Deep sea Syllidae (Annelida, Phyllodocida) from Southwestern Atlantic

Joao Miguel Matos Nogueira

We describe herein ten species of Syllidae from the Southern Brazil continental slope (700–2000 m deep), belonging to the genera Anguillosyllis, Exogone, Parexogone, Prosphaerosyllis, Sphaerosyllis and Syllis. Out of those, three species are new to science and six are formally reported for Brazil for the first time. Some synonymies are proposed and a taxonomic key for all described species of the genus Anguillosyllis is provided.

Zootaxa 4221 (4): 401–430 http://www.mapress.com/j/zt/ ISSN 1175-5326 (print edition) Article Copyright © 2017 Magnolia Press ZOOTAXA ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4221.4.1 http://zoobank.org/urn:lsid:zoobank.org:pub:F353EEB2-882D-464B-A2CC-F40606B58EDC Deep sea Syllidae (Annelida, Phyllodocida) from Southwestern Atlantic RÔMULO BARROSO1,5, PAULO CESAR DE PAIVA2, JOÃO MIGUEL DE MATOS NOGUEIRA3 & MARCELO VERONESI FUKUDA4 1 Departamento de Biologia, Pontifícia Universidade Católica do Rio de Janeiro, Av. Marquês de São Vicente, 255, Gávea, 22451-900, Rio de Janeiro, RJ, Brazil. 2 Laboratório de Polychaeta, Depto de Zoologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Avenida Carlos Chagas Filho 373, Bloco A, 21941-902, Ilha do Fundão, Rio de Janeiro, RJ, Brazil 3 Laboratório de Poliquetologia (LaPol), Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, travessa 14, n. 101, 05508-090, São Paulo, SP, Brazil 4 Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Juiz de Fora, Rua José Lourenço Kelmer, s/n, Martelos, 36036-330, Juiz de Fora, MG, Brazil 5 Corresponding author. E-mail: barroso.romulo@gmail.com Abstract We describe herein ten species of Syllidae from the Southern Brazil continental slope (700–2000 m deep), belonging to the genera Anguillosyllis, Exogone, Parexogone, Prosphaerosyllis, Sphaerosyllis and Syllis. Out of those, three species are new to science and six are formally reported for Brazil for the first time. Some synonymies are proposed and a taxonomic key for all described species of the genus Anguillosyllis is provided. Key words: Polychaeta, Anguillosyllis, Exogone, Parexogone, Prosphaerosyllis, Sphaerosyllis, Syllis Introduction Syllidae is one of the most diverse families of polychaetes, currently with approximately 700 species arranged into 74 genera (San Martín & Aguado 2014), found in several marine habitats at all depths and latitudes, although more diverse and abundant in shallow waters when compared with deep environments (Desbruyères & Segonzac 1997; San Martín 2003), in which syllids are frequently present although found in lower diversity and abundance (Böggemann & Purschke 2005; Lucas et al. 2012). However, as our knowledge about the deep-sea in general can still be considered as in its beginning, this apparent lower diversity may be shown, in the future, to be artefactual given the inadequacy of the sampling methods currently in use, since syllids have relatively small, fragile bodies (San Martín 2004). Thus, the list of about 90 species of syllids currently known to occur in the deep-sea (cf. Gil & Musco 2015) is expected to increase considerably as this large and plural environment is more thoroughly investigated. About 150 species of Syllidae have been reported from the Brazilian coast so far. Most of these records, however, come from shallow waters, mostly from the intertidal zone, and only a few species were collected below 200 m deep: Brevicirrosyllis cf. mariae (San Martín & Hutchings, 2006) (Fukuda et al. 2015); Eusyllis assimilis Marenzeller, 1875 (Fukuda et al. 2015); Exogone anomalochaeta Benham, 1921 (Barroso 2011); E. arenosa Perkins, 1981 (Attolini 2002); Odontosyllis cf. fulgurans (Audouin & Milne-Edwards 1833) (Fukuda & Nogueira 2006; Fukuda et al. 2013); Opisthodonta morena Langerhans, 1879 (Fukuda et al. 2015); O. russelli San Martín, López & Aguado, 2009 (Fukuda et al. 2015); Parexogone wolfi (San Martín, 1991) (Fukuda 2010; Barroso 2011); Perkinsyllis longisetosa (Hartmann-Schröder, 1965) (Fukuda & Nogueira 2013); Prosphaerosyllis isabellae (Nogueira, San Martín & Amaral, 2001) (Nogueira et al. 2001; Fukuda et al. 2009, Barroso 2011); Syllis aciculigrossa (San Martin, 1990) (Barroso 2011), S. sclerolaema Ehlers, 1901 (Fukuda 2010); and S. variegata Grube, 1860 (Attolini 2002). Accepted by A. Nygren: 23 Nov. 2016; published: 19 Jan. 2017 401 We found ten species of the Syllidae among material collected from the southeastern Brazilian continental slope, southwestern Atlantic, between 700–2000 m deep: Anguillosyllis palpata (Hartman, 1967), A. lanai sp. nov., Exogone anomalochaeta, Parexogone campoyi (San Martín, Ceberio & Aguirrezabalaga, 1996), P. minuscula (Hartman, 1953), P. wolfi, Prosphaerosyllis isabellae, Sphaerosyllis ceciliae sp. nov., Sphaerosyllis monicae sp. nov., and Syllis aciculigrossa. Out of those, three species are new to science and six are formally reported to Brazilian waters for the first time; some synonymies are also proposed. Material and methods The material analyzed for the present study came from 3 independent projects carried out off southeastern Brazil, all coordinated by the research center CENPES/PETROBRAS. Two of these projects sampled on the Campos Basin: ‘Campos Basin Deep-sea Environmental Project' (‘Oceanprof’—from 400 to 2.400 m depth), and ´Environmental heterogeneity in the Campos Basin' (‘HABITATS’—from 400 to 3.000 m depth); the last project, 'Environmental heterogeneity in the Espírito Santo Basin and northern region of the Campos Basin' (‘AMBES’— from 50 to 3.000 m depth), collected mostly in the Espírito Santo Basin, but also in the northern part of the Campos Basin. For all these projects, the sediment was collected with a 0.25m2 Box-corer (Ocean Instruments, Unsel Spade Corer MK I), samples were separate in 3 strata (0–2 cm, 2–5 cm and 5–10 cm) and sieved with 0.3–0.5 mm meshes. The specimens found were immediately preserved in 10% formalin and later rinsed in fresh water and stored in 70% ethanol. Identifications were based on morphological characters. Length of the specimens was measured from the tip of palps to the tip of pygidium, excluding anal cirri, or to the end of the fragment, in cases of incomplete specimens; width was measured at the proventricle level, excluding parapodia; measurements of the blades of compound chaetae are given in dorso-ventral sequence. Specimens were examined under stereomicroscope, compound light microscope and scanning electron microscope (SEM). For the latter, specimens were first dehydrated in a series of progressively increasing concentrations of ethanol (70–100%), critical point dried, covered with ~25 nm of gold and then examined and photographed at the Laboratório de Microscopia Eletrônica (IB/USP) and Laboratório de Microscopia Eletrônica (NEMA/PUC-Rio). Abbreviations for museum names are as follows: AM BMNH MNRJP MZUSP USNM ZUEC ZMH The Australian Museum, Sydney, Australia Natural History Museum, London, UK Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil National Museum of Natural History, Smithsonian Institution, Washington DC, USA Museu de Zoologia, Universidade Estadual de Campinas, Campinas, Brazil Zoologisches Museum der Universität Hamburg, Hamburg, Germany Results Family Syllidae Grube, 1850 Type genus. Syllis Savigny in Lamarck, 1818. Diagnosis. Small sized polychaetes, with nearly cylindrical to strongly dorso-ventrally flattened bodies. Palps well developed, from completely free from each other to completely fused. Prostomium typically with 2 pairs of eyes, sometimes also with 1 pair of anterior eyespots, and 3 antennae. Peristomium with 1–2 pairs of peristomial cirri. Smooth to articulated dorsal cirri, usually longer on anterior chaetigers; ventral cirri usually present, shorter than dorsal cirri. Compound chaetae as falcigers; modified falcigers frequently present as spiniger-like chaetae or, more 402 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. rarely, secondarily simple due to loss of blade or fusion to shaft; falciger blades unidentate or bidentate, distal teeth of variable sizes. Dorsal and ventral simple chaetae usually present, at least on posterior chaetigers. One to many aciculae of variable morphologies per parapodium, typically progressively fewer and stouter posteriorwards. Pygidium with 1 pair of anal cirri, usually longer than posterior dorsal cirri. Straight to sinuous pharynx on anterior segments, usually with anterior margin surrounded by papillae and central tooth and/or trepan; some species with unarmed pharynx; pharynx followed by muscular proventricle of variable size. Reproduction by epigamy or schizogamy, sometimes with incubation of eggs and embryos; a few species viviparous. Subfamily Exogoninae Langerhans, 1879 Genus Exogone Ørsted, 1845 Type species. Exogone naidina Ørsted, 1845. Diagnosis. Body short, slender. Palps well developed, completely or almost completely fused. Prostomium with 2 pairs of eyes, sometimes also with 1 pair of anterior eyespots, and 3 antennae. Peristomium with 1 pair of short, papilliform or ovate peristomial cirri. Compound chaetae as falcigers and, frequently, spiniger-like chaetae; falciger blades usually bidentate, subdistal tooth larger, blades partially fused to shafts or absent in some species; shafts of spiniger-like chaetae with triangular processes in some species. Dorsal simple chaetae present from anterior chaetigers, ventral simple chaetae usually only on posterior chaetigers. Aciculae commonly inflated distally, frequently hollow, with slightly curved tip. Pygidium with 1 pair of anal cirri, usually longer than dorsal cirri. Reproduction by epigamy, with ventral incubation of eggs and embryos. Remarks. The genus Exogone was revised by San Martín (1991) who established 3 sub-genera: Parexogone Mesnil & Caullery, 1918, Exogone Ørsted, 1845 and Sylline Claparède, 1864. Böggemann & Westheide (2004) raised Parexogone to the generic level, and recent studies on the Syllidae have abandoned the subgenera classification, considering Exogone as monophyletic and not recognizing E. (Exogone) and E. (Sylline) as separate subgenera (Aguado & San Martín 2009; Aguado et al. 2012, Paresque et al. 2014). This classification is followed herein. Exogone anomalochaeta Benham, 1921 Figures 1–2 Exogone anomalochaeta Benham, 1921: 24–26, pl. 3, figs. 11–13; Benham 1927: 62, pl. A, figs. 9–10. Exogone (Exogone) anomalochaeta. San Martín & Parapar 1997: 291; Fukuda 2010: 115–118, fig. 31; Barroso 2011: 68–70, fig. 1. Exogone heterosetosa. Monro 1939: 115–116, fig. 9 (not E. heteroseta McIntosh, 1885). Exogone (Parexogone) tridentata Hartman-Schröder 1993: 143–144, fig. Abb 20–21. Material examined. Project 'Oceanprof'. 22°02’50.81”S, 39°52’24.10”W, 0–2 cm, 1050 m deep: 1 spec. (MNRJP 1151), 14 Dec 2002; 22°27’31.1”S, 40°09’23.5”W, 0–2 cm, 749 m deep: 1 spec. (MNRJP 1152), 18 Jun 2003. Project ‘Habitats’. 21°22’58”S, 40°19’44”W, 53 m deep: 1 spec., 21 Jul 2009; 21°42’53”S, 40°10’16”W, 98 m deep: 1 spec. (MZUSP 2905), 9 Mar 2009, and 1 spec., 7 Jul 2009; 22°17’42”S, 40°26’59”W, 104 m deep: 1 spec., 23 Feb 2009, and 3 specs (MZUSP 2908), 4 Jul 2009; 22°23’22”S, 40°34’59”W, 110 m: 1 spec. (MZUSP 2909), 25 Jul 2009; 22°46’54”S, 41°3’33”W, 78 m deep: 1 spec., 22 Feb 2009, and 1 spec., 03 Jul 2009; 22°52’1”S, 40°57’29”W, 92 m deep: 1 spec., 22 Feb 2009, and 1 spec. (MZUSP 2907), 3 Jul 2009; 23°10’5”S, 41°3’6”W, 107 m deep: 2 specs (MZUSP 2906), 02 Jul 2009; 23°11’28”S, 41°0’49”W, 117 m deep: 1 spec. (MZUSP 2904), 21 Feb 2009. Project ‘AMBES’. 19°33’20”S, 39°2’36”W, 420 m deep: 1 spec. (MZUSP 2912), 11 Dec 2011; 19°36’26”S, 39°10’17”W, 392 m deep: 6 specs (MZUSP 2911), 14 Dec 2011; 19°49’37”S, 39°35’41”W, 410 m deep: 1 spec., 14 Jan 2012; 20°14’19”S, 39°48’36”W, 381 m deep: 1 spec. (MZUSP 2914), 9 Jan 2012, and 5 specs (MZUSP 2915), 19 Jun 2013; 20°15’36”S, 39°46’15”W, 1040 m deep: 1 spec. (MZUSP 2913), 9 Jan 2012; 20°35’16”S, 39°53’47”W, 410 m deep: 58 specs, 8 Jan 2012, and 27 specs, 18 Jun 2013; 20°36’2”S, 39°51’35”W, 1000 m deep: 18 specs, 8 Jan 2012; 21°4’43”S, 40°8’31”W, 1024 m deep: 3 specs, 30 Dec 2011. DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 403 FIGURE 1. Exogone anomalochaeta. (A) anterior body, dorsal view; (B–C) falcigers, anterior and mid- to posterior body chaetigers, respectively; (D, H) aciculae, anterior and posterior body chaetigers, respectively; (E–F) dorsal simple chaetae, anterior and posterior body chaetigers, respectively; (G) ventral simple chaeta. Additional material examined. Exogone anomalochaeta—Australia, Tasmania, Macquarie Island (54°37'S, 158°51'E): 17 specs (AM W617, syntypes), coll. H. Hamilton, det. W. Benham. Antartic, South Shetland Islands, Livingston Island: 2 specs (MNCN 16.01/1517), coll. & det. G. San Martín, 07 Feb 1994; 24 specs (MNCN 16.01/ 1519), coll. & det. G. San Martín, 19 Feb 1994. Exogone heterosetosa—Australia, Western Australia, Goss Passage, north end of Long Island (28°28'18"S, 113°46'18"E), 8 m deep: 10 specs (AM W27057), coll. C. Bryce, 22 May 1994, det. G. San Martín, 2000. Exogone tridentata—Antarctic, Antarctic Peninsula, close to Base General Bernardo O´Higgins Riquelme (63°00.49'S, 57°09.45'W), 97 m deep: 2 specs (ZMH P-21218, holotype; ZMH P– 21219, paratype), coll. G. Hartmann, M. Siegel & G. Hartmann-Schröder, 1991, det. G. Hartmann-Schröder, 1992. Description. Longest specimen examined 3.0 mm long, 0.3 mm wide, with 35 chaetigers. Palps narrower than anterior chaetigers, almost totally fused, with distal notch. Prostomium ovate, shorter than palps, with two pairs of eyes in open trapezoidal arrangement, almost transversally in line (Fig. 1A); antennae inserted close to each other, on middle of prostomium or slightly posteriorly; lateral antennae ovate, papilliform; median antenna digitiform, elongate, distally tapering, reaching around half length of palps (Fig. 2A–B, D). Peristomium shorter than subsequent segments, sometimes covering posterior part of prostomium, including posterior pair of eyes; peristomial cirri minute, rounded to ovate. Dorsal cirri ovate, longer than peristomial cirri, slightly shorter than lateral antennae, absent on chaetiger 2 (Figs 1A; 2A–C). Ventral cirri similar to dorsal cirri but shorter (Fig. 2C). Parapodial lobes conical (Fig 1A; 2A–C, G–I). Compound chaetae as falcigers only; anterior parapodia with 5–9 falcigers each, midbody with 4–5, posterior parapodia with 3 falcigers each; falcigers with distally swollen, spinulated shafts; blades small, with inverted dorso-ventral gradation in length, ~4–8 μm long along body, dorsalmost blades sometimes not reaching beyond tips of shafts (Fig. 2F), blades bidentate and spinulated, distal tooth shorter (Figs 1B–C; 2F–I). Dorsal simple chaetae present on all parapodia, subdistally inflated, with slightly more sigmoid and more acute tip on posterior body chaetigers (Figs 1E–F; 2G–I); ventral simple chaetae only present on posterior body, smooth, bidentate, tips resembling those of blades of ventralmost falcigers, with distal 404 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. FIGURE 2. Exogone anomalochaeta, SEM. (A) entire worm, dorsal view; (B–C) anterior body, dorsal and ventral views, respectively; (D) detail of prostomium and peristomium, right dorso-lateral view; (E) nuchal organ; (F) dorsalmost falciger, midbody chaetiger; (G–H) chaetae, chaetiger 1; (I) chaetae, midbody chaetiger. DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 405 tooth shorter (Fig. 1G). Anterior parapodia with 2 aciculae each, aciculae distally inflated, with apparently hollow tip, one straight, other slightly bent distally (Fig. 1D); from proventricle onwards, single acicula per parapodium, similar to anterior ones, progressively stouter (Fig. 1H). Pygidium with 1 pair of elongate anal cirri, much longer than dorsal cirri and up to twice as long as median antenna. Pharynx through 3 chaetigers, tooth close to anterior border; proventricle extending for 3.5–5 chaetigers, with ca. 20 rows of muscle cells (Fig. 1A). Remarks. The specimens herein examined match the original description, except for the length of the proventricle, which is longer in Brazilian specimens (3.5–5 chaetigers, as opposed to 2–3 chaetigers, as in the original description). In addition, although not mentioned in the text of the original description, Benham illustrated a specimen with dorsal cirri on chaetiger 2 (Benham 1921: Pl. 5, Fig. 11), while in Brazilian specimens, dorsal cirri are absent on chaetiger 2. Both these differences, however, could not be confirmed in the specimens examined from other localities, including the syntypes, all specimens being remarkably similar to the Brazilian ones. Based on specimens from the Kerguelen Islands, Monro (1939) synonymized this species with E. heterosetosa McIntosh, 1885. However, based on the illustrations provided (Monro 1939: Fig. 9), all the material examined by Monro belongs to E. anomalochaeta and not to E. heterosetosa, which is a fairly different species (see San Martín 2005). Exogone heterosetosa differs from E. anomalochaeta by having the median antenna longer than the combined length of prostomium and palps, by having falcigers with proportionally larger blades, and also by having spiniger-like compound chaetae, which are absent in E. anomalochaeta. So, we consider both E. anomalochaeta and E. heterosetosa as valid species. The type series of E. tridentata was also examined, but we could not find significant differences to E. anomalochaeta. Thus, E. tridentata is herein considered as a junior-synonymy of E. anomalochaeta. This is the first formal report of Exogone anomalochaeta out of the Southern Ocean, although the presence of this species in waters off southern Brazil was already known from two unpublished PhD thesis (Fukuda 2010; Barroso 2011). Geographic distribution and bathymetric range. Southern Ocean—Commonwealth Bay and Macquarie Island (type locality—from the intertidal zone to ~10 m deep) (Benhan 1921); South Shetland Islands, 19–184 m deep (San Martín & Parapar 1997). Brazilian specimens were collected from 749–1050 m deep. Genus Parexogone Mesnil & Caullery, 1918 Type species. Paedophylax hebes Webster & Benedict, 1884. Diagnosis. Body short, slender, palps totally fused or with only distal notch. Prostomium with 2 pair of eyes, sometimes also 1 pair of anterior eyespots; 3 antennae, frequently inserted away from each other. Peristomium with 1 pair of peristomial cirri. Dorsal cirri short, papilliform or ovate; ventral cirri smaller than dorsal ones. Falcigers with little variation along body, unidentate or bidentate, the latter with distal tooth larger than subdistal one. Some species with dorsalmost falcigers with remarkably longer blades, similar to spiniger-like chaetae. Dorsal simple chaetae with similar morphology throughout body, unidentate or bidentate. Reproduction by epigamy, with ventral incubation of eggs and embryos; some species viviparous. Remarks. See remarks of Exogone. Parexogone wolfi (San Martín, 1991) Figure 3 Exogone (Parexogone) wolfi San Martín, 1991: 726–727, fig. 6; 2003: 243–244, figs. 129–130; 2005: 111–113, fig. 64. Exogone sp. A. Uebelacker 1984: 30–37, fig. 30. Parexogone wolfi. Böggeman & Purschke 2005: 223–225, fig. 2. Material examined. Project 'Oceanprof'. 22°04’32.8”S, 39°54’11.4”W, 0–2 cm, 722 m deep: 1 spec. (MNRJP 1153), 30 Jun 2003; 22°04’33.9”S, 39°52’05.1”W, 0–2 cm, 1030 m deep: 1 spec. (MNRJP 1154), 30 Jun 2003; 22°04’43.3”S, 39°49’09.3”W, 0–2 cm, 1299 m deep: 1 spec. (MNRJP 1155), 25 Mar 2006; 22°35’03.7”S, 40°08’52.5”W, 0–2 cm, 1043 m deep: 2 specs (MNRJP 1156), 15 Jun 2003; 22°34’05.0”S, 40°00’12.6”W, 0–2 cm, 406 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. 1337 m deep: 1 spec. (MNRJP 1157), 15 Jun 2003; 22°26’28.8”S, 39°58’53.3”W, 0–2 cm, 1046 m deep: 2 specs (MNRJP 1158), 20 Jun 2003; 22°30’21.7”S, 39°56’53.7”W, 0–2 cm, 1353 m deep: 1 spec. (MNRJP 1159), 21 Jun 2003. Project ‘HABITATS’. 21°42’37”S, 40°8’59”W, 147 m deep: 1 spec., 9 Mar 2009; 22°31’7”S, 40°31’32”W, 139 m deep: 8 specs, 23 Feb 2009; 23°11’29”S, 41°0’49”W, 150 m deep: 5 specs, 21 Feb 2009; 23°13’2”S, 40°57’36”W, 724 m deep: 3 specs, 24 Jun 2008; 23°13’48”S, 40°55’56”W, 986 m deep: 39 specs, 8 May 2009; 23°15’11”S, 40°53’53”W, 1302 m deep: 18 specs, 9 May 2008; 23°36’14”S, 41°21’29”W, 142 m deep: 1 spec., 1 Mar 2009. Project ‘AMBES’. 19°54’4”S, 39°22’27”W, 1335 m deep: 6 specs, 27 Jun 2013; 21°4’14”S, 40°14’14”W, 141 m deep: 1 spec., 11 Jul 2013. Additional material examined. Parexogone wolfi—United States, Florida, off Port Everglades (26°00'54"N, 80°03'24"W), 188 m deep: 1 spec. (USNM 101326, holotype), det. G. San Martín, 1991; France, Bay of Biscay, Capbreton Canion (43°43'25"N, 02°18'17"W) 983–1032 m deep: 3 specs (MNCN 16.01/2948), coll. Campaña Capbreton 88, 8 Jul 1988, det. G. San Martín, Ceberio & Aguirrezabalaga. Description. Body thin, elongate; longest specimen examined 6.0 mm long, 0.2 mm wide, with 67 chaetigers. Palps completely fused, triangular. Prostomium ovate, shorter than palps, with two pairs of eyes in trapezoidal arrangement, anterior and posterior eyes sometimes close to each other, nearly coalescent; antennae inserted slightly separated from each other; lateral antennae inserted slightly anteriorly to median antenna, approximately in front of anterior eyes, ovate to digitiform; median antenna inserted on middle of prostomium or slightly anteriorly, up to 5 times longer than lateral ones, reaching beyond tip of palps or chaetiger 4, if directed posteriorly (Fig. 3A, J). Peristomium shorter than subsequent segments; peristomial cirri ovate to papilliform, shorter than lateral antennae (Fig. 3A, J). Dorsal cirri on all chaetigers, papilliform, larger than peristomial cirri but shorter than lateral antennae (Fig. 3A, J); ventral cirri similar to peristomial cirri. Compound chaetae as spiniger-like and falcigers. Anterior parapodia with up to 3 spiniger-like chaetae each, from midbody onwards 1–2 spiniger-like chaetae per parapodium; shafts of spiniger-like chaetae from anterior and mid-body chaetigers with long and thin distal spines; blades bidentate and spinulate, spines of almost uniform length, except for the distalmost spines, much longer than remaining and directed distally, reaching beyond level of distal tooth ("aristae"); blades 55–35 µm long on anterior parapodia, 80–58 µm long on midbody and 40–10 µm long on posterior parapodia (Fig. 3B–D, K). Anterior parapodia with 8–10 falcigers each, midbody with 3–4, posterior parapodia with 2–3 falcigers each; shafts of facigers distally spinulated, spines longer on anterior body; falciger blades bidentate and spinulate, teeth close to each other, distal tooth slightly larger, blades with distally directed aristae reaching beyond level of distal tooth, sometimes absent on posterior body; blades of falcigers 22–11 µm long on anterior parapodia, 28–11 μm on midbody, and 12–8 µm long on posterior parapodia (Fig. 3B–D, L). Dorsal simple chaetae present from proventricle level, sigmoid, distally tapering, bidentate and spinulated, aristae reaching beyond level of distal tooth, chaetae progressively stouter and more conspicuously bidentate towards posterior body (Fig. 3E–F); ventral simple chaetae only present on posterior body, bidentate, smooth or with short subdistal spines (Fig. 3G). Anterior parapodia with up to 3 aciculae each, distally inflated and slightly oblique, with apparently hollow tips (Fig. 3H); single acicula per parapodium from midbody onwards, similar in shape to those of anterior body chaetigers, but progressively stouter (Fig. 3I). Pygidium with 1 pair of elongate anal cirri as long as median antenna. Pharynx through 3–4 chaetigers, with conical tooth close to anterior border; proventricle extending for ~3 chaetigers, with ~16 rows of muscle cells (Fig. 3A). Remarks. Brazilian specimens have longer blades of spiniger-like chaetae, up to 80 μm long, similar to those of specimens from the Gulf of México and Florida (San Martín 1991), and Bay of Biscay (San Martín 2003), while Australian specimens have spiniger-like chaetae with shorter blades, up to 58 μm long (San Martín 2005). Furthermore, the specimens herein analysed have the proventricle extending through 3 chaetigers, while specimens from other localities have a shorter proventricle, extending through 2 chaetigers (San Martín 1991, 2005). The wide distribution and bathymetric range of this species, together with the discrepancies between descriptions of specimens from different localities, may indicate that this is in fact a complex of sibling species. Although already informally recorded in Brazil (Fukuda 2010; Barroso 2011), this is the first formal report of this species to Brazilian waters. Geographic distribution and bathymetric range. Atlantic Ocean—USA, Florida, 188 m deep (type locality) (San Martín 1991); Gulf of México, 34–106 m deep (Uebelacker 1984); Bay of Biscay, 1000 m deep (San Martín et al. 1996); Angola Basin, 3964–5497 m deep (Böggemann & Purschke 2005; Böggemann 2009). Indian Ocean— Australia, Western Australia, 8 m deep (San Martín 2005). Brazilian specimens were collected from 722–1353 m deep. DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 407 FIGURE 3. Parexogone wolfi. (A) anterior body, dorsal view; (B–D) compound chaetae, anterior, mid- and posterior body chaetigers, respectively; (E–F) dorsal simple chaetae, anterior and posterior body chaetigers, respectively; (G) ventral simple chaeta; (H–I) aciculae, anterior and posterior body chaetigers, respectively; SEM: (J) anterior end, right lateral view; (K) spiniger-like chaeta, midbody chaetiger; (L) falcigers, midbody chaetiger. Parexogone campoyi San Martín, Ceberio & Aguirrezabalaga, 1996 Figures 4–6 Exogone (Parexogone) campoyi San Martín et al., 1996: 252–253, Figs. 4–5; San Martín 2003: 244, Figs. 131–132. Material examined. Project ‘Oceanprof'’. 22°39’44.28”S, 40°15’44.41”W, 0–2 cm, 994 m deep: 1 spec. (MNRJP 1160), 19 May 2002; 22°43’17.37”S, 40°12’12.34”W, 2–5 cm, 1157 m deep: 1 spec. (MNRJP 1161), 15 May 2002; 22°10’53.4”S, 39°52’18.3”W, 0–2 cm, 1039 m deep: 1 spec. (MNRJP 1162), 01 Jul 2003; 22°11’04.39”S, 39°47’04.64”W, 0–2 cm, 1654 m deep: 1 spec. (MNRJP 1163), 25 Nov 2002; 22°11’16.63”S, 39°43’44.72”W, 0–2 cm, 1964 m deep: 1 spec. (MNRJP 1164), 25 Nov 2002; 21°57’15.5”S, 39°47’41.7”W, 2–5 cm, 1857 m deep: 1 spec. (MNRJP 1165), 28 Jun 2003; 21°52’59.2”S, 39°55’32.2”W, 0–2 cm, 751 m deep: 1 spec. (MNRJP 1166), 29 408 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. Jun 2003; 22°31’11.8”S, 40°15’12.1” W, 0–2 cm, 743 m deep: 1 spec. (MNRJP 1167), 18 Jun 2003; 22°35’03.7”S, 40°08’52.5”W, 0–2 cm, 1043 m deep: 1 spec. (MNRJP 1168), 15 Jun 2003; 22°27’31.1”S, 40°09’23.5”W, 0–2 cm, 749 m deep: 1 spec. (MNRJP 1169), 18 Jun 2003; 22°34’05.0”S, 40°00’12.6”W, 0–2 cm, 1337 m deep: 1 spec. (MNRJP 1170), 15 Jun 2003; 22°37’02.9”S, 39°56’20.1”W, 0–2 cm, 1945 m deep: 1 spec. (MNRJP 1171), 13 Jun 2003;. Project ‘HABITATS’. 22°19’57”S, 40°1’59”W, 707 m deep: 1 spec., 7 Jul 2008; 22°36’27”S, 40°22’29”W, 700 m deep: 3 specs, 25 Jun 2008; 22°49’22”S, 40°8’17”W, 1864 m deep: 1 spec., 11 May 2008; 23°13’46”S, 40°55’56”W, 986 m deep: 1 spec., 8 May 2008; 23°15’11”S, 40°53’48”W, 1302 m deep: 3 specs, 9 May 2008. Project ‘AMBES’. 19°3’29”S, 37°48’39”W, 1302 m deep: 2 specs, 30 Jan 2012; 19°34’20”S, 38°41’19”W, 449 m deep: 5 specs, 30 Jun 2013; 19°40’8”S, 39°7’22”W, 1035 m deep: 23 specs, 13 Dec 2011; 19°50’1”S, 39°26’30”W, 1055 m deep: 6 specs, 11 Jan 2012; 19°53’31”S, 39°32’56”W, 1023 m deep: 22 specs, 13 Jan 2012; 19°58’11”S, 39°31’38”W, 1300 m deep: 4 specs, 13 Jan 2012; 20°16’35”S, 38°27’18”W, 1902 m deep: 1 spec., 21 Dec 2011; 20°36’2”S, 39°51’35”W, 1000 m deep: 8 specs, 8 Jan 2012; 20°36’42”S, 39°49’25”W, 1333 m deep: 6 specs, 8 Jan 2012; 21°4’43”S, 40°8’31”W, 1023 m deep: 11 specs, 30 Dec 2011; 21°4’51”S, 40°4’14”W, 1300 m deep: 7 specs, 31 Dec 2011. FIGURE 4. Parexogone campoyi. (A, J) anterior body, dorsal view; (B) dorsal simple chaeta; (C) ventral simple chaeta; (D–E) aciculae, anterior and posterior body chaetigers, respectively; (F–I, K–L) falcigers, midbody chaetigers. DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 409 FIGURE 5. Parexogone campoyi, SEM. (A) entire worm (posteriorly incomplete, broken); (B–C) anterior body, dorsal and ventral views, respectively; (D) close up of the oral area. Description. Body filiform, longest specimen examined 4 mm long, 0.2 mm wide, with 39 chaetigers. Palps subpentagonal, totally fused, distally pointed (Figs. 4A, J; 5A–D). Prostomium ovate, with two pairs of eyes in trapezoidal arrangement; antennae inserted slightly separated from each other, approximately between posterior eyes; lateral antennae digitiform, reaching chaetiger 2, if directed posteriorly; median antenna about twice as long as lateral ones, reaching beyond tips of palps and chaetiger 4, if directed posteriorly (Fig. 4A). Peristomium shorter than subsequent segments, peristomial cirri ovate, shorter than lateral antennae (Figs 4A, J; 5B). Dorsal cirri digitiform, longer than peristomial cirri but shorter than lateral antennae, progressively thinner towards posterior body (Figs 4A; 5A–B); ventral cirri similar to dorsal cirri but slightly shorter (Fig. 5C–D). Anterior parapodia with 9–15 compound chaetae each, midbody with 5–7, posterior parapodia with 3–4 compound chaetae each; compound chaetae with smooth to slightly spinulated shafts, blades unidentate to sub-bidentate, with relatively coarse spines 410 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. and conspicuous dorso-ventral gradation in length (Figs 4F–I, K–L; 6A–E); blades 60–10 µm long on anterior body parapodia, 80–10 µm on midbody and 50–10 µm on posterior body parapodia. Dorsal simple chaetae present from midbody onwards, subdistally spinulated, sub-bidentate with acute, needle-like tip (Figs 4B; 6F–G); ventral simple chaetae only present on posterior chaetigers, slightly spinulated, sub-bidentate (Fig. 4C). Anterior body parapodia with 2 aciculae each, subdistally inflated, with sharp, acute, needle-like tip; from midbody onwards, single acicula per parapodium, with similar shape (Fig. 4D–E). Pygidium with 1 pair of thin, elongate anal cirri, longer than posterior dorsal cirri and median antenna. Pharynx through 3 chaetigers, with conical tooth close to anterior border; proventricle extending for 5 chaetigers, with ca. 15–17 rows of muscle cells (Fig. 4A). Remarks. Brazilian specimens match the original description of P. campoyi in all characters. Geographic distribution and bathymetric range. Atlantic Ocean—Bay of Biscay (type locality), 1000–1100 m deep (San Martín et al. 1996); North Sea—UK, Tynemouth (San Martín & Worsfold 2015). Brazilian specimens were collected from 743–1964 m deep. FIGURE 6. Parexogone campoyi, SEM. (A–B) parapodia, anterior body chaetigers; (C) falcigers, anterior body chaetigers; (D–E) parapodia and falcigers, midbody chaetigers; (F–G) dorsal simple chaetae, midbody chaetigers. Parexogone minuscula (Hartman, 1953) Figure 7 Exogone minuscula Hartman, 1953: 26–27, Fig. 5 A–F. Exogone obtusa Hartmann-Schröder & Rosenfeldt, 1988: 44–45, Figs 24–27. Parexogone cf. gambiae. Fukuda 2010: 163–165, Fig. 48 (not P. gambiae Lanera, Sordino & San Martín, 1994). DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 411 FIGURE 7. Parexogone minuscula. (A) anterior body, dorsal view; (B–C, F) falcigers, anterior, mid- and posterior body chaetigers, respectively; (D–E) dorsal simple chaetae, anterior and posterior body chaetigers, respectively; (G) ventral simple chaeta; (H) acicula; SEM: (I–K) falcigers. Material examined. Project 'Oceanprof'. 22°04’32.8”S, 39°54’11.4”W, 0–2 cm, 722 m deep: 2 specs (MNRJP 1172), 30 Jun 2003; 21°57’11.8”S, 39°56’04.2”W, 0–2 cm, 698 m deep: 3 specs (MNRJP 1173), 29 Jun 2003; 22°36’01.3”S, 40°21’43.7”W, 0–2 cm, 754 m deep: 4 specs (MNRJP 1174), 11 Jun 2003; 22°31’11.8”S, 40°15’12.1”W, 0–2 cm, 743 m deep: 1 spec. (MNRJP 1175), 18 Jun 2003; 22°27’31.1”S, 40°09’23.5”W, 0–2 cm, 749 m deep: 1 spec. (MNRJP 1176), 18 Jun 2003. Additional material examined. Exogone minuscula—Antarctica, South Shetland Islands, Livingston Island, off False Bay, 31 m deep: 1 spec. (USNM 46549), coll. W.L. Schmitt, Feb 1963, det. O. Hartman; Antarctica, Antarctica Peninsula, Palmer Archipelago, Anvers Island, Arthur Harbor, 40 m deep: 20 specs (USNM 46849), coll. J.S. Rankin, 17 Mar 1968, det. O. Hartman. Exogone obtusa—Antarctica, Antarctica Peninsula, near Joinville (63°30'S, 54°15'W), mud with gravel and stones, 220 m deep: 1 spec. (ZMH P-18965, holotype), coll. R/V 'Polarstern' st. 120, 21 Nov 1984, det. G. Hartmann-Schröder, 1987; Antarctica, Antarctica Peninsula, near Joinville (62°51'S, 56°00'W), 133 m deep: 4 specs (ZMH P-19165, paratypes), coll. R/V 'Polarstern' st. 158, 26 Nov 1984, det. G. Hartmann-Schröder & P. Rosenfeldt. Description. Body filiform, longest specimen examined 2.6 mm long, 0.2 mm wide, with 33 chaetigers. Palps 412 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. triangular, totally fused, distally blunt (Fig. 7A). Prostomium rectangular, with two pairs of eyes in trapezoidal arrangement; antennae inserted close to each other, on middle of prostomium or slightly posteriorly; lateral antennae digitiform, reaching chaetiger 2, if directed posteriorly; median antenna elongate, reaching chaetiger 4, if directed posteriorly (Fig. 7A). Peristomium shorter than subsequent segments with small, papilliform peristomial cirri. Dorsal cirri ovate, slightly larger than peristomial cirri, absent on chaetiger 2; dorsal cirri progressively longer towards posterior body (Fig. 7A). Ventral cirri shorter than dorsal cirri, larger than peristomial cirri. Anterior parapodia with 10–16 compound chaetae each, midbody with 5–6, posterior parapodia with 2–3 compound chaetae each; compound chaetae with subdistally spinulated shafts and spinulated and bidentate blades, with distal tooth rounded and larger; blades with tips progressively more strongly curved towards posterior body, especially in ventralmost chaetae; blades 50–15 μm long on anterior body, 23–15 μm long on midbody and 20–10 μm long on posterior body (Fig. 7B–C, F, I–K). Dorsal simple chaetae present from anterior body, slightly sigmoid, bidentate and subdistally spinulated, distal tooth slightly larger and rounded on anterior body, with tip progressively more acute towards posterior body (Fig. 7D–E); ventral simple chaetae present from midbody, more conspicuously sigmoid than dorsal simple chaetae, bidentate and subdistally spinulated (Fig. 7G). Parapodia throughout with single acicula each, with inflate, rounded and apparently hollow tip (Fig. 7H). Pygidium with long anal cirri, with length equivalent to length of 3 posteriormost chaetigers together. Pharynx through 4 chaetigers, with conical tooth slightly away from anterior border; proventricle through 4–5 chaetigers, with ca. 17–19 rows of muscle cells (Fig. 7A). Remarks. This species is readily recognizable by the morphology of the dorsal simple chaetae and compound chaetae, with the characteristic falcate, bidentate blades, more conspicuously in ventralmost falcigers from midbody onwards. We examined the type material of P. obtusa and could not find significant differences to P. minuscula, therefore considering the former as a junior-synonym of the latter species. Geographic distribution and bathymetric range. Atlantic Ocean—South Georgia, Moraine Fjord, 148 m deep (type locality); Falkland Islands, 12–40 m deep (Hartman 1953). Brazilian specimens were collected from 698–754 m deep. Genus Prosphaerosyllis San Martín, 1984a Type species. Sphaerosyllis xarifae Hartmann-Schröder, 1960, designated by San Martín (1984a). Diagnosis. Body short, covered dorsally and ventrally with numerous papillae, including on palps, cirri and parapodia. Prostomium with 3 antennae, 4 eyes and usually 1 pair of anterior eyespots. Peristomium with 1 pair of peristomial cirri. Dorsal cirri usually present on all chaetigers, absent on chaetiger 2 in a few species. Antennae, peristomial cirri, dorsal cirri throughout and anal cirri with spherical base (cirrophore) and pointed, distally blunt tip (cirrostyle) usually retractable within cirrophore, at least from midbody. Parapodial glands usually absent. Blades of falcigers usually unidentate and relatively short, with dorso-ventral gradation in length. Aciculae subdistally inflated and slightly curved, with acute tip. Pharynx long and wide, usually not surrounded by soft papillae on opening, pharyngeal tooth conical, usually away from anterior margin; proventricle massive, about same size as pharynx (Fukuda et al. 2009). Reproduction with dorsal brooding of eggs. Remarks. San Martín (1984a) recognized two subgenera of Sphaerosyllis based on morphology, Sphaerosyllis and Prosphaerosyllis. Although this division was not immediately accepted by some authors (Russell 1989; Kudenov & Harris 1995), San Martín (2003) raised Prosphaerosyllis to the generic level, taking into account the brooding system, which is different from one genus to another, and this proposal has been adopted by subsequent authors (Böggemann & Westheide 2004; San Martín 2005; Fukuda et al. 2009). In Brazil, three species of Prosphaerosyllis were already reported, P. brachycephala Fukuda, Yunda-Guarin & Nogueira, 2009, P. isabelllae and P. xarifae (Fukuda et al. 2009). Prosphaerosyllis isabellae (Nogueira, San Martín & Amaral, 2001) Sphaerosyllis isabellae Nogueira et al., 2001: 1777–1779, Fig. 1. Prosphaerosyllis isabellae. San Martín 2005: 68, Fig. 23; Fukuda et al. 2009: 1450–1453, Figs. 4–5. DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 413 Material examined. Project 'Oceanprof'. 22°40’57.84”S, 40°16’30.35”W, 0–2 cm, 1045 m, 1 spec. (MNRJP 1177), 22 Nov 2002. Diagnosis. Palps densely papillated. Dorsal cirri throughout with bulbous base, with iridescent inclusions, distal end with nearly spherical tip. A pair of papillae on each parapodium; aciculae subdistally bent and thick, with acute tip. Pharynx with pair of brown glands in chaetiger 1. Remarks. The specimen herein studied is similar to those of the original description. This is the deepest record of this species. Geographic distribution and bathymetric range. Atlantic Ocean—Brazil, Rio de Janeiro, 110–1045 m deep; São Paulo, shallow subtidal to 808 m deep (Nogueira et al. 2001; Fukuda et al. 2009); Pacific Ocean: Australia, 6– 30 m deep (San Martín 2005). The specimen found in Campos Basin was obtained from 1045 m deep. Genus Sphaerosyllis Claparède, 1863 Type species. Sphaerosyllis hystrix Claparède, 1863. Diagnosis. Small bodied specimens, body usually covered by numerous papillae. Palps totally fused or with distal notch. Prostomium with 3 antennae and 2 pairs of eyes, anterior eyespots usually absent. Peristomium with 1 pair of peristomial cirri. Antennae, peristomial and dorsal cirri with similar shape, with bulbous, inflate bases, and narrower, tapering tips. Dorsal cirri usually absent on chaetiger 2. Parapodial glands frequently present, with granular or fibrilar material. Compound chaetae as heterogomph falcigers, usually with short, unidentate blades. Aciculae frequently stout, subdistally bent at ~90°, with acute tips. Pharynx narrow, with opening surrounded by soft, rounded papillae, and conical central tooth at anterior margin; proventricle short, with few, coarse rows of muscle cells. Reproduction by epigamy, with ventral incubation of eggs and embryos. Remarks. The genus currently counts with 2 formal records for the Brazilian coast, S. annulata Nogueira, San Martín & Fukuda, 2004 and S. brasiliensis Nogueira, San Martín & Amaral, 2001, plus ~8 other species, from informal records (e.g., Fukuda 2010). Sphaerosyllis ceciliae sp. nov Figure 8–9 Type material. Holotype (MNRJP 1178): 22°26’28.8”S, 39°58’53.3”W, 0–2 cm, 1046 m deep, 20 Jun 2003. Paratypes: 19°53’31”S, 39°32’56”W, 1023 m deep: 48 specs (MZUSP 2964), 13 Jan 2012; 21°4’43”S, 40°8’31”W, 1024 m deep: 24 specs (MZUSP 2965), 30 Dec 2011; 22°40’57.84”S, 40°16’30.35”W, 0–2 cm, 1045 m deep: 2 specs (ZUEC POL 19881), 22 Nov 2002; 22°10’54.6”S, 39°48’59.5”W, 0–2 cm, 1336 m deep: 2 specs (MNRJP 1179), 25 Jun 2003; 22°04’33.9”S, 39°52’05.1”W, 0–2 cm, 1030 m deep: 2 specs (MNRJP 1180), 30 Jun 2003; 21°52’51.8”S, 39°48’12.5”W, 2–5 cm, 1372 m deep: 1 spec. (MNRJP 1181), 26 Jun 2003; 22°31’12”S, 40°15’11”W, 0–2 cm, 722 m deep: 1 spec. (MNRJP 1182), 22 Nov 2002; 22°26’28.5”S, 39°54’08.3”W, 0–2 cm, 1354 m deep: 2 specs (MNRJP 1183), 21 Jun 2003; 22°31’37.2”S, 39°55’14.5”W, 0–2 cm, 1630 m deep: 1 spec. (MNRJP 1184), 16 Jun 2003. Additional material. Project ‘AMBES’. 19°3’29”S, 37°48’39”W, 1302 m deep: 3 specs, 30 Jan 2012; 19°40’8”S, 39°7’22”W, 1035 m deep: 15 specs, 13 Dec 2011; 19°47’5”S, 39°3’11”W, 1258 m deep: 1 spec., 28 Jan 2012; 19°49’37”S, 39°35’41”W, 410 m deep: 1 spec., 14 Jan 2012; 19°50’1”S, 39°26’30”W, 1055 m deep: 10 specs, 11 Jan 2012; 19°52’56”S, 38°35’8”W, 1022 m deep: 10 specs, 17 Dec 2011; 19°54’5”S, 39°22’20”W, 1335 m deep: 1 spec., 11 Jan 2012; 19°58’11”S, 39°31’38”W, 1300 m deep: 5 specs, 13 Jan 2012; 20°4’8”S, 38°31’27”W, 1302 m deep: 2 specs, 21 Dec 2011; 20°15’36”S, 39°46’15”W, 1040 m deep: 30 specs, 9 Jan 2012; 20°17’41”S, 39°42’38”W, 1358 m deep: 2 specs, 9 Jan 2012; 20°36’2”S, 39°51’35”W, 1000 m deep: 49 specs, 8 Jan 2012, and 35 specs, 18 Jun 2013; 20°36’42”S, 39°49’25”W, 1333 m deep: 5 specs, 8 Jan 2012; 21°4’43”S, 40°4’12”W, 1295 m deep: 1 spec., 9 Jun 2013; 21°4’51”S, 40°4’14”W, 1300 m deep: 1 spec., 31 Dec 2011. 414 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. FIGURE 8. Sphaerosyllis ceciliae sp. nov. (A, I) anterior body, dorsal view; (B) midbody parapodium; (C, M) dorsal simple chaetae; (D) ventral simple chaeta; (E) acicula; (F–H, K–L) falcigers; (J) midbody parapodia, dorsal view.; N, posterior end, dorsal view. Description. Holotype with 40 chaetigers, 2.4 mm long and 0.2 mm wide; complete paratypes with 29–33 chaetigers, 1.4–1.9 mm long, 0.2–0.3 mm wide. Dorsum scattered with small papillae, more conspicuously dorsolaterally (Figs 8I–J, 9A–B), papillae more elongate towards posterior body (Fig. 9C). Palps slightly longer than prostomium, almost totally fused, with distal notch, median line well marked (Figs 8A, I; 9A–B), sometimes with tips ventrally bent. Prostomium rectangular, with 2 pairs of eyes in trapezoidal arrangement; lateral antennae inserted close to anterior border of prostomium, almost reaching tip of palps; median antenna inserted posteriorly DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 415 FIGURE 9. Sphaerosyllis ceciliae sp. nov., SEM. (A) anterior body, dorsal view; (B) detail of prostomium, peristomium and first chaetigers, dorsal view; (C) posterior body, dorsal view; (D) left-hand parapodia, chaetigers 1 and 2, dorsal view; (E) dorsal simple chaetae, posterior body chaetiger; (F) spiniger-like chaeta, chaetiger 3; (G) parapodium, chaetiger 1, dorsal view; (H) anterior to midbody chaetigers; (I) posterior body parapodia, ventral view. 416 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. to middle of prostomium, slightly longer than lateral antennae, reaching around same level as lateral antennae (Figs 8A, I; 9A–B). Peristomium slightly shorter than anterior body chaetigers, sometimes forming a fold covering posterior part of prostomium, especially middorsally, reaching base of median antenna (Figs 8I; 9A–B). Antennae, peristomial and dorsal cirri throughout with similar morphology, basally bulbous and with elongate, thin tip. Dorsal cirri of uniform length throughout, shorter than antennae (Figs 8A, I–J; 9A–C). Thin, digitiform ventral cirri, shorter than dorsal cirri, of uniform length throughout (Fig. 9I). From chaetiger 4 onwards, each parapodium with one gland containing fibrilar inclusions, 1 pair per chaetiger, located dorsally to bases of dorsal cirri; glands opening dorsally through prominent papilla, larger than other papillae (Fig. 8A, I–J). Anterior parapodia with 6–7 falcigers each, midbody with 4–5, posterior parapodia with 2–4 falcigers each; falcigers with distally spinulated shafts, with elongate spines; blades spinulated, with straight spines, approximately of uniform length along blade, more evident on dorsalmost and intermediate falcigers in each fascicle; blades bidentate to sub-bidentate, subdistal tooth much smaller, resembling an enlarged spine, more evident on dorsalmost chaetae; subdistal tooth progressively shorter ventralwards within each fascicle, ventralmost chaetae usually unidentate with a developed subdistal spine (Figs 8F–H, K–L; 9D, F–I); blades 45–20 μm long on anterior body, 50–20 μm on midbody, and 30–20 μm long on posterior body. Dorsal simple chaetae present from first chaetiger onwards, sigmoid, distally spinulate (Figs 8C, M; 9E); ventral simple chaetae present from midbody, sigmoid, smooth (Fig. 8D). Anterior body with 2 aciculae per parapodium, one distally bent at 90°, with acute tip, other straight, distally pointed; single acicula per parapodium in mid- and posterior body chaetigers, of first type; aciculae progressively stouter towards posterior body (Fig. 8B, E). Pygidium with 1 pair of anal cirri longer and thinner than dorsal cirri. Pharynx through 3–4 chaetigers, with conical tooth on anterior border; proventricle through 3 chaetigers, with ca. 15 rows of muscle cells (Fig. 8A). Remarks. Sphaerosyllis ceciliae sp. nov. is similar to S. boeroi Musco, Çinar & Giangrande, 2005, S. hystrix Claparède, 1863, S. parabulbosa San Martín & López, 2002, S. taylori Perkins, 1981, and S. thomasi San Martín, 1984b, due to the fibrilar nature of the inclusions within the parapodial glands. However, S. ceciliae sp. nov. differs from S. parabulbosa, S. taylori and S. thomasi on the morphology of the falcigers, remarkably longer bidentate/ sub-bidentate blades in the new species compared to these congegers; furthermore, S. thomasi and S. taylori present blades more conspicuously spinulated than in S. ceciliae sp. nov. Sphaerosyllis hystrix differs from S. ceciliae sp. nov. in having antennae and cirri throughout proportionally larger than S. ceciliae sp. nov., and the subdistal tooth/ spine of the falciger blades of S. ceciliae sp. nov. is more conspicuous and longer than in S. hystrix, although the spinulation of the blades is more prominent in the latter species. Sphaerosyllis boeroi, described from the Mediterranean Sea, is probably the most similar species to S. ceciliae sp. nov., but can be distinguished by the longer blades of the midbody falcigers (up to 62.5 μm long in S. boeroi, against up to 50 μm long in S. ceciliae sp. nov.), as well as by the more conspicuous spinulation of the falcigers blades. Furthermore, S. boeroi inhabits shallower environments (5–150 m), when compared to those where S. ceciliae sp. nov. was found (749–1903 m). The large papilla present above each parapodial gland seems to be connected to the gland, serving as a channel through which the inclusions are discharged to the environment. Many specimens were found with fibrilar components partially extruded through the openings on top of these papillae, and those components are similar to the ones observed by transparency inside the glands. Geographic distribution and bathymetric range. Sphaerosyllis ceciliae sp. nov. is only known from the Campos Basin (749–1903 m deep). Etymology. This species is dedicated to Cecilia Amaral, a famous Brazilian polychaetologist, for her essential contribution to the knowledge of Brazilian marine invertebrates, especially polychaetes, as well as her for continuous efforts regarding research and education of new polychaetologists. Sphaerosyllis monicae sp. nov. Figure 10 Material examined. Project 'Oceanprof'. 22°04’32.8”S, 39°54’11.4”W, 722 m deep: 1 spec. (ZUEC POL 19882, Holotype), 30 Jun 2003. Description. Body filiform, holotype 3 mm long, 0.2 mm wide, complete, with 47 chaetigers. Dorsum DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 417 scattered with few short, digitiform papillae, more evident dorso-laterally. Palps completely fused, distally rounded. Prostomium shorter than palps, rectangular, with two pairs of eyes in trapezoidal arrangement, anterior eyespots absent. Antennae all about same size, median antenna inserted slightly posteriorly to lateral ones (Fig. 10A). Peristomium covering the posterior part of prostomium, including part of posterior eyes. Antennae, peristomial and dorsal cirri throughout with similar morphology, basally bulbous, with elongate, thin tip. Dorsal cirri absent on chaetiger 2; thin, digitiform ventral cirri, shorter than dorsal cirri, of uniform length throughout. Parapodial lobes rectangular to distally rounded; parapodial glands absent. Anterior parapodia with 5–6 falcigers each, midbody with 4–5, posterior parapodia with 3–4 falcigers each; anterior body falcigers with subdistally spinulated shafts, especially dorsalmost ones; from midbody parapodia onwards, falcigers with smooth shafts; blades of dorsalmost and intermediate falcigers poorly spinulated, ventralmost falcigers with smooth blades; blades unidentate, except for some sub-bidentate dorsalmost falcigers on posterior body chaetigers, with prominent subdistal spine; blades 30–10 μm long on anterior body, 25–15 μm on midbody, and 15–10 μm long on posterior body (Fig. 10B–C). Dorsal simple chaetae on all chaetigers, subdistally spinulated (Fig. 10D); ventral simple chaetae present from midbody chaetigers, sigmoid, smooth (Fig. 10E). Parapodia all with single acicula each, distally bent at 90°, aciculae progressively stouter towards posterior body (Fig. 10F). Pygidium with 1 pair of anal cirri slightly larger than posterior body dorsal cirri (Fig. 10G). Pharynx through 2 chaetigers, with conical tooth near anterior border; proventricle extending for 3 chaetigers, with ca. 17 rows of muscle cells (Fig. 10A). Remarks. Sphaerosyllis monicae sp. nov. differs from all known species of Sphaerosyllis by the complete absence of both parapodial glands and papillae over the body and parapodia, as well as by the morphology of falcigers and aciculae. FIGURE 10. Sphaerosyllis monicae sp. nov. (A) anterior end and midbody, dorsal view; (B–C) falcigers, anterior and posterior body chaetigers, respectively; (D) dorsal simple chaeta; (E) ventral simple chaeta; (F) acicula; (G) posterior end, dorsal view. 418 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. Sphaerosyllis austriaca Banse, 1959, S. pirifera Claparède, 1868 and S. piriferopsis Perkins, 1981 are the most similar species to S. monicae sp. nov. due to the absence of parapodial glands. However, all these species have the body dorsally covered by papillae (San Martín 1984a, 2003), while in S. monicae sp. nov. the dorsal surface of the body is completely smooth. Sphaerosyllis bulbosa Southern, 1914, is also similar to Sphaerosyllis monicae sp. nov. regarding the absence of dorsal papillae. However, S. bulbosa presents much shorter falciger blades (up to 10 μm long), secondarily simple chaetae on midbody, apparently by the loss of blades and enlargement of shafts, and aciculae straight, subdistally enlarged (San Martín 2003). Although we only have one specimen, we prefer to describe it as a new species, given the unusual and unique set of characters it presents, and the difficulties in collecting more material from the locality where the specimen was found. Geographic distribution and bathymetric range. Sphaerosyllis monicae sp. nov. is only known from the Campos Basin, at 722 m deep. Etymology. This species is dedicated to Mônica Petti, a great Brazilian polychaetologist, for her relevant contribution to the knowledge of polychaetes. Subfamily Syllinae Grube, 1850 Genus Syllis Savigny in Lamarck, 1818 Type species. Syllis monilaris Savigny in Lamarck, 1818. Diagnosis. Body cylindrical of variable size, from relatively medium-sized to large, compared to most syllids. Subpentagonal prostomium, with 1 pair of palps totally free from each other or fused at bases, 3 antennae, 2 pairs of eyes and, sometimes, 1 pair of anterior eyespots. Peristomium usually shorter than segments, with 2 pairs of peristomial cirri; border between prostomium and peristomium usually with conspicuous nuchal organs, as 1 pair of ciliated grooves. Antennae, peristomial cirri, dorsal cirri throughout and anal cirri moniliform; ventral cirri short, digitiform. Parapodia with compound falcigers, some taxa with secondarily simple chaetae, due to fusion between shaft and blade, or loss of blade; spiniger-like chaetae frequently present; dorsal and ventral simple chaetae usually present on posterior chaetigers. Pharynx with large tooth on anterior border or slightly posteriorly to opening, trepan absent (Nogueira & San Martín 2002; Nogueira & Yunda-Guarin 2008). Reproduction by stolonization, with stolons with different morphologies, according to species. Some species viviparous. Remarks. The largest genus in the family, Syllis, as currently conceived, seems to be paraphyletic, according to phylogenetic analyses performed on the family (Aguado et al. 2007, 2012). However, until more detailed studies are made specifically on this genus, we follow herein the opinion of most authors (e.g., San Martín 2003; ÁlvarezCampos et al. 2015a, b), considering Typosyllis Langerhans, 1879 as a junior synonym of Syllis. Syllis aciculigrossa (San Martín, 1990) Figure 11 Pionosyllis aciculigrossa San Martín, 1990: 604–606, figs 10–11. Pionosyllis sp. B. Uebelacker 1984: 30–69, figs 30-61–30-62. Typosyllis aciculigrossa. Licher 1999: 36–37. Syllis aciculigrossa. Fukuda 2010: 246–250, fig. 70; Barroso 2011: 90–92, fig. 8. Material examined. Project 'Oceanprof'. 22°27’31.1”S, 40°09’23.5”W, 2–5 cm, 749 m deep: 1 spec. (MNRJP 1185), 18 Jun 2003. Project ‘AMBES’. 19°34’20”S, 38°41’18”W, 450 m deep: 8 specs, 9 Dec 2011; 19°36’26”S, 39°10’17”W, 392 m deep: 12 specs (MZUSP 2899), 14 Dec 2011; 19°49’7”S, 39°36’8”W, 158 m deep: 3 specs, 14 Jan 2012; 19°49’37”S, 39°35’41”W, 410 m deep: 1 spec., 14 Jan 2012; 20°14’17”S, 39°48’34”W, 395 m deep: 16 specs (MZUSP 2901), 19 Jun 2013; 20°35’16”S, 39°53’47”W, 410 m deep: 22 specs, 8 Jan 2012; 21°4’9”S, 40°13’7”W, 410 m deep: 2 specs, 30 Dec 2011. DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 419 FIGURE 11. Syllis aciculigrossa. (A) anterior body, dorsal view; (B) spiniger-like chaeta, posterior body chaetiger; (C–D) falcigers, anterior and posterior body chaetigers, respectively; (E–F) dorsal and ventral simple chaetae, respectively; (G–I) aciculae, anterior, mid- and posterior body chaetigers, respectively. Additional material examined. Pionosyllis aciculigrossa—United States, Florida, Gulf of México, off Panamá City (29°30'02"N, 86°24'59"W, 189 m deep): 1 spec. (USNM 65684, paratype), coll. BLM/MMS, Sep 1975, det. G. San Martín, 1989; United States, Texas, Gulf of Mexico (26°58'12"N, 96°33'W, 106 m deep): 1 spec. (USNM 89890, paratype), coll. BLM/MMS, 1976, det. G. San Martín, 1989. Description. Body slender, subcylindrical. Largest specimen analyzed incomplete, 23 mm long, 1 mm wide, with 134 chaetigers. Palps longer than prostomium, fused along basal third. Prostomium ovate, with 2 pairs of eyes in trapezoidal arrangement; median antenna inserted slightly backwards to middle of prostomium, with ~26 articles; lateral antennae inserted anteriorly to anterior pair of eyes, with ~22 articles each (Fig. 11A). Peristomium slightly shorter than anterior body chaetigers, dorsal peristomial cirri with ~20 articles each, ventral peristomial cirri shorter, with ~15 articles each. Dorsal cirri of chaetiger 1 about same length as median antenna, longer than remaining dorsal cirri, with ~25 articles each; dorsal cirri throughout with length equivalent to about half body width of corresponding chaetiger, articles progressively more elongate distalwards on each cirrus (Fig. 11A), and 420 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. also posteriorwards along body; anterior dorsal cirri with 13–17 articles, 8–13 articles per cirrus from proventricle level; ventral cirri shorter than parapodial lobes, digitiform. Parapodial lobes conical, progressively more slender and elongate towards posterior body. Anterior parapodia with 25–30 compound chaetae each, midbody with 7–15, posterior parapodia with ~8 compound chaetae each; 1–3 dorsalmost compound chaetae throughout spiniger-like (Fig. 11B), with unidentate, slightly spinulated blades, ~145 μm long on anterior body, 160–100 μm on midbody, 110–90 μm long on posterior body; falcigers on anterior body more slender than remaining, with apparently unidentate, spinulate blades (Fig. 11C), blades 60–20 μm long; from midbody onwards, falcigers with spinulated and bidentate blades, distal tooth slightly larger (Fig. 11D), blades 70–30 μm long on midbody, 50–32 μm long on posterior body. Dorsal simple chaetae only present on posterior body, thinner than shafts of falcigers, slightly sigmoid, bidentate, with short subdistal spines (Fig. 11E); ventral simple chaetae only present on posteriormost parapodia, slightly thinner than shafts of falcigers, sigmoid, smooth and bidentate, distal tooth slightly larger (Fig. 11F). Anterior parapodia with up to 7 aciculae each, one of which much thinner, remaining aciculae with rounded to blunt tips slightly protruding from parapodial lobes (Fig. 11G); from proventricle region, 2 distally tapering aciculae per parapodium (Fig. 11H), single acicula per parapodium from mid- to posterior body (Fig. 11I), with acute tips, progressively more distinctly protruding from parapodial lobes towards posterior body (Fig. 11G–I). Pharynx through ~8 chaetigers, with conical tooth close to anterior border, opening surrounded by crown of soft papillae; proventricle through 7–8 chaetigers, with ~36 rows of muscle cells (Fig. 11A). Remarks. The specimens herein studied agree with the paratypes examined, including in the presence of the apparently unidentate spiniger-like chaetae, which were not mentioned in the original description (San Martín 1990). Geographic distribution and bathymetric range. Atlantic Ocean—Gulf of México (type locality, 106–189 m deep) (San Martín 1990); Brazil, states of São Paulo (280 m deep), Rio de Janeiro (102–750 m deep), and Paraná (480 m deep) (Fukuda 2010). Syllidae Incertae Sedis Genus Anguillosyllis Day, 1963 Type species. Anguillosyllis capensis Day, 1963. Diagnosis. Body of relatively small size, with few chaetigers. Palps elongated, at least partially fused for most of their length. Prostomium with 3 antennae; eyes absent. Peristomium with 1 pair of ovate to digitiform peristomial cirri. Dorsal cirri smooth, long, filiform, frequently coiled over dorsum, absent on chaetiger 2. Ventral cirri digitiform, inserted medially to distally in parapodial lobes. Parapodial lobes relatively long, with pre- and postchaetal lobes. Compound chaetae heterogomph, with unidentate, slender blades. Pharyngeral tooth absent. Reproduction unknown (Aguado & San Martín 2008). Remarks. Anguillosyllis was recently redefined by Aguado & San Martín (2008), who recognized Braniella Hartman, 1965 as a junior-synonym and redescribed 2 of the 3 known species, all restricted to the deep sea. Aguado & San Martín (2008) considered three valid species in this genus, A. pupa (Hartman, 1956), A. palpata and A. capensis. We describe herein an additional species, A. lanai sp. nov., from material from off southeastern Brazil, raising to four the number of known species in this genus. All the species in this genus are quite similar to each other, differing mostly by the morphology of palps, presence/absence of dorsal cirri on chaetiger 2, details on the structure of the parapodial lobes, length of the blades of compound chaetae, and presence of parapodial glands. Also, the number of chaetigers in adults seems to be constant for each species, which is not a common feature for syllids. With the amendment to the description of A. palpata proposed herein (see below), it is possible that ‘absence of dorsal cirri on chaetiger 2’ should also be included in the diagnosis of the genus; the only exception being A. capensis, originally described as with dorsal cirri on all chaetigers (Day 1963), but the holotype of that species is currently in terrible shape, preventing confirmation of that character. Also, further descriptions of A. capensis do not bring clear statements on the matter, but the absence of dorsal cirri on chaetiger 2 is suggested by images (see Böggemann & Purschke 2005, Fig. 1; Aguado & San Martín 2008, Fig. 1). DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 421 Identification key to the species of Anguillosyllis 1. 2. 3. - Palps basally fused for 1/2–¾ of their lengths; complete specimens with up to 11 chaetigers. . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Palps totally fused along their lengths or with only distal notch; complete specimens with up to 10 chaetigers. . . . . . . . . . . . . 3 Parapodia bilobed, with postchaetal lobes elongate, digitiform. Blades of compound chaetae relatively short, 98–48 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. capensis Parapodia distally rounded to slightly bilobed, with posterior lobe more pronounced. Blades of compound chaetae elongate, 450–20 µm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. palpata Parapodial glands absent. Blades of compound chaetae relatively short, 95–10 mm long. . . . . . . . . . . . . . . . . . . . . . . . A. pupa Parapodial glands present. Blades of compound chaetae elongate, 170–10 mm long . . . . . . . . . . . . . . . . . . . . .A. lanai sp. nov. Anguillosyllis palpata (Hartman, 1967), amended Figure 12 Braniella palpata Hartman 1967: 55, pl. 14. Material examined. Project ‘Habitats’. 21°11’6”S, 40°12’17”W, 807 m: 3 specs, 29 Jun 2008; 22°20’50”S, 40°2’58”W, 705 m: 8 specs, 8 Jul 2008; 22°25’54”S, 39°53’55”W, 1299 m: 3 specs (MZUSP 2939), 29 May 2008; 22°33’33”S, 40°26’40”W, 393 m: 1 spec., 11 Jul 2008; 22°52’14”S, 40°5’8”W, 2455 m: 2 specs (MZUSP 2938), 22 May 2008; 22°59’52”S, 40°47’45”W, 689 m: 8 specs (MZUSP 2940), 24 Jun 2008; 23°1’33”S, 40°45’22”W, 964 m: 3 specs (MZUSP 2944), 16 Jan 2009; 23°12’59”S, 40°57’35”W, 761 m: 1 spec. (MZUSP 2943), 29 Jan 2009; 23°13’2”S, 40°57’36”W, 724 m: 12 specs, 24 Jun 2008; 23°18’28”S, 40°24’52”W, 2513 m: 2 specs (MZUSP 2498), 16 Fev 2009; 23°3’33”S, 40°41’53”W, 1285 m: 2 specs (MZUSP 2946), 17 Jan 2009; 23°39’20”S, 41°18’28”W, 701 m: 1 spec., 28 Jan 2009; 23°41’7”S, 41°16’4”W, 1017 m: 1 spec., 12 Jan 2009; 23°45’8”S, 41°11’51”W, 1319 m: 3 specs (MZUSP 2942), 13 Jan 2009; 23°45’18”S, 39°59’57”W, 2423 m: 1 spec., 6 Fev 2009; 24°1’24”S, 40°54’13”W, 2493 m: 1 spec., 17 Feb 2009; 24°29’15”S, 40°23’22”W, 2981 m: 1 spec., 19 May 2008; 24°29’21”S, 40°23’20”W, 3028 m: 1 spec., 5 Fev 2009. Project ‘AMBES’. 19°3’10”S, 37°45’28”W, 2954 m: 3 specs, 23 Jun 2013; 19°3’30”S, 37°48’46”W, 1347 m: 8 specs (MZUSP 2960), 1 Jul 2013; 19°34’20”S, 38°41’19”W, 449 m: 1 spec., 30 Jun 2013; 19°47’2”S, 39°3’14”W, 1251 m: 6 specs (MZUSP 2959), 26 Jun 2013; 19°53’31”S, 39°32’56”W, 1023 m: 12 specs (MZUSP 2934), 13 Jan 2012; 19°54’4”S, 39°22’29”W, 1335 m: 8 specs (MZUSP 2958), 27 Jun 2013; 20°48’37”S, 38°45’28”W, 2494 m: 1 spec. (MZUSP 2953), 15 Jun 2013; 20°53’28”S, 38°21’21”W, 3000 m: 1 spec. (MZUSP 2955), 12 Jun 2013; 21°4’43”S, 40°4’12”W, 1295 m: 9 specs (MZUSP 2927), 9 Jun 2013. Additional material examined. Braniella palpata—Argentina, Tierra del Fuego, Cape Horn, Drake Passage (56°06’29”S, 66°24’29”W, 384–494 m deep): 1 spec. (USNM 55505, holotype), coll. University of Southern California (Department of Zoology), ‘Eltanin R/V’, 18 Sep 1963, det. O. Hartman, 1967; Chile, Drake Passage (57°13’01”S, 70°57’W, 3770–3806 m deep): 1 spec. (USNM 55506, paratype), coll. University of Southern California (Department of Zoology), ‘Eltanin R/V’, 4 Nov 1962, det. O. Hartman, 1967; Antarctica, Weddell Sea (72°47’36”S, 30°29’42”W, 3697 m): 1 spec. (USNM 46847), coll. University of Connecticut, ‘Glacier II R/V’, 14 Mar 1969, det. O. Hartman; Antarctica, Coats Land (73°28’24”S, 30°26’24”W, 3111 m deep): 3 specs (USNM 46846), coll. University of Connecticut, ‘Glacier II R/V’, 13 Mar 1969, det. O. Hartman. Description. Usually of relatively small body size, complete specimens with 11 chaetigers, largest individual analysed 4.3 mm long, 0.5 mm wide. Palps elongate, distally acute, basally fused to 1/2–3/4 of their lengths (Fig. 12A–B, D–E). Prostomium ovate, conspicuously shorter than palps, wider than long (Fig. 12A, D); eyes absent, antennae ovate, broken or missing in most specimens, median antenna inserted approximately at middle of prostomium, lateral antennae inserted more anteriorly. Peristomium shorter than anterior chaetigers, frequently densely covered by cilia, with 1 pair of small, papilliform peristomial cirri. Dorsal cirri with short cirrophores and long, slender cirrostyles often coiled over dorsum, most cirri broken or missing; dorsal cirri absent on chaetiger 2. Ventral cirri papiliform to digitiform, progressively more pyriform towards posterior body, inserted at midlength of parapodial lobes, not reaching their tips (Fig. 12B, F). Parapodial lobes distally rounded to slightly bilobed, with posterior lobe more pronounced; dorsalmost chaetae emerging from tip of lobe, below semicircular flap located dorsally on distal half of parapodial lobe (Fig. 12F, H), chaetae progressively inserted more basally along lobe ventralwards, with numerous rows of chaetae, especially on anterior chaetigers (Fig. 12B, F, I). Anterior parapodia 422 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. FIGURE 12. Anguillosyllis palpata, SEM. (A) entire worm, dorsal view; (B) anterior body, ventral view; (C) intermediate compound chaeta, chaetiger 8 (mid- to posterior body); (D–E) close ups of the anterior end, dorsal and ventral views, respectively; (F, I) anteriormost and posteriormost parapodia, respectively, left-hand side, dorsal view; (G) ventralmost compound chaeta, chaetiger 6 (midbody); (H) close up of the tip of parapodial lobe of chaetiger 3, dorsal view; (J) close up of the tips of compound chaetae, anterior body. with 30+ compound chaetae each, posterior parapodia with ~10–15 chaetae each; chaetae heterogomph, with long, unidentate, finely spinulated to smooth blades (spinulation nearly inconspicuous under compound microscope); conspicuous dorso-ventral gradation in length, blades 320–30 μm, 450–30 μm and 350–20 μm long on anterior, mid- and posterior body chaetigers, respectively (Fig. 12C, G). Parapodia throughout usually with 2 aciculae each, aciculae away from each other at tip of parapodium, one usually extending anteriorly and other posteriorly to DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 423 chaetae, in dorsal side of parapodium, frequently with tips protruding from parapodial lobes (Fig. 12H); aciculae subdistally slightly enlarged, distally acute. Pharynx through 2–3.5 chaetigers, border surrounded by ~10 soft, distally rounded papillae, tooth absent. Proventricle barrel- to heart-shaped, through ~3 chaetigers (0.2–0.7 mm long), with ca. 12 rows of muscle cells. Remarks. In the original description, this species was illustrated as having dorsal cirri on chaetiger 2 (Hartman 1967, Plate 14), although this was not mentioned in the description, and it was considered as such by Aguado & San Martín (2008). This, however, could not be confirmed in any of the specimens herein analysed, including the holotype and paratypes. Therefore, we amend the description of the species, to state that dorsal cirri are absent on chaetiger 2. Anguillosyllis palpata differs from A. pupa as this latter species has completely or almost completely fused palps and proventricle with 20–25 rows of muscle cells, while in A. palpata the proventricle has 12 rows of muscle cells. Anguillosyllis palpata also differs from A. capensis, as this species has more rows of muscle cells in the proventricle (30 rows) and parapodial lobes with distinctly larger, elongate post-chaetal lobes (see Aguado & San Martín 2008, Fig. 1). Geographic distribution and bathymetric range. South Atlantic Ocean—Argentina (Cape Horn – type locality, 384–494 m deep); Brazil (Campos and Espírito Santo basins, 393–3028 m deep). South Pacific Ocean— Chile (3770–3806 m deep). Antarctic Ocean—Weddell Sea (3111–3697 m deep). Anguillosyllis lanai sp. nov. Figure 13–14 Type material. Project ‘Habitats’. Holotype (MNRJP 1186): 22°49’22”S, 40°8’19”W, 1864 m, 11 May 2008; Project ‘Ambes’: Paratype (ZUEC POL 19880): 19°42’01.8”S, 39°22’21.4”W, 1333 m.). Material examined. Project ‘Habitats’: 23°25’19”S, 40°35’37’W, 2491 m: 1 spec. (MZUSP 2947), 17 Feb 2009; 23°3’34”S, 40°41’55”W, 1285 m: 1 spec. (MZUSP 2945), 16 Jan 2009; 23°8’25”S, 40°36’40”W, 1954 m: 1 spec., 27 Jan 2009. Project ‘AMBES’: 19°3’13”S, 37°45’37”W, 2426 m: 3 specs (MZUSP 2919), 6 Dec 2011; 19°3’45”S, 37°47’28”W, 1928 m: 3 specs (MZUSP 2935), 29 Jan 2012; 19°3’55”S, 37°45’8”W, 2993 m: 1 spec. (MZUSP 2920), 5 Dec 2011; 19°3’55”S, 39°45’8”W, 2993 m: 2 specs (ZUEC 19884), 5 Dec 2011; 19°40’8”S, 39°7’22”W, 1035 m: 1 spec., 13 Dec 2011; 20°25’16”S, 39°27’20”W, 1918 m: 2 specs (MZUSP 2931), 7 Jan 2012; 20°29’3”S, 38°23’15”W, 2504 m: 3 specs (MZUSP 2923), 22 Dec 2011; 20°41’33”S, 39°35’14”W, 1914 m: 2 specs (MZUSP 2921), 27 Dec 2011; 20°49’23”S, 38°17’11”W, 2997 m: 2 specs (MZUSP 2922), 23 Dec 2011; 20°54’14”S, 38°56’10”W, 2519 m: 1 spec. (MZUSP 2930), 4 Jan 2012; 20°8’42”S, 39°7’29”W, 1922 m: 1 spec. (MZUSP 2933), 6 Jan 2012; 20°8’45”S, 39°7’31”W, 1927 m: 5 specs (MZUSP 2954), 16 Jun 2013; 21°36’42”S, 39°49’25”W, 1333 m: 6 specs (MZUSP 2926), 8 Jan 2012; 21°4’51”S, 40°4’14”W, 1300 m: 4 specs (MZUSP 2924 m), 31 Dec 2011; 21°6’30”S, 39°38’36”W, 1889 m: 2 specs (MZUSP 2925), 31 Dec 2011; 21°6’38”S, 39°38’31”W, 1889 m: 1 spec. (MZUSP 2928), 9 Jun 2013; 21°9’39”S, 38°52’7”W, 2502 m: 1 spec. (MZUSP 2929), 10 Jun 2013; 22°49’22”S, 40°8’19”W, 1864 m: 1 spec., 11 May 2008. Additional material examined: Anguillosyllis capensis—South Africa, off Cape Province (34º51’S, 23º41’E, 183 m deep): 1 spec. (BMNH 1963.1.29, holotype), coll. J.H. Day, 30 Nov 1960, det. J.H. Day, 1963. Braniella pupa—USA, off New England: 1 spec. (ZMH P-13585, paratype?), coll. Allan Hancock Foundation, det. Hartman & Fauchald, 1971; USA, off Delaware (38°44’36”N 73°03’06”W, 183 m deep): 1 spec. (USNM 56762), coll. “VIMS for BLM/MMS”, 19 Mar 1976, det. G.R. Gaston; USA, Georges Bank, Northern Slope (41°33’26”N 68°58’36”W, 117 m deep): 1 spec. (USNM 103505), coll. BLM/MMS, 27 Feb 1977, det. University of Delaware. Description. Fragile bodies, usually missing antennae and most cirri, complete specimens with 10 chaetigers, largest specimen examined complete, 2 mm long, 0.2 mm wide. Palps elongate, distally acute, completely fused or with only minute distal notch, occasionally with faint line of fusion. Prostomium ovate, broader than long; eyes absent, antennae cirriform to ovoid (Fig. 13A–B; 14A). Peristomium shorter than subsequent segments, with 1 pair of ovate, papilliform peristomial cirri, smaller than lateral antennae (Fig. 13A–B; 14A). Dorsal cirri long, slender, often coiled over dorsum, most cirri broken or missing, not present on chaetiger 2 (Fig. 13A, I), frequently only cirrophores left; parapodial glands as slightly swollen areas dorsally, close to bases of dorsal cirri, with granular 424 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. FIGURE 13. Anguillosyllis lanai sp. nov., SEM. (A) entire worm, dorsal view; (B–C) close ups of the anterior end, dorsal view, pharynx everted; (D) parapodium, dorsal view, arrow points to acicula; (E–F) falciger (left) and spiniger-like chaeta (right), anterior and midbody chaetigers, respectively; (G) compound chaetae; (H) granules of parapodial glands; (I) posterior end, dorsal view. DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 425 FIGURE 14. Anguillosyllis lanai sp. nov. (A) entire worm, dorsal view; (B) midbody parapodium, posterior view; (C–D) falcigers and spiniger-like chaetae, respectively; (E) tip of acicula; SEM: (F) left-hand parapodia, chaetigers 2–3, dorsal view; (G) close up of a parapodial gland, chaetiger 3. material of unknown nature, approximately rounded to polyhedral under SEM (Fig. 13A, H; 14A, F–G). Ventral cirri digitiform, inserted at mid-length of parapodial lobes, not reaching tip (Fig. 14B). Parapodia distally rounded to weakly bilobed, in such case posterior lobe progressively slightly larger towards posterior body, or only short, nearly inconspicuous posterior lobe present. Anterior and midbody parapodia with ~15 compound chaetae each, posterior body with ~5–10 compound chaetae each; chaetae heterogomph, with long, unidentate, finely spinulated 426 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. to smooth blades (spinulation nearly inconspicuous under compound microscope); conspicuous dorso-ventral gradation in length, blades 160–7 μm, 170–30 μm and 170–20 μm long on anterior, mid- and posterior body chaetigers, respectively (Fig. 13E–G; 14C–D). Parapodia throughout usually with 2 aciculae each, aciculae away from each other at tip of parapodium, one usually extending anteriorly and other posteriorly to chaetae, frequently with tips protruding from parapodial lobes (Fig. 13D); aciculae subdistally slightly enlarged, distally acute (Fig.14E). Pharynx through 1.5–2 chaetigers, border surrounded by ~10 soft, elongate, digitiform papillae (Fig, 13C), tooth absent. Proventricle barrel-shaped to heart-shaped, through 2.5–3 chaetigers (0.25–0.35 mm long), with ca. of 12 rows of muscle cells (Fig. 14A). Remarks. Anguillosyllis lanai sp. nov. has completely fused palps, parapodial lobes distally rounded, bilobed or with posterior lobe only, and structures near parapodial bases which we consider as parapodial glands, because their location is in agreement with parapodial glands found in other species of syllids. The species most similar morphologically to A. lanai sp. nov. is A. pupa, since both species have completely or nearly completely fused palps and ventral cirri inserted at mid-length of parapodial lobes. However, A. lanai sp. nov. differs from A. pupa as this latter species has a proventricle with 20–25 rows of muscle cells (against ~12 rows as in A. lanai sp. nov.); and post-chaetal lobes conspicuously larger than in A. lanai sp. nov. (see Hartman 1965, Pl. 8; Aguado & San Martín 2008, Fig. 2A–B). Anguillosyllis lanai sp. nov. also differs from A. capensis, since in that species the proventricle presents more rows of muscle cells (30 rows), and the parapodial lobes have distinctly larger, elongate post-chaetal lobes (see Aguado & San Martín 2008, Fig. 1). Finally, A. lanai sp. nov. differs from A. palpata because the latter species presents palps separated from each other for at least half of their length, longer blades of compound chaetae (up to 450 μm long, as opposed to 170 μm long, as in A. lanai sp. nov.), and by the adults of that species having 11, instead of 10 chaetigers, as in A. lanai sp. nov. Geographic distribution and bathymetric range. Anguillosyllis lanai sp. nov. was found in Campos and Espírito Santo basins, between 1035–2997 m deep. Etymology. This species is dedicated to Paulo Lana, responsible for an enormous increase in the knowledge of marine invertebrates, especially polychaetes, inspiring new generations of researchers. Conclusion This study showed how the deep fauna of southwestern Atlantic is still unknown, since most of the species herein identified were considered as new to science or new reports to the region. Taxonomic studies are fundamental to the understanding of the biota and the community as well as a fundamental tool to biogeographical and bathymetrical approaches. More taxonomic studies should reveal much more species of syllids inhabiting the deep region of southwestern Atlantic. Acknowledgements We are grateful to CENPES/PETROBRAS for providing the samples for this study, and to Enio Matos, Phillip Lenktaitis (IB/USP), and Karla Avelar (NEMA/PUC-Rio) for preparing and photographing specimens under SEM. We also thank all the museum staffs where the material analysed is deposited, namely Drs Stephen Keable (AM), Emma Sherlock and Andrew Cabrinovic (MNH), Aline Staskowian Benetti (MZUSP), Geoff Keel and Karen Osborn (USNM), Angelika Brandt (ZMH), and Tatiana Steiner (ZUEC). RB received a Ph. D. fellowship from Rio de Janeiro Research Foundation (FAPERJ proc. E-26/151.912/2007), MVF received a postdoc fellowship from the São Paulo Research Foundation (FAPESP proc. 2010/19424-7), JMMN and PCP receives a productivity grant from CNPq. Finally, we would like to thank Guillermo San Martín and Patrícia Álvarez-Campos, for the valuable comments and suggestions that greatly improved the final version of this paper. References Aguado, M.T., Nygren, A & Siddall, M.E. (2007) Phylogeny of Syllidae (Polychaeta) based on combined molecular analysis of nuclear and mitochondrial genes. Cladistics, 23, 552–564. DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 427 https://doi.org/10.1111/j.1096-0031.2007.00163.x Aguado, M.T. & San Martín, G. (2008) Re-description of some enigmatic genera of Syllidae (Phyllodocida: Polychaeta). Journal of the Marine Biological Association of the United Kingdon, 88, 35–56. https://doi.org/10.1017/S002531540800026X Aguado, M.T. & San Martín, G. (2009) Phylogeny of Syllidae (Annelida, Phyllodocida) based on morphological data. Zoologica Scripta, 38 (4), 379–402. https://doi.org/10.1111/j.1463-6409.2008.00380.x Aguado, M.T., San Martín, G. & Siddall, M.E. (2012) Systematics and evolution of syllids (Annelida, Syllidae). Cladistics, 28, 234–250. https://doi.org/10.1111/j.1096-0031.2011.00377.x Álvarez-Campos, P., Gil, J. & San Martín, G. (2015a) Unveiling the Rosetta Stone of syllids: Redescription and neotype designation of Syllis monilaris Savigny in Lamarck, 1818, type species of type genus of family Syllidae Grube, 1850 (Annelida). Zootaxa, 4040 (3), 317–330. https://doi.org/10.11646/zootaxa.4040.3.4 Álvarez-Campos, P., Riesgo, A., Hutchings, P. & San Martín, G. (2015b) The genus Syllis Savigny in Lamarck, 1818 (Annelida, Syllidae) from Australia. Molecular analysis and re-description of some poorly-known species. Zootaxa, 4052 (2), 297– 331. Attolini, F. (2002) Padrões de distribuição de poliquetas macrobentônicos na região de plataforma externa e talude superior entre Cabo Frio (RJ) e Cabo de Santa Marta (SC), costa sudeste do Brasil. PhD thesis. Instituto Oceanográfico, Universidade de São Paulo, São Paulo, 115 pp. Audouin, J.V. & Milne Edwards, H. (1833) Classification des Annélides et description de celles qui habitent les côtes de la France. Annales des sciences naturelles, 30, 411–425. https://doi.org/10.5962/bhl.part.8010 Banse, K. (1959) Über die Polychaeten-Besiedlung einiger submariner Höhlen. Ergebnisse der Österreichischen TyrrheniExpedition 1952, Teil XII. Pubblicazioni della Stazione Zoologica di Napoli, 30 (Supplement), 417–469. Barroso, R. (2011) Anelídeos poliquetas das famílias Amphinomidae, Opheliidae, Syllidae e Paraonidae de oceano profundo da Bacia de Campos, Sudeste do Brasil. PhD thesis. Museu Nacional da Universidade Federal do Rio de Janeiro, Universidade Federal do Rio de Janeiro, Rio de Janeiro, 174 pp. Benham, W.B. (1921) Polychaeta. Scientific Reports of the Australasian Antarctic Research Expedition, 1911–1914, C (Zoology and Botany), 6, 1–128. https://doi.org/10.5962/bhl.title.31708 Benham, W.B. (1927) Polychaeta. British Antarctic 'Terra Nova' Expedition Natural History Reports. Zoology, 7 (2), 47–182. Böggemann, M. (2009). Polychaetes (Annelida) of the abyssal SE Atlantic. Organisms Diversity & Evolution, 9, 252–428. Böggemann, M. & Purschke, G. (2005) Abyssal benthic Syllidae (Annelida: Polychaeta) from the Angola Basin. Organisms, Diversity & Evolution, 5, 221–226. https://doi.org/10.1016/j.ode.2004.11.006 Böggemann, M. & Westheide, W. (2004) Interstitial Syllidae (Annelida: Polychaeta) from Mahé (Seychelles). Journal of Natural History, 38 (4), 403–446. https://doi.org/10.1080/0022293021000033229 Claparède, E. (1863) Beobachtungen über Anatomie und Entwicklungsgeschichte wirbelloser Thiere an der Küste von Normandie angestellt. Wilhelm Engelmann, Leipzig, 120 pp. https://doi.org/10.5962/bhl.title.10030 Claparède, E. (1864) Glanures zootomiques parmi les annélides de Port-Vendres (Pyrénées Orientales). Mémoires de la Societé de Physique d´Histoire Naturelle de Genève, 17 (2), 463–600. Claparède, E. (1868) Les annélides chétopodes du Golfe de Naples. Mémoires de la Société de Physique et d'Histoire Naturelle de Genève, 19 (2), 313–584. Day, J.H. (1963) The polychaete fauna of South Africa. Part 8. New species and records from grab samples and dredgings. Trustees of the British Museum, 10, 383–443. https://doi.org/10.5962/bhl.part.20530 Desbruyeres, D. & Segonzac, M. (1997) Handbook of deep-sea hydrothermal vent fauna. IFREMER, Brest, 278 pp. Ehlers, E. (1901) Die Polychaeten des magellanischen und chilenischen Strandes. Ein faunistischer Versuch. Festschrift zur Feier des Hundertfünfzigjährigen Bestehens des Königlichen Gesellschaft der Wissenschaften zu Göttingen, Abhandlungen der Mathematisch-Physikalischen Klasse, 1901, 1–232. Fukuda, M.V. (2010) Contribuição ao conhecimento taxonômico dos silídeos Polychaeta: Syllidae) da região sudeste-sul do Brasil. PhD thesis, Instituto de Biociências, Universidade de São Paulo, São Paulo, 340 pp. Fukuda, M.V. & Nogueira, J.M.M. (2006) A new species of Odontosyllis Claparède, 1863 (Polychaeta: Syllidae: Eusyllinae), and description of Brazilian material of Odontosyllis cf. fulgurans (Audouin & Milne Edwards, 1834). Zoological Studies, 45 (2), 223–233. Fukuda, M.V. & Nogueira, J.M.M. (2013) First occurrence of Perkinsyllis San Martín, López & Aguado, 2009 (Polychaeta: Syllidae) in Brazil, with descriptions of two new species. Marine Biology Research, 9 (10), 972–989. https://doi.org/10.1080/17451000.2013.793807 428 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL. Fukuda, M.V., Nogueira, J.M.M., Paresque, K. & San Martín, G. (2013) Species of Odontosyllis Claparède, 1863 (Annelida: Polychaeta: Syllidae) occurring along the Brazilian coast. Zootaxa, 3609 (2), 142–162. https://doi.org/10.11646/zootaxa.3609.2.2 Fukuda, M.V., Nogueira, J.M.M. & San Martín, G. (2015) Eusyllinae and “Incertae sedis” syllids (Annelida: Syllidae) from South America, with a new species from Brazil and a new combination for a Peruvian species. Zootaxa, 3936 (4), 507– 537. https://doi.org/10.11646/zootaxa.3936.4.3 Fukuda, M.V., Yunda-Guarín, G. & Nogueira, J.M.M. (2009) The genus Prosphaerosyllis (Polychaeta: Syllidae: Exogoninae) in Brazil, with description of a new species. Journal of the Marine Biological Association of the United Kingdom, 89 (7), 1443–1454. https://doi.org/10.1017/S0025315409000095 Gil, J. & Musco, L. (2015) Syllidae Grube, 1850. In: Read, G. & Fauchald, K. (2016) World Polychaeta Database. Accessed through: Glover, A.G., Higgs, N. & Norton, T. (2016) World register of deep-sea species. Available from: http:// www.marinespecies.org/DeepSea/aphia.php?p=taxdetails&id=948 (accessed 10 November 2016) Grube, A.E. (1850) Die Familien der Anneliden. Archiv für Naturgeschichte, 16 (1), 249–364. Grube, A.E. (1860) Beschreibung neuer oder wenig bekannter Anneliden. Fünfter Beitrag. Archiv für Naturgeschichte, 26 (1), 71–118. Hartman, O. (1953) Non-pelagic Polychaeta of the Swedish Antarctic Expedition 1901-1903. Further Zoological Results of the Swedish Antarctic Expedition 1901–1903, 4 (11), 1–83. Hartman, O. (1956) Polychaetous annelids erected by Treadwell, 1891 to 1948, together with a brief chronology. Bulletin of the American Museum of Natural History, 109 (2), 239–310. Hartman, O. (1965) Deep-water benthic polychaetous annelids off New England to Bermuda and other North Atlantic areas. Occasional Papers of the Allan Hancock Foundation, 28, 1–378. Hartman, O. (1967) Polychaetous annelids collected by the USNS Eltanin and Staten Island cruises, chiefly from Antarctic Seas. Allan Hancock Monographs in Marine Biology, 2, 1–387. Hartmann-Schröder, G. (1960) Polychaeten aus dem Roten Meer. Kieler Meeresforschungen, 16, 69–125. Hartmann-Schröder, G. (1965) Teil II. Die Polychaeten des Sublitorals. In: Hartmann-Schröder, G. & Hartmann, G. (Eds.), Zur Kenntnis des Sublitorals der chilenischen Küste unter besonderer Berücksichtigung der Polychaeten und Ostracoden. Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut, 62, pp. 59–305. Hartmann-schröder, G. (1993) Die Polychaeten der "Polarstern" -Reise ANTX/1b zur Antarktischen Halbinsel und Isla de los Estados (Feuerland, Argentinien) 1991. Teil 1: Polynoidae bis Iphitimidae. Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut, 90, 127–150. Hartmann-Schröder, G. & Rosenfeldt, P. (1988) Die Polychaeten der Polarstern-Reise ANT III/2 in die Antarktis 1984. Teil 1: Euphrosinidae bis Chaetopteridae. Mitteilungen aus dem Hamburgischen zoologischen Museum und Institut, 85, 25–72. Kudenov, J.D. & Harris, L.H. (1995) Family Syllidae Grube, 1850. In: Blake, J., Hilbig, B.Y. & Scott, P.H. (Eds.), Taxonomic Atlas of the Benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel. Santa Barbara Museum of Natural History, Santa Barbara, pp. 1–97. Lamarck, J.B.M. (1818) Histoire Naturelle des animaux sans vertèbres, presentant les caractères generaux et particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres, et la citation des principales espèces qui s’y rapportent; précédés d’une introduction offrant la détermination des caractères essentiels de l’animal, sa distinction du végétal et des autres corps naturels, enfin l’exposition des principes fondamentaux de la zoologie. Tome cinquième. Deterville/Verdière, Paris, 612 pp. Lanera, P., Sordino, P. & San Martín, G. (1994) Exogone (Parexogone) gambiae, a new species of Exogoninae (Polychaeta, Syllidae) from the Mediterranean Sea. Bollettino di Zoologia, 61, 235–240. https://doi.org/10.1080/11250009409355891 Langerhans, P. (1879) Die Wurmfauna von Madeira [part I]. Zeitschrift für wissenschaftliche Zoologie, 32 (4), 513–592. Licher, F. (1999) Revision der Gattung Typosyllis Langerhans, 1879 (Polychaeta: Syllidae). Morphologie, Taxonomie und Phylogenie. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 551, 1–336. Lucas, Y., San Martín, G. & Parapar, J. (2012) Two new species of Syllidae (Annelida : Polychaeta) from DIVA-Artabria I project (cruise 2002) to deep areas off NW Spain. Zootaxa, 3589, 77–88. Marenzeller, E.V. (1875) Zur Kenntniss der adriatischen Anneliden. Zweiter Beitrag. (Polynoinen, Hesioneen, Syllideen). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Wien, Mathematisch-Naturwissenschaftliche Classe, 72 (1), 129–171. McIntosh, W.C. (1885) Report on the Annelida Polychaeta collected by H.M.S. Challenger during the years 1873-1876. Report on the Scientific Results of the Voyage of H.M.S.Challenger during the years 1872-76, Zoology, 12, 1–554. Mesnil, F. & Caullery, M. (1918) Sur l'organisation et la biologie d'un Syllidien [Exogone (Parexogone, n. s. g.) hebes Webst. et Ben. var. hibernica South.] habitant un sable compact. Bulletin de la Société Zoologique de France, 42 (Volume for 1917), 126–132. Monro, C.A. (1939) Polychaeta. B.A.N.Z. Antarctic Research Expedition Reports, Ser. B Zoology and Botany, 4 (4), 87–156. Musco, L., Çinar, M.E. & Giangrande, A. (2005) A new species of Sphaerosyllis (Polychaeta, Syllidae, Exogoninae) from the coasts of Italy and Cyprus (eastern Mediterranean Sea). Italian Journal of Zoology, 72, 161–166. https://doi.org/10.1080/11250000509356666 Nogueira, J.M.M. & San Martín, G. (2002) Species of Syllis Savigny in Lamarck, 1818 (Polychaeta: Syllidae) living in corals in DEEP SEA SYLLIDAE FROM SOUTHWESTERN ATLANTIC Zootaxa 4221 (4) © 2017 Magnolia Press · 429 the state of São Paulo, southeastern Brazil. Beaufortia, 52 (7), 57–93. Nogueira, J.M.M., San Martín, G. & Amaral, A.C.Z. (2001) Description of five new species of Exogoninae Rioja, 1925 (Polychaeta: Syllidae) associated with the stony coral Mussismilia hispida (Verrill, 1868) in São Paulo State, Brazil. Journal of Natural History, 35 (12), 1773–1794. https://doi.org/10.1080/00222930152667096 Nogueira, J.M.M., San Martín, G. & Fukuda, M.V. (2004) On some exogonines (Polychaeta, Syllidae, Exogoninae) from the northern coast of the State of São Paulo, southeastern Brazil Results of BIOTA/FAPESP/bentos marinho Project. Meiofauna marina, 13, 45–77. Nogueira, J.M.M. & Yunda-Guarin, G. (2008) A new species of Syllis (Polychaeta: Syllidae: Syllinae) from off Fortaleza, north-eastern Brazil. Journal of the Marine Biological Association of the United Kingdom, 88 (7), 1391–1399. https://doi.org/10.1017/S0025315408002099 Ørsted, A.S. (1845) Ueber die Entwicklung der Jungen bei einer Annelide und über die äusseren Unterschiede zwischen beiden Geschlechtern. Archiv für Naturgeschichte, 11 (1), 20–23. Paresque, K., Fukuda, M.V. & Nogueira, J.M.M. (2014) The genus Exogone (Polychaeta: Syllidae) from the Brazilian coast, with the description of a new species. Zootaxa, 3790 (4), 501–533. https://doi.org/10.11646/zootaxa.3790.4.1 Perkins, T.H. (1981) Syllidae (Polychaeta), principally from Florida, with descriptions of a new genus and twenty-one new species. Proceedings of the Biological Society of Washington, 93 (4), 1080–1172. Russel, D.E. (1989) Three new species of Sphaerosyllis (Polychaeta: Syllidae) from Mangrove habitats in Belize. Zoologica Scripta, 18, 375–380. https://doi.org/10.1111/j.1463-6409.1989.tb00132.x San Martín, G. (1984a) Descripción de una nueva especie y revisión del género Sphaerosyllis (Polychaeta: Syllidae). Cahiers de Biologie Marine, 25, 375–391. San Martín, G. (1984b) Estudio biogeográfico, faunístico y sistemático de los Poliquetos de la família Sílidos (Syllidae: Polychaeta) en Baleares. Publicaciones de la Universidad Complutense de Madrid, 187, 1–581. San Martín, G. (1990) Eusyllinae (Syllidae, Polychaeta) from Cuba and Gulf of Mexico. Bulletin of Marine Science, 46 (3), 590–619. San Martín, G. (1991) Grubeosyllis and Exogone (Exogoninae, Syllidae, Polychaeta) from Cuba, the Gulf of Mexico, Florida and Puerto Rico, with a revision of Exogone. Bulletin of Marine Science, 49 (3), 715–740. San Martín, G. (2003) Annelida Polychaeta II: Syllidae. In: Ramos, M.A. (Ed.), Fauna Ibérica. Vol. 21. Museo Nacional de Ciências Naturales, CSIC, Madrid, pp. 1–544. San Martín, G. (2004) Deep sea Syllidae from the Pacific Ocean, collected during cruises with the RV Sonne (Annelida, Polychaeta, Syllidae). Senckenbergiana biologica, 84 (1/2), 13–25. San Martín, G. (2005) Exogoninae (Polychaeta, Syllidae) from Australia with the description of a new genus and twenty-two new species. Records of the Australian Museum, 57, 39–152. https://doi.org/10.3853/j.0067-1975.57.2005.1438 San Martín, G. & Aguado, M.T. (2014) Family Syllidae. In: Schmidt-Rhaesa, A. (Chief Ed.), Phyllodocida: Nereidiformia. Handbook of Zoology, Annelida. A Natural History of the Phyla of the Animal Kingdom. Verlag Walter der Gruyter GmbH & Co., Zürich, pp. 1–52. San Martín, G., Ceberio, A. & Aguirrezabalaga, F. (1996) Exogone species (Polychaeta: Syllidae: Exogoninae) from the Capbreton Canyon (Bay of Biscay, NE Atlantic). Cahiers de Biologie Marine, 37, 249–258. San Martín, G. & Hutchings, P.A. (2006) Eusyllinae (Polychaeta, Syllidae) from Australia with the description of a new genus and fifteen new species. Records of the Australian Museum, 58, 257–370. https://doi.org/10.3853/j.0067-1975.58.2006.1466 San Martín, G. & López, E. (2002) New species of Autolytus Grube, 1850, Paraprocerastea San Martín & Alós, 1989, and Sphaerosyllis Claparède, 1863 (Syllidae, Polychaeta) from the Iberian Peninsula. Sarsia, 87 (2), 135–143. https://doi.org/10.1080/003648202320205210 San Martín, G., López, E. & Aguado, M.T. (2009) Revision of the genus Pionosyllis (Polychaeta: Syllidae: Eusyllinae), with a cladistic analysis, and the description of five new genera and two new species. Journal of the Marine Biological Association of the United Kingdom, 89 (7), 1455–1498. https://doi.org/10.1017/S0025315409003099 San Martín, G. & Parapar, J. (1997) “Errant” polychaetes of the Livingston Island shelf (South Shetlands, Antarctica), with the description of a new species. Polar Biology, 17, 285–295. https://doi.org/10.1007/PL00013370 San Martín, G. & Worsfold, T.M. (2015) Guide and keys for the identification of Syllidae (Annelida, Phyllodocida) from the British Isles (reported and expected species). ZooKeys, 488, 1–29. https://doi.org/10.3897/zookeys.488.9061 Southern, R. (1914) Clare Island Survey. Archiannelida and Polychaeta. Proceedings of the Royal Irish Academy, 31 (47), 1– 160. Uebelacker, J.M. (1984) Family Syllidae Grube, 1850. In: Uebelacker, J.M. & Johnson, P.G. (Eds.), Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Vol. IV. Barry A. Vittor & Associates, Metairie, pp. 1–151. [pp. 30-1–30-151] Webster, H.E. & Benedict, J.E. (1884) The Annelida Chaetopoda from Provincetown and Wellfleet, Massachusetts. Annual Report of the United States Commission of Fish and Fisheries, Washington, 1881, 699–747. 430 · Zootaxa 4221 (4) © 2017 Magnolia Press BARROSO ET AL.

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Deep sea Syllidae (Annelida, Phyllodocida) from Southwestern Atlantic

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