Abstract
Alitta virens species complex encompasses elongate nereidids appreciated commercially both in the fishing and aquaculture industries. This complex has been well studied in biological and ecological terms. Nevertheless, detailed taxonomic analyses have scarcely been addressed, to the extent that only a few species in the complex have been recognized as valid but with some difficulties: Alitta brandti Malmgren, 1865 (Sea of Okhotsk), A. grandis (Stimpson, 1853) (northeastern USA) and A. virens (Sars, 1835) (Norway). Whereas, other several species have typically been regarded as synonyms, including those originally described from the North Pacific Ocean: Nereis (Alitta) virens plenidentata Moore, 1909 (California, USA), Nereis dyamusi Izuka, 1912 (Japan) and Nereis foliata Baird, 1863 (Vancouver, Canada). In this study, an examination of the immature and epitoke type and non-type material available for the A. virens species complex from the North Pacific was carried out. Herein, the status of A. brandti as a valid species is reinforced being clearly distinct from A. virens and related species. Alitta dyamusi n. comb., A. plenidentata n. comb. and A. williami nom. nov. are reinstated and transferred to Alitta, the specific epithet of the latter species is a replacement name for Nereis foliata Baird, 1863, which is a junior, primary homonym of Nereis foliata Dalyell, 1853. Lectotypes for A. plenidentata n. comb. and A. williami nom. nov. are designated. The North Pacific species of the A. virens complex, excluding A. plenidentata n. comb. which has several distinctive features that differ from all the Alitta species, are characterized by having homogomph spinigers in both supracicular and subacicular neurochaetae, oral ring with a larger number of rows and paragnaths, and epitoke males with unmetamorphosed pygidium and epitoke-modified chaetae in both neuropodial fascicles. The morphology of epitoke females in the A. virens complex is described for the first time. Identification keys to atoke and epitoke species of this complex are also provided.
References
Abbott, D.P. & Reish, D.J. (1980) Polychaeta: the marine annelid worms. In: Morris, R.H., Abbott, D.P. & Haderlie, E.C. (Eds.), Intertidal invertebrates of California. Stanford University Press, Stanford, California, pp. 448–489.
Abdel-Moez, M.K. & Humphries, C.F. (1955) A description of a new nereid - Nereis southerni. Proceedings of the Royal Irish Academy. Section B: Biological, Geological, and Chemical Science, 57 (11), 147–153.
Armstrong, J.L., Armstrong, A.D. & Mathews, S.B. (1995) Food habits of estuarine staghorn sculpin, Leptocottus armatus, with focus on consumption of juvenile Dungeness crab, Cancer magister. Fishery Bulletin, 93, 456–470.
Augener, H. (1918) Polychaeta. Beitrage zur Kenntnis der Meeresfauna Westafrikas, 2 (2), 67–625.
Baird, W. (1863) Descriptions of several new species of worms belonging to the Annelida Errantia and Sedentaria or Tubicola of Milne Edwards. Proceedings of the Zoological Society of London, 1863, 106–110.
Bakalenko, N.I., Novikova, E.L., Nesterenko, A.Y. & Kulakova, M.A. (2013) Hox gene expression during postlarval development of the polychaete Alitta virens. EvoDevo, 4, 13.
http://dx.doi.org/10.1186/2041-9139-4-13
Bakken, T., Glasby, C.J. & Wilson, R.S. (2009) A review of paragnath morphology in Nereididae (Polychaeta). Zoosymposia, 2, 305–316.
Bakken, T. & Wilson, R.S. (2005) Phylogeny of nereidids (Polychaeta, Nereididae) with paragnaths. Zoologica Scripta, 34, 507–547.
http://dx.doi.org/10.1111/j.1463-6409.2005.00200.x
Banse, K. & Hobson, K.D. (1974) Benthic errantiate polychaetes of British Columbia and Washington. Bulletin of the Fisheries Research Board of Canada, 185, 1–111.
Bass, N.R. & Brafield, A.R. (1972) The life cycle of the polychaete Nereis virens. Journal of the Marine Biological Association of the United Kingdom, 52, 701–726.
http://dx.doi.org/10.1017/S0025315400021664
Berger, V.Y. & Khalaman, V.V. (1996) Non-spawning migrations of Nereis virens in the White Sea. Reports of the Academy of Sciences, 349 (5), 710–711. [in Russian]
Berkeley, E. & Berkeley, C. (1935) Some notes on the polychaetous annelids at Elkhorn Slough, Monterey Bay, California. American Midland Naturalist, 16 (5), 766–775.
Blake, J.A. (1975) Phylum Annelida: Class Polychaeta. In: Smith, R.I. & Carlton, J.T. (Eds.), Light's Manual: Intertidal Invertebrates of the Central California Coast. University of California Press, Berkeley, pp. 151–243.
Blake, J.A. & Ruff, R.E. (2007) Polychaeta. In: Carlton, J.T. (Ed.), The Light and Smith Manual. Intertidal Invertebrates from Central California to Oregon. University of California Press, Berkeley, pp. 309–410.
Blake, R.W. (1979) On the exploitation of a natural population of Nereis virens Sars from the north-east coast of England. Estuarine and Coastal Marine Science, 8, 141–148.
http://dx.doi.org/10.1016/0302-3524(79)90069-0
Bondarenko, S.K. (1997) The life cycle of Wardium fryei (Cestoda: Hymenolepididae). Parasitology, 31 (2), 142–156. [In Russian].
Brafield, A.E. & Chapman, G. (1967) Gamotogenesis and breeding in a natural population of Nereis virens. Journal of the Marine Biological Association of the United Kingdom, 47, 619–627.
http://dx.doi.org/10.1017/S0025315400035232
Buzhinskaja, G. N. (1985) Polychaeta of the shelf off south Sakhalin and their ecology. Academy of Sciences Zoological Institute Research of the Fauna of the Seas, 30 (38), 72–224. [In Russian].
Chamberlin, R.V. (1919) Pacific coast Polychaeta collected by Alexander Agassiz. Bulletin of the Museum of Comparative Zoology, 63 (6), 251–270.
Chambers, S. & Garwood, P. (1992) Polychaetes from Scottish waters. A guide to identification: Part 3 Family Nereidae. National Museums of Scotland, Edinburgh, 64 pp.
Clark, R.B. (1960) The fauna of the Clyde Sea area: Polychaeta. Scottish Marine Biological Assocation, Millport, 71 pp.
Clark, R.B. (1961) The origin and formation of the Heteronereis. Biological Reviews, 36, 199–236.
http://dx.doi.org/10.1111/j.1469-185X.1961.tb01584.x
Clark, R.B. & Tritton, D.J. (1970) Swimming mechanisms in nereidiform polychaetes. Journal of Zoology, 161 (2), pp. 257–271.
http://dx.doi.org/10.1111/j.1469-7998.1970.tb02040.x
Conde-Vela, V.M. & Salazar-Vallejo, S.I. (2015) Redescriptions of Nereis oligohalina (Rioja, 1946) and N. garwoodi González-Escalante & Salazar-Vallejo, 2003 and description of N. confusa sp. n. (Annelida, Nereididae). ZooKeys, 518, 15–49.
http://dx.doi.org/10.3897/zookeys.518.9564
Conde-Vela, V.M., Wu, X. & Salazar-Vallejo, S.I. (2018) Reevaluation and new species of Kainonereis Chamberlin, 1919 (Annelida: Polychaeta: Nereididae). Zoological Studies, 57, 6.
http://dx.doi.org/10.6620/ZS.2018.57-06
Creaser E.P. & Clifford, D.A. (1982) Life history studies of the sandworm, Nereis virens Sars, in the Sheepscot estuary, Maine. Fishery Bulletin, 80 (4), 735–743.
Creaser, Jr., E.P., Clifford, D.A., Hogan, M.J. & Sampson, D.B. (1983) A commercial sampling program for sandworms, Nereis virens Sars, and bloodworms, Glycera dibranchiata Ehlers, harvested along the Maine coast. NOAA Technical Report NMFS SSRF-767. U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, 53 pp.
Cunningham, J.T. (1887) On some points in the anatomy of Polychaeta. Quarterly Journal of Microscopical Science, 28, 239–278.
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, vol. 2. John van Voorst, London, 359 pp.
de Blainville, H. (1818) Mémoire sur la classe des Sétipodes partie des Vers á sang rouge de M. Cuvier, et des Annélides de M. de Lamarck. Bulletin des Sciences par la Société Philomatique de Paris, 1818, 78–85.
de Quatrefages, A. (1866) Histoire Naturelle des Annelés Marins et d’Eau Douce: Annélides et Géphyriens. Tome Premier. Librairie Encyclopédique de Roret, Paris, 588 pp.
Desrosiers, G., Caron, A., Olivier, M. & Miron, G. (1994) Cycle de développement d'une population intertidale de Nereis virens (Polychaeta Nereidae) de l'estuaire maritime du Saint-Laurent. Oceanologica Acta, 17 (6), 683–695.
Du Clos, K.T., Lindsay, S.M. & Jumars, P.A. (2013) Spatial distribution of Alitta virens and Clymenella torquata with respect to rigid boundaries in mud and sand. Journal of Marine Research, 71 (3), 211–226.
http://dx.doi.org/10.1357/002224013807719482
Ehlers, E. (1868) Die Borstenwürmer (Annelida Chaetopoda) nach Systematischen und Anatomischen Untersuchungen. Verlag von Wilhelm Engelmann, Leipzig, xx + 269, 269–748, 24 taf.
Ehlers, E. (1901) Die Polychaeten des magellanischen und chilenischen Strandes: Ein faunisticher Versuch. Fetschrift zur Feier des 150jährigen Bestehens der Königlichen Gesellschaft der Wissenschaften zu Göttingen. Weidmannsche, Berlin, 232 pp.
Evans, S.M. (1963) Behaviour of the polychaete Nereis in T-mazes. Animal Behaviour, 11 (2–3), 379–392.
http://dx.doi.org/10.1016/S0003-3472(63)80129-3
Fauvel, P. (1923) Polychètes errantes. Faune de France, 5, 1–488.
Fernald, R.L., Hermans, C.O., Lacalli, T.C., Wilson, Jr, W.H. & Woodin, S.A. (1987) Phylum Annelida, Class Polychaeta. In: Strathmann, M.F. (Ed.), Reproduction and development of marine invertebrates of the northern Pacific coast. University of Washington Press, Seattle, pp. 138–195.
Foster, J.W. & Stimpson, W. (1871) To the Friends and Correspondents of the Chicago Academy of Sciences. Chicago Academy of Science, October 30th, 1871, 4 pp. Available from: https://www.loc.gov/resource/rbpe.01802000/?sp=1 (Accessed 11 Mar. 2017)
Gilpin-Brown, J.B. (1958) The development and structure of the cephalic nerves of Nereis. Journal of Comparative Neurology, 109 (3), 317–348.
http://dx.doi.org/10.1002/cne.901090302
Glasby, C.J. (1999) The Namanereidinae (Polychaeta: Nereididae). Part 1, taxonomy and phylogeny. Records of the Australian Museum, Supplement, 25, 1–129.
Grube, A.E. (1851) Annulaten. Dr. A. Th. von Middendorff Reise in den Äussersten Norden und Osten Sibiriens während der Jahre 1843 und 1844, mit allerhöchster Genehmigung auf Veranstaltungder kaiserlichen Akademie der Wissenschaften zu St. Petersburg ausgeführt und in Verbindung mit vielen Gelehrten, Theil 2. Buchdruckerei der Kaiserlichen Akademie der Wissenschaften, 24 pp.
Grube, A.E. (1858) Annulata örstediana. Enumeratio Annulatorum, quae in itinere per Indiam occidentalem et Americam centralem annis 1845–1848 suscepto legit cl. A.S. Örsted, adjectis speciebus nonnullis a cl. H. Kröyero in itinere ad Americam meridionalem collectis, (Fortsættelse [continued]) [2. Familia Euniceae - F. Syllidea]. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 1857, 158–186.
Grube, E. (1870) Bemerkungen über anneliden des Pariser Museums. Archiv für Naturgeschichte, Berlin, 36 (1), 281–352.
Gustus, R.M. & Cloney, R.A. 1973. Ultrastructure of the larval compound setae of the polychaete Nereis vexillosa Grube. Journal of Morphology, 140 (3), 355–366.
Hartman, O. (1936) New species of polychaetous annelids of the family Nereidae from California. Proceedings of the United States National Museum, 83, 467–480.
Hartman, O. (1938) Brackish and fresh-water Nereidae from the northeast Pacific, with the description of a new species from central California. University of California Publications in Zoology, 43 (4), 79–82.
Hartman, O. (1948) The polychaetous annelids of Alaska. Pacific Science, 2 (1), 3–58.
Hartman, O. (1954) The marine annelids of San Francisco Bay and its environs, California. Occasional Papers of the Allan Hancock Foundation, 15, 1–20.
Hartman, O. (1956) Polychaetous annelids erected by Treadwell, 1891 to 1948, together with a brief chronology. Bulletin of the American Museum of Natural History, 109 (2), 239–310.
Hartman, O. (1959) Catalogue of the polychaetous annelids of the world. Part 1. Occasional Papers of the Allan Hancock Foundation, 23, 1–353.
Hartman, O. (1968) Atlas of the errantiate polychaetous annelids from California. Allan Hancock Foundation, University of Southern California, Los Angeles, 828 pp.
Hartman, O. & Reish, D.J. (1950) The marine annelids of Oregon. Oregon State Monographs Studies in Zoology, 6, 1–64.
Healy, E.A. & Wells, G.P. (1959) Three new lugworms (Arenicolidae, Polychaeta) from the north Pacific area. Proceedings of the Zoological Society of London, 133 (2), 315–355.
Herpin, R. (1923) Sur les rélations sexuelles chez Perinereis cultrifera Gr. Comptes Rendus de l'Académie des sciences de Paris, 176, 417–420.
Hiebert, T.C. (2015) Neanthes brandti. In: Hiebert, T.C., Butler, B.A. & Shanks, A.L. (Eds.), Oregon Estuarine Invertebrates: Rudys' Illustrated Guide to Common Species, 3rd ed. University of Oregon Libraries and Oregon Institute of Marine Biology, Charleston, Oregon, pp. 158–163.
Hylleberg, J., Nateewathana, A. & Bussarawit, S. (1986) Polychaetes of Thailand, Nereidae (Part 1), Perinereis and Pseudonereis, with notes on species of commercial value. Phuket Marine Biological Center Research Bulletin, 43, 1–22.
ICZN (International Commission on Zoological Nomenclature). (1999) International Code of Zoological Nomenclature, 4th ed. International Trust for Zoological Nomenclature (The Natural History Museum), London, 306 pp.
http://dx.doi.org/10.5962/bhl.title.50608
Imajima, M. (1972) Review of the annelid worms of the family Nereidae of Japan, with descriptions of five new species or subspecies. Bulletin of the National Science Museum, Tokyo, 15, 37–153.
Imajima, M. & Hartman, O. (1964) The polychaetous annelids of Japan. Occasional Papers of the Allan Hancock Foundation, 26 (1), 1–237.
Izuka, A. (1912) The errantiate polychaeta of Japan. Journal of the College of Science, Imperial University, Tokyo, 30, 1–262.
Jansonius, J. & Craig, J.H. (1971) Scolecodonts: I. Descriptive terminology and revision of systematic nomenclature; II. Lectotypes, new names for homonyms, index of species. Bulletin of Canadian Petroleum Geology, 19 (1), 251–302.
Johnson, A.M. (1972) John Keast Lord. Dictionary of Canadian Biography, vol. 10, University of Toronto/Université Laval. Available from: http://www.biographi.ca/en/bio/lord_john_keast_10E.html (Accessed 17 May 2017)
Johnson, H.P. (1901) The Polychaeta of the Puget Sound region. Proceedings of the Boston Society for Natural History, 29 (18), 381–437.
Khlebovich, V.V. (1963) Biology of Nereis virens in Kandalaksha Bay of White Sea. Proceedings of the White Sea Biological Station, Moscow State University, 2, 250–257. [in Russian]
Khlebovich, V.V. (1996) Fauna of Russia and neighboring countries. Polychaetous annelids. 3. Polychaetes of the family Nereididae of the Russian Seas and the adjacent waters. New Series 140. Russian Academy of Sciences, Zoological Institute, St Petersburg. Nauka Publishing House, Moscow, 222 pp. [in Russian]
Khlebovich, V.V. & Jirkov, I.A. (2001) Nereidae Lamarck, 1818. In: Jirkov, I.A. (Ed.), Polychaeta of the Arctic Ocean. Yanus-K, Moscow, pp. 222–235. [in Russian]
Khlebovich, V.V., Komendantov, A.Y. & Shklyarevich, G.A. (1980) Validity of Nereis virens, N. grandis and N. brandti (Annelida, Polychaeta) and variations in the number of their paragnaths. Zoological Journal, 59, 1617–1624. [in Russian]
Khlebovich, V.V. & Shklyarevich, G.A. (1977) About the origin of Nereis virens Sars in Kandalaksha Bay of White Sea. Zoological Journal, 58 (3), 464–467. [in Russian]
Kinberg, J.G.H. (1865) Annulata nova. Nereidum dispositio nova. Öfversigt af Kongelige Vetenskaps-Akademiens Förhandlingar, 22, 167–179.
Kozin, V.V. & Kostyuchenko, R.P. (2015) Vasa, PL10, and Piwi gene expression during caudal regeneration of the polychaete annelid Alitta virens. Development Genes and Evolution, 225 (3), 129–138.
http://dx.doi.org/10.1007/s00427-015-0496-1
Kulakova, M.A., Cook, C.E. & Andreeva, T.F. (2008) ParaHox gene expression in larval and postlarval development of the polychaete Nereis virens (Annelida, Lophotrochozoa). BMC Developmental Biology, 8, 61.
http://dx.doi.org/10.1186/1471-213X-8-61
Langdon, F.E. (1900) The sense-organs of Nereis virens, Sars. Journal of Comparative Neurology, 10 (1), 1–77.
http://dx.doi.org/10.1002/cne.910100102
Last, K.S. & Olive, P.J.W. (2004) Interaction between photoperiod and an endogenous seasonal factor in influencing the diel locomotor activity of the benthic polychaete Nereis virens Sars. Biological Bulletin, 206 (2), 103–112.
http://dx.doi.org/10.2307/1543541
Leuckart, R.S. (1847) Verzeichnis der zur Fauna Helgoland’s gehørenden wirbellosen Seethiere. In: Frey, H. & Leuckart, R.S. (Eds.), Beiträge zur Kenntnis Wirbelloser Thiere mit Besonderer Berücksichtigung der Fauna des Norddeutschen Meeres. Friedrich Vieweg und sohn, Braunschweig, pp. 136–168.
Levinsen, G.M.R. (1883) Systematisk-geografisk Oversigt over de nordiske Annulata, Gephyrea, Chaetognathi og Balanoglossi. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i København, 1882, 160–251.
Loi, T. (1980) Catalogue of the types of polychaete species erected by J. Percy Moore. Proceedings of the Academy of Natural Sciences, Philadelphia, 132, 121–149.
Lord, J.K. (1866) The naturalist in Vancouver Island and British Columbia, vol. 2. Richard Bentley, London, 375 pp.
MacGinitie, G.E. (1935) Ecological aspects of a California marine estuary. The American Midland Naturalist, 16 (5), 629–765.
MacGinitie, G.E. & MacGinitie, N. (1968) Natural History of Marine Animals. 2nd ed. McGraw-Hill, New York, 1968. 14 + 523 pp.
Malmgren, A.J. (1865) Nordiska Hafs-Annulater. Öfversigt af Konglia Vetenskaps-Akademiens förhandlingar, Stockholm, 22, 51–110.
Malmgren, A.J. (1867) Annulater polychaeta Spetsbergiae, Gronlandiae, Islandiae et Scandinaviae hactenus cognita. Öfversigt af Konglia Vetenskaps-Akademiens förhandlingar, Stockholm, 24, 127–235.
Malmgren, A.J. (1869) Ueber die Gattung Heteronereis (Erst.) und ihr Verhältniss zu den Gattungen Nereis (Gr.) und Nereilepas (Gr.). Archiv für Naturgeschichte, 35 (1), 58–61.
Manning, R.B. (1993) The scientific contributions of William Stimpson, an early American naturalist and taxonomist. In: Truesdale, F. (Ed.), History of Carcinology, A. A. Balkema Publishers, Rotterdam, pp. 109–117.
Mayer, A.G. (1918) Biographical memoir of William Stimpson, 1832–1872. National Academy of Sciences of the United States of America, Biographical Memoirs 8, 419–433.
McIntosh, W.C. (1910) A monograph of the British annelids, 2. Polychaeta. Syllidae to Ariciidae. Vol. 2. Ray Society of London, London, 291 pp.
Moore, J.P. (1903) Polychaeta from the coastal slope of Japan and from Kamchatka and Bering Sea. Proceedings of the Academy of Natural Sciences of Philadelphia, 55, 401–490.
Moore, J.P. (1904) New Polychaeta from California. Proceedings of the Academy of Natural Sciences of Philadelphia. 56: 484–503.
Moore, J.P. (1908) Some polychaetous annelids of the northern Pacific coast of North America. Proceedings of the Academy of Natural Sciences of Philadelphia, 60, 321–364.
Moore, J.P. (1909) Polychaetous annelids from Monterey Bay and San Diego, California. Proceedings of the Academy of Natural Sciences of Philadelphia, 61, 235–295.
Müller, O.F. (1806). Zoologia Danica seu animalium Daniae et Norvegiae rariorum ac minus notorum descriptiones et historia. N. Christensen, Havniae (Copenhagen). Volumen quartum: [1–5], pp. 1–46, pls. 121–160.
Nishi, E. & Tanaka, K. (2011) Phylum Annelida (Classes Polychaeta and Myzostomida): a catalogue of Akira Izuka’s type and non-type polychaete collection in the University Museum, the University of Tokyo. In: Ueshima, R. (Ed), Catalogue of invertebrate collection deposited in the Department of Zoology, the University Museum, the University of Tokyo (3). Material Reports, No. 90. The University Museum, The University of Tokyo, Tokyo, pp. 1–57.
Oglesby, L.C, Mangum, C.P., Heacox, A.E. & Ready, N.E. (1982) Salt and water balance in the polychaete Nereis virens. Comparative Biochemistry & Physiology Part A: Physiology, 73 (1), 15–19.
http://dx.doi.org/10.1016/0300-9629(82)90085-8
Oldfield, J.D. & Shaw, D.J.B. (2016) The development of Russian environmental thought. Scientific and geographical perspectives on the natural environment, 1880s–1960s. Routledg, London, 212 pp.
Olive, P.J.W. (1993) Management of the exploitation of the lugworm Arenicola marina and the ragworm Nereis virens (Polychaeta) in conservation areas. Aquatic Conservation: Marine and Freshwater Ecosystems, 3 (1), 1–24.
http://dx.doi.org/10.1002/aqc.3270030102
Olive, P.J.W. (1994) Polychaeta as world resource: a review of patterns of exploitation as sea angling baits and the potential for aquaculture based production. In: Dauvin, J.C., Laubier, L. & Reish, D.J. (Eds.), Actes de la 4ème Conférence internationale des Polychètes. Mémoires du Muséum National d'Histoire Naturelle, Angers, pp. 603–610.
Olive, P.J.W., Fletcher, J., Rees, S.W. & Desrosiers, G. (1997) Interactions of environmental temperature with photoperiod in determining age at maturity in a semelparous polychaete Nereis (Neanthes) virens Sars. Journal of Thermal Biology, 22 (6), 489–497.
http://dx.doi.org/10.1016/S0306-4565(97)00067-3
Olive, P.J.W., Rees, S.W. & Djunaedi, A. (1998) Influence of photoperiod and temperature on oocyte growth in the semelparous polychaete Nereis (Neanthes) virens. Marine Ecology Progress Series, 172, 169–183.
http://dx.doi.org/10.3354/meps172169
Oug, E., Bakken, T. & Kongsrud, J.A. (2014) Original specimens and type localities of early described polychaete species (Annelida) from Norway, with particular attention to species described by O.F. Müller and M. Sars. Memoirs of Museum Victoria, 71, 217–236.
http://dx.doi.org/10.24199/j.mmv.2014.71.17
Pamungkas, J. & Glasby, C.J. (2015) Taxonomy of reproductive Nereididae (Annelida) in multispecies swarms at Ambon Island, Indonesia. ZooKeys, 520, 1–25.
http://dx.doi.org/10.3897/zookeys.520.9581
Pettibone, M.H. (1956) Some polychaete worms of the families Hesionidae, Syllidae and Nereidae from the east coast of North America, West Indies, and Gulf of Mexico. Journal of the Washington Academy of Sciences, 46 (9), 281–294.
Pettibone, M.H. (1963) Marine polychaete worms of the New England region. 1. Aphroditidae through Trochochaetidae. Bulletin of the United States National Museum, 227, 1–356.
Rasmussen, E. (1973) Systematics and ecology of the Isefjord marine fauna (Denmark). Ophelia, 11 (1), 1–507.
http://dx.doi.org/10.1080/00785326.1973.10430115
Rathbun, R. (1884) Descriptive catalogue of the collection illustrating the scientific investigation of the sea and fresh waters. In: Goode, B.G. (Ed.), Descriptive catalogues of the collections sent from the United States to the International Fisheries Exhibition, London, 1883. Government Printing Office, Washington, D.C., 109 pp.
Read, G.B. (2007) Taxonomy of sympatric species of New Zealand Platynereis, with description of three new species additional to P. australis (Schmarda) (Annelida: Polychaeta: Nereididae). Zootaxa, 1558, 1–28.
http://dx.doi.org/10.11646/zootaxa.1558.1.1
Ricketts, E.F, Calvin, J. & Hedgpeth, J.W. (1985) Between Pacific Tides, 5th ed. Standford University Press, Standford, 680 pp.
Rostov, I.D., Yurasov, G.I., Rudyh, V.V., Moroz, V.V., Dmitrieva, E.V, Rostov, V.I., Nabiullin, A.A., Khrapchenkov, F.F. & Bunin, V.M. (2003) Oceanographic atlas of the Bering Sea, Sea of Okhotsk and Japan/East Sea. V. I. Il’ichev Pacific Oceanographic Institute, Vladivostok. Available from: http://pacificinfo.ru/data/cdrom/2/HTML/e_1_00.htm (Accessed 25 January 2018)
Ruppert, E.E. & Barnes, R.D. (1994) Invertebrate zoology, 6th ed. Saunders College Publishing, Fort Worth, USA, 12 +1056 pp.
Santos, C.S.G., Pleijel, F., Lana, P. & Rouse, G.W. (2006) Phylogenetic relationships within Nereididae (Annelida: Phyllodocida). Invertebrate Systematics, 19, 557–576.
http://dx.doi.org/10.1071/IS05001
Sars, M. (1835) Beskrivelser og Iagttagelser over nogle moerkelige 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 Forfatter. Bergen. Trykt paa Thorstein Hallagers Forlag hos Chr. Dahl, R. S. xii + 81 pp., 15 pls.
Sato, M. (2013) Resurrection of the genus Nectoneanthes Imajima, 1972 (Nereididae: Polychaeta), with redescription of Nectoneanthes oxypoda (Marenzeller, 1879) and description of a new species, comparing them to Neanthes succinea (Leuckart, 1847). Journal of Natural History, 47, 1–50.
http://dx.doi.org/10.1080/00222933.2012.743609
Sato, M. (2017) Nereididae (Annelida) in Japan, with special reference to life-history differentiation among estuarine species. In: Motokawa, M. & Kajihara, H. (Eds.), Species Diversity of Animals in Japan. Springer Japan, pp. 477–512.
http://dx.doi.org/10.1007/978-4-431-56432-4_19
Sato, M. & Nakashima, A. (2003) A review of Asian Hediste species complex (Nereididae, Polychaeta) with descriptions of two new species and a redescription of Hediste japonica (Izuka, 1908). Zoological Journal of the Linnean Society, 137, 403–445.
http://dx.doi.org/10.1046/j.1096-3642.2003.00059.x
Schroeder, P.C. & Hermans, C.O. (1975) Annelida: Polychaeta. In: Giese, A.C. & Pearse, J.S. (Eds.), Reproduction of marine invertebrates. Academic Press, London, pp. 1–213.
Snow, D.R. (1972) Some aspects of the life history of the nereid worm Nereis virens (Sars), on an intertidal mudflat at Brandy Cove, St. Andrews, N.B. M.Sc. thesis dissertation, McGill University, Montreal, 161 pp.
Snow, D.R. & Marsden, J.R. (1974) Life cycle, weight and possible age distribution in a population of Nereis virens (Sars) from New Brunswick. Journal of Natural History, 8 (5), 513–527.
http://dx.doi.org/10.1080/00222937400770431
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, 3 pls.
Stasek, C.R. (1966) Type specimens in the California Academy of Sciences, Department of Invertebrate Zoology. Occasional Papers of the California Academy of Sciences, 51, 1–38.
Sukhova N.G. & Tammiksaar, E. (2015) Aleksandr Fyodorovich Middendorff: on the bicentenary of his birth. 2nd edition. Nestor-History, Saint-Petersburg, 384 pp. [In Russian]
Sveshnikov, V.A. (1978) The morphology of polychaete larvae. Academy of Sciences of the USSR, Moscow, 152 pp. [In Russian]
Tammiksaar, E. & Stone, I.R. (2007) Alexander von Middendorff and his expedition to Siberia (1842–1845). Polar Record, 43 (226), 193–216.
http://dx.doi.org/10.1017/S0032247407006407
Toomsalu, M. (2016) Alexander Theodor von Middendorff – Doctor, explorer and promoter of cattle-breeding. Papers on Anthropology, 25 (2), 77–86.
http://dx.doi.org/10.12697/poa.2016.25.2.08
Treadwell, A.L. (1914) Polychaetous annelids of the Pacific Coast in the collections of the Zoological Museum of the University of California. University of California Publications in Zoology, 13 (8), 175–234.
Treadwell, A.L. (1939) New polychaetous annelids from New England, Texas and Puerto Rico. American Museum Novitates, 1023, 1–7.
Treadwell, A.L. (1941) Polychaetous annelids from the New England region, Porto Rico and Brazil. American Museum Novitates, 1138, 1–4.
Turnbull, F.M. (1876) On the anatomy and habits of Nereis virens. Transactions of the Connecticut Academy of Arts and Sciences, 3, 265–281.
Uschakov, P.V. (1950) Polychaetes from the Sea of Okhotsk. Studies of the Far Eastern Seas of the USSR, 2, 140–236. [In Russian]
Uschakov, P.V. (1955) Polychaete worms from the Far Eastern seas of the USSR. Academy of Sciences of the USSR, Moscow, 445 pp.
Ushakova, O.O. & Sarantchova, O.L. (2004) The influence of salinity on fertilization and larval development of Nereis virens (Polychaeta, Nereidae) from the White Sea. Journal of Experimental Marine Biology and Ecology, 301 (2), 129–139.
http://dx.doi.org/10.1016/j.jembe.2003.09.025
Villalobos-Guerrero, T.F. & Carrera-Parra, L.F. (2015) Redescription of Alitta succinea (Leuckart, 1847) and reinstatement of A. acutifolia (Ehlers, 1901) n. comb. based upon morphological and molecular data (Polychaeta: Nereididae). Zootaxa, 3919 (1): 157–178.
http://dx.doi.org/10.11646/zootaxa.3919.1.7
Vinogradov, K.A. (1948) On the question of the use of polychaetes as food for fish. Reports of the Academy of Sciences of the USSR, 60 (7), 1273–1276. [In Russian]
Vinogradov, K.A. (1950) On the finding of Nereis virens worms in the stomachs of Kamchatka red salmon and some other fishes. Nature, 3, 69. [In Russian]
von Marenzeller, E. (1879) Südjapanische Anneliden, 1 Amphinomea, Aphroditea, Lycoridea, Phyllodocea, Hesionea, Syllidea, Eunicea, Glycerea, Sternaspidea, Chaetopterea, Cirratulea, Amphictenea. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe classe der Kaiserlichen Akademie der Wissenschaften, 41, 109–154.
Watson, G.J., Murray, J.M., Schaefer, M. & Bonner, A. (2017) Bait worms: a valuable and important fishery with implications for fisheries and conservation management. Fish and Fisheries, 18 (2), 374–388.
http://dx.doi.org/10.1111/faf.12178
Yokouchi, K. (1985) Reproduction and larval ecology of the sandworm Neanthes virens Sars from southern Hokkaido. Bulletin of the Plankton Society of Japan, 32, 1–13.