The “ Spaghetti Project ”: the final identification guide to European Terebellidae (sensu lato) (Annelida, Terebelliformia) Author Lavesque, Nicolas 3E6771E7-1A94-4FD2-8E7B-36AD6ED8B446 Univ. Bordeaux, EPOC, UMR 5805, Station Marine d’Arcachon, Arcachon, France. nicolas.lavesque@u-bordeaux.fr Author Hutchings, Pat E83A37D3-33D8-4999-ACA6-8DFECAF05D11 Australian Museum Research Institute, Australian Museum, Sydney, Australia; Department of Biological Sciences, Macquarie University, Australia. pat.hutchings@Australian.Museum Author Londoño-Mesa, Mario H. 198696D8-8FB2-4F03-AFEB-0E0773CB6669 Grupo LimnoBasE y Biotamar, Instituto de Biología Facultad de Ciencias Exactas y Naturales, Universidad de Antioquia, Medellín, Colombia. hernan.londono@udea.edu.co Author Nogueira, João M. M. C40C8C12-619D-4EC2-8998-253708120D3F Laboratório de Poliquetologia, Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Brazil. nogueira@ib.usp.br Author Daffe, Guillemine BC283B5F-3757-4941-BEED-4004C6850912 Université de Bordeaux, Observatoire Aquitain des Sciences de l’Univers, UMS 2567 POREA, Pessac, France. guillemine.daffe@u-bordeaux.fr Author Nygren, Arne 46801B86-2D81-4702-A1D1-65E1C6C40FC1 Sjöfartmuseet Akvariet, Göteborg, Sweden; Institutionen för marina vetenskaper, Göteborgs Universitet, Göteborg, Sweden. maskmedmera@gmail.com Author Blanchet, Hugues 47011CCC-0911-4EB5-9EBA-9BAF5394D042 Univ. Bordeaux, EPOC, UMR 5805, Station Marine d’Arcachon, Arcachon, France. hugues.blanchet@u-bordeaux.fr Author Bonifácio, Paulo 19882300-C635-4CF2-A7CE-98EE01C87120 Independent researcher, Brest, France. bonif@me.com Author Broudin, Caroline 0CE07B72-5E59-4A7F-8F04-D77A1167CBAB Sorbonne Université, CNRS, Station Biologique de Roscoff, 29680 Roscoff, France. broudin@sb-roscoff.fr Author Dauvin, Jean-Claude 12B3072A-421D-47A0-82BE-10125016C8D9 Normandie Univ, UNICAEN, UNIROUEN, Laboratoire Morphodynamique Continentale et Côtière, CNRS UMR 6143 M 2 C, Caen, France. jean-claude.dauvin@unicaen.fr Author Droual, Gabin C3137160-D1C5-4996-91D7-A3F4E38E25DD Ifremer, DYNECO-LEBCO, Plouzané, France; Ifremer, EMH, Nantes, France. Gabin.Droual@ifremer.fr Author Gouillieux, Benoit D0BE2D38-05DC-4DE0-BEE2-D458ED5C3299 Univ. Bordeaux, EPOC, UMR 5805, Station Marine d’Arcachon, Arcachon, France. benoit.gouillieux@u-bordeaux.fr Author Grall, Jacques 74F9A220-94E5-4C80-B375-0FD4C894F804 Univ Brest, CNRS, IRD, OSU-IUEM, Plouzané, France. jacques.grall@univ-brest.fr Author Guyonnet, Benjamin 82664894-3735-4CA4-8443-D904052FE3CF Environnement, Auray, France. b.guyonnet@tbm-environnement.com Author Houbin, Céline 41D368C7-3084-4E8A-B6EF-24E184D4752E Sorbonne Université, CNRS, Station Biologique de Roscoff, 29680 Roscoff, France. houbin@sb-roscoff.fr Author Humbert, Suzie 4354F19C-8567-4713-8E3B-492B4025EF26 Univ. Bordeaux, EPOC, UMR 5805, Station Marine d’Arcachon, Arcachon, France. suzie.humbert@u-bordeaux.fr Author Janson, Anne-Laure 587C0411-5819-4FEB-AD5F-A0107D72A725 CNRS, MNHN, UMS 2006 Patrimoine Naturel, Station Marine de Dinard, Dinard, France. anne-laure.janson@mnhn.fr Author Jourde, Jérôme 746DEE71-E146-4355-B164-D55D9A971044 CNRS, La Rochelle Université, Littoral Environnement et Sociétés, UMR 7266 LIENSs, La Rochelle, France. jjourde@univ-lr.fr Author Labrune, Céline 9200C8BA-E199-46B8-8727-BCF01DBC383A CNRS, Sorbonne Université, Laboratoire d’Ecogéochimie des Environnements Benthiques, LECOB, Banyuls, France. celine.labrune@obs-banyuls.fr Author Lamarque, Bastien BE414235-3305-483C-8ED2-53AF7F68482A Univ. Bordeaux, EPOC, UMR 5805, Station Marine d’Arcachon, Arcachon, France. bastien.lamarque@u-bordeaux.fr Author Latry, Lise 9F307F07-9830-4AE7-A77C-397BA3B8BE19 Univ. Bordeaux, EPOC, UMR 5805, Station Marine d’Arcachon, Arcachon, France. lise.latry@u-bordeaux.fr Author Garrec, Vincent Le 00E7C229-D215-4035-96C5-43B072EFC28A Univ Brest, CNRS, IRD, OSU-IUEM, Plouzané, France. Vincent.Legarrec@univ-brest.fr Author Pelaprat, Corine AD117DA7-DFA1-4190-BA8B-4958AF593822 Benthos Identification, 33840 Escaudes, France; Stareso, Calvi, France. benthid@gmail.com Author Pezy, Jean-Philippe 9307F067-DF5F-4BF1-B7EF-9B876640B891 Normandie Univ, UNICAEN, UNIROUEN, Laboratoire Morphodynamique Continentale et Côtière, CNRS UMR 6143 M 2 C, Caen, France. jean-philippe.pezy@unicaen.fr Author Sauriau, Pierre-Guy C5012CB5-D12A-468F-9536-E0C08F9A91E6 CNRS, La Rochelle Université, Littoral Environnement et Sociétés, UMR 7266 LIENSs, La Rochelle, France. pierre-guy.sauriau@univ-lr.fr Author Montaudouin, Xavier De D90D06BE-E569-4B4B-98F4-AFB1E183C2A5 Univ. Bordeaux, EPOC, UMR 5805, Station Marine d’Arcachon, Arcachon, France. xavier.de-montaudouin@u-bordeaux.fr text European Journal of Taxonomy 2021 2021-12-14 782 1 108 156 http://dx.doi.org/10.5852/ejt.2021.782.1593 journal article 2940 10.5852/ejt.2021.782.1593 313d099b-3bb0-4272-b956-c466e3591188 2118-9773 5781605 510DE23F-4CE5-4DDF-B1E7-CA8346AA4F5F 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 ). Fig. 3. Diversity of Telothelepodidae Nogueira, Fitzhugh & Hutchings, 2013 and Thelepodidae Hessle, 1917 . A . Parathelepus collaris (Southern, 1914) , anterior end, frontal view (AM W.53063). B . Parathelepus collaris , anterior end, ventral view (NHMUK ANEA 1983.1696). C . Thelepus japonicus Marenzeller, 1884 , anterior end, lateral view (AM W.53073). D . Thelepus corsicanus Lavesque, Londoño-Mesa, Daffe & Hutchings, 2020 , anterior end, frontal view (AM W.53068). E . Streblosoma cabiochi Lavesque, Londoño-Mesa, Daffe & Hutchings, 2020 , anterior end, dorsal view (MNHN-IA- TYPE 2000). F . Thelepus japonicus , anterior end, ventral view (MNHN-IA-PNT 117). Abbreviations: Br = Branchiae; Bt = Buccal tentacles; Gp = genital papillae; Ll = Lower lip; Ul = Upper lip. Fig. 4. Diversity of Telothelepodidae Nogueira, Fitzhugh & Hutchings, 2013 and Thelepodidae Hessle, 1917 , SEM. A . Parathelepus collaris (Southern, 1914) , anterior end, frontal view (AM W.53063). B . Thelepus corsicanus Lavesque, Londoño-Mesa, Daffe & Hutchings, 2020 , anterior end, latero-frontal view (AM W.53069). C . Streblosoma cabiochi Lavesque, Londoño-Mesa, Daffe & Hutchings, 2020 , anterior end, lateral view (AM W.53066). D . Thelepus corsicanus , thoracic notochaetae (AM W.53069). E . Parathelepus collaris , abdominal uncini, (AM W.53063). F . Thelepus japonicus Marenzeller, 1884 , abdominal uncini, SG48. (SMA-NL-Thele08). Abbreviations: Br = Branchiae; Bt = Buccal tentacles; Lc = lateral crest; Ll = Lower lip; Vl = Ventral lobe of SG I. 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, Londoño-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, Londoño-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 ( Claparède, 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, Londoño-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