Scientia Marina, Vol 81, No 3 (2017)

Sea pen (Pennatulacea) aggregations on the northern Spanish shelf: distribution and faunal assemblages

Susana Ruiz-Pico
Instituto Español de Oceanografía de Santander , Spain

Alberto Serrano
Instituto Español de Oceanografía de Santander , Spain

Antonio Punzón
Instituto Español de Oceanografía de Santander , Spain

Álvaro Altuna
INSUB, Museo de Okendo , Spain

Olaya Fernández-Zapico
Instituto Español de Oceanografía de Santander , Spain

Francisco Velasco
Instituto Español de Oceanografía de Santander , Spain


The sea pens Funiculina quadrangularis, Pennatula aculeata, Pennatula phosphorea, Pteroeides spinosum and Veretillum cynomorium are known to be characteristic species of the habitat ‘pennatulacean communities on circalittoral and upper bathyal soft bottoms’, described recently on the northern Atlantic Spanish shelf under the terms of the EU directives. Distribution, abundance and environmental data collected from 1995 to 2010 in the Northern Spanish Shelf Groundfish Surveys in the Cantabrian Sea and off Galicia (SPNGFS) were analysed. Although low densities of sea pens were commonly found over the years, some aggregations were shown. Multivariate analysis separated the aggregations of F. quadrangularis and of Pennatula spp. found on circalittoral soft bottoms. In addition, evidence of aggregations of F. quadrangularis on upper bathyal sites was also found. Depth and longitude, which were closely related to organic matter and sediment particle size, were key factors in determining the distribution and abundance of these aggregations. However, very little information on distribution and density of the species P. spinosum and V. cynomorium was obtained. Details of community composition of F. quadrangularis and Pennatula spp. were studied and compared with those of the OSPAR habitat “sea pen and burrowing megafauna communities” in other areas. The results improve our knowledge concerning sea pen communities in the southern Bay of Biscay and contribute to a global assessment of the status of these communities in the northeast Atlantic.


sea pens; southern Bay of Biscay; abundance; distribution; habitat

Full Text:



Aguilar R., Torriente A., García S. 2009. Propuesta de Áreas Marinas de Importancia Ecológica. Zona galaico-cantábrica. Oceana-Fundación Biodiversidad, 252 pp.

Altuna Á. 2010. Listado de los cnidarios (Cnidaria) bentónicos del Golfo de Vizcaya y zonas próximas (Atlántico NE). Proyecto Fauna Ibérica. Museo Nacional de Ciencias Naturales, Madrid, 27 pp.

Baillon S., Hamel J.F., Wareham V.E., et al. 2012. Deep cold-water corals as nurseries for fish larvae. Front. Ecol. Environ. 10: 351-356.

Baillon S., Hamel J.F., Mercier A. 2014. Diversity, distribution and nature of faunal associations with deep-sea pennatulacean corals in the Northwest Atlantic. PloS ONE 9: e111519. PMid:25369515 PMCid:PMC4219758

Birkeland C. 1974. Interactions between a sea pen and seven of its predators. Ecol. Monogr. 44: 211-232.

Blanchet F.G., Legendre P., Borcard D. 2008. Forward selection of explanatory variables. Ecology 89: 2623-2632. PMid:18831183

BOE. 2011. Real Decreto 1629/2011, de 14 de noviembre, por el que se declara como Área Marina Protegida y como Zona Especial de Conservación el espacio marino de El Cachucho, y se aprueban las correspondientes medidas de conservación. BOE 295: 130084-130138.

Borcard D., Gillet F., Legendre P. 2011. Numerical Ecology with R. Springer, New York.

Clarke K.R., Gorley R.N. 2006. PRIMER v6: User Manual/Tutorial. PRIMER-E, Plymouth.

Connor D.W., Allen J.H., Golding N., et al. 2004. The marine habitat classification for Britain and Ireland, version 04.05. JNCC, Peterborough.

de Juan S.D., Cartes J.E., Demestre M. 2007. Effects of commercial trawling activities in the diet of the flat fish Citharus linguatula (Osteichthyes: Pleuronectiformes) and the starfish Astropecten irregularis (Echinodermata: Asteroidea). J. Exp. Mar. Biol. Ecol. 349: 152-169.

de Moura Neves B., Edinger E., Layne G.D., et al. 2015. Decadal longevity and slow growth rates in the deep-water sea pen Halipteris finmarchica (Sars, 1851) (Octocorallia: Pennatulacea): implications for vulnerability and recovery from anthropogenic disturbance. Hydrobiologia 759: 147-170.

DFO (Fisheries and Oceans Canada). 2010. Occurrence, susceptibility to fishing and ecological function of corals, sponges and hydrothermal vents in Canadian waters. DFO Can. Sci. Advis. Sec. Sci. Advis. Rep. 2010/041.

EU (European Union). 2008. Directive 2008/56/EC of the European Parliament and of the Council of 17 June 2008, Establishing a Framework for Community Action in the Field of Marine Environmental Policy (Marine Strategy Framework Directive). Official Journal of the European Union L164: 19-40.

Fari-a A.C., Herraiz I.G. 2003. Trends in catch-per-unit-effort, stock biomass and recruitment in the North and Northwest Iberian Atlantic Nephrops stocks. Fish. Res. 65: 351-360.

Fernández J.A., Bonache J., Gordillo E. 2005. Parque Nacional das Illas Atlánticas de Galicia. Recursos rurais. Serie cursos 2: 91-102.

Gili J.M., Pagès F. 1987. Pennatuláceos (Cnidaria, Anthozoa) recolectados en la plataforma continental catalana (Mediterráneo occidental). Misc. Zool. 11: 25-39.

Greathead C.F., Donnan D.W., Mair J.M., et al. 2007. The sea pens Virgularia mirabilis, Pennatula phosphorea and Funiculina quadrangularis: distribution and conservation issues in Scottish waters. J. Mar. Biol. Assoc. U.K. 87: 1095-1103.

Greathead C., González-Irusta J.M., Clarke J., et al. 2015. Environmental requirements for three sea pen species: relevance to distribution and conservation. ICES J. Mar. Sci. 72: 576-586.

Hughes D.J. 1998. Sea pens and burrowing megafauna (volume III). An overview of dynamics and sensitivity characteristics for conservation management of marine SACs. Scott. Assoc. Mar. Sci. (UK Marine SACs Project). 105 pp.

ICES. 2010. Manual for the International Bottom Trawl Surveys in the Western and Southern Areas Revision III. Agreed during the meeting of the International Bottom Trawl Survey Working Group 22-26 March 2010, Lisbon. Addendum 2: ICES CM 2010/SSGESST: 06. 58 pp.

ICES. 2012. Report of the ICES Advisory Committee 2012. Book 7, Bay of Biscay and Atlantic Iberian waters. ICES Advice, 2012. Book 7. 205 pp.

IEO. 2014a. Caracterización ecológica del banco de Galicia. Informe final del proyecto LIFE+ INDEMARES (LIFE07/ NAT/E/000732), subproyecto LIC. IEO. Coordinación: Fundación Biodiversidad, Madrid, 237 pp.

IEO. 2014b. Caracterización ecológica del área marina del sistema de ca-ones submarinos de Avilés. Informe final área LIFE+INDEMARES (LIFE07/NAT/E/000732). IEO. Coordinación: Fundación Biodiversidad, Madrid, 243 pp.

Jouanneau J.M., Weber O., Champilou N., et al. 2008. Recent sedimentary study of the shelf of the Basque country. J. Mar. Sys. 72: 397-406.

Kaiser M.J., Edwards D.B., Armstrong P.J., et al. 1998. Changes in megafaunal benthic communities in different habitats after trawling disturbance. ICES J. Mar. Sci. 55: 353-361.

Kaiser M.J., Clarke K.R., Hinz H., et al. 2006. Global analysis and recovery of benthic biota to fishing. Mar. Ecol. Prog. Ser. 311: 1-14.

Kenchington E., Lirette C., Cogswell A., et al. 2010. Delineating Coral and Sponge Concentrations in the Biogeographic Regions of the East Coast of Canada Using Spatial Analyses. DFO Canadian Scientific Advisory Secretariat Research Document 2010/041. iv + 207 pp. PMid:20824204 PMCid:PMC2930843

Kenchington E., Murillo F.J., Cogswell A., et al. 2011. WG on the ecosystem approach to fisheries management Serial No. N6005 NAFO SCR Doc. 11/75.

Kenchington E., Murillo F.J., Lirette C., et al. 2014. Kernel Density Surface Modelling as a Means to Identify Significant Concentrations of Vulnerable Marine Ecosystem Indicators. PLoS ONE 9: e109365. PMid:25289667 PMCid:PMC4188592

Langton R.W., Langton E.W., Theroux R.B., et al. 1990. Distribution, behavior and abundance of sea pens, Pennatula aculeata, in the Gulf of Maine. Mar. Biol. 107: 463-469.

López-González P.J., Gili J.M., Williams G.C. 2001. New records of Pennatulacea (Anthozoa: Octocorallia) from the African Atlantic coast, with description of a new species and a zoogeographic analysis. Sci. Mar. 65: 59-74.

López-Jamar E., Cal R.M., González G., et al. 1992. Upwelling and outwelling effects on the benthic regime of the continental shelf off Galicia, NW Spain. J. Mar. Res. 50: 465-488.

MacDonald D.S., Little M., Eno N.C., et al. 1996. Disturbance of benthic species by fishing activities: a sensitivity index. Aquat. Conserv. 6: 257-268.<257::AID-AQC194>3.0.CO;2-7

MAGRAMA (Ministerio de Agricultura, Alimentación y Medio Ambiente). 2012. Estrategia Marina para la demarcación marina noratlántica, parte IV: descriptores del buen estado ambiental, descriptor 1: biodiversidad. Evaluación inicial y buen estado ambiental.

Manuel R.L. 1981. British Anthozoa. Synopses of the British Fauna (New Series). No 18. Linnean Society Academic Press, Lon don, 241 pp.

NOAA. 2014. Deep Sea Coral Research and Technology Program 2014 Report to Congress. NOAA, Silver Spring, MD. 54 pp.

Oca-a A., Sánchez-Tocino L., López-González P.J. 2000. Consideraciones faunística y biogeográficas de los antozoos (Cnidaria: Anthozoa) de la costa de Granada (Mar de Alborán). Zool. Baetica 11: 51-65.

Olaso I. 1990. Distribución y abundancia del megabentos invertebrado en fondos de la plataforma cantábrica. Ph.D. thesis, Inst. Esp. Oceanogr.

OSPAR. 2000. Quality Status Report 2000. Region IV- Bay of Biscay and Iberian coast. OSPAR Commission, London, 123 pp.

OSPAR. 2008. Case Reports for the OSPAR List of Threatened and/or Declining Species and Habitats. OSPAR Commission, London, 261 pp.

OSPAR. 2010. Background document for seapen and borrowing megafauna communities. OSPAR Commission, London, 27 pp.

Palmer M.W. 1993. Putting things in even better order: the advantages of canonical correspondence analysis. Ecology 74: 2215-2230.

Pirtle J.L. 2005. Habitat-based assessment of structure-forming megafaunal invertebrates and fishes on Cordell Bank, California. Ph.D. thesis, Univ. Washington, 74 pp.

Porporato E.M., Mangano M.C., De Domenico F., et al. 2014. First observation of Pteroeides spinosum (Anthozoa: Octocorallia) fields in a Sicilian coastal zone (Central Mediterranean Sea). Mar. Biodiv. 44: 589-592.

Punzón A., Hernández C., Abad E., et al. 2010. Spanish otter trawl fisheries in the Cantabrian Sea. ICES J. Mar. Sci. 67: 1-13.

R Development Team. 2014. R: a Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria.

Sánchez F., Olaso I. 2004. Effects of fisheries on the Cantabrian Sea shelf ecosystem. Ecol. Model. 172: 151-174.

Sánchez F., Serrano A., Gómez Ballesteros A.M. 2009. Photogrammetric quantitative study of habitat and benthic communities of deep Cantabrian Sea hard grounds. Cont. Shelf Res. 29: 1174-1188.

Sardá R., Rossi S., Martí X., et al. 2012. Marine benthic cartography of the Cap de Creus (NE Catalan Coast, Mediterranean Sea). Sci. Mar. 76: 159-171.

Serrano A., Sánchez F., García-Castrillo G. 2006. Epibenthic communities of trawlable grounds of the Cantabrian Sea. Sci. Mar. 70: 149-159.

Thrush S.F., Dayton K. 2002. Disturbance to Marine Benthic Habitats by trawling and Dredging: Implications for Marine Biodiversity. Ann. Rev. Ecol. Syst. 33: 449-473.

Tixier-Durivault A., d'Hondt M.J. 1975. Les Octocoralliaires de la campagne Biaçores. Bull. Mus. Natn. Hist. Nat. Paris, Zool. 174: 1361-1433.

Vergnon R., Blanchard F. 2006. Evaluation of trawling disturbance on macrobenthic invertebrate communities in the Bay of Biscay, France: Abundance Biomass Comparison (ABC method). Aquat. Living Resour. 19: 219-228.

Williams G.C. 1990. The Pennatulacea of Southern Africa (Coelenterata, Anthozoa). Ann. S. Afr. Mus. Cape Town 99: 31-119.

Williams G.C. 2011. The Global Diversity of Sea Pens (Cnidaria: Octocorallia: Pennatulacea). PloS ONE 6: e22747. PMid:21829500 PMCid:PMC3146507

Williams G.C. 2014. Pennatulacea. In: Costello M.J., Bouchet P., et al. 2014. European Register of Marine Species. Accessed on 2014-11-21 at

Copyright (c) 2017 Consejo Superior de Investigaciones Científicas (CSIC)

Creative Commons License
This work is licensed under a Creative Commons Attribution 4.0 International License.

Contact us

Technical support