INTRODUCTION
Spionid polychaetes represent one of the most diversified and common polychaete families in themarine benthic communities along most continental shorelines (Blake Reference Blake1983, Reference Blake and Blake1996; Johnson, Reference Johnson, Uebelacker and Johnson1984). These animals can be found from shallow to deep waters. This family is well-distributed worldwide, and many species are cosmopolitan. In soft-bottom habitats they are often the most abundant, with one or two species dominating the benthic infaunal community. Scolelepis are typically sand dwellers, inhabiting intertidal sand beaches or subtidal flats (Johnson, Reference Johnson, Uebelacker and Johnson1984).
The genus Scolelepis was revised by Maciolek (Reference Maciolek1987), who established two subgenera, Scolelepis and Parascolelepis, based on the morphology of the hooded hooks. Subgenus Scolelepis De Blainville, 1828 includes species having unidentate to tridentate hooks, with a falcate or straight shaft, while subgenus Parascolelepis Maciolek, Reference Maciolek1987 includes species having multidentate hooks with a curved shaft. Delgado-Blas (Reference Delgado-Blas2006) described Scolelepis (S.) lighti and Scolelepis (S.) vosseae from the Gulf of Mexico and the Caribbean region. Currently, 56 species of Scolelepis are recognized, including 45 of the subgenus Scolelepis, and 11 species of subgenus Parascolelepis (Delgado-Blas, Reference Delgado-Blas2006). Few Scolelepis species have been reported for the Brazilian coast: S. (S.) chilensis Hartmann-Schröder, 1962; S. (S.) goodbodyi Jones, 1962; S. (S.) squamata Müller, 1806; and S. (Parascolelepis) texana Foster, 1971 (Amaral et al., Reference Amaral, Nallin and Steiner2006). This paper reports the first record of the genus Scolelepis on the north-east coast of Brazil, enlarging the geographical distribution of Scolelepis (S.) lighti from the Gulf of Mexico (Tamaulipas) to the Brazilian coast.
MATERIALS AND METHODS
The material was collected from intertidal sand beaches on the coast of João Pessoa city (Cabo Branco and Bessa beaches), in the State of Paraíba, during the first week of September 2007. Specimens were fixed in 10% formaldehyde solution and, after 24 hours, the material was washed in distilled water, and transferred to 70% ethanol for long-term preservation. The specimens were examined under a stereomicroscope. For a complete analysis of species, some specimens were prepared on permanent slides with pure glycerin or using contemporary techniques such as Gray & Wess (PVA) and Hoyer. The sections prepared under Gray & Wess or Hoyer had the muscle excess removed, and after that, were mounted with Gray & Wess on a slide for permanent conservation according to the methodology described by Brasil et al. (in preparation). To increase the contrast of some chaetae, some sections were hydrated, and then stained with Mallory coloring or immersed for three hours in alcian blue. Drawings of the notopodial and neuropodial chaetae were made using a camera lucida. The material has been deposited in the Invertebrate Collection of Paulo Young (Coleção de Invertebrados Paulo Young), at Universidade Federal da Paraíba (UFPB), under the abbreviation CIPY-POLY.
SYSTEMATICS
DIAGNOSIS
Prostomium conical, extended anteriorly to long, tapering point, posteriorly triangular, raised, occipital tentacle present or absent, branchiae beginning on chaetiger 2, partially fused to notopodial postchaetal lamellae, continuing to near posterior end, neuropodial postchaetal lamellae notched or separate on posterior chaetigers, neuropodial hooded hooks present in all kinds, pygidium with oval disc or multilobed appendages.
Key to Scolelepis (Scolelepis) from Brazilian coast
1. Notopodial hooded hooks present … … … … … … …2
— Notopodial hooded hooks absent … … … … … … … . 3
2. Peristomium distinct from prostomium, with low lateral wings; anterior notopodial lamellae oval, elongated, slightly ruffled; all hooks only tridentate … … … … … … … . … … … … … … … . . . (S.) lighti Delgado-Blas, 2006
—Peristomium distinct from prostomium, with low lateral wings; anterior notopodial lamellae oval, elongated, slightly ruffled; all hooks bidentate … … … …… … … … … . … … … … … S. (S.) chilensis Hartmann-Schröder, 1962
3. Peristomium long; hooks unidentate and bidentate; notochaetae of chaetiger 1 of one type: nearly straight, thin … … … … … … … . . . . S. (S.) squamata Müller, 1806
—Peristomium long; hooks uni-, bi-, and tridentate; notochaetae of chaetiger 1 of two types: (1) abruptly curved, very broad and (2) nearly straight, thin … … … … … … … … … … … … … . .S.(S.) goodbodyi Jones, 1962
Scolelepis (Scolelepis) lighti Delgado-Blas, 2006
EXAMINED MATERIAL
CIPY-POLY 926—939, 946—947, 962—964.
DESCRIPTION
Five complete specimens, 27 to 32 mm in length. Mature individuals were not observed. Specimens of S. (S.) lighti found herein are characterized by having a conical and inflated prostomium, without occipital antennae (Figure 1B,C). The caruncle is present on the prostomium, being conical and inflated, and projected posteriorly to chaetiger one (Figure 1A–C). Two pairs of small brown eyes are present, arranged in a transverse row. Lateral eyes kidney-shaped and inner ones rounded. There are big reddish spots above the eyes (Figure 1B). Narrow curved semicircular nuchal organs surround posterior margin of the prostomium. Peristomium long, distinct from chaetiger one, slightly inflated, forming well developed lateral wings (Figure 1A–C). Proboscis eversible, unarmed, with a sac-like form (Figure 1C). First chaetiger reduced, but with notopodial lamella well developed; notopodial postchaetal lamellae triangular and neuropodial postchaetal lamellae subtriangular, with capillary chaetae in both rami (Figures 1A–C & 2E). Branchiae present from chaetiger 2 to end of body (absent on last chaetigers); branchiae tapered, elongate, longest on anterior and middle part of body, partially fused to notopodial postchaetal lamellae (in the middle of branchiae). Branchiae of posterior chaetigers fused basally to lamellae (Figures 1A–C & 2). Branchiae with band of cilia along inner and outer edge, not seen with stereomicroscope. Anterior and middle notopodial postchaetal lamellae entire, with slightly ruffled edge (Figure 2). Posterior lamellae oval, with elongated pointed tip (Figure 2D–K). Low dorsal folds present on chaetigers 3–21, becoming low ciliated crests, continuing to end of body (Figure 2D–K). Neuropodial postchaetal lamellae on chaetiger 1 ellyptical (Figure 2D). Lamellae on chaetigers 2 oval (Figure 2E); on the following chaetigers the lamellae are rounded, with a slight notch in chaetigers 21–25 (Figure 2J,K). This notch becomes deeper posteriorly, dividing the lamellae into two lobes in chaetigers 22–29 (Figure 2K). On the following chaetigers the lower lobe remains triangular, being located ventrally to the hooks. Posterior hooks accompanied by two small, broad, limbate, finely tapered chaetae. Hooded hooks with two small apical teeth surmounting large main tooth and with long, curved, shaft (Figure 2B,C). Neuropodial hooded hooks beginning generally from chaetigers 10 to 14, numbering seven to nine per fascicle, but sometimes they occur from chaetigers 23 to 24, numbering four to five per fascicle. The variation in the starting point of the neuropodial hooks was directly related to body size. In larger body lengths, the neuropodial hooks usually begin in more posterior segments. Sabre chaetae absent. Notopodial capillary chaetae similar in morphology to those of the neuropodium, although more elongate, arranged in two rows, those of posterior row longest (Figure 2A). Notopodial hooded hooks absent, even in the five complete specimens with up to 72 chaetigers. Most specimens examined were broken in the middle of the body. The relatively small number of the chaetigers could be the reason for the absence of notopodial hooded hooks. Pygidium with ventral cushion (Figure 1D,E).
Scolelepis (S.) lighti. (A) CIPY-POLY 928 in lateral view, anterior and median regions; (B) CIPY-POLY 964 in dorsal view; (C) CIPY-POLY 963 in lateral view of anterior region; (D) CIPY-POLY 928 in ventral view of posterior region showing the ventral cushion; (E) CIPY-POLY 928 in lateral view of posterior region. Scale bars: A,B, 1 mm; C,D, 0.5 mm; E, 1 mm.
Scolelepis (S.) lighti Delgado-Blas, Reference Delgado-Blas2006. (A) Capillary chaeta; (B) tridentate hooks; (C) detail of tridentate hooks; (D) chaetiger 1; (E) chaetiger 2; (F) chaetiger 6; (G) chaetiger 7; (H) chaetiger 13; (I) chaetiger 20; (J) chaetiger 21; (K) chaetiger 22. Scale bars: A, 0.58 mm; B, 1.45 mm; C, 0.58 mm; D–K, 2.9 mm.
REMARKS
No Scolelepis species have been published before in the north-east region of Brazil. The material examined is similar to the descriptions of S. (S.) chilensis Hartman-Schröder, 1962, and of S. (S.) lighti Delgado-Blas, Reference Delgado-Blas2006, because both species have a similar form of prostomium, peristomium, organization of eyes, form and organization of lamellae, and form of pygidium. The illustrations provided by Hartmann-Schröder in 1962 and Blake (Reference Blake1983) show that the notopodial lamellae are less fused to the branchiae than in S. (S.) lighti. Therefore only the examination of type series using the same methodology could solve the question of a possible synonymy between the two species. Although Delgado-Blas (Reference Delgado-Blas2006) described S. (S.) lighti as having tridentate hooks, the apical teeth were not well illustrated. Studies of intraspecific and interspecific variation of S. (S.) chilensis and S. (S.) lighti have to be carried out to comprehend this characteristic. Consequently, the illustrations of hooded hooks presented in the description of S. (S.) chilensis and S. (S.) lighti are very similar, but no observations under a scanning electron microscope have been done to confirm that specimens from both species have tridentate hooded hooks.
Blake (Reference Blake1983) re-examined the type specimens of S. (S.) chilensis and considered the apical teeth of the hooded hooks fused by a ridge as being another distinguishing characteristic of S. (S.) chilensis, whereas S. (S.) squamata is characterized by having unidentate, bidentate and tridentate hooded hooks with distinct apical teeth. The specimens examined by us show tridentate hooded hooks with two distinct apical teeth. These characteristics reinforce the difference between S. (S.) chilensis and S. (S.) lighti. On the other hand, this character was not clearly illustrated in the type material of S. (S.) lighti.
Delgado-Blas (Reference Delgado-Blas2006) shows that S. (S.) lighti is most similar to S. (S.) squamata Müller, 1806, and that it differs in the form of the anterior and middle notopodial lamellae, the shape of the branchiae, and for having only tridentate hooded hooks. Another species was found along the Brazilian coast, S. (S.) goodbodyi Jones, 1962. This species could be easily distinguished from S. (S.) lighti and S. (S.) squamata by the presence of bands of pigmented cells in the first chaetigers, by the inflated shape of the peristomium, the lamellate form of the parapodia, and the segment in which neuropodial hooks begin (on chaetigers 10–14 in S. (S.) lighti, and on chaetigers 43–50 in S. (S.) squamata).
Recently, some specimens have been identified as S. (S.) cf. chilensis for the south and south-east of Brazil by Maria Emilia Diez and Vasily Radashesvky (personal communication), but no description or illustration has been provided for this material up to the present time.
GEOGRAPHICAL DISTRIBUTION
Scolelepis (S.) lighti is now distributed from the Gulf of Mexico (Tamaulipas, Tabasco, Quintana Roo) to Brazil (State of Paraíba, coast of João Pessoa city).
ACKNOWLEDGEMENTS
Many thanks to Anne Isabelley Gondim de Farias, Aline dos Santos Lima, Carmen Alonso Samiguel, Daniela Correia Granjeiro, Fabiolla Maria Alencar, Felipe Ferreira Campos, Igor Alex Binder d'Angelis, João Gilherme Barreto De Marchi, Laurivania Daniella Silva de Souza, Pablo Riul, Patricia Lacouth and Ricardo Silva Cardoso for helping to collect samples. We are grateful to Maria Emilia Diez and Paulo Cesar Paiva for discussions and manuscript revision. We are grateful to Programa de Pós-Graduação em Ciências Biológicas for laboratory material received. André Souza dos Santos and Dimítri de Araújo Costa acknowledge financial support from PIBIC/CNPq. Martin Lindsey Christoffersen acknowledges financial support from CNPq.
