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Annelida, Polychaeta, Chaetopteridae

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Annelida, Polychaeta, Chaetopteridae Zoological Journal of the Linnean Society, 2008, 152, 201–225. With 12 figures Description of a new species of Mesochaetopterus (Annelida, Polychaeta, Chaetopteridae), with redescription of Mesochaetopterus xerecus and an approach to the phylogeny of the family Downloaded from https://academic.oup.com/zoolinnean/article/152/2/201/2630849 by guest on 25 March 2021 DANIEL MARTIN1*, JOÃO GIL1, JOSEP CARRERAS-CARBONELL1 and MICHEL BHAUD2 1Centre d’Estudis Avançats de Blanes (CSIC), Carrer d’accés a la Cala Sant Francesc 14, 17300 Blanes (Girona), Catalunya, Spain 2Observatoire Océanologique de Banyuls, Université P. et M. Curie – CNRS, BP 44, 66650 Banyuls-sur-Mer, Cedex, France Received 28 March 2006; accepted for publication 21 May 2007 Mesochaetopterus rogeri sp. nov., a new large chaetopterid polychaete from the Mediterranean Sea, is described. The analyses of partial sequences from the nuclear 18S rRNA (643 bp) and the mitochondrial cytochrome oxidase I (577 bp) genes of representative individuals of all known chaetopterid genera indicated the initial assignment of the new species into Mesochaetopterus. These analyses also supported the monophyly of the family and revealed two well-supported clades: Chaetopterus/Mesochaetopterus and Spiochaetopterus/Phyllochaetopterus. Mesocha- etopterus rogeri sp. nov. is close to Mesochaetopterus xerecus, which is redescribed here from newly collected material. Mesochaetopterus rogeri sp. nov. was characterized as follows: (1) two long tentacles with dorsal transversal black bands of alternating widths (sometimes with two additional longitudinal light-brown bands); (2) region A with nine chaetigers (up to 12), with 13–19 modified chaetae in the fourth chaetiger; (3) region B with three flat segments, with accessory feeding organs in the second and third segments; (4) sandy straight tubes, 2.5-m long or more, vertically embedded in the sand. In the Bay of Blanes, M. rogeri sp. nov. occurs between 6- and 9-m deep (but also up to 30-m deep), in a patchy distribution (< 1 individual m-2), with maximum densities in April/June (likely to be the result of recruitment events), and minimum densities in September/November (likely to be a behavioural response to increasing sediment dynamics). Although it was originally thought that M. rogeri sp. nov. could be an introduced species, we argue that it is probably a native of the Mediterranean that has been overlooked by scientists up to now. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 201–225. ADDITIONAL KEYWORDS: 18S rRNA – behaviour – Chaetopterus – cytochrome oxidase I – distribution – genetics – Mediterranean Sea – Phyllochaetopterus – Spiochaetopterus. INTRODUCTION data on their presence were these incidental reports, sometimes accompanied by pictures of the living During recent years, scuba divers have occasionally specimens ‘in situ’ (Fig. 1A, B). The oldest known ref- reported the presence of ‘strange’ large worms, with a erence, a worm from Almería, on the south-western pair of long palps and a characteristic striped ‘black Iberian Penninsula, appeared in an encyclopaedia in and white’ colour pattern, in shallow sandy bottoms of 1979 labelled as ‘Spionidae’ (George & George, 1979: the Iberian Mediterranean shoreline, mainly along fig. 8, 353). the Catalan littoral. Up to now, the only available Routine monitoring of the brine discharge from the Blanes desalination plant in the shallow sandy bottoms facing the beach at Punta del Tordera *Corresponding author. E-mail: [email protected] (Catalan coast, north-western Mediterranean) © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 201–225 201 202 D. MARTIN ET AL. Downloaded from https://academic.oup.com/zoolinnean/article/152/2/201/2630849 by guest on 25 March 2021 © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 201–225 NEW MESOCHAETOPTERUS FROM THE MEDITERRANEAN 203 Figure 1. Mesochaetopterus rogeri sp. nov. A, B, Two typical positions of the protruded tentacles in living worms (underwater pictures taken by L. Dantart). C, Regions A, B, and C (preserved worm, morphotype with reddish bands). D, Ventral view of region A (preserved worm, morphotype without reddish bands). E, Detail of the anterior end (dorsal view). F, Detail of the anterior end (dorsal view, tentacles removed). Ar, region A; b1–b3, segments of region B; d1–d3, dorsal rami of region-B segments; v1–v3, ventral rami of region-B segments; Cr, region C. Scale bars are in cm. ᭣ revealed a stable population, allowing the collection of shoreline of the Mediterranean coast of the Iberian enough appropriate material. Peninsula (Fig. 2), from October 2001 to September In this study, we used a partial nuclear gene 2004. As a first collection method, we used a PVC Downloaded from https://academic.oup.com/zoolinnean/article/152/2/201/2630849 by guest on 25 March 2021 (18S rRNA) and a mitochondrial gene, cytochrome plate introduced obliquely into the sediment to cut oxidase I (COI), aiming to confirm the assignment of across the worm’s tube, trying to prevent the worms the Catalan specimens into the correct genus within from withdrawing deeper inside the tube. The body in the Chaetopteridae (Annelida, Polychaeta), as well Mesochaetopterus is divided into three differentiated as to assess the phylogenetic relationships within parts: the nine-to-ten anteriormost segments, the the four known genera of the family: Chaetopterus, two-to-three mid-body segments, and an undefined Mesochaetopterus, Spiochaetopterus, and Phyllocha- (but usually very numerous) number of posterior seg- etopterus. ments; these form the regions A, B, and C, respec- The Catalan specimens were revealed to be an tively. Using the PVC plate, the quick reaction of unknown species of Mesochaetopterus. Thus, they are the worm only allowed for the collection of anterior described as a new species, Mesochaetopterus rogeri fragments containing region A and, at most, part of sp. nov, in spite of a lack of data on the morpho- the region B (as well as part of the tube). Successive logy and chaetal arrangement of the posteriormost attempts using either irritating or anaesthetizing segments (because of the logistic difficulties of ex- compounds, and sucking or impelling pumps of tracting complete worms). They were compared with different types, did not result in collection of all known species of the genus, particularly with either entire specimens or larger fragments. The Mesochaetopterus xerecus Petersen & Fanta, 1969, last attempt, carried out in September 2004, involved which closely resembles the new species both in size the archaeological research vessel ‘Tethys’, from the and in having striped palps (Petersen & Fanta, Centre d’Arqueologia Submarina of the Museu 1969). In the case of M. xerecus, however, all material d’Arqueologia de Catalunya. The vessel is equipped was lost (including the types). For this study, new with powerful suction devices, which allowed us to material collected at Paranaguá Bay (Paraná, Brazil) excavate up to 2.5-m deep into the sediment. was used both in genetic and morphologic analyses. However, the tube extended deeper into the sediment, In addition, we provide a brief diagnosis of the soft and only a large fragment of a worm (with ten seg- body of the species, together with a full description ments of region C, in addition to regions A and B) was and pictures of its chaetal arrangement, which was collected after several hours of underwater working. poorly described and illustrated in the original Being the most complete available fragment, this description. specimen was selected as the holotype of the new The seasonal trends of the population in Punta species. del Tordera, habitat preferences, and behaviour of The newly collected specimens of M. xerecus were M. rogeri sp. nov., are also provided, and are com- found in the Ilha do Mel, Paranaguá Bay (Paraná, pared with the known traits of the other known Brazil). The worms were collected by researchers large-sized species of Mesochaetopterus, and are from the Centro de Estudos do Mar (Pontal do Sul, then discussed in an attempt to explain the pres- Brazil) from intertidal soft sediments at low tide ence of such a large unknown worm in the widely using a shovel. investigated shallow waters of the Mediterranean Sea. GENETIC ANALYSIS, DNA EXTRACTION AND SEQUENCING MATERIAL AND METHODS For the genetic analyses, all newly collected speci- SAMPLING PROCEDURE mens were preserved directly in 100% ethanol. Total All observations on living worms and sampling were genomic DNA was extracted from individuals using performed by scuba diving. Most sampling attempts the QIAamp DNA Minikit (Qiagen). A fragment of were carried out off Punta del Tordera on the Catalan 18S rRNA nuclear gene was amplified by polyme- © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 201–225 204 D. MARTIN ET AL. Downloaded from https://academic.oup.com/zoolinnean/article/152/2/201/2630849 by guest on 25 March 2021 Figure 2. Mesochaetopterus rogeri sp. nov., location of reports. A, Iberian Peninsula. B, Catalan coast. C, Blanes litoral. Key: , seasonal monitoring at the Punta del Tordera; 1, Almería; 2, Alicante; 3, Valencia. rase chain reaction (PCR) using newly designed pri- (Genotek), and 20–30 ng genomic DNA. PCR was mers (Chae18SF, 5′-AAACGGCTACCACATCCAAG- performed in a Primus 96 plus (MWG Biotech). PCR 3′; Chae18SR, 5′-AACTAAGAACGGCCATGCAC-3′). products were cleaned with the QIAquick PCR Puri- Cycle parameters consisted of first a denaturing step fication Kit (Qiagen) or ethanol precipitation, and at 94 °C for 2 min, followed by 35 cycles of 1 min at then sequenced with the BigDye Sequencing Kit ABI 94 °C, 1 min at 55 °C, and 1 min at 72 °C, and then a Prism. PCR products were purified by ethanol pre- final extension at 72 °C for 7 min. The mitochondrial cipitation and analysed on an ABI 3700 automatic COI gene was amplified using primers HCO2198 sequencer (Applied Biosystems) from the Serveis (5′-TAAACTTCAGGGTGACCAAAAAATCA-3′) and Cientifico-Tècnics of the Universitat de Barcelona.
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