taxonID	type	description	language	source
03A4A269793961161B11F9728A6D659A.taxon	description	Figs 1 B, 2	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269793961161B11F9728A6D659A.taxon	diagnosis	Diagnosis (after Hutchings et al. 2021 a; 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. 2 A – 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. 2 B, 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. 2 A – 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. 2 C). 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. 2 A – D). Branchiae absent. Notopodia, if present, from SG III (Fig. 2 A – 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. 2 A – D); notochaetae winged (Fig. 2 E) 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 1 C, 2 F – G). Nephridial and genital papillae usually present, at anterior bases of all notopodia, or only at anteriormost notopodia (Fig. 2 A). Pygidium smooth or with rounded ventral papilla.	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269793961161B11F9728A6D659A.taxon	discussion	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. 2 A, C), Hauchiella, Lysilla and Polycirrus (Fig. 2 B, D – G) are represented in European waters (Lavesque et al. 2020 b) (Table 1).	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269793961161B11F9728A6D659A.taxon	description	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 1 B, 2 F – 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. 2 B) or with a single medial lobe (Fig. 2 D). 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. 2 B – D).. NOTOCHAETAE. Two types of notochaetae can be present: winged chaetae as for P. glasbyi (Fig. 2 E) 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. 2 E). 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. 1 B); 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. 2 F) or the absence (as for P. arenivorus) of a main tooth above the main fang, and the number of rows of secondary teeth.	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792D610918B2FEA98C98652D.taxon	description	Figs 3 – 4	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792D610918B2FEA98C98652D.taxon	diagnosis	Diagnosis (after Nogueira et al. 2018; Hutchings et al. 2021 a, 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 one pair of dorso-lateral clusters, each with several rows of eyespots (Fig. 3 A); distal part at base of upper lip, frequently with low or erect mid-dorsal tongue-like process, fused to upper lip at variable degrees, with free distal lobe (s), or free from the base. Buccal tentacles of two types, short ones thin, uniformly cylindrical, long tentacles stouter and expanded at tips, slightly spatulate (Figs 3 A – B, F, 4 A). Peristomium forming lips and continuing dorsally at least for short extension, with dorso-lateral nuchal organs at margin with prostomium; lips expanded, upper lip distinctly elongate and narrow, undulated to convoluted; swollen lower lip extending across ventrum, cushion-like or segment-like, frequently deeply grooved (Figs 3 A – B, 4 A). Either SG I or SG II reduced, not forming complete ring in many species. Anterior segments glandular ventrally, smooth, discrete shields absent and frequently with glandular regions poorly developed in comparison to other families of Terebellidae s. l.; mid-ventral groove frequently extending from anterior segments. Two pairs of cirriform branchiae on SG II – III, each pair with simple thin, curled and relatively short filaments progressively tapering to tips (Figs 3 A, 4 A), originating from raised crests on anterior margins of SG II and III, or from specialised, apparently glandular, dorso-lateral cushion-like pads occupying from anterior margins to level of posterior bases of notopodia of those segments. Notopodia beginning from SG II or III, usually SG III, extending for at least 15 segments; notopodia as short cones, notochaetae originating from central core on top, distal lobes absent; notochaetae winged, sometimes with bulbous head and alimbate tips (bayonet-like chaetae), at least in anterior row of anterior thoracic segments. Neuropodia beginning posteriorly to notopodia, usually around SG VIII – XII; neuropodia in conjunction with notopodia as sessile tori, as distinctly low pinnules after notopodia terminate; neurochaetae in single row, as avicular uncini about as long as high, with short triangular heel directed posteriorly, wide and slightly curved base, and dorsal button near midlength of uncini, but closer to anterior margin (Fig. 4 E). Nephridial and genital papillae, if conspicuous, on SG V – VII, posterior to bases of notopodia.	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792D610918B2FEA98C98652D.taxon	discussion	Remarks This recent family was described by Nogueira et al. (2013) after conducting a comprehensive phylogenetic analysis. The members of this family were previously considered as Thelepodidae but differ in having a narrow and elongate upper lip, poorly developed neuropodia and anterior segments less glandular ventrally than in other thelepodids. In European waters, this family is represented by a single species, Parathelepus collaris (Figs 3 A – B, 4 A, E; Table 1), characterised by an expanded, tongue-like upper lip, by neuropodia poorly developed and beginning from SG XI.	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792D610C1B63F9248A6D6064.taxon	description	Figs 1 F, 3 – 4	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792D610C1B63F9248A6D6064.taxon	diagnosis	Diagnosis (after Hutchings et al. 2021 a, 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 3 D, F, 4 B). 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 3 D, F, 4 B – C). Segment 1 usually present all around, frequently with ventral lobe marginal to mouth (Figs 3 D, F, 4 B – C); SG II typically with anterior margin as protruding crest, at least ventrally (Figs 3 D – E, 4 B – 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 3 D F, 4 B); 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 3 C, E, 4 C), 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. 4 D), 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. 4 F). Nephridial and genital papillae usually present, on SG IV – VII, posterior to bases of notopodia or between parapodial lobes (Fig. 3 C).	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792D610C1B63F9248A6D6064.taxon	discussion	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. 2020 a) (Fig. 3 C).	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792D610C1B63F9248A6D6064.taxon	description	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 3 C, E, 4 C) 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. 3 E) 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. 1 F) 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. 4 C) while S. bairdi has a protruding crest (Nogueira 2019).	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A2697928610518C6FCE68C1A660A.taxon	description	Figs 1 C – D, 5 – 6	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A2697928610518C6FCE68C1A660A.taxon	diagnosis	Diagnosis (after Hutchings et al. 2021 a, most important diagnostic characters highlighted in bold) Transverse prostomium attached to dorsal surface of upper lip; basal part as thick crest, eyespots frequently present (Fig. 5 B), in short rows at each lateral sides, or extending transversely across basal part of prostomium. Buccal tentacles all usually uniformly cylindrical. Peristomium usually forming lips only; lips expanded, relatively short upper lip, hood-like, about as long as wide; swollen, usually button-like and mid-ventral lower lip. Segment I terminating laterally to ventro-laterally, partially fused to expanded lower lip, or developed, forming lobes of variable extension and position. Lobes on anterior segments frequently present, of variable length, sometimes extending to SGV – VII (Figs 5 B – D, 6 A – D). Anterior segments highly glandular ventrally, with discrete, smooth to corrugated, rectangular to trapezoidal mid-ventral shields extending from anterior segments until termination of notopodia, or near it; mid-ventral groove extending from termination of mid-ventral shields. Two to three pairs of branchiae usually present (Figs 5 A – D, 6 A – D), but three genera have a single pair and several are abranchiate; branchial filaments originating all together from single point on body wall, on either side of branchiferous segments, unbranched, or, more frequently, originating from conspicuous main stalk on either side of pair, branching from one to several levels, in plumose (spiraled), dichotomous, pectinate or arborescent arrangement. Notopodia beginning on SGII – V, SGIV in most genera, usually extending to mid-body, around SGXX, but sometimes present on fewer segments or extending more posteriorly for variable extension, rarely until near posterior end; first pairs of notopodia inserted dorso-laterally, progressively more laterally, then vertically aligned; cylindrical to rectangular notopodia. Notochaetae originating from central core on top, distal lobes absent; notochaetae distally winged, wings of variable length and width, or serrate, sometimes with wings at midlength, basally to a serrated blade; some more specialised types of notochaetae may be present (Fig. 5 E – G). Neuropodia beginning posteriorly to notopodia, on SGV – IX, usually on SGV; neuropodia in conjunction with notopodia as low, sessile ridges, sometimes continuing posteriorly until pygidium, but most taxa with rectangular to cylindrical or foliaceous neuropodial pinnules after notopodia terminate; neurochaetae as avicular uncini usually as long as high, with short triangular heel directed posteriorly, slightly curved and wide base, and dorsal button (Figs 1 C – D, 6 E – F); uncini arranged in double rows (Fig. 6 E) from around SGXI usually until termination of notopodia, but several genera with double rows.	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A2697928610518C6FCE68C1A660A.taxon	discussion	Remarks In European waters, the Terebellidae s. s. are represented by 19 genera and 44 species (Table 1). Four genera are represented only by a single species: Artacama Malmgren, 1866; Baffinia Wesenberg-Lund, 1950; Laphania Malmgren, 1866; Leaena Malmgren, 1866 and Stschapovella Levenstein, 1957. Eleven of them are represented by two European species each: Amphitritides Augener, 1922; Axionice Malmgren, 1866; Lanassa Malmgren, 1866; Lanice Malmgren, 1866; Loimia Malmgren, 1866; Nicolea Malmgren, 1866; Paramphitrite Holthe, 1976; Phisidia Saint-Joseph, 1894; Pistella Hartmann-Schröder, 1996; Proclea Saint-Joseph, 1894 and Terebella Linnaeus, 1767 (Lavesque et al. 2021). The genus Eupolymnia Verrill, 1900 is represented by four species and the two most diverse European genera are Amphitrite Müller, 1771, with ten species, and Pista Malmgren, 1866 with eleven (Lavesque et al. 2021). In our recent paper focusing on French Terebellidae s. s. we have confirmed the synonymy of Neoamphitrite with Amphitrite, as suggested by several authors (Jirkov 2020; Hutchings et al. 2021 a). In contrast, we consider that Amphitritides, Lanice Loimia and Paramphitrite are still valid genera (Read & Fauchald 2021), contrary to what was proposed by Jirkov (2020) (see details in Lavesque et al. 2021).	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A2697928610518C6FCE68C1A660A.taxon	description	Main morphological characters of European species BRANCHIAE. The number and shape of branchiae are very important to separate species of Terebellidae s. s. Typically, species 2 – 3 pairs of branchiae are present on SG II – III or II – IV, but members of some genera, as for Polycirridae, completely lack branchiae: Baffinia, Lanassa, Laphania, Leaena, Phisidia and Proclea. Terebella banksyi is characterised by having branchiae on discontinuous segments: SG II – III and V (Fig. 5 D). Generally branchiae are branching (dichotomous or arborescent), originating dorsolaterally from a main stalk (Figs 5 A – D, 6 A – D) or a single point on body wall, but some species have multiple unbranching branchial filaments, like Amphitrite cirrata or A. rzhavskyi. The presence or absence and the size of the branchial stem is important, like in Eupolymnia (Figs 5 A – B, 6 D). LOBES. Genera of Terebellidae s. s. are distinguished from each other by the presence (or absence) and morphology of anterior lobes, usually positioned laterally. These structures are flaps of tissues covering at least part of the preceding segment (Nogueira et al. 2010) (Figs 5 B – C, 6 A – D). They can be absent, as in Nicolea or Terebella, narrow, as in Paramphitrite birulai or large, as in Lanice and Eupolymnia (Fig. 5 A – C) and they also vary significantly in morphology and position, from ventral to dorso-lateral (Figs 5 B – C, 6 A – D). NEPHRIDIAL AND GENITAL PAPILLAE. Terebellids are characterised by the presence of papillae situated close to the notopodia or between parapodial lobes. The nephridial papillae occur from SG III – V, while genital papillae are present from SG VI onwards and are prominent only when the animals are mature (Fig. 5 C – D, 6 C). When they are visible, the morphology and the number of these papillae and their number permit the discrimination of species, as for Amphitrite or Terebella for example. NOTOPODIA AND NEUROPODIA. Terebellidae s. s. differ by the number of pairs of notopodia, the segment on which notopodia and neuropodia start and the morphology of both noto- and neurochaetae. Usually, notochaetae are present on 17 segments, beginning from SG IV, but several exceptions exist as for example for Lanassa (n <15) or Terebella (n> 25, often present to the pygidium). Notochaetae of Terebellidae are divided in two types: distally smooth as in Pista, Eupolymnia or Lanice, or distally serrated as in Amphitritides or Paramphitrite (Fig. 5 E), and each types are sub-divided in sub groups (Nogueira et al. 2010: table 4). Concerning the neurochaetae, each part of the uncinus (Fig. 1 C – D) differ greatly among the genera of Terebellidae and their morphology should be examined in detail. For example, members of the genus Pista have long-handled uncini, with the handle originating from the heel (Fig. 1 C), while uncini in most of the other genera have short-handles. Contrary to tendons which are soft and thin structures attached to uncini, handles are chitinous structures extended from the heel. Members of the genus Loimia are unique due to the presence of pectinate uncini, with teeth arranged vertically in a single row (Fig. 1 D), while other species have multiple transverse rows of secondary teeth above the main fang (Fig. 6 E – G). The dorsal button is generally well developed and situated about midway between the base of the main fang and the tip of the prow, but is inconspicuous in specimens of Lanice and can be closer to the tip of the prow, as in Eupolymnia gili or the base of the main fang, as for Artacama proboscidea. Finally, the prow and the heel vary in shape and can be distally rounded or pointed.	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792161381B3CFA388A6C6409.taxon	description	Figs 1 A, 7 – 8	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792161381B3CFA388A6C6409.taxon	diagnosis	Diagnosis (after Hutchings et al. 2021 a, 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 7 A – C, 8 A, 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 7 C, 8 C – 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 1 A, 7 D, 8 F), 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.	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792161381B3CFA388A6C6409.taxon	discussion	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. 2019 b; Parapar et al. 2020 a) (Table 1).	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792161381B3CFA388A6C6409.taxon	description	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 7 C, 8 C) and the absence or presence of eyespots. In Octobranchus, the species differ by the shape of the branchiae (Fig. 8 D) 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. 2019 a; Parapar et al. 2020 a, 2020 b). 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. 2020 a) 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 7 A – B, 8 A – 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. 7 B), and finally the presence and the shape of papillae situated on the margins of the branchial lamellae (Fig. 8 B) (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. 8 E), 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. (2020 b) for thoracic uncini (Fig. 8 F) and Parapar et al. (2020 a) 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. (2020 a), 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. 2020 a; Fig. 7 B).	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
03A4A269792161381B3CFA388A6C6409.taxon	conservation	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. 7 B) and the compact / striped pattern of the ventral part of anterior chaetigers (e. g., CH 4 (SG VI) white in T. ceneresi).	en	Lavesque, Nicolas, Hutchings, Pat, Londoño-Mesa, Mario H., Nogueira, João M. M., Daffe, Guillemine, Nygren, Arne, Blanchet, Hugues, Bonifácio, Paulo, Broudin, Caroline, Dauvin, Jean-Claude, Droual, Gabin, Gouillieux, Benoit, Grall, Jacques, Guyonnet, Benjamin, Houbin, Céline, Humbert, Suzie, Janson, Anne-Laure, Jourde, Jérôme, Labrune, Céline, 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). European Journal of Taxonomy 782 (1): 108-156, DOI: 10.5852/ejt.2021.782.1593, URL: http://dx.doi.org/10.5852/ejt.2021.782.1593
