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Type: Monograph
Published: 2021-09-13
Page range: 1-63
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Revision of the French Terebellidae sensu stricto (Annelida, Terebelliformia), with descriptions of nine new species

CNRS, Univ. Bordeaux, EPOC, UMR 5805, Station Marine d’Arcachon, 33120 Arcachon, France
Univ. Bordeaux, CNRS, INRAE, Univ. La Rochelle UMS 2567 POREA, 33615 Pessac, France
Grupo LimnoBasE y Biotamar, Instituto de Biología Facultad de Ciencias Exactas y Naturales, Universidad de Antioquia, Calle 70 #52-21, Medellín (Antioquia), Colombia.
Australian Museum Research Institute, Australian Museum, Sydney, Australia. Department of Biological Sciences, Macquarie University, North Ryde 2109, Australia
Annelida Spaghetti Project new species molecular morphology taxonomy systematics

Abstract

This work is the last of four papers of the Spaghetti Project, aiming to revise the species of terebellids, a.k.a. “spaghetti” worms, present in the European French waters. In this last paper the Terebellidae, sensu stricto, from French waters are revised based, on material available in the French marine stations, type materials stored in the MNHN collection and newly collected specimens. Nine new species are described using both morphological and molecular tools: Eupolymnia gili n. sp., E. lacazei n. sp., E. meissnerae n. sp., Lanice kellyslateri n. sp., Paramphitrite dragovabeci n. sp., Pista labruneae n. sp., P. miosseci n. sp., P. sauriaui n. sp., and Terebella banksyi n. sp. European species of Eupolymnia are distinguished mainly by the shape of the lateral lobes and the size of the branchial stems. The two species belonging to Lanice genus are distinguished by the fusion of the first ventral shields, the shape of both noto- and neuropodia, and the pigmentation of the upper lip. The two species of Paramphitrite are distinguished by the presence or absence of a medial dorsal gap between the pairs of branchiae, by the shape of the lateral lobes and the presence or absence of a nephridial papilla on segment 4. The different species of Pista are distinguished by the number of pairs of branchiae, the shape of the lateral lobes and uncini. Finally, the two species of Terebella are distinguished by the number of segments with nephridial and genital papillae and the segments on which the branchiae occur. An identification key for European species of Terebellidae sensu stricto is also provided.

 

References

  1. Annenkova, N.P. (1924) Neues über die Verbreitung einiger Arten der Polychaeten. Comptes Rendus de l’Académie des Sciences de Russie, Leningrad, 1924, 125–128.
    Arvanitidis, C. & Koukouras, A. (1995) Amphitritides kuehlmanni sp. nov. (Polychaeta, Terebellidae, Amphitritinae) from the Aegean Sea, with comments on the genus Amphitritides Augener. Ophelia, 40 (3), 219–227. https://doi.org/10.1080/00785326.1995.10430587
    Augener, H. (1922) Über littorale Polychaeten von Westindien. Sitzungsberichte der Gesellshaft naturforschender Freunde zur Berlin, 1922 (3–5), 38–63.
    Capa M. & Hutchings P. (2006) Terebellidae (Polychaeta) from Coiba National Park, Panamanian Pacific, including description of four new species and synonymy of the genus Paraeupolymnia with Lanicola. Zootaxa, 1375, 1–29. https://doi.org/10.11646/zootaxa.1375.1.1
    Carr, C.M., Hardy, S.M., Brown, T.M., Macdonald, T.A. & Hebert P.D.N. (2011) A Tri-Oceanic Perspective: DNA Barcoding Reveals Geographic Structure and Cryptic Diversity in Canadian Polychaetes. PLoS ONE, 6 (7), e22232. https://doi.org/10.1371/journal.pone.0022232
    Claparède, E. (1869) Les Annélides Chétopodes du Golfe de Naples. Seconde partie. Ordre IIme. Annélides Sédentaires (Aud. et Edw.). Mémoires de la Société de Physique et d’Histoire Naturelle de Genève, 20 (1), 1–225.
    Costa, O.G. (1841) Description de quelques Annelides nouvelles du Golfe de Naples. Annales des sciences naturelles, Paris, Ser. 2 zoologie, 16, 267–280.
    Costa, A. (1862) Descrizione di alcuni Anellidi del Golfo di Napoli. Annuario del Museo Zoologico della R. Università di Napoli, 1 (9), 82–90.
    Dalyell, J.G. (1853) The Powers of the Creator displayed in the Creation: or, observations on life amidst the various forms of the humbler tribes of animated nature with practical comments and illustrations, volume 2. John van Voorst. London, 359 pp. https://doi.org/10.5962/bhl.title.10022
    Day, J.H. (1963) The polychaete fauna of South Africa. Part 8: New species and records from grab samples and dredgings. Bulletin of the British Museum (Natural History), Series Zoology, 10 (7), 381–445.
    Delle Chiaje, S. (1828) Memorie sulla storia e notomia degli animali senza vertebre del Regno di Napoli, 3, 1–232 pp. https://doi.org/10.5962/bhl.title.10021
    Fauchald, K. (1984) Polychaete distribution patterns, or: can animals with Palaeozoic cousins show large-scale geographical patterns? In: Hutchings P. (ed.) Proceedings of the First International Polychaete Conference’, July 1983, Sydney, Australia. The Linnean Society of New South Wales: Sydney, 1–6.
    Fauvel, P. (1927) Polychètes Sédentaires. Addenda aux Errantes, Archiannélides, Myzostomaires. Faune de France 16. Lechevalier, Paris, 494 pp.
    Gaillande, D. (1970) Une polychète Terebellidae nouvelle des côtes de Provence : Pista mediterranea n. sp., Téthys 2 (2), 443–448.
    Geller, J., Meyer, C., Parker, M. & Hawk H. (2013) Redesign of PCR primers for mitochondrial cytochrome c oxidase subunit I for marine invertebrates and application in all-taxa biotic surveys. Molecular Ecology Resources, 13 (5), 851–861. https://doi.org/10.1111/1755-0998.12138
    Giard, A. (1878) Sur les Wartelia, genre nouveau d’Annélides, considérés à tort comme des embryons de Térébelles. Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences, Paris, 86 (18), 1147–1149.
    Gil, J. (2011) The European Fauna of Annelida Polychaeta. Ph.D. Dissertation, Departamento de Biologia Animal, Faculdade de Ciências, Universidade de Lisboa, Lisboa, xlii + 1554 pp.
    Grube, A.E. (1855) Beschreibungen neuer oder wenig bekannter Anneliden. Archiv für Naturgeschichte, Berlin, 21 (1), 81–136. https://doi.org/10.5962/bhl.part.13989
    Grube, A.E. (1860) Beschreibung neuer oder wenig bekannter Anneliden. Beitrag: Zahlreiche Gattungen. Archiv für Naturgeschichte, Berlin, 26, 71–118, pls. 1–3.
    Grube, A.E. (1878) Annulata Semperiana. Beiträge zur Kenntniss der Annelidenfauna der Philippinen nach den von Herrn Prof. Semper mitgebrachten Sammlungen. Mémoires l’Académie Impériale des Sciences de St. Pétersbourg, série 7, 25, 1–300.
    Hardy, S.M., Carr, C.M., Hardman, M., Steinke, D., Corstorphine, E. & Mah, C. (2011) Biodiversity and phylogeography of Arctic marine fauna: insights from molecular tools. Marine Biodiversity, 41, 195–210. https://doi.org/10.1007/s12526-010-0056-x
    Hartman, O. (1959) Catalogue of the Polychaetous Annelids of the World. Parts 1 and 2. Allan Hancock Foundation Occasional Paper, 23, 1–628.
    Hartmann-Schröder, G. (1996) Annelida, Borstenwϋrmer, Polychaeta. Die Tierwelt Deutschlands, 58, 1–648.
    Hessle, C. (1917) Zur Kenntnis der terebellomorphen Polychaeten. Zoologiska bidrag från Uppsala, 5, 39–258.
    Holthe, T. (1976) Paramphitrite tetrabranchia gen. et sp. nov. a new terebellid polychaete from western Norway. Sarsia, 60, 59–62. https://doi.org/10.1080/00364827.1976.10411303
    Holthe, T. (1986) Polychaeta, Terebellomorpha. Marine Invertebrates of Scandinavia, 7, 1–194.
    Hutchings, P. (1990) Terebellidae (Polychaeta) from the Hong Kong region. In: Morton, B. (Ed.), Proceedings of the Second International Marine Biological Workshop: The Marine Flora and Fauna of Hong Kong and Southern China. Hong Kong, University Press, pp. 377–412.
    Hutchings, P & Glasby, C.J. (1988) The Amphitritinae (Polychaeta: Terebellidae) from Australia. Records of the Australian Museum, 40 (1), 1–60. https://doi.org/10.3853/j.0067-1975.40.1988.150
    Hutchings, P. & Kupriyanova, E. (2018) Cosmopolitan polychaetes – fact or fiction? Personal and historical perspectives. Invertebrate Systematics, 32, 1–9. https://doi.org/10.1071/IS17035
    Hutchings, P. & Lavesque, N. (2020) I know who you are, but do others know? Why correct names are so important. Zoosymposia, 19, 151–163. https://doi.org/10.11646/zoosymposia.19.1.16
    Hutchings, P., Carrerette, O., Nogueira, J.M.N.N., Hourdez, S. & Lavesque N. (2021a) The Terebelliformia- Recent developments and future directions. Diversity 13, 60. https://doi.org/10.3390/d13020060
    Hutchings, P., Nogueira, J.M.N. & Carrerette, O. (2021b) Terebellidae Johnston, 1846. In: Schmidt-Rhaesa, A. Hr., Beutel, R.G., Glaubrecht, M., Kristensen, N.P., Prendini, L., Purschke, G., Richter, S., Westheide, W. & Leschen, R.Z.E. (Ed.), Handbook of Zoology. A Natural History of the Phyla of the Animal Kingdom. Walter de Gruyter & Co, Berlin, pp. 1–64.
    Jirkov, I.A. (2001) Polychaeta of the Arctic Ocean (In Russian) Polikhety severnogo Ledovitogo Okeana. Yanus-K Press, Moscow, 632 pp.
    Jirkov, I.A. (2020) Review of the European Amphitrite (Polychaeta: Terebellidae) with description of two new species. Invertebrate Zoology, 17 (4) 311–360. https://doi.org/10.15298/invertzool.17.4.01
    Jirkov, I.A. & Leontovitch, M.K. (2017) Review of genera within the Axionice/Pista complex (Polychaeta, Terebellidae), with discussion of the taxonomic definition of other Terebellidae with large lateral lobes. Journal of the Marine Biological Association of the United Kingdom, 97 (5), 911–934. https://doi.org/10.1017/S0025315417000923
    Jirkov, I.A., Ravara, A. & Cunha, M.R. (2018) Amphitrite fauveli sp.n. (Polychaeta: Terebellidae) from the Bay of Biscay and the Gulf of Cadiz (NE Atlantic). Invertebrate Zoology, 15 (1), 85–91. https://doi.org/10.15298/invertzool.15.1.06
    Johnston, G. (1846) An index to the British annelides. Annals and Magazine of Natural History, Series 1, 16 (108 Supplement), 433–462. https://doi.org/10.1080/037454809495980
    Keferstein, W. (1862) Untersuchungen über niedere Seethiere. Zeitschrift für wissenschaftliche Zoologie, 12 (1), 1–147.
    Labrune, C., Lavesque, N., Bonifácio, P. & Hutchings, P. (2019) A new species of Pista Malmgren, 1866 (Annelida, Terebellidae) from the Western Mediterranean Sea. Zookeys, 838, 71–83. https://doi.org/10.3897/zookeys.838.28634
    Langerhans, P. (1880) Die Wurmfauna von Madeira. III. Zeitschrift für wissenschaftliche Zoologie, 34 (1), 87–143.
    Langerhans, P. (1881) Ueber einige canarische Anneliden. Nova Acta der Kaiserlichen Leopold-Carolin Deutschen Akademie der Naturforscher, Halle, 42 (3), 95–124.
    Langerhans, P. (1884) Die Wurmfauna von Madeira, IV. Zeitschrift für Wissenschaftliche Zoologie, 40 (2), 247–285.
    Lavesque, N., Bonifácio, P., Londoño-Mesa, M.H., Le Garrec, V. & Grall, J. (2017a) Loimia ramzega sp. nov., a new giant species of Terebellidae (Polychaeta) from French waters (Brittany, English Channel). Journal of the Marine Biological Association of the United Kingdom, 97 (5), 935–942. https://doi.org/10.1017/S0025315417000571
    Lavesque, N., Daffe, G., Bonifácio, P. & Hutchings, P. (2017b) A new species of the Marphysa sanguinea complex from French waters (Bay of Biscay, NE Atlantic) (Annelida, Eunicidae). Zookeys, 716, 1–17. https://doi.org/10.3897/zookeys.716.14070
    Lavesque, N., Hutchings, P., Daffe, G., Nygren A. & Londoño-Mesa, M.H. (2019a) 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
    Lavesque, N., Daffe, G., Grall, J., Zanol, J., Gouillieux, B. & Hutchings P. (2019b) 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
    Lavesque, N., Londoño-Mesa, M.H., Daffe, G. &. Hutchings, P. (2020a) 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
    Lavesque, N., Londoño-Mesa, M.H., Daffe, G. &. Hutchings, P. (2020b) Revision of the French Polycirridae (Annelida, Terebelliformia), with descriptions of eight new species. Zootaxa, 4869 (2), 151–186. https://doi.org/10.11646/zootaxa.4869.2.1
    Lavesque, N., Hutchings, P., Abe, H., Daffe, G., Gunton, L.M. & Glasby, C.J. (2020c) Confirmation of the exotic status of Marphysa victori Lavesque, Daffe, Bonifácio & Hutchings, 2017 (Annelida) in French waters and synonymy of Marphysa bulla Liu, Hutchings & Kupriyanova, 2018. Aquatic Invasions, 15 (3), 355–366. https://doi.org/10.3391/ai.2020.15.3.01
    Leidy, J. (1855) Contributions towards a knowledge of the marine Invertebrate fauna of the coasts of Rhode Island and New Jersey. Journal of the Academy of Natural Sciences of Philadelphia, 3 (2), 135–152.
    Linnaeus, C. (1767) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio duodecima reformata, 1 (2), 533–1327 + 37 pp.
    Loia M., Nicoletti L. & La Porta B. (2017) First record of genus Paramphitrite (Polychaeta: Terebellidae) in Mediterranean Sea. Marine Biodiversity Records, 10, 11. https://doi.org/10.1186/s41200-017-0113-2
    Malm, A.W. (1874) Annulata i hafvet utmed Sveriges westkust och omkring Göteborg. Göteborgs Königlich vetenskaps - och vitterhetssamhälles handlingar. [Zoologiska observationer. VII.]. Göteborgs Königlich vetenskaps - och vitterhetssamhälles handlingar, 14, 67–105.
    Malmgren, A.J. (1866) Nordiska Hafs-Annulater. Öfversigt af Kongiliga Veteskaps-Akademiens Förhandlingar, 22, 355–410.
    Marenzeller, E. (1884) Südjapanische Anneliden. II. Ampharetea, Terebellacea, Sabellacea, Serpulacea. Denkschriften der Akademie der Wissenschaften, Wien, 49 (2), 197–224.
    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. https://doi.org/10.5962/bhl.title.6513
    McIntosh, W.C. (1922) Notes from the Gatty Marine Laboratory, St. Andrews. No. XLIV. 1. On new and rare Polychaeta, Gephyrea, etc., from various regions. 2. Recent additions to the British marine Polychaeta (continued). Annals and Magazine of Natural History, Series 9, 9 (49), 1–30. https://doi.org/10.1080/00222932208632638
    Meyer, A.H. (1912) Die Amphicteniden, Ampharetiden und Terebelliden der Nord- und Ostsee. Inaugural-Dissertation zur Erlangung der Doktorwürde der hohen philosophischen Fakultät der Königlichen Christian-Albrechts-Universität zu Kiel. Heider Anzeiger, Heide, 1–68.
    Mikac, B. & Hutchings, P. (2017) One new species of Pista Malmgren, 1866 (Annelida: Terebellidae) and one new species of Pistella Hartmann-Schröder, 1996 (Annelida: Terebellidae) from the Adriatic Sea (Mediterranean). Journal of the Marine Biological Association of the United Kingdom, 97 (05), 943–953. https://doi.org/10.1017/s0025315417000868
    Montagu, G. (1813) Descriptions of several new or rare animals, principally marine, found on the south coast of Devonshire. Transactions of the Linnean Society of London, 11, 18–21. https://doi.org/10.1111/j.1096-3642.1813.tb00035.x
    Montagu, G. (1819) Descriptions of five British species of the genus Terebella. Transactions of the Linnean Society of London, 12 (2), 340–344. https://doi.org/10.1111/j.1095-8339.1817.tb00231.x
    Moore, J.P. (1907) Descriptions of new species of Polychaeta from the southeastern coast of Massachusetts. Proceedings of the Academy of Natural Sciences of Philadelphia, 58, 501–508.
    Müller O.F. (1771) Von Würmern des süssen und salzigen Wassers. Mumme & Faber, Copenhagen, 200 pp. https://doi.org/10.5962/bhl.title.14428
    Müller, O.F. (1776) Zoologicae Danicae Prodromus, seu Animalium Daniae et Norvegiae indigenarum characteres, nomina et synonyma imprimis popularium. Hallageriis. Havniae [Copenhagen], 282 pp. https://doi.org/10.5962/bhl.title.13268
    Nogueira, J.M.M., Hutchings, P. & Fukuda, M.V. (2010) Morphology of terebelliform polychaetes (Annelida: Polychaeta: Terebelliformia), with a focus on Terebellidae. Zootaxa, 2460, 1–185. https://doi.org/10.11646/zootaxa.2460.1.1
    Nogueira, J.M.M., Harris, L., Hutchings, P. & Fukuda, M.V. (2011) Four terebellines (Polychaeta, Terebellidae) with problematic taxonomic histories. Zootaxa, 2995 (1), 1–26. https://doi.org/10.11646/zootaxa.2995.1.1
    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/IS12062.
    Ørsted, A.S. (1844) Zur Classification der Annulaten mit Beschreibung einiger neuer oder unzulänglich bekannter Gattungen und Arten. Archiv für Naturgeschichte, Berlin, 10 (1), 99–112.
    Pallas, P.S. (1766) Miscellanea Zoologica quibus novae imprimis atque obscurae animalium species describunture et observationibus iconibusque illustrantur. Apud Petrum van Cleef, Hague Comitum, 224 pp. https://doi.org/10.5962/bhl.title.69851
    Palumbi, S.R. (1996) Nucleic acid II: The polymerase chain reaction. In: Hillis, D.M., Moritz, G. & Mable, B. (Eds.), Molecular systematics. Sinauer Associates, Sunderland, pp. 205–247.
    Parapar, J., Besteiro, C. & Urgorri, V. (1991) Primera cita en el litoral ibérico de Paramphitrite tetrabranchia Holte, 1976 (Polychaeta, Terebellidae). Miscellània Zoològica, 15, 63–68.
    Pearson, T.H. (1969) Scionella lornensis sp. nov., a new terebellid (Polychaeta: Annelida) from the west coast of Scotland, with notes on the genus Scionella Moore, and a key to the genera of the Terebellidae recorded from European waters. Journal of Natural History, 3 (4), 509–516. https://doi.org/10.1080/00222936900770441
    Quatrefages, A.de. (1866) Histoire naturelle des Annelés marins et d’eau douce. Annélides et géphyriens. Volume 2. Librairie Encyclopédique de Roret, Paris, 794 pp. https://doi.org/10.5962/bhl.title.122818
    Rambaut, A. (2007) FigTree. Available from: http://tree.bio.ed.ac.uk/software/figtree/(accessed 12 July 2021) https://doi.org/10.1007/BF00351456
    Read, G. & Fauchald, K. (Eds) (2021). World Polychaeta Database. Terebellidae Johnston, 1846. Available from: http://www.marinespecies.org/aphia.php?p=taxdetails&id=982 (accessed 12 July 2021)
    Risso, A. (1826) Histoire naturelle des principales productions de l’Europe méridionale et particulièrement de celles des environs de Nice et des Alpes Maritimes. Volume 4. Levrault, Paris, 439 pp. https://doi.org/10.5962/bhl.title.58984
    Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574. https://doi.org/10.1093/bioinformatics/btg180.
    Saphronova, M.A. (1988) On Cosmopolitan Distribution of Pista cristata (Polychaeta, Terebellidae). Zoologicheskii zhurnal, 67 (6), 888–897.
    Saint-Joseph, A. (1894) Les Annélides polychètes des côtes de Dinard. Annales des sciences naturelles, zoologie, Paris, Série 7, 17, 127–272.
    Saint-Joseph, A. (1898) Les Annélides polychètes des côtes de France (Manche et Océan). Annales des sciences naturelles, zoologie, Paris, Série 8, 5, 209–260.
    Sars, M. (1863) Geologiske og zoologiske Iagttagelser, anstillede paa en Reise i en Deel af Trondhjems Stift i Sommerren 1862. Nyt Magazin for Naturvidenskaberne, Christiania, 12 (3), 253–340.
    Sars, M. (1865) Fortsatte Bidrag til Kundskaben om Norges Annelider. Forhandlinger i Videnskabs-Selskabet i Christiania, 1864, 5–20.
    Savigny, J.C. (1822) Système des annélides, principalement de celles des côtes de l’Égypte et de la Syrie, offrant les caractères tant distinctifs que naturels des Ordres, Familles et Genres, avec la Description des Espèces. Description de l’Égypte ou Recueil des Observations et des Recherches qui ont été faites en Égypte pendant l’Expédition de l’Armée Française, publié par les Ordres de sa Majesté l’Empereur Napoléon le Grand, Histoire Naturelle, Paris, 1 (3), 1–128. https://doi.org/10.5962/bhl.title.66284
    Schmarda, L.K. (1861) Neue Wirbellose Thiere: Beobachted und Gesammelt auf einer Reise um die Erdr 1853 bis 1857. In Turbellarien, Rotatorien und Anneliden. Leipzig, Verlag von Wilhelm Engelmann. Erster Band, Zweite Hälfte 2, 164 pp.
    Sjölin, E., Erseus, C. & Källersjö, M. (2005) Phylogeny of Tubificidae (Annelida, Clitellata) based on mitochondrial and nuclear sequence data. Molecular Phylogenetics and Evolution, 35, 431–441. https://doi.org/10.1016/j.ympev.2004.12.018.
    Southward, E.C. (1956) On some Polychaeta of the Isle of Man. Annals and Magazine of Natural History, Series 12, 9 (100), 257–270. https://doi.org/10.1080/00222935608655812
    Ssolowiew, M. (1899) Polychaeten-Studien I. Die Terebelliden des Weissen Meeres. Annuaire du Musée Zoologique de l’Académie Impériale des Sciences de St. Pétersbourg, 4 (2), 179–220.
    Stiller, J., Tilic, E., Rousset, V., Pleijel, F. & Rouse, G.W. (2020) Spaghetti to a Tree: A Robust Phylogeny for Terebelliformia (Annelida) Based on Transcriptomes, Molecular and Morphological Data. Biology, 9, 73. https://doi.org/10.3390/biology9040073
    Stimpson, W. (1853) Synopsis of the marine Invertebrata of Grand Manan: or the region about the mouth of the Bay of Fundy, New Brunswick. Smithsonian Contributions to Knowledge, 6, 1–66.
    Thunberg, C.P. (1793) Tekning och Beskrifning på en stor Ostronsort ifrån Japan. Kongliga Vetenskaps Academiens Nya Handlingar, 14 (4–6), 140–142.
    Verrill, A.E. (1900) Additions to the Turbellaria, Nemertina, and Annelida of the Bermudas, with a revision of the New England genera and species. Transactions of the Connecticut Academy of Arts and Sciences, 10, 595–671. https://doi.org/10.5962/bhl.part.7035
    Wesenberg-Lund, E. (1950) Polychaeta. Danish Ingolf-Expedition, 4 (14), 1–92.

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