Molecular identity,morphology and taxonomy of the Rhynchospio glutaea complex with a key to Rhynchospio species (Annelida,Spionidae)

Rhynchospio glutaea (Ehlers, 1897 Ehlers, E. 1897. Polychaeten. Ergebnisse der Hamburger Magalhaensischen Sammelreise 1892/93. L. Friederichsen & Co., Hamburg, Germany. [Google Scholar]), Rhynchospio arenincola Hartman, 1936 Hartman, O. 1936. New species of Spionidae (Annelida Polychaeta) from the coast of California. University of California Publications in Zoology 41, 45–52. [Google Scholar] and Rhynchospio arenincola asiatica Chlebovitsch, 1959 Chlebovitsch, V.V. 1959. Species of Polychaeta worms from the Kurile Islands, which are new or recorded for the first time in the USSR fauna. Zoologichesky Zhurnal 38, 167–181. [Google Scholar], originally described from Strait of Magellan, California, and South Kurile Islands respectively, appear similar to each other in adult morphology. These species and subspecies have been considered by some authors as subjective synonyms and are here referred to as members of the R. glutaea complex. Sequence data of four gene fragments (2465 bp in total) of the mitochondrial 16S rDNA, nuclear 18S and 28S rDNA, and Histone 3 have shown that R. glutaea complex individuals from the South West Atlantic (Argentina), North East Pacific (British Columbia and Oregon) and North West Pacific (South Korea) were genetically distant and not conspecific. These data also indicate that R. arenincola from North America and R. aff. asiatica from South Korea are more closely related to each other, and both are closer to R. glutaea from South America than to R. nhatrangi from Vietnam: nhatrangi (glutaea (arenincola-aff. asiatica)). Adults of the R. glutaea complex are hermaphrodites and the arrangement of gametes is suggested to be a crucial reproductive character for distinguishing these species. Based on this character, two species of the complex are apparent in the North West Pacific: R. asiatica Chlebovitsch, 1959 Chlebovitsch, V.V. 1959. Species of Polychaeta worms from the Kurile Islands, which are new or recorded for the first time in the USSR fauna. Zoologichesky Zhurnal 38, 167–181. [Google Scholar] stat. nov. from the Kurile Islands (not analysed here) and an unnamed species from the mainland coast of Asia (here referred to as R. aff. asiatica). Adult morphology of R. asiatica stat. nov. is briefly described and illustrated. The lectotype and the type locality of the species on Iturup Is. are established for the first time. An identification key is provided to the 10 currently recognized Rhynchospio species.     

Indo-Pacific Syllidae (Annelida,Phyllodocida) share an evolutionary history

The geographic distribution of three intriguing genera of Syllidae (Annelida, Phyllodocida), Alcyonosyllis, Paraopisthosyllis and Megasyllis, is restricted to the Pacific Ocean, Indian Ocean and the Red Sea. In this study new material of several species in all three genera is identified, and the distributions of the species and genera refined; four species of Alcyonosyllis were found to have increased ranges. Additionally, one new species of Paraopisthosyllis is described. Paraopisthosyllis pardus sp. nov. is characterized by having long bidentate bladed-chaetae, segments covered by small papillae, and a colour pattern consisting in dark red-brown antennae and dorsal cirri and several transversal dark red-brown lines per segment. The three genera share several striking morphological characteristics, such as alternation in the arrangement of dorsal cirri, wide segments with secondary annuli and bright, contrasting colour patterns. Alcyonosyllis species are found in association with other organisms, most noticeably anthozoans, whereas members of Paraopisthosyllis and Megasyllis are free living. Molecular information (sequences of DNA) from Alcyonosyllis species (including the type species, A. phili), and the type of Megasyllis (M. corruscans) is included herein for the first time in a phylogenetic analysis. A phylogenetic analysis performed through different methods (Maximum parsimony and Maximum likelihood) using sequences of three genes (18S, 16S and COI) reveals that all the three genera form a monophyletic group within Syllidae, with several synapomorphies, and a common ancestor probably from the Indo-Pacific. Their geographic distribution pattern, the relationships between these genera and the rest of syllids, and the symbiosis in Alcyonosyllis are discussed.     

支序分类CLADISTICS在浮游多毛类眼蚕科ALCIOPIDAE系统演化研究中的应用

本论文运用支序分类学,借助于计算机技术,详细探讨了浮游多毛类眼蚕科(Alciopidae)九属的系统演化关系,并同前人的进化分类学和数值分类学研究结果进行了比较,对眼蚕科各属的演化途径提出了新的见解。     

Determination of food sources of marine invertebrates from a subtidal sand community using analyses of fatty acids and stable isotopes

The fatty acid compositions and stable isotope ratios of carbon, nitrogen, and sulfur were analyzed in the bivalve mollusks Mactra chinensis, Pandora pulchella, Felaniella usta, and Megangulus zyonoensis, the polychaete Chaetopterus cautus, and the main sources of organic matter in a subtidal sand bottom community in Vostok Bay (Sea of Japan). The fatty acid composition and stable isotope ratios of some bivalves is likely to be indicative of substantial inputs from benthic microalgae and an important role of microbial food chains. Only the filter-feeding polychaete C. cautus showed similarity in these characteristics to suspended particulate organic matter synthesized by phytoplankton. It is suggested that the contribution of benthic microalgae to the diet of a consumer organism, inferred solely from the carbon stable isotope analysis, can be significantly overestimated.     

A multigene framework for polychaete phylogenetic studies

Producing a robust phylogenetic reconstruction for Polychaeta using either morphological or molecular data sets has proven very difficult. There remain many conflicts between morphological analyses and hypotheses based on DNA data, the latter principally derived from 18S rRNA sequences. For the present study a data set covering a broad range of polychaete diversity was assembled, including 38 new sequences from 21 species. Besides available 18S rRNA data, five additional gene segments were examined: the D1 and D9-10 expansion regions of 28S rRNA, histone H3, snU2 RNA and cytochrome c oxidase subunit I. Maximum parsimony, maximum likelihood and Bayesian analyses were conducted.Annelida and Mollusca were reciprocally monophyletic in maximum likelihood analyses, but Polychaeta included a cephalopod in maximum parsimony analyses, and a patellogastropod in Bayesian analyses. When rooted on the Mollusca, optimal topologies from maximum likelihood analyses showed a recognisable basal group of taxa, including Oweniidae, Chaetopteridae and Amphinomidae. The six studied phyllodocidan families plus Orbiniidae (as the sister group of the scale-worms) formed the next most basal group. All analyses support the inclusion of Echiura, Clitellata and Siboglinidae within polychaetes. Bayesian analyses show Echiura as the sister group of Capitellidae, in agreement with previous 18S rRNA results, In contrast, Echiura formed the sister group to Trichobranchidae in maximum likelihood and maximum parsimony analyses.Supra-familial groupings consistently recovered within Polychaeta in the analyses are: (i) Terebellida without Ampharetidae; (ii) Scolecida (excepting Orbiniidae); (iii) Eunicidae, Lumbrineridae and Clitellata; and (iv) “Cirratuliformia” (including Sternaspidae) plus Sabellidae, Serpulidae and Spionidae.     

Changes in Resistance to Salinity and Temperature in Some Species of Polychaetes from the White Sea in Early Ontogenesis

We studied the effects of different salinities on plankton larvae of some polychaetes in the White Sea. It has been found that the salinity resistance of Alitta virens (Nereidae) increases during ontogenesis. Successful fertilization and further larval development in this species occur at the salinity of 22 to 34; embryos taken into the experiment at the stage of 32 blastomeres, trochophores, and early nektochaetes could survive and normally develop at the salinity of 16–32, 14–45, and 12–45 respectively. The rate of settling and metamorphosis in late nektochaetes of A. virens at normal or lowered (down to 14) salinity is dependent on temperature in the range of 5 to 23°C. It is found that the larvae of Harmothoe imbricata (Polynoidae) show the greatest salinity resistance at the stage of nektochaeta, whose lower limit of salinity is 14. Later larval stages of these species can survive in a wide range of salinity due to the development of a provisory nephridial system. The eurybionty of larvae of Spirorbis spirorbis ready for metamorphosis was higher than that in the larvae of Circeus spirillum (Spirorbidae). Under salinity reduced down to 10 the larvae of S. spirorbis die in 8–14 days, whereas more stenohaline larvae of C. spirillum die by the 3-rd day of the experiment. At water temperatures under 5°C the survival of S. spirorbis was the highest at three examined values of salinity, whereas C. spirillum showed the highest survival only under normal salinity.     

Where did Marenzelleria spp. (Polychaeta: Spionidae) in Europe come from?

Abundant populations of Marenzelleria spp. were reported for the first time in the North Sea during the late 1970s and then in the Baltic Sea in 1985. Genetic analysis by means of allozyme electrophoresis and sequencing of a segment of mitochondrial 16srDNA showed that two different genetic types or sibling species of Marenzelleria were present in Europe. Marenzelleria Type I is found only in the North Sea, whereas Type II has been found in both the North Sea and the Baltic Sea. The North Sea animals (Type I) correspond to Type I specimens found in North American coastal waters between Nova Scotia and Cape Henlopen (Delaware), while Marenzelleria Type II from the Baltic Sea correspond to Marenzelleria type II animals from the Arctic (Tuktoyaktuk Harbor, Northwest Territories, Canada), New Hampshire and coastal waters between Chesapeake Bay southward to the Ogeechee River (Georgia). Human mediated introduction (by shipping) and natural range expansion are discussed as the possible causes of these virtually simultaneous invasions by two sibling species. Marenzelleria Type I appears to colonize habitats with a higher salinity and/or in which salinities tend to fluctuate considerably. The osmolality of the coelomic fluid after acclimation to various salinities between 0.25 and 18 is the same for both sibling species/genetic types. Although the two types do not differ in respect of hyperosmoregulation (<10), differences may exist in their cell volume regulation or its time course in the almost isoosmotic range at salinities >10.     

Lateral organs in sedentary polychaetes (Annelida) – Ultrastructure and phylogenetic significance of an insufficiently known sense organ

Lateral organs are sense organs visible as densely ciliated pits or papillae between the noto‐ and the neuropodia in certain taxa of sedentary polychaetes. Ultrastructural studies in about 10 species of the following taxa Maldanidae, Opheliidae, Orbiniidae, Paraonidae, Magelonidae, Spionidae, Poecilochaetidae and Terebellidae have been designed to evaluate whether these organs are homologous among polychaetes. In spite of great external diversity, the investigations revealed an overall ultrastructural similarity. Differences between species investigated mainly concern the size of the organs as well as the number and arrangement of cells. The organs comprise supportive cells and uniciliated penetrative sensory cells. Their dendrites are closely arranged and thus their cilia may resemble multiciliated cells. There are two types of sensory cells: one type possesses no or mainly thin microvilli of which usually only a few reach the cuticular surface, and in the other type the cilium is consistently surrounded by 10 strong microvilli, which form a pore‐like opening in the cuticle. Further differences occur in the structure of the rootlet system. Basally, a retractor muscle attaches to the organ. The systematic significance of these organs within Annelida is discussed with respect to the conflicting phylogenetic hypotheses explaining the relationships of annelid taxa.     

Spatial and temporal changes in epibenthic communities at deep,hard bottom aquaculture sites in Newfoundland

This study explores spatio-temporal changes in epibenthic communities due to salmon aquaculture at deep (>30 m), hard-bottom dominated sites in Newfoundland (Canada). Using a drop-camera approach, we followed changes with production at two aquaculture sites, as well as potential recovery processes at two fallow sites, comparing their epibenthic composition with nearby non-production sites. Multivariate analysis revealed that aquaculture production led to rapid changes in epibenthic communities, as evidenced primarily by the presence of two visual indicators: bacterial mats and opportunistic polychaetes. Due to low taxon richness and abundances, as well as variability among sites, no clear intermediate successional stages were apparent. Beggiatoa-like mats and opportunistic polychaetes appeared rapidly once aquaculture production began; these indicators did not increase in spatial extent during the survey and were typically observed up to 100 m, and occasionally up to 160 m from cages. After 15 months of fallowing, Beggiatoa-like mats and opportunistic polychaetes remained present at sites and were occasionally accompanied by few other taxa, suggesting incomplete recovery.     

A complex species complex: The controversial role of ecology and biogeography in the evolutionary history of Syllis gracilis Grube, 1840 (Annelida,Syllidae)

The cryptic diversity in the polychaete Syllis gracilis Grube, 1840, in the Mediterranean Sea was examined with an integrative morpho-molecular approach. Individuals of S. gracilis were collected at eleven Mediterranean localities to provide an insight into the role of brackish environments in inducing cryptic speciation. The examination of morphological features combined with a molecular genetic analysis based on a partial sequence of the 16S rRNA gene highlighted discrepancies between morphological and molecular diversity. Morphological data allowed to identify a morphotype with short appendages occurring in coralline algae communities and another one with long appendages observed in brackish-water environments and Sabellaria reefs. Multivariate analyses showed that sampling localities were the greatest source of morphological divergence, suggesting that phenotypic plasticity may play a role in local adaptations of S. gracilis populations. Molecular data showed the occurrence of four divergent lineages not corresponding to morphological clusters. Different species delimitation tests gave conflicting results, retrieving, however, at least four separated entities. Some lineages occurred in sympatry and were equally distributed in marine and brackish-water environments, excluding a biogeographic or ecological explanation of the observed pattern and suggesting instead ancient separation between lineages and secondary contact. The co-occurrence of different lineages hindered the identification of the lineage corresponding to S. gracilis sensu stricto. The discrepancy between morphological and molecular diversity suggests that different environmental and biogeographic features may interact in a complex and unpredictable way in shaping diversity patterns. An integrative approach is needed to provide a satisfactory insight on evolutionary processes in marine invertebrates.