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Family Thelepodidae Hessle, 1917 Figs 1F, 3���4 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 frequently present, in short lateral rows, or extending transversely across basal part of prostomium, usually progressively more spaced towards dorsal mid-line, with mid-dorsal gap or not; distal part of base of upper lip short, from nearly indistinct to shelf-like. Buccal tentacles all uniformly thin and cylindrical, to slightly spatulate distally (Figs 3D, F, 4B). Peristomium forming lips, sometimes also complete annulation, with dorso-lateral nuchal organs as ciliated grooves; lips expanded, relatively short upper lip, hood-like, about as long as wide; swollen, button-like, mid-ventral lower lip (Figs 3D, F, 4B���C). Segment 1 usually present all around, frequently with ventral lobe marginal to mouth (Figs 3D, F, 4B���C); SG II typically with anterior margin as protruding crest, at least ventrally (Figs 3D���E, 4B���C); lobes on following anterior segments sometimes present. Anterior segments highly glandular ventrally, smooth to highly corrugated between neuropodia within pairs, discrete shields absent (Figs 3D F, 4B); mid-ventral groove frequently extending from anterior segments with notopodia. Two to three pairs of branchiae, on SG II���III or II���IV, each pair with simple thin, curled and relatively short filaments progressively tapering to tips (Figs 3C, E, 4C), leaving mid-dorsal gap or not between filaments within pairs; branchial filaments originating directly from the body wall or from specialised dorsolateral cushion-like pads. Notopodia beginning on SG II���III, usually extending to mid-body, at least, sometimes until near posterior end; cylindrical to rectangular, distally bilobed notopodia, notochaetae originating between lobes; most taxa with winged notochaetae only, with wings of variable width (Fig. 4D), distally serrated notochaetae sometimes also present; bayonet-like and pinnate chaetae both absent. Neuropodia beginning posteriorly to notopodia, on SG IV���VI, typically on SG V; neuropodia in conjunction with notopodia as fleshy, swollen ridges, as raised rectangular to cylindrical pinnules after notopodia terminate; neurochaetae as avicular uncini frequently longer than high, with short triangular heel directed posteriorly, distinctly curved and wide base, and dorsal button near anterior margin of uncini, or within anterior third of distance between anterior margin of uncini and base of main fang (Fig. 4F). Nephridial and genital papillae usually present, on SG IV���VII, posterior to bases of notopodia or between parapodial lobes (Fig. 3C). Remarks A comprehensive phylogenetic analysis conducted by Nogueira et al. (2013) permitted the elevation of the previous Thelepodinae subfamily to Thelepodidae family level, as they represented a separate clade from other terebellids. This family is represented in European waters by three genera Euthelepus McIntosh, 1885 (a single species), Streblosoma Sars, 1872 (seven species) and Thelepus Leuckart, 1849 (nine species) (Table 1). Among these species, Thelepus japonicus Marenzeller, 1884, native from Japan, is considered as a non-indigeneous species in French waters, probably introduced with oyster transfers (Lavesque et al. 2020a) (Fig. 3C). Main morphological characters of European species BRANCHIAE. Both in Thelepus and Streblosoma genera, the number of pairs of branchiae varies between two (e.g., Streblosoma lindsayae or Thelepus nucleolata) and three (e.g., Streblosoma hutchingsae or Thelepus setosus). Branchiae in Thelepodidae are always cirriform (Figs 3C, E, 4C) but the number of branchial filaments varies among the species with for example 5���10 filaments on the second and third pairs of branchiae for Streblosoma cabiochi (Fig. 3E) and only three or less filaments for Streblosoma intestinale. Finally, the size of the medial dorsal gap separating the pairs of branchiae is a good diagnostic character. This gap is for example inconspicuous for T. parapari and wide for Thelepus cincinnatus (Nogueira 2019). PRESENCE OF EYESPOTS. The eyespots are very useful in differentiating species of Streblosoma and Thelepus for which they can be absent (e.g., Thelepus davehalli or Streblosoma hutchingsae) or present (e.g.m Thelepus corsicanus or Streblosoma nogueirai). Also, the arrangement of the eyespots, if in a continuous line, or leaving a medial gap is of taxonomic importance (Nogueira et al. 2010). START AND EXTENSION OF NOTOPODIA. The segment with the first appearance of notopodia permits the discrimination between the genus Streblosoma, for which notopodia begin on the second segment, and Euthelepus and Thelepus for which it begins on the third segment. These notopodia also extend for a variable number of segments, sometimes present only on the anterior half of the body (e.g., T. corsicanus) or present until the end of the body (T. japonicus). SHAPE OF NEUROPODIA AND UNCINI. In most of the species, the uncini start on SGV which could correspond to CH 3 (as in Thelepus) or CH 4 (as in Streblosoma). The uncini are arranged habitually in single rows but some have uncini forming loops (C-shaped arrangement) from mid thorax onwards. This last character is found for example in S. nogueirai. Between species, the uncini differ in the development of the prow (e.g., well developed in T. triserialis), the shape of the base (e.g., strongly curved in S. cabiochi), the position of the dorsal button (e.g., far from anterior margin in S. bairdi or in a terminal position for T. japonicus (Fig. 1F) and number of secondary of teeth. CREST AND LATERAL LOBES. The presence of lateral lobes on SG II���IV allows the separation of the genus Euthelepus from other genera of the family. The presence of lateral crests on SG II (= thick anterior margin) is an important character within the Streblosoma genus. For example, S. cabiochi has a very low crest on SG II (Fig. 4C) while S. bairdi has a protruding crest (Nogueira 2019). Key to European species of Thelopodidae (after Lavesque et al. 2020a ) 1. Notopodia from SG II (i.e., first branchiferous segment), start of uncini from CH 4.............................................................................................................................................................2 (Streblosoma) ��� Notopodia from SG III (i.e., second branchiferous segment), start of uncini from CH 3.................. 8 2. Two pairs of branchiae................................................................................................................................................................ Streblosoma lindsayae Lavesque, Londo��o-Mesa, Daffe & Hutchings, 2020 ��� Three pairs of branchiae.................................................................................................................... 3 3. Uncini arranged in C-shaped loops from mid thorax....................................................................... 4 ��� Uncini always in straight rows......................................................................................................... 6 4. Notopodia not extending to posterior body...................................................................................... 5 ��� Notopodia until posterior body................. Streblosoma pseudocomatus Lezzi & Giangrande, 2019 5. Eyespots absent.............................................. Streblosoma hutchingsae Lezzi & Giangrande, 2019 ��� Eyespots present................................................. Streblosoma nogueirai Lezzi & Giangrande, 2019 6- Branchiae on SG III and SG IV with 3 or less filaments on each side.......................................................................................................................... Streblosoma intestinale M. Sars in G.O. Sars, 1872 ��� Branchiae on SG III and SG IV with 5���10 filaments on each side.................................................. 7 7. Absence of prostomial process, presence of lateral crest on SG II, absence of branchial cushion............................................. Streblosoma cabiochi Lavesque, Londo��o-Mesa, Daffe & Hutchings, 2020 ��� Presence of prostomial process, absence of lateral crest on SG II, presence of branchial cushion................................................................................... Streblosoma bairdi (Malmgren, 1866) 8. Lateral lobes on SG II���IV................................................. Euthelepus setubalensis McIntosh, 1885 ��� Lateral lobes on SG I only.............................................................................................. 9 (Thelepus) 9. Two pairs of branchiae.................................................................................................................... 10 ��� Three pairs of branchiae................................................................................................................. 15 10. Uncini in a single row throughout...................................................................................................11 ��� Uncini in loops from SG XIV.............................................. Thelepus nucleolata (Clapar��de, 1870) 11. Notopodia present on 50���66% of body length............................................................................... 12 ��� Notopodia present on at least 90% of body length......................................................................... 13 12. Eyespots absent................................................................................ Thelepus davehalli Jirkov, 2018 ��� Eyespots present.............. Thelepus corsicanus Lavesque, Londo��o-Mesa, Daffe & Hutchings, 2020 13. Uncini of CH 1 with one tooth above main fang............................................................................ 14 ��� Uncini of CH 1 with two teeth above main fang............................. Thelepus parapari Jirkov, 2018 14. Eyespots present................................................................. Thelepus cincinnatus (Fabricius, 1780) ��� Eyespots absent................................................................................. Thelepus marthae Jirkov, 2018 15. Prow of uncini well developed; notch between the prow and dorsal button of the uncini well marked......................................................................................... Thelepus triserialis (Grube, 1855) ��� Prow of uncini poorly developed; notch between the prow and dorsal button of the uncini poorly marked............................................................................................................................................ 16 16. Notopodia present on about 60% of the body length............. Thelepus setosus (Quatrefages, 1866) ��� Notopodia present until end of the body length.................... Thelepus japonicus Marenzeller, 1884, 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 124-129, DOI: 10.5852/ejt.2021.782.1593, http://zenodo.org/record/5781605, {"references":["Hessle C. 1917. Zur Kenntnis der terebellomorphen Polychaeten. Zoologiska bidrag fran Uppsala 5: 39 - 258. Available from https: // www. biodiversitylibrary. org / page / 38891407 [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.","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.","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].","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]","Leuckart R. 1849. Zur Kenntnis der Fauna von Island. Archiv fur Naturgeschichte 15 (1): 149 - 208.","Marenzeller E. 1884. Sudjapanische Anneliden. II. Ampharetea, Terebellacea, Sabellacea, Serpulacea. Denkschriften der Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe 49 (2): 197 - 224.","Lavesque N., Londono-Mesa M. H., Daffe G. & Hutchings P. 2020 a. A revision of the French Telothelepodidae and Thelepodidae (Annelida, Terebelliformia), with descriptions of three species and first European record of a non-indigenous species. Zootaxa 4810 (2): 305 - 327. https: // doi. org / 10.11646 / zootaxa. 4810.2.4","Nogueira J. M. M. 2019. Redescriptions of Streblosoma bairdi (Malmgren, 1866) and Thelepus cincinnatus (Fabricius, 1780), based on types and material from type localities. Zootaxa 4544 (3): 419 - 428. https: // doi. org / 10.11646 / zootaxa. 4544.3.7","Nogueira J. M. M., Hutchings P. & Fukuda M. V. 2010. Morphology of terebelliform polychaetes (Annelida: Polychaeta: Terebelliformia), with a focus on Terebellidae. Zootaxa 2460 (1): 1 - 185. https: // doi. org / 10.11646 / zootaxa. 2460.1.1","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","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].","Claparede E. 1870. Les annelides chetopodes du Golfe de Naples. Supplement. Memoires de la Societe de Physique et d'Histoire naturelle de Geneve 20 (2): 365 - 542. https: // doi. org / 10.5962 / bhl. title. 2142","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].","Quatrefages A. de. 1866. Note sur la Classification des Annelides. Annales des Sciences Naturelles 5: 253 - 296."]}

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). ZooKeys 859: 1 - 15. https: // doi. org / 10.3897 / zookeys. 859.34117","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","Lavesque N., Hutchings P., Daffe G., Nygren A. & Londono-Mesa M. H. 2019 a. A revision of the French Trichobranchidae (Polychaeta), with descriptions of nine new species. Zootaxa 4664 (2): 151 - 190. https: // doi. org / 10.11646 / zootaxa. 4664.2.1","Parapar J., Martin D. & Moreira J. 2020 b. On the diversity of Terebellides (Annelida, Trichobranchidae) in West Africa, seven new species and the redescription of T. africana Augener, 1918 stat. prom. Zootaxa 4771 (1): 1 - 61. https: // doi. org / 10.11646 / zootaxa. 4771.1.1.","Nygren A., Parapar J., Pons J., Meissner K., Bakken T., Kongsrud J. A., Oug E., Gaev D., Sikorski A., Johansen R. A., Hutchings P., Lavesque N. & Capa M. 2018. A megacryptic species complex hidden among one of the most common annelids in the North East Atlantic. PLoS One 13 (6): e 0198356. https: // doi. org / 10.1371 / journal. pone. 0198356","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].","Malm A. W. 1874. Annulata i hafvet utmed Sveriges westkust och omkring Goteborg. Goteborgs Koniglich vetenskaps - och vitterhetssamhalles handlingar [Zoologiska observationer. VII.] 14: 67 - 105.","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","Parapar J., Moreira J. & Helgason G. V. 2011. 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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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The tanaidacean crustacean, Zeuxo holdichi was first described from Arcachon Bay, France, by Bamber in 1990. Since then, this species has been recorded several times in Europe, from Iberian Peninsula to Germany. In this present study, we report new records of this species along the northern coasts of France bordering the English Channel. It is hypothesized that Z. holdichi is being spread because of aquaculture activities.

A unique specimen of the decapod Pachygrapsus marmoratus, the marbled crab, was reported for the first time in August 2014 from the intertidal zone on the Calvados coast (western part of the Bay of Seine, Eastern English Channel). However, information is still insufficient to ascertain the origin and mode of introduction of this species into the Eastern English Channel. Among the different hypotheses, the presence of the marbled crab could be related to shipping traffic, while the occurrence of mild winters in the Bay of Seine over several years could provide favorable conditions for the permanent establishment of this Mediterranean species. Supplementary records will be necessary to confirm that this species is becoming established in the Bay of Seine, such as already observed along the western coast of the Cotentin peninsula (western English Channel).

Le règlement européen relatif aux espèces exotiques envahissantes (EEE) (1143/20141) fournit une liste d’EEE réglementées à l’échelle européenne. A ce jour, sur les 66 EEE de cette liste, seulement deux sont des espèces non indigènes (ENI) marines. L’objectif du présent travail est de proposer une liste d’espèces à réglementation à l’échelle nationale (selon les articles L411-5 dit de Niveau 1 et L411-6 dit de Niveau 2 du Code de l’Environnement2), suivant une nouvelle procédure d’analyse de risques décrite sommairement ci-dessous.