Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2018-09-05
Page range: 245–278
Abstract views: 268
PDF downloaded: 4

New species of Pseudonereis Kinberg, 1865 (Polychaeta: Nereididae) from the Atlantic Ocean, and a review of paragnath morphology and methodology

El Colegio de la Frontera Sur, Unidad Chetumal, Departamento de Sistemática y Ecología Acuática, Av. Centenario km 5.5, Chetumal, Quintana Roo, 77014, Mexico.
Annelida amphiamerican nereidid paragnath pharyngeal arrangement pharynx


Pseudonereis gallapagensis Kinberg, 1865 and P. variegata (Grube & Kröyer in Grube, 1858) are the only two species of this genus commonly recorded along Atlantic American coasts, but their type localities are in the Eastern Pacific, and their morphology differs. Two new Pseudonereis species are described from Eastern Mexico: P. brunnea sp. n. from the Gulf of Mexico, and P. citrina sp. n. from the Caribbean Sea, previously confused with P. gallapagensis. In order to facilitate comparisons, descriptions based on specimens from near the type locality for P. gallapagensis (Peru and Ecuador), and topotypes for P. variegata (Valparaiso, Chile), are included. Based on these comparisons and current descriptions, the synonymies of Nereis ferox Hansen, 1882 described from Brazil with P. variegata, and of Pseudonereis formosa Kinberg, 1865 described from Hawaii with P. gallapagensis, are rejected. Consequently, both are regarded as distinct species and revised diagnoses are provided for them. The record of P. ferox from the Gulf of Guinea proved to be an undescribed species, and is herein described as P. fauveli sp. n. The number of paragnath rows in nereidid pharynx areas VII–VIII has been interpreted in several ways, leading to confusion; an alternative method to determine the number of bands and rows is proposed. The midventral region, the division of areas VII–VIII in furrow and ridge regions, and the description of the arrangement based on the pattern of paragnaths in such regions, are proposed. Further, the terms shield-shaped and pointed (P-bars) bars are redefined, and a new term, crescent-shaped bars, is proposed for paragnaths in the areas VI in some Pseudonereis and Perinereis species. A key for all Pseudonereis species is also included.



  1. Augener, H. (1918) Polychaeta. In: Michaelsen, W. (Ed.), Beiträge zur kenntnis der Meeresfauna Westafrikas. L Friederichsen & Co., Hamburg, pp. 67–625.

    Augener, H. (1927) Polychaeten von Curaçao. Bijdragen tot de Dierkunde, 25 (1), 39–82.

    Augener, H. (1934) Polychaeten aus den zoologischen Museen von Leiden und Amsterdam. IV (schluss). Zoologische Mededeelingen, 17 (8), 67–160.

    Bakken, T. (2007) Revision of Pseudonereis (Polychaeta, Nereididae). Zoological Journal of the Linnean Society, 150, 145–176.

    Bakken, T. & Wilson, R.S. (2005) Phylogeny of nereidids (Polychaeta, Nereididae) with paragnaths. Zoologica Scripta, 34 (5), 507–547.

    Bakken, T., Glasby, C.J. & Wilson, R.S. (2009) A review of paragnath morphology in Nereididae (Polychaeta). Zoosymposia, 2, 305–316.

    Chamberlin, R.V. (1919) The Annelida Polychaeta. Plates. Memoirs of the Museum of Comparative Zoology at Harvard College, 48, 1–80.

    Chambers, S. & Garwood, P. (1992) Polychaetes from Scottish waters. A guide to identification. 3. Family Nereidae. National Museums of Scotland, Edinburgh, 65 pp.

    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.

    Day, J.H. (1967) A monograph on the polychaeta of Southern Africa. Part I. Errantia. Trustees of the British Museum (Natural History), London, 878 pp.

    Darbyshire, T. (2014) Intertidal and nearshore Nereididae (Annelida) of the Falkland Islands, southwestern Atlantic, including a new species of Gymnonereis. ZooKeys, 427, 75–108.

    de León-González, J.Á. (1997) Poliquetos de la familia Nereidae (Annelida: Polychaeta) de los litorales Mexicanos: Sistemática, Biogeografía y alimentación. Tesis doctoral. Facultad de Ciencias Biológicas, Universidad Autónoma de Nuevo León. San Nicolás de los Garza Nuevo León, 298 pp.

    de León-González, J.Á. (2009) Nereididae Lamarck, 1818. In: de León-González, J.Á., Bastida-Zavala, J.R., Carrera-Parra, L.F., García-Garza, M.E., Peña-Rivera, A., Salazar-Vallejo, S.I. & Solís-Weiss, V. (Eds.), Monterrey, México, pp. 325–354.

    de León-González, J.Á. & Solís-Weiss, V. (1998) The genus Perinereis (Polychaeta: Nereididae) from Mexican littoral waters, including the description of three new species and the redescriptions of P. anderssoni and P. elenacasoae. Proceedings of the Biological Society of Washington, 111 (3), 674–693.

    de León-González, J.Á., Solís-Weiss, V. & Ochoa-Rivera, V. (1999) Nereidids (Polychaeta) from the Caribbean Sea and adjacent coral islands of the southern Gulf of Mexico. Proceedings of the Biological Society of Washington, 112 (4), 667–681.

    de León-González, J.Á., Solís-Weiss, V. & Valadez-Rocha, V. (2001) Two new species of Platynereis (Polychaeta: Nereididae) from Eastern Mexican shores. Proceedings of the Biological Society of Washington, 114 (2), 389–395.

    Dean, H.K. (2012) A literature review of the Polychaeta of the Caribbean Sea. Zootaxa, 3596, 1–86.

    Díaz-Díaz, Ó.F. & Liñero-Arana, I. (2002) Poliquetos asociados a substratos artificiales sumergidos en la costa nororiental de Venezuela IV. Phyllodocida (Annelida: Polychaeta). Boletín del Instituto Oceanográfico de Venezuela, 41, 25–37.

    Dueñas-Ramírez, P.R. & Quiros-Rodríguez, J.A. (2012) Ocurrence of Pseudonereis gallapagensis Kinberg, 1865 (Annelida: Polychaeta) in the Colombian Caribbean coast. Revista Colombiana de Ciencia Animal, 4 (2), 454–457.

    Ehlers, E. (1901) Die Polychaeten des magellanischen und chilenischen Strandes. Ein faunistischer Versuch. Abhandlungen der Mathematisch-Physikalischen Klasse, 1901, 1–232.

    Fauchald, K. (1977a) Polychaetes from intertidal areas in Panama, with a review of previous shallow-water records. Smithsonian Contributions to Zoology, 221, 1–81.

    Fauchald, K. (1977b) The polychaete worms: Definitions and keys to the orders, families and genera. Natural History Museum of Los Angeles County, Science Series, 28, 1–190.

    Fauchald, K. & Reimer, A.A. (1975) Clave de poliquetos panameños con la inclusión de una clave para todas las familias del mundo. Boletín del Instituto Oceanográfico de la Universidad de Oriente, 14 (1), 71–94.

    Fauvel, P. (1914) Annélides polychètes de San-Thomé (Golfe de Guinée) recueillies par M. Ch. Gravier. Archives de Zoologie Expérimentale et Générale, 54, 105–155.

    Fauvel, P. (1932) Annelida polychaeta of the Indian Museum, Calcutta. Memoirs of the Indian Museum, 12 (1), 1–261.

    Glasby, C.J., Wei, N.-W.V. & Gibb, K.S. (2013) Cryptic species of Nereididae (Annelida: Polychaeta) on Australian coral reefs. Invertebrate Systematics, 27, 245–264.

    Gravier, C. (1909) Annélides polychètes recueillis a Payta (Pérou) par M. le Dr Rivet, membre de la Mission géodésique de l'Équateur. Archives de Zoologie Expérimentale et Générale, 4e Série, 10, 617–659.

    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 den Naturhistorisk Forening for 1857, 1858, 158–186.

    Grube, A.E. (1859) 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, (Slutning [conclusion]) [3. Familia Ariciaea - F. Sipunculacea and Index]. Videnskabelige Meddelelser fra den Naturhistorisk Forening for 1858, 1859, 105–120.

    Grube, A.E. (1868) Anneliden. Reise der Oesterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858 and 1859. Zoologischer Theil, 2 (3), 1–44.

    Hansen, A. (1882) Recherches sur les Annélides Recueillies par M. le professeur Édouard Van Beneden pendant son Voyage au Brésil et à La Plata. Memoires Couronnes et Memoires des Savants Etrangers publies par L'Academie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique, 44 (3), 1–29.

    Hartman, O. (1938) Nomenclatorial changes involving types of polychaetous annelids of the family Nereidae in the United States National Museum. Journal of the Washington Academy of Sciences, 28, 13–15.

    Hartman, O. (1940) Polychaetous annelids. Part II. Chrysopetalidae to Goniadidae. Allan Hancock Pacific Expeditions, 7 (3), 173–287.

    Hartman, O. (1944) Polychaetous Annelids. Allan Hancock Atlantic Expedition, 3, 1–33.

    Hartman, O. (1948) The marine annelids erected by Kinberg with notes on some other types in the Swedish State Museum. Arkiv för Zoologi, 42A (1), 1–137.

    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 I. Allan Hancock Foundation Publications, Occasional Paper, 23 (1), 1–353.

    Hartmann-Schröder, G. (1962a) Zweiter Beitrag zur polychaetenfauna von Peru. Kieler Meeresforschungen, 18, 109–147.

    Hartmann-Schröder, G. (1962b) Zur Kenntnis der Nereiden Chiles (Polychaeta errantia), mit Beschreibung epitoker Stadien einiger Arten und der Jugendentwicklung von Perinereis vallata (Grube). Zoologischer Anzeiger, 168 (11–12), 389–441.

    Hartmann-Schröder, G. (1992) Zur Polychaetenfauna der Polynesischen Inseln Huahiné (Gesellschaftsinseln) und Rangiroa (Tuamotu-Inseln). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 89, 49–84.

    Horst, R. (1889) Note XXXIV. Contribution towards the knowlegde of the annelida polychaeta. On species of Nereis, belonging to the sub-genus Perinereis. Notes from the Leyden Museum, 11, 161–186.

    Ibárzabal, D.R. (1986) Lista de especies de poliquetos bentónicos cubanos. Reporte de Investigación del Instituto de Oceanología, 45, 1–17.

    Kinberg, J.G.H. (1865) Annulata Nova (Nereidum Dispositio Nova. Leonnatidea, Nereidea, Aretidea, Pisenoidea, Niconidea). Öfversigt af Kongelige Vetenskaps-Akademiens Förhandlingar, 22 (2), 167–179.

    Kinberg, J.G.H. (1910) Kongliga svenska fregatten Eugenies resa omkring jorden under befäl af C. A. Virgin. åren 1851-1853. Vetenskapliga iakttagelser. Zoologi. IV. Annulater. Almqvist & Wikssells, Uppsala & Stockholm, 46 pp. [pp. 33–78]

    Liñero-Arana, I. & Reyes-Vásquez, G. (1979) Nereidae (Polychaeta, Errantia) del Golfo de Cariaco, Venezuela. Boletín del Instituto Oceanográfico de la Universidad de Oriente, 18 (1–2), 3–12.

    Maggenti, M.A.B., Maggenti, A.R. & Gardner, S.L. (2008) Dictionary of Invertebrate Zoology. Zea E-Books, Lincoln, Nebraska, 379 pp.

    Monro, C.C.A. (1933) The Polychaeta Errantia collected by Dr. Crossland at Colón, in the Panama Region, and the Galapagos Islands during the expedition of the S.Y. “St. George.” Proceedings of the Zoological Society of London, 103 (1), 1–96.

    Núñez, J. (2004) Familia Nereididae. In: Ramos, M.Á. (Ed.), Fauna Ibérica. Museo Nacional de Ciencias Naturales, CSIC, Madrid, pp. 293–390.

    Perkins, T.H. & Savage, T. (1975) A bibliography and checklist of polychaetous anelids of Florida, the Gulf of Mexico, and the Caribbean Region. Florida Marine Research Publications, 14, 1–62.

    Read, G.B. (2007) Taxonomy of sympatric New Zealand species of Platynereis, with description of three new species additional to P. australis (Schmarda). Zootaxa, 1558, 1–28.

    Read, G. (2018) Pseudonereis variegata (Grube, 1857). In: Read, G. & Fauchald, K. (Eds.), World Polychaeta database. Available from: (accessed 24 July 2018)

    Ramírez-Hernández, A., Hernández-Alcántara, P. & Solís-Weiss, V. (2015) Nereis alacranensis, a new species of polychaete (Annelida, Nereididae) from Alacranes Reef, southern Gulf of Mexico, with a key to Nereis from the Grand Caribbean. Zootaxa, 4012 (1), 156–166.

    Rioja, E. (1960) Estudios anelidológicos. XXIV. Adiciones a la fauna de anélidos poliquetos de las costas orientales de México. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, 31, 289–316.

    Rozbaczylo, N. & Bolados, J. (1980) Nereidos de Iquique, Chile (Polychaeta: Nereidae). Boletín del Museo Nacional de Historia Natural, Chile, 37, 205–224.

    Salazar-Vallejo, S.I. (1996) Lista de especies y bibliografía de poliquetos (Polychaeta) del Gran Caribe. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, 67 (1), 11–50.

    Salazar-Vallejo, S.I. (2000) Biogeografía marina del Gran Caribe. Interciencia, 25 (1), 7–12.

    Salazar-Vallejo, S.I. & Jiménez-Cueto, M.S. (1997) Neréididos (Polychaeta) del Caribe mexicano con una clave para las especies del Gran Caribe. Revista de Biología Tropical, 44 (45), 361–377.

    Salazar-Vallejo, S.I. & Eibye-Jacobsen, D. (2012) Annulata örstediana: publication dates, composition and annotated taxonomic list, with some comments on Hemipodus (Polychaeta: Glyceridae). Revista de Biología Tropical, 60 (3), 1391–1402.

    Santos, C.S.G., Pleijel, F., Lana, P.D.C. & Rouse, G.W. (2005) Phylogenetic relationships within Nereididae (Annelida: Polychaeta). Invertebrate Systematics, 19 (6), 557–576.

    Schmarda, L.K. (1861) Neue wirbellose thiere beobachtet und gesammelt auf einer reise um die erde 1853 bis 1857. Erster Band. Turbellarian, Rotatorien un Anneliden. Zweite Hälfte. Wilhelm Engelmann, Leipzig, 164 pp.

    Stearn, W.T. (1992) Botanical Latin. 4th Edition. David & Charles, Devon, 546 pp.

    Stimpson, W. (1856) Description of some new marine invertebrates. Proceedings of the Academy of Natural Sciences of Philadelphia, 7 (10), 385–395.

    Treadwell, A.L. (1923) Two new species of polychaetous annelids of the genus Nereis from Brazil. Revista do Museu Paulista, 13, 1–10.

    Vanegas-Espinosa, V., Díaz-Díaz, Ó.F. & Liñero-Arana, I. (2007) Nereididae Lamarck, 1818 (Annelida: Polychaeta) de la Costa Occidental de Venezuela. Boletín del Instituto Oceanográfico de Venezuela, 46 (2), 119–127.

    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.

    Villalobos-Guerrero, T.F. & Tovar-Hernández, M.A. (2013) Una especie nueva de Pseudonereis (Polychaeta: Nereididae) de Mazatlán, golfo de California, incluyendo una clave para las especies del mundo. Revista Mexicana de Biodiversidad, 84 (3), 774–781.

    Willey, A. (1904) Littoral Polychaeta from the Cape of Good Hope. Transactions of the Linnean Society of London. 2nd Series: Zoology, 9 (6), 255–268.