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Microspio moorei (Gravier, 1911) Figures 2A–G; 3A–K Mesospio moorei Gravier, 1911a: 100–105, Plates VII, figs 80–83, VIII, 84–86.— Gravier, 1911b: 313.— Augener, 1932: 39–40.— Hartman, 1966: 17, Plate IV, figs 1–3.— Bellan, 1975: 789.— Blake, 1983: 241.— Sicinski et al. 2011: 35, Table 1: 37. Microspio moorei Foster, 1971: 35.— Maciolek, 1990: 1113–1115, Table 1. Microspio cf. moorei Petti et al. 2006: 166, Table 1.— Sicinski et al. 2011: 37. Microspio sp. Barbosa et al. 2010: 1158, Table 1. Material examined. All samples collected in Fildes Bay, King George Island, South Shetland Islands, Antarctic Peninsula; 62º12’31,32”S 58º57’45,86”W: UDEA: CEMUA: ANNE:001594 (8 specimens); 0.3 m depth, low tide. Coll. M. Londoño & I. Fonseca. Feb. 24, 2017. UDEA: CEMUA: ANNE:001595 (7); 0.3 m depth, low tide. Coll. M. Londoño & I. Fonseca. Feb. 24, 2017. UDEA: CEMUA: ANNE:001596 (2); 1.3 m depth low tide. Coll. M. Londoño & I. Fonseca. Feb. 26, 2017. UDEA: CEMUA: ANNE:001597 (9); 1.3 m depth low tide. Coll. M. Londoño & I. Fonseca. Feb. 24, 2017. UDEA: CEMUA: ANNE:001598 (3); 1.3 m depth low tide. Coll. M. Londoño & I. Fonseca. Feb. 24, 2017. Description. Complete specimens with 4.1–12.3 mm long and 0.5–1.5 mm wide with 36–62 segments. In life and in alcohol, prostomium, peristomium, caruncle, and dorsum of first five chaetigers dark, subsequent segments with diminishing pigmentation, palps dark (Fig. 2A, B), ventral surface of first five segments with dark pigmentation, subsequent segments with pigmentation decreasing gradually, concentrated along midline up to chaetiger 12 (Fig. 2C); in life, specimens pink with visible blood vessel running inside the branchiae (Fig. 2D); ventral epidermal glands absent. Prostomium broadly rounded and tapered anteriorly (Figs 2A, 3A), posteriorly narrow, tapered in a narrow caruncle reaching the base of chaetiger 2 (Fig. 3A), with slightly elevated keel near base of palps (Fig. 3A). Occipital tentacle absent. Two pairs of black eyespots arranged in trapezoid, anterior pair larger, crescent-shaped, widely spaced; posterior pair smaller, rounded, closely spaced (Figs 2A, B, D, 3A). Peristomium long, collar-like, partially enveloping prostomium and extending around base of palps, not forming lateral wings (Figs 2A, 3A, B), separated from chaetiger 1. Palps long, thick, extending to chaetigers 8–11; palps longitudinally grooved, with dark brown pigment along both sides, except basally (Figs 2B, D, E, 3B); palpal sheath short, smooth, fused to anterior base of palps (Fig. 2B, D). Nuchal organs with medial ciliary bands around caruncle, extending to chaetiger 2, then turning laterally, with small gap between this and the second lateral band. From chaetiger 3, dorsum with two transverse rows of ciliated patches; the first row extending between branchial bases; the second row widely separated from the first, near segmental groove (Figs 2A, 3A), transverse rows of ciliated patches visible (Fig. 2A) up to around chaetiger 22. Branchiae from chaetiger 2 to almost posterior end; the first pair of branchiae slightly shorter and thinner or as long as those on following chaetigers (Figs 2A, D, 3A, C); longest through mid-body region, reaching dorsal midline (Fig. 2A, E), then becoming very small; short posteriorly (Fig. 2F); branchiae partly fused at the base with notopodial postchaetal lamellae anteriorly (Figs 2A, B, 3A), increasingly separate from lamellae posteriorly, flattened, robust, elongate, distally rounded (Figs 2A, B, 3A), with long cilia on inner margin. Notopodial postchaetal lamellae triangular, short on chaetiger 1; lamellae on chaetigers 2–8 small, subtriangular with rounded ventral edge (Figs 2A, E, 3A, C); thereafter becoming oval and slightly decreasing in size throughout the body (Fig. 2F). Notopodial prechaetal lamellae very short, rounded on chaetiger 1, robust, subtriangular on chaetigers 2–9 (Figs 2A, B, 3A, C); subsequent lamellae progressively decreasing in size, becoming round and smaller (Fig. 2F). Neuropodial postchaetal lamellae small, triangular on chaetiger 1 (Fig. 2E); subtriangular on chaetiger 2 (Figs 2E, 3B); subsequent neuropodial lamellae large, rounded, wider (Figs 2E, 3C), up to end of the body (Fig. 2F). Neuropodial prechaetal lamellae absent. Notopodial capillary chaetae on chaetiger 1 longer, thinner and alimbate, arranged in one row; capillary chaetae from chaetiger 2 arranged in two rows; both rows with slightly granulated, striated, unilimbate chaetae (Fig. 3D); posterior row with very long and pointed chaetae. All chaetigers with an additional superior fascicle; anterior chaetigers with 4–7 long, granulated capillary chaetae (Fig. 3E); middle chaetigers with short, thin, smooth and alimbate chaetae; posterior chaetigers with slender, smooth, long and alimbate chaetae (Fig. 3K). Neuropodial capillaries of chaetigers 1–3 arranged in one row; capillaries long, smooth, unilimbate; capillaries of subsequent chaetigers arranged in two rows, capillaries of both rows of same length, most dorsal capillaries stout, slightly granulated, striated and unilimbate (Fig. 3F); capillaries of ventral region slender, slightly granulated, unilimbate (Fig. 3G); inferior fascicle with 4–6 long, smooth, thin capillaries (Fig. 3H) in position of sabre chaetae usually present in most anterior chaetigers, around chaetiger 13 with granulate and long sabre chaetae (Fig. 3I), up to 3 per fascicle. Neuropodial hooded hooks (Fig. 3J) from chaetigers 14–17; up to 11 hooks per fascicle, accompanied by granulated, unilimbate capillaries in first chaetiger with hooks, thereafter only hooks. All hooks bidentate, with small tooth above main tooth (Fig. 3J). Pygidium long, with four short highly glandular digitate lobes surrounding the anal opening (Fig. 3K). Methyl Green staining pattern. Body destains fairly rapidly; stain is retained briefly on anterior-most end of body. Anterior parapodial lamellae initially stain deeply but rapidly lose the stain. Variation. The segments where the hooded hooks started varied from segment 14 in small specimens to segment 17 in the longest specimens. A significant positive linear regression was found between the chaetiger where the hooded hooks first appeared and the body length (R 2 = 0.22474; Permutation p = 0.0031) (Fig. 4A). Specimens with an average length of 6.3 mm (SD±0.94) showed hooks starting on chaetiger 14, and as the body length increased, hooks first appeared on segments up to chaetiger 17 in individuals with a length close to 11 mm (SD±1.17). Even though the statistical analysis was not significant (R 2 = 0.10305; Permutation p = 0.0653), it was also observed that in individuals with greater body width, hooded hooks first appeared on posterior segments, in such a way that specimens with a diameter greater than 1.1 mm (SD±0.30) had hooks starting in chaetiger 17 (Fig. 4B). A multivariate regression analysis supports these results and shows that as the polychaetes grow in length and width, the chaetiger where the hooded hooks first appear is progressively later (R 2 = 0.2235; p(regr) = 0.01907). Discussion. Specimens herein described become the only additional material that has been used for taxonomic purpose, since none of the recent material used for ecological studies from the type locality, Admiralty Bay, and identified as Microspio moorei (Sicinski 2004; Pabis & Sicinski 2010a, b), Microspio cf. moorei (Petti et al. 2006), Microspio sp. (Barbosa et al. 2010), and Mesospio moorei (Sicinski et al. 2011), was available for checking their identity. Taxonomic information given by Hartman (1966) and Blake (1983) are based on the description by Gravier (1911a), and comparative notes by Maciolek & Blake (2021) are based on the description by Blake (1983). The quantity of individuals obtained from Fildes Bay, adjacent to the type locality, were sufficient to evaluate the relationship between the segment where different types of chaetae first appear and the body length and number of segments presented by each complete individual. The original description, based on the holotype with 16 mm in length, considered chaetiger 15, where hooded hooks appear for the first time, as a character with taxonomic importance (Gravier, 1911a); nevertheless, regression analyses from additional material herein studied indicate that this character is size dependent, so increasing body length leads to hooded hooks occurring more posteriorly on the body (Fig. 4). More analyses on ontogenetic development are needed to assess the physiological bases of this morphological variability in chaetation. Type locality: Admiralty Bay, King George Island, South Shetland Islands, Antarctic Peninsula. Distribution: This species has been identified only in the South Shetland Islands, Antarctic Peninsula, in Admiralty Bay by Gravier (1911a, b), Sicinski (2004), Pabis & Sicinski (2010a, 2010b), and in its different inlets (lagoons or fjord-like shaped bays), Mackellar, Martel, and Ezcurra (Barbosa et al. 2010), in Deception Island by Augener (1932), and in Fildes Bay, King George Island, in this study. From 0.3 m (this research) to 30 m depth (sensu Augener 1932)., Published as part of Fonseca-González, Idalyd, Londoño-Mesa, Mario H. & Delgado-Blas, Víctor H., 2022, Redescription of Microspio moorei (Gravier, 1911) (Annelida: Spionidae) with inclusion of a taxonomic key for all the species of the genus, pp. 334-344 in Zootaxa 5120 (3) on pages 337-341, DOI: 10.11646/zootaxa.5120.3.2, http://zenodo.org/record/6389455, {"references":["Gravier, C. (1911 a) Annelides polychetes recueillis par la seconde expedition antarctique francaise (1908 - 1910). Deuxieme Expedition Antarctique Francaise (1908 - 1910) Comamandee par le Dr. Jean Charcot, 1, 1 - 165, 12 plates.","Gravier, C. (1911 b) Expedition Antarctique Francaise du \" Pourquoi-Pas \", dirigee par le Dr. J. - B. Charcot (1908 - 1910). Especes nouvelles d'annelides polychetes. Bulletin du Museum National d'Histoire Naturelle, 17 (5), 310 - 316.","Augener, H. (1932) Antarktische und antiboreale Polychaeten nebst einer Hirudinee. Scientific Results of the Norwegian Antarctic Expeditions 1927 - 1928 et sqq., instituted and financed by Consul Lars Christensen. Det norske videnskaps-akademi i Oslo, 9, 1 - 85. https: // doi. org / 10.2307 / 4080428","Hartman, O. (1966) Polychaeta Myzostomidae and Sedentaria of Antarctica. In: Antarctic Research Series. Vol. 7. American Geophysical Union, Washington, D. C., pp. 1 - 158, 46 plates., 5 charts. https: // doi. org / 10.1029 / AR 007","Blake, J. A. (1983) Polychaetes of the family Spionidae from South America, Antarctica and adjacent seas and islands. In: Kornicker, L. (Ed.), Biology of The Antarctic Seas. Vol. 14. Antarctic Research Series 39. American Geophysical Union, Washington, D. C., pp. 205 - 287. https: // doi. org / 10.1029 / AR 039 p 0205","Sicinski, J., Jazdzewski, K., De Broyer, C., Presler, P., Ligowski, R., Nonato, E. F., Corbisier, T. N., Petti, M. A. V., Brito, T. A. S., Lavrado, H. P., Blazewicz-Paszkowycz, M., Pabis, K., Jazdzewska, A. & Campos, L. S. (2011) Admiralty Bay Benthos Diversity - A census of a complex polar ecosystem. Deep Sea Research II, 58, 30 - 48. https: // doi. org / 10.1016 / j. dsr 2.2010.09.005","Foster, N. M. (1971) Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea. Studies on the Fauna of Curacao and other Caribbean Islands, 36, 1 - 183.","Maciolek, N. J. (1990) A redescription of some species belonging to the genera Spio and Microspio (Polychaeta: Annelida) and description of three new species from the northwestern Atlantic Ocean. Journal of Natural History, 24, 1109 - 1141. https: // doi. org / 10.1080 / 00222939000770701","Petti, M. A. V., Nonato, E. F., Skowronski, R. S. P. & Corbisier, T. N. (2006) Bathymetric distribution of the meiofaunal polychaetes in the nearshore zone of Martel Inlet, King George Island, Antarctica. Antarctic Science, 18, 163 - 170. https: // doi. org / 10.1017 / S 0954102006000186","Barbosa, L. S., Soares-Gomes, A. & Paiva, P. C. (2010) Distribution of polychaetes in the shallow, sublittoral zone of Admiralty Bay, King George Island, Antarctica in the early and late austral summer. Natural Science, 2 (10), 1155 - 1163. https: // doi. org / 10.4236 / ns. 2010.210143","Sicinski, J. (2004) Polychaetes of Antarctic sublittoral in the proglacial zone (King George Island, South Shetland Islands). Polish Polar Research, 25 (1), 67 - 96.","Pabis, K. & Sicinski, J. (2010 a). Polychaete fauna associated with holdfasts of the large brown alga Himantothallus grandifolius in Admiralty Bay, King George Island, Antarctic. Polar Biology, 33, 1277 - 1288. https: // doi. org / 10.1007 / s 00300 - 010 - 0816 - x","Maciolek, N. J. & Blake, J. A. (2021) New species of Microspio Mesnil, 1896 (Annelida: Spionidae), with additions to the description of M. pigmentata (Reish, 1959) and comparative notes on all members of the genus. Proceedings of the Biological Society of Washington, 134 (1), 265 - 293. https: // doi. org / 10.2988 / 0006 - 324 X- 134.1.265","Pabis, K. & Sicinski, J. (2010 b). Distribution and diversity of polychaetes collected by trawling in Admiralty Bay: An Antarctic glacial fiord. Polar Biology, 33 (2), 141 - 151. https: // doi. org / 10.1007 / s 00300 - 009 - 0692 - 4"]}

Family Trichobranchidae Malmgren, 1866 Figs 1A, 7���8 Diagnosis (after Hutchings et al. 2021a, most important diagnostic characters highlighted in bold) Transverse prostomium attached to dorsal surface of upper lip; basal part as thick crest, eyespots sometimes present; distal part at base of upper lip or extending along lip. Buccal tentacles of two types, uniformly cylindrical and expanded at tips, spatulate. Peristomium forming lips, sometimes also a ventral lobe, as an extension of the lower lip; lips expanded, circular upper lip, distal margin folded or convoluted; lower lip button-like, usually continuing by ventral lobe, or expanded, forming large scoop-shaped process (Figs 7A���C, 8A, C���D). Segment I usually short, frequently only visible ventrally; anterior margin of anterior segments with lobes as low, even-length collars covering posterior margins of preceding segments, at least ventrally; ventro-lateral or lateral lobes on anterior segments sometimes present. Anterior segments poorly glandular ventrally, smooth, discrete shields absent; midventral groove extending from posterior segments with notopodia. Two to four pairs of branchiae, beginning from SGII, each pair with single, thick and elongate, tapered or foliaceous filament, or two pairs fused in single four lobed structure originating mid-dorsally between SGII���III or II���IV (Figs 7C, 8C���D). Notopodia beginning from SGIII���VI, typically terminating at SGXX; short, conical notopodia, chaetae emerging from central core on top, distal lobes absent; narrowly-winged notochaetae in both rows throughout. Neuropodia beginning on same segment as notopodia or slightly posteriorly, rarely beginning before notopodia; sessile neuropodia until termination of notopodia, neurochaetae emerging directly from body wall, as rectangular to foliaceous pinnules after termination of notopodia; thoracic neurochaetae as acicular uncini (Figs 1A, 7D, 8F), sometimes with small hood or beard below main fang; avicular abdominal uncini, with secondary teeth in rows on top and laterally to main fang. Nephridial papillae on SGIII usually present, other papillae sometimes present on SGVI and SGVII, but reduced to inconspicuous in most taxa. Pygidium smooth to slightly crenulate, sometimes bilobed. Remarks In the past, the Trichobranchidae family was considered to be a subfamily of Terebellidae (Fauvel 1927; Day 1967; Garrafoni & Lana 2004), but recent phylogenetic analyses support the hypothesis of a valid family (Glasby et al. 2004; Nogueira et al. 2013). The family includes only three genera, i.e., Octobranchus Marion & Bobretzky, 1875, Terebellides Sars, 1835, and Trichobranchus Malmgren, 1866. For Trichobranchus and Octobranchus, only three species of each occur in Europe. The genus Terebellides is very speciose and is represented in Europe by 19 species, 13 of them described in the last two years (Lavesque et al. 2019b; Parapar et al. 2020a) (Table 1). Main morphological characters for European species The number of branchiae is the best character to discriminate the different genera, with Terebellides having a single large branchia, Trichobranchus with two or three pairs of branchiae and finally Octobranchus with four pairs. Trichobranchus species are easy to differentiate based on the number of branchiae (two vs three) (Figs 7C, 8C) and the absence or presence of eyespots. In Octobranchus, the species differ by the shape of the branchiae (Fig. 8D) and the number of secondary teeth above the main fang of the uncini. Regarding Terebellides species, recent studies highlighted that several characters are very important for identification to the species level (Lavesque et al. 2019a; Parapar et al. 2020a, 2020b). However, as many cryptic species occur at a small geographical scale (Nygren et al. 2018), which currently are confirmed only by molecular analyses (Parapar et al. 2020a) much more work needs to be done to resolve all the species present. BRANCHIAE. Even if Terebellides branchiae seem to be very similar within the genus (Figs 7A���B, 8A���B), several morphological characters permit the discrimination of species, such as the presence of a fifth anterior branchial lobe (e.g., T. europaea), the degree of fusion of both upper and lower lobes (e.g.. not fused on T. ceneresi), the presence of long terminal filaments (e.g., in T. shetlandica) or short posterior processes (Fig. 7B), and finally the presence and the shape of papillae situated on the margins of the branchial lamellae (Fig. 8B) (e.g., T. lilasae). NOTOCHAETAE FROM FIRST CHAETIGER. The size of notochaetae of the first chaetiger varies between species. For most of the species, these chaetae are of a similar size compared to those of the following chaetigers. However, they can be absent or much shorter (e.g., T. ceneresi) or much longer (e.g., T. mediterranea). PRESENCE OF GENICULATE CHAETAE ON ONE OR TWO CHAETIGERS. The geniculate chaetae are exclusive to members of Terebellides and they are typically present on CH 6 (SG VIII) only (Fig. 8E), but in some species they are present on two chaetigers, as for example in T. bigeniculatus. UNCINI DENTICULATION. The different types of uncini follow the classifications provided by Parapar et al. (2020b) for thoracic uncini (Fig. 8F) and Parapar et al. (2020a) for abdominal uncini. These classifications are based on the ratio between the length of the main fang (rostrum) and the crest of secondary teeth (capitium), and the size and number of the secondary teeth. THORACIC CILIATED PAPILLAE. Following the recent study of Parapar et al. (2020a), the absence or the presence of thoracic ciliated papillae allow for the discrimination of Terebellides species. These papillae are situated dorsally to the thoracic notopodia (see for example Parapar et al. 2020a; Fig. 7B). METHYL GREEN PATTERN. The colouration of Terebellides specimens prior to identification is essential. Indeed, MG staining highlights the presence and the shape of the glandular region of the third thoracic chaetiger (e.g., undulating glandular region present and in members of T. gentili, oval for T. lilasae Fig. 7B) and the compact/striped pattern of the ventral part of anterior chaetigers (e.g., CH 4 (SG VI) white in T. ceneresi). Key to European species of Trichobranchidae (after Lavesque et al. 2019a and Parapar et al. 2020a) 1. One large branchia consisting of a stem and four lobes with transverse lamellae.....5 (Terebellides) ��� Two or three pairs of branchiae........................................................................... 2 (Trichobranchus) ��� Four pairs of branchiae........................................................................................... 4 (Octobranchus) 2. Two pairs of branchiae...................................................................................................................... 3 ��� Three pairs of branchiae, eyespots present................................................................................................................................................................................. Trichobranchus glacialis Malmgren, 1866 3. Eyespots absent......................................................................... Trichobranchus roseus Malm, 1874 ��� Eyespots present.................................................................................................................................... Trichobranchus demontaudouini Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 4. Pairs of branchiae of different shapes; abdominal uncini with three rows of secondary teeth above the main fang..................................................... Octobranchus floriceps Kingston & Mackie, 1980 ��� All pairs of branchiae similar; abdominal uncini with two rows of secondary teeth above the main fang..................................................................................... Octobranchus lingulatus (Grube, 1863) ��� Bases of branchiae covered by dorso-lateral lobes, abdominal uncini with two rows of secondary teeth above the main fang.............................. Octobranchus sikorskii (Leontovich & Jirkov. 2001) 5. Geniculate acicular chaetae on CH 5 (SG VII) and CH 6 (SG VIII)............................................................................................................. Terebellides bigeniculatus Parapar, Moreira & Helgason, 2011 ��� Geniculate acicular chaetae on CH 6 (SG VI) only........................................................................... 6 6. Branchial lamellae without marginal papillae.................................................................................. 7 ��� Branchial lamellae with marginal papillae..................................................................................... 15 7. Lower branchial lobes with long filaments....................................................................................... 8 ��� Lower branchial lobes with or without short projections................................................................. 9 8. Glandular region on CH 3 (SG V) present; branchial lamellae pointed; notochaetae from CH 1 longer than following ones; dorsal papillae absent............................................................................................................... Terebellides parapari Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 ��� Glandular region on CH 3 (SG V) absent; branchial lamellae rounded; all notochaetae equal-sized; dorsal papillae present........................ Terebellides shetlandica Parapar, Moreira & O���Reilly, 2016 9. Ventral white band present on CH 4 (SG VI) after MG staining..................................................... 10 ��� No distinct pattern on CH 4 (SG VI) after MG staining...................................................................11 10. Large species (> 30 mm); 5 th branchial lobe present; notochaetae of CH 1 (SG III) similar to following ones; main fang of thoracic uncini straight.................................... Terebellides gracilis Malm, 1874 ��� Small species (Terebellides ceneresi Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 11. First notopodia and notochaetae longer than following ones............................................................................................................................... Terebellides mediterranea Parapar, Mikac & Fiege, 2013 ��� First notopodia and notochaetae similar or shorter than following ones........................................ 12 12. Large-sized species (> 50 mm); dorsal rounded projections on CH 1��� CH 5 conspicuous............... 13 ��� Small-sized species (Terebellides kongsrudi Parapar, Capa, Nygren & Moreira, 2020 and Terebellides bakkeni Parapar, Capa, Nygren & Moreira, 2020 complex ��� Abdominal uncini of type 2 (capitium of about same length as main fang, capitium complex composed of a first row of 4(5) denticles and a variable number of teeth in two more rows)..................................................................................................................... Terebellides stroemii Sars, 1835 14. Glandular region on CH 3 (SG V) and 5 th branchial lobe both absent................................................................................................................................................... Terebellides atlantis Williams, 1984 ��� Glandular region on CH 3 (SG V) and 5 th branchial lobe both present............................................................................ Terebellides gralli Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 15. Glandular region on CH 3 (SG V) rounded or oval......................................................................... 16 ��� Glandular region on CH 3 (SG V) otherwise.................................................................................. 17 16. Glandular region on CH 3 (SG V) staining in white, branchial lamellae with rounded papillae, CH 1��� 3 without conspicuous dorsal projection....................................................................................................................... Terebellides lilasae Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 ��� Glandular region on CH 3 (SG V) staining in blue, branchial lamellae with conical papillae, CH 1���3 with conspicuous dorsal projection................................................................................................................................ Terebellides bonifi Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 17. Most branchial lamellae with marginal papillae............................................................................. 18 ��� Only anterior branchial lamellae with marginal papillae................................................................ 19 18. Branchial lamellae with digitiform papillae, upper lip elongated; MG staining pattern as compact bands from CH 1���5.................................................................................................................................................... Terebellides resomari Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 ��� Branchial lamellae with widely spaced, small and elongated digitiform papillae; MG staining pattern leaving white stripes from CH 1���5................................................................................................................................ Terebellides gentili Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 19. Thoracic uncini type 1 (main fang vs capitium length ratio 2(3)/1; capitium with 2(3) large teeth, following ones much smaller).................................................................................................................................................................. Terebellides ronningae Parapar, Capa, Nygren & Moreira, 2020 ��� Thoracic uncini type 3 (main fang vs. capitium length ratio 1/1; capitium with 4(5) mid-sized teeth, following ones slightly smaller)..................................................................................................... 20 20. Deep-water species, mostly found below 200 m deep.............................................................................................................................. Terebellides norvegica Parapar, Capa, Nygren & Moreira, 2020 ��� Shallow-water species, mostly found above 100 m deep.................................................................................. Terebellides europaea Lavesque, Hutchings, Daffe, Nygren & Londo��o-Mesa, 2019 and Terebellides scotica Parapar, Capa, Nygren & Moreira, 2020 complex, Published as part of Lavesque, Nicolas, Hutchings, Pat, Londo��o-Mesa, Mario H., Nogueira, Jo��o M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonif��cio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, C��line, Humbert, Suzie, Janson, Anne-Laure, Jourde, J��r��me, Labrune, C��line, Lamarque, Bastien, Latry, Lise, Garrec, Vincent Le, Pelaprat, Corine, Pezy, Jean-Philippe, Sauriau, Pierre-Guy & Montaudouin, Xavier De, 2021, The " Spaghetti Project ": the final identification guide to European Terebellidae (sensu lato) (Annelida, Terebelliformia), pp. 108-156 in European Journal of Taxonomy 782 (1) on pages 136-141, DOI: 10.5852/ejt.2021.782.1593, http://zenodo.org/record/5781605, {"references":["Malmgren A. J. 1866. Nordiska Hafs-Annulater. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar 22: 355 - 410. Available from https: // www. biodiversitylibrary. org / part / 244483 [accessed 8 Nov. 2021].","Hutchings P., Nogueira J. M. N. & Carrerette O. 2021 a. Terebellidae Johnston, 1846. In: Schmidt- Rhaesa A. Hr., Beutel R. G., Glaubrecht M., Kristensen N. P., Prendini L., Purschke G., Richter S., Westheide, W. & Leschen R. Z. E. (eds) Handbook of Zoology. A Natural History of the Phyla of the Animal Kingdom: 1 - 64. Walter de Gruyter & Co, Berlin.","Fauvel P. 1927. Polychetes Sedentaires. Addenda aux Errantes, Archiannelides, Myzostomaires. Faune de France 16, Lechevalier, Paris.","Day J. H. 1967. A Monograph on the Polychaeta of Southern Africa. Part 2. Sedentaria. Trustees of the British Museum (Natural History), London. https: // doi. org / 10.5962 / bhl. title. 8596","Glasby C. J., Hutchings P. & Hall K. 2004. Assessment of monophyly and taxon affinities within the polychaete clade Terebelliformia (Terebellida). Journal of the Marine Biological Association of the United Kingdom 84: 961 - 971. https: // doi. org / 10.1017 / S 0025315404010252 h","Nogueira J. M. M., Fitzhugh K. & Hutchings P. 2013. The continuing challenge of phylogenetic relationships in Terebelliformia (Annelida: Polychaeta). Invertebrate Systematics 27: 186 - 238. https: // doi. org / 10.1071 / IS 12062.","Marion A. F. & Bobretzky N. V. 1875. Etude des Annelides du Golfe de Marseille. Annales des Sciences Naturelles, Sixieme Serie 2: 1 - 106. Available from https: // www. biodiversitylibrary. org / page / 33155516 [accessed 8 Nov. 2021].","Sars M. 1835. Beskrivelser og Iagttagelser over nogle maerkelige eller nye i Havet ved den Bergenske Kyst Levende Dyr af Polypernes, Acalephernes, Radiaternes, Annelidernes og Molluskernes classer, med en kort Oversigt over de hidtil af Forfatteren sammesteds fundne Arter og deres Forekommen. T. Hallager, Bergen. https: // doi. org / 10.5962 / bhl. title. 13017","Lavesque N., Daffe G., Grall J., Zanol J., Gouillieux B., Hutchings P. 2019 b. Guess who? On the importance of using appropriate name: case study of Marphysa sanguinea (Montagu, 1813). 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Family Polycirridae Malmgren, 1866 Figs 1B, 2 Diagnosis (after Hutchings et al. 2021a; most important diagnostic characters highlighted in bold) Transverse prostomium attached to dorsal surface of upper lip; basal part usually as thick horse-shoe shaped crest, eye spots absent; distal part either as another thick crest, with flaring distal lobes, with or without mid-dorsal process, or extending along upper lip until near anterior margin of lip; prostomium frequently extending ventrally, terminating laterally to mouth (Fig. 2A���D). Buccal tentacles of two types at least, short ones thin, uniformly cylindrical, long tentacles stouter, expanded at tips to variable degrees, distally spatulate (Fig. 2B, D) or more specialised. Peristomium forming lips; lips expanded, upper lip large, frequently circular and convoluted, folded into three lobes; swollen lower lip, only midventral or cushion-like across ventrum, sometimes extending posteriorly for a few segments (Fig. 2A��� D). Segment I reduced, frequently only visible ventrally, sometimes completely hidden. Segment II distinctly narrower than following segments, constricting body posteriorly to ���lips head���; SG II usually with rectangular or pentagonal mid-ventral shield at beginning of mid-ventral groove, sometimes extending anteriorly through SG I until near posterior margin of lower lip (Fig. 2C). Anterior segments highly glandular ventrally, frequently papillose or tessellated, with paired ventro-lateral pads separated from each other within pairs by mid-ventral groove extending from SG II���IV to posterior body (Fig. 2A���D). Branchiae absent. Notopodia, if present, from SG III (Fig. 2A���D), extending for variable number of segments, usually few; bilobed, elongate notopodia, post-chaetal lobes sometimes longer, notochaetae originating between lobes along all extension of notopodia, separating lobes from base on ventral side of notopodia (Fig. 2A���D); notochaetae winged (Fig. 2E) and/or pinnate, wings of variable width. Neuropodia, if present, located posteriorly to notopodia, frequently from posterior thoracic segments or only on abdomen; neurochaetae as acicular spines or avicular uncini, of two types, and arranged in a single row (Figs 1C, 2F���G). Nephridial and genital papillae usually present, at anterior bases of all notopodia, or only at anteriormost notopodia (Fig. 2A). Pygidium smooth or with rounded ventral papilla. Remarks This family was previously considered as a subfamily of Terebellidae (Polycirrinae Malmgren, 1866), but was recently raised to familial level after a comprehensive phylogenetic analysis showed the monophyly of this group (Nogueira et al. 2013). Polycirridae is represented by six genera (Amaeana Hartman, 1959; Biremis Polloni, Rowe & Teal, 1973; Enoplobranchus Verrill, 1879; Hauchiella Levinsen, 1893; Lysilla Malmgren, 1866 and Polycirrus Grube, 1850), distinguished from each other by the presence/ absence of noto- and neuropodia, and if present, the type of neurochaetae. Only Amaeana (Fig. 2A, C), Hauchiella, Lysilla and Polycirrus (Fig. 2B, D���G) are represented in European waters (Lavesque et al. 2020b) (Table 1). Main morphological characters of European species PARAPODIA. The parapodia of the members of this family are extremely important to separate the different genera. The genus Hauchiella is characterised by the absence of parapodia and Lysilla by the absence of neuropodia only. The neuropodia of members of Amaeana are characterised by the presence of spines, while those of Polycirrus bear avicular uncini (Figs 1B, 2F���G). Within the genus Polycirrus, the number and location of segments with notopodia and/or neuropodia are of important taxonomic value. Particularly, some species have uncini present only on abdominal segments, i.e., on segments without notopodia, and others have uncini starting before the end of the thorax, on segments bearing also notopodia. SHAPE OF THE LIPS. As for other terebellids, polycirrids have a peristomium with well-defined upper and lower lips. The upper lip is large and can be trilobed (Fig. 2B) or with a single medial lobe (Fig. 2D). Generally, the upper lip is trilobed but the lobes differ in size and shape and lateral lobes can be reduced or well developed. The shape and the size of the lower lip is also highly variable between species. This lip can be rectangular, squared, rounded or subtriangular, swollen or not, longer than wide or wider than long (Fig. 2B���D). . NOTOCHAETAE. Two types of notochaetae can be present: winged chaetae as for P. glasbyi (Fig. 2E) and/ or pinnate as for P. plumosus. The winged notochaetae have wings of different width which are often conspicuous under light microscope but appear hirsute under SEM (Fig. 2E). UNCINI SHAPE AND DENTICULATION. In Polycirrus two types of uncini are present: Type 1 with a short occipitum (back) and a straight to slightly convex base (Fig. 1B); and Type 2 with a long occipitum and a concave base (Glasby & Hutchings 2014). To date, all described European species have Type 1 uncini. The denticulation of uncini is also helpful in separating species, with the presence (as for P. catalanensis) (Fig. 2F) or the absence (as for P. arenivorus) of a main tooth above the main fang, and the number of rows of secondary teeth. Key to European species of Polycirridae (after Lavesque et al. 2020b) 1. Parapodia absent (no chaetae)............................................. Hauchiella tribullata (McIntosh, 1869) ��� Parapodia present.............................................................................................................................. 2 2. Only notopodia present....................................................................................................... 3 (Lysilla) ��� Notopodia and neuropodia present................................................................................................... 4 3. Notochaetae with smooth tips, 6 pairs of thoracic papillae............... Lysilla loveni Malmgren, 1866 ��� Notochaetae with plumose tips, 9 pairs of thoracic papillae............ Lysilla nivea Langerhans, 1884 4. Neuropodia with spines..................................................................................................5 (Amaeana) ��� Neuropodia with avicular uncini..................................................................................6 (Polycirrus) 5. Upper lip without lobe, lower lip rounded, long achaetous region.......................................................................................................... A. gremarei Lavesque, Hutchings, Daffe & Londo��o-Mesa, 2020 ��� Upper lip with trilobed, lower lip rectangular, short achaetous region......................................................................................................................................................... Amaeana trilobata (Sars, 1863) 6. With 28 or more segments with notochaetae.................................................................................... 7 ��� With 22 or fewer segments with notochaetae................................................................................... 8 7. With 29 segments with notopodia, neuropodia from SG XII, lower lip longer than wide, uncini without a main tooth above the main fang........................... Polycirrus arenivorus (Caullery, 1915) ��� With 46 segments with notopodia, neuropodia from SG XIV, lower lip longer than wide, uncini with a main tooth above the main fang............................................. Polycirrus aurantiacus Grube, 1860 ��� With 28 segments with notopodia, neuropodia from SG XV, lower lip wider than long, uncini with a main tooth above the main fang........................................................................................................................................... Polycirrus gujanensis Lavesque, Hutchings, Daffe & Londo��o-Mesa, 2020 8. Neuropodia beginning before SG VIII............................................................................................. 9 ��� Neuropodia beginning between SG IX and SG XII....................................................................... 10 ��� Neuropodia beginning after SG XIII.............................................................................................. 14 9. Upper lip trilobed, lower lip wider than long, uncini with 2 rows of teeth above the main tooth.......................................................................................... Polycirrus asturiensis Cepeda & Lattig, 2016 ��� Upper lip with single medial lobe, lower lip longer than wide, uncini with 1 row of teeth above the main tooth........................... Polycirrus idex Lavesque, Hutchings, Daffe & Londo��o-Mesa, 2020b 10. Uncini without a main tooth about the main fang.............. Polycirrus norvegicus Wollebaek, 1912 ��� Uncini with a main tooth about the main fang................................................................................11 11. Lower lip subtriangular, pointed towards mouth............................................................................ 12 ��� Lower lip oval or oblong................................................................................................................ 13 12. With 12 or 13 segments with notopodia, lower lip longer than wide......................................................................................................................................... Polycirrus denticulatus Saint-Joseph, 1894 ��� With 16 segments with notopodia, lower lip wider than long........................................................................................................................................................... Polycirrus elisabethae McIntosh, 1915 13. With 18 or more segments with notopodia, lower lip oval, ventro-lateral pads not separated by a large mid-ventral groove............................................................................................................................................................... Polycirrus glasbyi Lavesque, Hutchings, Daffe & Londo��o-Mesa, 2020 ��� Fewer than 18 segments with notopodia, lower lip oblong, ventro-lateral pads separated by a large midventral groove................ Polycirrus readi Lavesque, Hutchings, Daffe & Londo��o-Mesa, 2020 14. With 16 or more segments with notopodia..................................................................................... 15 ��� Fewer than 16 segments with notopodia........................................................................................ 17 15. Neuropodia beginning from SG XIV���XVI.................................................................................... 16 ��� Neuropodia beginning from SG XVIII���XX....................... Polycirrus plumosus (Wollebaek, 1912) 16. Upper lip elongated, uncini with a main tooth above the main fang, ventro-lateral pads well developed..................... Polycirrus nogueirai Lavesque, Hutchings, Daffe & Londo��o-Mesa, 2020 ��� Upper lip semicircular, uncini without a main tooth above the main fang, ventro-lateral pads poorly defined................................................................................................ Polycirrus arcticus Sars, 1865 17. Neuropodia beginning from SG XIV, uncini with four teeth above the main fang arranged in single vertical series; lower lip large, shield-like, wider than long......... Polycirrus latidens Eliason, 1962 ��� Neuropodia beginning from SG XV or after, secondary teeth of uncini not as above................... 18 18. Upper lip trilobed, lower lip subtriangular pointed toward mouth............................................................................................................................................................... Polycirrus medusa Grube, 1850 ��� Upper lip with a single median lobe, lower lip not subtriangular.................................................. 19 19. Upper lip with thick medial lobe, uncini with two small lateral teeth above the main tooth, lower lip rectangular longer than wide................................................................................................................................................ Polycirrus catalanensis Lavesque, Hutchings, Daffe & Londo��o-Mesa, 2020 ��� Upper lip with elongated triangular medial lobe, uncini with two rows of teeth above the main tooth, lower lip oval and wider than long.................................................................................................................................................... P. pennarbedae Lavesque, Hutchings, Daffe & Londo��o-Mesa, 2020, Published as part of Lavesque, Nicolas, Hutchings, Pat, Londo��o-Mesa, Mario H., Nogueira, Jo��o M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonif��cio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, C��line, Humbert, Suzie, Janson, Anne-Laure, Jourde, J��r��me, Labrune, C��line, Lamarque, Bastien, Latry, Lise, Garrec, Vincent Le, Pelaprat, Corine, Pezy, Jean-Philippe, Sauriau, Pierre-Guy & Montaudouin, Xavier De, 2021, The " Spaghetti Project ": the final identification guide to European Terebellidae (sensu lato) (Annelida, Terebelliformia), pp. 108-156 in European Journal of Taxonomy 782 (1) on pages 112-123, DOI: 10.5852/ejt.2021.782.1593, http://zenodo.org/record/5781605, {"references":["Malmgren A. J. 1866. Nordiska Hafs-Annulater. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar 22: 355 - 410. Available from https: // www. biodiversitylibrary. org / part / 244483 [accessed 8 Nov. 2021].","Hutchings P., Nogueira J. M. N. & Carrerette O. 2021 a. Terebellidae Johnston, 1846. 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A new species of Pista Malmgren, 1866 (Annelida, Terebellidae) from the Western Mediterranean Sea. ZooKeys 838: 71 - 83. https: // doi. org / 10.3897 / zookeys. 838.28634","Gaillande D. 1970. Une polychete Terebellidae nouvelle des cotes de Provence: Pista mediterranea n. sp. Tethys 2 (2): 443 - 448.","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 1873 - 76. Zoology 12: 1 - 554. Available from https: // www. biodiversitylibrary. org / page / 50688432 [accessed 8 Nov. 2021].","Pearson T. H. 1969. Scionella lornensis sp. nov., a new terebellid (Polychaeta: Annelida) from the west coast of Scotland, with notes on the genus Scionella Moore, and a key to the genera of the Terebellidae recorded from European waters. Journal of Natural History 3 (4): 509 - 516. https: // doi. org / 10.1080 / 00222936900770441","Linnaeus C. 1767. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio duodecima reformata. Typis Ioannis Thomae von Trattner, Wien [Vindobonae]. https: // doi. org / 10.5962 / bhl. title. 156772","Lezzi M. & Giangrande A. 2019. New species of Streblosoma (Thelepodidae, Annelida) from the Mediterranean Sea: S. pseudocomatus sp. nov., S. nogueirai sp. nov. and S. hutchingsae sp. nov. Journal of Natural History 52 (43 - 44): 2857 - 2873. https: // doi. org / 10.1080 / 00222933.2018.1556357","Sars G. O. 1872. Diagnoser af nye Annelider fra Christianiaforden, efter Professor M. Sar's efterladte Manuskripter. Forhandlinger i Videnskabs-Selskabet i Christiania 1871: 406 - 417. Available from https: // biodiversitylibrary. org / page / 44067540 [accessed 8 Nov. 2021]","Fabricius O. 1780. Fauna Groenlandica, systematice sistens, Animalia Groenlandiae occidentalis hactenus indagata, quoad nomen specificum, triviale, vernaculumque synonyma auctorum plurium, descriptionem, locum, victum, generationem, mores, usum, capturamque singuli prout detegendi occasio fuit, maximaque parte secundum proprias observations. Impensis Ioannis Gottlob Rothe, Copenhagen et Leipzig [Hafniae et Lipsiae]. https: // doi. org / 10.5962 / bhl. title. 13489","Grube A. E. 1855. Beschreibungen neuer oder wenig bekannter Anneliden. Archiv fur Naturgeschichte 21 (1): 81 - 136. Available from https: // doi. org / 10.5962 / bhl. part. 13989 [accessed 8 Nov. 2021].","Kingston P. F. & Mackie A. S. Y. 1980. Octobranchus floriceps sp. nov. (Polychaeta: Trichobranchidae) from the northern North Sea with a re-examination of O. antarcticus Monro. Sarsia 65: 249 - 254. https: // doi. org / 10.1080 / 00364827.1980.10431487","Grube A. E. 1863. Beschreibung neuer oder wenig bekannter Anneliden. Sechster Beitrag. Archiv fur Naturgeschichte 29: 37 - 69. Available from https: // doi. org / 10.5962 / bhl. part. 9306 [accessed 8 Nov. 2021].","Williams S. J. 1984. The status of Terebellides stroemi (Polychaeta; Trichobranchidae) as a cosmopolitan species, based on a worldwide morphological survey, including description of new species. In: Hutchings P. A. (ed.) Proceedings of the First International Polychaete Conference, Sydney, Australia, 1984: 118 - 142. The Linnean Society of New South Wales, Sydney, Australia.","Parapar J., Capa M., Nygren A. & Moreira J. 2020 a. To name but a few: descriptions of five new species of Terebellides (Annelida, Trichobranchidae) from the North East Atlantic. ZooKeys 992: 1 - 58. https: // doi: 10.3897 / zookeys. 992.55977","Parapar J., Moreira J. & Helgason G. V. 2011. Taxonomy and distribution of Terebellides (Polychaeta, Trichobranchidae) in Icelandic waters, with the description of a new species. Zootaxa 2983 (1): 1 - 20. https: // doi. org / 10.11646 / zootaxa. 2983.1.1","Parapar J., Mikac B. & Fiege D. 2013. Diversity of the genus Terebellides (Polychaeta: Trichobranchidae) in the Adriatic Sea with the description of a new species. Zootaxa 3691 (3): 333 - 350. https: // doi. org / 10.11646 / zootaxa. 3691.3.3","Parapar J., Moreira J. & O'Reilly M. 2016. A new species of Terebellides (Polychaeta: Trichobranchidae) from Scottish waters with an insight into branchial morphology. Marine Biodiversity 46 (3): 211 - 225. https: // doi. org / 10.1007 / s 12526 - 015 - 0353 - 5","Sars M. 1835. Beskrivelser og Iagttagelser over nogle maerkelige eller nye i Havet ved den Bergenske Kyst Levende Dyr af Polypernes, Acalephernes, Radiaternes, Annelidernes og Molluskernes classer, med en kort Oversigt over de hidtil af Forfatteren sammesteds fundne Arter og deres Forekommen. T. Hallager, Bergen. https: // doi. org / 10.5962 / bhl. title. 13017","Glasby C. J. & Hutchings P. 2014. Revision of the taxonomy of Polycirrus Grube, 1850 (Annelida: Terebellida: Polycirridae). Zootaxa 3877 (1): 1 - 117. https: // doi. org / 10.11646 / zootaxa. 3877.1.1"]}

Loimia ramzega Lavesque, Bonifácio, Londoño-Mesa, Le Garrec & Grall, 2017 Figure 2B Loimia ramzega Lavesque et al. 2017a: 935–942, figs 2–5. Diagnosis. ( for a detailed description, see Lavesque et al. 2017a). Large sized animals with live specimens up to 650 mm long. Three pairs of arborescent branchiae on SG II–IV, two pairs of lateral lobes on SG I and III; first pair more ventral, second pair more developed and lateral, but oblique, with undulating edge, sixteen ventral shields from SG II (fused on SG II–III); uncini pectinate with five or six teeth in vertical row (Fig. 2B), pygidium with about 14 long conical papillae. Type locality. France, English Channel, Brittany, Plouguerneau, Lilia beach; 48°36’38”N 04°36’24”W. Type material. Holotype MNHN-IATYPE 1788, four paratypes MNHN-IATYPE 1789 to 1992. NMW. Z.2017.002.0001, one paratype, and six individuals. CEMUA-POLY-TERE-0100, one paratype. Distribution. Only known from the French coasts: Bay of Biscay (Bay of Brest, Grall pers. obs.), English Channel (Lavesque et al. 2017a), southern North Sea (Dunkerque, Lavesque pers. obs.). Habitat. Sandy beaches, intertidal ( Lavesque et al. 2017a), shallow waters, fine sands (this study). Remarks. Lomia ramzega is very distinct from other European species of Loimia (see Lavesque et al. 2017a), but surprisingly quite similar to Terebella gigantea Montagu, 1819. Despite the very sparse original description given by Montagu (1819) and the absence of type material, several morphological details can be found in this paper. These two species have a very large size (650 mm long for L. ramzega, 406 mm for T. gigantea) and approximately the same type locality (Brittany, English Channel, France for L. ramzega and Devon, English Channel, UK, for T. gigantea). Moreover, as Montagu wrote “with seventeen pairs of exserted fasciculi” in the description, this species probably does not belong to the genus Terebella genera which generally have notopodia until the end of the body, or on more than 25 segments (Hutchings et al. 2021b). However, Montagu’s description lacks some taxonomic details, particularly concerning uncini and shape of lateral lobes, which are very important in the genus Loimia. Montagu described the branchiae as relatively short, while they are very long on L. ramzega, and that ventral shields are brown, while they are blood red on L. ramzega. Finally, the tubes of the five Terebella species described by Montagu (1819) in his paper are “extremely delicate”, while a membrane (together with shells fragments and gravels) allows the tube of L. ramzega to “maintain a hard consistency”. Later, McIntosh (1922) moved T. gigantea to the genus Amphitrite. Even if several taxonomic characteristics, like the shape of the branchiae, the size of the animals, or the presence of lateral lobes are similar with L. ramzega, the shape of the uncini is very different (McIntosh 1922, plate CXXV, Fig. 10b). In that paper, McIntosh also suggested that A. gigantea could be the same as Terebella edwardsii Quatrefages, 1866, but this last species is different and valid as Amphitrite edwardsii (this study). Finally, we suggest that T. gigantea should be considered as a nomen dubium considering that the species probably does not belong to the genus Terebella and that there is no known type material. Montagu appears to have never designated and deposited type material, and the original descriptions lacks important information, so it should be a nomen dubium until a neotype is designated and described., Published as part of Lavesque, Nicolas, Daffe, Guillemine, Londoño-Mesa, Mario H. & Hutchings, Pat, 2021, Revision of the French Terebellidae sensu stricto (Annelida, Terebelliformia), with descriptions of nine new species, pp. 1-63 in Zootaxa 5038 (1) on pages 35-36, DOI: 10.11646/zootaxa.5038.1.1, http://zenodo.org/record/5502867, {"references":["Lavesque, N., Bonifacio, P., Londono-Mesa, M. H., Le Garrec, V. & Grall, J. (2017 a) Loimia ramzega sp. nov., a new giant species of Terebellidae (Polychaeta) from French waters (Brittany, English Channel). Journal of the Marine Biological Association of the United Kingdom, 97 (5), 935 - 942. https: // doi. org / 10.1017 / S 0025315417000571","Montagu, G. (1819) Descriptions of five British species of the genus Terebella. Transactions of the Linnean Society of London, 12 (2), 340 - 344. https: // doi. org / 10.1111 / j. 1095 - 8339.1817. tb 00231. x","Hutchings, P., Nogueira, J. M. N. & Carrerette, O. (2021 b) Terebellidae Johnston, 1846. In: Schmidt-Rhaesa, A. Hr., Beutel, R. G., Glaubrecht, M., Kristensen, N. P., Prendini, L., Purschke, G., Richter, S., Westheide, W. & Leschen, R. Z. E. (Ed.), Handbook of Zoology. A Natural History of the Phyla of the Animal Kingdom. Walter de Gruyter & Co, Berlin, pp. 1 - 64.","McIntosh, W. C. (1922) Notes from the Gatty Marine Laboratory, St. Andrews. No. XLIV. 1. On new and rare Polychaeta, Gephyrea, etc., from various regions. 2. Recent additions to the British marine Polychaeta (continued). Annals and Magazine of Natural History, Series 9, 9 (49), 1 - 30. https: // doi. org / 10.1080 / 00222932208632638","Quatrefages, A. de. (1866) Histoire naturelle des Anneles marins et d'eau douce. Annelides et gephyriens. Volume 2. Librairie Encyclopedique de Roret, Paris, 794 pp. https: // doi. org / 10.5962 / bhl. title. 122818"]}

Amphitrite figulus (Dalyell, 1853) Figure 4 Terebella figulus Dalyell, 1853: 191–197, pl. XXVII figs 1–2, pl. XXVIII figs 1–2. Amphitrite johnstoni.— Fauvel 1927: 248–249, fig. 85, a–g. Neoamphitrite figulus.— Holthe 1986: 100–101, fig. 42. Amphitrite figulus.— Jirkov 2020: 330, fig. 1c, 14–15. Other synonym. Amphitrite stimpsoni Meyer 1912. Material examined. MNHN-IA-PNT 127, posteriorly incomplete, English Channel, Dieppe, 49°55’5”N 1°04’28”E, intertidal, May 2020, posterior part used for molecular analysis. SMA-DIEP-Tere-02, posteriorly incomplete, English Channel, Dieppe, 49°55’5”N 1°04’28”E, intertidal, May 2020, posterior part used for molecular analysis. AM W.53324, complete, English Channel, Dieppe, 49°55’5”N 1°04’28”E, intertidal, May 2020, few parapodia used for SEM (plot SMA-DIEP-Tere-04). Description. Moderate-sized specimens, complete one being 47.9 mm long and 4.7 mm wide, for about 155 segments. Prostomium at base of upper lip, without eyespots, distal part forming shelf-like tentacular membrane from which numerous filiform, wrinkled and deeply grooved buccal tentacles originate (Fig. 4B, D). Peristomium forming lips; upper lip thick, hood-like, convoluted, broader than high; lower lip swollen, broader than high (Fig. 4D). Segment I clearly visible, forming protruding lobe below lower lip. Segments II–III with small ventro-lateral lobes, SG IV without lateral lobes (Fig. 4A–C). Three pairs of dichotomous branchiae, on SG II–IV, with wide medial gap; first pair the longest, situated slightly more dorsally; with numerous long filaments, arising from short stems (Fig. 4A–C). Dorsum of anterior chaetigers tessellated. Eleven ventral shields, rectangular, broader than long, present on SG III–XIII (Fig. 4A); absence of mid-ventral groove. Notopodia short, rectangular, present on SG IV–XXVII (n=24). Notochaetae almost straight, medially winged with wings of same width, and distally serrated (Fig. 4E–F); two rows of chaetae, those of anterior row less than half as long as those of posterior row. Neuropodia beginning from SG V, with uncini arranged in single rows on SG V–X, uncini in double rows on SG XI–XXV or XXVI, in a face-to-face arrangement, and in single rows again from SG XXVI–XXVII; thoracic neuropodia as low ridges, situated latero-ventrally (Fig. 4B–C); abdominal neuropodia raised from body and displaced more laterally (Fig. 4A). Uncini avicular, with short triangular heel, with distally pointed prow, large pointed to digitiform dorsal button inserted halfway between base of main fang and tip of prow, convex base; and crest with five rows of secondary teeth above main fang (Fig. 4G–H). Fifteen pairs of small globular nephridial and genital papillae present on SG III–XVII (Fig. 4B–C), first pair situated above base of second pair of branchiae, second pair below first notopodia and slightly displaced dorsally, subsequent pairs between noto- and neuropodia, slightly displaced dorsally. Nephridial papillae (first three pairs) larger than genital ones (from SG VI), last pairs difficult to observe. Pygidium rounded. Type locality. Probably North Sea coast of Scotland (Gil 2011). Type material. Could not be traced (Holthe 1986, Jirkov 2020). Distribution. In Europe, from Norwegian Sea to Aegan Sea (Gil 2011) and White Sea (GenBank accession number: HM417784) (Fig. 26). In France, from North Sea to Bay of Biscay (Fauvel 1927; Jirkov 2020, this study, Resomar database). Also recorded from Canada, Japan, Gulf of Mexico and Sea of Okhotsk (Gil 2011) but all these records have to be considered as doubtful. Mediterranean records could also correspond to misidentifications of Amphitrite rubra (Risso, 1826) (see Jirkov 2020). Habitat. In empty shells in deep water, among rocks in shallow intertidal pools, on mud or sandy mud, among Zostera, Fucus or Laminaria, on mussel and oyster banks (Gil 2011), in mud (this study), upper sublittoral to intertidal (this study, Jirkov 2020). This range of different habitats/depths suggests that these records may represent more than one species. Remarks. The original description is not sufficiently informative according to current standards, consisting of a mixture of behavioural considerations and morphological data. Specimens examined in our study match those of the description provided by Fauvel (1927, specimens from French coasts), and the description of Amphitrite johnstoni Malmgren, 1866, by Holthe (1986, specimens from Scandinavia) and by Jirkov (2020, specimens from White Sea and UK, and also from the Western Pacific). Obviously a detailed review of this species is required including the re-examination of material from all the above localities. Genus Amphitritides Augener, 1922 Type-species: Terebella gracilis (Grube, 1860), by subsequent designation. Diagnosis. (after Hutchings et al. 2021b). Transverse prostomium attached to dorsal surface of upper lip; basal part as a thick crest, eyespots may be present; distal part shelf-like. Buccal tentacles usually all uniformly cylindrical. Peristomium forming lips and continuing dorsally for a short extension, not forming a complete annulation; lips expanded, relatively short upper lip, hood-like, about as long as wide, distal margin rounded, slightly undulated; narrow, rectangular, mid-ventral lower lip. Segment I conspicuous all around, dorsally narrow, ventrally developed, with mid-ventral lobe marginal to mouth; additional lobes on anterior segments absent. Anterior segments highly glandular ventrally, with discrete, smooth to slightly corrugated, rectangular to trapezoidal shields. Two pairs of arborescent branchiae, on SG II–III, with short main stems. Rectangular to conical notopodia beginning on SG IV, extending for a variable number of segments; notochaetae all medially-winged and finely serrated distally, with basally bulbous wings. Neuropodia beginning on SG V, as low, sessile ridges throughout; neurochaetae as shorthandled avicular uncini, in completely separate double rows, beak to beak arrangement, from SG XI until posterior body. Nephridial papillae on SG III, genital papillae on some anterior segments, beginning from SG VI, between parapodial lobes or at anterior bases of notopodia. Remarks. Recently, Jirkov (2020) proposed the synonymisation of Amphitritides with Amphitrite, arguing that characters such as the number of pairs of neuropodia with double rows of uncini is not unique to species of Amphitritides, but can be also present in species of Amphitrite. This is a character highly significant in terms of convergence in the family, and so it could be exhibited by some other genera besides these two. On the other hand, the structure of the lateral lobes is poorly understood in the whole family, and there is a great deal of misinterpretations regarding the shape, structure and orientation of the lateral lobes vs. the lateral ridges. Our suggestion is that Amphitritides has no lobes (except mid-ventral one on SGI, as described in the diagnosis), and which clearly differentiates it from Amphitrite, which has lobes on anterior segments, of variable length, usually on SG II–IV. It is clear, however, that the diagnosis of a genus is not based on a unique character that distinguishes it from the rest, but on a combination of characters, which defines the genus. It is also clear that a major revision of this group of genera, based on type material, if available, and fresh topotype material, is required in order to solve the relations within this group using both morphological and molecular data., Published as part of Lavesque, Nicolas, Daffe, Guillemine, Londoño-Mesa, Mario H. & Hutchings, Pat, 2021, Revision of the French Terebellidae sensu stricto (Annelida, Terebelliformia), with descriptions of nine new species, pp. 1-63 in Zootaxa 5038 (1) on pages 13-16, DOI: 10.11646/zootaxa.5038.1.1, http://zenodo.org/record/5502867, {"references":["Dalyell, J. G. (1853) The Powers of the Creator displayed in the Creation: or, observations on life amidst the various forms of the humbler tribes of animated nature with practical comments and illustrations, volume 2. John van Voorst. London, 359 pp. https: // doi. org / 10.5962 / bhl. title. 10022","Fauvel, P. (1927) Polychetes Sedentaires. Addenda aux Errantes, Archiannelides, Myzostomaires. Faune de France 16. Lechevalier, Paris, 494 pp.","Holthe, T. (1986) Polychaeta, Terebellomorpha. Marine Invertebrates of Scandinavia, 7, 1 - 194.","Jirkov, I. A. (2020) Review of the European Amphitrite (Polychaeta: Terebellidae) with description of two new species. Invertebrate Zoology, 17 (4) 311 - 360. https: // doi. org / 10.15298 / invertzool. 17.4.01","Meyer, A. H. (1912) Die Amphicteniden, Ampharetiden und Terebelliden der Nord- und Ostsee. Inaugural-Dissertation zur Erlangung der Doktorwurde der hohen philosophischen Fakultat der Koniglichen Christian-Albrechts-Universitat zu Kiel. Heider Anzeiger, Heide, 1 - 68.","Gil, J. (2011) The European Fauna of Annelida Polychaeta. Ph. D. Dissertation, Departamento de Biologia Animal, Faculdade de Ciencias, Universidade de Lisboa, Lisboa, xlii + 1554 pp.","Risso, A. (1826) Histoire naturelle des principales productions de l'Europe meridionale et particulierement de celles des environs de Nice et des Alpes Maritimes. Volume 4. Levrault, Paris, 439 pp. https: // doi. org / 10.5962 / bhl. title. 58984","Malmgren, A. J. (1866) Nordiska Hafs-Annulater. Ofversigt af Kongiliga Veteskaps-Akademiens Forhandlingar, 22, 355 - 410.","Augener, H. (1922) Uber littorale Polychaeten von Westindien. Sitzungsberichte der Gesellshaft naturforschender Freunde zur Berlin, 1922 (3 - 5), 38 - 63.","Grube, A. E. (1860) Beschreibung neuer oder wenig bekannter Anneliden. Beitrag: Zahlreiche Gattungen. Archiv fur Naturgeschichte, Berlin, 26, 71 - 118, pls. 1 - 3.","Hutchings, P., Nogueira, J. M. N. & Carrerette, O. (2021 b) Terebellidae Johnston, 1846. In: Schmidt-Rhaesa, A. Hr., Beutel, R. G., Glaubrecht, M., Kristensen, N. P., Prendini, L., Purschke, G., Richter, S., Westheide, W. & Leschen, R. Z. E. (Ed.), Handbook of Zoology. A Natural History of the Phyla of the Animal Kingdom. Walter de Gruyter & Co, Berlin, pp. 1 - 64."]}

Key to European species of Terebellidae (sensu stricto) Based on Gil 2011; Jirkov 2020; Hutchings et al. 2021b. 1A. Peristomiun ventrally forming a larg conical process.................... Artacama proboscidea Malmgren, 1866 ** 1B. Absence of peristomial ventral process................................................................ 2 2A (1B). Notochaetae on more than 25 segments, body uniform throughout.......................................... 3 2B. Notochaetae on 25 or fewer segments, thorax and abdomen clearly defined................................... 5 3A (2A). Branchiae absent................................................... Baffinia hesslei (Annenkova, 1924) ** 3B. Branchiae present...................................................................... 4 (Terebella) 4A. Branchiae on SG II–IV, five pairs of nephridial and genital papillae............. Terebella lapidaria Linnaeus, 1767 4B. Branchiae on SG II–III and V, 12 pairs of nephridial and genital papillae................... Terebella banksyi n. sp. 5A (2B). Absence of branchiae............................................................................. 6 5B. Presence of branchiae............................................................................ 12, Published as part of Lavesque, Nicolas, Daffe, Guillemine, Londoño-Mesa, Mario H. & Hutchings, Pat, 2021, Revision of the French Terebellidae sensu stricto (Annelida, Terebelliformia), with descriptions of nine new species, pp. 1-63 in Zootaxa 5038 (1) on pages 56-57, DOI: 10.11646/zootaxa.5038.1.1, http://zenodo.org/record/5502867, {"references":["Gil, J. (2011) The European Fauna of Annelida Polychaeta. Ph. D. Dissertation, Departamento de Biologia Animal, Faculdade de Ciencias, Universidade de Lisboa, Lisboa, xlii + 1554 pp.","Jirkov, I. A. (2020) Review of the European Amphitrite (Polychaeta: Terebellidae) with description of two new species. Invertebrate Zoology, 17 (4) 311 - 360. https: // doi. org / 10.15298 / invertzool. 17.4.01","Hutchings, P., Nogueira, J. M. N. & Carrerette, O. (2021 b) Terebellidae Johnston, 1846. In: Schmidt-Rhaesa, A. Hr., Beutel, R. G., Glaubrecht, M., Kristensen, N. P., Prendini, L., Purschke, G., Richter, S., Westheide, W. & Leschen, R. Z. E. (Ed.), Handbook of Zoology. A Natural History of the Phyla of the Animal Kingdom. Walter de Gruyter & Co, Berlin, pp. 1 - 64.","Malmgren, A. J. (1866) Nordiska Hafs-Annulater. Ofversigt af Kongiliga Veteskaps-Akademiens Forhandlingar, 22, 355 - 410.","Annenkova, N. P. (1924) Neues uber die Verbreitung einiger Arten der Polychaeten. Comptes Rendus de l'Academie des Sciences de Russie, Leningrad, 1924, 125 - 128.","Linnaeus, C. (1767) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio duodecima reformata, 1 (2), 533 - 1327 + 37 pp."]}

Amphitritides gracilis (Grube, 1860) Figures 5 and 6 Terebella gracilis. Grube 1860: 99. Amphitrite gracilis.— Saint-Joseph 1894: 198–201, pl. VIII, figs. 224; Fauvel 1927: 252–253, fig. 87, a–e; Jirkov 2020: 332– 334, figs 2B, 16–17. Amphitritides gracilis.— Holthe 1986: 104–106, fig. 45; Arvanitidis & Koukouras 1995: Table 1. Other synonyms. Physelia scylla Quatrefages, 1866; Terebella gelatinosa Keferstein, 1862; Terebella laevirostris Claparède, 1869. Material examined. MNHN-IA-PNT 128, posteriorly incomplete, Northeastern Atlantic, Bay of Biscay, Bay of Brest, 48°21’54”N 4°26’00”W, depth 5 m, February 2013. SMA-BR-Amphi-01, posteriorly incomplete, Northeastern Atlantic, Bay of Biscay, Bay of Brest, 48°21’54”N 4°26’00”W, depth 5 m, February 2013, mounted for SEM. Description. Small specimens, all incomplete, largest specimen 14.7 mm long and 1.9 mm wide, for 33 segments. Prostomium at base of upper lip without eyespots, distal part forming a shelf-like tentacular membrane from which long, thin and non-grooved buccal tentacles originate (few remaining) (Fig. 5B), continuing ventrally as a thin lobe (Fig. 5C). Peristomium forming lips; upper lip, hood-like, broader than high; lower lip swollen, very glandular and ridged, slightly broader than high (Figs 5A, C; 6A–B). Segment I dorsally narrow, more developed ventrally, forming protruding lobe below ventral lobe of prostomium and lower lip (Fig. 5C). Segments II–IV without lateral lobes (Figs 5A–B; 6A–B). Ventral shields on SG II–XII (n=11), rectangular, wide and thin, shields separated by deep incision (Fig. 5C); replaced by groove posteriorly. Two pairs of short, arborescent branchiae on SG II–III, both pairs with short, thick main stem, with dichotomous ramifications starting from close to base and ending with short wrinkled filaments (Fig. 5D); the two pairs inserted dorso-laterally, at the level of the following notopodia (Fig. 6A–B). Notopodia beginning from SG IV and extending for 19 segments. Notopodia well-developed, approximately rectangular, all similar in size, first two pairs aligned more dorsally, laterally aligned from SG VI (Figs 5A–B; 6A– B). Notochaetae in two rows with symmetrical limbation and curved serrated tips, first row shorter (Fig. 5E–F). Neuropodia starting from SG V, thoracic neuropodia as long lateral ridges slightly raised from surface of body, not reaching ventral groove (Figs 5B–C; 6A–B); abdominal neuropodia as short fleshy ridges, close to mid ventral groove. Uncini arranged in double rows from SG XI until end of body, in a face-to-face arrangement, with rows completely separated from each other. Uncini avicular, with short triangular heel, distally pointed prow downwardly directed, short dorsal button inserted halfway between base of main fang and tip of prow, convex base, and main fang surmounted by crest of five rows of secondary teeth (Figs 5G; 6C–D). Nephridial and genital papillae on SG III and SG VI–XII, inserted laterally and slightly anteriorly to branchial stem on SG III, and anteriorly to base of notopodia on SG VI–XII (NP). Pygidium not seen. Type locality. Scilly Isles, off southwest England, approx. 49°56’02”N 6°18’44”W. Type material. ZMB Q.5045, 2 syntypes. Distribution. Along all the French coasts (Fauvel 1927, this study, Resomar database). In Europe, from Scotland to Black Sea (Gil 2011; Jirkov 2020). Also recorded from Ivory Coast (Holthe 1986), but this record is considered to be doubtful. Habitat. Shallow waters, in maërl (rhodolith) beds (this study), soft muddy bottoms and sandy mud, under stones and among Posidonia, Zostera and algae, eulittoral to 80 m (Fauvel 1927; Holthe 1986; Gil 2011). Remarks. Our material corresponds closely to the recent description by Jirkov (2020), including the variation in the number of nephridial papillae and notopodia (specimens from UK and Black Sea). Two morphological differences can be perceived between the present description and the data by Arvanitidis & Koukouras (1995: Table 1, based on the original descriptions and French specimens from the “Collection des Polychètes d’Angers”). Indeed, our specimens, and those of Jirkov, do not show any eyespots (which may fade with time when stored in alcohol), and they also show “rugose” branchial tips, i.e. wrinkled (as in A. kuehlmanni Arvanitidis & Koukouras, 1995). This last character was not described by Jirkov (2020) and could be related to method of fixation. Genus Eupolymnia Verrill, 1900 Type-species: Terebella danielsseni Malmgren, 1866, by monotypy. Diagnosis. (after Hutchings et al. 2021b). Prostomium transverse, attached to dorsal surface of upper lip; basal part as a thick crest, eyespots usually present; distal part shelf-like. Buccal tentacles all uniformly cylindrical. Peristomium restricted to expanded lips; relatively short upper lip, hood-like, wider than long, distal margin rounded, frequently undulated; lower lip button-like, mid-ventrally, almost completely covered by lobes of SG I. Segment I conspicuous all around, dorsally narrow, with pair of low ventro-lateral lobes connected to each other by mid-ventral lobe marginal to mouth. Segments II–IV with pairs of progressively shorter and more laterally inserted lobes, those on SG II ventro-lateral and frequently connected to each other by low collar-like lobe across ventrum.Anterior segments highly glandular ventrally, with discrete rectangular shields, anterior shields frequently corrugated. Three pairs of branchiae, on SG II–IV, each one with a single short and thick main stem, dichotomously branching to short distal filaments. Conical to roughly rectangular notopodia from SG IV, extending 17 segments, until SG XX; notochaetae all narrowly-winged, wings slightly broader basally on one side and with smooth tips. Neuropodia beginning on SG V, as low, sessile ridges in conjunction with notopodia as conical to rectangular pinnules posteriorly; neurochaetae as short-handled avicular uncini, in completely intercalated double rows in face-to-face arrangement from SG XI until end of notopodia. Nephridial and genital papillae present, from SG II or III, extending for few anterior segments, between parapodial lobes or equivalent position on anterior segments. Pygidium crenulate to papillate. Remarks. Currently, only two valid species (but with long lists of synonymized species) of this genus occur in European waters, both without any extant type material: Eupolymnia nesidensis (Delle Chiaje, 1828) (described from Gulf of Naples, Italy) and Eupolymnia nebulosa (Montagu, 1819) (described from Cornwall, UK). The main problem with these two species is that in spite of being widely referred to in the taxonomic literature and used in dichotomous keys, we do not believe that there is a good recent morphological description of any of them as both of the original descriptions are brief. For example, following Holthe (1986), E. nebulosa would have “anterior branchiae with long stems” and a “buccal segment forming a broad cushion-like lip”, but this author only observed specimens from Norway. After examination of several specimens from near the type locality (Cornwall, UK), Hutchings & Glasby (1988) concluded that this species has “branchiae with virtually no main stalk” and “peristomium expanded as a discrete narrow elongated raised ventral collar”, though, knowing that there are other species present in the area (as shown in this study) it is possible that the studied material could also belong to a different species. Capa & Hutchings (2006) after observing two specimens sampled close from the type locality (AM W.200882 and BMNH ZK 1950.6.6.21) provided more details concerning this species: presence of 10 ventral shields, branchiae with no stalk and few branches, small spherical lateral lobes on SG II and IV, bilobed on SG III, dental formula of uncini MF:2:3 with dorsal button in the middle of uncini. This species has been widely reported because of its particular “strawberry” colour (e.g. Fauvel 1927) but at least two other species (and probably more) seem to exhibit this pattern of red colour with white spots (see below). Concerning E. nesidensis, Delle Chiaje (1828) briefly wrote that the species was yellow, with three pairs of red arborescent branchiae and with parapodia bearing chaetae. There is also a variety, differing from the former by its green colour, the yellowish-red branchiae and the thinner body (Delle Chiaje, 1828). Until a neotype from the Gulf of Naples is selected and described, it is recommended that this name should not be used., Published as part of Lavesque, Nicolas, Daffe, Guillemine, Londoño-Mesa, Mario H. & Hutchings, Pat, 2021, Revision of the French Terebellidae sensu stricto (Annelida, Terebelliformia), with descriptions of nine new species, pp. 1-63 in Zootaxa 5038 (1) on pages 16-19, DOI: 10.11646/zootaxa.5038.1.1, http://zenodo.org/record/5502867, {"references":["Grube, A. E. (1860) Beschreibung neuer oder wenig bekannter Anneliden. Beitrag: Zahlreiche Gattungen. Archiv fur Naturgeschichte, Berlin, 26, 71 - 118, pls. 1 - 3.","Saint-Joseph, A. (1894) Les Annelides polychetes des cotes de Dinard. Annales des sciences naturelles, zoologie, Paris, Serie 7, 17, 127 - 272.","Fauvel, P. (1927) Polychetes Sedentaires. Addenda aux Errantes, Archiannelides, Myzostomaires. Faune de France 16. Lechevalier, Paris, 494 pp.","Jirkov, I. A. (2020) Review of the European Amphitrite (Polychaeta: Terebellidae) with description of two new species. Invertebrate Zoology, 17 (4) 311 - 360. https: // doi. org / 10.15298 / invertzool. 17.4.01","Holthe, T. (1986) Polychaeta, Terebellomorpha. Marine Invertebrates of Scandinavia, 7, 1 - 194.","Arvanitidis, C. & Koukouras, A. (1995) Amphitritides kuehlmanni sp. nov. (Polychaeta, Terebellidae, Amphitritinae) from the Aegean Sea, with comments on the genus Amphitritides Augener. Ophelia, 40 (3), 219 - 227. https: // doi. org / 10.1080 / 00785326.1995.10430587","Quatrefages, A. de. (1866) Histoire naturelle des Anneles marins et d'eau douce. Annelides et gephyriens. Volume 2. Librairie Encyclopedique de Roret, Paris, 794 pp. https: // doi. org / 10.5962 / bhl. title. 122818","Keferstein, W. (1862) Untersuchungen uber niedere Seethiere. Zeitschrift fur wissenschaftliche Zoologie, 12 (1), 1 - 147.","Claparede, E. (1869) Les Annelides Chetopodes du Golfe de Naples. Seconde partie. Ordre IIme. Annelides Sedentaires (Aud. et Edw.). Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve, 20 (1), 1 - 225.","Gil, J. (2011) The European Fauna of Annelida Polychaeta. Ph. D. Dissertation, Departamento de Biologia Animal, Faculdade de Ciencias, Universidade de Lisboa, Lisboa, xlii + 1554 pp.","Verrill, A. E. (1900) Additions to the Turbellaria, Nemertina, and Annelida of the Bermudas, with a revision of the New England genera and species. Transactions of the Connecticut Academy of Arts and Sciences, 10, 595 - 671. https: // doi. org / 10.5962 / bhl. part. 7035","Malmgren, A. J. (1866) Nordiska Hafs-Annulater. Ofversigt af Kongiliga Veteskaps-Akademiens Forhandlingar, 22, 355 - 410.","Hutchings, P., Nogueira, J. M. N. & Carrerette, O. (2021 b) Terebellidae Johnston, 1846. In: Schmidt-Rhaesa, A. Hr., Beutel, R. G., Glaubrecht, M., Kristensen, N. P., Prendini, L., Purschke, G., Richter, S., Westheide, W. & Leschen, R. Z. E. (Ed.), Handbook of Zoology. A Natural History of the Phyla of the Animal Kingdom. Walter de Gruyter & Co, Berlin, pp. 1 - 64.","Delle Chiaje, S. (1828) Memorie sulla storia e notomia degli animali senza vertebre del Regno di Napoli, 3, 1 - 232 pp. https: // doi. org / 10.5962 / bhl. title. 10021","Montagu, G. (1819) Descriptions of five British species of the genus Terebella. Transactions of the Linnean Society of London, 12 (2), 340 - 344. https: // doi. org / 10.1111 / j. 1095 - 8339.1817. tb 00231. x","Hutchings, P & Glasby, C. J. (1988) The Amphitritinae (Polychaeta: Terebellidae) from Australia. Records of the Australian Museum, 40 (1), 1 - 60. https: // doi. org / 10.3853 / j. 0067 - 1975.40.1988.150","Capa M. & Hutchings P. (2006) Terebellidae (Polychaeta) from Coiba National Park, Panamanian Pacific, including description of four new species and synonymy of the genus Paraeupolymnia with Lanicola. Zootaxa, 1375, 1 - 29. https: // doi. org / 10.11646 / zootaxa. 1375.1.1"]}

Pista labruneae n. sp. Figure 18 Material examined. Holotype. MNHN-IA-TYPE 2031, posteriorly incomplete, Mediterranean Sea, Gulf of Lion, Redit F 20, 42°46’44”N 3°3’21”E, depth 20 m, August 2010. Paratypes. AM W.53329, posteriorly incomplete, Mediterranean Sea, Gulf of Lion, Redit F 20, 42°29’20”N 3°8’19”E, depth 17 m, October 2020. MNHN-IA-TYPE 2032, posteriorly incomplete, Mediterranean Sea, Gulf of Lion, Beauduc, 43°24’47”N 4°30’7”E, depth 14 m, 2012. Description. Holotype posteriorly incomplete, 22.3 mm long (5.8 mm) and 1.5 mm wide (0.5 mm), for 29 segments. Transverse prostomium attached to dorsal surface of upper lip; basal part without eyespots; distal part of prostomium convoluted. Buccal tentacles filiform and deeply grooved (Fig. 18B–F). Peristomium forming lips, upper lip hood-like, large, wider than long, rectangular, with convoluted margin, slightly pointed anteriorly; lower lip swollen, crescent-shaped, wider than long (Fig. 18B–C). Segment I conspicuous, with one pair of rounded lateral lobes, connected to each other by a thin and smooth membrane with rounded ventral indentation, surrounding lower lip (Fig. 18B–C, E). Segment II with one pair of large ventro-lateral lobes, trapezoidal, with long ventral base and rounded margins, connected to each other by large mid-ventral crenulated crest. Segment III with one pair of auricular dorso-lateral lobes, much shorter than those of SG II, connected to each other by thin, slightly crenulated mid-ventral crest. Segment IV without lateral lobes. Dorsal anterior margins of SG III as protruding crest (Fig. 18D). A single plume-shaped branchia present on SG II inserted mid-dorsally; with conspicuous, annulated basal stem, branches arranged in spiral around the main stem, with short filaments (Fig. 18A, C–E). Smooth mid-ventral shields present on SG V–XIX, rectangular shields, becoming progressively longer and wider posteriorly. Notopodia beginning on SG IV, extending until SG XX; notopodia short, rectangular, first four pairs shorter, inserted progressively more laterally, then longitudinally aligned (Fig. 18C–D). Broadly-winged notochaetae in two rows, broader on one side (Fig. 18G), with first row shorter. Neuropodia present from SG V, as low, almost sessile ridges until end of notopodia, as low rectangular pinnules thereafter (Fig. 18 C–D). Neurochaetae as long-handled uncini on SG V–X, with well-developed handles originating from heel, progressively thinner; uncini arranged in partially intercalated double rows on SG XI–XX in a faceto-face arrangement. Avicular uncini with slightly convex base, large rounded heel, distally rounded prow, dorsal button inserted halfway between base of main fang and prow, and well-developed pointed main fang, surmounted by crest with 5 rows of numerous and progressively shorter secondary teeth above the main fang (Fig. 18H–I). Genital papillae on SG VI–VII, tubular, situated dorsally behind notopodia. Pygidium unknown. Etymology. This species is dedicated to Céline Labrune, from the Banyuls-sur-Mer Arago Laboratory, for her friendship with NL for about 15 years and for providing the type material of eight new species since the beginning of the “ Spaghetti Project ”, including this new one. Habitat. Fine sand, depth 17–20 m. Type locality. Gulf of Lion, Western Mediterranean Sea, Gulf of Lion. 42°46’44”N 3°3’21”E. Distribution. Only known from the type locality, but probably present in Italy and other parts of the Mediterranean Sea (see below). Remarks. Among the European species, P. labruneae n. sp. is easily recognizable by the presence of a single branchia situated mid dorsally on SG II. The type material was carefully checked to ensure that only a single branchia was ever present, and that another one had been lost but no evidence of a branchial scar was ever found, so we are confident that only a single branchia was present. Moreover, specimens with only a single branchia are regularly found in this area (Labrune pers. com.) confirming that only a single branchia is present. Pista labruneae n. sp. differs from P. colini recently described from the same area (Labrune et al. 2019). First, P. colini has one pair of branchiae instead of a single mid-dorsal branchia. The two species differ also by the number and shape of lateral lobes. The lateral lobes on SG II of P. labruneae n. sp. are larger than those of P. colini. Finally, P. colini has neurochaetae as long-handled avicular uncini on SG V–VI, short-handled thereafter while P. labruneae n. sp. has long-handled uncini at least until SG X. Pista labruneae n. sp. has probably been previously confused and misidentified in the Mediterranean Sea as P. unibranchia Day, 1963, a species from South Africa also described as having a single branchia. However, as underlined recently by Langeneck et al. (2020), the presence of this last species as a non-indigenous species is doubtful. Moreover, P. unibranchia is characterised by the absence of long-handled uncini on SGV (Day 1963) and should be transferred to the genus Pistella Hartmann-Schröder, 1996. In addition to the different shape of uncini, Pista labruneae n. sp. differs from P. unibranchia by the absence of lateral lobes on SG IV.

Polycirrus Grube, 1850 Type-species. Polycirrus medusa Grube, 1850, by monotypy., Published as part of Lavesque, Nicolas, Hutchings, Pat, Daffe, Guillemine & Londoño-Mesa, Mario H., 2020, Revision of the French Polycirridae (Annelida, Terebelliformia), with descriptions of eight new species, pp. 151-186 in Zootaxa 4869 (2) on page 158, DOI: 10.11646/zootaxa.4869.2.1, http://zenodo.org/record/4418481, {"references":["Grube, A. E. (1850) Die Familien der Anneliden. Archiv fur Naturgeschichte Berlin, 1691, 249 - 364."]}

Polycirrus catalanensis n. sp. Figures 4–5, Table 2. Material examined: Holotype: MNHN-IA-TYPE 2007, one complete specimen, Mediterranean Sea, Gulf of Lion, Reserve St Troc, 42°29’01”N, 3°08’48”E, 31 m depth, 2017. Paratypes: AM W.53112, one complete specimen, Mediterranean Sea, Gulf of Lion, Reserve St Troc, 42°29’01”N, 3°08’48”E, 31 m depth, 2017. MNHN-IA-TYPE 2008, two complete specimens, Mediterranean Sea, Gulf of Lion, Reserve St Troc, 42°29’01”N, 3°08’48”E, 31 m depth, 2017. AM W.53113, one complete specimen, Mediterranean Sea, Gulf of Lion, Reserve St Troc, 42°29’01”N, 3°08’48”E, 31 m depth, 2017, mounted for SEM. Description. Small species, holotype 11.5 (6.2–7.8) mm long and 0.5 (0.3–0.7) mm wide. Transverse prostomium attached to dorsal surface of base of upper lip; basal part as thick crest across dorsum, extending laterally and dorsally, covering SG1 laterally and terminating lateral to lower lip. Buccal tentacles lost. Peristomium forming lips; upper lip, longer than wide, not elongated, with a single median and very thick convoluted lobe only (Figs 4 A–D; 5A); lower lip rectangular with rounded corners, longer than wide, very glandular and ridged (Figs 4 A–D; 5A). SG1 and 2 reduced, SG1 visible dorsally, SG2 visible dorsally and laterally (Fig. 4C); body slightly broader until SG6, mid-body inflated, posterior part tapering (Fig. 4 A–B, D). Ventro-lateral inflated pads present from SG3 to SG10, well-defined (Figs 4 A–D; 5A). Anterior mid-ventral groove present from SG3 (Figs 4D, 5A). Notopodia from SG3, extending for 15 (13) segments, until SG17 (SG15); distinctly elongate, rectangular, first two pairs slightly shorter, bilobed, postchaetal lobe digitiform, larger than prechaetal one (Fig. 5 A–B). Narrowly winged notochaetae in two rows, distal ends whip-like (Fig. 5 B–C), with wings inconspicuous under light microscopy (Fig. 4E). Neuropodia from SG17 (SG15–16); uncini with short occipitum and slightly convex base (Type 1) (Fig. 4F), crest with single row of secondary teeth, with a sharp elongate central tooth almost reaching tip of main fang, and two small lateral teeth on each side (Fig. 5D), subrostral process present as low protuberance (Fig. 4F). Nephridial and genital papillae not seen. Pygidium rounded with pointed tip (Fig. 4D). Etymology. Species name refers to the Catalan Sea (north-western Mediterranean), where the species was found. Habitat. Coastal heterogeneous sands, 30 m depth. Type locality. Banyuls-Cerbere natural reserve, Catalan Sea, Mediterranean Sea, France. Distribution. Only known from type locality. Remarks. Polycirrus catalanensis n. sp. is characterized by the presence of notopodia extending over 13–15 segments which is similar to P. denticulatus (12–13 segments), P. pennarbedae n. sp. (12–13 segments) and P. norvegicus (14–20 segments) (Table 2). Among these species, only P. pennarbedae n. sp. shows almost the same range in the first appearance of neuropodia (from SG15–16 for P. pennarbedae n. sp., from SG15–17 for P. catalanensis n. sp.) (Table 2). However, P. catalanensis n. sp. differs from P. pennarbedae n. sp. by the shape of uncini (two rows of teeth above the main tooth and the absence of subrostral process for P. pennarbedae n. sp., a single row of secondary teeth above the main tooth and the presence of subrostral process for P. catalanensis n. sp.), the shape of upper lip (elongated triangular medial lobe for P. pennarbedae n. sp., very thick single medial lobe for P. catalanensis n. sp.), the shape of lower lip (oval and wider than long for P. pennarbedae n. sp., rectangular and longer than wide P. catalanensis n. sp.) and the presence of a large mid-ventral grove on anterior segments for P. catalanensis n. sp. (not visible for P. pennarbedae n. sp.) (Table 2). Polycirrus catalanensis n. sp. differs from P. denticulatus by the first appearance of neuropodia (from SG15–17 for P. catalanensis n. sp., from S12 for P. denticulatus), by he shape of lower lip (rectangular for P. catalanensis n. sp., subtriangular, pointing toward mouth for P. denticulatus), the shape of notochaetae (with wings inconspicuous for P. catalanensis n. sp., subdistally expanded for P. denticulatus following Glasby & Hutchings, 2014) and dental formula of uncini (MF:3 for P. catalanensis n. sp., MF:1: α for P. denticulatus). Another species occurring in the French Mediterranean Sea, P. medusa, differs from P. catalanensis n. sp. by the presence of notopodia on 12 segments (instead of 13–15 for P. catalanensis n. sp.), by the type of notochaetae (absence of pinnate chaetae for P. catalensis n. sp., but present for P. medusa), by uncini with complete transverse series of secondary teeth above main tooth (instead of a crest with a very long tooth and two small lateral teeth for P. catalanensis n. sp.), by the presence of trefoiled upper lip with thin margin (instead of upper lip with single medial lobe with thick margin for P. catalanensis n. sp.) and by the presence of a subtriangular lower lip, pointing towards the mouth (instead of a rectangular one for P. catalanensis n. sp.) (Table 2)., Published as part of Lavesque, Nicolas, Hutchings, Pat, Daffe, Guillemine & Londoño-Mesa, Mario H., 2020, Revision of the French Polycirridae (Annelida, Terebelliformia), with descriptions of eight new species, pp. 151-186 in Zootaxa 4869 (2) on pages 158-160, DOI: 10.11646/zootaxa.4869.2.1, http://zenodo.org/record/4418481, {"references":["Glasby, C. J. & Hutchings, P. (2014) Revision of the taxonomy of Polycirrus Grube, 1850 (Annelida: Terebellida: Polycirridae). Zootaxa, 3877 (1), 1 - 117. https: // doi. org / 10.11646 / zootaxa. 3877.1.1"]}

Amaeana gremarei n. sp. Figures 2–3 Material examined: Holotype: MNHN-IA-TYPE 2006, incomplete, Mediterranean Sea, Gulf of Lion, 42°31’36”N, 3°23”47’E, 110 m depth, June 2019, posterior parapodia mounted for SEM. Paratypes: AM W.53110, incomplete, Mediterranean Sea, Gulf of Lion, A90, 42°32’39”N, 3°16’03”E, 90 m depth, April 2018. AM W.53111, incomplete, Mediterranean Sea, Gulf of Lion, Reserve St Troc, 42°29’01”N, 3°08’48”E, 31 m depth, 2017, mounted for SEM. Description. Large species, holotype incomplete, 16.4 (8.0–22.2) mm long and 2.2 (2.0–2.8) mm wide. Prostomium at base of upper lip (Fig. 2D), both basal and distal parts developed, basal part as thickened crest (Fig. 2C), distal part with large lateral lobes and also oval mid-dorsal process (Fig. 2C); prostomium covering SG1 laterally and terminating laterally to lower lip, near mouth (Fig. 2D). Only few buccal tentacles remaining, short ones thin, uniformly cylindrical, intermediate ones (only on paratype AM W.53110) distally broader, spatulate (Fig. 2 C–D). Peristomium restricted to lips, upper lip almost circular, wider than long, single lobe only (Figs 2 C–D; 3A); lower lip large, almost rounded, as long as wide (Figs 2 C–D; 3A). Body progressively broader until SG7–8, then gradually tapering to narrower uniformly cylindrical posterior body, beginning from SG13–14 (Fig. 2A); achaetous gap between termination of notopodia and beginning of neuropodia, corresponding to SG13–16, with poorly marked segmentation, longer than region with notopodia (Fig. 2A). Segments biannulated, SG1 short, visible dorsally and ventrally, laterally covered by expanded prostomium (Figs 2D; 3A); SG 2 narrower and shorter than following segments, with large pentagonal to hexagonal mid-ventral shield at beginning of mid-ventral groove (Figs 2D; 3A), extending anteriorly through segment 1 until near ventral edge of lower lip. Ventrum highly glandular, covered by small papillae on SG2–11, less conspicuous on S13, then smooth body wall, with paired longitudinal crests bordering mid-ventral groove through posterior body (Figs 2 C–D; 3A). Ventro-lateral pads on SG3–13; with transverse ridges but without longitudinal groove (Figs 2D; 3A). Notopodia from SG3, extending through 10 segments, until segment 12 (Fig. 2A); distinctly elongate, rectangular with rounded distal part, with equal lobes (Fig. 3C). Notochaetae in both rows with narrow limbs (light microscopy), with fine hairs under SEM, emerging from the ventral face of notopodia to distal part (Fig. 3 C–D). Neuropodia present from SG17 (hard to see exactly), laterally to mid-ventral groove, on outer margins of longitudinal crests (Fig. 2A). Neurochaetae up to 8–9 thin, long spines, with all tips broken (Fig. 3B). Nephridial and genital papillae at anterior bases of first nine notopodia (Fig. 2B). Pygidium unknown. Etymology. This species is dedicated to Antoine Grémare for his many contributions to benthic ecology and his important support to NL and Céline Labrune, who sent us type material. Habitat. Coastal heterogeneous sands, 30–100 m depth. Type locality. Gulf of Lion, Mediterranean Sea, France. Distribution. Only known from type locality. Remarks. Among the valid species of this genus, a single species is known from European waters: A. trilobata. Amaeana gremarei n. sp. is similar to this species by both having 10 pairs of notopodia. However, Amaeana gremarei n. sp. differs in having a circular upper lip without distinct lobes (instead of upper lip with three lobes for A. trilobata), by a rounded lower lip, wider than long (instead of rectangular broad one as found in A. trilobata), by nephridial papillae on SG3–11 (instead of SG3–12 for A. trilobata), by a long achaetous region (instead of a short one as occurs in A. trilobata) and by ventro-lateral pads on SG3–13 (instead of on SG3–12 for A. trilobata)., Published as part of Lavesque, Nicolas, Hutchings, Pat, Daffe, Guillemine & Londoño-Mesa, Mario H., 2020, Revision of the French Polycirridae (Annelida, Terebelliformia), with descriptions of eight new species, pp. 151-186 in Zootaxa 4869 (2) on pages 155-158, DOI: 10.11646/zootaxa.4869.2.1, http://zenodo.org/record/4418481

Polycirrus readi n. sp. Figures 18–19, Table 2. Material examined: Holotype: MNHN-IA-TYPE 2019, one complete specimen, Mediterranean Sea, Corsica Cape, 42°44’23”N, 9°28’41”E, 60 m depth, May 2019. Paratypes: AM W.53126, one complete specimen, Mediterranean Sea, Corsica Cape, 43°01’18”N, 9°24’30”E, 18 m depth, May 2019, posterior part used for molecular analysis, mounted for SEM. Additional material: MNHN-IA- PNT 124, one complete specimen, Mediterranean Sea, Calvi, 42°34’48”N, 8°43’45”E, 40 m depth, August 2011. Description. Small specimens except MNHN-IA- PNT 124, holotype 8.3 (4.1–14.5) mm long and 0.6 (0.4–0.9) mm wide. Transverse prostomium attached to dorsal surface of base of upper lip; basal part as thick crest across dorsum, extending laterally and dorsally, covering SG1 laterally and terminating lateral to lower lip. Buccal tentacles of two types, long and thin tentacles uniformly cylindrical, long and thick ones deeply grooved (Figs 18 A–D; 19A). Peristomium forming lips; upper lip elongate, longer than wide, with single median lobe only, triangular, convoluted (Figs 18B, D; 19A); lower lip oblong, longer than wide, very glandular, ridged (Figs 18B, D; 19A). SG1 and 2 reduced, SG1 visible dorsally, SG2 dorsally and laterally; body slightly broader until SG6, then tapering until SG10, then of relatively uniform width (larger than anterior part) until mid-body, posterior end tapering (Fig. 18A). Ventro-lateral inflated pads well-defined from SG3 to SG9 (SG10), smooth, with transverse ridges on largest specimen, subsequent ones less conspicuous until SG12 (SG15) (Figs 18 A–D; 19A). Large anterior midventral groove from S3, present until end of body as a stripe (Figs 18B, D; 19A). Notopodia from SG3, extending for 17 (13) segments, until SG19 (SG15); distinctly elongate, rectangular, first pair slightly shorter, bilobed, postchaetal lobe conical with rounded tip, larger than prechaetal one (Fig. 19B). Winged notochaetae of two different lengths (Figs 18E; 19 B–C). Neuropodia beginning from SG10 (SG9); uncini with short occipitum and slightly convex base (Type 1), crest with single elongate and sharp tooth on first row above main fang, with one additional rows of 4–5 short, irregularly sized teeth (Fig. 19D), subrostral process present as a protuberance (Fig. 18F). Nephridial and genital papillae not seen. Pygidium rounded with pointed tip. ...Continued on the next page Etymology. This species is dedicated to Dr. Geoff Read for his outstanding contribution to WoRMS website and for sharing his immense knowledge on the annelida.net forum. Habitat. 16–60 m depth, among rocks and dead leaves of Posidonia sp. Type locality. Corsica Cape, Mediterranean Sea. Distribution. Known from type locality only. Remarks. Polycirrys readi n. sp. is characterized by the presence of notopodia extending on 13–17 segments and neuropodia from SG9–10, which is similar to P. norvegicus (notopodia on 14–20 segments neuropodia from SG10–12) and P. elisabethae (notopodia on 16 segments and neuropodia from SG11) (Table 2). However, P. readi n. sp. differs from P. norvegicus by the presence of a mid-ventral groove from S3 (instead of S4 for P. norvegicus), of notopodia with conical postchaetal lobe (instead of digitiform lobe for P. norvegicus) and of uncini with a main tooth always present and secondary row of teeth above it (instead of teeth arranged in a single transverse row, with or without main tooth for P. norvegicus). Finally, the two species are described from very different geographical areas (North Sea for P. norvegicus, Mediterranean Sea for P. readi n. sp.). Polycirrus readi n. sp. differs from P. elisabethae by the shape of lower lip (oblong and longer than wide for P. readi n. sp., subtriangular and wider than long for P. elisabethae), by the shape of ventro-lateral pads (very inflated for P. readi n. sp., discrete for P. elisabethae), by the appearance of the mid-ventral groove (from S3 for P. readi n. sp., from S 4 P. elisabethae) and by the dental formula of uncini (secondary row with 4–5 teeth for P. readi n. sp., with 11–15 teeth for P. elisabethae).

Polycirrus idex n. sp. Figure 12–13, Table 2. Material examined: Holotype: MNHN-IA-TYPE 2015, one complete specimen, Mediterranean Sea, Corsica Cape, 42°44’42”N, 9°28’00”E, 16 m depth, May 2019, posterior part used for molecular analysis. Paratype: AM W.53127, one complete specimen, Mediterranean Sea, Corsica Cape, 43°01’18”N, 9°24’30”E, 18 m depth, May 2019, mounted for SEM, posterior part used for molecular analysis. Description. Small specimen 4.6 (4.3) mm long and 0.6 (0.7) mm wide. Transverse prostomium attached to dorsal surface of base of upper lip; basal part as thick crest across dorsum, extending laterally and dorsally, covering SG1 laterally and terminating lateral to lower lip; distal part extending along upper lip, terminating subdistally. Buccal tentacles of two types, long grooved annulated and uniformly cylindrical ones, short thick grooved and distally enlarged ones (Figs 12 A–B; 13A). Peristomium forming lips; upper lip elongate, longer than wide, with single median lobe only (Figs 12 B–C; 13A); lower lip oblong, slightly longer than wide, very glandular, ridged (Figs 12C; 13A). SG1 and 2 reduced, SG2 visible ventrally (Figs 12C; 13A); body slightly broader until SG7. Ventro-lateral inflated pads well-defined from SG3 to SG9 (SG8), less defined on SG10 (SG9), smooth (Figs 12C; 13A). Large anterior mid-ventral groove from S3, present until end of body as a stripe (Figs 12C; 13A). Notopodia from SG3, extending for 13 (14) segments, until SG15 (SG16); not very elongate, rectangular, first pair slightly shorter, bilobed, postchaetal lobe conical with rounded tip, larger than prechaetal one (Fig. 18B). Winged notochaetae in two rows (Figs 12D; 13 B–C). Neuropodia beginning from SG8 (SG7); uncini with short occipitum and slightly convex base (Type 1), crest with single elongate and sharp tooth on first row above main fang, with one additional row of 4–5 short, irregularly sized teeth (Fig. 13D), subrostral process present as small tooth (Fig. 13D). Nephridial and genital papillae not seen. Pygidium rounded. Etymology. This species is named after IDEX (Excellence Initiative of Bordeaux University), which funded two grants for NL and GD to spend 9 months at the Australian Museum (Sydney). “Idex” is an unmodifiable noun in apposition. Habitat. 18 m depth, among rocks. Type locality. Corsica Cape, Mediterranean Sea. Distribution. Only known from type locality. Remarks. With 14 segments with notopodia and neuropodia starting on SG7–8, P. idex n. sp. differs from all other European species (Table 2). The closest species is P. asturiensis (notopodia on 10 segments and neuropodia starting on SG6) but this last species has a trefoiled upper lip (instead of a single lobe only for P. idex n. sp.), a lower lip wider than long (instead of longer than wide for P. idex n. sp.) and uncini showing two rows of secondary teeth above the main tooth (instead of a single row of secondary teeth for P. idex n. sp.).

Polycirrus nogueirai n. sp. Figures 14–15, Table 2. Material examined: Holotype: MNHN-IA-TYPE 2016, incomplete, NE Atlantic, Bay of Biscay, Arcachon Bay, Jacquets, 44°43’21”N, 1°11’15”W, intertidal, December 2018. Paratypes: AM W.53121, one incomplete specimen, NE Atlantic, Bay of Biscay, Arcachon Bay, Jacquets, 44°43’21”N, 1°11’15”W, intertidal, December 2018, mounted for SEM. MNHN-IA-TYPE 2017, one incomplete specimen, NE Atlantic, Bay of Biscay, Arcachon Bay, Jacquets, 44°43’21”N, 1°11’15”W, intertidal, December 2018. AM W.53122, one incomplete specimen, NE Atlantic, Bay of Biscay, Arcachon Bay, Jacquets, 44°43’21”N, 1°11’15”W, intertidal, December 2018. Description. Small species, holotype incomplete, 9.1 (4.5–11.0) mm long and 0.9 (0.6–0.8) mm wide. Transverse prostomium attached to dorsal surface of base of upper lip; basal part as thick crest across dorsum, extending laterally, covering SG1 laterally and terminating lateral to lower lip. Buccal tentacles of two types, long and thin tentacles uniformly cylindrical, long and thick ones deeply grooved, with pointed tips (Fig. 14 A–C). Peristomium forming lips; upper lip elongate, longer than wide, with a single median lobe only (Figs 14B, D; 15A); lower lip rounded and inflated, as wide as long, very glandular and ridged (Figs 14B, D; 15A). SG1 and 2 reduced, SG1 visible dorsally, SG2 dorsally, laterally and ventrally (Fig. 15A); body with relatively uniform width until SG20, gently tapering after (Fig. 14A, C). Ventro-lateral inflated pads present from SG3 to SG13 (SG14), well-defined, smooth and transversely ridged until SG10, subsequent ones less conspicuous (Figs 14 B–D; 15A). Midventral groove with small discrete shields from SG3 to SG10, continuing as a stripe until end of body (Figs 14B, D; 15A). Notopodia from SG3, extending for 16 (19–21) segments, until SG18 (SG21–23); distinctly elongate, rectangular, first pair slightly shorter, bilobed, postchaetal lobe digitiform, larger than prechaetal one (Fig. 15B). Winged notochaetae of two lengths, distal ends whip-like (Figs 14E; 15 B–C). Neuropodia beginning from SG14 (SG16); uncini with short occipitum and slightly convex base (Type 1) (Fig. 14F), crest with single elongate and sharp tooth on first row above main fang, with two additional rows of shorter, irregularly sized teeth at base (Fig. 15D), absence of subrostral process (Fig. 14F). Nephridial and genital papillae present on segments 3–8. Pygidium unknown. Etymology. This species is dedicated to João Nogueira for his great contribution to taxonomy of terebellids. Habitat. Intertidal, muddy sands, sheltered waters. Type locality. Arcachon Bay, Bay of Biscay, NE Atlantic Ocean, France. Distribution. Only known from type locality. Remarks. Polycirrus nogueirai n. sp. is characterized by the presence of notopodia on 16–21 segments, which is similar to P. glasbyi n. sp. (18–22 segments), P. plumosus (17–19 segments), P. readi n. sp. (13–17 segments), P. elisabethae (16 segments) and P. norvegicus (14–20 segments) (Table 2). Among these species, only P. plumosus shows almost the same range in appearance of neuropodia (from SG18–22 for P. plumosus, from SG14–16 for P. nogueirai n. sp.). However, P. nogueirai n. sp. differs from P. plumosus by the presence of one type of notochaetae only (instead of two for P. plumosus), uncini with a median tooth in first row above main fang (absent for P. plumosus), the shape of lower lip (protruding above venter and longer than wide for P. plumosus, rounded and as wide as long for P. nogueirai n. sp.) and by the shape of ventro-lateral pads (poorly defined for P. plumosus, inflated for P. nogueirai n. sp.).

Polycirrus cf. denticulatus Saint-Joseph, 1894 Figures 6–7, Table 2. Material examined: AM W.53114, incomplete, NE Atlantic, Bay of Biscay, Bay of Brest, 48°21’28”N, 4°26’38”W, 7 m depth, May 2018, mounted for SEM, some parapodia used for molecular analysis. AM W.53115, incomplete, NE Atlantic, Bay of Biscay, Bay of Brest, 48°21’28”N, 4°26’38”W, 7 m depth, May 2018, mounted for SEM, some parapodia used for molecular analysis. MNHN-IA-PNT 121, incomplete, English Channel, Morlaix Bay, 48°41’04”N, 3°54’58”W, intertidal, February 2016. SMA_ BR _ Poly _17, incomplete, English Channel, 48°35’50”N, 4°37’27”W, intertidal, February 2015. SMA_ BR _ Poly _14, incomplete, English Channel, 48°35’50”N, 4°37’27”W, intertidal, February 2015. Description. Transverse prostomium attached to dorsal surface of base of upper lip; basal part as thick semicircular crest across dorsum, extending lateral and posteriorly (Fig. 6B), covering SG1 laterally and terminating lateral to lower lip. Distal part extending along lip, terminating subdistally. Few buccal tentacles remaining, short and thick, cylindrical (Figs 6A; 7A). Peristomium forming lips; upper lip comprising single medial lobe only, margin of lobe thickened and overturned (Figs 6 A–C; 7A). Outer lower lip shield-like, subtriangular and pointing toward mouth, longer than wide, ridged and grooved, extending posteriorly to segment 3 (Figs 6A; 7A). SG1 and 2 reduced, SG1 visible dorsally and laterally, SG2 visible dorsally, laterally and ventrally (Fig. 6A); body slightly broader until SG5, gently tapering until SG10, then of relatively uniform width, cylindrical (Fig. 6B). Ventro-lateral inflated pads well-defined from SG3 to SG10, smooth and transversely ridged, subsequent ones less conspicuous until mid-body (Figs 6A, C; 7A). Deep mid-ventral groove from SG3 (Figs 6A; 7A). Notopodia from SG3, extending for 12–13 segments, until SG14–15. Notopodia more-or-less rectangular, lobes slightly triangular (distally rounded) (Fig. 7 B–C). Narrowly-winged notochaetae of two distinct lengths, slightly expanded subdistally, with fine tips (Figs 6 D–E; 7B–C). Neuropodia beginning from SG12; type of uncini not seen (specimen used for SEM), crest with single elongate and sharp tooth on first row above main fang, with at least one additional row of shorter, irregularly sized teeth at base (exact number of additional rows not visible) (Fig. 7D). Nephridial and genital papillae present on segments 3–8, ventral to bases of notopodia (Fig. 6C). Pygidium unknown. Habitat. Shallow waters, in maerl (rhodolith) beds, seagrass beds (this study) and among laminarians, algae, serpulids, and hydrozoa (Gil 2011). Type locality. Brittany, France (exact locality problematic). Distribution. NE Atlantic, English Channel, Mediterranean Sea (Fauvel 1927; Gil 2011), Adriatic Sea (Mikac 2015). Remarks. French specimens, sampled close to the potential type locality, match almost completely with the recent redescription of type material by Glasby & Hutchings (2014). However, few minor variations were observed: mid-ventral groove appearing on segment 3 (instead of segment 4), notochaetigerous segments 12–13 (instead of 10, but type material is broken). Finally, we also document the first segment on which neuropodia begin. As commented by Glasby & Hutchings (2014), doubts exist about the location of the type material. Indeed, the holotype seems to have been sampled in Concarneau (Brittany, Bay of Biscay) while in the original description Mr le Baron de St. Joseph (1894) reported the type locality in Dinard (Brittany, English Channel). Moreover, the redescription of the holotype does not fit exactly the type description (Glasby & Hutchings 2014). Consequently, until a neotype is collected and described, we prefer to identify these specimens as Polycirrus cf. denticulatus., Published as part of Lavesque, Nicolas, Hutchings, Pat, Daffe, Guillemine & Londoño-Mesa, Mario H., 2020, Revision of the French Polycirridae (Annelida, Terebelliformia), with descriptions of eight new species, pp. 151-186 in Zootaxa 4869 (2) on page 161, DOI: 10.11646/zootaxa.4869.2.1, http://zenodo.org/record/4418481, {"references":["Saint-Joseph, A. (1894) Annelides Polychetes des cotes de Dinard. Troisieme Partie. Annales des Sciences naturelles Zoologie et Paleontologie, 17, 1 - 395.","Gil, J. (2011) The European Fauna of Annelida Polychaeta. Ph. D. Dissertation, Departamento de Biologia Animal, Faculdade de Ciencias, Universidade de Lisboa, Lisboa, xlii + 1554 pp.","Fauvel, P. (1927) Polychetes Sedentaires. Addenda aux Errantes, Archiannelides, Myzostomaires. Faune de France 16. Lechevalier, Paris, 494 pp.","Mikac, B. (2015) A sea of worms: polychaete checklist of the Adriatic Sea. Zootaxa, 3943 (1), 1 - 172. https: // doi. org / 10.11646 / zootaxa. 3943.1.1","Glasby, C. J. & Hutchings, P. (2014) Revision of the taxonomy of Polycirrus Grube, 1850 (Annelida: Terebellida: Polycirridae). Zootaxa, 3877 (1), 1 - 117. https: // doi. org / 10.11646 / zootaxa. 3877.1.1"]}

Polycirrus gujanensis n. sp. Figures 10–11, Table 2. Material examined: Holotype: MNHN-IA-TYPE 2013, incomplete, NE Atlantic, Bay of Biscay, Arcachon Bay, Matelle, 44°39’51”N, 1°06’51”W, intertidal, May 2010, some parapodia mounted for SEM. Paratypes: MNHN-IA-TYPE 2014, one specimen, incomplete, NE Atlantic, Bay of Biscay, Arcachon Bay, Matelle, 44°39’51”N, 1°06’51”W, intertidal, May 2010. AM W.53119, one specimen, incomplete, NE Atlantic, Bay of Biscay, Arcachon Bay, Dispute, 44°42’17”N, 1°06’48”W, intertidal, August 2016. AM W.53120, one specimen, incomplete, NE Atlantic, Bay of Biscay, Arcachon Bay, Matelle, 44°39’51”N, 1°06’51”W, intertidal, May 2010, mounted for SEM. Description. Large species, holotype incomplete (41 segments), 23.2 mm long and 2.2 mm wide. Transverse prostomium attached to dorsal surface of base of upper lip; basal part as thick semicircular crest across dorsum, extending lateral and posteriorly (Fig. 10D), covering SG1 laterally and terminating lateral to lower lip. Buccal tentacles of two types, short tentacles uniformly cylindrical, long ones deeply grooved, with pointed tips (Fig. 10 A–D). Peristomium forming lips; upper lip elongate, longer than wide, highly convoluted, folded into three lobes, lateral ones well-developed (Figs 10B; 11A); lower lip oval and inflated, wider then long, very glandular and ridged (Figs 10B; 11A). SG1 and 2 reduced, only visible dorsally (Fig. 10 C–D); body slightly broader until SG8, gently tapering until SG10, then of relatively uniform width, cylindrical, tapering posteriorly (Fig. 10A). Ventro-lateral inflated pads well-defined from SG3 to SG12, smooth and transversely ridged, subsequent ones less conspicuous until mid-body (Figs 10 A–C; 11A). Deep midventral groove from SG3 to end of the body, with small discrete shields from SG3 to midbody (Figs 10B; 11A). Notopodia from SG3, extending for 28 segments, until SG30; distinctly elongate, rectangular, first two pairs slightly shorter, bilobed, postchaetal lobe larger than prechaetal one (Fig. 11B). Narrowly-winged notochaetae in two rows, with wings inconspicuous under light microscopy (Figs 10E; 11 B–C). Neuropodia beginning from SG15; uncini with short occipitum and slightly convex base (Type 1) (Fig. 10F), crest with single elongate and sharp tooth on first row above main fang, with two additional rows of shorter, irregularly sized teeth at base (Fig. 11D), subrostral process present as small tooth (Fig. 10F). Nephridial and genital papillae present on segments 3–8, anterior and ventral to bases of notopodia. Pygidium unknown. Etymology. The species name refers to the city of Gujan-Mestras, off where the species was sampled. Habitat. Intertidal, medium sands, sheltered waters. Type locality. Arcachon Bay, Bay of Biscay, NE Atlantic Ocean, France. Distribution. Only known type locality. Remarks. Polycirrus gujanensis n. sp. is characterized by the presence of 28 pairs of notopodia, and neuropodia from SG15, which is different from all European species, excepting P. arenivorus (29 pairs of notopodia, neuropdia from SG12) (Table 2). However, P. gujanensis n. sp. differs from this latter species by the shape of lower lip (oval and wider than long for P. gujanensis n. sp., oblong and longer than wide for P. arenivorus), by the upper lip with three well-developed lobes (instead of medial lobe well developed only in P. arenivorus) and by uncini with an elongate tooth above the main fang (this tooth absent in P. arenivorus).

Polycirrus pennarbedae n. sp. Figures 16–17, Table 2. Material examined: Holotype: MNHN-IA-TYPE 2018, one complete specimen, NE Atlantic, Bay of Biscay, Glenan, 47°44’06”N, 4°0’01”W, 7 m depth, March 2015. Paratypes: AM W.53123, one complete specimen, NE Atlantic, Bay of Biscay, Glenan, 47°43’43”N, 3°58’04”W, 5 m depth, March 2015. AM W.53124, one complete specimen, NE Atlantic, Bay of Biscay, Bay of Brest, 48°18’57”N, 4°22’40”W, 2 m depth, May 2017, mounted for SEM. Additional material: MNHN-IA- PNT 122, one complete specimen, NE Atlantic, Bay of Biscay, Morlaix Bay, 48°42’44”N, 3°57’04”W, 4 m depth, February 2016. MNHN-IA- PNT 123, one complete specimen, NE Atlantic, Bay of Biscay, Morlaix Bay, 48°42’40”N, 3°57’03”W, 4 m depth, February 2016. AM W.53125, one complete specimen, NE Atlantic, Bay of Biscay, Morlaix Bay, 48°42’40”N, 3°57’03”W, 4 m depth, February 2016. Description. Small species, holotype 11.5 (7.2–10.0) mm long and 0.7 (0.5–1.0) mm wide. Transverse prostomium attached to dorsal surface of base of upper lip; basal part as thick crest across dorsum, extending laterally and dorsally, covering SG1 laterally and terminating lateral to lower lip. Buccal tentacles of two types, thin tentacles uniformly cylindrical, long and thick ones deeply grooved (Fig. 16C). Peristomium forming lips; upper lip elongate, longer than wide, triangular shape, comprising medial lobe only, very convoluted (Figs 16 A–C; 17A); lower lip oval and inflated, wider than long, very glandular and ridged (Figs 16 A–B; 17A). SG1 and 2 reduced, SG1 visible dorsally, SG2 dorsally and laterally; body slightly broader until SG6, then of relatively uniform width until mid-body, posterior end tapering (Fig. 16A). Ventro-lateral inflated pads present from SG3 to SG10, smooth (Figs 16 A–C; 17A). Anterior mid-ventral groove present until end of body as a stripe, shallow and narrow between anterior ventral shields (Figs 16 A–C; 17A). Notopodia from SG3, extending for 12 (13) segments, until SG14 (SG15); distinctly elongate, rectangular, first two pairs slightly shorter, bilobed, postchaetal lobe digitiform, larger than prechaetal one (Fig. 17B). Narrowly winged notochaetae of two different lengths, with wings inconspicuous under light microscopy (Figs 16E; 17 B–C). Neuropodia beginning from SG15 (SG16); uncini with short occipitum and slightly convex base (Type 1) and with a very long prow (Fig. 16F), crest with single elongate and sharp tooth on first row above main fang, with two additional rows of shorter, irregularly sized teeth at base (Fig. 17D), absence of subrostral process (Fig. 16F). Nephridial and genital papillae not seen. Pygidium rounded with ventral pointed tip. Etymology. The species name was chosen by V. Le Garrec who sent us the type material. It refers to Penn-Ar-Bed which is the Breton name for Finistère (Finis Terrae in latin), the French department situated on western part of Brittany. The term “ pennarbedae ” is the genitive of the noun “Penn-Ar-Bed”. Habitat. Coastal maerl (rhodolith) beds in 2–7 m depth. Type locality. Glenan archipelago, Bay of Biscay, NE Atlantic Ocean, France. Distribution. Bay of Biscay (Glenan archipelago, Bay of Brest), English Channel (Morlaix Bay). Remarks. Polycirrus pennarbedae n. sp is characterized by 12–13 pairs of notopodia and neuropodia starting from SG14–15, which is similar to P. medusa (12 segments with notopodia, neuropodia from SG15), P. latidens (12 segments with notopodia, neuropodia from SG14) and P. catalanensis (13–15 segments with notopodia, neuropodia from SG15–17) (Table 2). Polycirrus pennarbedae n. sp. differs from P. medusa by the presence of a triangular upper lip with elongated median lobe (instead of trefoiled upper lip for P. medusa), the presence of a lower lip oval, wider than long (instead of upper lip subtriangular and pointing towards the mouth), by the presence of one type of notochaetae (instead of two types for P. medusa) and the ventro-lateral pads not separated by a large mid-ventral groove (large mid-ventral groove on anterior segments for P. medusa) (Table 2). Polycirrus pennarbedae n. sp. differs from P. latidens by having a narrow and shallow mid-ventral groove between ventro-laterals pads (instead of a well-defined mid-ventral groove present from SG4 for P. latidens), the presence of a lower lip oval (instead of oblong lower lip P. latidens). Polycirrus latidens is also characterized by uncini with teeth above main fang, arranged in single vertical series (instead of crests of teeth as present in P. pennarbedae) (Table 2). Finally, P. catalanensis n. sp. differs from P. pennarbedae n. sp. by the shape of uncini (two rows of teeth above the main tooth for P. pennarbedae n. sp., a single row of secondary teeth above the main tooth for P. catalanensis n. sp.), the shape of upper lip (elongated triangular medial lobe for P. pennarbedae n. sp., in contrast a very thick single medial lobe present in P. catalanensis n. sp.), the shape of lower lip (oval and wider than long for P. pennarbedae n. sp., rectangular and longer than wide P. catalanensis n. sp.) and the presence of a large mid-ventral grove on anterior segments for Polycirrus catalanensis n. sp. (narrow and shallow for P. pennarbedae n. sp.) (Table 2).

Key to European species of Polycirridae 1A. Parapodia absent (no chaetae)............................................ Hauchiella tribullata (McIntosh, 1869) 1B. Parapodia present..................................................................................... 2 2A. Only notopodia present........................................................................... 3 (Lysilla) 2B. Notopodia and neuropodia present......................................................................... 4 3A. Notochaetae with smooth tips, 6 pairs of thoracic nephridial papillae......................... L. loveni Malmgren, 1866 3B. Notochaetae with plumose tips, 9 pairs of thoracic nephridial papillae....................... L. nivea Langerhans, 1884 4A. Neuropodia with spines...................................................................... 5 (Amaeana) 4B. Neuropodia with avicular uncini.............................................................. 6 (Polycirrus) 5A. Upper lip without lobe, lower lip rounded, long achaetous region................................. A. gremarei n. sp. 5B. Upper lip with three lobes, lower lip rectangular, short achaetous region....................... A. trilobata (Sars, 1863) 6A. 28 or more segments with notochaetae.................................................................... 7 6B. 22 or less segments with notochaetae...................................................................... 8, Published as part of Lavesque, Nicolas, Hutchings, Pat, Daffe, Guillemine & Londoño-Mesa, Mario H., 2020, Revision of the French Polycirridae (Annelida, Terebelliformia), with descriptions of eight new species, pp. 151-186 in Zootaxa 4869 (2) on page 183, DOI: 10.11646/zootaxa.4869.2.1, http://zenodo.org/record/4418481, {"references":["McIntosh, W. C. (1869) On the structure of the British nemerteans, and some new British annelids. Transactions Royal Society of Edinburgh, 25, 249 - 252. https: // doi. org / 10.1017 / S 0080456800035262","Malmgren, A. J. (1866) Nordiska Hafs-Annulater. Ofversigt af Kongiliga Veteskaps-Akademiens Forhandlingar, 22, 355 - 410.","Langerhans, P. (1884) Die Wurmfauna von Madeira. IV. Zeitschrift fur wissenschaftliche Zoologie, 40, 247 - 285.","Sars, M. (1863) Geologiske og zoologiske lagttagelser, anstillede paa en Reise i en Deel af Trondhjems stift i Sommeren 1862. Nyt magazin for naturvidenskaberne, 12, 253 - 340."]}

Genus Amaeana Hartman, 1959 Type-species. Polycirrus trilobatus Sars, 1863, designated by Hartman (1959)., Published as part of Lavesque, Nicolas, Hutchings, Pat, Daffe, Guillemine & Londoño-Mesa, Mario H., 2020, Revision of the French Polycirridae (Annelida, Terebelliformia), with descriptions of eight new species, pp. 151-186 in Zootaxa 4869 (2) on page 155, DOI: 10.11646/zootaxa.4869.2.1, http://zenodo.org/record/4418481, {"references":["Hartman, O. (1959) Catalogue of the polychaetous annelids of the world. Part II. Occasional Papers of the Allan Hancock Foundation, 23, 355 - 628.","Sars, M. (1863) Geologiske og zoologiske lagttagelser, anstillede paa en Reise i en Deel af Trondhjems stift i Sommeren 1862. Nyt magazin for naturvidenskaberne, 12, 253 - 340."]}

Genus Octobranchus Marion & Bobretzky, 1875 Type species: Terebella lingulata Grube, 1863 (Octobranchus lingulatus (Grube, 1863) by monotypy).

Genus Trichobranchus Malmgren, 1866 Type species Trichobranchus glacialis Malmgren, 1866 by monotypy., Published as part of Lavesque, Nicolas, Hutchings, Pat, Daffe, Gullemine, Nygren, Arne & Londo��o-Mesa, Mario H., 2019, A revision of the French Trichobranchidae (Polychaeta), with descriptions of nine new species, pp. 151-190 in Zootaxa 4664 (2) on page 181, DOI: 10.11646/zootaxa.4664.2.1, http://zenodo.org/record/3384444, {"references":["Malmgren, A. J. (1866) Nordiska Hafs-Annulater. Ofversigt af Kongiliga Veteskaps-Akademiens Forhandlingar, 22, 355 - 410."]}

Terebellides bonifi n. sp. Figures 3–4, Table 2 Type material: Holotype: MNHN-IA-TYPE 1859, gravid, incomplete, Mediterranean Sea, Gulf of Lion, A90, 42°32’39”N, 3°16’03”E, 90 m depth, April 2018. Paratypes: MNHN-IA-TYPE 1860, one specimen, complete (posterior part used for molecular analysis), Mediterranean Sea, Gulf of Lion, A90, 42°32’39”N, 3°16’03”E, 90 m depth, April 2018; MNHN-IA-TYPE 1861, one specimen, complete, Mediterranean Sea, Gulf of Lion, A90, 42°32’39”N, 3°16’03”E, 90 m depth, April 2018. MNHN-IA-TYPE 1862, one specimen, gravid, complete, Mediterranean Sea, Gulf of Lion, A90, 42°32’39”N, 3°16’03”E, 90 m depth, April 2018, mounted for SEM. All type specimens fixed with 96% alcohol. Description. Relatively large species, with incomplete holotype 29.8 mm long (22.2– 24.3) and 1.8 mm (1.8–2 mm). Body tapering posteriorly with segments becoming increasingly shorter and more compacted towards pygidium. Prostomium compact; eyespots absent; large upper lip surrounding mouth with many buccal tentacles (Fig. 3 B–D). Buccal tentacles of 2 types, uniformly cylindrical and with expanded tips, spatulate (Fig. 3 C–D). Lower lip forming an expanded structure below upper lip (Fig. 2B, D). SG 1 and 2 short, only visible ventrally; following segments with lobes as ventral collars (Fig. 2B). Lateral lappets on SG 3–6 (TC 1–4), continuing ventrally in TC 1–5, largest on TC 1 and declining in size posteriorly (Figs 3B, D & 4A). Conspicuous dorsal rounded projection on TC 1–3 (Figs 3B, D & 4A). Presence of oval glandular lateral region on TC 3 (Fig. 3D). Branchiae arising as a single structure from TC 1, reaching TC 4 (TC 3), consisting of single elongate and annulated stalk placed mid-dorsally (Figs 3 A–D & 4A), 2 pairs of lobes, fused for about 1/2 of length, lower pair thinner. Upper lobes with about 45 tightly packed lamellae (Figs 3 A–D & 4A–B). Anterior lamellae of each branchia with short conical papillae, visible under stereomicroscope (Figs 3C & 4 A–B). Distal region of upper lobes with short projections, lower lobes with long projections (Fig. 3 C–D). Anterior branchial projection (5 th lobe) present (Figs 3 A–B, D & 4A). Eighteen pairs of thoracic notopodia (SG 3–20). First notopodium slightly longer than subsequent notopodia; notochaetae from TC 1 about same size as ones from subsequent notopodia, and transversally aligned (Fig. 3D). All notochaetae simple capillaries, arranged in two rows. Neuropodia present as sessile pinnules from TC 6 (SG 8) to pygidium; uncini arranged in single rows from TC 7. First thoracic neuropodium (TC 6) provided with 5–7 sharply bent acute tipped, geniculate chaetae. All subsequent thoracic neuropodia with about 12–15 uncini per torus arranged in one irregular row. Uncini as shafted denticulate hooks provided with long, thin and pointed main fang. Three or four teeth above the main fang, surmounted by a row of three to five short denticles and an upper crest of several minute denticles (Fig. 4C). About 32 abdominal neuropodia as erect pinnules, paddle-shaped with entire margin provided with about 40 uncini; uncini with four pointed teeth above main fang, surmounted by a row of three short pointed teeth and an upper crest of minute teeth (Fig. 4D). Two pairs of globular nephridial papillae, located latero-posteriorly to base of each notopodium of TC 4 and TC 5. Pygidium crenulated, as a funnel-like depression. Methyl green staining pattern. First 4 TC stain solid; TC 5 and TC 11 with distinct stripes; ventral faces of lobes stained dark blue; glandular region blue (Fig. 3 C–D). Etymology. The species is named after Paulo Bonifácio, alias Bonif, for his friendship and for being an excellent and fascinating worms’ teacher to NL. Habitat. Mud, about 90 m depth. Type locality. Western Mediterranean Sea (Gulf of Lion), France (Fig. 1). Distribution. Only known from type locality. Remarks. Terebellides bonifi n. sp. is characterized by the papillar projections pointing over the edge of the branchial lamellae. By this characteristic, T. bonifi n. sp. is similar to T. europea n. sp., T. gentili n. sp., T. lilasae n. sp., T. resomari n. sp. and T. stroemii from Adriatic Sea (Parapar et al. 2013) (see T. lilasae n. sp. remarks). Among these species, T. bonifi n. sp. is similar to T. europea n. sp., in having papillar projections on anterior lamellae only, but differs from this species by the presence of an oval glandular region on TC 3 (instead of undulating one for T. europea n. sp.) and presence of rounded conspicuous dorsal projection on TC 1–3. The presence of oval glandular region T. bonifi n. sp. is similar of that present in T. lilasae n. sp. and T. stroemii from Adriatic. However, these species differ by the absence of papillar projections on posterior branchial lamellae in T. bonifi n. sp. (presence on most of the lamellae T. lilasae n. sp. and T. stroemii from Adriatic), and by the shape of papillar projections, conical for T. bonifi n. sp., and rounded for T. lilasae n. sp. and T. stroemii from Adriatic. Molecular results, based on material available for this study, show that T. bonifi n. sp. appears to be absent from Northern waters (Fig. 23).

Terebellides ceneresi n. sp. Figures 5–6, Table 2 Type material: Holotype: MNHN-IA-TYPE 1863, complete, Northeast Atlantic Ocean, Bay of Biscay, P34, 43°33’31’’N, 1°43’49’’W, 112 m depth, May 2018; Paratypes: MNHN-IA-TYPE 1864, one specimen, complete, Northeast Atlantic Ocean, Bay of Biscay, P36, 43°34’58’’N, 1°42’01’’W, 126 m depth, May 2018; AM W.51400, one specimen, complete, Northeast Atlantic Ocean, Bay of Biscay, P37, 43°33’31”N, 1°43’49”W, 129 m depth, May 2018; MNHN-IA-TYPE 1865, one specimen, complete, Northeast Atlantic Ocean, Bay of Biscay, P36, 43°35’0”N, 1°42’02”W, 125 m depth, May 2018. Additional material examined: SMA _BAN_11, one specimen, complete (posterior part used for molecular analysis), Mediterranean Sea, Gulf of Lion, A90, 42°32’39”N, 3°16’03”E, 90 m depth, April 2018, mounted for SEM; SMA _BAN_20, one specimen, complete, Mediterranean Sea, Gulf of Lion, Ibis 183, 42°30’30”N, 3°09’06”E, 40 m depth, December 2018; SMA _BAN_03, one specimen, complete, gravid, Mediterranean Sea, Gulf of Lion, A90, 42°32’40”N, 3°16’05”E, 90 m depth, August 2010, mounted for SEM; SMA _BR_34, incomplete, Northeast Atlantic Ocean, Brittany, Bay of Brest, ZC, 48°18’55” N, 4°21’53” W, 5 m depth, May 2018; AM W.51401, one specimen, complete, gravid, Mediterranean Sea, Gulf of Lion, A90, 42°32’40”N, 3°16’05”E, 90 m depth, August 2010; AM W.51402, one specimen, incomplete, Mediterranean Sea, Gulf of Lion, A90, 42°32’39”N, 3°16’03”E, 90 m depth, April 2018. Description. Small species, holotype 14.2 mm long (14.2–16.5 mm) and 1.1 mm (0.8–1.1 mm). Body tapering posteriorly with segments becoming increasingly shorter and more compacted towards pygidium. Prostomium compact; eyespots absent; large upper lip surrounding mouth with many buccal tentacles (Fig. 5B, D). Buccal tentacles of 2 types, short cylindrical and very long tentacles, spatulate (Fig. 5D). Lower lip forming an expanded structure below upper lip. SG 1 and 2 short, only visible ventrally; following segments with lobes as ventral collars (Fig. 5D). Lateral lappets on SG 3–7 (TC 1–5), continuing ventrally, largest on TC 1–4 and declining in size posteriorly (Figs 5 A–C). No conspicuous dorsal rounded projection on anterior chaetigers. Glandular lateral region absent on TC 3 (Fig. 5C). Branchiae arising as a single structure from TC 1, reaching TC 7 (TC 8), consisting of a single elongate and annulated stalk placed mid-dorsally (Figs 5 B–C & 6A), two pairs of lobes, not fused, lower pair narrower (Fig. 6A). Upper lobes with about 25 tightly packed lamellae (Figs 5A & 6A). Both sides of branchial lamellae provided with several parallel rows of cilia and tufts of cilia on outer edge (Fig. 6 A–B). Branchiae provided without any papillar projections pointing over the edge of the branchial lamellae (Fig. 6B). Distal region of lower lobes with short terminal pointed projections (Fig. 6A). Anterior branchial projection (5 th lobe) absent (Fig. 6A). Eighteen pairs of thoracic notopodia (SG 3–20). First notopodium on TC 1 well-developed, same size as on subsequent notopodia, notopodia placed in a straight line; notochaetae from TC 1 much smaller (or absent) than following notochaetae (Figs 5 A–C & 6A). All notochaetae simple capillaries, arranged in two rows. Neuropodia present as sessile pinnules from TC 6 (SG 8) to pygidium; uncini arranged in single rows from TC 7. First thoracic neuropodium (TC 6) provided with about five to six sharply bent acute tipped, geniculate chaetae. All subsequent thoracic neuropodia with 7–10 uncini per torus arranged in one irregular row. Uncini as shafted denticulate hooks provided with long, thin and pointed main fang appearing bent terminally giving an ‘eagle head’ appearance (Fig. 6C). Three or four teeth above the main fang, surmounted by a row of four to five short denticles and an upper crest of several smaller denticles (Fig. 6C). About 30 abdominal neuropodia as erect pinnules paddle-shaped with entire margin provided with about 20 uncini; each with four to six pointed teeth above main fang, surmounted by a row of four or five short pointed teeth and an upper crest of minute teeth (Fig. 6D). Nephridial papillae not seen. Pygidium blunt, as a funnel-like depression. Methyl green staining pattern. First 10 TC stain solid, except TC 4 much lighter (Fig. 5D); TC 11 and TC 12, stain with distinct stripes, fading towards posterior thoracic region (without staining, first four chaetigers slightly lighter than following ones) (Fig. 5D). Etymology. The species is named in honour of the CNRS (French National Center for Scientific Research) that celebrates, in 2019, 80 years since its foundation. Habitat. Mud to sandy mud, from 90 to 140 m depth, coastal maerl (rhodolith) beds. Type locality. Near Capbreton Canyon, Bay of Biscay, Northeast Atlantic Ocean, France. Distribution. Northeast Atlantic Ocean (Bay of Biscay), Western Mediterranean Sea (Gulf of Lion), France (Fig. 1). Remarks. Terebellides ceneresi n. sp. is very similar to specimens of T. gracilis from the Adriatic Sea (Parapar et al. 2013), in being of small size, without rounded dorsal projection or glandular region in CH 3, the absence of notochaetae on first chaetiger in smaller specimens and short notochaetae on first chaetiger for larger ones and in having the same very distinctive MG staining pattern. The two species differ only by the presence of an ‘eagle head’ appearance of thoracic uncini of T. ceneresi. However, T. ceneresi n. sp. is clearly different from the holotype of T. gracilis (Parapar et al., 2011). Indeed, T. gracilis is a larger species (32 mm vs 16 mm for gravid specimens of T. ceneresi n. sp.), with a 5 th lobe present (absent in T. ceneresi n. sp.), with up to 22 uncini in thoracic chaetigers (instead of 8 in T. ceneresi n. sp.), with posterior region of lobes with pointed projections (instead of absence or very short projections in T. ceneresi n. sp.), with MG compact colouration from CH 1 to CH 10 (instead of whitish bands from CH 1 to CH 4, especially on CH 4 in T. ceneresi n. sp.) (Table 2). Molecular results, based on material available for this study, show that T. ceneresi n. sp. appears to be absent from Northern waters (Fig. 23).

El concepto de Indicador Biológico ha sido usado sin mayor precaución al momento de emplear ciertos organismos en programas de monitoreo ambiental, lo cual genera confusión. Uno de los objetivos de este trabajo fue resaltar la importancia en la consolidación de una definición precisa de este concepto a partir de información disponible, proponiendo una definición para el mismo. Las características ecológicas de los poliquetos permiten que, al estar en contacto permanente con diferentes tipos de contaminantes, respondan bioacumulando, disminuyendo o aumentando su abundancia, según sea la especie, hecho que posiciona este tipo de organismos como potenciales indicadores de contaminación marina. En este artículo se presenta de manera concreta un análisis de la literatura disponible para poliquetos en el campo de los indicadores biológicos, resaltando cómo éstos han sido usados en diferentes metodologías, con ejemplos a internacionales, así como una selección especial para Colombia. De los resultados más sobresalientes se encontró que Capitella capitata es la especie más estudiada al estar asociada con ambientes contaminados a causa del incremento de materia orgánica y es la única especie reportada en el país como indicador biológico usando las técnicas clásicas de bioindicación. Finalmente, se reitera la importancia de iniciar investigaciones sobre los aspectos ecológicos, ecotoxicológicos y bioensayos de laboratorio con otras especies de poliquetos para validar cuáles especies y por qué pueden ser consideradas como indicadores biológicos para el país.

La familia Terebellidae ha sido poco estudiada en Colombia; la información disponible ha sido reciente y solo en forma de listas faunísticas, las cuales suman 17 especies en el nororiente del Caribe colombiano. Los estudios taxonómicos de terebélidos que incluyan descripciones detalladas son prácticamente nulos. Esta investigación tuvo como objetivo evaluar taxonómicamente la diversidad de los terebélidos en Isla Fuerte (Caribe colombiano), así como aumentar el conocimiento de las especies hacia el suroccidente de la región. Se analizaron los organismos recolectados en aguas someras de siete playas, asociados a roca coralina y algas calcáreas. En total, seis géneros y diez especies de terebélidos fueron identificados. Eupolymnia rullieri, Lanicola carus, Pista palmata y Terebella verrilli ya habían sido registradas previamente para el Caribe colombiano. Polycirrus sp.1, Polycirrus sp.2 y Pista sp. son especies que aún no se describen debido a la falta de más material. Loimia minuta, Pista cetrata y Polycirrus angeli se encontraron como nuevos registros para la región. Por lo tanto, el número de especies de la familia para Caribe colombiano se aumenta a 20 y se amplía el ámbito de distribución de las últimas tres especies para el país. Se ofrece información relevante en cuanto al patrón de tinción con verde de metilo y a la variación morfológica de los ejemplares encontrados para cada especie. The family terebellidae has been poorly studied in colombia. The only available information on the family presented is in term of lists, which comprise 17 species occurring in the northeastern Colombian Caribbean. Taxonomic studies including detailed descriptions are none. This research focused on taxonomic evaluation of terebellids diversity from Isla Fuerte, Colombian Caribbean, as well as on the increasing the species knowledge for the southwestern region. Specimens from coralline rock and calcareum algae were collected and analyzed from seven beaches around the island. Six genera and ten species were found: Eupolymnia rullieri, Lanicola carus, Pista palmata, and Terebella verrilli have been already identified in the Colombian Caribbean; Polycirrus sp.1, Polycirrus sp.2 and Pista sp. remain undescribed due the lack of enough material justifing new species category. Finally, Loimia minuta, Pista cetrata and Polycirrus angeli are new records for Colombia. After this research, the number of recorded terebellid species increased to 20, and the species distributions were enlarged. Staining patterns and morphologic variation for each species are provided.

Los sipúnculos, "gusanos cacahuate" o "gusanos maní", son invertebrados marinos con gran importancia, tanto en el proceso de disolución de rocas, como por ser una importante fuente de alimento para otras especies marinas. Se realizó una búsqueda en la literatura de todas las familias, géneros y especies registradas para el Gran Caribe. Esta región comprende desde Carolina del Sur, en Estados Unidos, hasta el litoral norte de Brasil, incluyendo el Golfo de México, Antillas Mayores y Menores, y las costas de Centro y Sur América, dentro de esos límites. La revisión incluyó la mayor cantidad de información publicada, tanto electrónica como en papel, la cual se recopiló en forma de listado en orden alfabético y no filogenético. Se diferenciaron especies que no son de la región, así como las sinonimias, para ofrecer una idea acerca de los problemas taxonómicos que presenta el grupo. Se encontraron 84 nombres de especies, distribuidas en 14 géneros y seis familias. De los 84 nombres encontrados, sólo 23 nombres corresponden a especies válidas; 10 nombres no tienen información alguna. De esta forma, se comienza a organizar este grupo a través de la estandarización y organización sistemática preliminar. Por lo tanto se espera que este listado sea un aporte importante para el conocimiento del grupo en el Caribe y otras regiones cercanas. Sipunculans, or "peanut worms", are marine invertebrates with high importance in rock dissolution processes, and because they are an important source of food for other marine species. A search of all literature about recorded families, genera and species for the Grand Caribbean was done. The region comprises between the limits from South Carolina, in the United States, to the Northern littoral of Brazil, including the Gulf of Mexico, Major and Lesser Antilles, as well as Center and South American coasts. The revision includes as much published information as possible, both electronically and in hard copy, and was organized and listed in alphabetical and not phylogenetic order. Species that are not from the region where differentiated such as synonymies, in order to offer an idea about the taxonomic problems that the group has. 84 specie names where found, divided into 14 genera and 6 families. From the 84 names that where found, only 23 names correspond to valid species; 10 out of the 84 names that where found do not have sufficient information. In these terms, the group is organized through the standardized and systematic preliminary order. It is expected that this list becomes an important source of knowledge for sipunculans in the Caribbean and other close regions.

Nereididae Blainville, 1818, es una de las familias de gusanos poliquetos errantes más representativas, con cerca de 535 especies descritas en 43 géneros, y de gran importancia biológica y económica. El objetivo de esta investigación fue realizar un análisis taxonómico de los neréidido asociados a las raíces de mangle rojo, Rhizophora mangle, en el archipiélago de San Andrés, Providencia y Santa Catalina. El archipiélago forma parte de la Reserva de Biosfera Seaflower. San Andrés es la isla más importante en el archipiélago, con 27 km² de área emergida, mientras que isla Providencia, localizada a 95 km al norte de San Andrés, posee 17 km²; ambas islas tienen buen desarrollo de bosque de manglar, compuesto principalmente por R. mangle. Los neréididos, depositados en la Colección de Invertebrados Marinos de la Universidad de Antioquia (CEMUA), fueron identificados con la ayuda de claves actualizadas, mediante disección y extracción de estructuras con valor taxonómico. Se encontraron cuatro especies de poliquetos de la familia Nereididae: Neanthes acuminata, Neanthes succinea, Neanthes sp., y Platynereis mucronata. De las especies identificadas, N. succinea y N. acuminata habían sido registradas para el Caribe colombiano, y solo N. succinea había sido registrada para el archipiélago. Consideraciones sistemáticas, ecológicas y biogeográficas fueron adicionadas para cada especie. The family Nereididae Blainville, 1818, is one of the most representative group of errant polychaete worms, with about 535 described species in 43 genera, with great biological and ecological importance. The aim of this research was to make a taxonomic analysis of the nereidids associated to red mangrove roots, Rhizophora mangle, in the San Andrés, Old Providence and Santa Catalina Archipelago. This archipelago is part of the Seaflower Biosphere Reserve. San Andrés is the largest island, having 27 km² of landmass area, while Old Providence Island, located 95 km to the north of San Andrés, has 17 km²; both islands have well developed mangrove forests, composed mainly by R. mangle. The nereidids, deposited in the Coleccion Estuarina y Marina de la Universidad de Antioquia (CEMUA), were identified with updated keys, by performing dissections and extracting parts with taxonomic value. Four species of polychaetes from the family Nereididae were found: Neanthes acuminata, Neanthes succinea, Neanthes sp., and Platynereis mucronata. Only N. succinea and N. acuminata had been reported from the Colombian Caribbean, being N. succinea the only species previously reported from the archipielago. In addition, systematic, ecological and biogeographical considerations were added for each species.

Pista hataam sp. nov. Figs 3A–K Material examined: Type material: Holoype: USNM 1157001, Off Quintana Roo, UMML 22.1030, R/V “Pillsbury”, Sta. 577, Yucatan Basin, Mexican Caribbean (19°52’N 85°29’W), 22.V.1967; 4413 m depth. Description: Holotype incomplete, 32 segments, 14 mm long, thorax 10 mm long, 1.1 mm wide. Tentacular membrane long, with well defined edge; long base with two lateral groups of small, brown eyespots. Tentacles short in preserved condition. Upper lip long, with ventral edges folded; lower lip long, swollen, not covered by ventral edge of first lateral lappets (Fig. 3A). Four pairs of lateral lappets; from segment 1, almost as long as upper lip, projected forwards and dorsally near branchiae bases, with lateral edges rounded, well developed, slender margins and swollen bases (Fig. 3B); ventral edge U-shaped. Second pair well developed, but not as long as those on first and third segment, with rounded lateral edges projecting between other lappets; connected to the first ventral shield. Third pair longer dorsally, closes to the branchiae bases, and shorter ventrally, directed forwards, with rounded ventral margin, not connected to ventral shield; dorsal margin well developed. Fourth pair the shortest, a thin edge below third pair between base of first notopodium and ventral shield; ventrally connected to ventral shield. Dorsally, segment 4 with free edge developed, carrying two separated, lateral and rounded short lobes projecting anteriorly, with mid-dorsal notch between them. Segments 5–6 with short ventral lappet, between neuropodium and ventral shield. Two pairs of nephridial papillae on segments 6–7, swollen, placed dorsally and posteriorly to notopodium. Fifteen ventral shields from segment 2; first shield swollen, tessellated, separated from second thinner ventral shields. Fourth, fifth, and sixth shields slightly decreasing in size; thereafter of similar size, separated from each other. Two pairs of branched branchiae on segments 2–3 (Fig. 3C); first pair missing, scars visible; stalks short, thick, wrinkled, ending in four secondary stalks, arising alternately from the main stem; secondary stalks with dichotomous tips. Notopodia small, from segment 5; 16 notochaetigers; chaetae of one size (Fig. 3D), lanceolate, asymmetrically bilimbate, distally acute (Fig. 3E). Neuropodia from segment 5; uncini in segments 5–10 in single rows (Fig. 3F–H), MF:5:4–5:3–4:1–2, PP and PF thick, long, slightly tapering, finishing in rounded tip; Oc long, slightly convex; Cp with small teeth over a very long MF; USr and Lsr concave, SrP rounded; SrA triangular, reaching the MF tip; AP and AF absent; Bs slightly curved. Uncini in segments 11–20 in intercalated double rows, face to face (Figs 3I,J); MF:4–5:4:2–3:2, PP thick; PF short, thick, slightly tapering, finishing in rounded tip; Oc long, convex; Cp with small teeth over stout MF; USr and LSr slightly concave, divided by short, acute SrP; SrA triangular, reaching MF tip; AP and AF folded under convex Bs. Abdominal uncini in single rows (Fig. 3K), smaller; MF:4:4–5, PP rounded, PF long, thin; Oc short, slightly convex; Cp with two series of teeth decreasing in size above short MF, as long as twice the longest secondary tooth; Sr straight, SrP acute; SrA not seen; AP and AF absent. Pygidium missing. Staining pattern by methyl green: Edge of tentacular membrane stains deeply. First pair of lateral lappet stains in two bands, one lighter, external and running parallel to edge, the other darker, at the base of lappet. Second and third pairs of lateral lappets stain completely, but ventral shield is lighter. Ventral shields from segments 4–15 stain dark, in trapezoids. Notopodial glandular tissue absent, thus, dorsally, segment 4 and center of segments 5–6, stain weakly. Tentacles and branchiae do not stain. Discussion: This specimen differs from the other species of Pista reported here in having a particular staining pattern, in having a pair of short dorsal lobes at the edge of segment 4, separated by membranous tissue, with a mid-dorsal notch, and in having all thoracic uncini with long PP. Pista cetrata from Florida, is, perhaps, the closest species, having two laterodorsal lobes in segment 4 as well, but separated by a more swollen membrane. Nevertheless, the presence in P. cetrata of uncini in single rows, with a longer PP, and a blind cavity or internal pocket in the third pair of lateral lappets are the main characters to distinguish these two species. Pista hataam sp. nov., differs from P. lizae sp. nov., in that the latter has short second pair of lateral lappets, uncini only from the first three uncinigers with long posterior process, and segment 4 with only one long and rounded mid-dorsal lobe. Finally, P. hataam sp. nov., differs from the other species of Pista described in the Caribbean, in the depth in which this material was collected (4413 m). Etymology: This species is named using the compound Mayan word ha (water)-taam (deep), due to the depth where the specimen was collected. Type locality and Distribution: Yucatan Basin, Mexican Caribbean. Found at 4413 m depth., Published as part of Londoño-Mesa, Mario H., 2012, New Species of Pista Malmgren, 1866 (Polychaeta: Terebellidae) from the Caribbean, pp. 39-48 in Zootaxa 3317 (1) on pages 45-46, DOI: 10.11646/zootaxa.3317.1.3