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. " E 70/ C ;-.1 1 (0./'-/<) THESE DE DOCTORAT EN SCmNCES DE j L'UNIVERSITE DE BRETAGNE OCCIDENTALE Spécialité : Océanologie Biologique présentée par Virginie TILOT La structure des assemblages mégabenthiques d'une . province à nodules polymétalliques de l'océan Pacifique tropical Est DEPARTEMENT DE L'ENVIRONNEMENT PROFOND DIRECITON DES RECHERCHES OCEANIQUES IFREMER 1CENTRE DE BREST BP70 29280 PLOUZANE FRANCE Dépôt légal Brest 1992 Nouvelle série , Abstract The structure of megabenthic assemblages of an abyssal polymetallic nodule province located in the tropical East Pacific ocean, between the Clarion and Clipperton fracture zones, is investigated at different levels. On a qualitative level, the identification, the ethology, the taxonomic richness and the faunal composition c1assified by functional groups are described over the totality of the area thus creating a work of reference and an annotated photographic atlas with the contribution of worldwide specialists for each phylum. On basis of a collection of more than 200 000 photographs and 55 hours of videos of the seafloor, a taxonomic richness of 240 taxons with 46 different echinoderms is recorded. Cnidaria is the most diversified phylum on the Clarion-Clipperton area with 59 different taxons. The trophic group of suspension feeders is the most represented. On a quantitative level, the particularly weil described site of NIXO 45 (1300oo'W/13001O'W, 13°56'N/14°08'N) lying at a mean depth of 4950 m, is chosen for the evaluation of its faunal density and composition, cIassified by phyla and by trophic and functional groups, within different edaphic conditions. Suspension feeders are more numerous than detritus feeders, carnivores or necrophagous feeders whatever the edaphic facies. The total faunal abundance is more important on facies C+10% anù facies C: with a slupe >15". Higiler ùensilies of suspension feedels are observed on facies ü (no nodules) composed of old sediments (from Oligocene to middle Miocene) and on facies c+ with a slope >15". Meanwhile detritus feeders are more abundant on facies C+lO%. The similarities among taxons and edaphic facies are emphasized by Reciprocal Averaging and enable us to differentiate preferential habitats and "faunal facies" ranked according to an edaphic gradient. The quantitative analysis is a1so set at a specific level when assessing the spatial heterogeneity of the population distributions. Comparisons are drawn from the data collected by other "in situ observation submersibles". The manned submersible "Nautile" explored one of the facies of the NIXO 45 site, the towed submersible "R.A.I.E." sampled three facies over the NIXO 41 site and the "Deep Tow instrumentation system" was used to sample pictures of 3 facies found on the Echo 1 site. The megafaunal biomass, arranged by trophic and functional groups is assessed and annotated for the three study sites. These data are confronted with the biomasses of megafauna, macrofauna and meiofauna sampled at the DaMES site C (Hecker and Paul, 1979) and with those from the Meriadzek Terrace in the bay of Biscay (Mahaut, 1991). The role of Iimiting factors (edaphic heterogeneity, currents) and of biotic factors in the structuration of epibenthic assemblages is displayed and discussed (1300 00'W1 \30° lO'W, \3°56'NI14°08'N). A particular type of biological activity is illustrated by echiuran Bonellids and its consequential effects at the sediment-water interface are analyzed. Environmental impact asscssment studics are presented and compensatory measures recommended . THESE de DOCTORAT de L'UNIVERSITE DE BRETAGNE OCCIDENTALE spécialité : Oceanographie Biologique présentée par Virginie TILOT La structure des assemblages mégabenthiques d'une province à nodules polymétalliques de l'océan Pacifique tropical Est Thèse soutenue le 7 juillet 1992 Composition du jury : Président: A. Guilcher Rapporteurs: F. Baguet M. Hoffert D.Reyss Membres: M. G1emarec J.P. Lenoble M. Sibuet C. Smith Tout ce que le coeur désire peut toujours se réduire à la figure de l'eau Paul Claudel, Positions et Propositions. II, p, 235, A la harpe de Meloe' qui me permit d'évider l'écheveau de mes pensées A la viole de Pasiphaë' qui sanctionna mes envolées AVANT·PROPOS Spiritus Dei ferebatur super aquas .. , Et creavit Deus... omnem animam viventem et motabilem. quaru produxerant aquae in speeies suas... Genèse. chap. 1. vers 2 et 21 Ce travail n'aurait pu se faire sans l'accueil, le soutien et les précieux conseils de nombreuses personnes qui m'ont aidé à jeter les filets sur le domaine de l'environnement profond et m'ont permis de découvrir la beauté d'un monde peuplé de formes rares et étranges ainsi que de m'interroger sur leurs comportements et leur organisation au sein d'un environnement de nodules polymétalliques se révélant aussi diversifié que la faune qui s'y abrite. Merci à toutes les personnes qui m'ont permis de réparer les mailles de mes pensées et de ramener mes données à l'horizon d'une science encore balbutiante dans son intervention en milieu profond. Pour le professeur Fernand Baguet qui m'a fait découvrir le monde fascinant de l'océanographie par son précieux enseignement et ses recherches éclairant de bioluminescence l'obscurité de mes connaissances. Pour m'avoir accueilli dans son laboratoire et fait découvrir un univers nouveau, m'avoir guidé dans mes recherches et fait profiter de sa très grande connaissance des sciences marines, j'exprime ma plus profonde reconnaissance et admiration à Myriam Sibuet qui me dirigea lout au long de ma thèse en me communiquant sa passion. Je tiens à témoigner ma profonde reconnaissance à Monsieur le Professeur Michel Giemarec qui a accepté de me conseiller dans la rédaction de cette thèse en me faisant bénéficier de ses précieux conseils et de son charisme bien connu par tous ceux qui l'entourent. Je tiens vivement à remercier toutes les personnes du département de l'environnement profond de L'IFREMER, pour leur chaleureux soutien et leurs nombreux conseils, Anne-Marie Alayse, Georges Barbier, Pierre Chevaldonne, Philippe Crassous, Daniel Desbruyères, Anne Godfroy, Alexis Kripounoff, Christine Lamy, Michel Marchand, Violaine Martin, Martine Morvan, Marie-Josée Rousic et Annick Vangrieshem. A toute l'équipe des Géosciences marines pour leur accueil et leur soutien efficace et chaleureux. Je pense tout particulièrement à Gérard Auffret. Claude Augris, Félix Avedik, Jean-Marie Auzende, Serge Berne. Daniel Bideau, Jean-François Sourillet, Pierre Cochonat, Gilbert Roch, Louis Geli, Roger Hekinian, Eliane Le Drezen, Gilles Lericolais, Raymond Le Suavé, Tania Marsset, Marie­ Michèle Pedel, Philippe Saget, Bruno et Elizabeth Savoye, Jean-Claude Sibuet, Nicole Uchard et Michel Voisset. J'exprime ma très profonde gratitude aux professeurs Hjalmar Thiel, Craig Smith et à George Wilson, Erdogan Ozturgut qui, en m'invitant à participer à leurs missions à la mer, m'ont permis d'acquérir une expérience professionnelle d'une richesse incomparable et de partager une aventure scientifique exceptionnelle. Ce travail a été rendu possible grâce aux contrats de consultant que m'ont passé Denis Tytgat aux Communautés Européennes, Daniel Reyss et Guy Herruoin à IFREMER et Erdogan Ozturgut à la c NOAA, Department of Commerce, U.S.A. à qui j'exprime ici ma plus profonde gratitude. Je tiens aussi à remercier tout particulièrement mon amie Marie-Laure Mahaut qui fit preuve d'une remarquable disponibilité pour rattraper mes bévues à mes balbutiements sur les ordinateurs et pour alimenter de longues discussions souvent passionnées. Toute mon amitié aussi à Annie Birroleau et à Pascal Gente pour leur soutien précieux ainsi qu'à DidierJolivet, co-thésard toujours aussi disponible dont j'admire le travail et la bonne humeur Je n'aurai pas pu rassembler autant de documents pour ma recherche biblographique sans l'aide efficace et si compétente des membres du service de la Bibliothèque et tout particulièrement Annick Salaun, Véronique Gautier, Jacqueline Prad'homme. Grâce à elles j'ai pu savourer le plaisir de consulter les premiers documents océanographiques et d'admirer les aquarelles d'expéditions mémorables. J'ai beaucoup appris aussi dans le monde de la taxinomie grâce à l'équipe du Centre de TRI d'IFREMER, Michel Segonzac, Patrick Briand et Joelle Galeron. envers qui je tiens à exprimer, ici, ma plus profonde gratitude pour leur sympathie et leur connaissance de la systématique. Ce travail n'aurait pas pu se faire sans la lourde correspondance échangée avec les nombreux spécialistes (dont les noms sont repris en annexe) qui ont si gentiment apportés leurs avis et leurs connaissances à mes hypothèses d'identification faunistique. Par eux, j'ai découvert les méandres de la systématique dans une ambiance de franche amitié qui semble caractériser le monde de la biologie abyssale et je me suis passionnée à essayer de découvrir les indices qui permettraient d'identifier des formes encore inconnues à la science. J'exprime toute ma sympathie à Gérard Auffret pour m'avoir accordé une oreille attentive et bienveillante durant toute cette thèse. C'est à lui que je dois ma première campagne en eaux hauturières et les joies de découvrir le travail en équipe à bord d'un navire océanographique. Je dois énormément à Philippe Gros et à Pierre Watremez pour leur disponibilité et leur maîtrise des techniques statistiques. Toute ma reconnaissance se porte à Gérard Vincent qui m'a ouvert les portes de son laboratoire photographique et initié aux mystères de j'image. Enfin je suis trés t1attée de faire examiner cette
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    1 Zitteliana An International Journal of Palaeontology and Geobiology Series B/Reihe B Abhandlungen der Bayerischen Staatssammlung für Paläontologie und Geologie 29 Paläontologie im Blickpunkt 80. Jahrestagung der Paläontologischen Gesellschaft 5. – 8. Oktober 2010 in München Programm und Kurzfassungen – Program and Abstracts München 2010 Zitteliana B 29 118 Seiten München, 1.10.2010 ISSN 1612-4138 2 Editors-in-Chief/Herausgeber: Gert Wörheide, Michael Krings Mitherausgeberinnen dieses Bandes: Bettina Reichenbacher, Nora Dotzler Production and Layout/Bildbearbeitung und Layout: Martine Focke, Lydia Geissler Bayerische Staatssammlung für Paläontologie und Geologie Editorial Board A. Altenbach, München B.J. Axsmith, Mobile, AL F.T. Fürsich, Erlangen K. Heißig, München H. Kerp, Münster J. Kriwet, Stuttgart J.H. Lipps, Berkeley, CA T. Litt, Bonn A. Nützel, München O.W.M. Rauhut, München B. Reichenbacher, München J.W. Schopf, Los Angeles, CA G. Schweigert, Stuttgart F. Steininger, Eggenburg Bayerische Staatssammlung für Paläontologie und Geologie Richard-Wagner-Str. 10, D-80333 München, Deutschland http://www.palmuc.de email: [email protected] Für den Inhalt der Arbeiten sind die Autoren allein verantwortlich. Authors are solely responsible for the contents of their articles. Copyright © 2010 Bayerische Staassammlung für Paläontologie und Geologie, München Die in der Zitteliana veröffentlichten Arbeiten sind urheberrechtlich geschützt. Nachdruck, Vervielfältigungen auf photomechanischem, elektronischem oder anderem Wege sowie
  • Experimental Neoichnology of Crawling Stalked Crinoids

    Experimental Neoichnology of Crawling Stalked Crinoids

    Swiss Journal of Palaeontology https://doi.org/10.1007/s13358-018-0158-9 (0123456789().,-volV)(0123456789().,-volV) REGULAR RESEARCH ARTICLE Experimental neoichnology of crawling stalked crinoids 1,2 3 4 1,2 5 Krzysztof R. Brom • Kazumasa Oguri • Tatsuo Oji • Mariusz A. Salamon • Przemysław Gorzelak Received: 13 June 2018 / Accepted: 5 July 2018 Ó The Author(s) 2018 Abstract Stalked crinoids have long been considered sessile. In the 1980s, however, observations both in the field and of laboratory experiments proved that some of them (isocrinids) can actively relocate by crawling with their arms on the substrate, and dragging the stalk behind them. Although it has been argued that this activity may leave traces on the sediment surface, no photographs or images of the traces produced by crawling crinoids have been available. Herein, we present results of neoichnological experiments using the shallowest species of living stalked crinoid, Metacrinus rotundus, dredged from Suruga Bay (near the town of Numazu, Shizuoka Prefecture, * 140 m depth). Our results demonstrate that isocrinids produce characteristic locomotion traces, which have some preservation potential. They are composed of rather deep and wide, sometimes weakly sinuous, central drag marks left by the stalk and cirri, and short, shallow scratch marks made by the arms. Based on the functional morphology and taphonomy, it has been argued that the ability to autotomize the stalk and relocate had already evolved in the oldest stem-group isocrinids (holocrinids), likely in response to increased benthic predation pressure during the so-called Mesozoic marine revolution. Our data show that this hypothesis may be corrob- orated in the future by ichnological findings, which may provide more direct proof of active locomotion in Triassic holocrinids.