BULLETIN ZOOLOGICAL MUSEUM UNIVERSITY OF AMSTERDAM. Vol. 52, no. 7. UniversidadAutdnoma de Madrid. Abstract. are: S. beneliahuae (Campoy

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1 UAM, Beaufortia BULLETIN ZOOLOGICAL MUSEUM UNIVERSITY OF AMSTERDAM Vol. 52, no. 7 November 29, 2002 Species Syllis Savigny in Lamarck, 1818 (Polychaeta: Syllidae) living in corals in the state São Paulo, SoutheasternBrazil João Miguel de Matos Nogueira & Guillermo San Martin *Departamento de Apologia, Institute de Biociencias, Universidade de Sao Paulo USP, R. do Matao, travessa 14, n 101, , SP, Brazil nogueira@ib.usp.br **Departamento de Biologia, Unidad de Apologia, UniversidadAutdnoma de Madrid Canto Blanco, 28049, Madrid, Spain. guillermo.sanmartin@uam.es Abstract Eleven species Syllis Lamarck, 1818 found in living coral heads Mussismilia hispida (Verrill, 1868) in the State São Paulo (Brazil) are reported and described, based on detailed analysis under stereo, light, and scanning electron microscopes. Comments on the identity each species are provided. The species are: S. beneliahuae (Campoy & Alquézar, 1982), S. corallicola Verrill, 1900, S. glandulata n. sp., S. gracilis Grube, 1840, S. hyllebergi (Licher, 1999), S. lutea (HartmannSchröder, 1960), S. maryae San Martín, 1992, S. prolifera Krohn, 1852, S. pseudoarmillaris n. sp., S. truncata Haswell, 1920, and S. tyrrhena (Licher & Kuper, 1998). Syllis glandulata n. sp. and S. pseudoarmillaris n. sp. are herein described. Except for S. gracilis, S. prolifera, and S. corallicola, this is the first report for all those species from Brazilian waters. Syllis hyllebergi is reported for the first time after the original description. INTRODUCTION Syllidae (Nogueira et al., 2001b), together with Coral heads Mussismilia hispida (Verrill, 1868) three redescriptions (Nogueira, 2002; Radashevsky & Nogueira, in press; Steiner et al., 2002). Prior to the present study, ten species Syllis proved to support a rich and diverse assem very blage polychaetes, dominatedmainly by syllids had been reported for Brazilian waters, most and spionids (Nogueira, 2000). A taxonomic analysis this assemblage in them, however, dissertations and thesis not formally published, in the sense the IGZN (see is currently being carried out and, until present, it has Table 1). led to the description several new taxa Besides not having been formally registered, (Nogueira & Amaral, 2000, 2001; Nogueira & most the species reported by those studies have Ten Hove, 2000; Nogueira & Rizzo, 2001; also not been fully described. On the other hand, Nogueira et al., 2001a, b, in press; Nogueira & taxonomists all around the world are presently KnightJones, 2002), some them belonging to coming to the conclusion that many the old 57

2 Table 1: Species Syllis previously reported for Brazilian waters, the localities in which they were found, and the main references; those references marked with * have not been formally published; those taxa marked with * are doubtful records (see Remarks S. pseudoarmillaris n. sp. for explanation). Species Localities (states) References Syllis amica Syllis gracilis Sao Paulo Rio de Janeiro Sao Paulo Souza, 1989* Attolini, 1997* Hansen, 1882 (as Syllis brevicirris); Morgado and Amaral, 1985; Rullier and Amoureux, 1979 Espirito Santo, Rio dejaneiro, and Rio Grande do Sul Rullier and Amoureux, 1979 Syllis corallicola fas Syllis (Typosyllis) tigrinoides] Syllis prolixa [as Syllis [Typosyllis) prolixa] Syllis armillaris (as Typosyllis armillaris)* Bahia Rio Grande do Sul Rio Grande do Norte, Pernambuco, and Bahia Alagoas, Augener, 1931 Orensanz and Gianuca, 1974 Rullier and Amoureux, 1979 Syllis ehlersioides (as Typosyllis ehlersioides) Rio Grande do Sul Borzone, 1988* Syllisfasciata (as Typosyllis fasciata) Syllis hyalina (as Typosyllis hyalina)* Syllis prolifera (as Typosyllis prolifera) Syllis variegata (as Typosyllis variegata) Sao Paulo Rio Grande do Norte, Bahia Sao Paulo Parana Alagoas, Sergipe Rio de Janeiro Sao Paulo Rio Grande do Norte, Pernambuco, Bahia, Rio de Janeiro Alagoas, Sergipe Sao Paulo Souza, 1989* Rullier and Amoureux, 1979 Paiva, 1996 Lana, 1981*, 1984* Nonato and Luna, 1970 Attolini, 1997*; Machado, 1989* Souza, 1989* Rullier and Amoureux, 1979 Nonato and Luna, 1970 Morgado and Amaral, 1985; Duarte and Nalesso, 1996 Parana Lana, 1981*, 1984* herein treated. 'well known species', priorly considered to be cosmopolitans, are in fact complexes sibling species (for example, see Martin et al., (in press), MATERIALS AND METHODS review ' Haplosyllis spongicola ), which can only be distinguished if researchers have access to Twelve colonies Mussismilia hispida (Verrill, 1868) were collected in two islands f the coast descriptions the features different populations. the State Sao Paulo, Ilha dos Alcatrazes In the present paper, all species Syllis found in colonies M. hispida are described, based on analysis under stereo, light and scanning eletron microscopes, except for S. truncata Haswell, 1920, (26 06'S 45 42'W) and Laje de Santos (24 19'S 46 11'W), on 17 March 1996 and 4 December 1996 respectively, by means SCUBA diving, in shallow water (310 m deep). which has not been analysed under SEM due to Corals were detached from the substrate with the shortage specimens and their poor state the aid a chisel, put inside plastic bags and preservation. Intraspecific variation within all fixed with 4% formalin. In the lab, corals were decalcified with formaldehyde and formic acid taxa was evaluated and information is incorporated in the descriptions. solution, and the polychaetes sorted and stored in Except for S. corallicola Verrill, 1900, S. gracilis 70% ethanol, before identification. Grube, 1840 and S. prolifera Krohn, 1852, this is the first report for Brazilian waters for all species Observations and measurements were made using Nomarsky interferencecontrast optics. 58

3 Body Line drawings were made with a drawing tube, REMARKS. Syllis is one the most controversial genera among polychaetes. Based mainly on the morphology the chaetae, it was in four split from specimens mounted on glycerine jelly permanent slides. Observations with scanning electron microscope (SEM) subgenera by Langerhans (1879), which were were made after criticalpoint drying and coating with 25 nm gold, at Laboratorio de Microscopia Eletronica, Instituto later raised to generic level: Ehlersia Langerhans, 1879, Haplosyllis Langerhans, 1879, Syllis Savigny de Biociencias, Universidade de Sao Paulo (IB in Lamarck, 1818, and Typosyllis Langerhans, USP) and Laboratorio de Microscopia Eletronica, Instituto de Biologia, Universidade Estadual de Campinas (IBUNICAMP), both in Brazil Ehlersia is characterized by having pseudospinigerous chaetae together with much short Specimens are deposited at Museu de Historia er falcigers. According to San Martin (1992), the Natural (MHN), Brazil, and Museo Nacional de Ciencias Naturales de Madrid (MNCNM), length the blades falcigers should be regarded as a specific character, instead a generic Spain, and, in case the two new species here one. Therefore, we do not accept Ehlersia as a described, also at the Invertebrate Reference Collection the Florida State Board Conservation (FSBC I). valid taxon, and consider all its species as belonging to Syllis, the only exception being E. ferrugina Langerhans, 1879, whose assignment to Syllinae was doubtful (see San Martin, 1992), and TAXONOMIC PART has been proposed as belonging to Eusyllinae by Nunez & San Martin (1996). Family Syllidae Grube, 1850 Subfamily Syllinae Grube, 1850 Haplosyllis is a clearly defined with the genus, Genus Syllis Savigny in Lamarck, 1818 autapomorphy possessing only thick truly simple chaetae, together with the absence capillary dorsal and ventral simple chaetae. The major problems on the systematics the TYPE SPECIES. Lamarck, 1818 Syllis monilaris Savigny in group, however, arise when separating Syllis from Typosyllis. Some authors, such as Langerhans (1879), Kudenov & Harris (1995), and Licher DESCRIPTION. cylindrical, elongated, with (1999), among several others, consider Syllis as only having simple, thick chaetae, typical S. gracilis Grube, 1840, in midbody chaetigers, and large number segments. Prostomium with one pair palps, joined basally, projecting downwards. One median antenna and one pair lat the remaining species, which do not present that eral antennae present; most the species with particular type chaeta, as Typosyllis, although sometimes they state that this separation eyes, two posterior pairs and commonly one additional pair anterior ocular spots. Peristomium is artificial but keep it for practical reasons, due to the with two pairs tentacular cirri, the superior usually longer. Antennae, tentacular, dorsal and anal cirri moniliform throughout. Ventral cirri usually shorter than parapodial lobes. Compound chaetae heterogomph, blades falcigers variable length, from lost or fused to the shaft (simple, thick chaetae), to very elongated (pseudospinigerous chaetae); some posterior parapodia with dorsal and ventral capillary simple chaetae. Acicula present, usually more than large number species the group (see Kudenov & Harris, 1995: 82). However, other characters show similarity between S. S. gracilis, armillaris, S. and the hyalina two new species herein described, such as the shape dorsal cirri and the dorsal and ventral simple chaetae; besides this, at morphology least S. armillaris and S. hyalina have the same kind stolon as S. gracilis (Estape & San Martin, 1991). Further, small specimens S. gracilis have only true compound chaetae and the fusion one in anterior chaetigers, progressively diminishing in number per parapodium towards pygidium, until, in most species, one single aciculum shafts and blades occurs progressively as the animals grow, leaving a very shallow sulcus, however persists in each posterior parapodium (San Martin, 1984, 1992). true compound chaetae are always present, at least in anterior chaetigers (see 'Description' S. 59

4 ... gracilis). Finally, there are species such as S. magellanica Augener, 1918, which present superior chaetae similar to those S. gracilis, but with distal one; posterior chaetigers each with aciculum subdistally inflated, with short and oblique tip S. hyllebergi shaft and blade still distinct, together with true 4a. Simple, thick chaetae absent 5 compound chaetae, in midbody chaetigers, clearly demonstrating that reduction the blades followed by fusion shafts and blades occurred progressively along the evolution these species. Further discussions on the subject can be seen in San Martin (1992 and in press). For all the above, consider we Syllys as derived b. Adults only with simple, thick chaetae in midbody chaetigers; juveniles with shafts fused to some partially blades in some midbody parapodia S. gracilis 5a. From midbody, dorsal cirri and alternating long short, which at least the longest exceeds body width b. From dorsal cirri about the midbody, same length, usually shorter than body width 8 6 from the same ancestral stock as the group named by Licher (1999) as ' Typosyllis armillaris complex', rendering Typosylis paraphyletic if both are considered. genera Therefore, we see no reason to keep Typosyllis as a valid taxon at this moment, and consider Syllis as defined above, presenting compound chaetae as falcigers with variable length the blades, which may be very long or reduced until the point being lost or fused with the shaft. Obviously, such statement requires further cladistic support, calling for an urgent analysis the whole group, Syllis and Typosyllis. We believe 6a. Posterior acicula subdistally inflated, with pointed oblique tip 7 b. Posterior acicula distally inflated, rounded and hollow S. prolifera 7a. Anterior dorsal cirri with dark inclusions; blades falcigers with both teeth about the same length, or subdistal tooth slightly shorter, and with large rounded gap in between; in large specimens, anterior dorsum with transversal dark red lines, leaving one pair dorsal white circles per segment b. Anterior dorsal cirri with light, S. corallicola iridescent inclusions; blades falcigers with long subdistal spines, reaching or surpassing the level subdistal tooth; from midbody, subdistal tooth falcigers larger than distal that only after such study is performed, it will be one S. lutea possible to answer the question whether both taxa are valid or not. KEY TO THE SPECIES OF SYLLIS FOUND IN LIVING CORALS IN THE STATE OF SAO PAULO 8a. Posterior acicula subdistally inflated, with pointed oblique tip; dorsal chaetae bidentate simple 9 b. Posterior acicula curved at subdistally right angle; dorsal simple chaetae straight, distally truncate 10 9a. Blades falcigers with dorsoventral the gradation length the blades conspicuous throughout; parapodia with dorsal glands, containing granular material.. Pseudospinigerous chaetae present la. 2 b. Pseudospinigerous chaetae absent 4 2a. Anterior parapodia with 2 acicula each; dorsal simple S. glandulata n. sp. b. Blades falcigers very short in midbody parapodia, with dorsoventral gradation the the length blades reduced or absent; parapodia without dorsal chaetae distally truncate, bifid; posterior chaetigers with acicula distally curved at right or subdistally glands S. pseudoarmillaris n. sp. inflated with short and oblique tip 3 10a. Body long and very slender; appendages short b. Anterior parapodia with 34 very thin acicula each; dorsal simple chaetae thin and curved, distally biden throughout,except for antennae, tentacular and dorsal cirri chaetiger 1; falcigers with sharp teeth tate; posterior chaetigers each with aciculum subdis S. tyrrhena tally inflated with short and oblique tip.. S. beneliahuae b. Body thick; dorsal cirri slightly longer than body width, with dark inclusions; falcigers with blunt 3a. Tip pseudospinigers clearly bidentate, with teeth teeth S. truncata close to each other and about the same size; falcigers with subdistal tooth shorter than distal one through out; posterior parapodia each with aciculum distally curved at right angle S. maryae b. Tip pseudospinigers bidentate (appearing rounded, unidentate to subbidentate at highest magnification light microscope), with teeth rounded; posterior falcigers with subdistal tooth larger and stronger than Syllis beneliahuae (Campoy & Alquézar, 1982) Figs. 12 Langerhansia beneliahui Campoy & Alquezar, 1982: 60

5 Our 124, 3. fig. Syllis beneliahuae; San Martin, 1984: , Pis. 9091; Capaetal., 2001: 107. Typosyllis beneliahuae; Licher, 1999: 4748, figs. 17B, 22, see also for synonymy. distal tooth hooked; cutting edges with spines all along their length, stronger in the basal half (Figs. 1DE, GH, 2CG, JL), with about four to nine chaetae, which one to two are pseudospinigers and three to five falcigers; from midbody, fewer chaetae present on each parapodium: usually one MATERIAL. 21 specimens, all from Ilha dos Alcatrazes. Three specimens deposited at MHN (MHNBPO 02/13) and two specimens deposited at MNCNM (16.01/8725 and pseudospiniger and three to five falcigers. Blades pseudospinigers with teeth about the same size, distally pointed (Figs. IE, G, 2F), blades 16.01/8726). Six specimens retained. examined under SEM not increasing in length towards midbody, and measuring, on average, 31, 48, and 35 mm, on anterior, midbody and posterior chaetigers, DESCRIPTION. Body long and slender, without pigmentation the patterns; longest complete specimen examined counting on 98 chaetigers and respectively. Blades falcigers with dorsoventral gradation in length (Figs. ID, H, 2G), not evident anteroposteriorly; blades measuring, on average, measuring about 7.9 mm in length and 0.21 mm 22, 18, and 13 (im on anterior parapodia; 23, 18, in width, at the level proventriculum; longer and 13 incomplete specimens have also been studied. on midbody; and, on posterior parapodia, 20, 16, and 12 um. Pseudosponigerous Prostomium subpentagonal to ovate, broader chaetae absent in last chaetigers. Dorsal and ventral simple chaetae thin, bidentate, with subdistal than long, with four posterior eyes in wide trapezoidal arrangement (Fig. 1A). Lateral antennae on anterior border prostomium, with 1016 articles, median antenna originating on posterior prostomium, with 1623 articles, about twice the length lateral pair in most specimens (Fig. 1A). spines, and about the same size, but ventral chaetae slightly thicker (Figs. IF, J, 2MN); both types simple chaetae only present on posterior chaetigers. Acicula subdistally inflated, with short acuminate tip; three to four thin acicula present Palps longer than prostomium, ventrally directed in each anterior parapodium (Fig. 1C), two acic (Figs. 2AB). Peristomium dorsally reduced, superior pair cirri slighdy shorter than median ula in each midbody parapodium (Fig. II), and single aciculum, with the same but shape, thicker, antenna, inferior pair about half the length in each posterior parapodium (except for one superior cirri, with about 714 articles (Figs. 1A, 2AB). specimen, which keeps two acicula per parapodium until pygidium). Dorsal cirri containing pairs fibrillar spiral inclusions, dark in longer specimens; cirri REMARKS. specimens differ from those chaetiger 1 much longer than following cirri and examined by Licher (1999) by having blades pseudospinigers remarkably shorter in midbody and Another difference posterior chaetigers. between our specimens and those studied by Licher (1999) is the presence two acicula in each in posterior parapodium, the specimens median antenna, with 1937 articles, depending on the size the specimen. From midbody, parapodia alternating long and short cirri, the longest about twice the length the shortest (Fig. 2H). Ventral cirri digitiform throughout, not exceeding parapodial lobes (Figs. 2AB, H). Anal cirri examined by Licher, one them is thin and very with about 820 articles and one ventral papilla in frequently hidden by the stouter aciculum (see between (Figs. IB, 21). Licher, 1999: 47), while in our specimens there is Pharynx long, extending through 811 segments, with one tooth on anterior margin. Proventriculum with abouthalf the length the only one stout aciculum in each posterior parapodium, except for one as specimen, mentioned above. In to regard that particular specimen, it is worth to mention that, besides being the only one pharynx, occupying 4.57 chaetigers and counting on 3035 rows musclecells (Fig. 1A). with two acicula in each posterior parapodia, it is Compound chaetae falcigers and pseudospinigcrs, both with bidentate blades, teeth about the same length, subdistal tooth triangular, also aberrant in the length the blades posterior falcigers, being very similar to those midbody parapodia, and lacking dorsal and ven 61

6 Fig. 1. Syllis beneliahuae. A, anterior end, dorsal view. B, posterior end. C, acicula, anterior chaetiger. D, falcigers, anterior chaetiger. E, pseudospinigerous chaeta, anterior chaetiger. F, dorsal simple chaeta. G, pseudospinigerous chaeta, midbody chaetiger. H, falcigers, midbody chaetiger. I, acicula, midbody chaetiger. J, ventral simple chaeta (scale bars: A = 250 μm; B = 100 μm; CJ= 10 μm). 62

7 Fig. 2. Syllis beneliahuae, scanning blades detail midbody, superior the tip ventral and inferior electron micrographs. A, falcigers, respectively, anterior anterior pseudospinigerous chaeta, midbody chaetiger. G, view. I, posterior end, arrow end, chaetiger. E, blades indicates papilla between lateral view. anterior B, blade superior anal end, ventral view. CD, falciger, midbody chaetiger. F, intermediary falcigers, midbody chaetiger. H, cirri. JL, blades superior, intermediate and inferior falcigers, respectively, posterior chaetigers. M, dorsal simple chaeta. N, ventral simple chaeta (scale bars: A, F, I = 100 μm; B = 200 tral simple posterior chaetae; it μm; CD, is possibly JN = 3 μm; E = 4 regenerating Pacific (Capa et al., Mediterranean Sea end. Caribbean DISTRIBUTION. μm; G = 5 μm; H = Most probably circumtropical, with several records in the Atlantic and one in the (Cuba), 2001). (Spain), and Pacific Ocean: Panama first report Atlantic Canary southeastern (Coiba Island). for the Brazilian 1 μm). Ocean: Islands, Brazil. This is the coast. 63

8 Body Our Syllis maryae San Martin, Figs. Syllis maryae San Martin, 1992: , 9. fig. Typosyllis maryae; Licher, 1999: MATERIAL. 15 specimens, all from Ilha dos Alcatrazes. Three specimens deposited at MHN (MHNBPO 03/13), and 2 specimens deposited at MNCNM (16.01/8718 and 16.01/8729). 410 compound chaetae, including 12 pseudospinigers; from midbody, parapodia each bearing 12 pseudospinigerous and 34 falcigers (Fig. 4H). Pseudospinigerous chaetae absent in last chaetigers; blades increasing in length towards posterior chaetigers, evidendy bidentate, with teeth about the same size (Figs. 3D, G, 4E, G, H); blades measuring, on average, 40 am, in anterior parapodia, 59 jim, in midbody, and 59 in posterior segments. Falcigers with blades with conspicuous spines all along the cutting edge, spines DESCRIPTION. elongated, without distinct thicker basally, progressively diminishing in size pigmentation; the longest complete specimen with 73 chaetigers and measuring about 5.4 mm towards the tip; blades bidentate, distal tooth hooked and subdistal tooth triangular, teeth in length and 0.16 mm in width, at the level about the same size, or subdistal tooth shorter, proventriculum. and distinct gap in between (Figs. 3D, G, 4DF, Prostomium broader than subrectangular, long, with two posterior pairs eyes in trapezoidal arrangement (Fig. 3A). Lateral antennae originating near anterior border prostomium, with 711 articles; median antenna on posterior prostomium, between posterior with about eyes, 1119 articles (Figs. 3A, 4A). Palps longer than prostomium, distally rounded (Figs. 3A, 4AB). Peristomium dorsally narrower than following HI, KL, OQ). Blades falcigers measuring, on average, 19, 17, and 10 (am, on anterior parapodia; on midbody, blades, on average, measuring 20, 15, and 9 p.m; on posterior body, blades width 18, 16, and 10 Dorsal and ventral simple chaetae usually present only in posteriormost jam. chaetigers, dorsal chaetae distally truncated (Fig. 3F); ventral chaetae bidentate, with short subdistal spines (Figs. 3H, 4N), slightly shorter than dor segments, superior pair cirri about the same sal chaeta. Anterior parapodia each with two aci length as lateral antennae, or slightly longer, with cula, thin, two types: subdistally inflated, with 817 articles; inferior pair slightly shorter, with 8 short oblique tip, and distally curved at right 11 articles (Figs. 3A, 4AB). Dorsal cirri chaetiger 1 much longer than following cirri and antennae, with about 1922 angle, with short and acuminate tip (Fig. 3C); posterior parapodia each with single aciculum, thicker, the second type (Fig. 3E). articles (Figs. 3A, 4AB). Cirri containing fibrillar spiral inclusions throughout, iridescent. Midbody REMARKS. specimens differ from those cirri shorter, with 514 articles; in some specimens, alternating long and short cirri for a short extension (Fig. 4J). Ventral cirri short and digitiform throughout, up to the same length as parapodial lobes (Fig. 4B). Anal cirri with about 1012 articles and one papilla in between (Fig. 3B, 4M). Pharynx long, extending for 611 segments, with one big tooth slighdy posterior to the anterior margin, which is surrounded by a crown about 10 st papillae (Figs. 3A, 4BG). Proventriculum extending through 47 chaetigers, with rows musclecells. Anteriormost parapodia bearing only falcigers with blades superior pair progressively more described by San Martin (1992), and Licher (1999) by having: considerably shorter antennae and cirri throughout the body, not alternating long and short dorsal cirri, except for short extension in a few specimens; slighdy longer pseudospinigers in midbody parapodia, and much shorter inferior falcigers; presence dorsal simple chaetae far more posteriorly; and pharynx much longer. Licher (1999) considers postventricular dorsal as the most characteristic feature this glands species, however such structures were absent in all Brazilian specimens. Similar structures occur in several species, such as S. amica and S. pulvinata Langerhans, 1881 (San Martin, in press), but in elongated (Figs. 4DE), becoming pseudospinigerous chaetae from chaetiger 6 onwards. all them, including S. maryae, only a few speci Anterior parapodia, from chaetiger 6, each with mens present this character. Therefore, we 64

9 Fig. 3. Syllis maryae. A, anterior end, dorsal view. B, posterior end. C, acicula, anterior chaetiger. D, compound chaetae, anterior body. E, aciculum, posterior chaetiger. F, dorsal simple chaeta. G, compound chaetae, midbody. H, ventral simple chaeta (scale bars: A = 200 μm B = 50 μm; CH = 10 μm). 65

10 Fig. 4. Syllis maryae, scanning electron micrographs. A, anterior end, dorsal view. B, anterior end, ventral view. C, detail proboscis. D, blade superiormost falciger, chaetiger 1. E, blades compound chaetae, chaetiger 8. F, detail the blades falcigers, chaetiger 8. G, tip pseudospinigerous chaeta, midbody chaetiger. HI, blades compoundchaetae, midbody chaetigers. J, midbody, dorsal view. KL, blades superior and inferior falcigers, respectively, midbody chaetiger. M, posterior end, lateral view, arrow indicates papilla between anal cirri. N, ventral simple chaeta. OQ, blades superior, intermediate, and inferior falcigers, respectively, posterior chaetigers (scale bars: A, B and M = 100 μm; C = 20 μm; D, H and I = 5 μm; E = 10 μm; F and K = 4 μm; J = 200 μm; L = 2 μm; G and NQ = 3 μm). believe such a great importance should not be each anterior parapodia, in contrast to two in S. attributed to it. maryae, which are also thin, but much stronger Syllis maryae and S. beneliahuae are very similar at first glance, but they can be distinguished because S. beneliahuae has: (1) three to four thin acicula on than those Syllis beneliahuae; (2) posterior acicula subdistally inflated with short and oblique tip, while in S. maryae acicula are bent at right angle; 66

11 Body Western (3) dorsal simple chaetae curved and bidentate in and several terminal papillae (Fig. 5A). contrast to truncate condition in S. maryae; and (4) body more elongate and proportionally much Proventriculum occupying 49 segments, 35 rows musclecells (Fig. 5A). with 29 thinner, with dorsal cirri containing pairs dark Each anterior parapodium with about 59 inclusions in each article, and alternating cirri different lengths throughout, which the longest compound chaetae, midbody with 56, and posterior chaetigers with 45 each, including 12 are about twice the length the shortest. pseudospinigerous chaetae, present in all but the posteriormost parapodia; in some specimens DISTRIBUTION. Atlantic: North Carolina, Caribbean Sea (SalazarValejo, 1996) and State Sao Paulo. This is the first for report this species on the Brazilian coast. Syllis hyllebergi (Licher, 1999) n. comb. Figs. 56 Typosyllis hyllebergi Lichcr, 1999: 7678, fig. 34. pseudospinigerous chaetae are absent for a long part the body. Blades falcigers bidentate, all with cutting edges conspicuously spinulated, with long spines basally, decreasing progressively in length towards the tip (Figs. 5D, E, 6AE, HJ), pseudospinigerous chaetae with uniform fringe spines all along their length (Figs. 5D, E). Blades falcigers anterior parapodia with teeth closely together, about the same size (Figs. 5D, 6AB); from midbody, gap between teeth progressively increasing, while subdistal tooth becomes MATERIAL. 13 specimens, all from Ilha dos Alcatrazes. stronger, triangular, with large base (Figs. 5E, 6C Three specimens deposited at MHN (MHNBPO 04/13) and 2 deposited at MNCNM (16.01/8730 and 16.01/8731). Two specimens examined under SEM not preserved. E, HJ). Falcigers with blades about the same length throughout the body, with slight dorsoventral gradation; 15, 13, and 11 am, on average, in each anterior parapodia, and 17, 13, and 11 DESCRIPTION. short and proportionally pm from midbody. Blades pseudospinigerous robust, without remarkable colour patterns; chaetae with distal tooth remarkably rounded longest specimen examined about 3.7 mm long, 0.24 mm wide, at the level proventriculum, with 41 chaetigers. Prostomium subpentagonal to ovate, broader throughout (Figs. 5D, E, 6F, G), blades increasing in length towards posterior midbody, measuring, on average, about 45 jam in anterior parapodia, 58 jam in midbody, and 60 pm in posterior parapodia. Dorsal simple chaetae present than long, with four posterior eyes in wide trapezoidal arrangement, and two anterior eyespots. from midbody, shortly after proventriculum; thick, distally Antennae short, lateral in pair originating front anterior eyes, with about 10 articles, and median truncate and bifid, spinulated on distal third (Figs. 5F, 6L). Ventral simple chaetae only on posterior antenna in the middle prostomium, longer, parapodia, distally curved, bidentate, with teeth with 1415 articles (Fig. 5A). Palps longer than well separated (Figs. 5H, 6K). Two acicula on prostomium. Peristomium narrower than following chaetigers, superior pair cirri about twice each anterior parapodia, one which thick, subdistally inflated, with oblique tip, the other the length inferior pair; superior cirri with articles, inferior with 67 (Fig. 5A). Dorsal cirri short and containing iridescent inclusions throughout the body, those straight with slightly oblique tip (Fig. 5B); posterior chaetigers each with single aciculum, with the same shape anterior thicker aciculum (Fig. 5G). chaetiger 1 longer than following cirri, with 1216 articles; remaining cirri with 613 articles, usually alternating long and short in midbody. Ventral REMARKS. Brazilian specimens show slight variation from those described by Licher (1999) by having a shorter pharynx, fewer chaetae per cirri digitiform, about the same length as parapodial lobes (Figs. 5A, C). 12 articles (Fig. 5C). Anal cirri with about 10 parapodium, especially on anterior chaetigers, and by presenting one pair anterior eyespots. Pharynx short, extending for 37 chaetigers, Possibly, those features are all sizedependent, with one big tooth posterior to anterior margin, since our specimens are much shorter than those 67

12 100 Atlantic Fig. 5. Syllis hyllebergi. A, anterior end, dorsal view. B, acicula, anterior chaetiger. C, posterior end. D, compound chaetae, anterior chaetiger, from left to right, inferiorto superior chaetae.e, compound chaetae, posterior chaetiger, from left to right, superior to inferior chaetae. F, dorsal simple chaeta. G, aciculum, posterior chaetiger. H, ventral simple chaeta (scale bars: AC = μm; B and DH: 10 μm). described by Licher; the holotype counts on 63 Syllis gracilis Grube, 1840 chaetigers and measures 8.8 mm in length and Figs mm in width. In recent studies (Nogueira, in prep.), this Syllis gracilis Grube, 1840: species was common in samples from the inter Syllis (Syllis) gracilis; Rullier & Amoureux, 1979: tidal zone, in rocky shores, in greater amount 162; Morgado & Amaral: 1985: 221. than sublitorally, in corals. MATERIAL specimens, from de Laje Santos and DISTRIBUTION. Ocean: Mediterranean Sea (Israel, Cyprus, and Suez Canal), and State Sao Paulo. Indian Ocean: Red Sea (Gulf 127 from Ilha dos Alcatrazes. Three specimens deposited at MHN (MHNBPO 05/13) and 2 specimens deposited at MNCNM (16.01/8716 and 16.01/8717). Five specimens examined under SEM not preserved. Akaba). This is the first record for this species following the original description, and the first record for Brazil and southern Adantic. DESCRIPTION. Body elongate and robust, usually with dorsal transversal bands dark pigmentation until the end proventriculum, two rows 68

13 Fig. 6. electron Syllis hyllebergi, scanning midbody falcigers. F, tip blades chaetiger. HJ, chaeta. L, dorsal pseudospinigerous chaeta, superior, intermediate, chaetae simple (scale blades micrographs. AB, = 3 pigmentation in each anterior chaetiger (Fig. 7B), and one row per proventriculum studied in chaetiger (Fig. 7C). measuring 9.9 width, 72 at but observed, Prostomium posterior 7A). mm the level chaetigers (longer were eyes from the in incomplete studied in subpentagonal, in trapezoidal Antennae short, front anterior eyes, longest specimen length, by 0.33 mm proventriculum, and with and not The beginning lateral with specimens detail). with four mid arrangement (Fig. pair originating 1015 broad in 11 articles Dorsal dorsally slightly pos nar μm; falcigers, chaeta, posterior posterior chaetigers. K, C ventral simple = 5 μm). following segments; superior peristoslightly about 1218 than longer articles, inferior antennae, cirri with shorter, with 8 (Figs. 7A, 8AB). cirri chaetiger 1 antenna, or short about those throughout, the same longer (Figs. 7 A, length 8AB). as median From mid body, cirri shorter, with 69 articles, with broader articles basally Ventral cirri tubercular prostomium, slightly longer, with 1420 articles. Peristomium cirri same terior border near than rower mial 4 = the antenna originating B and L blades chaetigers. CE, G, tip pseudospinigerous falcigers, respectively, mm; articles, and the median anterior falcigers, midbody chaetiger. and inferior bars: A and DK length longer one and very as midbody than dorsal (Fig. 81). about the lobes in anterior (Fig. with up 81). Anal body, cirri to 16 articles and in between (Figs. 7D, 8M). cirri, papilla pointed throughout, parapodial from st ventral distally short 69

14 Fig. 7. Syllis gracilis. A, anterior end, dorsal view (dorsal pigmentation omitted). B, chaetiger 3, dorsal view, showing pigmentation. C, chaetiger 15, dorsal view, showing pigmentation. D, posterior end. E, acicula, anterior chaetiger. F, aciculum, posterior chaetiger. G, simple, thick chaeta. H, falcigers, anterior chaetiger. I, dorsal simple chaeta. J, falcigers, posterior chaetiger. K, ventral simple chaeta (scale bars: A and D [scale at the left Fig. D] = 400 BC μm; [scale at the left Fig. C] = 100 μm; EK [scale at the right side Fig. F] = 10 μm). Pharynx long, extending through 714 seg length the blades and wide gap between distal ments, with large anterior tooth (Fig. 7A). and subdistal teeth; distal tooth the blades Proventriculum shorter, occupying 410 segments hooked, subdistal tooth shorter; cutting edges and with 4161 rows musclecells (Fig. 7A). with a fringe numerous and short spines, Anterior parapodia bearing only compound falcigers, 57 per parapodium, blades bidentate, diminishing in length towards tip (Figs. 7H, 8G E). From mibdody, two simple, thick chaetae per with acentuate dorsoventral gradation in the parapodium, originated by strong reduction 70

15 Fig. 8. Syllis gracilis, scanning electron micrographs. A, anterior end, dorsal view. B, anterior end, ventral view. C, falcigers, anterior chaetiger. D, superior falcigers, anterior chaetiger. E, intermediate falciger, anterior chaetiger. F, G, simple, thick chaetae. G, tip thick chaeta. simple, H, dorsal simple chaeta. I, midbody, lateral view. J, ventral simple chaeta. K, L, falcigers, posterior chaetigers. M, posterior end, arrow indicates the papilla between anal cirri (scale bars: A and B = 200 μm; C and F = 10 μm; D, E, G, H, J and L = 5 μm; I = 100 μm; K = 3 μm; M = 60 μm). the blades and fusion to the shafts; chaetae with few and very short spines near the base the fused blade, and shallow sulcus the along line fusion (Figs. 7G, 8FG). Posterior parapodia with 24 compound falcigers, with blades bidentate, as on anterior parapodia, but shorter and with 71

16 Cosmopolitan. dorsoventral gradation in the length the blades less if pronounced, present (Figs. 7J, 8K the pharynx some specimens; the longest specimen examined in detail with 55 chaetigers and measuring 56 mm in length, by 5.9 mm in width, L). Blades falcigers measuring, on average, about 28, 21, and 14 fim, in each anterior parapodium, and 15, 13, 11 am, in posterior body. Dorsal and ventral simple chaetae on last present chaetigers, dorsal chaetae subidentate, with short subdistal spinulation (Figs. 71, 8H); ventral at the level proventriculum, but longer specimens have been collected. Prostomium ovate, with four posterior eyes in trapezoidal arrangement and, in some specimens, two anterior eyespots (Fig. 9A). Lateral pair antennae on the anterior border prostomium, chaetae shorter, distinctly bidentate, with short with 1320 articles, median antenna longer, with spines (Figs. 7K, 8J). Anterior parapodia each 2028 articles, inserted on posterior prostomium usually with 23 acicula, subdistally inflated with (Figs. 9A, 10A). Peristomium sligthtly narrower than following segments, superior pair cirri very short oblique tip (Fig. 7E); posterior parapodia each with single aciculum, the same shape, but stronger (Fig. 7F). longer than antennae, with 1933 articles, inferior pair shorter, with 1116 articles (Figs. 9A, 10A REMARKS. C). Dorsal cirri long throughout, those This is a well known species, reported worldwide, recognised by the sim presence chaetiger 1 longer than following cirri, with 2332 ple, thick chaetae, originated by the fusion articles (Figs. 9A, 10AB). From midbody, cirri shafts and blades (see San martin, 1984, 1992; in usually alternating long and short (Fig. 1OE), the press), and the very short and distally tapering dorsal cirri from midbody. Due to asexual reproduction followed by incomplete regeneration, longest exceeding body width. Ventral cirri digitiform, up to the same length parapodial lobes (Figs. 10BC, Q). Anal cirri about the same length specimens with simple, thick chaetae extending as dorsal cirri, with 1024 articles (Figs. 10Q). until posterior end are common. Pharynx short, usually with red pigmentation inside, and extending for 36 segments (Fig. 9A), DISTRIBUTION. Along the with sharp tooth near the anterior margin, which Brazilian coast, S. gracilis has been identified in the states Espirito Santo (Rullier & Amoureux, 1979), Rio de Janeiro (Rullier & Amoureux, 1979; Attolini, 1997), Sao Paulo (Morgado & is surrounded by a crown ten st and triangular papillae (Fig. 10D). Prostomium about the same length as pharynx, with about 2529 rows musclecells (Fig. 9A). Amaral, 1985; Nogueira, 2000), and Rio Grande do Sul (Rullier & Amoureux, 1979). Compound chaetae all falcigers, blades bidentate, with distal tooth hooked, subdistal tooth shorter and with wide gap in between (Figs. 9B Syllis prolifera Krohn, 1852 Figs. 910 C, F, 10FN); cutting edges the blades superior chaetae with spines about the same thickness throughout, progressively diminishing in length Syllis prolifera Krohn, 1852: 6675, PL 3, fig. 1. towards the tip (Figs. 10FG), intermediate and Typosyllis prolifera; Nonato & Luna, 1970: 69; inferior chaetae with spines remarkably longer Licher, 1999: , figs. 17 S, 61, see also for synonymy. MATERIAL. 15 specimens, all from Ilha dos Alcatrazes. Three specimens deposited at MHN (MHNBPO 06/13), and 2 specimens deposited at MNCNM (16.01/8727 and 16.01/8728). Four specimens examined under SEM not preserved. and coarser in proximal half (Figs. 10FIN). Anterior parapodia each with 610 chaetae, midbody parapodia each with 611, and posterior parapodia with 37. Blades measuring, on average, 32, 20, and 14 jm, in anterior segments, 30, 25, and 18 (im in midbody, and 24, 20, and 17 im in posterior ones. Dorsal simple chaetae present from midbody, immediately after the proven DESCRIPTION. Body long and robust, without triculum in some specimens, long and thick, dis distinct colour patterns, except for dark inclusions inside dorsal cirri, and red pigmentation inside tally bifid (Figs. 9E, 100); ventral simple chaetae from shorter and thinner than posterior body, dorsal chaetae, distally curved, strongly bidentate, 72

17 This Fig. 9. Syllis prolifera. A, anterior end, dorsal view. B, falcigers, anterior chaetiger. C, falcigers, midbody chaetiger. D, aciculum, posterior chaetiger. E, dorsal simple chaeta. F, falcigers, posterior chaetiger. G, ventral simple chaeta (scale bars: A = 100 μm; BG = 10 μm). with teeth about the same size and short subdistal spines (Figs. 9G, 10P). Acicula distally inflated and to hollow; up three thin acicula in each ante Martin (1984) points out the fact that in S. prolifera this feature is on highly dependent the contraction the body due to fixation. rior parapodium, decreasing in number towards DISTRIBUTION. Cosmopolitan. In Brazil, it has posterior end until only a single aciculum persists in each posterior parapodium (Fig. 9D). been identified in the states Sergipe and Alagoas (Nonato & Luna, 1970), Rio de Janeiro (Machado, 1989; Attolini, 1997), and Sao Paulo REMARKS. species is easily recognised by (Souza, 1989). the short pharynx and proventriculum, the long and dark cirri, and, mostly, the shape acicula. Our with specimens agree the descriptions already provided in other studies (San Martin, Syllis corallicola Verrill, 1900 Figs ; Licher, 1999), except for the length the Syllis (Typosyllis) corallicola Verrill, 1900: 603. pharynx, which is much shorter than in the description by Licher (1999). However, San Typosyllis corallicola; Licher, 1999: , fig. 54, see also for synonymy. 73

18 10. Fig. Syllis prolifera, scanning electron micrographs. A, anterior end, dorsal view. B, anterior end, lateral view. C, anterior end, ventral view. D, detail proboscis. E, midbody, dorsal view. FI, blades superior, intermediates, and inferior falcigers, respectively, anterior chaetigers. JL, blades superior, intermediate and inferior falcigers, respectively, midbody chaetigers. MN, blades superior and inferior falcigers, respectively, posterior chaetigers. O, dorsal simple chaetae. P, ventral simple chaeta. Q, posterior end (scale bars: A, C and E = 200 B and D = 100 μm; μm; FG, JM and 0 = 4 μm; HI, N, O, P = 3 μm; Q = 150 μm). 74

19 83 Body Our Atlantic MATERIAL. specimens, 10 from Laje de Santos, and 73 parapodia each bearing 58 (Fig. 12G). In anteri from Ilha dos Alcatrazes. Three specimens deposited at MHN (MHNBPO 07/13) and 2 specimens deposited at MNCNM (16.01/8713 and 16.01/8723). Five specimens examined under SEM not preserved. or parapodia, blades measuring, on average, 37, 28, and 18 fim; in midbody, 31, 25, and 19 im; in posterior chaetigers, 29, 22, and 16 fj.m. Dorsal and ventral simple chaetae present on posteriormost chaetigers, dorsal chaetae subidentate, with DESCRIPTION. long and robust, usually short subdistal spines (Figs. 11G, 12J); ventral simple chaetae about the same length as dorsal with pigmentation until near the end proventriculum, as dark transversal bands, leaving one chaetae, strongly bidentate, with short subdistal pair unpigmented dorsolateral circles per chaetiger (' ( variegata type'; Fig. 11G); the longest spines (Figs. 11F, 12 MN). Acicula subdistally inflated, with short oblique tip, distally rounded; 46 thin acicula in anterior chaetigers (Fig. 1 ID), specimen examined with 93 chaetigers, and measuring about 21 mm in length by 0.87 mm in 24 in each midbody chaetiger, and one single width, at the level proventriculum. aciculum in posterior body (Fig. 1 IE). Prostomium subrectangular, with four eyes in trapezoidal arrangement. Antennae long, lateral REMARKS. specimens are much shorter pair inserted on anterior margin prostomium, with about 1420 articles, median antenna originating on posterior prostomium, between posterior eyes and with about 2430 articles (Figs. 11A, 12A). Peristomium almost the same size as than those studied and by Jones (1962) Licher (1999), among others, but agree very well with the descriptions provided for the species. Jones (1962) referred to the prostomium as being posteriorly indented, but, with the aid the SEM, we following segments, with one pair dorsolateral could observe that indentation is actually on the ciliated pits on anterior margin (Figs. 12AB); superior cirri with about 1831 articles, inferior pair shorter, with 817 articles (Figs. 11A, 12A). Dorsal cirri very long throughout, containing anterior margin peristomium, due to the dorsolateral ciliated pits (Figs. 12AB), and, at least in our specimens, much less pronounced than represented in Jones (1962: 181, Licher fig. 28). dark inclusions, those chaetiger 1 much longer (1999: 119; figs. 54GI) points out to a 'knob' on than following cirri, about the same length as median antenna, or with longer, 2345 articles (Figs. 11A, 12A). From midbody, cirri alternating long, with about 1933 articles, and short, with the top the shaft posterior falcigers, however we did not observe it in Brazilian specimens, even with the aid SEM. In recent studies (Nogueira, in prep.), this 1224 articles, both types exceeding body width. species was very common in samples from the intertidal zone, in rocky shores, in greater amount Ventral cirri shorter than parapodial lobes, digitiform (Figs. than sublitorally, in corals. 12C, OP). Anal cirri longer than posterior dorsal cirri, with about 1028 articles and one ventral st papilla in between (Figs. 1 IB, DISTRIBUTION. Ocean: Mediterranean 120P). Sea, Bermuda, Caribbean Sea and Brazil. This is the second report for Brazilian waters, and the Pharynx long, extending through 715 segments (Fig. 11 A); anterior border with big tooth first for the State Sao Paulo; the previous and 56 dorsal papilla (Figs. 12CD). Proventriculum about half the length pharynx, with occurrence was detected by Augener (1931) from about 3850 rows musclecells (Fig. 11A). material collected in the State Bahia, northeastern Brazil, and described as Syllis (Typosyllis) Blades falcigers bidentate, teeth triangular, about the same size and with large and rounded to triangular gap in between (Figs. 11H, 12EI, tigrinoides (Table 1). Syilis lutea (HartmannSchröder, 1960) KL); cutting edges with fringe numerous Figs spines all along. Gradation the the length blades evident dorsoventrally and anteroposteriorly. Anterior parapodia each with 713 falcigers, midbody parapodia with 711, and posterior Typosyllis lutea HartmannSchroder, 1960: 81, figs. 3841; Licher, 1999: , figs. 17 H, 79, see also for synonymy. 75

20 Fig. 11. Syllis corallicola.a, anteriorend, dorsal view (dorsal pigmentation omitted). B, posterior end, dorsal view. C, chaetiger 14, dorsal view, showing pigmentation. D, acicula, anterior chaetiger. E, aciculum, posterior chaetiger. F, ventral simple chaeta. G, dorsal simple chaeta. H, falcigers, midbody chaetiger (scale bars: A = 300 μm; B and C = 200 μm; DH = 20 μm). 76

21 Fig. 12. nucal mediate mediate arrows organ, ventral MATERIAL. MHN Ilha 25 dos (MHN MNCNM 10 = μm; simple C = Bl'C) examined under 11 08/13) and (16.01/8721 SEM not chaetae. from Three and view. end, D, chaetigers. J, dorsal simple OP, posterior end, and P Laje de 15 μm; E = Santos specimens 2 anterior lateral and deposited specimens deposited 16.01/8722); 6 = arrows pigmentation; examined 46 view, arrows indicate proboscis. EF, 8 μm; F 4 μm; N = 14 about 4.7 mm at the at specimens in = inferior anal 2 incomplete specimens were and inter superior and (scale μm; O = length by 0.27 superior detail B, organ. 100 mm but inter posterior falcigers, cirri proventriculum, Prostomium blades between papilla HM level blades KL, nucal blades bars: A 150 = μm). in width, much at longer also collected. subrectangular to ovate, broader preserved. Body the with chaeta. and posed indicate than DESCRIPTION. dorsal detail chaetiger. G, falcigers, posterior midbody chaedger. HI, midbody 60 μm; D specimens, Alcatrazes. micrographs. A, C, anterior end, cilia. anterior falcigers, respectively, B and G from indicate falcigers, respectively, chaetigers. MN, μm; electron Syllis corallicola, scanning long and robust, without longest complete chaetigers and specimen measuring line. long, with in wide two pairs posterior eyes dis trapezoidal arrangement, nearly Antennae short, anterior border articles; median lateral prostomium, antenna pair inserted with about originating more in on 1219 poste 77

22 Fig. 13. Syllis lutea. A, anterior body, dorsal view. B, posterior end. C, falcigers, anterior chaetiger. D, acicula, anterior chaetiger. E, acicula, midbody chaetiger. F, aciculum, posterior chaetiger. G, ventral simple chaeta. H, falcigers, midbody chaetiger. I, dorsal simple chaeta. J, falcigers, posterior chaetiger (scale bars: A = 200 μm; B = 100 μm; DJ = 10 μm). 78

23 14. Fig. Syllis lutea, scanning electron micrographs. A, anterior end, dorsal view. B, anterior end, ventral view. C, falcigers, anterior chaetiger. DF, blades inferior, superior and intermediate falcigers, respectively, anterior chaetiger. G, midbody, lateralview. H, falcigers, midbody chaetiger. I, blade superior falciger, midbody chaetiger. JK, blades intermediate chaete, midbody parapodia. LM, blades inferior and superior falcigers, respectively, posterior parapodia. NO, blades intermediate chaetae, posterior parapodia. P, dorsal simple chaeta. Q, ventral simple chaeta. R, posterior end, ventral view (scale bars: A = 150 μm; B, G and R = 100 μm; C and E = 5 μm; D, F, JM, O, Q = 3 μm; H = 10 μm; I, N = 4 μm; P = 2 μm). 79

24 15 Our Long riorly, between eyes, longer than lateral antennae, bidentate, with subdistal tooth larger with 1833 articles (Figs. 13A, 14A). Palps short, and subdistal bristles (Figs. 13G, 14Q). Acicula subdistally broader than long, completely separated from inflated, with short and oblique tip; up to four each other (Figs. 14AB). Peristomium dorsally reduced, superior tentacular cirri about pair twice as long as inferior pair (Figs. 14AB); superior cirri with 1528 articles, inferior pair with 8 thin acicula in each anterior chaetiger (Fig. 13D), two in midbody, thicker (Fig. 13E), and solitary aciculum in each posterior chaetiger (Fig. 13F). 13 articles. REMARKS. specimens agree with the Dorsal cirri chaetiger 1 longer than the fol redescription by Licher (1999), except for being lowing cirri, with 1925 articles (Figs. 13A, 14A). slightly smaller in all measurements. Cirri long and thin throughout, containing iridescent inclusions. From midbody, alternating long DISTRIBUTION. Circumtropical. First report for and short dorsal cirri, the longest exceeding body the Brazilian coast. width and with about twice the length short cirri (Fig. 14G). Ventral cirri thin, up to the same Syllis glandulata n. sp. length parapodial lobes, broader basally in Figs anterior chaetigers (Fig. 14B), more elongated and tapering from midbody onwards (Fig 14H). MATERIAL. specimens, which 13 from Laje de Anal cirri longer than posterior dorsal cirri, with about 17 articles and one st in papilla between (Figs. 13B, 14R). Pharynx long, extending through 513 segments, with big tooth located on anterior margin Santos (24 19'S, 46 1 and 2 from Ilha l'w) dos Alcatrazes (26 06'S, 45 42'W). Type series: 6 4 specimens, from Laje de Santos and 2 from Ilha dos Alcatrazes; holotype and paratype 1 at deposited MHN (holotype: MHNBPO 77/0; paratype 1: MHNBPO 77/1); paratypes 34 deposited at MNCNM (16.01/8719 and 16.01/8720); paratypes 56 (Fig. 13A). Proventriculum occupying 513 seg deposited at FSBC I (66261 and 66263). Holotype and ments, with about 3258 musclecells rows (Fig. 13A). Blades falcigers bidentate, cutting edges with paratypes 1, 4, and 5 from Laje de Santos; paratypes 2 and 3 from Ilha dos Alcatrazes. Four examined specimens under SEM not preserved. spines thicker and longer basally, progressively diminishing in size distally; a few subdistal bristles DESCRIPTION. species, without remarkable extend until the level subdistal tooth, or even color patterns; holotype complete, in good condi surpass it (Figs. 13C, H,J, 14CF, HP). Blades tions, measuring 15.4 mm in length, 0.33 mm in falcigers with distinct dorsoventral gradation in width, with 106 chaetigers. length (Figs. 13C, H, J, 14C, H), but without anteroposterior gradation. About 10 compound Prostomium subpentagonal to ovalate, with four chaetae in each anterior parapodium, 56 from eyes in trapezoidal arrangement and three antennae about the same size, or median antenna midbody. In anterior chaetigers, blades fal slightly longer, with 1014broad articles (1014 in cigers with teeth about the same size, distal tooth holotype); lateral antennae inserted in front hooked (Figs. 13C, 14CF); blades measuring, on anterior pair eyes, median antennae between average, about 32, 21, 15 itu. From midbody posterior pair eyes, near posterior end pros onwards, subdistal tooth larger, strongly triangular, more evident on inferior chaetae (Figs. 13H, tomium (Figs. 15A, 16A). Palps longer than prostomium, separate for most the extension, J, 14IP); blades, on average, measuring about 32, 23, 18 (am, in median chaetigers, and 30, 23, 18 except basally. Peristomium conspicuous, shorter than following dorsal segments; pair tentacular fim posteriorly. Dorsal simple chaetae present cirri about the same size as antennae, or longer, from posterior midbody, about the same length as ventral pair shorter (Fig. 15A). the shafts superior falcigers, or slighdy shorter, subidentate, with short subdistal spinulation (Figs. Dorsal cirri with broad articles, especially basally, posteriorly tapering; those from chaetiger 1 131, 14P). Ventral simple chaetae present in posterior chaetigers, thicker than dorsal chaetae, and much longer than antennae, and those from following chaetigers, with 1123 articles (19 shorter than shafts inferior falcigers; strongly in holotype; Figs. 15A, 16A); from proventricular level, 80

25 Fig. 15. Syllis glandulata n. sp. A, holotype, anterior end, dorsal view. B, holotype, posterior end. C, acicula, anterior chaetiger. D, acicula, midbody chaetiger. E, aciculum, posterior chatiger. F, holotype, parapodium, chaetiger 80. G, ventral simple chaeta. H, falcigers, anterior chaetiger, from left to right, inferior to superior chaetae. I, falcigers, midbody chaetiger. J, dorsal simple chaeta. K, falcigers, posterior chaetiger (scale bars: A = 92 μm; B = 200 μm; CE and GK = 10 μm; F = 50 μm). 81

26 Fig. 16. Syllis glandulata n. sp., scanning electron micrographs. A, anterior body, dorsal view. B, falcigers, anterior chaetiger. C, superior falcigers, anterior chaetiger. D, falcigers, midbody chaetiger. E, falcigers, posterior chaetiger. F, dorsal simple chaeta. G, ventral simple chaeta. H, detail the tip dorsal simple chaeta. I, detail ventral simple chaeta (scale bars: A = 150 μm; B = 30 μm; C = 10 μm; D = 20 μm; E, F = 8 μm; G = 6 μm; H and I = 3 μm). dorsal cirri short, distally tapering (Fig. 15F), with 512 articles (812 in holotype), not alternating in 15H, I, K, 16BE), although some specimens have posterior blades remarkably shorter; blades bidentate, with conspicuous spines, teeth length. Pygidium semicircular, with two long anal about the same size and with wide gap in cirri, with 816 articles (16 in holotype), and short between (Figs. 15H, I, K, 16BE); compound papillae in between (Fig. 15B). chaetae similar throughout. Blades falcigers Pharynx extending through 510 segments (Fig. measuring, on average, in anterior parapodia, 32 (xm, 22 am and 15 (am; in midbody, 34 _im, 24 am and 15 jam; and in posterior parapodia, 30 (im, 15A), bearing a big tooth on anterior border, surroundedby a crown st papillae; proventriculum usually shorter than pharynx, with about fim and 15 (am (holotype with blades measur 34 broad rows musclecells (holotype with ing 33 jim, 23 pi and 16 am in anterior pharynx extending for 10 segments, proventriculum for 6 segments, with 33 rows musclecells). chaetigers, 33 (am, 25 (un and 17 (am in midbody, and 32 (am, 23 jam and 15 (am in posterior chaetigers). Dorsal simple chaetae relatively long, Parapodia along the whole body bearing conspicuous dorsal glands, with big and rounded with subdistal spines, subdistal tooth resembling cells containing granular material (Fig. 15AB, F); an enlarged spine (Figs. articles dorsal cirri with inclusions similar 15J, 16F, H); ventral simple chaetae shorter, curved, with subdistal spines, material. Parapodial lobes triangular bidentate, with teeth about the same size (Figs. to subrectangular, with 58 compound chaetae anteriorly (Fig. 16B), 67 in midbody (Fig. 16D), and 35 15G, 16G, I); both types simple chaetae present only in posteriormost chaetigers. Anterior posteriorly; strong gradation size the blades parapodia with 34 short and thin acicula, subdistally inflated, with acuminate, oblique tip (Fig. evident dorsoventrally, but not anteroposteriorly (Figs. 15C); median parapodia each with two acicula, 82

27 The Short acicula rounded. about the same shape as anterior ones, but thicker (Fig. 15D), posteriormost parapodia with single aciculum (Fig. VARIATION. 15E). Specimens with parapodial glands ETYMOLOGY. specific name glandulata refers to the parapodial glands, present in all chaetigers most the adults this species. poorly developed, or even inconspicuous, were also observed, and they were distinguished from S. pseudoarmillaris n. sp. (see below) by having Syllis pseudoarmillaris n. sp. Figs longer and gently tapering dorsal cirri, conspicuous dorsoventral gradation the length blades falcigers throughout the body, and by the number acicula in anterior parapodia. REMARKS. Syllis glandulata n. sp. fits in a group MATERIAL. Syllis pseudoarmillaris: State Sao Paulo: 40 specimens, 29 from Laje de Santos and 11 from Ilha dos Alcatrazes. Type series: 7 specimens, 3 from Laje de Santos, including the holotype, and 4 from Ilha dos Alcatrazes; holotype and paratypes 12 deposited at MHN (holotype: MHN BPO 09/0; paratypes: MHNBPO 09/12); paratypes 34 Syllis called by Licher (1999) as' Typosyllis armillaris complex', characterized by falcigers short, bidentate, with secondary tooth shorter than distal one, and posterior acicula subdistally inflated, with oblique acuminate tip. This group comprises, according to Licher (1999) 10 species, besides S. glandulata n. sp.: S. armillaris (Muller, 1776); S. cras deposited at MNCNM (16.01/8858, both specimens together); paratypes 56 deposited at FSBC I (66260 and 66262). Holotype and paratypes 2 and 5 from Laje de Santos, paratypes 1, 3, 4,and 6 from Ilha dos Alcatrazes. Four specimens examined under SEM not preserved. Syllis armillaris: Spain: 20 specimens from the Mediterranean (Isla de Alboran, Almeria) and 1 specimen from the Atlantic (Cabo de Finisterre, A Coruna). sicirrata (Treadwell, 1925); S. ehlersioides (Marenzeller, 1890); S. hyalina Grube, 1863; S. DESCRIPTION. species, without remark krohni Ehlers, 1864; S. neglecta Grube, 1870; S. stellaepolaris HartmannSchroder, 1993; valida S. Grube, 1857, and S. violacea Grube, 1870; although not included by Licher (1999), we believe S. gracilis and S. magellanica also belong to this group. Syllis glandulata n. sp. differs from all these species because the parapodial glands, a able color patterns; holotype complete, in good condition, measuring 10.9 mm in length, 0.32 mm in width, at the level proventriculum, with 96 chaetigers. Prostomium subpentagonal to ovalate, with four eyes in wide trapezoidal arrangement and three antennae, which the median antenna is longer, character unknown in all other species this with 1321 broad articles (21 in holotype); lateral genus. antennae inserted in front anterior pair The species most similar to S. glandulata n. sp. eyes, with 812 articles (10 in holotype), median are S. armillaris and S. hyalina. S. glandulata n. sp. antenna between posterior pair eyes, differs from S. armillaris, besides for the absence near posterior end prostomium (Figs. 17A, 18A). Palps parapodial glands in the latter, because S. armil longer than prostomium, separate for most laris has: (1) blades midbody falcigers minutely bidentate, with subdistal tooth very reduced, resembling a while in spine, S. glandulata n. sp. both teeth are conspicuous, with nearly the same their length, except basally. Peristomium conspicuous, shorter than following segments; dorsal pair tentacular cirri shorter than median antenna, ventral pair usually about the same size as lateral size; (2) dorsal cirri abruptly tapering near tip, antennae (Figs. 17A, 18AB). instead the gently tapering condition present in S. glandulata n. sp.; (3) tips while in S. glandulata n. sp. acicula rounded, they are acuminate. Dorsal cirri with broad articles, especially basally, distally tapering; those from chaetiger 1 usually with the same length as median antennae, Syllis hyalina differs from S. glandulata n. sp., besides the absence parapodial glands, much longer than those from the following chaetigers, with 923 articles (22 in holotype; because it has: (1) more chaetae per parapodium; Figs. 17A, 18A); from proventricular level, dorsal (2) falcigers bidentate, with subdistal tooth conspicuous, but much shorter than distal tooth; (3) dorsal cirri abruptly tapering near tip; (4) tips cirri short, broader at midlength and abruptly tapering (Fig. 17B); cirri containing iridescent inclusions, initially with eight articles, shortly pos 83

28 Fig. 17. Syllis pseudoarmillaris n. sp. A, holotype, anterior end, dorsal view. B, holotype, parapodium, chaetiger 21. C, holotype, posterior end. D, acicula, anterior chaetiger. E, falcigers, anterior parapodium. F, aciculum, posterior chaetiger. G, dor sal simple chaeta. H, falcigers, midbody. I, ventral simple chaeta (scale bars: A = 200 μm; B = 25 μm; C = 100 μm; DI = 10 μm). 84

29 terior to proventriculum, decreasing in number articles towards posterior body; posterior cirri with 35 articles (Fig. I7C). Pygidium semicircular, with two anal cirri, with 716 (16 in holotype) abruptly tapering, and blades falcigers in midbody chaetigers, strongly triangular and bidentate, nearly without dorsoventral gradation in length the blades. The species most similar to articles, and short papilla in between (Fig. 17C). S. pseudoarmillaris n. sp. is S. armillaris (Miiller, 1776), but they are can be distinguished because S. armillaris is larger, with dorsal cirri slightly longer and with blades midbody falcigers unidentate or subidentate (San Martin, 1984; Pharynx extending through 59 segments (Fig. 17A), usually containing red pigmentation inside, bearing one big tooth at the anterior border, surrounded by a crown st papillae; proventriculum shorter, with around 4050 rows muscle Licher, 1999), while in S. pseudoarmillaris n. sp. the cells (holotype with pharynx extending for 6.5 secondary tooth falcigers is well marked. segments, proventriculum for 4 segments, with 42 rows musclecells). Parapodial lobes to triangular subrectangular, Syllis hyalina is another very similar species, however it has dorsal cirri than S. armillaris, it has more chaetae per parapodium, even longer and with 68 compound chaetae anteriorly (Fig. 18C), falcigers with blades much longer, 2550 (im in 35 from midbody (Figs. 18D, F), last chaetigers with 25 dorsal and falcigers, plus simple ventral chaetae (Fig. 18F); strong gradation in the size the blades evident only in anterior chaetigers anterior body, 2025 jam in and midbody, 2530 (am in posterior body (Licher, 1999). Finally, S. pseudoarmillaris n. sp. shares similar morphology dorsal cirri with S. glandulata n. (Figs. 17E, 18C), very slight from midbody (Figs. 17H, 18D, F); blades bidentate, with spines on the sp., described above, but the latter has cirri slightly longer and gently tapering towards tip, dorsoventral gradation in length the blades cutting edges, and subdistal tooth always conspicuous and only slightly shorter than distal tooth, with wide gap in between (Figs. 17E, H, 18CF); falcigers with proportionally broader bases at falcigers well marked throughout, and more acicula per anterior parapodium, besides the parapodial glands, characteristic for S. glandulata n. sp. midbody (Figs. 18D, E). Blades falcigers mea Rullier & Amoureux (1979) identified S. armil suring, on average, in anterior parapodia, 29 (am, 20 am and 14 (am; in midbody, 19 jam, 17 (am and 13 (am; and in posterior parapodia, measuring 16 laris and S. hyalina (as Typosyllis armillaris and T. in hyalina, respectively) northeastern Brazil (Table 1), however, in their brief very diagnosis for both species, the authors said the identifications were jam, 15 jam and 12 (am (holotype with blades measuring 31 chaetigers, (im, 23 (am and 15 jam in anterior 18 jam, 16 (am and 13 am in midbody, based mostly in the shape dorsal cirri, short and 17 (am, 15 jam and 11 (am in posterior and fusiform in both taxa, and in the morphology falcigers, clearly bidentate in S. hyalina and chaetigers). Dorsal simple chaetae relatively long, unidententate in S. armillaris, with some blades with subdistal spines, subdistal tooth resembling almost fused with the shaft (Rullier & Amoureux, an enlarged spine (Figs. 17G, 18FG); ventral simple chaetae shorter, curved, with subdistal spines, 1979: 161). We suspect none those species really occurs in Brazil. In the case S. hyalina, the bidentate, with teeth about the same size (Figs. specimens examined by Rullier & Amoureux 171, 18F, FI); dorsal and ventral simple chaetae (1979) are possibly S. glandulata n. sp., while those present only on posterior chaetigers, usually identified as S. armillaris cannot belong to this appearing in the same chaetiger. Anterior para species, due to the fusion between shafts and podia with 23 short and thin acicula, subdistally blades mentioned by the authors. Instead, they inflated, with acuminate, oblique tip (Fig. 17D); probably belong to the taxon named by Ben median parapodia each with 12 acicula, about Eliahu (1977: 9) as S. armillaris type 3', which is the same shape as anterior ones, but thicker, pos possibly S. magellanica, a very common species in terior parapodia with single aciculum (Fig. 17F). the State Sao Paulo (Nogueira, pers. observ.). REMARKS. Syllis pseudoarmillaris n. sp. is charac ETYMOLOGY The specific name pseudoarmillaris terized by the morphology dorsal cirri from refers to the great similarity between this species midbody, with broad articles at midlength, then and S. armillaris. 85

30 Brazil: n. sp., scanning electron micrographs. A, anterior end, dorsal view. B, anterior ventral Fig. 18. Syllis pseudoarmillaris end, view. C, falcigers, anterior chaetiger. D, compound chaetae, midbody parapodium. E, detail the blade superior falciger. F, chaetae, posterior parapodium, large indicates dorsal arrow simple chaeta, small arrow indicates ventral simple chaeta. G, dorsal simple chaeta. H, ventral simple chaeta (scale bars: AB = 150 μm; C, GH = 10 D = 20 E = 5 F = 15 μm; μm; μm; μm). Syllis tyrrhena (Licher & Kuper, 1998) n. comb. chaetigers. Prostomium subpentagonal to ovalate, broader Figs than long (Figs. 19A, 20A), with two pairs eyes disposed in trapezoidal arrangement; one pair Typosyllis tyrrhena Licher & Kuper, 1998: , figs. 14; Licher, 1999: 140, , fig. 62. anterior eyespots may be present in short specimens. Antennae long, the lateral pair with about 713 articles, inserted in the anterior border MATERIAL specimens, from Laje de Santos and 21 from Ilha dos Alcatrazes. Three specimens deposited at MHN (MHNBPO and 12/13) 2 specimens depositedat MNCNM (16.01/8714 and 16.01/8715). Four specimens examined under SEM not preserved. Spain: 2 specimens, prostomium, and median antenna originated more posteriorly, between posterior and eyes, about twice the length lateral pair in all specimens with 1533 analysed, articles (Figs. 19A, MediterraneanSea, Isla de Alboran, Almeria, deposited at 20A). Palps longer than prostomium, separated MNCNM (16.01/6701 and 16.01/6702). for most their extension (Fig. 19A). Peristomium distinctly reduced dorsally (Figs. DESCRIPTION. Body very elongated and thin, with long segments, lacking special pigmentation 19A, 20A), superior pair tentacular cirri about the same length as lateral antennae, or slightly longer, and about twice the length inferior patterns; the longest specimen examined measuring about 7 mm in length by 0.1 mm in width, at the level proventriculum, and counting on 49 pair; superior pair with about 817 articles, inferior pair with 59 articles (Figs. 19A, 20B). 86

31 Fig. 19. Syllis tyrrhena. A, anterior end, dorsal view. B, posterior end. C, falcigers, anterior chaetiger. D, aciculum, anterior chaetiger. E, falcigers, midbody chaetiger. F, dorsal simple chaeta. G, falcigers, posterior chaetiger. H, ventral simple chaeta. I, aciculum, posterior body (scale bars: AB = 140 = μm; CI 10 μm). 87

32 Fig. 20. Syllis tyrrhena, scanning stomial cirri detail missing). C, anterior chaetiger. F, ventral electron micrographs. A, anterior end, dorsal view. B, anterior end, lateral view figure A, showing proboscis. D, falcigers, simple chaeta. G, midbody, lateral anterior chaetiger. E, blades (superior peri superior falcigers, H, falcigers, posterior midbody chaetiger. I, view. detail posterior midbody chaetiger, showing intermediate and inferior falcigers. J, superior falcigers, midbody chaetiger. K, poste (scale bars: A = 60 Dorsal cirri μm; following cirri, superior falciger, posterior chaetiger. M, B, G, K = chaetiger with median antenna, or 1 about μm; C 100 much the slightly = μm; D = longer than same shorter 15 length (Figs. 19A, 20AB). From midbody, dorsal cirri short throughout, in distally tapering, posterior body in with 68 articles, some even specimens (Figs. 20G, K). Ventral cirri shorter than parapodial digitiform than 88 (Figs. posterior 20B, dorsal G, K). cirri, shorter Anal cirri lobes, longer with about 69 articles 10 inferior chaeta, EF, J, μm; and one LN st Pharynx about seven thin tooth C). = 3 μm; papilla long H 5 μm; a and thin, the anterior crown I in between dorsal = (Figs. chaeta 19B, 20K). extending through margin, which is short papillae (Figs. much pharynx, occupying and with about 2537 simple μm). 19A), with sharp and Proventriculum broader than 19A). = chaetigers (Fig. at rounded by 20A, midbody chaetiger. N, 4 rior end, dorsal view. L, rows shorter two sur 19A, and chaetigers musclecells (Fig.

33 Our Brazil: Atlantic Falcigers with bidentate blades and short spines DISTRIBUTION. Ocean: Mediterranean all along the cutting edges; teeth sharp, subdistal Sea (Tyrrhenian Sea and Isla de Alboran, Spain), one shorter (Figs. 19C, E, G, 20DE, HJ, LM). and State Sao Paulo. This is the first report for Length the blades graded in both directions, dorsoventral and anteroposterior (Figs. 19C, E, Brazilian waters and for beyond the Mediterranean Sea. G, 20D, H), although less accentuated dorsoventrally from midbody (Fig. 20H). In anterior parapodia, blades, on average, measuring about 25, Syllis truncata Haswell, 1920 Fig and 11 (am; in midbody, 18, 14 and 10 jam; in posterior chaetigers, 14, 11 and 9 Dorsal pm. simple chaetae usually from present midbody chaetigers, immediately posterior to proventricu Syllis (Typosyllis) truncata Haswell, 1920: 9496, PI. X, figs Typosyllis truncata; Licher, 1999: , fig. 71. lum in some specimens, distally truncate, with Typosyllis subterranea HartmannSchroder, 1962: short subdistal spines (Figs. 19F, 20N); ventral 9395, figs simple chaetae present only in last chaetigers, shorter and thinner than dorsal chaetae, distally MATERIAL. 3 specimens, all from Ilha dos curved, bidentate, and with very short subdistal spines (Figs. 19H, 20F). One to two acicula per parapodium, distally curved at right angle, and Alcatrazes. Two specimens deposited at MHN (MHNBPO 13/12), and 1 deposited at MNCNM (16.01/8724). Panama: 2 specimens, from Coiba Island (Eastern Pacific). with very short and sharp tip (Fig. 19D); single aciculum from midbody in all specimens, thicker (Figs. 191). DESCRIFI'ION. Body robust, without distinct patterns pigmentation; the largest complete specimen with 83 chaetigers and measuring about 5 REMARKS. specimens differ from the mm in length by 0.22 mm in width, at the level descriptions by Licher & and Licher Kuper (1998) proventriculum. (1999) in some aspects. Prostomium subrectangular, with two pairs First all, in our specimens dorsal cirri chaetiger 1 are much longer eyes in trapezoidal arrangement, and one pair than following cirri, while in the descriptions provided by those authors the difference is very slight; in contrast, dorsal cirri from midbody and posterior parts the body are much shorter in anterior eyespots; antennae short and relatively thick, the lateral pair originating on the anterior border prostomium, with 1213 articles, median antenna inserted in the center prostomium, Brazilian specimens, since Licher (1999) described midbody cirri with 1316 articles, and about the same length, with 1314articles (Fig. 21 A). Palps short, completely separated. posterior ones with 1014 articles. Additionally, Peristomium slightly shorter than following seg the blades falcigers are much shorter in the specimens described by Licher & Kuper (1998) and Licher (1999), 916 j_un on anterior body, 69 jam on midbody, and 612 (xm on posterior chaetigers, according to Licher (1999). However, we consider the general aspect the body, very thin and with and elongate segments, the small number chaetae per parapodium, as well as the morphology the blades falcigers, acicula, and dorsal and ventral simple chaetae, as the most characteristic features this species, and our specimens fit with the descriptions provided. Moreover, we could see no morphological ments, superior pair peristomial cirri almost twice as long as inferior pair, about the same length as antennae; superior pair with 1216 articles, inferior pair with 78 articles (Fig. 21A). Dorsal cirri thick, containing many iridescent inclusions. Those 1 chaetiger much longer than following cirri, with 1420 articles (Fig. 21 A); from midbody, cirri not evidently alternating long and short, about the same length as body width, or slightly longer, and counting on 913 articles. Ventral cirri digitiform, about the same length as parapodial lobes. Anal cirri with about 1114 articles. differences between our specimens collected in Spain, even in regard length the blades. and material to cirri and Pharynx long, extending for 810 segments, with big tooth on the anterior margin. Proventriculum broader than pharynx, extending 89

34 Fig. 21. Syllis truncata. A, anterior end, dorsal view. B, acicula, anterior chaetiger. C, falcigers, anterior chaetiger. D, falcigers, midbody chaetiger. E, aciculum, midbody. F, dorsal simple chaeta. G, falcigers, posterior chaetiger. H, ventral simple chaeta (scale bars: A = 130 μm; BH = 10 μm). 90

35 Our The through 56 chaetigers, and with about 2030 rows musclecells (Fig. 21A). emend the current diagnosis Syllis, considering Typosyllis as a junior synonym. Furthermore, the Falcigers with bidentate blades, subdistal tooth geographic range several species is enlarged, to shorter; blades with short spines all along include Brazil and, in some cases, southern the cutting edge (Figs. 21C, D, G). Blades falcigers Atlantic. with strong dorsoventral gradation in length, but In regard to the extension the geographical not anteroposterior (Fig. 21C, D, G). Blades superior falcigers measuring about 2729 im, distribution the species, very unlikely that species we agree that it seems from Pacific Ocean, or even from northern Atlantic can also be present in the southern Atlantic. However, after careful intermediate about 1522 jam, and inferior falcigers about 912 (im; blades posteriormost falcigers shorter, especially the superior chaetae. examination under stereo, light, and scanning Simple chaetae only present in posteriormost electron microscope, we could find no significant chaetigers; dorsal simple chaetae distally truncate differences between our specimens and the (Fig. 2IF), ventral simple chaetae thinner, distally descriptions provided in the literature (San Martin, 1984, 1992; Licher, 1999). curved and bidentate (Fig. 21H). Anterior parapodia each with two very thin acicula, in all specimens, subdistally curved at right angle, with ACKNOWLEDGEMENTS short and sharp tip (Fig. 21B); from midbody, single aciculum per parapodium, thicker (Fig. 2IE). This work was supported by the State Sao Paulo Research Foundation (FAPESP), within the REMARKS. specimens differ from those BIOTA/FAPESP Biodiversity Virtual described by Licher (1999) by having: (1) antennae and cirri shorter throughout, with fewer articles; (2) from midbody, dorsal cirri with nearly uniform length, not evidentiy alternating long and short cirri; (3) fewer chaetae per parapodium; InstituteProgram ( and projects 01/ and 01 /036350, and by CNPq (Conselho Nacional de Desenvolvimento Cientifico e Tecnologico). Besides, we extend our gratitude to the Departamento de IB Zoologia, and (4) two acicula in anterior parapodia, instead UNICAMP, especially Dr Cecilia Amaral, for one single aciculum throughout the whole body. However, our specimens are also much assistance and support; to the coordinates the Postgraduate Course IBUSP while this study was being conducted, Drs Erika Schlenz and shorter than those Licher (1999) and it is possible that these differences are due to size. Fabio Lang da Silveira; to Dr Erika Schlenz, The analysis species from the Pacific (Coiba again, for the translation several texts from German; to Maria Teresa Aguado, from LAM, Island, Pacific side Panama), revealed no significant differences to our specimens. for drawing figure 15A; to Maria Capa, from The reduced amount specimens observed, UAM, for kindly loaning us specimens S. trun and their poor state preservation, prevented us cata from Panama; to Drs Tom Perkins, Frank from examining this species under SEM. Licher, and two anonymous referees for the very useful comments and suggestions; to the staff DISTRIBUTION. Pacific Ocean: Australia and the Laboratorio de Microscopia Eletronica, IB Chile; Adantic Ocean: State Sao Paulo. This is USP, technician Enio Mattos, and Laboratorio de the first for this report species in Brazil, and, for one doubtful identification in the except Mediterranean Sea, the first occurrence in the Atlantic. Microscopia Eletronica, IBUNICAMP, technicians Adriane Sprogis Lima; to Fausto Pires de Campos and Antonia Maria de and Tatiana Neves the IF (Instituto Florestal, Brazil) for permitting us to collect on both islands, which are CONCLUSION protected areas. During the study various species Syllis found in living colonies Mussismilia hispida, we discovered two new species, and provided more data to 91

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