* Corresponding author : Sophie Arnaud-Haond, address :

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1 Deep Sea Research Part II: Topical Studies in Oceanography November 2017, Volume 145, Pages 110-119 http://dx.doi.org/10.1016/j.dsr2.2015.07.013 http://archimer.ifremer.fr/doc/00274/38530/ 2015 Elsevier Ltd. All rights reserved. Achimer http://archimer.ifremer.fr Two pillars of cold-water coral reefs along Atlantic European margins: Prevalent association of Madrepora oculata with Lophelia pertusa, from reef to colony scale Arnaud-Haond Sophie 1, 2, *, Van Den Beld Inge 1, Becheler Ronan 1, Orejas C. 3, Menot Lenaick 1, Frank N. 4, 5, Grehan A. 6, 7, Bourillet Jean-Francois 8 1 Ifremer, Centre de Bretagne, Laboratoire Environnement Profond, CS 10070, 29280 Plouzané, France 2 Ifremer, Station de Sète, Laboratoire Ressources Halieutiques de Sète, Avenue Jean Monnet CS 30171, 34203 Sète Cedex, France 3 Instituto Español de Oceanografía, Centro Oceanográfico de Baleares, Moll de Ponent s/n, 07015 Palma de Mallorca, Islas Baleares, Spain 4 Institut für Umweltphysik, University Heidelberg, Im Neuenheimerfeld 229, 69120 Heidelberg, Germany 5 Laboratoire des Sciences du Climat et de L Environnement, unité mixte CEA-CNRS-UVSQ, Bât. 12 Avenue de la Terrasse, 91190 Gif-sur-Yvette, France 6, 7 Earth and Ocean Sciences, School of NaturalSciences, National University of Ireland,Galway, Ireland, 2Ryan Institute, NationalUniversity of Ireland, Galway, Ireland 8 Ifremer, Centre de Bretagne, Physical resources and sea floor ecosystems Department, CS 10070, 29280 Plouzané, France * Corresponding author : Sophie Arnaud-Haond, email address : Sophie.Arnaud@ifremer.fr Abstract : The scleractinian coral Lophelia pertusa has been the focus of deep-sea research since the recognition of the vast extent of coral reefs in North Atlantic waters two decades ago, long after their existence was mentioned by fishermen. These reefs where shown to provide habitat, concentrate biomass and act as feeding or nursery grounds for many species, including those targeted by commercial fisheries. Thus, the attention given to this cold-water coral (CWC) species from researchers and the wider public has increased. Consequently, new research programs triggered research to determine the full extent of the corals geographic distribution and ecological dynamics of Lophelia reefs. The present study is based on a systematic standardised sampling design to analyse the distribution and coverage of CWC reefs along European margins from the Bay of Biscay to Iceland. Based on Remotely Operated Vehicle (ROV) image analysis, we report an almost systematic occurrence of Madrepora oculata in association with L. pertusa with similar abundances of both species within explored reefs, despite a tendency of increased abundance of L. pertusa compared to M. oculata toward higher latitudes. This systematic association occasionally reached the colony scale, with twin colonies of both species often observed growing next to each other when isolated structures were occurring off-reefs. Finally, several false chimaera were observed within reefs, confirming that colonial structures can be coral bushes formed by an accumulation of multiple colonies even at the inter-specific scale, with no need for self-recognition mechanisms. Thus, we underline the importance of the hitherto underexplored M. oculata in the Eastern Please note that this is an author-produced PDF of an article accepted for publication following peer review. The definitive publisher-authenticated version is available on the publisher Web site.

2 Atlantic, re-establishing a more balanced view that both species and their yet unknown interactions are required to better elucidate the ecology, dynamics and fate of European CWC reefs in a changing environment. Keywords : Lophelia pertusa, Madrepora oculata, false-chimaera colonies, cold water corals (CWC), Bay of Biscay, Ireland, Iceland. Please note that this is an author-produced PDF of an article accepted for publication following peer review. The definitive publisher-authenticated version is available on the publisher Web site.

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