On the taxonomic status of the European Scolelepis (Scolelepis) squamata (Polychaeta: Spionidae), with description of a new species from southern Europe

Victor  Surugiu

In order to clarify taxonomic problems relating to the identity of Scolelepis (Scolelepis) specimens from the Black Sea, the identified material was compared with specimens of Scolelepis cirratulus (Delle Chiaje, 1829) from the Mediterra-nean, of S. squamata (Abildgaard, in O.F. Müller, 1806) from the North Sea and the Atlantic coast of Spain, and with the syntypes of S. mesnili (Bellan & Lagardère, 1971) from the Atlantic coast of France. The examination of a large number of specimens (both juveniles and adults) reveals that the currently accepted morphological differences distinguishing all species show size-related patterns, suggesting that they all belong to one species. Therefore, this study supports the view that Scolelepis cirratulus and Scolelepis mesnili are junior synonyms of Scolelepis (Scolelepis) squamata. As a result of the re-assessment of the species limits of Scolelepis (Scolelepis) squamata, a new species, Scolelepis (Scolelepis) neglecta sp. nov., is distinguished and described from the Cantabrian coast of Spain. It inhabits shallow sublittoral fine sands and was earlier misidentified and reported from the Atlantic coast of Europe and the Mediterranean Sea as Scolelepis squa-mata, Scolelepis mesnili, Scolelepis cantabra (Rioja, 1918), or Dispio uncinata Hartman, 1951. The new species is characterized by having a trilobate prostomium with an acuminate medial portion, a short peristomium with well-developed dorso-lateral wings, short palps with two longitudinal bands of elevated lobes, neuropodial postchaetal lamellae notched from chaetigers 14–41, and strongly curved bidentate neuropodial hooded hooks with a slight constriction on the shaft starting from chaetigers 19–49. The morphology, diagnostic characters and ecology of both Scolelepis (Scolelepis) squa-mata and Scolelepis (Scolelepis) neglecta sp. nov. are discussed. Introduction The genus Scolelepis represents a group of spionid polychaetes that share the presence of a pointed prostomium, branchiae beginning from chaetiger 2, the lack of ciliated grooves on palps, and a cushion-like pygidium without cirri. The genus Scolelepis (sensu lato), currently comprising around 85 species, is commonly divided into two subgenera, Scolelepis Blainville, 1828 (sensu stricto) with approximately 73 species and Parascolelepis Maciolek, 1987 with 12 species (Sikorski & Pavlova 2015; Meißner & Götting 2015; Read 2016). The diagnostic characters that have proven to be relevant in distinguishing species of Scolelepis (sensu lato) are those related to the shape of the prostomium, the presence and size of lateral wings on the peristomium, the shape of the basal palpal sheaths, the palp ciliation pattern, the presence or absence of notochaetae on chaetiger 1, the shape of postchaetal lamellae of chaetiger 1, the degree of branchial fusion, the shape and the distribution of hooded hooks, the nature of notopodial capillaries, the ratio of the length of the hood to main fang, the shape of the anterior and posterior neuropodial postchaetal lamellae, the shape of the posterior notopodial postchaetal lamellae, and the shape of the pygidium

Zootaxa 4161 (2): 151–176 http://www.mapress.com/j/zt/ ISSN 1175-5326 (print edition) Article Copyright © 2016 Magnolia Press ZOOTAXA ISSN 1175-5334 (online edition) http://doi.org/10.11646/zootaxa.4161.2.1 http://zoobank.org/urn:lsid:zoobank.org:pub:B36A20C2-A9DA-4F35-89A3-2153F88673BC On the taxonomic status of the European Scolelepis (Scolelepis) squamata (Polychaeta: Spionidae), with description of a new species from southern Europe VICTOR SURUGIU Universitatea “Alexandru Ioan Cuza” Iaşi, Facultatea de Biologie, B-dul Carol I, nr. 20A, 700507 Iaşi, România, fax: +40 232 201472. E-mail: vsurugiu@uaic.ro Abstract In order to clarify taxonomic problems relating to the identity of Scolelepis (Scolelepis) specimens from the Black Sea, the identified material was compared with specimens of Scolelepis cirratulus (Delle Chiaje, 1829) from the Mediterranean, of S. squamata (Abildgaard, in O.F. Müller, 1806) from the North Sea and the Atlantic coast of Spain, and with the syntypes of S. mesnili (Bellan & Lagardère, 1971) from the Atlantic coast of France. The examination of a large number of specimens (both juveniles and adults) reveals that the currently accepted morphological differences distinguishing all species show size-related patterns, suggesting that they all belong to one species. Therefore, this study supports the view that Scolelepis cirratulus and Scolelepis mesnili are junior synonyms of Scolelepis (Scolelepis) squamata. As a result of the re-assessment of the species limits of Scolelepis (Scolelepis) squamata, a new species, Scolelepis (Scolelepis) neglecta sp. nov., is distinguished and described from the Cantabrian coast of Spain. It inhabits shallow sublittoral fine sands and was earlier misidentified and reported from the Atlantic coast of Europe and the Mediterranean Sea as Scolelepis squamata, Scolelepis mesnili, Scolelepis cantabra (Rioja, 1918), or Dispio uncinata Hartman, 1951. The new species is characterized by having a trilobate prostomium with an acuminate medial portion, a short peristomium with well-developed dorso-lateral wings, short palps with two longitudinal bands of elevated lobes, neuropodial postchaetal lamellae notched from chaetigers 14–41, and strongly curved bidentate neuropodial hooded hooks with a slight constriction on the shaft starting from chaetigers 19–49. The morphology, diagnostic characters and ecology of both Scolelepis (Scolelepis) squamata and Scolelepis (Scolelepis) neglecta sp. nov. are discussed. Key words: Scolelepis (S.) squamata, Scolelepis (S.) neglecta sp. nov., taxonomy, palp ciliary pattern, ontogenetic variation, chaetal replacement, nuchal organs Introduction The genus Scolelepis represents a group of spionid polychaetes that share the presence of a pointed prostomium, branchiae beginning from chaetiger 2, the lack of ciliated grooves on palps, and a cushion-like pygidium without cirri. The genus Scolelepis (sensu lato), currently comprising around 85 species, is commonly divided into two subgenera, Scolelepis Blainville, 1828 (sensu stricto) with approximately 73 species and Parascolelepis Maciolek, 1987 with 12 species (Sikorski & Pavlova 2015; Meißner & Götting 2015; Read 2016). The diagnostic characters that have proven to be relevant in distinguishing species of Scolelepis (sensu lato) are those related to the shape of the prostomium, the presence and size of lateral wings on the peristomium, the shape of the basal palpal sheaths, the palp ciliation pattern, the presence or absence of notochaetae on chaetiger 1, the shape of postchaetal lamellae of chaetiger 1, the degree of branchial fusion, the shape and the distribution of hooded hooks, the nature of notopodial capillaries, the ratio of the length of the hood to main fang, the shape of the anterior and posterior neuropodial postchaetal lamellae, the shape of the posterior notopodial postchaetal lamellae, and the shape of the pygidium (Mesnil 1896; Pettibone 1963; Light 1977, 1978; Maciolek 1987; Dauer 1987; Blake 1996; Eibye-Jacobsen 1997; Delgado-Blas 2006; Williams 2007; Rocha & Paiva 2012; Zhou 2014; Meißner & Götting 2015). To date, twelve species of the genus Scolelepis (sensu lato) have been reported in European waters: Accepted by J. Williams: 15 Jul. 2016; published: 5 Sept. 2016 151 Scolelepis (Scolelepis) squamata (Abildgaard, in O.F. Müller, 1806) [=Nerine cirratulus (Delle Chiaje, 1829)] S. (S.) foliosa (Audouin & Milne Edwards, 1833) S. (S.) longirostris (Quatrefages, 1843) [species inquirendae] S. (S.) bonnieri (Mesnil, 1896) S. (S.) cantabra (Rioja, 1918) S. (S.) mesnili (Bellan & Lagardère, 1971) S. (S.) laonicola (Tzetlin, 1985) [=S. matsugae Sikorski, 1994] S. (S.) finmarchicus Sikorski & Pavlova, 2015 S. (Parascolelepis) tridentata (Southern, 1914) S. (P.) gilchristi (Day, 1961) S. (P.) korsuni Sikorski, 1994 S. (P.) burkovskii Sikorski, 1994 Among these species, S. (S.) laonicola, S. (S.) finmarchicus, S. (P.) korsuni, and S. (P.) burkovskii are confined to northern European waters (Sikorski & Pavlova 2015). In southern Europe, S. (S.) squamata is the most problematic due to the substantial degree of variation of its morphological characters and is reported to have a cosmopolitan distribution. S. (S.) squamata was originally described (as Lumbricus squamatus) from the German island of Helgoland (then a Danish possession) in the Otto Friedrich Müller’s Zoologia Danica (1806). It is worth mentioning that O.F. Müller died in 1784 and that his fourth volume of the Zoologia Danica was actually edited and completed by P.C. Abildgaard, M. Vahl, J. Rathke, and H.S. Holten (Oug et al. 2014). Peter Christian Abildgaard also died in 1801, but from the contents of the volume it is clear that he was responsible for the description of Lumbricus squamatus as well as for other twelve species. Therefore, according to Art. 50.1 of the International Code of Zoological Nomenclature (ICZN 1999), the authorship must be attributed to P.C. Abildgaard. The original description contains only three sentences in Latin and it seems that Abildgaard did not designate any type material (Danny EibyeJacobsen, pers. comm., Sep 2013; Oug et al. 2014). In 1822 Delle Chiaje illustrated, and in 1829 described, Scolelepis cirratulus (as Lumbricus cirratulus) from the Gulf of Naples, Mediterranean Sea, Italy. The description is also very brief and rather general. The type material of Delle Chiaje could not be located and most likely was not designated (Maria Cristina Gambi, pers. comm., Mar 2014). Though generally recognized as junior synonym of S. (S.) squamata, the latter was widely reported, as Nerine cirratulus, not only for populations in southern European waters (Claparède 1869; Bellan & Lagardère 1971; Marinov 1977), but also for those in northern European waters (Mesnil 1896; Saint-Joseph 1898; McIntosh 1915; Fauvel 1927; Joyner 1962). In reviewing the genus Scolelepis, Pettibone (1963) lumped together, without providing any justification, several species within Scolelepis (Scolelepis) squamata, including Lumbricus cirratulus, Malacoceros longirostris Quatrefages, 1843 from France, Nerine agilis Verrill, 1873 from New Jersey, Spio acuta Treadwell, 1914 from California, Nerine capensis McIntosh, 1925 from South Africa, Nerine minuta Treadwell, 1939 from Texas, and Nerinides goodbodyi Jones, 1962 from Jamaica. This synonymization expanded the character states of the species and extended the distribution range from Europe to South Africa, Madagascar, China, North and South America, leading to greater taxonomic confusion and inconsistency. Based on slight morphological and ecological differences, Bellan & Lagardère (1971) distinguished a new species from the Atlantic coast of Europe, Scolelepis mesnili (as Nerine mesnili). They also stressed that Scolelepis cirratulus from the Mediterranean might be regarded as a separate species from that inhabiting the Atlantic coasts of France, which was previously assigned to Scolelepis squamata. Foster (1971) and Light (1977, 1978) indicated that in the North American populations of Scolelepis squamata there is a great intraspecific variability of characters such as the number of neuropodial hooded hooks, the presence or absence of the notopodial hooded hooks, and the shape of the neuropodial postchaetal lamellae. Recent studies by MacCord & Amaral (2005), Delgado-Blas (2006), and Zhou et al. (2009) have resurrected some species previously synonymised with Scolelepis squamata. This has led to some authors stressing the urgent need for a re-assessment of this taxon (Delgado-Blas 2006; Radashevsky 2012; Meißner & Götting 2015). While resolving taxonomic problems pertaining to the identity of Scolelepis (Scolelepis) squamata reported from the European seas, a new species of Scolelepis has been distinguished and is herein described. 152 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU Materials and methods Scolelepis specimens were collected from littoral and shallow sublittoral sediments along the Romanian and Spanish coasts between June 2010 and August 2015. A PVC-hand-held sediment corer, 10-cm in diameter, was pushed to a depth of 10 cm into the sediment, or sediment was collected with a spade. Samples were carefully washed in the field with sea water through a 0.5 mm mesh sieve and polychaetes retained on the mesh were handpicked with live-insect handling forceps. Immediately, the worms were narcotized with 3.5% magnesium chloride solution in sea water for 10–20 minutes. After a preliminary examination under a stereomicroscope to notice colour and movement patterns, animals were fixed for several days in 10% formaldehyde solution in sea water or for 30 min in 4% glutaraldehyde. Afterwards all material was transferred to 70% ethanol for long-term storage. In addition to the freshly collected material, specimens deposited in museum collections were also examined, including: Muséo Nacional de Ciencias Naturales, Madrid, Spain (MNCN), Muséum National d’Histoire Naturelle, Paris, France (MNHN), “Grigore Antipa” National Museum of Natural History, Bucharest, Romania (MNINGA), Senckenberg Museum, Frankfurt, Germany (SMF), and Zoologische Sammlung der Universität, Rostock, Germany (ZSRO). For light microscopy (LM), specimens were rinsed in tap water and examined under a Nikon SMZ800 stereomicroscope. Chaetal arrangement of small worms and parapodia or other parts of large-bodied specimens were examined under a compound microscope by mounting the relevant tissue in polyvinyl lactophenol for 2–14 days. In order to reveal the methyl green staining pattern, specimens were bathed in a saturated solution of methyl green in 70% ethanol for at least 60 seconds and afterwards de-stained for a few seconds in clean 70% ethanol. Maximum length (excluding palps) and width (including parapodia but excluding chaetae) of worms were measured to the nearest 0.01 mm using a calibrated ocular micrometer mounted on a stereomicroscope. Illustrations were prepared from photographs taken with a Leica DFG500 camera coupled either to a Leica M250A stereomicroscope, or to a Leica DM750 compound microscope. Final plates were compiled with Adobe® Photoshop® software. Graphs were prepared in Microsoft® Excel® and edited in Adobe® Photoshop®. A series of correlation analyses was carried out using morphometric variables observed in specimens of Scolelepis cirratulus from the Black Sea, the Mediterranean Sea, and Morocco, S. squamata from the North Sea and the Atlantic coast of Spain and France, and S. mesnili from the Atlantic coasts of France. Variables included body width and total number of chaetigers (both treated as independent), palp length (referring to the number of the chaetiger they reach back to), hooks in noto- and neuropodia (referring to the numbers of the first hook-bearing chaetigers) and the number of teeth on the hooks. Prior to correlation analyses, the Shapiro-Wilk normality test (W) was applied to the underlying variables. Because the dependent variables were not normally distributed (all P < 0.05), the strength of correlations was measured by calculating the non-parametric Spearman’s rank correlation coefficient (rs) and its significance has been tested by applying a two-tailed Student’s t-test using PAST 3.12 software. Correlations were considered statistically significant at P < 0.05. For scanning electron microscopy (SEM), selected formalin- or glutaraldehyde-fixed specimens were transferred from 70% ethanol to distilled water for a few hours in order to allow the ethanol in the tissues to be replaced with distilled water. After cutting the specimens transversally into fragments, they were sonicated in distilled water for a variable time (between 1.5 and 9 min) at 43 kHz to remove hoods from the hooks, dehydrated in a graded ethanol series, critical-point dried in carbon dioxide with an EMS 850 critical-point dryer, mounted on stubs, coated with a thin layer (30 nm) of gold with an EMS 550X sputter coater and examined with a Vega Tescan SBH scanning electron microscope at an accelerating voltage of 30 kV. All collected specimens have been deposited at the “Grigore Antipa” National Museum of Natural History (MNINGA), Bucharest, Romania. Information about samples is given below along with descriptions of specimens. The number of specimens in each sample is given in parentheses in the ‘Material examined’ sections after the museum abbreviation and registration number. Complete specimens are indicated by “cs”, while anterior, middle, and posterior fragments are designated as “af”, “mf”, and “pf”, respectively. TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 153 Systematics Spionidae Grube, 1850 Scolelepis Blainville, 1828 Scolelepis Blainville, 1828. Type species: Lumbricus squamatus Abildgaard, in O.F. Müller, 1806, by monotypy. Scolelepis (Scolelepis) squamata (Abildgaard, in O.F. Müller, 1806) Figures 1–4 Lumbricus squamatus Abildgaard, in O.F. Müller, 1806: 39, pl. CLV, figs. 1–5.—Grube 1850: 317. Lumbricus cirratulus Delle Chiaje 1822: pl. 64, figs. 16, 20, 21.—1829: 177–178, 196.—Quatrefages 1865: 449. Scolelepis squamosa.—Blainville 1828: 492–493. Nerine cirratulus.—Claparède 1869: 66–70, pl. XXIV, fig. 1.—Mesnil 1896: 152–165, pl. IX, figs. 1–24 (synonymy).—SaintJoseph 1898: 349–352, pl. XX, fig. 164 (synonymy).—McIntosh 1915: 148–154, pl. XC, fig. 2, pl. XCI, fig. 4, pl. XCVII, figs. 2–2c, pl. XCVIII, fig. 1, pl. XCIV, figs. 4–6, 12, pl. CV, figs. 1–1d (synonymy).—Fauvel 1927: 36, fig.11,g–n.— Bellan & Lagardère 1971: 577–578, fig. 3.—Marinov 1977: 150, pl. XXI, fig. 1,a–e. Scolecolepis squamata.—Söderström 1920: 218. Scolecolepis squammata [sic].—Dörjes 1971: 213, 215, fig. 4. Partim. Scolelepis squamata.—Light 1977: 73–75, figs. 2a–f, 3a–b. Partim.—Kisseleva 2004: 248–249, fig. 97. Nerine mesnili Bellan & Lagardère 1971: 572–576, figs. 1–2. New Synonymy. Scolelepis mesnili.—Capaccioni-Azzati 1991: 57–59, fig. 3. Scolelepis squamatus.—Sikorski 2001: 316. Scolelepis (Scolelepis) squamata.—Delgado-Blas 2006: 76–79, fig. 1A–Q. Material examined. Type material. Syntypes of Nerine mesnili Bellan & Lagardère, 1971: ATLANTIC OCEAN: FRANCE: Charente Maritime, south-western shore of l’Île d’Oléron, Seulières Bay, Chaucre beach, Rémige beach, fine infralittoral sand, 19 Jul 1970, coll. F. Lagardère, original label ‘AF 908’, MNHN POLY TYPE 0582 (3 cs, 1 af, in good condition). Non-type material. ATLANTIC OCEAN: SWEDEN: Kattegat, Gullmarfjord, bay at the station Fiskebäkskil, eulittoral, 16 Aug 2009, leg. P. Wolf, ZSRO-P 2011 (1 cs, 10 af, fixed in denatured alcohol); NORTH SEA: GERMANY: Weser Estuary, mouth of the Weser, km 113, 6.5 m depth, 3 Apr 1991, leg. F. Gosselck, original label ‘ROS-P 108’, ZSRO-P 108 (1 af, 1 pf); FRANCE: Saint-Malo, 1982, leg. M.C. Gambi, LBE Ischia (2 af, 2 mf); SPAIN: Cantabria, Santander, Peña Vieja, Oct 1891, coll. Estación Biológica, MNCN 16.01/370 partim (9 af, poorly preserved with damaged chaetae); Cantabria, Santander, coll. Estación Biológica, MNCN 16.01/368 (2 af, 11 mf, 2 pf); Cantabria, Asturias, Gijón, Ensenada de San Lorenzo, 43°32'28.14"N, 05°38'43.01"W, upper eulittoral, fine and medium sand, 21 Apr 2015, leg. V. Surugiu, no No. (4 af, in good condition, fixed in glutaraldehyde); Cantabria, Asturias, Luanco, Playa La Ribera, 43°36'48.70"N, 05°47'28.09"W, upper eulittoral, fine sand with organic load, 22 Apr 2015, leg. V. Surugiu, MNINGA PLY 0064 (1 af, 1 pf); Cantabria, Asturias, Bañugues, 43°37'51.24"N, 05°48'29.41"W, upper tidal zone, fine sand, 24 Apr 2015, leg. V. Surugiu, no No. (2 cs, 10 af, 1 mf, 1 pf); Galicia, Pontevedra, ría de Pontevedra, Marín, leg. E. Rioja, MNCN 16.01/369 (4 cs, 25 af, 5 pf, in good condition); Galicia, Pontevedra, ría de Vigo, Playa de Barra, leg. J.M. Viéitez, MNCN 16.01/372 (4 af, 5 mf, 1 pf); Galicia, Coruña, Ría de Ferrol, Laxe, intertidal, coarse sand, 22 Jan 1988, leg. J. Parapar, original label: ‘21096220188-1’, MNCN 16.01/12568 partim (2 af, 1 mf); MOROCCO: Bou Regreg estuary, mouth of Bou Regreg, right bank of Plage des Barcassiers, Salé, station 4, 1984, leg. P. Gillet (10 af, in good condition). MEDITERRANEAN SEA: FRANCE: Corsica, Bonifacio, leg. J.M. Pérès, SMF 14294 (1 af, 1 mf); TYRRHENIAN SEA: ITALY: Gulf of Napoli, 1885, don. W. Kükenthal, MNINGA 9482/71 (2 af, 2 pf); off RomeFiumicino, from off the mouth of the Tiber River, 1978, leg. M.C. Gambi, LBE Ischia (1 af, 1 mf, poorly preserved); Campania, N. of Napoli, Gulf of Gaeta, Licola, 1966–1968, leg. J. Dörjes, SMF 12914 partim (40+, in good condition). BLACK SEA: ROMANIA: Original label: ‘Marea Neagră (=Black Sea)’, coll. M. Băcescu, MNINGA 30.289 (18 cs, 73 af, 7 pf, poorly preserved); Constanţa, Modern beach (open water), 44°10'44.32"N, 28°39'37.89"E, 0.5 m, fine sand, 16 Jun 2010, leg. J. Runge, ZSRO-P 2132 (1 cs, 19 af, 19 mf, 1 pf, stiff and fragile material fixed in EtOH); Constanţa, Plaja Modern, 44°10'43.38"N, 28°39'36.21"E, 0.5 m, in fine sand with 154 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU sparse shell debris, 21 Jun 2013, leg. V. Surugiu, MNINGA PLY 0060 (7 cs, 6 af, in good condition); Cap Midia— Digul de Nord, 44°20'32.8"N, 28°41'27.6"E, 0.5–0.8 m, 29 Jun 2014, fine sand with shells, leg. V. Surugiu, MNINGA PLY 0061 (22 cs, 11 af, 2 pf, in good condition); Năvodari—Digul de Sud, 44°19'25.4"N, 28°38'21.4"E, 0.5–0.8 m, 29 Jun 2014, fine sand with shells, leg. V. Surugiu, MNINGA PLY 0062 (8 cs, 1 mf, in good condition); Mangalia, 43°48'52.98"N, 28°35'21.84"E, 0.9–1.3 m, fine sand, 6 Aug 2015, leg. V. Surugiu, MNINGA PLY 0063 (1 cs, 2 af, 1 pf, in good condition, fixed in glutaraldehyde). Description. Body long and thin, tapered posteriorly, kidney-shaped in cross section (Fig. 3G), up to 63 mm long and 3.70 mm wide (at chaetigers 20–30) for 140 chaetigers. Colour in life dusky-greenish anteriorly and darkgrey posteriorly, branchiae with red blood vessels, prostomium and pygidium white-yellowish. Colour in alcohol opaque off-white, without pigmentation. Prostomium fusiform extending anteriorly to a long, tapering point, posteriorly as a narrow, tapered, attached and raised caruncle, extending to posterior margin of chaetiger 1 (Fig. 1A). Occipital antenna absent. Two pairs of dark-brown eyes, arranged trapezoidally just in front of palp bases; anterior eyes slightly larger, usually reniform and set further apart than posterior rounded eyes. Nuchal cilia in U-shaped pattern on sides of caruncle, posterior to palp bases (Fig. 1A). Peristomium long, distinct from chaetiger 1, forming low lateral wings, without furrow (Fig. 1A). Eversible proboscis sac-like, inflated. Palps non-deciduous, long, in largest specimens reaching chaetigers 22–24 (Fig. 4A); ciliation of palps consisting of two compact longitudinal bands of transverse rows of short non-motile cilia with the long rows (on lateral side) 45–55 µm in length, the short rows (on medial side) 35–40 µm in length and a distance of 2–4 µm separating them (Fig. 1B); ratio of long to short rows is approximately 1.15–1.50. Rows of mucussecreting cells, represented by tubular, distinctly reticulated necks with mucus gland opening at the rim, present proximal to transverse ciliary rows (Fig. 1D–E); rows extending to distal ends of palps. Palp sheaths elongate, slightly rugose, fused to bases of palps (Fig. 1C). Branchiae from chaetiger 2 to last 1–2 chaetigers, elongate, gently tapered, best developed by chaetigers 8–10; ciliation along inner edge of branchiae; tips of branchiae glandular, without cilia. In anterior chaetigers, branchiae fused to notopodial postchaetal lamellae for more than two thirds of their length, with pointed tips free (Fig. 1A); on posterior chaetigers branchiae fused only basally to postchaetal lamellae (Fig. 1G). Chaetiger 1 with small lamellae, both shifted more dorsally than in following chaetigers, notopodial lamellae auricular, slightly elongated and rounded, neuropodial lamellae smaller and cordate; capillary chaetae present in both rami, fewer than in following chaetigers; dorsal capillaries twice as long as ventral ones, but shorter and thinner than those of following notopodia; ventral capillaries more numerous, arranged in two rows. Dorsal transverse ciliation as primary ciliary rows (nototrochs sensu Radashevsky (2012)) on mid-part of each chaetiger, continuous with ciliation along inner edge of branchiae (Fig. 1A); short rows of cilia on outer edge of branchiae between tips of notopodial lamellae and subdistal portion of each branchia (Fig. 1G). In anterior chaetigers (to about chaetigers 20–40) secondary transverse ciliary bands as dorsal rows of cilia between adjacent chaetigers (Fig. 1A). Low dorsal transverse folds uniting branchiae across dorsum and bearing nototrochs on middle (from around chaetigers 25–56) and posterior chaetigers (Fig. 1G). Notopodial postchaetal lamellae well developed from chaetiger 2 (Fig. 1A); on anterior chaetigers, notopodial postchaetal lamellae very elongated, narrow, attached almost along entire length to branchiae, outer margin folded, entire or notched in superior 1/3–1/2 from chaetiger 5, free tips bluntly-rounded in adults or sharply pointed in juveniles and small individuals; in middle and posterior chaetigers, notopodial postchaetal lamella becoming gradually shorter, wider and more separated from branchiae, elongate, with rounded margins in large specimens, sub-triangular, with pointed tips in small specimens, with lower portion directed towards neuropodial postchaetal lamella (Figs. 1F, 3G). Neuropodial postchaetal lamellae in anteriormost three or four chaetigers slightly cordate; in following chaetigers semi-oval and entire, then becoming larger, subrectangular, with rounded margins; slight notch developing at chaetigers 14–42; the notch becoming deeper in middle chaetigers, dividing lamella into two separate lobes, the upper lobe being twice as large in size as compared to the lower triangular lobe, which is located ventral to neurochaetae. In posterior chaetigers, gap between lobes gradually enlarges, setting both lobes further apart, upper lobe becoming narrow, rounded, with upper portion elongated and directed towards ventral portion of notopodial postchaetal lamella, lower part reduced to papilliform cirrus (Fig. 1F). Lateral ciliated organs (Figs. 1F, 2A) present between notopodial and neuropodial postchaetal lamellae from chaetiger 1 to posteriormost chaetigers. TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 155 FIGURE 1. Scolelepis (Scolelepis) squamata (Abildgaard, in O.F. Müller, 1806), SEM micrographs. A, voucher specimen (Black Sea, Cap Midia, MNINGA PLY 0061); B–E, voucher specimen (Spain, Gijón); F, G, voucher specimen (Black Sea, Mangalia, MNINGA PLY 0063). A, anterior end, dorsal view, palps missing, showing nuchal organs and dorsal primary and secondary transverse ciliary bands; B, middle section of left palp, frontal-lateral view, showing ciliation pattern; C, basal part of right palp, lateral view, showing tightly fused, slightly rugose sheath; D, close-up view of long transverse rows of cilia on palp, showing arrangement of cilia and mucus secreting cells; E, detail of necks of mucus secreting cells; F, middle chaetigers, left lateral view (arrowheads indicate lateral ciliated organs); G, middle chaetigers, dorsal view, showing the primary ciliary band on low transverse folds. c—cilia; n—neck of gland cells; nu—nuchal organ; pc—dorsal primary transverse ciliary band; sc— dorsal secondary transverse ciliary band. Scale bars: A, F, G = 500 μm; B = 20 μm; C = 200 μm; D = 10 μm; E = 2 μm. 156 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU FIGURE 2. Scolelepis (Scolelepis) squamata (Abildgaard, in O.F. Müller, 1806), SEM micrographs. A, E, F, voucher specimen (Black Sea, Mangalia, MNINGA PLY 0063); B–D, voucher specimen (Black Sea, Cap Midia, MNINGA PLY 0061). A, close-up view of the lateral ciliated organ; B, neuropodial hooded hooks, lateral view (hoods removed); C, tip of neuropodial hooded hook, antero-lateral view, showing notched apical tooth above the main fang (hood removed); D, close-up view of the tip of the hooded hook, dorso-lateral view, showing scale-like structures on the surface of the hood; E, notopodium of a posterior chaetiger, left lateral view, showing hooded hooks alternating with capillaries and a bundle of long dorsal superior capillaries; F, posterior end, dorsal view, showing terminal anus and cushion-like pygidium (somewhat damaged ventrally by sonication). Scale bars: A, B = 20 μm; C = 10 μm; D = 5 μm; E = 100 μm; F = 500 μm. Anterior chaetigers with only bilimbate capillaries in both rami; number of capillaries decreasing towards posterior end. Notopodial capillaries elongate, with narrow limbation, arranged in double rows (up to 18 capillaries per row) plus a dorsal superior tuft; those in anterior row fibrous, those in posterior row uniformly granulated (Fig. 3A), dorsal superior fascicle with longer and thinner capillaries (numbering up to 4). Neuropodial capillaries similar in morphology to those of notopodia, although somewhat shorter, with granulated cores (when viewed in direct light) or appearing fibrous (when viewed in reflected light) due to presence of hollow longitudinal tubes in cores of chaetal shafts (Fig. 3B), also arranged in double irregular rows (up to 15 capillaries per row) plus a ventral inferior fascicle of up to 5 capillaries; capillaries of anterior row shorter and broader, those of posterior row longer and narrower. Neuropodial hooded hooks first present in posterior row from chaetigers 14–43 (Fig. 4B), numbering up to 13 per fascicle, accompanied in middle and posterior chaetigers by 0–3 slender alternating capillaries above the hooks and by 0–1 shorter slender capillaries in ventral inferior tuft. In juveniles and small worms hooded hooks with TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 157 bluntly rounded main fang surmounted by two smaller apical teeth placed side by side (actually a broad notched apical tooth), angle between main tooth and shaft slightly less than 90° (Fig. 3C–D), in medium-sized worms hooded hooks with main fang surmounted by a single broad tooth, angle between main tooth and shaft slightly more than 90° (Fig. 2B–C), in very large worms hooks take the form of falcate spine with very narrow hood (Fig. 3E); shafts long, slightly curved, with tubular core and a minute constriction (Fig. 3C). Hood elongated, more than 5 times the length of the main fang, with fine scale-like structures on its surface (Fig. 2D). Ventral sabre setae absent. Notopodial hooded hooks first present from chaetigers 18–98 (Fig. 4B), numbering up to 5 per row, accompanied by 2–5 short alternating capillaries and 2–4 long capillaries in dorsal superior tuft (Fig. 2E). Notopodial hooded hooks similar in morphology to those of neuropodia, although more elongate (Fig. 3F). Shaft long, almost straight, hood long. One to three prepygidial achaetous segments. Pygidium cushion-like, with entire ventral margin (Fig. 2F). Methyl green staining pattern. Intense blue pigmentation on margins of both notopodial and neuropodial postchaetal lamellae, branchial tips and pygidium. Prostomium, peristomium, and palps with uniformly dispersed speckles. Two speckled transversal stripes ventrally on each segment, tapering from sides to venter, interrupted on either side of mid-ventral nervous chain, anterior stripe wider and more intensely stained. Remarks. Bellan & Lagardère (1971) differentiated Scolelepis mesnili from Scolelepis cirratulus based on the body size, the length of palps, the number of the first chaetiger bearing hooded hooks in the neuropodia and notopodia, the shape of postchaetal lamellae, and some ecological preferences. The present study indicated that the length of palps increases generally with the size of the worm (Fig. 4A). A moderate correlation coefficient is due to the different degree of contraction of the palps. Generally, palps in juveniles and small individuals (less than 1.0 mm in width) commonly reach chaetigers 10–15, while those in adult individuals (more than 2.0 mm width) can reach chaetiger 24. It has also been shown that hook-bearing chaetigers appear later with increasing body size, in both neuro- and notopodia (Fig. 4B), suggesting that as age increases there is a gradual replacement of hooks by capillaries in anterior hook-bearing chaetigers. In the literature there are other reports of such chaetal replacement (Mesnil 1896; Hutchings et al. 1998; Radashevsky & Fauchald 2000; Radashevsky 2012). In the neuropodia, hooded hooks appear in the posterior row, with the first hooks being ventral in position (the last ones not yet replaced by capillaries); however, on following chaetigers the newest hooks emerge in the dorsalmost position. In the notopodia, the newest hooks were in the ventralmost position. However, in posterior hook-bearing chaetigers the arrangement of chaetae in double rows becomes indistinct. In individuals with regenerated posterior segments, notopodial hooded hooks appeared on the same chaetiger with the first occurrence of neuropodial hooded hooks. In separating S. mesnili from S. cirratulus, Bellan & Lagardère (1971) also stressed that their populations are never mixed, those of the former inhabiting the lower intertidal horizons of the beach, whereas the latter are restricted to the upper intertidal. However, Richards (1970) demonstrated that juveniles of ‘Scolelepis squamata’ from Barbados aggregate in the lower levels of the intertidal zone compared to the adults. Contrary to this report of segregation of juveniles from adults, there is also strong evidence of their co-occurrence in the same sample (Gillet 1988; Junoy & Viéitez 1990; present study). The number of apical teeth in both noto- and neuropodial hooks decreases from two in juvenile and small individuals (less than 1.0 mm width) to one in medium-sized individuals (1.1–2.0 mm width), 0–1 in large individuals (2.1–3.0 mm width), and no apical teeth at all in exceptionally large individuals (more than 3.1 mm width) (Fig. 4C). This fact cannot, however, be attributed to the wear of the teeth with the age, as suggested by Foster (1971), because in individuals having hooded hooks of two sorts (i.e., 1 and 2 or 0 and 1 apical teeth), the hooded hooks with a smaller number of teeth occur in more posterior chaetigers and not in the older anterior ones. Thus, it appears that the hooks with smaller number of teeth are the more newly formed ones as they are closest to the prepygidial growth zone. Also the angle of main fang to shaft increases from slightly acute in small individuals (Fig. 3C, F) to slightly obtuse in large individuals (Fig. 2B) and finally reaches a shape of a falcate spine in very large individuals (Fig. 3E). In exceptionally large specimens from the Cantabrian coast of Spain all hooks were in the form of falcate (acicular) spines. The angle between the main fang and the shaft increases as well within the same individual from anterior to posterior chaetigers. The same finding has been reported for Scolelepis viridis Blake & Kudenov, 1978, which has heavy unhooded acicular spines in the notopodia of the middle and posterior chaetigers, whereas the hooks in the neuropodia and anterior notopodia are hooded and bear an apical tooth above the main fang (Blake & Kudenov 1978). 158 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU FIGURE 3. Scolelepis (Scolelepis) squamata (Abildgaard, in O.F. Müller, 1806), LM micrographs. A–D, F, H, voucher specimen (France, Île d’Oléron, MNHN POLY 0582); E, G, voucher specimen (Spain, Pontevedra, MNCN 369). A, capillary notochaetae from chaetiger 11; B, capillary neurochaetae from chaetiger 11; C, neuropodial hooded hooks of a small individual, from chaetiger 38, lateral view; D, neuropodial hooded hooks of a small individual, from chaetiger 45, front-on view, showing notched apical tooth above the main fang with bidentate appearance; E, neuropodial hooded hook of a very large individual, from chaetiger 104, lateral view; F, notopodial hooded hooks of a small individual, from chaetiger 48, lateral view; G, cross section of chaetiger 104, anterior view; H, egg with honeycomb-ornamented shell. Scale bars: A–D, F = 50 μm; E, H = 100 μm; G = 1 mm. TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 159 FIGURE 4. Scolelepis (Scolelepis) squamata, morphometric characteristics. A, relationship between the body-size (referring to maximum anterior width in millimetres) and the palp length (in chaetiger number) (y = 7.6755 + 3.5999x; rs = 0.593; P < 0.0001; n = 54). B, relationship between the total number of chaetigers and the first chaetiger on which neuropodial hooded hooks (triangles, y = 10.698 + 0.22969x; rs = 0.957; P < 0.0001; n = 51) and the notopodial hooded hooks (circles, y = 9.2712 + 0.55917x; rs = 0.948; P < 0.0001; n = 51) are present (black symbols—specimens from the Black Sea; white symbols—the Mediterranean Sea specimens from near type locality; red symbols—syntypes of Scolelepis mesnili; blue symbols—specimens from the Atlantic coast of Spain and France; green symbols—specimens from the North Sea and Kattegat). C, relationship between the body-size (referring to maximum anterior width in millimetres) and the number of teeth above main fang (y = 2.3168 + (−0.57224x); rs = −0.877; P < 0.0001; n = 104). 160 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU Thus, this study confirms Mesnil’s (1896) observation that in the adults of S. (S.) squamata (as Nerine cirratulus) the hooks are bidentate, with bluntly tapered teeth and the notopodial postchaetal lamellae are rounded, whereas in juvenile specimens (Nerine mesnili fide Bellan & Lagardère 1971) hooks are tridentate, with sharper teeth and notopodial postchaetal lamellae are pointed. Many of the descriptions of Scolelepis squamata (Foster 1971; Light 1977; Maciolek 1987; Blake 1996; Delgado-Blas 2006) were based on a small number of specimens. The examination of a large number of specimens previously identified as Scolelepis cirratulus from the Black Sea, the Mediterranean Sea (including specimens from a population close to the type locality of Lumbricus cirratulus), and Morocco, of Scolelepis squamata from the North Sea and the Atlantic coast of Spain and France, as well as of syntypes of Scolelepis mesnili showed the same patterns of size-related variability of some important taxonomic characters (Fig. 4A–C). All correlations were very strong (with exception of correlation between the body width and the palp length which was moderate) and were statistically very highly significant (P < 0.0001). All specimens examined in the present study agree well with the descriptions and illustrations of Scolelepis squamata provided by Mesnil (1896), Sikorski (2001), and DelgadoBlas (2006) and are therefore considered as conspecific. Because Scolelepis lack the median ciliated food groove on palps, all captured particles are transported to the everted pharynx by contraction of the entire palp into a tightly coiling arrangement (Dauer 1983). The rows of nonmotile cilia on palps are believed to represent trichoid sensillae used in mechanoreception and possibly in chemoreception. Proximal to these cilia are present projecting rows of tubular mucus-secreting cells which play role in retention of food particles on the palp surface. Though the North American ‘Scolelepis squamata’ are considered to be identical with the European Scolelepis (S.) squamata, the former seems to be a distinct species because it exhibits on the palps two transverse rows of cilia of different length, with the long rows 65–75 μm in length and the short rows 29–31 μm in length and a distance of 10–12 μm separates these rows (according to measurements from Fig. 2C of Dauer 1983). The ratio of long to short rows in the North American specimens is approximately 1.23–1.3. The European populations also have two rows of subequal lengths, with the long rows 45–55 µm in length and the short rows 35–40 µm in length and a distance of 2–4 µm between them (Fig. 1B). As well, Maciolek (1987) noticed that in the North American specimens in the middle chaetigers the notched neuropodia form an equally bilobed lamella, whilst in the European specimens the lower portion is always much smaller than the upper portion. Light (1977) reports that specimens from San Francisco Bay may or may not have the neuropodial notch in the middle or posterior chaetigers. Furthermore, in all European material examined the notopodial hooded hooks were always present, while Foster (1971) and Light (1977, 1978) reported that in most specimens these are absent. Delgado-Blas (2006) proposed that these authors mixed more than one species, including S. (S.) minuta and S. (S.) acuta. In this light, all records of ‘Scolelepis squamata’ from both coasts of North and South America should be critically re-evaluated. Habitat and ecology. Solelepis (S.) squamata inhabits fine (125–300 µm median grain size), clean sands, sometimes with an admixture of shells, from the upper intertidal to the shallow subtidal zone of exposed beaches (Bellan & Lagardère 1971; present study). The worms live in vertical non-permanent tubes made from loose sand and mucus. The abundance of this species on the Romanian coast of the Black Sea may reach 4,100 ind. m–2. Small Mytilus galloprovincialis Lamarck, 1819 shells were observed in the digestive tube. Reproduction. Scolelepis (S.) squamata is gonochoristic. Elliptical oocytes (170 × 140 µm) were observed in the coelom (along the dorsal blood vessel and in parapodial expansions) from chaetigers 19–36. The eggs have a thick, reticulated membrane and peripheral vesicles (Fig. 3H). Distribution. Atlantic Ocean: from Iceland (Sikorski 2001) and the Norwegian Sea (Hartmann-Schröder 1996) in the north to Morocco (Bellan & Lagardère 1971; Gillet 1988) and Tunisia (Pérez-Domingo et al. 2008) in the south, from the Madeira Archipelago in the west (Bellan & Lagardère 1971) to the Western Mediterranean (Lardicci 1990; Capaccioni-Azzati 1991), the Aegean Sea (Çinar et al. 2014), and the Black Sea (Kisseleva 2004; Surugiu 2005; Kurt Şahin & Çinar 2012) in the east. Scolelepis (Scolelepis) neglecta, new species Figures 5–8 Scolecolepis squammata [sic].—Dörjes 1971: 213, 215, fig. 4. Partim. Not Abildgaard, in O.F. Müller, 1806. Nerinides cantabra.—Viéitez 1976: 240. Not Rioja, 1918. TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 161 Pseudomalacoceros cantabra.—Viéitez 1981: 247, 254, tab. 1. Not Rioja, 1918. Scolelepis squamata.—Parapar 1991: 188–189, 990. Partim. Not Abildgaard, in O.F. Müller, 1806. Material examined. Type material. Holotype: ATLANTIC OCEAN: SPAIN: Cantábria, Asturias, sedimentary area between Cabo Vidio and Cabo de Peñas, st. 6 D 27, 25 m depth, fine sand, 1998, leg. R. Acuña Castroviejo, MNCN 16.01/3852 (1 af). Paratypes: same data as holotype, st. 2 D 59, 25.6 m depth, MNCN 16.01/3851 (1 af); st. D 84, MNCN 16.01/3854 (1 af); st. 7 D 53, 32 m depth, MNCN 16.01/3855 partim (1 af on SEM stub); st. 4 D 72, 8.4 m depth, MNCN 16.01/3856 (1 af); st. 9 D 16, 28 m depth, MNCN 16.01/3857 (1 af); st. 8 D 18, 32 m depth, MNCN 16.01/ 3858 (1 af, 1 mf); st. 7 D 53, 32 m depth, MNCN 16.01/3859 (2 rehydrated af); st. 8 D 19, MNCN 16.01/3860 (2 af); Galicia, Pontevedra, Ría de Vigo, beach of Meira, Oct 1975, leg. J.M. Viéitez, MNCN 16.01/17364 (2 af); Galicia, Coruña, Ría de Ferrol, Laxe, intertidal, coarse sand, 22 Jan 1988, leg. J. Parapar, MNCN 16.01/12568 partim (4 af). Non-type material. ATLANTIC OCEAN: SPAIN: Gulf of Biscaya, Cantabrian coast, Santander, Peña Vieja, Oct 1891, coll. Estación Biológica, MNCN 16.01/370 partim (2 af); Galicia, Pontevedra, Ría de Vigo, Playa de Meira, St. 13, 42°17'04"N, 8°43'08"W, Aug 1972, leg. J.M. Viéitez, MNCN 16.01/373 (1 dehydrated af). MEDITERRANEAN SEA: TYRRHENIAN SEA: ITALY: Campania, N. of Napoli, Gulf of Gaeta, Licola, 1966–1968, leg. J. Dörjes, SMF 12914 partim (~30 af, juveniles only). BLACK SEA: ROMANIA: Năvodari—Digul de Sud, 44°19'25.4"N, 28°38'21.4"E, 0.5–0.8 m, fine sand with shells, 29 Jun 2014, leg. V. Surugiu, MNINGA PLY 0059 (1 af, in good condition). Diagnosis. Prostomium anteriorly with medial portion elongated into an acute point and with rounded anterolateral lobes; prostomium with transverse furrow at level of palp insertions; caruncle subtriangular, slightly inflated, attached to dorsum. Occipital antenna absent. Peristomium short, separated from prostomium by distinct groove, forming well-developed dorso-lateral wings. Palps short, with smooth basal sheaths and with two widely spaced longitudinal bands of elevated lobes bearing transverse rows of cilia. Low dorsal, segmentally arranged, transverse ciliated bands present throughout body. Chaetiger 1 with foliaceous notopodial postchaetal lamellae, with chaetae in both rami. Anterior chaetigers with obtuse, subtriangular prechaetal notopodial lamellae. Branchial tips always free from notopodial postchaetal lamellae. Bidentate neuropodial hooded hooks from chaetiger 19–49; shafts of hooks strongly curved and slightly constricted at bend. Type locality. SPAIN, Cantabria, Asturias, sedimentary area between Cabo Vidio and Cabo de Peñas, 25 m. Description. All specimens incomplete; longest anterior fragment 51 mm for 93 chaetigers and widest 2.75 mm. Holotype 24 mm long, 1.42 mm wide (at chaetiger 15) for 61+ chaetigers, widest anteriorly (around chaetigers 7–21), then gradually tapering to posterior end; body suboval in cross section. Colour in alcohol yellowbrown, without pigmentation. Prostomium anteriorly trilobate, medially acuminate, antero-laterally rounded, posteriorly extended as a short, bluntly rounded, attached, slightly raised caruncle, extending to posterior margin of chaetiger 1 (Figs. 5A, 6A), with transverse constriction in posterior third. Occipital antenna absent. Eyes usually absent, but four faded, subcuticular eyes in a trapezoidal arrangement observed in a few specimens. Nuchal organs a J-shaped inconspicuous structure, on posterior lateral margins of caruncle, just behind bases of palps (Fig. 5A). Peristomium short, distinctly separated from prostomium by a shallow groove, forming well-developed dorsolateral wings, indistinctly separated from chaetiger 1 (Fig. 5A–B). Eversible proboscis sac-like, inflated. Palps deciduous, present only in six examined specimens; when present, palps very short, tapering (Figs. 5B, 6B), reaching at most to chaetiger 5; ciliation of palps consisting of two frontal longitudinal distinctly separated bands of transverse rows of short, non-motile cilia, both on elevated 12-µm-tall lobes of approximately equal width, those on medial side approximately 14–14.3 µm wide, those on lateral side subdivided into two closely applied rows of approximately 7 µm wide each (14 µm in total with a distance less than 1 µm between them) (Fig. 7A–B). Ratio of long to short rows approximately 1.20–1.25 with a distance of 11–15 µm separating them. Palp sheaths rounded, short, smooth, tightly adhering to lateral and abfrontal bases of palps (Fig. 7C). Holotype with only left palp present (fell off during the study), reaching back chaetiger 4. Branchiae from chaetiger 2 to end of fragments, best developed by chaetigers 5–20; ciliation along inner edge of branchiae; tips of branchiae acuminate, glandular, devoid of cilia. In anterior chaetigers, branchiae fused to notopodial postchaetal lamellae for approximately half of their length (Figs. 5A, B, D, E, 7E); on last chaetigers, branchiae fused only basally to postchaetal lamellae (Figs. 5F–H, 7F). 162 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU Chaetiger 1 well developed, capillary chaetae present in both rami, fewer than in following chaetigers; notopodial lamellae elongate, digitiform, tapered subdistally; neuropodial lamellae shorter, acute-ovate (Fig. 5B– C); notopodial capillaries with indistinct double rows and a dorsal superior tuft of thin and long capillaries numbering up to 10, longer than ventral ones and as long as, or slightly shorter than those of dorsal superior tuft on following chaetigers; neuropodial capillaries more numerous, arranged in two distinct rows, 7–9 per row, plus a ventral inferior tuft, all shorter than those of following chaetigers. Dorsal transverse ciliation as primary ciliary bands on mid-part of each chaetiger (nototrochs), continuous with ciliation of branchiae; additional short row of cilia on outer edge of branchiae between tips of notopodial lamellae and subdistal portion of branchiae. In anterior chaetigers (to about chaetiger 26–30) secondary dorsal transverse rows of cilia located on anterior part of chaetigers also present (Fig. 5A). In middle and posterior chaetigers dorsal surface smooth with low transverse folds uniting branchiae (from about chaetiger 25). Ventral surface smooth with a shallow median groove (Fig. 7D). Notopodial postchaetal lamellae well-developed from chaetiger 2 (Fig. 5A–B, D), on anterior chaetigers elongated, narrow, attached to branchiae, with outer margin smooth, entire, with free tips acuminate (Fig. 5D–E), in middle and posterior chaetigers becoming gradually shorter, wider and attached only at base of branchiae, with lower portion directed ventrally towards neuropodial postchaetal lamella (Figs. 5F–H, 7F). Notopodial prechaetal lamellae subtriangular, with stubby (subobtuse) tip, best developed from chaetiger 2 to around chaetiger 29 (Fig. 5B, 7D), then becoming lower and broader inconspicuous folds. Neuropodial postchaetal lamellae in anteriormost 2–5 chaetigers acute, with a small median knob (mammilate), as long as wide, in following chaetigers becoming progressively more rounded, semicircular, entire, wider than long (Fig. 5B, D); slight notch developing in inferior 1/3 of lamella on chaetigers 14–41, in adults (usually wider than 1.5 mm) on chaetigers 35–41 (Figs. 5F, 8); notch becoming deeper in middle chaetigers, dividing lamella into two separate lobes, upper lobe being more than twice as large as lower subtriangular lobe, lower lobe located at level of ventral inferior bundle of capillaries (Fig. 5F–H). In posteriormost chaetigers gap between lobes wider, setting lobes further apart, upper lobe becoming narrow, rounded, with upper portion elongated and directed towards ventral portion of notopodial postchaetal lamella, lower part reduced to triangular cirrus (Fig. 5G–H). Neuropodial prechaetal lamellae absent. Anterior chaetigers with capillaries only, arranged in three distinct groups in both noto- and neuropodia. Notopodial capillary chaetae elongate, narrow, arranged in double rows (7–11 per row) and a dorsal superior tuft of 2–10 longer (up to ~twice the branchiae length) and thinner capillaries; capillaries of anterior row unilimbate, with fibrose cores, of approximately same length and width as those of posterior row; capillaries of posterior row unilimbate, with uniformly granulated cores (Fig. 6C). Number and length of notopodial capillaries gradually decreasing towards posterior end. Neuropodial capillaries similar in morphology to those of notopodia, although shorter and broader, with narrow limbation, with granulated cores (when viewed in direct light) or appearing fibrous (when viewed in reflected light), also arranged in double vertical rows (6–16 per row) plus a ventral inferior bundle of 1–4 long, unilimbate capillaries in position of sabre chaetae (Fig. 6E); capillaries of anterior row (Fig. 6D) slightly shorter and broader than those of posterior row. Neuropodial hooded hooks first present in posterior row of chaetigers 19–49, in adults in chaetigers 40–49 (Fig. 8), up to 13 per fascicle, accompanied in middle and posterior chaetigers by 0–4 alternating slender capillary chaetae and by 0–7 shorter slender limbate capillaries in ventral inferior tuft. Hooded hooks with bluntly rounded main fang surmounted by a smaller bluntly rounded apical tooth, angle between main tooth and shaft 85–95°; shaft long, with small constriction and bend just below hood insertion (Fig. 6F–G). Hood elongate, more than 5 times length of main fang, with apical-rostral slit-like opening. Ventral sabre setae absent. Notopodial hooded hooks not observed. Pygidium unknown. Methyl green staining pattern. Most intense blue staining on peristomial wings, on margins of both notopodial and neuropodial postchaetal lamellae, on tips of notopodial prechaetal lamellae, on tips of branchiae and on superior margins of basal sheaths of palps (Fig. 6A–B). Prostomium and peristomium with uniformly dispersed speckles forming indistinct longitudinal stripes. Ventral side uniformly speckled on each chaetiger, with unclear transversal stripes. TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 163 FIGURE 5. Scolelepis (Scolelepis) neglecta sp. nov. A, B, holotype (MNCN 3852); C–H, paratype (MNCN 12568). A, anterior end, dorsal view (palps missing, left notopodial lamella and branchia of chaetiger 5 missing); B, anterior end, lateral view (left palp present only); C, parapodium of chaetiger 1, anterior view; D, parapodium of chaetiger 2, anterior view; E, parapodium of chaetiger 15, anterior view; F, parapodium of chaetiger 42, posterior view; G, parapodium of chaetiger 50, anterior view; H, parapodium of chaetiger 56, anterior view. Scale bars: A–H = 1 mm. 164 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU FIGURE 6. Scolelepis (Scolelepis) neglecta sp. nov., A, B, LM micrographs of formaldehyde-fixed specimens stained with a solution of methyl green in alcohol, H, I, LM micrographs. A, C–G, holotype (MNCN 3852); B, paratype (MNCN 3860); H, I, paratype (MNCN 12568). A, anterior end, dorsal view; B, anterior end, lateral view; C, capillary notochaeta from posterior row of chaetiger 9; D, capillary neurochaeta from anterior row of chaetiger 13; E, ventral inferior capillary from chaetiger 33. F, neuropodial hooded hook from chaetiger 50, lateral view; G, neuropodial hooded hook from chaetiger 59, front-on view; H, egg envelope ornamentation; I, coelomic egg with thick papillate membrane and peripheral vesicles. Scale bars: A–B = 1 mm; C–I = 0.1 mm. TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 165 FIGURE 7. Scolelepis (Scolelepis) neglecta sp. nov., SEM micrographs. A–F, paratype (MNCN 3855). A, distal tip of left palp with elevated lobes, frontal-lateral view (arrowheads indicate faint splitting of lateral rows into two smaller lobes); B, distal section of right palp, frontal view, showing elevated lobes; C, basal part of right palp, lateral view (arrowhead shows smooth basal palpal sheath); D, chaetigers 13–25 (from top right to bottom left), ventro-lateral view (arrowheads indicate prechaetal notopodial lamellae; arrow indicates ventral median groove); E, chaetigers 16–21 (from top right to bottom left), dorso-lateral view (arrowheads indicate lateral ciliated organs); F, chaetigers 29–32 (from right to left), lateral view (note ventrally directed inferior part of notopodial postchaetal lamella and dorsally directed superior part of neuropodial postchaetal lamella). Scale bars: A, B = 50 μm; C = 100 μm; D = 1 mm; E = 0.5 mm; F = 0.2 mm. Remarks. Scolelepis (S.) neglecta sp. nov. most closely resembles S. (S.) kudenovi Hartmann-Schröder, 1981, described from Western Australia, and S. (S.) angulata Zhou, 2014, recently described from the South China Sea and the Yellow Sea. The three species are morphologically similar in the following characteristics: the shape of the prostomium (somewhat truncate, with a conical medial tip and subdistal lateral angles anteriorly and with a transversal constriction in the posterior third), the caruncle extending to the posterior margin of chaetiger 1, short palps (never exceeding chaetiger 5), the presence of anterior prechaetal notopodial lamellae, and bidentate hooded hooks. With regard to palp ciliation pattern, the new species is somewhat similar to S. (S.) hutchingsae Dauer, 1985 in having distinctly separated long and short rows, with the short rows on elevated lobes (Williams 2007). However, S. (S.) neglecta sp. nov. presents a new pattern in having both rows on elevated lobes. Thus, for those eleven species of Scolelepis that have been so far examined with SEM for the palp ciliary pattern, six distinct morphological types have been recognized (Williams 2007; Zhou 2014; present study). It is likely that further investigation of palp morphology for other species will greatly increase the number of morphological types to which they belong. The new species is similar to the Australian specimens of S. (S.) kudenovi. However, S. (S.) neglecta sp. nov. differs from S. (S.) kudenovi by the shape of subdistal anterolateral lobes on prostomium, which are broadly 166 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU rounded in the former and bluntly tapered in the latter, the presence of well-developed peristomial wings in the former, and the shape of the neuropodial hooded hooks, which have smaller, bluntly rounded teeth instead of sharp teeth with an upright apical tooth. Also, specimens of S. (S.) kudenovi appear to be much more slender than those of S. (S.) neglecta sp. nov. (Table 1). Specimens of S. (S.) kudenovi reported by Imajima (1992) from Japan seem to belong to a species distinct from the Australian material (Meißner & Götting 2015). Scolelepis (S.) neglecta sp. nov. differs from S. (S.) angulata in the shape of the prostomium, the lack of pigmentation on the posterior caruncle, the shape of the peristomium, which has a smooth contour in S. (S.) neglecta sp. nov. and a wavy contour in S. (S.) angulata, the shape of the neuropodial postchaetal lamellae of anteriormost chaetigers, which are ovate in S. (S.) neglecta sp. nov. and rectangular in S. (S.) angulata, the extension of branchiae, which are present on at least 79 chaetigers (the longest anterior fragment), while in S. (S.) angulata the branchiae are limited to first 43–51 chaetigers, and by ciliation on palps (absent in S. (S.) angulata). There are also differences in the habitat preferences: S. (S.) angulata mainly inhabits tidal flats with saltmarsh cordgrass (Spartina alterniflora), whilst S. (S.) neglecta sp. nov. mainly inhabits fine sublittoral sands. Scolelepis (S.) neglecta sp. nov. has been confused in the past with S. (S.) squamata, S. (S.) mesnili, S. (Scolelepis) cantabra, and Dispio uncinata Harman, 1951. Thus, individuals from Cantabria were identified by G. San Martín either as Dispio uncinata and subsequently revised by O. Díaz as Scolelepis mesnili (unpublished study by Centro de Investigaciones Submarinas, 1998, Guillermo San Martín, pers. comm., Dec 2015), or as Scolelepis squamata by G. San Martín and J.M. Viéitez. The material from Galicia was identified as Scolelepis cantabra by J.M. Viéitez (1976, 1981) or as Scolelepis squamata by Parapar (1991). Also, the material from Licola, the Mediterranean Sea, was identified as Scolelepis squamata by Dörjes (1971). The new species differs from S. (S.) squamata by the shape of the prostomium (with anterolateral lobes and with a transverse furrow in front of the caruncle in S. (S.) neglecta sp. nov. and without anterolateral lobes and without a transverse furrow in front of the caruncle in S. (S.) squamata), by having a shorter peristomium, by having well-developed lateral peristomial wings which are separated from the prostomium by a distinct grove, by the shape of basal sheaths (smooth in S. (S.) neglecta sp. nov. and slightly rugose in S. (S.) squamata), by the ciliation pattern of palps, by the chaetiger 1 having much longer notopodial postchaetal lamellae and capillaries, by the shape of anterior postchaetal lamellae (with entire, smooth margins in S. (S.) neglecta sp. nov. and with folded, notched margins in S. (S.) squamata), and by the presence of a distinct prechaetal lamellae. Also, in S. (S.) neglecta sp. nov., the branchiae on anterior chaetigers are less fused to the notopodial postchaetal lamellae (about 1/2 as against to 2/3 in S. (S.) squamata). Palps are usually deciduous in S. (S.) neglecta sp. nov. and non-deciduous in S. (S.) squamata. The eggs in S. (S.) squamata are characterized by 12 peripheral vesicles (Joyner 1962), whereas in Scolelepis (S.) neglecta sp. nov. the eggs have 27–31 peripheral vesicles. The new species can be easily distinguished from S. (S.) cantabra by the absence of occipital antenna (which is present in S. cantabra fide Maciolek (1987)), by the presence of notochaetae in chaetiger 1, and by the presence of a notch in the neuropodial postchaetal lamellae. Scolelepis (S.) neglecta sp. nov. has been confused also with Dispio uncinata because of the large notopodial postchaetal lamellae on chaetiger 1, which resembles a lamella joined with branchia. At first glance it was difficult to discern whether on chaetiger 1 there is a large lamella or a small branchia completely fused to the notopodial lamella. However, the vascular core of the branchia, which can be detected on chaetiger 2, was not observed on chaetiger 1. Also, anterior notochaetae are very long, resembling those of Dispio uncinata. There are, however, major differences in most other characters between these two species. Etymology. The species name “neglecta” (derived from the Latin nominative singular feminine adjective neglecta meaning disregarded, neglected, or ignored) refers to the fact that the species has in the past been misidentified and overlooked, without being recognized as a distinct species. Habitat and ecology. The species inhabits fine sublittoral sands with an admixture of shells at depths ranging from 0.5 to 32 m. Short palps could indicate that this species is dependent on habitats rich in organic matter (Dauer 2000). It usually occurs together with Aponuphis bilineata (Baird, 1870), Goniada maculata Örsted, 1843, Hilbigneris gracilis (Ehlers, 1868), Magelona papillicornis F. Müller, 1858, Nephtys cirrosa Ehlers, 1868, Nephtys hombergii Savigny in Lamarck, 1818, Nothria conchylega (Sars, 1835), Onuphis eremita Audouin & Milne Edwards, 1833, Owenia fusiformis Delle Chiaje, 1844, Sigalion mathildae Audouin & Milne Edwards in Cuvier, 1830, and Sthenelais limicola (Ehlers, 1864) (Dörjes 1971; Viéitez 1976, 1981; Centro de Investigaciones Submarinas, unpublished study). TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 167 Reproduction. Oocytes were observed in the coelomic cavity from chaetiger 40 to the end of fragments. The eggs are ellipsoid, flattened disks, 200–230 µm × 150–170 µm in diameter. They have a thick papillated vitelline membrane, 27–31 peripheral vesicles, opaque yolky cytoplasm, and a centrally positioned hyaline nucleus (Fig. 6H–I). Occurrence. Scolelepis (S.) neglecta sp. nov. is known so far only from the Cantabrian and Galician coasts of Spain (Viéitez 1976, 1981, as Scolelepis cantabra; Parapar 1991, as Scolelepis squamata), from the Tyrrhenian Sea (Dörjes 1971, as Scolelepis squamata), and the Black Sea (present study). FIGURE 8. Scolelepis (Scolelepis) neglecta sp. nov., relationship between the body-size (referring to the maximum anterior width) and the first chaetiger on which a notch is present in the neuropodial postchaetal lamellae (triangles, holotype empty symbol, y = 13.28 + 10.106x; rs = 0.931; P < 0.0001; n = 36), and between the body-size (referring to the maximum anterior width) and the first chaetiger on which neuropodial hooded hooks are present (circles, holotype empty symbol, y = 15.673 + 11.97x; rs = 0.846; P < 0.0001; n = 36). Conclusion Scolelepis is generally a speciose and taxonomically difficult genus. Some of the taxonomic characters traditionally used in distinguishing different species of Scolelepis (Scolelepis), such as the shape and the distribution of hooded hooks, the shape of notopodial postchaetal lamellae and the first appearance of the notch in the postchaetal neuropodial lamellae, proved to be highly size-dependent. Such meristic characters can be useful only on statistical bases. An unequivocal identification of the representatives of these spionid polychaetes requires the use of a set of morphological features. Taxonomic characters useful in differentiating species of Scolelepis (Scolelepis) are the shape of the prostomium, the presence and the size of lateral wings on peristomium, and the shape of postchaetal lamellae on chaetiger 1. Due to the extreme fragility of these worms, morphology of the pygidium remains unknown for some rarely encountered species. Examination under SEM of the palp morphology seems to be very promising for the identification of Scolelepis (Scolelepis) species. However, integrative approaches including molecular analyses (as have been completed in other groups of spionids; e.g., Meißner et al. 2011; Sato-Okoshi & Abe 2013; Radashevsky et al. 2016) are necessary to clarify the taxonomy for most of the species within this genus. 168 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA TABLE 1. Taxonomic characters of selected species in the genus Scolelepis (Scolelepis). Zootaxa 4161 (2) © 2016 Magnolia Press · Species * Maximum length (mm) Maximum width (mm) Maximum number of chaetigers Prostomium anteriorly Posterior margin of prostomium (caruncle) neglecta sp. nov. 1 51+ 2.75 93+ Trilobate, median lobe sharply pointed, anterolateral lobes rounded Subtriangular, slightly inflated, attached to dorsum; transverse furrow in front; extending to chaetiger 2 angulata Zhou, 2014 2 12.9 0.9 74 Trilobate, median lobe pointed, short anterolateral angles Triangular, dark coloured; horizontal gap near anterior margin of chaetiger 1; extending to anterior margin of chaetiger 2 kudenovi Hartmann-Schröder, 1981 Australia 3, 4 42+ 1.0 123+ Trilobate, median lobe sharply pointed, anterolateral lobes tapered Subulate, slightly inflated, attached to dorsum; transverse furrow in front; extending to end of chaetiger 1 kudenovi Hartmann-Schröder, 1981 Japan 5 45 1.5 81 Conical, extended to a long, tapering point Subulate, slightly inflated, attached to dorsum; transverse furrow in front; extending to end of chaetiger 1 squamata (Abildgaard, in O.F. Müller, 1806) 1, 6, 7, 9, 11 70 3.7 200 Conical, extended to a long, tapering point Narrow, tapered, raised and attached to dorsum; extending to chaetiger 2 cantabra (Rioja, 1918) 8, 9 100 5 200+ Truncate or conical, more or less pointed Tapering, detached from dorsum as an occipital antenna; extending to middle of chaetiger 3 bonnieri (Mesnil, 1896) 6, 9, 10 60 4 90 Conical, pointed or bluntlytipped Pointed, detached from dorsum; extending to chaetiger 2 foliosa (Audouin & Milne Edwards, 1833) 6, 10, 11 160 9 250 Truncate, with small medial conical protuberance Extending to end of chaetiger 1 ……continued on the next page 169 170 · Zootaxa 4161 (2) © 2016 Magnolia Press TABLE 1. (continued) Species * Occipital antenna Peristomium Palps reaching chaetiger (basal palpal sheaths) Shape of notopodial postchaetal lamellae of chaetiger 1 neglecta sp. nov. 1 Absent Short, separated from prostomium by a distinct groove, forming well-developed dorso-lateral wings 5 (thickened, smooth) Digitiform angulata Zhou, 2014 2 Absent Long, distinct from chaetiger 1, with welldeveloped lateral wings 3 (thickened) Triangular kudenovi Hartmann-Schröder, 1981 Australia 3, 4 Absent Short, without lateral wings 5 (thickened) Subulate, long and slender kudenovi Hartmann-Schröder, 1981 Japan 5 Absent Short, separated from chaetiger 1, forming lateral wings 20 (lacking basal sheaths) Small lanceolate squamata (Abildgaard, in O.F. Müller, 1806) 1, 6, 7, 9, 11 Absent Long, distinct from chaetiger 1, with low lateral wings 6–25 (rugose) Small, auricular cantabra (Rioja, 1918) 8, 9 Present Short, indistinct from chaetiger 1, inflated, forming conspicuous lateral wings Long, thin and very deciduous Small, rounded bonnieri (Mesnil, 1896) 6, 9, 10 Absent ? Long and stout Rounded foliosa (Audouin & Milne Edwards, 1833) 6, 10, 11 Present (small) Large, surrounding prostomium, without lateral wings Stout (lacking basal sheaths) Oval ……continued on the next page SURUGIU TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA TABLE 1. (continued). Zootaxa 4161 (2) © 2016 Magnolia Press · Species* Shape of neuropodial postchaetal lamellae of chaetiger 1 Notochaetae on chaetiger 1 Branchial fusion on anterior chaetigers Branchial fusion on posterior chaetigers Prechaetal notopodial lamellae neglecta sp. nov. 1 Acute Long Fused for approx. 50% of length Fused for less than 30% Subtriangular, stubby, from chaetiger 2 to about chaetiger 29 angulata Zhou, 2014 2 Rectangular Long At most fused for 80% of length Fused for less than 20% Thick and round, from chaetiger 2 to posterior chaetigers kudenovi Hartmann-Schröder, 1981 Australia 3, 4 Subulate Long Fused for approx. 50% of length Fused for less than 25% Triangular, indistinct, present until about chaetiger 40 kudenovi Hartmann-Schröder, 1981 Japan 5 Lanceolate Long Fused for approx. 50% of length Fused for 30% Rounded squamata (Abildgaard, in O.F. Müller, 1806) 1, 6, 7, 9, 11 Small, cordate Short Fused for approx. 66% of length Fused for less than 25% Absent cantabra (Rioja, 1918) 8, 9 Semi-circular Absent Completely fused to notopodial lamellae in anterior 25–30 chaetigers Fused for less than 20% ? bonnieri (Mesnil, 1896) 6, 9, 10 Rounded Present Fused for approx. 75% of length Fused for approx. 75% of length ? foliosa (Audouin & Milne Edwards, 1833) 6, 10, 11 Rounded Short Completely fused to notopodial lamellae in anterior 56–67 chaetigers Almost completely separated Well-developed to chaetiger 78–84 ……continued on the next page 171 172 · Zootaxa 4161 (2) © 2016 Magnolia Press TABLE 1. (continued) Species* Shape of anterior notopodial postchaetal lamellae Shape of posterior notopodial postchaetal lamellae Neuropodial notch starting on chaetiger Neuropodial hooded hooks from chaetiger neglecta sp. nov. 1 Very elongated, narrow, attached to branchiae, with the outer margin smooth, entire, with free tips acuminate Triangular, more separated from branchiae, with lower portion directed downward 14–41 19–49 angulata Zhou, 2014 2 Long, narrow Round 29–36 32–48 kudenovi Hartmann-Schröder, 1981 Australia 3, 4 Tapered with lower portion rounded, slightly folded Triangular, more separated from branchiae 33–48 38–44 kudenovi Hartmann-Schröder, 1981 Japan 5 Elongated, narrow, attached to branchiae, with the outer margin smooth, entire, with free tips acuminate Flag-like, with upper and lower corners pointed 24–27 26–30 squamata (Abildgaard, in O.F. Müller, 1806) 1, 6, 7, Very elongated, narrow, attached to branchiae, with the outer margin ruffled, entire or notched, with free tips rounded (adults) or pointed (juveniles) Elongated with rounded margins (adults) or triangular (juveniles), more separated from branchiae, with lower portion directed downward 14–42 14–43 cantabra (Rioja, 1918) 8, 9 Wide, bordering branchiae, free margin folded Divided into an apical rounded, flag-like process and an inferior ovoid mamelon Absent 20–45 bonnieri (Mesnil, 1896) 6, 9, 10 Very elongated, narrow, attached to branchiae, with the outer margin smooth, entire, with free tips sharply pointed Triangular, with superior tips sharply pointed 30 21–36 foliosa (Audouin & Milne Edwards, 1833) 6, 10, 11 Folded or ruffled Rounded 70–93 43–68 9, 11 ……continued on the next page SURUGIU TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA TABLE 1. (continued) Zootaxa 4161 (2) © 2016 Magnolia Press · Species* Notopodial hooded hooks from chaetiger Number of teeth above main fang of hooded hooks Number of neuropodial hooded hooks per chaetiger Shape of pygidium Other features neglecta sp. nov. 1 ? 1 1–13 Unknown Eyes usually absent or faint angulata Zhou, 2014 2 Absent 1 5–14 Cushion-like Branchia continuing to chaetiger 43–51 kudenovi Hartmann-Schröder, 1981 Australia 3, 4 98 1 1–14 Unknown Very upright apical tooth kudenovi Hartmann-Schröder, 1981 Japan 5 56–62 1 17 Ventral, entire cushion squamata (Abildgaard, in O.F. Müller, 1806) 1, 6, 7, 9, 11 18–98 0–2 1–13 Cushion-like Eyes arranged in a trapezium, anterior pair larger cantabra (Rioja, 1918) 8, 9 Absent 1 5–16 Multilobate Eyes arranged in a trapezium, anterior pair smaller bonnieri (Mesnil, 1896) 6, 9, 10 30–74 0–1 20–28 Cushion-like Eyes arranged in a transverse row foliosa (Audouin & Milne Edwards, 1833) 6, 10, 11 50–74 0–1 10–25 Cushion-like, with oval dorsal plate Eyes absent. Branchiae absent in posterior region * References: 1. Present study; 2. Zhou (2014); 3. Hartmann-Schröder (1981); 4. Meißner & Götting (2015); 5. Imajima (1992); 6. Mesnil (1896); 7. Delgado-Blas (2006); 8. Rioja (1918); 9. Fauvel (1927); 10. Hartmann-Schröder (1996); 11. Sikorski (2001). 173 Acknowledgements I am much indebted to Víctor Hugo Delgado-Blas (Universidad de Quintana Roo, Chetumal, México), to Danny Eibye-Jacobsen (Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark), to Nancy J. Maciolek (ENSR Marine & Coastal Center, Woods Hole, USA), to Karin Meißner (Forschungsinstitut Senckenberg, Hamburg, Germany), and to Vasily I. Radashevsky (A.V. Zhirmunsky Institute of Marine Biology, Vladivostok, Russia) for critical appraisal of an earlier draft of this manuscript and whose invaluable suggestions greatly improved the quality of the paper. My sincere gratitude also to Jason D. Williams (Hofstra University, USA) and two anonymous reviewers for critical comments and editing the manuscript. I would like to thank Melanya Stan (MNINGA), Andreas Bick (ZSRO), Dieter Fiege (SMF), Tarik Meziane (MNHN), Javier Sánchez Almazán (MNCN), and José Manuel Viéitez Martín (Universidad Alcalá de Henares, Madrid, Spain) for arranging the loans of specimens. Patrick Gillet (Institut de Recherche Fondamentale et Appliquée, Université Catholique de l’Ouest, Angers, France) and Maria Cristina Gambi (Stazione Zoologica “Anton Dohrn”, Napoli, Italy) are kindly acknowledged for providing specimens for the present study. Many thanks to Nuria Anadón, Alexandra Richter and Andrés Arias (Universidad de Oviedo, Spain) and to Alex Strugariu (Universitatea “Alexandru Ioan Cuza” din Iaşi, Romania) for assistance in sample collection, to Chris J. Glasby (Museum & Art Gallery Northern Territory, Darwin, Australia) for correcting the English during his stay in Bucharest, and to Guillermo San Martín Peral (Universidad Autónoma de Madrid, Spain) for valuable information on the provenance of some Spanish material. Many thanks to Irina Gostin, Magda Dascălu, and Lucian Fusu for their assistance with SEM and LM studies, respectively. References Bellan, G. & Lagardère, F. (1971) Nerine mesnili, n. sp. Spionidien méconnu des plages sableuses de la Province Lusitaniene. Bulletin de la société Zoologique de France, 96 (4), 571–579. Blainville, H. de (1828) Dictionnaire des Sciences naturelles, dans lequel on traite méthodiquement des différents êtres de la nature, considérés soit en eux-mêmes, d'après l'état actuel de nos connaissances, soit relativement à l'utilité qu'en peuvent retirer la médicine, l'agriculture, le commerce et les arts. Suivi d'une biographie des plus célèbres naturalistes. Volume 57, F.G. Levrault, Strasbourg & Paris, 628 pp. Blake, J.A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J.A., Hilbig, B. & Scott, P.H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 81–223. Blake, J.A. & Kudenov, J.D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171–280. Capaccioni-Azzati, R. (1991) Lumbrineris cingulata (Polychaeta, Lumbrineridae) et Scolelepis mesnili (Polychaeta, Spionidae) en Méditerranée. Vie et Milieu, 41 (1), 55–60. Çinar, M.E., Dağli, E. & Kurt Şahin, G. (2014) Checklist of Annelida from the coasts of Turkey. Turkish Journal of Zoology, 38 (6), 734–764. http://dx.doi.org/10.3906/zoo-1405-72 Claparède, E. (1869) Les Annélides Chétopodes du Golfe de Naples. Seconde partie. Mémoires de la Société de Physique et d’Histoire Naturelle de Genève, 20 (1), 1–225, pls. 17–31. Dauer, D.M. (1983) Functional morphology and feeding behavior of Scolelepis squamata (Polychaeta: Spionidae). Marine Biology, 77, 279–285. http://dx.doi.org/10.1007/BF00395817 Dauer, D.M. (1985) A new species of Scolelepis (Polychaeta: Spionidae) from Lizard Island, Australia. Proceedings of the Biological Society of Washington, 98, 678–681. Dauer, D.M. (1987) Systematic significance of the morphology of spionid polychaete palps. Bulletin of the Biological Society of Washington, 7, 41–45. Dauer, D.M. (2000) Functional morphology and feeding behavior of Spio setosa (Polychaeta: Spionidae). Bulletin of Marine Science, 67 (1), 269–275. Delgado-Blas, V.H. (2006) Partial revision of Scolelepis (Polychaeta: Spionidae) from the Grand Caribbean region, with the description of two new species and a key to species recorded in the area. Contributions to Zoology, 75 (1–2), 75–97. Delle Chiaje, S. (1822) Memorie sulla storia e notomia degli animali senza vertebre del Regno di Napoli. Figure. Vol. 1, Napoli, 227 pp., 109 pls. http://dx.doi.org/10.5962/bhl.title.10021 174 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU Delle Chiaje, S. (1829) Memorie sulla storia e notomia degli animali senza vertebre del Regno di Napoli. Vol. 4, Stamperia della Societa Tipografica, Napoli, 214 pp. Dörjes, J. (1971) Der Golf von Gaeta (Tyrrhenisches Meer). IV. Das Makrobenthos und seine küstenparallele Zonierung. Senckenbergiana maritima, 3, 203–246. Eibye-Jacobsen, D. (1997) A new species of Scolelepis (Polychaeta: Spionidae), highly abundant on the sand beaches of western Phuket Island, Thailand. Bulletin of Marine Science, 60 (2), 240–251. Fauvel, P. (1927) Polychètes sédentaires. Addenda aux Errantes, Archiannélides, Myzostomaires. Faune de France 16, Paul Lechevalier, Paris, 494 pp. Foster, N.M. (1971) Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea. Studies on the Fauna of Curaçao and other Caribbean Islands, 36, 1–183. Gillet, P. (1988) Structure des peuplements intertidaux d'Annélides Polychètes de l’estuaire du Bou Regreg (Maroc). Bulletin d'Ecologie, 19 (1), 33–42. Grube, A.E. (1850) Die Familien der Anneliden. Archiv für Naturgeschichte, 16 (1), 249–364. Hartmann-Schröder, G. (1981) Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden. Teil 6: Die Polychaeten der tropisch-subtropischen Westküste Australiens (zwischen Exmouth im Norden und Cervantes im Süden). Mitteilungen aus dem Hamburger Zoologischen Museum und Institut, 78, 19–96. Hartmann-Schröder, G. (1996) Die Tierwelt Deutschlands. Vol. 58. Annelida, Borstenwürmer, Polychaeta. 2nd ed. Neubearbeitete Auflage, Gustav Fischer Verlag, Jena, 648 pp. Hutchings, P.A., Frouin, P. & Hily, C. (1998) Two new species of Spionidae (Polychaeta) from Tahiti, French Polynesia. Proceedings of the Biological Society of Washington, 111 (4), 799–806. Imajima, M. (1992) Spionidae (Annelida, Polychaeta) from Japan, 8. The genus Scolelepis. Bulletin of the National Science Museum Tokyo (Japan). Series A. Zoology, 18, 1–34. International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. Fourth edition. International Trust for Zoological Nomenclature, London, i–xxix + 1–306. Joyner, A. (1962) Reproduction and larval life of Nerine cirratulus (Delle Chiaje) Family Spionidae. Proceedings of the Zoological Society of London, 138 (4), 655–666. http://dx.doi.org/10.1111/j.1469-7998.1962.tb05338.x Junoy, J. & Viéitez, J.M. (1990) Macrozoobenthic community structure in the Ría de Foz, an intertidal estuary (Galicia, Northwestern Spain). Marine Biology, 107, 329–339. http://dx.doi.org/10.1007/BF01319833 Kisseleva, M.I. (2004) Polychaetes (Polychaeta) of the Azov and Black Seas. Kola Science Centre of the Russian Academy of Science Press, Apatity, 409 pp. [in Russian] Kurt Şahin, G. & Çinar, M.E. (2012) A check-list of polychaete species (Annelida: Polychaeta) from the Black Sea. Journal of the Black Sea/Mediterranean Environment, 18 (1), 10–48. Avaliable from: http://dergipark.ulakbim.gov.tr/jbme/article/ view/5000144348 (Accessed 2 Sept. 2016) Lardicci, C. (1990) Censimento dei Policheti dei Mari Italiani: Spionidae Grube, 1850. Atti della Societa Toscana di Scienze Naturali, Memorie, Serie B, 96, 121–152. Light, W.J. (1977) Spionidae (Annelida: Polychaeta) from San Francisco Bay, California: A revised list of nomenclatural changes, new records, and comments on related species from the northeastern Pacific Ocean. Proceedings of the Biological Society of Washington, 90 (1), 66–88. Light, W.J. (1978) Invertebrates of the San Francisco Bay Estuary System. Family Spionidae (Polychaeta: Annelida). The Boxwood Press, Pacific Grove, California, 211 pp. MacCord, F.S. & Amaral, A.C.Z. (2005) Morphometric analysis of two species of Scolelepis (Polychaeta: Spionidae). Journal of Marine Biological Association of the United Kingdom, 85, 829–834. http://dx.doi.org/10.1017/S002531540501177X Maciolek. N.J. (1987) New species and records of Scolelepis (Polychaeta: Spionidae) from the east coast of North America. Proceedings of the Biological Society of Washington, 7, 16–40. Marinov, T. (1977) Fauna na Bŭlgaria. Mnogochetinesti chervey (Polychaeta). Vol. 6, Bŭlgarskata Akademia na Naukite, Sofia, 258 pp. [in Bulgarian] McIntosh, W.C. (1915) A monograph of the British Annelids. Volume III. Part I. Text. Polychaeta, Opheliidae to Ammocharidae. Ray Society, London, 368 pp., Pls. 88–106. Meißner, K. & Götting, M. (2015) Spionidae (Annelida: ‘Polychaeta’: Canalipalpata) from Lizard Island, Great Barrier Reef, Australia: the genera Malacoceros, Scolelepis, Spio, Microspio, and Spiophanes. Zootaxa, 4019 (1), 378–413. http://dx.doi.org/10.11646/zootaxa.4019.1.15 Meißner, K., Bick, A. & Bastrop, R. (2011) On the identity of Spio filicornis (O.F. Müller, 1776)—with the designation of a neotype, and the description of two new species from the North East Atlantic Ocean based on morphological and genetic studies. Zootaxa, 2815, 1–27. Mesnil, F. (1896) Études de morphologie externe chez les Annélides. I. Les Spionidiens des côtes de la Manche. Bulletin Scientifique de la France et de la Belgique, 29, 110–292, pls. 7–15. Müller, O.F., Abildgaard, P.C., Vahl, M., Holten, J.S. & Rathke, J. (1806) Zoologiae Danica seu Animalium Daniae et TAXONOMIC STATUS OF SCOLELEPIS SQUAMATA Zootaxa 4161 (2) © 2016 Magnolia Press · 175 Norvegiae rariorum ac minus notorum. Descriptiones et historia. Volume IV. Volume Atlas. N. Christensen: Havniae [Copenhagen], 1–2, 46 pp., pls. 121–160. Oug, E., Bakken, T. & Kongsrud, J.A. (2014) Original specimens and type localities of early described polychaete species (Annelida) from Norway, with particular attention to species described by O.F. Müller and M. Sars. Memoirs of Museum Victoria, 71, 217–236. Parapar, J. (1991) Anélidos Poliquetos bentónicos de la ría de Ferrol (Galicia). PhD-thesis, University of Santiago de Compostela, 1104 pp. Pettibone, M.H. (1963) Revision of some genera of polychaete worms of the family Spionidae, including the description of a new species of Scolelepis. Proceedings of the Biological Society of Washington, 76, 89–104. Pérez-Domingo, S., Castellanos, C. & Junoy, J. (2008) The sandy beach macrofauna of Gulf of Gabès (Tunisia). Marine Ecology, 29 (Suppl. 1), 51–59. http://dx.doi.org/10.1111/j.1439-0485.2007.00201.x Quatrefages, A. de (1865) Histoire naturelle de annelés marins et d’eau douce. Annélides et Géphyriens. Vol. I, Librairie Encyclopédique de Roret, Paris, 588 pp. Radashevsky, V.I. (2012) Spionidae (Annelida) from shallow waters around the British Islands: an identification guide for the NMBAQC Scheme with an overview of spionid morphology and biology. Zootaxa, 3152, 1–35. Radashevsky, V.I. & Fauchald, K. (2000) Chaetal arrangement and homology in spionids (Polychaeta: Spionidae). Bulletin of Marine Science, 67, 13–23. Radashevsky, V.I., Malyar, V.V., Pankova, V.V. & Nuzhdin, S.V. (2016) Molecular analysis of six Rhynchospio Hartman, 1936 species (Annelida: Spionidae) with comments on the evolution of brooding within the group. Zootaxa, 4127 (3), 579–590. http://dx.doi.org/10.11646/zootaxa.4127.3.10 Read, G. (2016) Scolelepis Blainville, 1828. In: Read, G. & Fauchald, K. (Eds.), World Polychaeta database. Available from: http://www.marinespecies.org/polychaeta/aphia.php?p=taxdetails&id=129623 (Last accessed on 14 March 2016) Richards, S.L. (1970) Spawning and reproductive morphology of Scolelepis squamata (Spionidae: Polychaeta). Canadian Journal of Zoology, 48, 1369–1379. http://dx.doi.org/10.1139/z70-234 Rioja, E. (1918) Adiciones a la fauna de anélidos del Cantábrico. Revista de la Real Academia de Ciencias Exactas, Físicas y Naturales de Madrid, 17, 54–79. Rocha, M.B. & Paiva, P.C. de (2012) Scolelepis (Polychaeta: Spionidae) from the Brazilian coast with a diagnosis of the genus. Zoologia, 29 (4), 385–393. http://dx.doi.org/10.1590/S1984-46702012000400011 Sato-Okoshi, W. & Abe, H. (2013) Morphology and molecular analysis of the 18S rRNA gene of oyster shell borers, Polydora species (Polychaeta: Spionidae), from Japan and Australia. Journal of the Marine Biological Association of the United Kingdom, 93 (5), 1279–1286. http://dx.doi.org/10.1017/S002531541200152X Saint-Joseph, A. de (1898) Les Annélides Polychètes des côtes de France (Manche et Océan). Annales des Sciences Naturelles, Zoologie, 8 (5), 209–464. Sikorski, A.V. (2001) Spionidae Grube, 1850. In: Jirkov, I.A. (Ed.), Polychaeta of the Arctic Ocean. Yanus—K Press, Moscow, pp. 271–332. Sikorski, A.V. & Pavlova, L.V. (2015) New species of Scolelepis (Polychaeta, Spionidae) from the Norwegian coast and Barents Sea with a brief review of the genus. Fauna norvegica, 35, 9–19. http://dx.doi.org/10.5324/fn.v35i0.1666 Söderström, A. (1920) Studien über die Polychaetenfamilie Spionidae. Inaugural-Dissertation. Almquist & Wicksells, Uppsala, 286 pp. Surugiu, V. (2005) Inventory of inshore polychaetes from the Romanian coast (Black Sea). Mediterranean Marine Science, 6, 51–73. http://dx.doi.org/10.12681/mms.193 Viéitez, J.M. (1976) Ecología de Poliquetos y moluscos de la playa de Meira (Ría de Vigo). I. Estudio de las comunidades. Investigación Pesquera, 40 (1), 223–248. Viéitez, J.M. (1981) Estudio de las comunidades bentónicas de dos playas de las rías de Pontevedra y Vigo (Galicia, España). Boletín. Instituto Español de Oceanografía, 6 (331), 242–258. Williams, J.D. (2007) New records and description of four new species of spionids (Annelida: Polychaeta: Spionidae) from the Philippines: the genera Dispio, Malacoceros, Polydora, and Scolelepis, with notes on the palp ciliation patterns of the genus Scolelepis. Zootaxa, 1459, 1–35. Zhou, J., Weiwei, J. & Xinzheng, L. (2009) A new species of Scolelepis (Polychaeta: Spionidae) from sandy beaches in China, with a review of Chinese Scolelepis species. Zootaxa, 2236, 37–49. Zhou, J. (2014) A new species of Scolelepis (Polychaeta: Spionidae) from Chinese seas. Raffles Bulletin of Zoology, 62, 490– 495. 176 · Zootaxa 4161 (2) © 2016 Magnolia Press SURUGIU

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