Four new species and further records of Dorvilleidae (Annelida, Polychaeta) from deep-sea organic substrata, NE Atlantic

  • Ascensão Ravara CESAM - Centro de Estudos do Ambiente e do Mar, Departamento de Biologia, Universidade de Aveiro, Campus de Santiago, 3810-193 Aveiro, Portugal
  • Helena Wiklund Department of Marine Sciences, University of Gothenburg, Carl Skottbergsgata 22B, 413 19 Gothenburg, Sweden
  • Marina R. Cunha CESAM - Centro de Estudos do Ambiente e do Mar, Departamento de Biologia, Universidade de Aveiro, Campus de Santiago, 3810-193 Aveiro, Portugal
Keywords: Ophryotrocha, Parougia, mud volcanoes, Gulf of Cadiz, Estremadura Spur


Eight species of Ophryotrocha and one of Parougia were identified from organic substrata (wood and alfalfa) sampled at the Gulf of Cadiz and Western Iberian Margin (NE Atlantic). Morphological examination and molecular phylogenetic analyses, based on the nuclear gene H3 and the mitochondrial gene 16S, indicate the presence of four species new to science: Ophryotrocha chemecoli sp. nov., O. nunezi sp. nov., O. geoffreadi sp. nov. and Parougia ougi sp. nov. The geographic and/or bathymetric distribution is extended for four previously known species: O. cantabrica, O. hartmanni, O. mammillata and O. scutellus. Another species may also have its distribution extended, pending the molecular confirmation of its identity: O. lipscombae. Full descriptions and figures are given for all the new species and, when justified, also for the previously known ones. The original description of O. scutellus is amended. New DNA sequences are given for eight of the nine species studied here. The ecology and geographic distribution of the reported species is discussed.


Åkesson B. 1973. Reproduction and larval morphology of five Ophryotrocha species (Polychaeta, Dorvilleidae). Zoologica Scripta 2: 145–155.

Åkesson B. & Rice S.A. 1992. Two new Dorvillea species (Polychaeta, Dorvilleidae) with obligate asexual reproduction. Zoologica Scripta 21 (4): 351–362.

Bennett B.A., Smith C.R., Glaser B. & Maybaum H.L. 1994. Faunal community structure of a chemoautotrophic assemblage on whale bones in the deep northeast Pacific Ocean. Marine Ecology Progress Series 108: 205–223.

Blake J.A. 1985. Polychaeta from the vicinity of deep-sea geothermal vents in the eastern Pacific. I: Euphrosinidae, Phyllodocidae, Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae and Maldanidae. Bulletin of the Biological Society of Washington 6: 67–101. Available from [accessed 27 May 2020].

Blake J.A. & Hilbig B. 1990. Polychaeta from the vicinity of deep-sea hydrothermal vents in the Eastern Pacific Ocean. II. New species and records from the Juan de Fuca and Explorer Ridge systems. Pacific Science 44 (3): 219–253. Available from [accessed 27 May 2020].

Brito M.C. & Nuñez J. 2003. Three new interstitial dorvilleids (Annelida: Polychaeta) from the Cymodocea nodosa meadows of the Canary Islands. Hydrobiologia 496 (1): 27–34.

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: e22232.

Chamberlin R.V. 1919. The Annelida Polychaeta [Albatross Expeditions]. Memoirs of the Museum of Comparative Zoology at Harvard College 48: 1–514. Available from [accessed 27 May 2020].

Claparède E. & Mecznikow E. 1869. Beiträge zur Kenntnis der Entwicklungsgeschichte der Chaetopoden. Zeitschrift für wissenschaftliche Zoologie 19: 163–205. Available from [accessed 27 May 2020].

Colgan D.J., Ponder W.F. & Eggler P.E. 2000. Gastropod evolutionary rates and phylogenetic relationships assessed using partial 28S rDNA and histone H3 sequences. Zoologica Scripta 29 (1): 29–63.

Cunha M.R., Matos F.L., Génio L., Hilário A., Moura C.J., Ravara A. & Rodrigues C.F. 2013. Are organic falls bridging reduced environments in the deep sea? - Results from colonization experiments in the Gulf of Cádiz. PLoS One 8 (10): e76688.

Dahlgren T., Åkesson B., Schander C., Halanych K.M. & Sundberg P. 2001. Molecular phylogeny of the model annelid Ophryotrocha. The Biological Bulletin 201: 193–203.

Danise S., Dominici S., Glover A.G. & Dahlgren T.G. 2014. Molluscs from a shallow-water whale-fall and their affinities with adjacent benthic communities on the Swedish west coast. Marine Biology Research 10: 3–16.

Eilertsen M.H., Kongsrud J. A., Alvestad T., Stiller J., Rouse G.W. & Rapp H.T. 2017. Do ampharetids take sedimented steps between vents and seeps? Phylogeny and habitat-use of Ampharetidae (Annelida, Terebelliformia) in chemosynthesis-based ecosystems. BMC Evolutionary Biology 17: e222.

Goffredi S.K., Johnson S., Tunnicliffe V., Caress D., Clague D., Escobar E., Lundsten L., Paduan J.B., Rouse G.W., Salcedo D.L., Soto L.A., Spelz-Madero R., Zierenberg R. & Vrijenhoek R. 2017. Hydrothermal vent fields discovered in the southern Gulf of California clarify role of habitat in augmenting regional diversity. Proceedings of the Royal Society B. Biological Sciences 284: 20170817.

Hilbig B. & Blake J.A. 1991. Dorvilleidae (Annelida: Polychaeta) from the U.S. Atlantic slope and rise. Description of two new genera and 14 new species, with generic revision of Ophryotrocha. Zoologica Scripta 20 (2): 147–183.

Hilbig B. & Fiege D. 2001. A new species of Dorvilleidae (Annelida: Polychaeta) from a cold seep site in the northeast Pacific. Proceedings of the Biological Society of Washington 114 (2): 396–402. Available from [accessed 27 May 2020].

Huth W. 1933. Ophryotrocha-Studien. I. Zur Cytologie der Ophryotrochen. Zeitschrift für Zellforschung und mikroskopische Anatomie, Berlin 20: 309–381.

Josefson A.B. 1975. Ophryotrocha longidentata sp.n. and Dorvillea erucaeformis (Malmgren) (Polychaeta, Dorvilleidae) from the west coast of Scandinavia. Zoologica Scripta 4 (2–3): 49–54.

Jumars P.A. 1974. A generic revision of the Dorvilleidae (Polychaeta), with six new species from the deep Pacific. Zoological Journal of the Linnean Society, London 54 (2): 101–135. Available from [accessed 27 May 2020].

Katoh K., Misawa K., Kuma K.I. & Miyata T. 2002. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research 30 (14): 3059–3066. Available from [accessed 27 May 2020].

Kearse M., Moir R., Wilson A., Stones-Havas S., Cheung M., Sturrock S., Buxton S., Cooper A., Markowitz S., Duran C., Thierer T., Ashton B., Mentjies P. & Drummond A. 2012. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28 (12): 1647–1649.

Levin L.A., Ziebis W., Mendoza G.F., Growney V.A., Tryon M.D., Brown K.M., Mahn C., Gieskes J.M. & Rathburn A.E. 2003. Spatial heterogeneity of macrofauna at northern California methane seeps: Influence of sulfide concentration and fluid flow. Marine Ecology Progress Series 265: 123–139.

Levin L.A., Ziebis W., Mendoza G.F., Bertics V.J., Washington T., Gonzalez J., Thurber A.R., Ebbe B. & Lee R.W. 2013. Ecological release and niche partioning under stress: lessons from dorvilleid polychaetes in sulfidic sediments at methane seeps. Deep-Sea Research II 92: 214–233.

Ligges U. & Mächler M. 2003. Scatterplot3d – an R Package for Visualizing Multivariate Data. Journal of Statistical Software 8 (11): 1–20.

Lu H. & Fauchald K. 2000. Ophryotrocha lipscombae, a new species and a possible connection between ctenognath and labidognath-prionognath eunicean worms (Polychaeta). Proceedings of the Biological Society of Washington 113 (2): 486–492. Available from [accessed 11 Feb. 2021].

Marenzeller E. 1902. Südjapanische Anneliden. 3. Aphroditea, Eunicea. Denkschriften der Akademie der Wissenschaften, Wien 72: 563–582. Available from [accessed 27 May 2020].

Mercier A.J., Bailon S. & Hamel J.-F. 2014. Life history and seasonal breeding of the deep-sea annelid Ophryotrocha sp. (Polychaeta: Dorvilleidae). Deep-Sea Research I 91: 27–35.

Miura T. 1997. Two new species of the genus Ophryotrocha (Polychaeta, Iphitimidae) from Kagoshima Bay. Bulletin of Marine Science 60 (2): 300–305.

Núñez J., Riera R. & Maggio Y. 2014. A new Ophryotrocha species (Polychaeta: Dorvilleidae) from circalittoral seabeds of the Cantabrian Sea (north-east Atlantic Ocean). Journal of the Marine Biological Association of the United Kingdom 94 (1): 115–119.

Oug E. 1978. New and lesser known Dorvilleidae (Annelida, Polychaeta) from Scandinavian and northeast American waters. Sarsia 63 (4): 285–303.

Oug E. & Pleijel F. 2015. Dorvilleidae. In: Nygren A. & Pleijel F. (eds) Ringmaskar Havs-borstmaskar, Annelida: Polychaeta: Aciculata: 270–285. ArtDatabanken, SLU, Uppsala.

Palumbi S.R. 1996. Nucleic acids II: The polymerase chain reaction. In: Hillis D.M., Mable B.K. & Moritz C. (eds) Molecular Systematics: 205–247. Sinauer Associates, Sunderland, MA.

Parenti U. 1961. Ophryotrocha puerilis siberti, O. hartmanni and O. baccii nelle acque di Roscoff. Cahiers de Biologie marine 2: 437–445.

Paxton H. 2004. Jaw growth and replacement in Ophryotrocha labronica (Polychaeta, Dorvilleidae). Zoomorphology 123: 147–154.

Paxton H. 2009. Phylogeny of Eunicida (Annelida) based on morphology of jaws. Zoosymposia 2: 241–264.

Paxton H. & Åkesson B. 2007. Redescription of Ophryotrocha puerilis and O. labronica (Annelida, Dorvilleidae). Marine Biology Research 3: 3–19.

Paxton H. & Davey A. 2010. A new species of Ophryotrocha (Annelida: Dorvilleidae) associated with fish farming at Macquarie Harbour, Tasmania, Australia. Zootaxa 2509 (1): 53–61.

Paxton H. & Morineaux M. 2009. Three species of Dorvilleidae (Annelida: Polychaeta) associated with Atlantic deep-sea reducing habitats, with the description of Ophryotrocha fabriae, new species. Proceedings of the Biological Society of Washington 122 (1): 14–25.

Paxton H., Wiklund H., Alexander F. & Taboada S. 2017. Is the Antarctic Ophryotrocha orensanzi (Annelida: Dorvilleidae) a circumpolar non-specialized opportunist? Systematics and Biodiversity 15 (2): 105–114.

Pfannenstiel H.D. 1975. Ophryotrocha natans n. sp. (Polychaeta, Dorvilleidae): ein simultanzwitter mit acht männlichen segmenten aus dem Golf von Aqaba. Zoologischer Anzeiger 195 (1–2): 1–7.

Pleijel F., Jondelius U., Norlinder E., Nygren A., Oxelman B., Schander C., Sundberg P. & Thollesson M. 2008. Phylogenies without roots? A plea for the use of vouchers in molecular phylogenetic studies. Molecular Phylogenetics and Evolution 48: 369–371.

Posada D. 2008. jModelTest: Phylogenetic model averaging. Molecular Biology and Evolution 25: 1253–1256.

Prevedelli D., N’Siala G.M. & Simonini R. 2005. The seasonal dynamics of six species of Dorvilleidae (Polychaeta) in the harbour of La Spezia (Italy). Marine Ecology 26: 286–293.

Ravara A., Marçal A.R., Wiklund H. & Hilário A. 2015. First account on the diversity of Ophryotrocha (Annelida, Dorvilleidae) from a mammal-fall in the deep-Atlantic Ocean with the description of three new species. Systematics and Biodiversity 13 (6): 555–570.

R Core Team 2020. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from [accessed 11 Feb. 2021].

Read G. & Fauchald K. (eds) (2020). World Polychaeta database. Parougia Wolf, 1986. Accessed through World Register of Marine Species. Available from [accessed 12 Nov. 2018].

Ronquist F., Teslenko M., van der Mark P., Ayres D.L., Darling A., Höhna S., Larget B., Liu L., Suchard M.A. & Huelsenbeck J.P. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542.

Salvo F., Wiklund H., Dufour S.C., Hamoutene D., Pohle G. & Worsaae K. 2014. A new annelid species from whalebones in Greenland and aquacultire sites in Newfoundland: Ophryotrocha cyclops, sp. nov. (Eunicida: Dorvilleidae). Zootaxa 3887 (5): 555–568.

Sjölin E., Erséus C. & Källersjö M. 2005. Phylogeny of Tubificidae (Annelida, Clitellata) based on mitochondrial and nuclear sequence data. Molecular Phylogenetics and Evolution 35: 431–441.

Smith C.R. 2006. Bigger is better: the role of whales as detritus in marine ecosystems. In: Estes J.A., DeMaster D.P., Brownell R.L. Jr, Doak D.F. & Williams T.M. (eds) Whales, Whaling and Ocean Ecosystems: 286–301. University of California Press, Berkeley, CA.

Struck T.H., Purschke G. & Halanych K.M. 2006. Phylogeny of Eunicida (Annelida) and exploring data congruence using a partition addition bootstrap alteration (PABA) approach. Systematic Biology 55(1): 1–20.

Taboada S., Wiklund H., Glover A.G., Dahlgren T.G., Cristobo J. & Avila C. 2013. Two new Antarctic Ophryotrocha (Annelida: Dorvilleidae) described from shallow-water whale bones. Polar Biology 36: 1031–1045.

Taboada S., Bas M. & Avila C. 2015. A new Parougia species (Annelida, Dorvilleidae) associated with eutrophic marine habitats in Antarctica. Polar Biology 38 (4): 517–527.

Taboada S., Bas M., Leiva C., Garriga M., Sardá R. & Avila C. 2016. Life after death: shallow-water Mediterranean invertebrate communities associated with mammal bones. Marine Ecology 37: 164–178.

Taboada S., Leiva C., Bas M., Schult N. & McHugh D. 2017. Cryptic species and colonization processes in Ophryotrocha (Annelida, Dorvilleida) inhabiting vertebrate remains in the shallow-water Mediterranean. Zoologica Scripta 46 (5): 611–624.

Thornhill D.J., Struck T.H., Ebbe B., Lee R.W., Mendoza G.F., Levin L.A. & Halanych K.M. 2012. Adaptative radiation in extremophilic Dorvilleidae (Annelida): diversification of a single colonizer or multiple independent lineages? Ecology and Evolution 2 (8): 1958–1970.

Vedenin A.A., Kokarev V.N., Chikina M.V., Basin A.B., Galkin S.V. & Gebruk A.V. 2020. Fauna associated with shallow-water methane seeps in the Laptev Sea. PeerJ 8: e9018.

Voight J.R. 2015. Xylotrophic bivalves: aspects of their biology and the impacts of humans. Journal of Molluscan Studies 81 (2): 175–186.

Webster H.E. & Benedict J.E. 1884. The Annelida Chaetopoda from Provincetown and Wellfleet, Massachusetts. Annual Report of the United States Commission of Fish and Fisheries, Washington. 1881: 699–747.

Wiklund H., Glover A.G. & Dahlgren T.G. 2009. Three new species of Ophryotrocha (Annelida: Dorvilleidae) from a whale-fall in the North-East Atlantic. Zootaxa 2228 (1): 43–56.

Wiklund H., Altamira I.V., Glover A.G., Smith C.R., Baco A.R. & Dahlgren T.G. 2012. Systematics and biodiversity of Ophryotrocha (Annelida, Dorvilleidae) with descriptions of six new species from deep-sea whale-fall and wood-fall habitats in the north-east Pacific. Systematics and Biodiversity 10 (2): 243–259.

Wolf P.S. 1986. Three new species of Dorvilleidae (Annelida: Polychaeta) from Puerto Rico and Florida and a new genus for dorvilleids from Scandinavia and North America. Proceedings of The Biological Society of Washington 99: 627–638. Available from [accessed 11 Feb. 2021].

Yen N.K. & Rouse G.W. 2020. Phylogeny, biogeography and systematics of Pacific vent, methane seep, and whale-fall Parougia (Dorvilleidae: Annelida), with eight new species. Invertebrate Systematics 34 (2): 200–233.

Zhang D.-S., Zhou Y.-D., Wang C.-S. & Rouse G. 2017. A new species of Ophryotrocha (Annelida, Eunicida, Dorvilleidae) from hydrothermal vents on the Southwest Indian Ridge. ZooKeys 687: 1–9.

How to Cite
Ravara, A. ., Wiklund, H., & Cunha, M. R. (2021). Four new species and further records of Dorvilleidae (Annelida, Polychaeta) from deep-sea organic substrata, NE Atlantic. European Journal of Taxonomy, 736(1), 44-81.
Research article