When molecules support morphology: Phylogenetic reconstruction of the family Onuphidae (Eunicida, Annelida) based on 16S rDNA and 18S rDNA.

Onuphid polychaetes are tubicolous marine worms commonly reported worldwide from intertidal areas to hadal depths. They often dominate in benthic communities and have economic importance in aquaculture and recreational fishing. Here we report the phylogeny of the family Onuphidae based on the combined analyses of nuclear (18S rDNA) and mitochondrial (16S rDNA) genes. Results of Bayesian and Maximum Likelihood analyses supported the monophyly of Onuphidae and its traditional subdivision into two monophyletic subfamilies: Onuphinae and Hyalinoeciinae. Ten of 22 recognized genera were monophyletic with strong node support; four more genera included in this study were either monotypic or represented by a single species. None of the genera appeared para- or polyphyletic and this indicates a strong congruence between the traditional morphology-based systematics of the family and the newly obtained molecular-based phylogenetic reconstructions. Intergeneric relationships within Hyalinoeciinae were not resolved. Two strongly supported monophyletic groups of genera were recovered within Onuphinae: ((Onuphis, Aponuphis), Diopatra, Paradiopatra) and (Hirsutonuphis, (Paxtonia, (Kinbergonuphis, Mooreonuphis))). A previously accepted hypothesis on the subdivision of Onuphinae into the Onuphis group of genera and the Diopatra group of genera was largely rejected.

Two new species of Marphysa (Annelida, Eunicidae) from southern Australia.

Two new species of Marphysa Quatrefages, 1866 are described from the southeast coast of Australia. With the presence of only compound spinigers and the branchiae present over many chaetigers, Marphysa baudini n. sp. belongs to the Sanguinea-group. This species has ventral cirri with an inflated base and digitiform tip and thick and wide anodont pectinate chaetae, with 3-5 internal long and thick teeth. With the presence of only compound falcigers, Marphysa davidattenboroughi n. sp., belongs to the "Aenea-group". This species is characterised by the presence of a bilobed prostomium, a single pair of pygidial cirri and by the presence of thick, and wide anodont pectinate chaetae with 4-6 long internal and thick teeth.

A closer look at the taxonomic and genetic diversity of endemic South African Marphysa Quatrefages, 1865.

The current study investigates the final unresolved cosmopolitan species of Marphysa in South Africa, Marphysa corallina, collected from KwaZulu Natal, Eastern and Western Cape provinces, together with another species collected from northern KwaZulu Natal. Morphological and genetic data prove that M. corallina, originally described from Hawaii, does not occur in South Africa. The curvature of the inner base on maxilla I, the elevated inner base of maxilla II, and the ventral cirrus as a transverse welt with a rounded tip allow us to identify it as a new species of Treadwellphysa, T. izinqa sp. nov. (common name: brown wonderworm). Characteristic traits include the basal reddish and distal golden colour of the subacicular hook, the ear-shaped postchaetal lobe, and tridentate falcigers which is reported for the first time for the genus. This species is harvested as bait on the south coast of SA, although less frequently than the more common blood wonderworm, Marphysa haemasona Quatrefages, 1866, and can be distinguished by its more uniform brown colouration and white-tipped antennae. A second species, Marphysa mzingazia sp. nov., is characterized by red eyes, six branchial filaments extending to the posterior end, the golden aciculae in posterior chaetigers, weakly bidentate yellow/brown subacicular hooks, and the presence of similar sized spinigers along the body. A molecular analysis based on cytochrome oxidase I fragments confirm both taxa as different species. A key for all South African species of Marphysa is included.

A new species of giant Eunice (Eunicidae, Polychaeta, Annelida) from the east coast of Australia.

A new giant species is described from New South Wales, Australia. Eunicedharastii sp. nov. differs from described Australian species and is most similar to E.aphroditois (Pallas, 1788), E.flavopicta Izuka, 1912, and E.kinbergi Ehlers, 1868. The unique combination of features that characterizes the new species is irregular articulated prostomial appendages; antennae reaching back beyond chaetiger 4; branchiae starting at chaetiger 10, initially button-shaped and distinctly longer than notopodial cirri where best developed; dorsal fleshy knobs on anterior chaetal lobes; notopodial cirri pendulous, abrupt tapering from inflated bases; bidentate compound falcigerous chaetae with both teeth directed laterally, distal tooth much shorter than proximal tooth in median and posterior chaetigers; and dark bidentate subacicular hooks starting at chaetiger 58, tapering to a small head with both teeth directed distally, and proximal tooth much larger than minute and spur-like distal tooth. This new species lives in sandy sediments in coastal waters 1-8 m deep. It is highly mobile and not easy to collect, which may explain why it was not described before.

Phylogeny of the bristle worm family Eunicidae (Eunicida, Annelida) and the phylogenetic utility of noncongruent 16S, COI and 18S in combined analyses.

The bristleworm family Eunicidae is distributed worldwide and well-known for the large size of many of its species, its hard jaws and its economic importance in the bait industry. Monophyly of Eunicidae has been contradicted in previous molecular and morphological phylogenetic analyses focusing on Eunicida or on its type genus, Eunice. The current study focused on the phylogeny of the family per se combining mitochondrial and nuclear genes in different analyses. It recovered well-resolved phylogenetic hypotheses supporting the monophyly of Eunicidae and Palola, the only monophyletic genus among the genera for which we tested their monophyly (Eunice, Marphysa, Palola, Lysidice and Nematonereis). Four other stable clades containing the type species of different genera, or species deemed very similar to them, were recovered within Eunicidae. These clades may represent monophyletic redefinitions of current genera, except for Eunice, and of previous synonymized genera. Evolution of the number of peristomial cirri and prostomial appendages in Eunicidae happened by independent step-by-step reduction in the opposite order of their ontogenetic development, suggestive of sequential heterochrony. All three genes were informative, however, at different levels within the combined trees. 16S and COI were important in recovering a monophyletic Eunicidae and relationships within the family, while 18S was important in the resolution of basal eunicidan relationships, monophyly of Onuphidae and basal relationships within this family. Moreover, results of congruence tests (SH and WRST using PABA) indicate that hidden support is picked up in the combined analyses, which is not revealed in the single gene analyses. Further supporting the idea that congruence is not a requirement for combining different partitions.

Eunice sensu lato (Annelida: Eunicidae) from Australia: description of seven new species and comments on previously reported species of the genera Eunice, Leodice and Nicidion.

Species traditionally considered to belong to Eunice are now, also, distributed in two other genera Leodice and Nicidion recently resurrected to reconcile Eunicidae taxonomy with its phylogenetic hypothesis. In Australia, Eunice species have been reported from all seas and sum up to 22 species. In this study, we propose 10 new combinations for traditional Eunice species, which should be moved to Leodice; describe seven new species, four of Eunice and three of Leodice; and comment on species previous recorded from the Australian Coast. Previous records of E. denticulata, E. filamentosa, E. grubei, E. indica, E. longicirris, E. microprion, E. paupera, E. tridentata, E. tubifex and E. vittata from Australia are considered dubious. Eunice tribranchiata, originally described from Australian specimens, is considered here to be an indeterminable species. This study also includes descriptions of Eunice, Leodice and Nicidion as well as key to these genera and their species recorded from Australia.

Morphological and molecular systematic review of Marphysa Quatrefages, 1865 (Annelida: Eunicidae) species from South Africa.

A vast polychaete fauna is hidden behind complexes of cryptic and pseudo-cryptic species, which has greatly hindered our understanding of species diversity in several regions worldwide. Among the eunicids, Marphysa sanguinea Montagu, 1813 is a typical example, recorded in three oceans and with various species considered its junior synonyms. In South Africa, specimens previously misidentified as M. sanguinea are now known as Marphysa elityeni Lewis & Karageorgopoulos, 2008. Of the six Marphysa Quatrefages, 1865a species recorded from the same region, three have their distributions restricted to South Africa while the others are considered to have worldwide distributions. Here, we evaluated the taxonomic status of the indigenous M. elityeni and investigated the presence of the widespread species Marphysa macintoshi Crossland, 1903 and Marphysa depressa Schmarda, 1861 in South Africa using morphological and molecular data. Our results reveal that M. elityeni is a junior synonym of Marphysa haemasoma, a species previously described from South Africa which is herein reinstated as a valid species. Both M. macintoshi and M. depressa are not present in South Africa and their status as being distributed worldwide deserves further investigation. Marphysa durbanensis Day, 1934 and the new species described here, M. sherlockae n. sp., had been misidentified as M. macintoshi and M. depressa respectively. Thus, the number of Marphysa species with distributions restricted to South Africa increased from three to five. This study reiterates the importance of implementing an integrated taxonomic framework to unravel local biodiversity.

Correction: Digging the diversity of Iberian bait worms Marphysa (Annelida, Eunicidae).

[This corrects the article DOI: 10.1371/journal.pone.0226749.].

Digging the diversity of Iberian bait worms Marphysa (Annelida, Eunicidae).

During a visit to polychaete-rearing facilities in the vicinity of Bay of Cádiz (SW Iberian Peninsula, Atlantic Ocean), we sampled two populations of Marphysa (Annelida, Eunicidae) originally occurring at nearby intertidal soft bottoms, one being more than twice as long as the other at the same age. We analysed them using partial sequences of two mitochondrial genes, 16S rDNA and Cytochrome Oxidase I, and classical morphological observations. Our molecular results confirmed that the two populations corresponded to two different species, with PTP species delimitation values ranging from 0.973 (long-bodied species) to 0.999 (short-bodied species). Morphologically, the short-bodied species resembles the recently redescribed M. sanguinea (Montagu, 1813), but differs mainly in having some parapodia with two subacicular hooks (one bidentate and one unidentate) and three types of pectinate chaetae, Two isodont present all along the body, and one particularly large anodont asymmetric appearing only from mid-posterior parapodia. The long-bodied species resembles Marphysa aegypti Elgetany, El-Ghobashy, Ghoneim and Struck, 2018 both in size and in having very robust, unidentate subacicular hooks (single in most parapodia, two-both similar in size and form-in some posterior parapodia), but differs, among other features, in the maxillary formula, the number of acicula per parapodia and the number and shape of pectinate chaetae. Accordingly, we are here fully illustrating and formally describing the two Iberian populations as Marphysa gaditana sp. nov. (short-bodied) and Marphysa chirigota sp. nov. (long-bodied) and we are emending the description of M. aegypti based on our revision of the type material. Also, we discuss on the distribution of the species of the sanguinea-group and on the relevancy of taxonomically robust studies when dealing with species of commercial interest having the potential of being globally spread through human activities, as well as on the misunderstandings caused by the incorrect use of the "cosmopolitan species" concept.

Pilargidae (Annelida: Phyllodocida) from coastal and deep waters of the Southwestern Atlantic, with descriptions of two new species.

We describe seven species of Pilargidae from the coast and deep sea (300-1400 m deep) of southeastern Brazil, belonging to the genera Ancistrosyllis McIntosh, 1878, Cabira Webster, 1879, Glyphohesione Friedrich, 1950, Hermundura Müller, 1858, and Pilargis Saint-Joseph, 1899. Of those, two are new species (Glyphohesione campensis sp. nov. and Pilargis falconae sp. nov.) and two are new records of Ancistrosyllis for Brazil. Glyphohesione campensis sp. nov. is distinguished from its congeners by fewer neurochaetae, the anteriormost appearance of first notopodial spines, and intermediate relative length of dorsal and ventral cirri. Pilargis falconae sp. nov. is diagnosed by the presence of parapodial glands from the first chaetiger on the dorsal surface and from the sixth chaetiger on the ventral surface; and neurochaetae smooth capillaries and limbates with unidentate tips. Keys to pilargid species reported from, or likely to occur in, Brazilian waters are provided.

Guess who? On the importance of using appropriate name: case study of Marphysasanguinea (Montagu, 1813).

The common bait worm Marphysasanguinea (Montagu, 1813), originally described from the south coast of England, is the type species of the genus. This species has been widely reported from all around the world and has been considered as cosmopolitan until recently. This is partly because the original description was very brief and poorly illustrated, and also because all species superficially look similar. In order to clarify the situation, M.sanguinea was redescribed and a neotype was designated by Hutchings and Karageorgpoulos in 2003. Recently, specimens from Cornwall, close to the type locality, were sampled, examined morphologically, and used to obtain COI gene sequences for this species. Molecular results permitted us to confirm the identity and presence of M.sanguinea along the French coasts and to highlight the presence of inaccurate sequences of this species on GenBank. Use of this "false" cosmopolitan species at a worldwide scale by many biologists is also discussed in this paper.

Species of Eunicidae (Annelida, Errantia, Eunicida) from rhodolith beds off the states of Paraíba and Pernambuco, northeastern Brazilian coast.

Members of one species of Leodice Lamarck, 1818, one species of Lysidice Lamarck, 1818, and another of Palola Gray in Stair, 1847 were found among material from rhodolith beds, collected off the states of Paraíba and Pernambuco, northeastern Brazilian coast. Of those species, one is new to science and the other two are new records for the state of Pernambuco, although their occurrence on neighbouring areas was already known. Leodice calcaricola sp. nov. resembles morphologically L. miurai (Carrera-Parra Salazar-Vallejo, 1998) as members of both species share similar body size, moniliform prostomial appendages, similar numbers of branchial filaments (1-3 per branchia), and, especially, by having falcigers with bidentate blades on anterior chaetigers, and tridentate on posterior ones. These species differ, however, since specimens of L. calcaricola sp. nov. have distinctly shorter antennae and palps, homodont and heterodont pectinate chaetae, three types of aciculae and two subacicular hooks per midbody parapodium. The other two species which members were found on the rhodolith beds are Lysidice cf. unicornis (Grube, 1840) and Palola brasiliensis Zanol, Paiva Attolini, 2000. We describe herein new species to science and provide a redescription for L. cf. unicornis because the Brazilian specimens of this species have never been formally described.

The use of metabarcoding for meiofauna ecological patterns assessment.

Marine meiofauna comprises up to 22 phyla. Its morphological identification requires time and taxonomists' expertise, and molecular tools can make this task faster. We aim to disentangle meiofaunal diversity patterns at Araçá Bay by applying a model selection approach and estimating the effectiveness of metabarcoding (18S rDNA) and morphological methods for estimating the response of meiofauna diversity in small-scale interactions with environmental variables. A rarefaction curve indicated that ten samples were sufficient for estimating the total number of meiofauna OTUs in a tidal flat. In both approaches, richness was predicted by mean sand percentage, sediment sorting, and bacteria concentration. Nematode genera composition differed significantly between approaches, the result of taxonomic mismatch in the genetic database. The similarity between the model selected for diversity descriptors, the richness of nematode genera and meiofauna composition emphasized the utility of predictive models for metabarcoding estimates to detect small-scale interactions of these organisms.

One new species and two redescriptions of Marphysa (Eunicidae, Annelida) species of the Aenea-group from Australia.

Species of Marphysa belonging to the Aenea-group have compound falcigers as the subacicular chaetae along the entire body. Around 30 species belong to this group, among these, two have originally been described based on Australian specimens, M. bifurcata and M. sessilobranchiata. Here, we redescribe both species based on type and non-type material and describe a species new to science, M. pseudosessiloa n. sp., including intraspecific variation. Informative features for the identification of these species are relative length of prostomial appendages in relation to prostomium, shape of notopodial cirri, left MxIV plate, left and right MxIV attachment lamellae and placement of branchial stem. A taxonomic key to species of Marphysa currently reported from Australia is also provided.

New and previously known species of Oenonidae (Polychaeta: Annelida) from Lizard Island, Great Barrier Reef, Australia.

The family Oenonidae consists of Eunicida species with prionognath jaws. Its Australian fauna had been reported to comprise six species belonging to Arabella, Drilonereis, and Oenone. This study provides descriptions for four new species, redescriptions for three species (two previously recorded and a new record, Drilonereis cf. logani) and diagnoses for the genera recorded from Australia. Currently, eleven species of oenonids, distributed in three genera, are known for the Australian coast. On Lizard Island, this family shows low abundance (19 specimens collected) and high richness (seven species). Our results suggest that despite the increasing accumulation of information, the biodiversity of the family is still poorly estimated.

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

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

O caramujo exótico invasor Achatina fulica (Stylommatophora, Mollusca) no Estado do Rio de Janeiro (Brasil): situação atual / The exotic invasive snail Achatina fulica (Stylommatophora, Mollusca) in the State of Rio de Janeiro (Brazil): current status

O caramujo africano invasor Achatina fulica foi introduzido no Brasil a partir de Estados do Sul e Sudeste em pelo menos três ocasiões. Atualmente sua distribuição já abrange 24 dos 26 estados e o Distrito Federal. Populações densas dessa espécie vêm causando incômodos à populações humanas, danos à jardins e pequenas plantações, além de atuarem como transmissoras de duas zoonoses (angiostrongilíase abdominal e meningoencefalite eosinofílica) e outras parasitoses de interesse veterinário. No presente estudo, apresentamos novas ocorrências de A. fulica no Estado do Rio de Janeiro (Brasil) assim como dados sobre as larvas de nematódeos de interesse médico e veterinário encontradas em exemplares dessa espécie coletados neste estado. A. fulica está presente em 26 municípios adicionais em relação aos dados obtidos em 2006, ano do último levantamento, representando um aumento de 50 por cento no número de municípios infestados. Apenas nove dos 92 municípios do estado ainda não registram a presença dessa espécie. Larvas dos nematódeos Aelurostrongylus abstrusus, Rhabditis sp. e Strongyluris-like, todas com importância veterinária, foram encontradas em exemplares de A. fulica de onze municípios. A rápida dispersão de A. fulica evidencia a necessidade de planos de manejo contínuos e eficientes em todo o país visando o controle efetivo dessa invasão. A erradicação de A. fulica é pouco provável devido ao nível atual de infestação.

The invasive African snail Achatina fulica was introduced in Brazil through South and Southeast States in at least three separate occasions. A. fulica is currently present in 24 out of 26 Brazilian States and in the Federal District. Dense populations of A. fulica are nuisance to human populations and pest to gardens and small crops. Such populations also act in the transmission of two zoonosis (abdominal angiostrongyliasis and eosinophilic meningitis) as well as other parasitosis of veterinary importance. Here, we report new records of A. fulica and of nematode larvae of medical and veterinary importance found in this snail in the state of Rio de Janeiro (Brazil). A. fulica is present in 26 additional municipalities of the state when compared to the information obtained in 2006, year of the last census. This result represents an increase of 50 percent in the number of municipalities infested. Only nine out of the 92 municipalities of the state have not yet registered the presence of this invasive species. Nematode larvae of Aelurostrongylus abstrusus, Rhabditis sp. e Strongyluris-like, all of veterinary importance, have been registered in A. fulica individuals from eleven municipalities of the state. The rapid range expansion of A. fulica calls attention for the need of a general and continuous management plan throughout the country in order to effectively control the invasion. The current level of infestation makes a full eradication unlikely.