Sabellaria and Lygdamis (Polychaeta: Sabellariidae) from reefs off northeastern Brazil including a new species of Sabellaria

Zootaxa 3881 (2): 125–144 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press

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ZOOTAXA

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http://dx.doi.org/10.11646/zootaxa.3881.2.2 http://zoobank.org/urn:lsid:zoobank.org:pub:CE5F4D90-4AC0-44F1-8C29-0BC5E92043C8

Sabellaria and Lygdamis (Polychaeta: Sabellariidae) from reefs off northeastern Brazil including a new species of Sabellaria ANDRÉ SOUZA DOS SANTOS1,2,3, ANA CLAUDIA DOS SANTOS BRASIL2 & MARTIN LINDSEY CHRISTOFFERSEN1 1

Laboratório de Invertebrados Paulo Young, Departamento de Sistemática e Ecologia, Centro de Ciências Exatas e da Natureza, Universidade Federal da Paraíba Campus I, CEP 58.059-900, João Pessoa, Paraíba, Brazil. 2 Laboratório de Polychaeta, Instituto de Biologia, Departamento de Biologia Animal, Universidade Federal Rural do Rio de Janeiro, CEP 23.851-970, Seropédica, Rio de Janeiro, Brazil. 3 Laboratório de Ecossistemas Costeiros, Departamento de Biologia, Centro de Ciências Biológicas e da Saúde, Universidade Federal de Sergipe, CEP 49.100-000, Aracaju, Sergipe, Brazil. *Corresponding author: André Souza dos Santos ([email protected])

Abstract Members of the polychaete taxon Sabellariidae Johnston, 1865 are known to live in sand tubes cemented onto rocky substrata, mollusk shells, or sea grasses. Of 37 known Sabellaria species, only nine were reported for the Brazilian coast, in all cases being associated with aggregates of other species. The genus is considered cosmopolitan. Herein we describe for the first time an aggregate of sabellariids composed by Sabellaria nanella and Sabellaria wilsoni. In addition, we describe a new species of Sabellaria. Lygdamis are represented by solitary species. None of the 17 known species were previously reported from the southwestern Atlantic Ocean. We report Lygdamis rayrobertsi for the first time in the South Atlantic. Key words: Taxonomy, Sabellariidae, Sabellaria, Lygdamis, Brazil

Introduction Members of the polychaete taxon Sabellariidae Johnston, 1865 are known as sand-mansion or honeycomb worms. They live in sand tubes cemented onto rocky substrata (Bailey-Brock 1985; Hutchings 2000; Rouse & Pleijel 2003; Bailey-Brock et al. 2007), mollusk shells, or sea grasses (Uebelacker & Johnson 1984; Santos et al. 2011). Some species are known for building solitary tubes (Eckelbarger 1977a), or extensive reefs (Hutchings 2000; Rouse & Pleijel 2001). They occur from subtidal to abyssal depths (up to 4825 meters) (Eckelbarger 1977b; Uebelacker & Johnson 1984; Kirtley 1994). As natural “surf zone engineers” sabellariids offer a number of ecological benefits to marine benthic communities (Kirtley 1994; Nishi et al.2010; Capa et al. 2012), because the complex habitat structure of sabellariid reefs sustain a high biodiversity (Kirtley 1994). As a result, studies of sabellariid ecology, reproductive biology and phylogeny have been attracting biologists and geologists over the years (e.g., Kirtley & Tanner 1968; Pawlik 1988; Hendrick & Foster-Smith 2006; Gruet & Lana 1988; Ayata et al. 2009; Cullotoy et al. 2010; Nishi et al. 2010; Capa et al. 2012). Hutchings (2000) described the sabellariids as polychaetes with compact bodies divided into distinct sections: head, thorax, abdomen and pygidium. The head consists of an operculum with numerous golden paleae which almost close the tube. The prostomium is indistinct from the peristomium. The peristomium is visible only as lips around of the mouth. Paired palps are positioned in the central area of the prostomium, and nuchal organs are present at the bases of the palps. The thoracic region is composed of segments with short and cylindrical neuropodia, notopodia as tori, and elongate dorsal branchiae. The abdomen is composed by a neuropodium that is short and cylindrical, a notopodium that is reduced to a torus, and prolonged gills positioned dorsally. The abdomen is also indicated by the presence of chaetal inversion, where uncini are notopodial and neurochaetae are capillaries. The first revision of the family was provided by Kirtley (1994). All information about sabellarids was brought together, the taxonomy was reorganized based on the literature and the examination of a great number of species Accepted by J. Williams: 7 Aug. 2014; published: 4 Nov. 2014

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was accomplished. This work was considered a major revision of sabellariids, two subfamilies were created (Sabellariinae and Lygmaniinae (spelled Lygdamiinae therein)) and new genera and species were organized into taxonomic keys. The genus Sabellaria Savigny, 1818 presents a mid-ventral indentation on the anterior end of the opercular stalk and crown, which is divided into symmetrical halves. Members of the genus are differentiated from other sabellariine genera by having the opercular paleae organized into three conspicuous concentric rows and for being divided into symmetrical halves (Kirtley 1994; Nishi et al.2004; Santos et al. 2011; Capa et al.2012). Sabellaria is a cosmopolitan genus that usually inhabits shallow waters, but it has been found in 0–254 m (Kirtley 1994). Some species are known as reef-builders, capable of colonizing wide areas along the coast, where hydrodynamic conditions and sediment type support the aggregates. Sometimes individuals are found to be solitary, in variable kinds of substrata (Calline et al.1992; Kirtley 1994; Pandolfi et al.1998; Wood 1999; Nishi et al.2004). Presently, 37 species of Sabellaria are known world-wide (Kirtley 1994; Nishi & Kirtley 1999; Nishi & Kato 2002; Nishi et al.2004; Nishi et al.2010; Santos et al.2011). Of these only nine were reported for the Brazilian coast: S. alcocki (Gravier 1906); S. bella Grube 1870; S. bellis Hansen1882; S. corallinea Santos et al.2011; S. floridensis Hartman 1944; S. gracilis Hartman, 1944, S. nanella Chamberlin 1919; S. pectinata Fauvel 1913 and S. wilsoni Lana & Gruet 1989 (Santos et al.2011). The genus Lygdamis Savigny, 1818 presents a mid-ventral indentation on the anterior end of the opercular stalk and crown, which is divided into symmetrical halves that are completely free. Members of the genus are differentiated from other lygdamiine genera by having numerous straight and short opercular paleae without geniculae, organized into two conspicuous concentric rows and for being divided into symmetrical halves (Kirtley 1994). Species of Lygdamis are known to be solitary. Of the 17 known species, five species of Lygdamis were previously reported from the Atlantic Ocean: L. gilchristi (McIntoshi 1924); L. kirkegaardi Kirtley 1994; L. muratus (Allen 1904); L. rayrobertsi Kirtley 1994; and L. wirtzi (Nishi & Núñez1999). Herein we describe for the first time an aggregate of Sabellaria species along the Brazilian coast, with S. nanella and S. wilsoni. We also describe a new species of Sabellaria and provide the first occurrence of Lygdamis rayrobertsi for the South Atlantic Ocean.

MATERIAL AND METHODS Study area The Rio Grande do Norte continental shelf extends 30–40 km from the shore; the slope begins at 50–60 m depth and the topology is irregular, with an average gradient of 1:1,000 (Testa 1997). A series of submerged projections and banks exist alongside the shelf, constituting a very rough relief. The most common facies are composed mainly of calcareous algae, such as Lithothamnion Heydr, Lithophyllum Phil, and Halimeda J.V. Lamour (Mabesoone & Coutinho 1970). Siliciclastic sands are present in shallow waters near the coast, while carbonate sands are present offshore; mud occurs at the mouths of the rivers and in the channels on the shelf (Vital et al. 2005). Sampling strategy Sampling was carried out on the Rio Grande do Norte continental shelf during four campaigns: July 21–30, 2002 (Campaign 1), May 12–30, 2003 (Campaign 2), November 14–23, 2003 (Campaign 3), and May 17–31, 2004 (Campaign 4), by the CENPES/PETROBRAS Project "Potiguar Basin Environmental Assessment". The Potiguar Basin is an area of oil and gas exploitation where the research center (CENPES) of the Brazilian oil company PETROBRAS is carrying out an extensive program of environmental assessment. Benthic samples were collected at up to 43 stations along nine transects perpendicular to the Rio Grande do Norte coast (l4º–5º S36º–37º W), at depths ranging from 2 to 100 m (Fig. 1). Samples were collected with Corer and Van-Van dredging in 3–77.5 meters deep between May 21 and 27, 2004. After the extraction from their tubes, specimens were fixed in 4% formalin without anesthesia and were transferred to 70% ethanol. Specimen study Specimens were examined using a Zeiss stereoscopic dissecting microscope. For a better analysis of species, some specimens were prepared on semi-permanent slides with pure glycerin or using a permanent solution Gray and

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Wess (PVA) (Humason 1979). In the sections prepared with Gray and Wess the muscle excess was removed. After that, sections were mounted on Gray and Wess in a slide for permanent conservation. Drawings of the notopodial and neuropodial chaetae were made using of a camera lucida with an OLYMPUS BX41compound microscope. Specimens were photographed using a Nikon G-7 digital camera and later edited with Adobe Photoshop CS software. For examination using SEM, specimens were critical point dried, covered with 50 nm of gold, and examined under SEM JEOL JSM-6390LV at the Laboratório de Microscopia Eletrônica, Departamento de Invertebrados, Museu Nacional, Rio de Janeiro (MN/UFRJ). The nomenclature adopted follows Kirtley (1994), Lechapt and Kirtley (1998), and Santos et al. (2011). Type material was deposited in the collection Poliquetas (POLY), Coleção de Invertebrados Paulo Young (CIPY), Departamento de Sistemática e Ecologia, Universidade Federal da Paraíba, Coleção de Poliquetas da Universidade Federal de Pernambuco (POLY-UFPE), Museu de Zoologia, Universidade Federal de São Paulo (MZUSP), Museu de Zoologia, Universidade Estadual de Campinas (ZUEC), and Museu Nacional do Rio de Janeiro (MN/UFRJ).

FIGURE 1. Map of the Guamaré Bay and sampling stations.

RESULTS Systematics Key to species of Sabellariidae found in Guamaré Bay 1. 2. 3. -

Three parathoracic segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Four parathoracic segments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lygdamis rayrobertsi Kirtley 1994 Obovate medium and inner paleae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sabellaria guamare sp. n. Straight, pilose or denticulate outer paleae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Outer paleae with 3–6 dial teeth on each side of spatulate medium plume, middle paleae short, quadrangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sabellaria nanella Chamberlin 1919 Outer paleae with 3–4 dial teeth on each side of spatulate and pilose medium plume, middle paleae long, narrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sabellaria wilsoni Lana & Gruet 1989

Family SABELLARIIDAE Johnston, 1865 Subfamily Lygdamiinae Kirtley, 1994 Type genus. Lygdamis Savigny, 1818

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Diagnosis. Sabellariid polychaetes with four parathoracic segments. Discussion. Kirtley (1994) created this subfamily to group genera that show four parathoracic segments. The original spelling (Lygmaniinae, lapsus calami) is corrected herein to Lygdamiinae, because, according to the Code of Zoological Nomenclature, one of the family-group names must be based on the type genus Lygdamis.

Genus Lygdamis Savigny, 1818 Lygdamis Kirtley, 1994: 116. Type species. Lygdamis indicus Kinberg, 1867, by monotypy. Type locality. Bangka Straits, Java, Indonesia. Diagnosis. Operculum with separated lobes, distal end sloped posteriorly, opercular paleae straight, smooth, outer ones flat and inner ones cylindrical, spine-like (Capa et al. 2012). Description. Operculum longer than wide, lobes completely separated, distal end sloped posteriorly (oblique to longitudinal axis). Numerous opercular papillae around edge of lobes. Outer paleae in semicircles, straight, with flat blades, lateral and distal margins smooth. Inner opercular chaetae in one row along inner margin, opercular lobes with straight and cylindrical or slightly flattened blades. One pair of nuchal spines with bent tips (hooks) and without limbations. Palps similar in size to operculum. Median organ elongate at the dorsal junction of lobes of opercular stalk. Tentacular filaments compound (branching); buccal flaps absent. Neuropodia of segment 1 with conical cirrus, with or without capillary chaetae. Segment 2 with three triangular-shaped lobes between noto- and neuropodia, in some species rounded and small. Four parathoracic segments with notochaetae consisting of lanceolate and capillary chaetae and neurochaetae similar in shape but smaller. Branchiae from segment 2 to midabdominal segments (Kirtley 1994; Hutchings et al. 2012).

Lygdamis rayrobertsi Kirtley, 1994 Figure 2 A–B Lygdamis rayrobertsi Kirtley, 1994: 134, figs. 7.15.1–7.15.3. Lygdamis indicus.—Uebelacker, 1984: 49-5, fig. 49-2 (a–h).

Material examined. Total: 1 specimen. State of Rio Grande do Norte, Guamaré City: Project Caracterização Ambiental da Bacia Potiguar e Monitoramento Ambiental dos Emissários Submarinos do Pólo Industrial de Guamaré. St. BPOT24DG (05º3.561’0’’S 036º28.51’0’’W)[POLY 00291]: 1 complete specimen, 8 mm long, on 21/05/2004, 77.5 m. Additional material examined. Lygdamis indicus (Kinberg 1887). Mozambique, Mozambique Channel, 24º40’S 35º38’E [LACM-AHF POLY 2472]: 1 complete specimen collected on crustacean carapaces in coarse shelly sand, 18 August 1964, 360 m depth. Lygdamis indicus. International Indian Ocean Expedition, cruise 7, R/V ANTON BRUUN, St. AB 370G. Republic of Palau, Koror State, Malakal Island, entrance to Malakal Harbor, 35 m, Coral Reef Research Foundation, 963 L.H. [LACM-AHF POLY 2473]: 1 complete spec. collected in 1987. Lygdamis laevispinis Grube, 1870. St. AB 3706 [LACM-AHF POLY]: 2 complete specs. Lygdamis splendidus Lechapt & Kirtley, 1998. Pacific Ocean, South of New Caledonia, Biocal St. 66º (24’S55º18’E) [LACM-AHF POLY]: 1 complete spec. collected on 03/09/1985. Diagnosis. 28 pairs of long outer paleae, straight, with anterior end a curved tapered apex; 12 pairs of long inner paleae, straight, with anterior end a tapered apex. Body poorly pigmented; few brown eyespots along the body. Diagnosis. 28 pairs of long outer paleae, straight, with anterior end a curved tapered apex; 12 pairs of long inner paleae, straight, with anterior end a tapered apex. Body poorly pigmented; few brown eyespots along the body.

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FIGURE 2. Lygdamis rayrobertsi Kirtley, 1994 (POLY 00291): (A) Body in ventro-lateral view; (B) Body in dorso-lateral view. Scale bars: A–B, 1 mm.

Remarks. Five species of Lygdamis were previously reported from the Atlantic Ocean: L. gilchristi (McIntoshi, 1922), L. kirkegaard Kirtley, 1994, L. muratus (Allen, 1904), L. rayrobertsi, and L. wirtzi Nishi & Núñez, 1999. Of these species, only L. rayrobertsi and L. wirtzi possess curved outer paleae. Both species are very similar but differ in the form of the inner palea, which is straight, with distal end terminating in a tapered apex in L. rayrobertsi, while it is straight, with anterior end terminating in a curved tapered apex in L. wirtzi. Lygdamis rayrobertsi was previously recorded in Caribbean waters while L. wirtzi inhabits European waters.

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Lygdamis rayrobertsi is characterized by the form of inner and outer paleae, but the number of pairs of paleae were not indicated in the original description. The specimens of L. rayrobertsi identified herein are very similar to the previously known material, in spite of their geographical distances. The examined specimens represent the first reported occurrence of the species along the Brazilian coast. Other species of Lygdamis that were recorded in the Atlantic Ocean (L. gilchristi, L. kirkegaardi, and L. muratus), have long, straight outer and inner paleae with anterior end terminating in a tapered apex. These characters suggest that they could represent a single species. This problem was commented on by Kirtley (1994) who did not synonymize L. gilchristi and L. muratus because he had not examined the type material. At the same time, Kirtley (1994) described L. kirkergaardi from the North Atlantic, which has the same distribution as L. muratus. These two species also have a similar morphology (number of outer and inner paleae, and form of paleae) and need to be revised in the future. In the original descriptions of L. muratus and L. gilchristi, and in other subsequent publications, the body was not illustrated, neither were the form of the lateral palps or of the median organ. Only the form of the outer and inner paleae were figured. Lygdamis rayrobertsi inhabits sand and rocky substrates, or any hard surface such as rocks and crustacean carapaces belonging to Panulirus argus Latreille, 1804, or polychaete tubes (Kirtley 1994). The species has been found in depths of 2–73 meters and was collected at 77,5 meters in Guamaré Bay. Lygdamis rayrobertsi was originally described from the Gulf of Mexico, along the coast of Florida in Palm Beach (Kirtley 1994), and now is reported for Brazil.

Subfamily SABELLARIINAE Kirtley, 1994 Genus Sabellaria Lamarck, 1812 Sabellaria Kirtley, 1994: 45–46; Nishi et al. 2010: 7.

Type species. Sabella alveolata Linnaeus, 1767, by monotypy, collected in France. Emended diagnosis. Operculum completely divided into two symmetrical lobes, inner paleae in three symmetrical, semicircular rows (with mid and inner paleae), and three parathoracic segments (Santos et al. 2011). Emended description. Operculum length similar to maximum width, completely divided into two symmetrical lobes; distal disc flat, perpendicular to longitudinal axis. Numerous conical opercular papillae around operculum. Outer paleae numerous, in semicircles; geniculate, with flat blades, lateral edge smooth, and distal margin smooth or denticulate, sometimes with a midline plume. Inner opercular paleae of various shapes, in two concentric rows. Middle paleae strongly geniculate, blades excavated and margins smooth, pointing outwards; innermost paleae strongly geniculate, with short concave blades and smooth margins, directed inwards. 3–6 pairs of straight nuchal spines often present. Compound (branching) tentacular filaments in series of rows; buccal flaps absent. Palps similar or shorter than operculum. Median organ at dorsal junction of lobes of opercular stalk conspicuous in some species and small or absent in others, with eyespots on lateral margins. Neuropodia of segment 1 with one pair of conical cirri and capillary chaetae. Segment 2 with one triangular-shaped lobe on both sides of building organ. Thoracic branchiae present. Three parathoracic segments. Parathoracic notochaetae lanceolate and capillaries alternating; neurochaetae similar in shape but smaller (Capa et al. 2012). Remarks. Nishi et al. (2010) and Santos et al. (2011) have recently described new species of Sabellaria and provided a table with chaetal characteristics of the 36 species from around the world. They highlight the variability of certain features within the genus, such as the different geometry and superficial ornamentation of the paleal thecae (especially in the middle row), the presence or absence of a plume or median tooth in the outer paleae, and the presence or absence of nuchal spines. According to Capa et al. (2012) the large intraspecific variation regarding these features, together with the inconsistency of the presence or absence of a median organ among Sabellaria species (Kirtley, 1994), compromise the assessment of the monophyly of Sabellaria. Therefore, the species analyzed before were not examined observing the form and position of the medium organ. The presence of a medium organ was also reported for some other genera of Sabellariidae and for the genus Megalomma (family Sabellidae) (Tovar and Salazar-Vallejo 2008).

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According to Capa et al. (2012) all species of Sabellaria share the presence of an operculum completely divided into two symmetrical lobes, inner paleae being arranged into two symmetrical, semicircular rows (with mid and inner paleae), and with three parathoracic segments. Branchiae on second (thoracic) segment are present (contrary to diagnosis of Nishi et al. 2010). Sabellaria was supposed to share the arrangement of the inner paleae in two concentric rows with Phragmatopoma and Neosabellaria. However, we observed that species of Sabellaria have three concentric rows of inner paleae, while Phragmatopoma has the inner paleae under the middle paleae.

Sabellaria nanella Chamberlin, 1919 Figures 3A–C Sabellaria nanella Chamberlin 1919: 261, plate 2, figures 5–7; Johansson 1927: 98; Hartman 1938: 16, figures 8–10; 1944: 340, plate 30, figures 18–20; Rullier & Amoureux 1979: 188; Lana & Gruet 1989: 243; Kirtley 1994: 71, figures 4.26.1–4. 26. 2; Lana & Bremec 1994: 216, figures 1–2; Santos et al. 2011: 6, figure 4 (a–c); Lomonaco et al. 2011: figure 1.

Material examined. Total: 10 specimens. State of Rio Grande do Norte, Guamaré City: Project Caracterização Ambiental da Bacia Potiguar e Monitoramento Ambiental dos Emissários Submarinos do Polo Industrial de Guamaré. St. BPOT20DG (05º0’0’’S 036º27’0’’W) [POLY 00292]: 1 complete spec. 21/05/2004, 4.3 m, 4 mm long. St. BPOT24DG (04º40’0’’S 036º28.57’0’’W) POLY 00293]: 1 complete spec. 21/05/2004, 4.3 m, 3 mm long. St. BPOT8DG (04º25’0’’S 036º23’0’’W) [POLY 00294]: 1 complete spec. 21/05/2004, 5 m, 3 mm long. St. BPOT10DG (05º3.1’0’’S 036º29.77’0’’W) [POLY 00295]: 1 complete spec. 27/05/2004, 5 m, 3 mm long. St. BPOT25DG (05º20’0’’S 036º40’0’’W) [POLY 00296]: 1 complete spec. 21/05/2004, 5 meters of depth, 3 mm long. St. E06DG (05º3.561’0’’S 036º28.57’0’’W) [POLY 00297]: 1 complete spec. 21/05/2004, 3 mm long. St. E10DG (05º3.561’0’’S 036º24.5’0’’W) [POLY 00298]: 1 complete spec. 21/05/2004, 6 m, 3 mm long. St. E15DG (05º3.561’0’’S 036º28.57’0’’W) [POLY 00299]: 1 complete spec. 21/05/2004, 10 m, 3 mm long. St. E22DG (05º0’0’’S 036º23’0’’W) [POLY 00300]: 1 complete spec. 21/05/2004, 12 m, 3 mm long. St. E23DG (05º0’0’’S 036º23.57’0’’W) [POLY 00301]: 1 complete spec. 21/05/2004, 12.3 mm long. Additional material examined. Brazil, Paraíba State. Total: 2 specimens. St. B10 (7°01´02´´S34º47´55´´W) 1 complete spec. 03/2007 [CIPY-POLY 1025].St. B10 (7°01´02´´S34º47´55´´W) 1 complete spec. 03/2007 [CIPYPOLY 1026]. Diagnosis. Outer paleae semi-cylindrical or chaliciform, concave, geniculate, with series of small lateral teeth on thecal margin; thecal margin curved towards interior of theca or straight, with one-two points (Figure 3A). Middle paleae short, strongly geniculate, square-shaped, with slight central depression (Figure 3B), inner series of paleae strongly geniculate, elongate, with anterior ends pointed (Figure 3C). Remarks. Sabellaria nanella is an uncommon species because it is the only taxon that has this kind of organization of the opercular crown: outer palea chaliciform, medium paleae quadrangular, inner paleae geniculate. In 12 specimens examined no morphological variations were observed. The specimens examined have six pairs of nuchal hooks positioned after a short digitiform medium organ, as described in Santos et al.(2011). Kirtley (1994) considered the occurrence of S. nanella in Brazil to be a result of bioinvasion, possibly resulting from ballast water. In the majority of cases, the occurrences of S. nanella along the Brazilian coast were associated with aggregates of Phragmatopoma caudata Morch, 1883. The first report of this species along the Brazilian coast was in the State of Paraíba by Young (1989), in crevices of the coral Mussismilia hispida (Verrill 1901) from Quebra Quilha Reef, at Tambaú Beach, João Pessoa, Paraíba. In the present study the specimens were found from 4, 5–12 m. Sabellaria nanella was originally described from the Indo-West Pacific, but has also been reported from the eastern Pacific, from Ecuador (Kirtley 1994), and along the Atlantic Ocean, from the coast of Ceará (Kirtley 1994), Paraíba (Santos et al.2011; Lomonaco et al.2011), Bahia (Lana & Bremec 1994), Espírito Santo, Rio de Janeiro (Rullier & Amoureux 1979; Lana & Bremec 1994), São Paulo, and Uruguay (Lana & Bremec 1994).

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FIGURE 3. Sabellaria nanella Chamberlin, 1979 (POLY 292): (A) outer palea; (B) middle palea; (C) inner palea. Scale bars: A–C, 0.1 mm.

Sabellaria wilsoni Lana & Gruet, 1989 Figure 4A–C Saballaria wilsoni Lana & Gruet 1989: 239, figures 1–21; Kirtley 1994: 80, figure4.33; Lana & Bremec 1994: 216, figures 1–2; Santos et al. 2011: 9, fig. 6 (a–c); Lomonaco et al. 2011: fig. 1.

Material examined. Total: Four specimens. State of Rio Grande do Norte, Guamaré city: Projects Caracterização Ambiental da Bacia Potiguar and Monitoramento Ambiental dos Emissários Submarinos do Pólo Industrial de Guamaré. St. BPO19DG (05º3.561’0’’S036º28.57’0’’W) [POLY 00302]: 1 complete spec. 27/05/2004, 4.3 m, 6 mm long. St. BPOT20DG (05º0’0’’S 036º27’0’’W) [POLY 00303]: 1 complete spec. 21/05/2004, 5 m, 7 mm long. Est. BPOT24DG (05º20’0’’S 036º40’0’’W) [POLY 00304]: 1 complete spec. 21/05/2004, 5 m, 8 mm long. St. E07DG (05º3’1,74’’S 036º28.57’0’’W) [POLY 00305]: 1 complete spec. 21/05/2004, 6.3 m, 8 mm long. Additional material examined. Sabellaria wilsoni Lana & Gruet (1989). Type series: Holotype: Paraná State, Ponta Grossa, Paranaguá Bay, 25°25’06”S48°28’00”W, collected on 24/05/1982. 10 m [MCBM BPO 241]. Paratype: Ponta Grossa, Paranaguá Bay, 25°25’06”S48°28’00”W, 15 specs, 24/05/1982. 10 m [MCBM BPO 242]. Paratype: Ponta do Maciel, Paranaguá Bay, 25°33’S48°25’W, 16.5 m, 1 spec. 28/05/1982 [MCBM BPO 243]. Paratype: Ponta do Maciel, Paranaguá Bay: 25°33’S48°25’W. 1 spec. 05/06/1986, 16.5 m [MCBM BPO 244]. Paratype: Ponta da Pita, Antonina Bay, 25°26’30”48°31’W, 1 spec.,03/07/1986, 6 m [MCBM BPO 245]. Paraíba State: Total: 2 specs, 03/2009. 7 St. B10 (7°01´02´´S34º47´55´´W) [CIPY-POLY 1027]. 7°01´02´´S34º47´55´´W (Station B10), 1 spec. [CIPY-POLY 1028].

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FIGURE 4. Sabellaria wilsoni Lana & Gruet, 1989 (POLY 302): (A) outer palea; (B) middle palea; (C) inner palea. Scale bars: A–C, 0.1 mm.

Diagnosis: Outer paleae with flat, oblong, thecate blades; teeth and medial plume on distal margin of blades penicillate; 3–4 distal teeth on each side of dominant tooth, progressively smaller toward outer margins (Figure 4A). Middle paleae organized in 9–12 pairs on each side; strongly geniculate, with elongated blades distally blunted, curved toward inner series of opercular crown (Figure 4B). Inner series of paleae alveolate, strongly geniculate on proximal upper surface of blades, with thecal bands weakly excavate, slanting upward from horizontal plane; blade thicker and wider toward base, ending in bluntly rounded point; distal dentition on margin or, in some specimens, a series of sharp hooked tips (Figure 4C). Remarks.The material examined here (Figures 4A–C) agrees with the descriptions of Lana & Gruet (1989) and Santos et al. (2011). Sabellaria wilsoni is known from mixohaline and polyhaline enviroments (estuaries and

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shallow habitats on the continental shelf), living in sandy or muddy sediments (sometimes with shells). The specimens were found at 15 m, similar to the type series collected at 10 and 16,5 m. Recently, Santos et al. (2011) found S. wilsoni living near other sabellariid species, as in our new material. Sabellaria wilsoni was originally described from the coast of Paraná. The species has been reported only from the Atlantic Ocean, from French Guyana to Argentina, and possibly from the Gulf of Mexico (Lana & Gruet 1989; Bremec & Lana 1994; Santos et al.2011; Lomonaco et al. 2011).

Sabellaria guamare sp. n. Figures 5A–F, 6A–F, 7A–M and 8A–D, Tables 1 and 2. Material examined. Total: 22specimens. Type series: State of Rio Grande do Norte, Guamaré City: Projects Caracterização Ambiental da Bacia Potiguar and Monitoramento Ambiental dos Emissários Submarinos do Pólo Industrial de Guamaré. St. BPOT24DG (05º3.561’0’’S036º28.57’0’’W), 77,6 meters of depth. Holotype [CIPYPOLY 1689]: 1 complete spec.21/05/2004, 55 mm long. Paratypes: Paratype 1, [CIPY-POLY 1690]: 1 complete spec.21/05/2004 in a good state of preservation. Paratype 2 [CIPY-POLY 1691]: 1complete spec.21/05/2004. Paratype 4 [CIPY-POLY 1692]: 1incomplete spec.21/05/2004. Paratype 5 [POLY 1693]: 1 incompletespec.21/05/ 2004. Paratype 6, [CIPY-POLY 1694]: 1 incomplete spec.21/05/2004. Paratype 7 [POLY 1695]: 1 incomplete spec.21/05/2004. Paratype 8 [POLY 1696]: 21/05/2004. Paratype 9[CIPY-POLY 1697]: 21/05/2004. Paratype 10 [CIPY-POLY 1698]: 1 incomplete spec. 21/05/2004. Paratype 11 [xxx-POLY 1699]:1 incomplete spec. 21/05/ 2004. Paratype 12 [CIPY-POLY 1700]: 1incomplete spec. 21/05/2004. Paratype 13 [CIPY-POLY 1701]: 1incomplete spec.21/05/2004. Paratype 14 [CIPY-POLY 1702]: 1incomplete spec. 21/05/2004. Paratype 15 [POLY 1703]: 1 incompletespec.21/05/2004. Paratype 16 [CIPY--POLY 1704]: 1 incomplete spec.21/05/2004. Paratype 17 [CIPY-POLY 1705]: 1 incomplete spec.21/05/2004. Paratype 18 [CIPY-POLY 1706]: 1 incomplete spec.21/05/ 2004. Paratype 19 St. BPOT25DG. (05º20’0’’S 036º40’0’’W)[MN-POLY 1707]: 1 incomplete spec.19/05/2004, 5 m. Paratype 20 St. BPOT25DG (05º20’0’’S 036º40’0’’W). [MN-POLY 1708]: 1 incomplete spec.19/05/2004, 5 m. Paratype 21 St. E16DG (05º3.561’0’’S 036º29’0’’W)[MN-POLY 1709]: 1 complete spec.21/05/2004, 10 m, 3 mm long. Stubs for SEM. Paratype 3 [CIPY-POLY 1710]: 1 incomplete spec.21/05/2004.Paratype 4 [CIPY-POLY 1711]: 1 incomplete spec.21/05/2004.Paratype 12 [CIPY-POLY 1712]: 1 incomplete spec.21/05/2004.Paratype 13 [CIPYPOLY 1713]: 1 incomplete spec.21/05/2004. Diagnosis. This is a unique species of Sabellaria in possessing a prolonged form of the medium cirrus. This structure shows 3 small prolongations with extremity gauged, and small black eyespots agglomerated in some parts of the medium organ. The organization of the opercular crown is also exclusive. The opercular crown is organized into: outer paleae with small teeth (five to six along each distal margin), distal teeth present, with 2–3 distal teeth on each side of dominant tooth, progressively smaller toward outer margins on either side of dominant tooth. Middle paleae alveolate, strongly geniculate, with falciform blades lightly blunted, the inner portion positioned toward inner series of opercular crown; weakly excavate inner face of thecal bands, at lower portion; one to two large teeth on each side of dominant tooth. Description. Holotype. Complete specimen in good conditions with 45 chaetigers, total length 55 mm, length of peduncle 3 mm, width of peduncle 3 mm, length of thorax 2.5 mm, width of thorax 2 mm, and length of cauda 3 mm (Figures 5A, B; 6A, B and 7K, M). Anterior end of opercular stalk and crown with light mid-ventral indentation, divided into symmetrical halves (Figures 5C and 6C). Opercular paleae yellow-gold with vitreous luster organized in rows surrounding prostomium and peristomium in three apparent concentric rows differing in shapes and length. Outer paleae forming 28–30 pairs on each side, with flattened, oblong, thecate blades; plume with three pairs of teeth on side of dominant tooth (Figures 5C, D and 7A). Medial teeth denticulate. Medial plume on distal margin of blades with serrate distal teeth, with1 distal tooth on each side of dominant tooth; these progressively smaller toward outer margins on either side of dominant tooth (Figure 8A, B). Blades bent slightly outward from longitudinal axis of shaft; distal lateral margins directed posteriorly; inner surface of blade with many fine closely spaced striations (Figure 8C, D). Each striation consists of series of small teeth (occurring along a limited region).Twelve pairs of middle paleae on each side of the symmetrical halves. Middle paleae alveolate, strongly geniculate, with falciform blades slightly blunt, curved 90° (Figure 7B), with axis towards inner series on

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FIGURE 5. Sabellaria guamare n. sp., holotype (CIPY-POLY 1689) (A) body in ventro-lateral; (B) body in dorso-lateral view; (C) opercular peduncle; (D) detail of opercular peduncle; (E) abdominal region showing the stomach; (F)posterior region. Legends: BR, branchia; BO, building organ, FT, compound feeding tentacles; NH, Nuchal hooks; OP, opercular peduncle; PA, parathoracic region; PI, pygidium; ST, stomach; TH, thorax; TO, tori;. Scale bars, A–C, E and F, 5 mm; D, 1 mm.

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FIGURE 6. Sabellaria guamare n. sp., holotype (CIPY-POLY 1689) (A) body in ventro-lateral view; (B) body in dorsal-lateral view; (C) opercular peduncle. Paratype, (CIPY-POLY 1690) (D) ventral view without tentacles; (E) medium organ in peduncle; (F) detail of medium organ. Legends: ABS, abdominal segment; BR, branchia; BO, building organ; FT, compound feeding tentacles; MO, medium organ; VP, ventral pads. Scale bars: A–C, 5 mm; D and E, 2 mm; F, 0.5 mm.

opercular crown; weakly excavate innerface of thecal bands; with one to two divisions in apical region, and with one to two distal teeth in each side of dominant tooth and series of tiny extensions. Distal ends of paleae almost erect, tapering to distal tips, with weak excavation on inner face of thecal bands (Figure 5). Paleae of inner series strongly geniculate; proximal upper teeth along margin with thecal bands weakly excavate (Figure 7C), slanting

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FIGURE 7. Sabellaria guamare n. sp., holotype (CIPY-POLY 1689) (A) outer palea; (B) medium paleae; (C)inner paleae; (D) nuchal hooks; (E) thoracic chaeta; (F) lanceolate notopodial pharathoracic chaeta; (G) spiralete notopodial parathoraic chaeta; (H) lanceolate neuropodial pharathoraic chaeta; (I) spiralete neuropodial pharathoraic chaeta; (J) neuropodial abdominal chaeta; (K) dorso-lateral view; (L) nototopodial abdominal chaeta (M) ventro-lateral view. Scale bars: A, 0.1 mm; B, 0.1 mm; C, 0.1 mm; D, 0.3 mm; E, 0.1 mm; F, 0.3 mm; G, 0.3mm; H, 0.03 mm; I, 0.03 mm; J, 0.1 mm; L, 0.03mm; K, 5 mm; M, 5 mm. Legends: BR, branchia; CO, crown; CP, paired cirrus; O, eyespots; BO, building organ; PE, peduncle; PG, pygidium; PR, parapodium; PT, pharathoracic region; TO, thoracic region; UN, uncini.

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upward from horizontal plane; blade thicker and wider toward base, terminating distally in bluntly rounded point, series of distal teeth along margin, occurring along a limited region (Figure 7C, D). Opercular peduncle with 3 pairs of dorsal (nuchal) spines on either side of mid-line (Figure 7D). Anterior margin of peduncle with flattened big spaced palps in series attached to peristomium (Figures 5A–C and 6C). Peduncular bases with a series of 6 longitudinal rows of compound feeding tentacles (12–15 in transverse rows) on ventral and dorsal margins of buccal cavity attached to peristomium (Figure 5C, D). Prostomium fused with peristomium and limited to a region situated in front of the mouth (Figure 5C), bearing pair of grooved prehensile tentacles with conspicuous medium cirrus, with prolonged form positioned in front of grooved buccal palps; small black eyespots agglomerated in some parts of the medium organ (Figure 6, F). Peristomium lips beginning anterior to the mouth, ending at the base of the compound feeding tentacles (Figure 5D). Thoracic region consists of a U-shaped building organ, and two segments with spiraled capillary chaetae (Figure 7E). Thoracic segments with neuropodium only, without notopodium, and conical palps located on each side of opercular peduncle (ure 5A, B, and E). Cirri of first thoracic segment triangular, bent toward ventral side of body, quite similar to buccal cirri. Cirri of second thoracic segment triangular, elongate, bent toward dorsal side of body, on each side of the mid-ventral indentation of operculum. Parathoracic region with 3 segments and small biramous parapodia (Figure5A, B, and E). Notopodium and neuropodium with shaves of chaetae with lanceolate tips (“oar-shaped”) (Figure 7F, H) and thin spiraled capillary chaetae (Figure 7G, I) with slightly curved tips positioned between chaetae with lanceolate tips. Abdomen with 18–20 segments and tori in numerous segments (Figure 6C); dorsal digitiform branchiae in the five anterior abdominal segments, being reduced in size at segment 15 and disappearing in last segment (Figure 5F) tori with chaetae positioned in a single row. Notopodial uncini bipectinate, with 6 rows of superposed teeth, neurochaetal conical cirri arising from base of bundle in first abdominal segment; cirri diminishing in size posteriorly and disappearing on posterior-most segments (Figures 7J,L). Caudal region septate, composed of numerous achaetous segments. Cauda bent towards ventrum (Figures 5A, B and 6C). 16–20 anal septations. Eyespots brownish to reddish present in each expansion of the filaments of compound feeding tentacles (or branchial filaments). Black pigmented eyespots in opercular peduncle along inner face of serial conic palps, extended to midventral indentation; peristomium between compound feeding tentacles, medium cirrus in base of upper edge; parathoracic region with a large stain composed of numerous eyespots in ventral side of third setiger; few eyespots in caudal region. Red glandular spots on abdominal region, on paired cirri, and on neuropodial pads of chaetigers. Red eyespots form “C”-shaped pads in first abdominal chaetigers. Eyespots increase in number and concentration beyond abdominal chaetiger 10. Variations.The specimens examined present great variations in the number of eyespots. Some specimens do not have the brownish to reddish eyespots, or have them in lower concentrations than in other samples. The specimens found at the depths of 5–7 m are generally more pigmented than the specimens collected at 7–77 meters. These differences could represent sexual dimorphisms because we observed that specimens collected in 77 meters have a lot of black eyespots and others only a few. Lomonaco et al. (2011) observed that males of Sabellaria nanella and S. wilsoni are more pigmented than females. The number of abdominal segments does not vary incomplete animals as shown below in Table 1. Type locality. Brazil: Rio Grande do Norte State: Guamaré Bay. Etymology.The species is named after Guamaré Bay, the type locality of this species. Remarks. Sabellaria guamare n. sp. belongs to the group of the honeycombed species of Nishi et al. (2010) (Table 2). This group of species of Sabellaria possesses three kinds of paleae. These species have some distal teeth that are paired in relation to the central teeth. The species of this group always have honeycomb bed medium paleae, light or deeply excavated, that could be bent outwards. The inner paleae are honeycombed or expanded anteriorly (Figure 8 C), extremely geniculate, and could be reflected outwards or sometimes could be curved into an angle of 45°. Among the alveolate species, only S. bellis Hansen 1882, S. guamare n. sp., and S. moorei (Monro 1933), possess the outer paleae with 2–3 big teeth on each side of the dominant teeth. Only S. guamare n. sp. shows a rotated medium honeycombed palea. Therefore, the dominant teeth of S. guamare n. sp. are longer and have the distal end with small teeth of similar size. In S. moorei and S. bellis they are triangular and decreasing in size. S. guamare sp. n. constructs solitary tubes made by mud sand on sand substrata, but some specimens were found constructing their tubes around the tubes of other polychaetes, such as Loimia sp., S. wilsoni Lana and Gruet, 1989, and S. nanela Chamberlin, 1919. S. guamare sp. n. inhabits depths from three to 77 meters. This species lives

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2.5x2.5

2.0x2.0

3.0x2.5

3.0x2.0

2.5x1.5

1.5x1.0

5.0x3.0

3.0x2.0

2.0x1.0

2.0x1.0

2.0x1.0

2.0x1.0

3.0x2.0

Paratype 10 2.0

Paratype 11 2.5

Paratype 12 -

Paratype 13 -

Paratype 14 1.0

Paratype 15 3.0

Paratype 16 2.0

Paratype 17 1.0

Paratype 18 1.0

Paratype 19 1.0

Paratype 20 1.0

Paratype 21 2.0

3.0x2.5

Paratype 6 2.5

Paratype 9 2.5

4.0x2.0

Paratype 5 2.0

2.0x2.0

4.0x2.0

Paratype 4 2.0

3.0x1.5

3.0x1.5

Paratype 3 1.5

Paratype 8 1.5

2.5x2.0

Paratype 2 2.0

Paratype 7 -

2.5x2.5

2.5

Paratype1

5.0x3.0

3.0

Holotype

2.5x2.5

1.0x1.0

1.0x1.0

1.0x1.0

1.0x1.0

2.0x1.5

2.5x2.5

1.5x1.0

-

1.5x2.0

2.0x2.0

1.0x2.0

1.5x2.0

2.0x1.5

1.5x1.5

1.5x2.0

2.0x1.5

1.0x1.0

1.5x1.5

2.0x1.5

2.0x2.0

2.5x2.0

(mm))

2.0

1.0

1.0

1.0

1.0

1.5

2.5

1.0

-

2.0

2.0

2.0

2.0

1.5

1.5

2.0

1.5

1.0

1.5

1.5

2.5

2.0

(mm)

1

9

7

24

24

5

4

23

17

4

40

40

24

30

30

-

-

40

40

abdominal segments

Peduncle (length x width Parathoracic region (length x width Abdomen width Number of

operculum (mm) (mm))

Diameter of

TABLE 1. Variation in morphological features of S. guamare n. sp. type series.

8.0

8.0

8.0

19.0

19.0

7.0

6.0

20.0

-

-

13.0

10.0

20.0

8.00

19.0

-

17.0

18.0

-

-

21.0

55.0

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

complete spec.

complete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

incomplete spec.

complete spec.

complete spec.

Body length (mm) Additional data

in estuarine areas, and is the only gregarious sabellariid that doesn’t use any hard substrata to construct their sand tubes. This species is known only from the type locality.

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TABLE 2. Alveolate species of Sabellaria that have three kinds of paleae. Species

Nuchal spine

Outer paleae

Middle paleae

Inner paleae

Type locality

Distribution

Reference

Europe, Kirtley, Mediterranean, 1994; This Western Africa study

S. alveolata present (Linnaeus, 1767)

4–6 distal deep excavation, alveolate, teeth lateral to concave, geniculate, and a plume reflected outward concave

British Isles

S. bellis Hansen, 1882

present

2–3 teeth in each side of plume

geniculate, blade thicker, tip rounded

Rio de Janeiro, Brazil to Brazil Argentina

Kirtley, 1994; This study

S. fissendis Grube, 1870

not 3–4 distal alveolate, observed teeth lateral to light excavation, a plume concave

alveolate, geniculate, vertically pointed

Chile

Chile

Kirtley, 1994

geniculate blade thicker, tip rounded

Guamaré Bay

Guamaré Bay

This study

Andaman Sea

Kirtley, 1994

alveolate, light excavation, concave

S guamarei sp. n. present

2–3 teeth in each side of plume

alveolate, light excavation, concave

S. grueti Kirtley, 1994

present

3–4 distal alveolate, teeth lateral to concave a plume

geniculate, elongate

Burma, Andaman Sea

S. ishikawi (Okuda, 1938)

present

6–7 distal teeth

geniculate, elongate

San Thomé, West Africa Gulf of Guinea

Kirtley, 1994

S. javanica Auguener, 1934

present

a single apical alveolate, sigmoid tooth light excavation, concave

geniculate, elongate

Soerabaia

Java

Kirtley, 1994

S. moorei (Monro, 1933)

present

2–3 teeth in each side of plume

alveolate, light excavation, concave

geniculate blade thicker, tip rounded

Panama

Panama

Kirtley, 1994; This study

S. myriansis Parab & Gaikward, 1990

not 3–4 teeth in observed each side of plume, tip curved

alveolate, light excavation, concave

distal tip geniculate, elongate blade

West coast of India

West coast of India

Kirtley, 1994

alveolate, concave

DISCUSSION According to Kirtley (1994), the genera and species are defined based on the configuration in the opercular stalk and crown, the number and kinds of paleae of the opercular crown, differences in the geometry and superficial ornamentation of the thecae and paleae, presence of nuchal hooks or spines, presence or absence of a median organ (or aboral cirrus), form of the oral feeding tentacles, shape and relative length of the prostomial tentacles, shape and size of the building organ, shape and number of cirri, details of associated chaetal bundles on the first and second chaetigerous segments, number of parathoracic segments, and presence or absence of dorsal branchiae on some abdominal segments. This work shows that the paleae of some taxa are composed of series of small teeth, and Kirtley named these structures tiny extensions (Figure 8A–D). These tiny extensions should be observed for other sabellariids as well. The major illustration provided in the revision proposed by Kirtley (1994) and subsequent articles shows a majority of paleae with structures described as lines, tiny extensions or striations. Our results show that the paleae of all species of Sabellaria examined in this study, as well as species described by Nishi et al. (2010) and Santos et al. (2011), always have a series of teeth and the palea of each species has its own form. Consequently, if Kirtley (1994) and other authors had examined and compared the paleae of different taxa with SEM, they could have concluded that the striations are in fact a series of teeth and could be a new character used in the taxonomy of Sabellaria. The major abundance and diversity of species was concentrated around the depth of 77.5 meters. At this depth more than 25 samples of S. guamare sp. n. were found with few specimens of S. nanella and S. wilsoni, and one of L. rayrobertsi. At other depths the concentrations were lower. S. nanella was common in the material examined, this species occupying a great extension of the bay. The other species were found concentrated in small areas of Guamaré Bay.

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FIGURE 8. Sabellaria guamare n. sp.,paratype (CIPY-POLY 1710). (A) medium paleae; (B) detail of medium paleae; (C) inner series; (D) detail of inner paleae. Scale bars: A, 0.1 mm; B, 0.02 mm; C, 0.1 mm; D, 0.01 mm.

ACKNOWLEDGEMENTS We are grateful to Dr. José Roberto Botelho de Souza for loaning the material analyzed in this study, the Brazilian Petroleum Company (PETROBRAS), and to CENPES/PETROBRAS, for authorizing the publication of this material. The authors also thank Dr. Paulo da Cunha Lana, from MCEM-BPO, for the loan of the type series of Sabellaria wilsoni. We acknowledge Leslie Harris and Dr. Kirk Fitzhugh for help and for hosting the first author during a visit to Los Angeles and the Natural History Museum of Los Angeles Country. Programa de Pós Graduação em Ciências Biológicas, Federal University of Paraíba, Coordenação de Aperfeiçoamento de Pesquisa do Ensino Superior and Conselho Nacional de Pesquisa are acknowledged for financial support to the first author from 2004 to 2008. The Conselho Nacional de Desenvolvimento Científico e Tecnológico is also thanked for the productivity grant to the last author.

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