eusyllinae was clearly a polyphyletic group. The genera Anoplosyllis Claparède, 1868, Astreptosyllis Kudenov & Dorsey, 1982 , Streptosyllis Webster & Benedict, 1884, Streptospinigera Kudenov, 1983 and Syllides Örsted, 1845 comprise a well-supported monophyletic group, which we classified as a new subfamily: Anoplosyllinae n. subfam. Our results indicated high levels of homoplasy in the morphological characteristics that traditionally used to differentiate groups, such as the fusion of palps and the presence of nuchal epaulettes. Considering the reproductive modes, schizogamy has appeared twice in the family as the derived condition evolving from epigamy, and Exogoninae may be divided into two monophyletic groups based on the brood system.     

Platynereis nadiae sp. n. (Polychaeta: Nereididae) from Italian coasts

A new species, Platynereis nadiae sp. n. (Polychacta: Ncrcididac), from thc Mediterranean Sea is described. It is clearly distinguished from the other Platynereis species by the distribution and the morphology of the homogomph falciger setae and by the structure of thc tcntacular cirri.     

Phylogeny of Aphroditiformia (Polychaeta) based on molecular and morphological data

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.     

Reproductive effort,fecundity and energy allocation in two species of the genus Perinereis (Polychaeta: Nereididae)

Summary

The reproductive effort in terms of fecundity and energy allocation was studied in two species of semelparous polychaetes belonging to the genus Perinereis, living in the same environment, with different reproductive modalities. There is a great individual variability both in terms of reproductive effort and fecundity. Fecundity varied from 4080 to 15000 oocytes in P. rullieri and from 7000 to 26000 in P. cultrifera; no linear relationship was found between oocyte number and total jaw length utilised as size index. The energy content of germinal and somatic tissues was determined by Differential Scanning Calorimeter (DSC). The reproductive effort was calculated as RE = E_G/(E_G + E_S) where E_G is the total energy in germinal tissues and E_S is the total energy in somatic tissues. Reproductive effort is very high with mean values of 0.62 for P. rullieri and 0.79 for P. cultrifera. The different amounts of energy allocated in germinal tissues can be attributed to the different reproductive modalities—P. rullieri reproduces in the atokous phase whereas P. cultrifera has conserved epitoky in its life-cycle. The lack of correlation between reproductive effort and size index strongly suggests that reproductive allocation does not increase with age. In semelparous species the variability in fecundity and reproductive effort observed cannot be interpreted in terms of a trade-off between fecundity and survival as in iteroparous species. In fact, in semelparous an individual allocates all available resources to reproduction and then dies.     

Epitoky in Nereis (Neanthes) virens (Polychaeta: Nereididae): a story about sex and death

The nereidid Nereis (Neanthes) virens undergoes drastic behavioural, morphological and physiological changes during its sexual maturation (epitoky). This metamorphosis prepares benthic worms for a brief pelagic existence devoted to mating although in N. virens only mature males leave their burrows to swarm. After spawning, individuals of both sexes die. Specific adjustments of energy metabolism pathway allowing higher muscular activity and swimming capacity remain to be eluded. This study compared atokous worms (immature) and epitokous (mature) swimming males and benthic females of N. virens to detect metabolic changes that could occur during epitoky. Epitokous males showed significantly higher electron transport system, citrate synthase and aspartate aminotransferase activities (p<0.01) and significantly lower lactate dehydrogenase activity (p<0.01) compared to atokous worms and epitokous females. There was no difference in antioxidant enzyme capacities between epitokes and atokes. Lipase and trypsin activities were significantly lower (p<0.01) in epitokous males. The enzymatic changes observed are likely related to the metabolic adjustments required to support higher swimming abilities. Maintenance of antioxidant capacities could be related to protection of germinal tissues more than long term survival, since N. virens die after spawning.     

A review of Asian Hediste species complex (Nereididae, Polychaeta) with descriptions of two new species and a redescription of Hediste japonica (Izuka, 1908)

Three Asian species of the genus Hediste (Nereididae, Polychaeta, Annelida), including two new species, are reviewed. Hediste japonica (Izuka, 1908) was redescribed based on recently discovered type material and on additional specimens newly collected from the Ariake Sea in Japan and the west coast of Korea. Two new species, H .  diadroma and H. atoka , were described based on specimens collected from Japan. These three species are morphologically similar, but distinguishable by some differences in parapodial and chaetal morphology, at least at the sexually mature stage. The presence of homogomph falcigers and the absence of heterogomph spinigers are unique characteristics of H. japonica among this genus. Inconspicuous epitokous metamorphosis occurred as a slight enlargement of the eyes and parapodial ligules in mature adults of H. diadroma sp. nov. and H. japonica in combination with their reproductive swarming and pelagic larval development. The epitoke-specific sesquigomph spinigers were added in mature adults of H. diadroma sp. nov., while the number of ordinary homogomph spinigers increased in some adults of H. japonica. Epitokous metamorphosis in the Hediste species was compared with that in other nereidid genera. No epitokous metamorphosis was observed in H. atoka sp. nov., corresponding to its reproduction without swarming and to its development without a planktonic larval stage. Significant differences in paragnath numbers in the maxillary ring of the proboscis were found among the three species, suggesting different feeding habits. In Japan, the present distribution of H. japonica is restricted to muddy tidal flats in the inner part of the Ariake Sea, probably due to recent human impact, while the other two species are widespread.  © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society, 2003, 137, 403−445     

Phylogeny of benthic Phyllodocidae (Polychaeta) based on morphological and molecular data

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.     

Phylogeny of the mud worm genus Marenzelleria (Polychaeta, Spionidae) inferred from mitochondrial DNA sequences

The taxonomy of the cryptic morphospecies of the mud worm genus Marenzelleria is particularly difficult and the phylogenetic relationship within the genus is unknown. Herein we reconstructed the phylogeny of all five species of this genus using sequence data of three mitochondrial genes (16SrDNA, cytochrome b, cytochrome oxidase subunit I) from 104 specimens out of 26 populations. For the three invasive species of the genus, Marenzelleria neglecta, M. viridis and M. arctia , individuals from native populations as well as from recently invaded populations were included. Nuclear 18S rDNA sequences were used to evaluate the appropriate outgroup taxon among several spionid polychaete species. The results supported the monophyly of Marenzelleria , and Malacoceros fuliginosus was found to be a suitable outgroup for the analysis of the mitochondrial gene segments. All phylogenetic reconstructions revealed a basal position of M. arctia and M. wireni , which have primarily Arctic distribution, with M. arctia obtaining the most basal position. Together with the present-day distribution of the species, this indicates an origin of the genus in the Arctic region. The relationship of the species M. neglecta , M. viridis and M. bastropi could not be resolved sufficiently due to genealogical discordance that might reflect relatively young cladogenetic events.     

Physiological and antioxidant enzyme responses to acute and chronic exposure of Laeonereis acuta (Polychaeta, Nereididae) to copper

Chronic (14 days) and acute (48 h) copper effects on the antioxidant defense system and some physiological variables of Laeonereis acuta (Polychaeta, Nereididae) were evaluated. In both assays, two nominal copper concentrations (chronic: C₁=31.25 and C₂=62.50 μg/l; acute: A₁=250 and A₂=500 μg/l) and one control group (C_c and A_c=0 μg/l) were tested. End points analyzed were antioxidant enzyme activity (catalase, CAT; superoxide dismutase, SOD; and glutathione S-transferase, GST), oxygen consumption, metahemoglobin concentration, and lipid peroxidation (LPO). In the chronic assay, CAT activity was significantly higher in worms exposed to both concentrations of copper tested (C₁=3.36±0.07 U CAT/mg protein; C₂=4.06 0.32 U CAT/mg protein) than in control worms (C_c=2.16±0.39 U CAT/mg protein). SOD activity was also increased in the two copper-exposed groups (C₁=16.85±4.22 U SOD/mg protein; C₂=38.19±4.31 U SOD/mg protein) than in control group (C_c=3.54±0.46 U SOD/mg protein). However, GST activity was increased only in worms exposed to the higher copper concentration (C₂=0.022±9.10⁻⁴ U GST/mg protein) when compared to the other groups tested (C_c=0.012±3.10⁻³ U GST/mg protein; C₁=0.016±9.10⁻⁴ U GST/mg protein). None of the physiological variables analyzed (oxygen consumption, metahemoglobin concentration, and lipid peroxidation) was affected by chronic copper exposure. In the acute assay, only GST activity was induced in worms exposed to copper. This induction was observed only in the A₁ group (0.027±2.10⁻³ U GST/mg protein) when compared to A_c (0.017±2.10⁻³ U GST/mg protein) or A₂ (0.016±7.10⁻⁴ U GST/mg protein) groups. On the other hand, lipid peroxidation was higher in A₂ (481.9±49.2 nmol CHP/g ww) than in control worms (A_c=337.9±25.0 nmol CHP/g ww). Oxygen consumption was higher in worms acutely exposed to the lower copper concentration tested (A₁=0.27±0.04 mg O₂/g ww/h) than in the higher concentration (A₂=0.14±0.01 mg O₂/g ww/h). Changes in the swimming behavior of copper-exposed animals in both assays and edemas in the body wall of worms acutely exposed to copper were also observed. Results suggest that copper exposure favors reactive oxygen species generation and that enzymatic defense system is induced under chronic exposure, preventing oxygen consumption changes and lipid peroxidation and metahemoglobin formation. However, in acutely exposed worms, in spite of a transient peak of GST activity, no induction of antioxidant enzymes occurs, leading to morphological and physiological changes.     

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.