Second completely revised edition
Editors:
Daniel Desbruyères, Michel Segonzac & Monika Bright
Handbook of Deep-SeaHydrothermalVent Fauna
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Katalog / Publication: Denisia 18
ISSN 1608-8700
Zugleich Kataloge der Oberösterreichischen Landesmuseen, N.S. 43
ISBN 10 3-85474-154-5 / ISBN 13 978-3-85474-154-1
Erscheinungsdatum / Date of delivery: 6. April 2006
Medieninhaber und Herausgeber / Copyright: Land Oberösterrreich, Biologiezentrum der Oberösterreichische Landesmuseen, J.-W.-Klein-Str. 73, 4040
Linz, Austria
URL / Internet: http://www.biologiezentrum.at
E-Mail: [email protected]
Wissenschaftliche Redaktion / Scientific editors: Daniel Desbruyeres, Michel Segonzac & Monika Bright
Redaktionelle Betreuung / Editorial assistance: Dr. Erna Aescht
Layout, Druckorganisation /
Layout, printing organisation: Eva Rührnößl
Umschlag-, Plakatgestaltung / Cover, placard: Eva Rührnößl
Druck / Printing: Plöchl-Druck, Werndlstraße 2, 4240 Freistadt
Bestellung / Ordering: http://www.biologiezentrum.at/biowww/de/biblio/index.html oder / or [email protected]
Das Werk einschließlich aller seiner Teile ist urheberrechtlich geschützt. Jede Verwertung außerhalb der en-
gen Grenzen des Urheberrechtsgesetzes ist ohne Zustimmung des Medieninhabers unzulässig und strafbar.
Das gilt insbesondere für Vervielfältigungen, Übersetzungen, Mikroverfilmungen sowie die Einspeicherung
und Verarbeitung in elektronischen Systemen. Für den Inhalt der Abhandlungen sind die Verfasser verant-
wortlich. Schriftentausch erwünscht!
All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any me-
ans without prior permission from the publisher.
We are interested on an exchange of publications.
Umschlagfoto / Cover: Riesenbartwurm Riftia pachyptila. Photo: M. Bright, Layout: E. Rührnößl.
Zitiervorschlag für das Buch / The complete book may be referenced as follows:
Desbruyères D.l, Segonzac M. & M. Bright (Editors; 2006): Handbook of Deep-Sea Hydrothermal Vent Fauna.
Second completely revised edition. Denisia 18: 544 pp.
Zitiervorschlag für Einzelarbeiten / Single taxa or contributions may be referenced as follows:
Author(s) of taxon (2006): Species. Denisia 18: page(s).
e.g. Desbruyères D. (2006) Riftia pachyptila Jones, 1981 „giant tubeworm“. Denisia 18: 262-263.
Ausstellung / Exhibition: Heiß und giftig – Oasen der Tiefsee / Hot and toxic – Oases in the deep sea
Ort / Address: Biologiezentrum der Oberösterreichischen Landesmuseen, J.-W.-Klein-Str. 73, 4040 Linz, Austria
Zeitraum / Period: 7. April bis 1. Oktober 2006
Konzept, Organisation und Gestaltung /
Concept, organization, design: Mag. Stephan Weigl, Dr. Monika Bright, Johannes Rauch
Ausstellungstechnik, Mitarbeit /
Exhibition techniques, collaboration: Jürgen Plass, Roland Rupp, Franz Meindl, Bruno Tumfart, Erwin Kapl, Josef Schmidt, Roland Zarre
Museumspädagogik / Museum education: Mag. Sandra Kotschwar
Leihgeber / Lenders: Dr. Monika Bright (Univ. Wien), Dr. Bernd Moser (Joanneum, Graz), Dr. Michel Segonzac (Ifremer, Frank-
reich), Dr. Werner Tufa (Univ. Marburg/Lahn), Dr. Janet R. Voight (The Field Museum of Natural History, Chi-
cago, USA), Romain Chausse (Muséum National d’Histoire Naturelle, Paris, Frankreich)
Impressum
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Abstract, Résumé, Zusammenfassung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8Photo credits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Contributors and adresses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Milestones in the discovery of hydrothermal-vent faunas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Hydrothermal vent meiofauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27Hydrothermal vent parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Species descriptions
Granuloreticulosa, Foraminifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Porifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35Demospongiae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37Hexactinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Cnidaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .48Hydrozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48Scyphozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64Anthozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Mollusca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75Solenogastres . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75Polyplacophora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80Gastropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Patelligastropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83Vetigastropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86Neomphalina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104Neritimorphoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121Caenogastropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124Heterobranchia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135Prosobranchia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138Nudibranchia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Bivalvia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141Heterodonta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142Protobranchia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149Pteriomorphia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Cephalopoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
Nematoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Acanthocephala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Nemertini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Contents
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Annelida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183Polychaeta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Eunicida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .186Phyllodocida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194Sabellida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253Scolecida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266Spionida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272Terebellida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Oligochaeta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297
Arthropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .298Arachnida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298Pycnogonida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .301Crustacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
Ostracoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307Copepoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316Cirripedia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 356Leptostracta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .369Cumacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .370Tanaidacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .372Isopoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379Amphipoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382Ephausidacae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409Decapoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 410
Caridea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .410Astacidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .433Anomura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434Brachyura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455
Echinodermata Crinoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475Asteroidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477Echinoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 479Holothuroidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480Ophiuroidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481
Chaetognatha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 486
Hemichordata, Enteropneusta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487
Chordata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 489Chondrichthyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 489Osteichthyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 493
Major known deep-sea hydrothermal vent fields . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 513Abbreviations, ridges, back-arc basins and vent fields . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 518Species list . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 519Schematic drawings of major taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 532Genus index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 542Author index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 544
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The second extensively expanded edition of the “Hand-book of Deep-Sea Hydrothermal Vent Fauna“ gives on over-view of our current knowledge on the animals living at hydro-thermal vents. The discovery of hydrothermal vents and pro-gresses made during almost 30 years are outlined. A brief intro-duction is given on hydrothermal vent meiofauna and parasites.Geographic maps and a table of mid-ocean ridges and back-arcbasins with the major known hydrothermal vent fields, their lo-cation and depth range and the most prominent vent sites areprovided. Higher taxa are presented individually with informa-tion on the current taxonomic and biogeographic status, thenumber of species described, recommendations for fixation, and
schematic drawings, which aim to help non-specialists to iden-tify the animals. 86 authors contributed with their expertise tocreate a comprehensive database on animals living at hydro-thermal vents, which contains information on the morphology,biology, and geographic distribution of more than 500 current-ly described species belonging to one protist and 12 animalphyla. It comprises also the largest collection of more than1000 pictures of hydrothermal vent animals taken in situ withsubmersibles, in vivo after collection, and with various dissec-tion, light, and scanning electron microscopes after fixationand preparations.
La seconde édition, considérablement étendue et amélioréedu « Handbook of Deep-Sea Hydrothermal Vent Fauna » don-ne un panorama de notre connaissance sur les animaux qui vi-vent autour des sources hydrothermales océaniques. Les avan-cées faites depuis leurs découvertes il y a trente ans environ,donnent lieu à une mise en perspective rédigée par des acteursdes campagnes océanographiques. Une brève introduction estproposée sur la méiofaune et les parasites, puis chaque taxon su-périeur est présenté avec des rappels sur la taxinomie, des in-formations sur sa distribution géographique, des représentationsschématiques de la morphologie permettant à des non-spécia-listes de comprendre la nomenclature, des informations sur lenombre d’espèces présentes, des conseils pour la fixation et laconservation des échantillons. Des cartes de distribution des
champs hydrothermaux le long des rides océaniques et dans lesbassins arrière-arc sont proposées assorties d’un tableau quidonne leur position géographique, leur profondeur et les sur-noms des principaux sites. 86 auteurs ont contribué à la créa-tion d’une base de données sur les animaux présents dans cetécosystème surprenant, en donnant des informations sur lamorphologie, la biologie et la distribution géographique de plusde 500 espèces appartenant à 12 phylums animaux. Cette basede données, présentée sous forme de fiches individuelles, re-groupe une série unique de plus de 1000 illustrations compor-tant dessins au trait, photographies des organismes dans leurmilieu, vues d’animaux isolés après récoltes et micro-photogra-phies prises au microscope électronique à balayage.
Abstract
Résumé
Die zweite, wesentlich erweiterte Auflage des „Handbookof Deep-Sea Hydrothermal Vent Fauna“ gibt einen Überblicküber den momentanen Stand der Forschung. Die Entdeckungder Hydrothermalquellen, Fortschritte der letzten beinahe 30Jahre, sowie Meiofauna und Parasiten der Hydrothermalquellenwerden kurz umrissen. Alle ozeanischer Rücken und Back-ArcBasins mit ihren bekannten Hydrothermalquellfeldern, ihre ge-naue Lage und Tiefe, sowie ihre benannten Fundstellen werdenin geographischen Karten und einer Tabelle gezeigt. HöhereTaxa werden individuell dargestellt mit Information über denmomentanen taxonomischen und biogeographischen Stand,
die Anzahl der beschriebenen Arten, Methoden zur Fixierungund schematischen Zeichnungen, die es dem Nicht-Spezialis-ten die Identifizierung erleichtern sollen. Über 80 Autoren ga-ben ihren Beitrag zur Erstellung einer umfassenden Datenbankmit mehr als 500 Tieren aus einem Protisten-Stamm und 12Tierstämmen und Informationen zu deren Morphologie, Biolo-gie und Verbreitung. Darüber hinaus werden die Tiere derHydrothermalquellen in mehr als 1000 Bildern, Unterwasser-aufnahmen von bemannten und unbemannten U-Booten, invivo Aufnahmen und diversen licht- und elektronenmikrosko-pischen Aufnahmen, dargestellt.
Zusammenfassung
Key words: Hydrothermal vents, deep sea, mid-ocean ridge, back-arc basin, hydrothermal vent fauna.
5
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Deep-sea hydrothermal vents and their associated faunawere discovered in 1977. The presence of these rich oases of li-fe in the food-limited environment of basaltic Mid-OceanicRidges fostered the interest of the marine biologists to explorethe deep sea using submersibles with a focus on fragmented ha-bitats. During the intervening three decades, a large number ofcruises using deep submergence vehicles (HOV or ROV) weredevoted to the biological study of the hydrothermal communi-ties on the Pacific, Atlantic, and Indian Ridges as well as inBack-Arc Basins and Volcanic Arcs of the Pacific Rim of Fire.These series of cruises were mainly devoted to the understan-ding (1) the distribution, the phylogeny, and the dispersal ofthis specialized fauna which is restricted to this fragmented “ex-treme” habitat along the world’s oceanic ridges and (2) to theniche characterization and the study of the physiological adap-tations of the organisms (or symbiotic associations) living inthe mixing zone between the superheated fluid and the colddeep-sea water. This endeavour remains unique in its extentand intensity in the history of biological exploration of theworld’s oceans since the early studies of the deep-sea by the pio-neering expeditions of the nineteenth century. However, be-cause of logistic constrains, this accomplishment is still far fromproviding an exhaustive overview of the composition and dis-tribution of hydrothermal vent fauna; some Ridges (South At-lantic and the three Indian Ridges) still deserve obviously mo-re attention and the study of remote targets as high latitude rid-ges (Arctic and Antarctic Seas) or isolated basins (South Sand-wich and Andaman Sea) would bring valuable information onthe biogeography of the vent fauna. Multiyear surveys on theEast Pacific Rise and the Juan de Fuca Ridge have also demon-strated that the hydrothermal environment is submitted totemporal instability due to volcanic and tectonic events; thisinstability greatly affects the patterns of the ecosystem deve-lopment and produces a variation in specific composition ofcommunities only detected by temporal series of sampling.
Reliable morphological identification of collected speci-mens presently remains the inescapable base of most of the bi-ological work conducted at vents on animals, including the mo-lecular approach. Because of the taxonomic novelty of deep-sea hydrothermal vent communities, an impressive descriptive
work has been conducted for three decades producing morethan 500 fully identified species of which descriptions orrecords are dispersed throughout the scientific literature. Nev-ertheless the list of animal species living at vents is still evolv-ing every week; new species are described from additional sam-ples in known locations and from new locations, descriptionsare revisited, sibling species are detected based on moleculartaxonomy, and new combinations are proposed based on stud-ies of large series and molecular information. The technology ofimaging in situ greatly improved in the last few years givingconsiderable information on the shape, colors, behavior, andmicro-distribution of living animals. The dialogue betweenclassical and molecular taxonomy brought new insights andclarifies the biogeographic patterns of vent communities as wellas their relationships with other deep-sea reduced habitats(cold seeps, food falls, and minimum oxygen zones).
In the foreword of the first edition of the “Handbook ofDeep-Sea Hydrothermal Vent Fauna” (Ifremer Editions, 1997),we proposed to publish periodic addenda to the first editionbased on new published observations and we invited contribu-tors to help us by sending information and illustration. The re-sults far exceeded our hopes and we realized soon that a new is-sue of the handbook would be fully justified. At the same time,the Biologiezentrum der Oberösterreichischen Landesmuseen(Biology Center of the Upper Austrian Museums) was prepar-ing an exhibition on deep-sea hydrothermal vent life plannedfor April 2006 and was looking for authors willing to publish areview on vent biology. It was the perfect opportunity for us torestart the collection of contributions from the worldwide net-work of taxonomists involved in the study of hydrothermalvent samples. We want to thank here all the contributors whoparticipated in the wording of this new issue.
This book, published under the auspices of InterRidge andCensus of Marine Life/ChEss, is an overview of the present tax-onomic knowledge of the deep-sea hydrothermal vent fauna,intended for scientists and submersible pilots working at sea,but also for students and the general public curious about thelife in the ocean.
D. Desbruyères, M. Segonzac & M. Bright
Foreword
« Toute partie dans la chaise qui ne se voyait pas était toutaussi parfaitement faite que ce qu‘on voyait.
C‘est le principe même des cathedrales. »
Charles Péguy (1873–1914)
“All the elements of the chair that are hidden from vieware as perfectly crafted as the visible ones. This ist the very
principle of the cathedrals.“
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Collection of animals as well as in situ photographs and vi-deos of animals from a deep-sea environment are mainly achie-ved through the use of manned and unmanned submersibles.Valuable sampled animals are often a by-product of large bulksamples, fixation and storage sometimes are less than optimalfor all taxa, and the time for taking pictures while on board isoften scarce. Also, it is often due to pure luck that in situphotographs and videos are taken at the decision of chief scien-tists, pilots, and researchers on board as they must interrupttheir ongoing tasks and use valuable bottom time to documentanimals in situ. Good quality pictures taken on the bottom pro-vide a unique set of information on the micro-distribution, be-havior, and biological interactions. Even those photographs,which are in low resolution or out of focus, are highly valuablebecause they give us an impression of where these animals oc-cur and how they behave naturally.
We stress our belief that our successes in understanding thefauna living at hydrothermal vents will be highly influenced bythe way we approach solving these issues. So we recommend fortaxonomy, ecology and outreach purposes, that researchers ta-ke the time on the bottom to video and photograph, that scien-tists onboard research vessels take high quality photographs ofanimals before preservation as well as follow instructions forproper fixation and storage, and that they get into contact withtaxonomists for identification and appropriate procedures forvoucher specimens. Moreover, we hope that more taxonomistswill be invited to research cruises to better support our goal oflearning more about hydrothermal vents and their inhabitants.Recent experiences demonstrated that the presence of taxono-mists on cruises improved by far our knowledge of the faunalcomposition of vent communities and fostered interactionswith other disciplines.
D. Desbruyères, M. Segonzac & M. Bright
Remarks
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We would like to thank all the contributors whose know-ledge of the fauna and taxonomic expertise allowed us to revi-se and supplement the second edition of this book. We are gra-teful to Gerhard Aubrecht, the head/director of the Biologie-zentrum der Oberösterreichischen Landesmuseen (BiologyCenter of the Upper Austrian Museums), for making this pu-blication possible and for his continuous support throughoutthe entire process. Special thanks to Erna Aescht (BiologyCenter) for the editing, Eva Rührnößl (Linz) for the page-ma-king, Heidemarie Grillitsch (University Vienna) for the sche-matic drawings, Ingrid Kolar (University Vienna) for checkingthe numerous references, and Patrick Briand (Ifremer) for hishelp with the pictures, Violaine Martin (Ifremer) for many dra-wings, and Philippe Crassous (Ifremer) for his technical exper-tise with the SEM preparations.
We express our heartful thanks to the chief scientists Fern-ando Barriga, Nadine Le Bris, Charles R. Fisher, FrancoiseGaill, Anne Godfroy, Jun Hashimoto, S. Kim Juniper, FrançoisLallier, Marvin D. Lilley, Pierre-Marie Sarradin, Tim Shank,Verena Tunnicliffe, Karen Von Damm, Cindy L. Van Doverand Robert C. Vriejenhoek, who allowed us to have access tosamples, data, and in situ pictures. We are also grateful to themany contributors who shared their pictures with us.
This book was supported by the Federal Government ofUpper Austria, the Biology Center of the Upper Austrian Mu-seums, Ifremer, ChEss (especially the coordinators of the ChEssproject Eva Ramirez-Llodra and Maria Baker), InterRidge, theAustrian Science Foundation, and the Faculty of Life Sciences,University of Vienna.
D. Desbruyères, M. Segonzac & M. Bright
Acknowledgements
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Auger, V. – Canadian Scientific Submersible Facility, CanadaBiscoito, M. – Museu Municipal do Funchal, PortugalBlake, J. – ENSR Marine & Coastal Center, USABlaston, P. – University of Otago, New ZealandBriand, P. – Ifremer, Brest, FranceBright, M. – University of Vienna, AustriaBurreson, E. – College of William and Mary, Williamsburg,
USAChan, T. Y. – National Taïwan Ocean University, TaïwanCosel, R. von – Muséum National d’Histoire Naturelle, Paris,
FranceCunha, M.R. – Centro das Zonas Costeiras e do Mar, Universi-
dad, PortugalFautin, D. – Natural History Museum and Biodiversity Rese-
arch Center, Kansas, USADejouannet, J.-F. – IRD (Institut Recherche Développement)
Paris, FranceDrukker-Brammall, P. – Smith Institute of Ichthyology, Gra-
hamstown, South AfricaDugornay O. – Ifremer, Brest, FranceEncarnação, H. – Museu Municipal do Funchal, PortugalFifis, A. – Ifremer, Brest, FranceFisher, C. R. – The Pennsylvania State University, USAGlover, A. – The Pennsylvania State University, USAGuinot, D. – Muséum National d’Histoire Naturelle, Paris,
FranceHarrison-Nelson, E. – National Museum of Natural History,
Smithsonian Institution, USAHashimoto, J. – JAMSTEC, Nagasaki, JapanHeemstra, E. – Smith Institute of Ichthyology, Grahamstown,
South AfricaHessler, R.R. – Scripps Oceanographic Institution, USAHourdez, S. – Station Biologique Roscoff, FranceIvanenko, V. N. – Moscow State University, RussiaJuniper, K. S. – Université du Québec, Montréal, CanadaKlitz, K. – National Museum of Natural History, Smithsonian
Institution, USAKornicker, L.S. – National Museum of Natural History, Smith-
sonian Institution, USAKudenov, J.D. – University Alaska, Anchorage, USALe Goff, A. – Muséum National d’Histoire Naturelle, Paris,
FranceLindsay, S. C – Australian Museum, Syndney, AustrialiaLopez-Gonzalez, P. – Universidad de Sevilla, Spain
Lozouet, P. – National Muséum d’Histoire Naturelle, Paris,France
Lutz, R. A. – Rutgers University, USAMaddocks, R. M. – University of Houston, USAMaestrati, P. – Muséum National d’Histoire Naturelle, Paris,
FranceMarcus, J. – University of Victoria, CanadaMartin, V. – Ifremer, Brest, FranceMcLean, J. – Museum of Natural History, Los Angeles, USAMills, S.W. – Woods Hole Oceanographic Institution, USAMiura, T. – JAMSTEC, Nagasaki, JapanMullineaux, L. – Woods Hole Oceanographic Institution, USANg, N. K. – National University of Singapore, Republic of Sin-
gaporeOkata, Y. – JAMSTEC, Nagasaki, JapanPleijel, F. – University of Göteborg, SwedenRouse, G. – University of Adelaide, AustraliaRoux, M. – Université de Reims, FranceSagalevitch, A.M. – Shirshov Institute of Oceanology, Mos-
cow, RussiaSaldanha, L. † – LMG, Lisbonne, PortugalShagin, A. – PMGE (Mining Intitute, St Petersburg), RussiaSmith Jr, K. – Scripps Institution of Oceanography, USASouthward, E.C. – Plymouth Marine Laboratory, United King-
domStöhr, S. – Swedish Museum of Natural History, SwedenTakeda, M. – JAMSTEC, JapanTivey, M. – Woods Hole Oceanographic Institution, USATsuchida, S. – JAMSTEC, JapanTunnicliffe, V. – University of Victoria, CanadaVan Dover, C.L. – College of William and Mary, Williamsburg,
USAVerschelde, D. – Universiteit Gent, BelgiumVrijenhoek, R. – Monterey Bay Aquarium Research Institute,
USAWarén, A. – Swedish Museum of Natural History, SwedenWilson, G.D.F. – Australian Museum, Syndney, AustrialiaYoung, C.M. – Origon Institute of Marine Biology, USAZierenberg, R. – University of California, Davis, USA
Photo credits
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Almeida, Armando J. – Laboratório Marítimo da Guia, IMAR,Faculdade de Ciências da Universidade de Lisboa, Estradado Guincho, 2750-642 Cascais, Portugal [email protected]
Baba, Keiji – Kumamoto University, Faculty of Education, 2-40-1 Kurokami, Kumamoto 860-8555, [email protected]
Bamber, Roger – Environment: Coastal & Marine, The NaturalHistory Museum, Cromwell Road, London SW7 5BD, Uni-ted Kingdom [email protected]
Barthélémy, Roxane – Université de Provence, Biologie Ani-male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé-dex 3, France [email protected]
Bartsch, Ilse – Biologische Anstalt Helgoland, Zentrale Ham-burg, Notkestrae 31, D-22607 Hamburg, [email protected]
Bellan-Santini, Denise – Université d’Aix-Marseille 2, StationMarine d’Endoume-Luminy, Rue de la Batterie des Lions,13007 Marseille, France [email protected]
Biscoito, Manuel – Museu Municipal do Funchal, Rua da Mou-raria, 31, 9000 Funchal, Madeira, Portugal [email protected]
Blake, James A. – ENSR Marine & Coastal Center, 89 WaterStreet, Woods Hole, MA 02543, USA [email protected]
Böggemann, Markus – Habsburgerallee 37, 60385 Frankfurt amMain, Germany [email protected]
Bouchet, Philippe – Département Systématique et Évolution,Muséum National d’Histoire Naturelle, Taxonomie-Collec-tion Mollusques, 55 rue Buffon, 75231 Paris, France [email protected]
Boury-Esnault, Nicole – Université d’Aix-Marseille 2, Centred’Océanologie de Marseille, Station Marine d’Endoume -Luminy, Rue de la Batterie des Lions, 13007 Marseille, Fran-ce [email protected]
Briand, Patrick – Département «Etude des écosystèmes pro-fonds», Ifremer, Centre de Brest, BP 70, 29280 Plouzané,France [email protected]
Bright, Monika – Department of Marine Biology, University ofVienna, Althanstr. 14, A-1090 Vienna, [email protected]
Buron, de Isaure – Division of Natural Sciences and Enginee-ring, University of South Carolina Spartanburg, Spartan-burg, SC 29303, USA [email protected]
Burreson, Eugene M. – Department of Environmental andAquatic Animal Health, Virginia Institute of Marine Scien-ce, College of William and Mary, PO. Box 1346, GloucesterPoint, VA 23062 USA [email protected]
Casanova, Bernadette – Université de Provence, Biologie Ani-male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé-dex 3, France [email protected]
Casanova, Jean-Paul – Université de Provence, Biologie Ani-male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé-dex 3, France [email protected]
Causse, Romain – Muséum National d’Histoire Naturelle, La-boratoire d’Ichthyologie, 43, rue Cuvier, 75231 Paris cedex05, France [email protected]
Cohen, Daniel M. – California Academy of Sciences, GoldenGate Park, San Francisco, CA 95521, USA [email protected]
Corbera, Jordi – Carrer Gran, 90, 08310 Argentona, Spain [email protected]
Cosel, von Rudo – Département Systématique et Évolution,Muséum National d’Histoire Naturelle, Taxonomie-Collec-tion Mollusques, 55 rue Buffon, 75231 Paris, France [email protected]
Cunha, Marina R. – Centro das Zonas Costeiras e do Mar, Uni-versidad, Departamento de Biologia, 3810-193 Aveiro, Por-tugal [email protected]
Daly, Marymegan – Dept. Evolution, Ecology and OrganismalBiology, Ohio State University, 315 Kinnear RD, ColumbusOH 43212, USA [email protected]
Defaye, Danielle – Département Milieux et Peuplements Aqua-tiques, Muséum National d’Histoire Naturelle, 61, rue deBuffon, 75005 Paris, France [email protected]
Desbruyères, Daniel – Département «Etude des écosystèmes pro-fonds», Ifremer, Centre de Brest, BP 70, 29280 Plouzané,France [email protected]
Gebruk, Andrey V. – Laboratory of Ocean Benthic Fauna, P.P.Shirshov Institute of Oceanology, Nakhimovsky Pr., 36,Moscow 117997, Russia [email protected]
Geistdoerfer Patrick – Laboratoire d’Ichthyologie, 43, rue Cu-vier, 75231 Paris, France [email protected]
Gonzalez, Angel – Ecología y Biodiversidad Marina (ECOBIO-MAR), Instituto de Investigaciones Marinas (CSIC), C/Eduardo Cabello 6, 36208 Vigo, Spain [email protected]
Contributors and addresses
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Guerra, Angel – Ecología y Biodiversidad Marina (ECOBIO-MAR), Instituto de Investigaciones Marinas (CSIC), C/Eduardo Cabello 6, 36208 Vigo, Spain [email protected]
Guinot, Danièle – Département Milieux et Peuplements Aqua-tiques, Muséum National d’Histoire Naturelle, 61, rue deBuffon, 75005 Paris, France [email protected]
Haney, Todd – Department of Organismic Biology, Ecology, andEvolution, University of California Los Angeles, 621 Char-les E. Young Drive South, Box 1606, Los Angeles, Califor-nia 90095-1606, USA [email protected]
Harrison-Nelson, Elisabeth – Department of Invertebrate Zoo-logy, National Museum of Natural History, Smithsonian In-stitution, MRC 163, Washington, D. C. 20013-7012, [email protected]
Hashimoto, Jun – Japan Marine Science and Technology Cen-ter (JAMSTEC) 2-15, Natsushima-Cho, Yokosuka, 237, Ja-pan [email protected]
Holzmann, Maria – Department of Paleontology, University ofVienna, Althanstr. 14, A-1090 Vienna, Austria [email protected]
Hourdez, Stephan – Station Biologique, 29680 Roscoff, [email protected]
Hurtado, Luis – University of Arizona, 310 Biosciences West,P.O. Box 210088, Tucson, AZ, 85721-0088, USA [email protected]
Ivanenko, Viatcheslav N. – Department of Invertebrate Zoolo-gy, Biological Faculty, Moscow State University, Moscow119899, Russia [email protected]
Jaume, Damian – IMEDEA (CSIC-UIB), Instituto Mediterra-neo de Estudios Avanzados, C/ Miquel Marques 21, 07190-Esporles (Mallorca, Illes Balears), Spain [email protected]
Jones, Diana S. – Western Australian Museum, Locked Bag 49.WELSHPOOL DC, Western Australia 6986, Australia [email protected]
Justine, Jean-Lou – Institut Développement Recherche (IRD),Nouméa, Nouvelle-Calédonie, France [email protected]
Komai, Tomoyuki – Natural History Museum and Institute Chi-ba, 955-2, Aoba-cho, Chuo-ku, Chiba 260-8682, Japan [email protected]
Kornicker, Louis S. – Department of Invertebrate Zoology, Na-tional Museum of Natural History, Smithsonian Institution,MRC 163, Washington, D. C. 20013-7012, USA [email protected]
Kudenov, Jerry D. – Department Biological Sciences, Universi-ty Alaska Anchorage, 3211 Providence Drive, Anchorage,Alaska 99508, USA [email protected]
Larsen, Kim – Department of Invertebrates, Zoological Mu-seum, Universitetsparken 15, 2100 KBH, Denmark [email protected]
Lemaitre, Rafael – Smithsonian Institution, Department of Sys-tematic Biology, NMHN, MRC 163, PO Box 37012, Was-hington, DC 20013-7012, USA [email protected]
Lopez-Gonzalez, Pablo – Departamento de Fisiologia y BiologiaAnimal, Facultad de Biologia, Universidad de Sevilla, Rei-na Mercedes, 6, 41012 Sevilla , Spain [email protected]
Lutz, Richard A. – Center for Deep-Sea Biology & Biotechno-logy, Rutgers University, Dudley Road, New Brunswick, NJ08903-0231, USA [email protected]
Macpherson, Enrique – CSIC/CEAB, Camí Santa Bàrbara, s/n,17300 Blanes, Girona, Spain [email protected]
Maddocks, Rosalie F. – Department of Geosciences, Room 312Science & Research Bldg. 1, University of Houston, Hous-ton, TX 77204-5007, USA [email protected]
Mah, Christopher – Department. of Invertebrate Zoology, Na-tional Museum of Natural History, MRC-163, PO Box37012, Smithsonian Institution, Washington DC 20013,USA [email protected]
Miura, Tomoyuki – Faculty of Fisheries, Kagoshima University,4-50-20, Shimoarata, Kagoshima, 890 [email protected]
Möller, Peter R. – Zoological Museum, University of Copenha-gen Universitetsparken 15, 2100 KBH, Denmark [email protected]
Moreau, Xavier – Université de Provence, Biologie Animale(Plancton), 3, place Victor Hugo, 13331 Marseille, Cédex 3,France [email protected]
Myers, Alan A. – Department of Zoology, Ecology and PlantScience, National University of Ireland, Cork, Republic ofIreland [email protected]
Newman, William A. – Scripps Institution of Oceanography, LaJolla, CA 92093-0202, USA [email protected]
Ng, Ngan Kee – Department of Biological Sciences, NationalUniversity of Singapore, 14, Science Drive 4, Singapore117543, Republic of Singapore [email protected]
Nielsen, Jorgen G. – Department of Invertebrates, ZoologicalMuseum, Universitetsparken 15, 2100 KBH, [email protected]
Okutani, Takashi – Tokyo University of Fisheries, Konan 4-5-7,Minato-ku, Tokyo 108, Japan [email protected]
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Pleijel, Fredrik – Tjärnö Marine Biological Laboratory, Depart-ment of Marine Ecology, Göteborg University, SE-452 96Strömstad, Sweden [email protected]
Pugh, Phillip R. – School of Ocean and Earth Science, NationalOceanography Centre, European Way, Southampton, SO143ZH, United Kingdom [email protected]
Ramirez-Llodra, Eva – Institut de Ciències del Mar, CMIMA-CSIC, Passeig Marítim de la Barceloneta 37-49, E-08003Barcelona, Spain [email protected]
Rodríguez, Estefanía – Departamento de Fisiologia y BiologiaAnimal, Facultad de Biologia, Universidad de Sevilla ReinaMercedes, 6, 41012 Sevilla, Spain [email protected]
Roux, Michel – Laboratoire des Sciences de la Terre, Centre deRecherches Agronomiques, 2 esplanade Roland Garros,51100 Reims, France [email protected]
Salvini-Plawen, von Luitfried – Department of EvolutionaryBiology, University of Vienna, Althanstr. 14, A-1090 Vien-na, Austria [email protected]
Schein, Elisabeth – Laboratoire des Sciences de la Terre, Centrede Recherches Agronomiques, 2 esplanade Roland Garros,51100 Reims, France [email protected]
Segonzac, Michel – Département «Etude des écosystèmes pro-fonds», Ifremer, Centre de Brest, BP 70, 29280 Plouzané,France [email protected]
Senz, Wolfgang – Department of Evolutionary Biology, Univer-sity of Vienna, Althanstr. 14, A-1090 Vienna, Austria
Southward, Alan J. – Plymouth Marine Laboratory, CitadelHill, Plymouth, PL1 2PB, United [email protected]
Southward, Eve C. – Plymouth Marine Laboratory, Citadel Hill,Plymouth, PL1 2PB, United Kingdom [email protected]
Stöhr, Sabine – Swedish Museum of Natural History, Depart-ment of Invertebrate Zoology, Box 50 007, Stockholm SE,10405 Sweden [email protected]
Tsuchida, Shinji – Marine Ecosystems Research Department, Ja-pan Marine Science and Technology Center, 2-15 Natsushi-ma-cho, Yokosuka, Kanagawa, 237-0061, Japan [email protected]
Tunnicliffe, Verena – School of Earth and Ocean Sciences, andDepartment of Biology, University of Victoria, VictoriaV8W 2Y2, Canada [email protected]
Tyler, Paul A. – Department of Oceanography, SouthamptonOceanography Centre, Southampton, SO14 3ZH, UnitedKingdom [email protected]
Vacelet, Jean – Station Marine d’Endoume, Rue de la Batteriedes lions, 13007 Marseille, France [email protected]
Van Dover, Cindy Lee – Biology Department, The College ofWilliam & Mary Williamsburg, VA 23187, [email protected]
Verschelde, Dominick – Universiteit Gent, Museum voor Dier-kunde, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium [email protected]
Vervoort, Willem – National Museum of Natural History, P.O.Box 9517, 230 RA Leiden, The Netherlands [email protected]
Villanueva, Roger – Instituto de Ciencias del Mar (CSIC)Insti-tuto de Ciencias del Mar (CSIC), Paseo Juan de Borbon s/nE-08039 Barcelona, Spain [email protected]
Voight, Janet R. – Department of Zoology, The Field Museum ofNatural History, Roosevelt Road at Lake Shore Drive, Chi-cago, IL 60605, USA [email protected]
Vrijenhoek, Robert C. – Monterey Bay Aquarium Research In-stitute, 7700 Sandholdt Road, Moss Landing, CA 95039-9644, USA [email protected]
Warén, Anders – Swedish Museum of Natural History, Depart-ment of Invertebrate Zoology, Box 50 007, Stockholm, S-10405 Sweden [email protected]
Wilson, George D. F. – Australian Museum, 6 College Street,Sydney NSW 2010, Australia [email protected]
Yamaguchi, Toshiyuki – Chiba University, Marine BiosystemsResearch Center 1-33, Yayoi-cho, Inage, Chiba 263-8522,Japan [email protected]
Young, Craig – Oregon Institute of Marine Biology, Universityof Oregon, Charleston, OR 97420, USA [email protected]
Zekely, Julia – Department of Marine Biology, University ofVienna, Althanstr. 14, A-1090 Vienna, [email protected]
Zibrovius, Helmut – Université d’Aix-Marseille 2, Station Ma-rine d’Endoume, Rue de la Batterie des Lions, 13007 Mars-eille, France [email protected]
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Seafloor Spreading and Hot Springs
Any account of the discovery of hydrothermal-vent faunasmust begin with the geology of seafloor spreading centers. Thesymmetry of magnetic anomalies on either side of the mid-oceanridges that girdle the globe and the correspondence of the anom-aly patterns with the pattern of magnetic reversals on Earth con-firmed the process of seafloor spreading and led to general ac-ceptance of plate tectonic theory in the early 1960s (VINE &MATTHEWS 1963). The bathymetric relief of mid-ocean ridgeswas understood to be a consequence of the thermal buoyancy ofhot rock in volcanic systems. Conductive heat loss was expect-ed to be greatest at the axis of these linear volcanoes and to di-minish along transects away from the ridge crest, but heat-flowmeasurements collected by placing a vertical array of thermistorsinto seafloor sediments consistently documented a heat-deficitnear the ridge axis. This heat deficit suggested that conductiveheat loss was not the only operative mode of cooling (STEIN etal. 1995). Geologists hypothesized that the convective heatdriven by cells of seawater percolating into the crust and subse-quent heating and buoyancy-driven flux of fluids out of the crustwould account for the missing heat. Thus the presence of hotsprings on the seafloor with temperatures as great as 300°C wasanticipated (TALWANI et al. 1971; LISTER 1972; WOLERY & SLEEP
1976), and the first unequivocal evidence of warm-water, buoy-ant plumes was collected by May 1976 using the Scripps Institu-tion of Oceanography Deep-Tow vehicle (WEISS et al. 1977).While systematic geophysical studies predicted the existence ofhydrothermal vents on the seafloor, the discovery of their atten-dant chemosynthetic ecosystems was unimagined.
Strange Animals at Hydrothermal Vents onthe Galapagos Spreading Center
The study of chemosynthetic ecosystems in the deep seadates back to a Black and white photograph of large white clamshells lying within cracks in a pavement of Black basalt that wascaptured by the Deep-Tow camera system at 13:20:39 (GMT) on29 May 1976 at 0°47‘84‘‘N, 86°09‘18‘‘W (Galapagos SpreadingCenter, eastern Pacific Ocean; LONSDALE 1977). Given that bi-valves are generally suspension feeders, the diet of the clams wasinferred to have been organic particulates concentrated by bot-tom-water currents (convection cells) induced by hydrothermalactivity (LONSDALE 1977; ENRIGHT et al. 1981).
During Alvin dives at the Galapagos Spreading Center in1977, geologist Jack Corliss first described vent mussels (al-though he erred in his initial identification). His difficulty infinding the words to describe what was before him, a vista noone had ever seen before, is evident:
“They are abalone shells. They are shells. They are bigshells. They are living. … attached shellfish….”
The wonder in his voice is captured on the audio record ofthe dive.
Further into the dive series, geochemist John Edmond firstsounds incredulous as he provided the first description of giantworms, and then frustrated as his observations were limited bythe green light of the thallium iodide bulb on Alvin:
“There are big ones [worms] out there. Looks like an Indian[Native American] headdress. There are four of them in a row,right outside my view port. I wonder what color they are. Youcan’t tell. They may be red.”
Vent animals were useful as flux indicators of hydrothermalactivity for geologists prospecting for vents, but geologists alsoappreciated the need for an explanation of the tremendous bio-mass of animals and of the means by which species could bemaintained at vents in the face of inevitable local extinctions.Following the first Alvin dive series to Galapagos vents, micro-biologist John Baross postulated that the millimolar concentra-tions of hydrogen sulfide in vent fluids were a source of reducedsulfur for free-living chemolithoautotrophic, sulfur-oxidizingbacteria (cited in CORLISS et al. 1979). Chemolithoautotrophsare primary producers that use the energy from the oxidation ofreduced compounds like hydrogen sulfide to yield ATP. ThatATP is then used to fix inorganic carbon (CO2) into organiccarbon. This contrasts with photosynthetic processes, where itis light energy that is harvested to produce ATP. Subsequentbiochemical steps in the production of organic carbon can beidentical in chemoautotrophs and photoautotrophs. Discoveryof vents thus led us to understand for the first time that com-plex food webs could be dependent on microbial chemosyn-thetic primary production. The initial dives to deep-sea ventsalso sparked the hypothesis that life on Earth may have origi-nated at submarine hot springs (CORLISS et al. 1980).
The triptych of charismatic vent organisms – clams (Calyp-togena magnifica), mussels (Bathymodiolus thermophilus), and gi-
Milestones in the discovery of hydrothermal-vent faunas
C.L. VAN DOVER, M. BISCOITO, A. GEBRUK, J. HASHIMOTO, V. TUNNICLIFFE,P. TYLER & D. DESBRUYÈRES Denisia 18 (2006) 13–25
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ant tubeworms (Riftia pachyptila) – has sustained scientific inter-est since their discovery (BALLARD 1977) and the first field in-vestigations by biologists (GRASSLE et al. 1979). These organ-isms, together with “dandelions” (siphonophores: Thermopaliataraxaca), “spaghetti worms” (enteropneusts: Saxipendium coro-natum), limpets, crabs, amphipods, polychaetes, and other ani-mals collected from Galapagos vents in the earliest collections(HESSLER & SMITHEY 1983), established the existence of a faunaendemic to and specialized for life in chemosynthetic ecosys-tems. They provided a baseline against which all other vent fau-nas would be compared. Our understanding of species zonation,of food webs, and even of succession of megafaunal species atvents in the eastern Pacific dates back to reports from these firstgeological and biological expeditions to the Galapagos Spread-ing Center.
Early Investigations on the East Pacific Rise
Early opportunities for ecological comparisons came with dis-covery and description of vent fields at 21°N [RISE Program in1979 (Rise Project Group 1980); Oasis Expedition (HESSLER etal. 1985)] and 13°N [Biocyatherm (1982) and Biocyarise (1984)Expeditions; DESBRUYÈRES et al. 1982, LAUBIER & DESBRUYÈRES
1985] on the nearby northern East Pacific Rise (NEPR). Similar-ities between the invertebrate faunas at Galapagos and NEPRvent fields indicated that, despite the restricted, insular nature ofthe benthic vent communities, exchange between populationson the two ridge axes was sufficient to sustain relatively largespecies ranges (Rise Project Group 1980). The NEPR vent fieldsincluded a habitat – the warm-to-hot (20 to >110 °C), leaky sur-faces of Black smoker chimneys – not present in Galapagos ventfields. At least some of the difference between Galapagos andNEPR species lists derives from the addition of species adapted tothe warmer waters of the chimney habitat (DESBRUYÈRES &LAUBIER 1980, DESBRUYÈRES et al. 1982; FUSTEC et al. 1987), in-cluding the Pompeii worms (Alvinella pompejana and A. caudata)and a brachyuran crab (Cyanagraea praedator). Faunal similaritiesalong the East Pacific Rise are now known to extend from thenorthern limit of the ridge system (Guaymas Basin) to 19°S (JU-NIPER et al. 1990; BLACK et al. 1994; GEISTDOERFER et al. 1995).Vents in Guaymas Basin are exceptional in that they are one ofthe few localities along the mid-ocean ridge system where hy-drothermal activity and volcanic eruptions take place in associa-tion with thick layers of pelagic and terrigenous sediment ratherthan bare basalt (EINSELE et al. 1980). The nominal transition ofthe southern East Pacific Rise to the Pacific Antarctic Ridge oc-curs south of the Easter and Juan Fernandez microplates, i.e., at~37°S. This region lies at the boundary between the Indo-Pacif-ic and Antarctic marine biogeographic provinces (VINOGRADO-VA 1979); the region also appears to be a boundary region forsome vent taxa (HURTADO et al. 2004). Bathymodiolid musselsand bythograeid crabs from 32°S, for example, are sister speciesto their northern counterparts (GUINOT & HURTADO 2003; WON
et al. 2003b), but gene flow in alvinellid polychaetes (Alvinella
pompejana) and commensal polynoid polychaetes (Branchipolynoesymmytilida) is unimpeded across the microplates (HURTADO etal. 2004).
Samples from American and French cruises to Galapagos,21°N, and 13°N vent sites between 1979 and 1985 became thefoundation for a noteworthy series of studies, in which the re-markable anatomy, physiology, and trophic ecology of manyrepresentative species of vent organisms was established (seeJONES & BRIGHT 1985 for an exhaustive bibliography for thisperiod). The important role of endosymbiotic, sulfur-oxidizingbacteria in the nutrition of tubeworms (Riftia pachyptila), for ex-ample, was established in 1981 through morphological (JONES
1981), ultra-structural (CAVANAUGH et al. 1981), and biochem-ical studies (FELBECK et al. 1981). The paradox of large animalsliving in waters with sulfide concentrations normally toxic tometazoans was also largely resolved by 1985, with descriptionsof detoxification mechanisms, including symbiont consumptionof sulfide, sulfide-binding proteins in tubeworms (ARP & CHIL-DRESS 1983; POWELL & SOMERO 1983) and clams (ARP et al.1984), and high activities of sulfide-oxidizing enzymes in super-ficial cell layers (POWELL & SOMERO 1986).
Northeast Pacific Vents
The 1983 exploration of hydrothermal vents at AxialSeamount on the Juan de Fuca Ridge (CASM 1985; TUNNI-CLIFFE et al. 1985) and subsequent studies of vent faunas on theExplorer (TUNNICLIFFE et al. 1986) and Gorda Ridges (VAN
DOVER et al. 1990) in the northeast Pacific provided the firstevidence that the hydrothermal-vent fauna was not globallycosmopolitan at the species level (CASM 1985). Faunal al-liances between the EPR and the NE Pacific vent systems arerecognized at the level of genus and higher (TUNNICLIFFE 1988),although several major taxonomic groups commonly found athydrothermal vents on the EPR are so far conspicuously absent(e.g., alvinocarid shrimp, lysianassid amphipods) at NE Pacificvents. Some alliances between the East Pacific Rise and NE Pa-cific vent faunas may reflect the paleotectonic history of theEast Pacific Rise and NE Pacific ridge system, which were oncepart of a single continuous ridge system before the override ofthe North American Plate (TUNNICLIFFE 1988).
Subsequent explorations along the Juan de Fuca Ridge re-vealed vent assemblages in a wide variety of venting conditions:extensive Black smoker fields at Endeavour (e.g., SARRAZIN etal. 1997), sedimented sulfide mounds at Middle Valley (e.g., JU-NIPER et al. 1992), and recent eruptive lavas of the Cleft Seg-ment (e.g., TSURAMI & TUNNICLIFFE 2001). From ExplorerRidge to Gorda Ridge, eight major vent fields provide biologistswith settings that vary in depth, age, substratum and the rela-tive importance of volcanism versus tectonism. In recent years,studies of community dynamics on sulfide edifices of Juan deFuca vents have given us a view of temporally and spatiallyshifting species populations in response to physical and chemi-
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cal changes in the environment and to biological interactions(e.g., SARRAZIN & JUNIPER 1999). The proximity of NE Pacificvent sites to coastal ports has fostered studies that examine tem-poral change and community comparisons.
Discoveries in the Atlantic
Discovery of shrimp- (Rimicaris exoculata) dominated ventfields at TAG (36°N; RONA et al. 1986) and Snake Pit in 1985(23°N; ODP Leg 106 Scientific Party 1986; MEVEL et al. 1989)on the Mid-Atlantic Ridge (MAR) highlighted the fact thatsimilar types of organisms might be found at hydrothermal ventsthroughout the world’s oceans (e.g., alvinocarid shrimp, bathy-modioliform mussels, siphonostome copepods), but that thespecies, and even many genera, are different from one oceanbasin to another. Major taxonomic groups familiar from EPRvents (e.g., vestimentiferan tubeworms, alvinellid polychaetes,stalked barnacles) are so far unknown at the MAR vents. Incontrast to the dynamic succession of macrofaunal species ob-served in some NE Pacific vent habitats (e.g., SARRAZIN & JU-NIPER 1999) and of megafaunal invertebrates at EPR vents (e.g.,SHANK et al. 1998), communities at MAR vent fields are re-markably stable on decadal time scales (COPLEY et al. 1997,1999).
The visual impact of vent megafauna is different betweenMAR and eastern Pacific vents: motile shrimp at MAR ventswere described by John Edmond as looking “like maggotsswarming on a hunk of rotten meat”; sessile worms and bivalvesat EPR and NE Pacific vents lend the sites a more garden-likeserenity, as evoked by names like Rose Garden and Garden ofEden. There are differences as well in the details of the bacter-ial-invertebrate symbioses implicated in the nourishment of thedominant megafauna between the two ocean basins. Episym-bionts likely contribute to the nutrition of rimicarid shrimp onthe MAR (VAN DOVER et al. 1988; GAL’CHENKO 1989; GEBRUK
et al. 1992; SEGONZAC et al. 1993), whereas tubeworms, clams,and mussels that dominate EPR vents rely on endosymbionts.MAR mussels have dual endosymbionts (methanotrophs andthiotrophs), while EPR mussels have only thiotrophic en-dosymbionts in their gills (FIALA-MEDIONI 1984; LE PENNEC &HILY 1984). Discovery of shrimp swarms at Mid-Atlantic Ridgevents ultimately led to the description of modified eyes in Rim-icaris exoculata and other alvinocarid shrimp, eyes that areadapted for detecting dim sources of light (VAN DOVER et al.1989; WHITE et al. 2002).
Western Pacific Explorations
Hydrothermal vents in Manus Basin were discovered in1985 (BOTH et al. 1986), the same year that vents on the Mid-Atlantic Ridge were reported (RONA et al. 1986). Further at-tention was brought to the southwestern Pacific in 1987 withthe first description of vent faunas from the Mariana back-arcspreading center (HESSLER et al. 1988, HESSLER & LONSDALE
1991). Back-arc spreading centers are zones of extension ofoceanic crust between active and remnant volcanic arcs associ-ated with subduction zones. Because back-arc systems tend tobe short, relatively young, and isolated (i.e., separated by largedistances from the nearly continuous mid-ocean ridge systems),fauna distinct from that of EPR vents, but evolved to occupysimilar niches was postulated (HESSLER & LONSDALE 1988).Dense clusters of large, “hairy gastropods” (Alviniconcha hessleri)with chemoautotrophic endosymbionts in their gills (STEIN etal. 1988) proved to be one of the most distinctive species, be-longing to a new genus and having trophic attributes not previ-ously known within the Gastropoda.
Although Mariana vent organisms belonged to undescribedspecies, more than half of the genera found there were alreadyknown from vent fields in the eastern Pacific. A mussel in thegenus Bathymodiolus dominated the biomass; other familiar an-imal types included polychaete worms in the genus Paralvinellaand limpets in the genus Lepetodrilus. Affinities of some Mari-ana taxa were shared with species known from Atlantic hy-drothermal vents – for example, shrimp in the genus Chorocaris.There were no surprises in terms of fidelity to a particular nicheor microhabitat in genera shared between Mariana and othermid-ocean ridge vents; the implication is that speciation haslargely been passive within these genera, a consequence of iso-lation by distance and barriers to dispersal, rather than of adap-tive radiations into new niches. The Mariana studies providedthe first incontestable evidence for faunal interchange and forbarriers or filters to dispersal of species between Pacific back-arcbasins and the mid-ocean ridge system (HESSLER & LONSDALE
1988). They also provided compelling evidence for the poten-tial for discovery of unanticipated taxa (in this case, the hairygastropods) as new geographic regions are explored.
Subsequent explorations of other southwestern Pacificback-arc systems [Manus (AUZENDE et al. 1997; HASHIMOTO etal. 1999); North Fiji and Lau Basins (HASHIMOTO et al. 1989;DESBRUYÈRES et al. 1994); New Ireland Basin (HERZIG et al.1994)] demonstrated that some species have restricted distribu-tions while other species, such as provannid gastropods, areshared among basins. Molecular techniques have now distin-guished four species of alviniconchid snails: Alviniconcha hessleriat Mariana vents, two Alviniconcha species in the Manus andNorth Fiji Basins, and a fourth Alviniconcha species at IndianOcean vents (KOJIMA et al. 2003; OKUTANI et al. 2004). In con-trast to the basin-scale differentiation of alviniconchid gas-tropods in the southwestern Pacific, Ifremeria nautilei (synony-mous with Olgaconcha tufari) so far is only known to inhabitvent sites in the Manus, North Fiji, and Lau Basin region (DES-BRUYÈRES et al. 1994; HASHIMOTO et al. 1999). The relativelynarrow distributional range of Ifremeria nautilei might be attrib-uted its lower dispersal ability compared to that of Alviniconchaspecies (KOJIMA et al. 2000, 2001). Ifremeria nautilei is also theonly known gastropod that supports dual symbioses, withmethane- and sulfur-oxidizing bacteria in gill bacteriocytes(GAL’CHENKO et al. 1992).
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Japanese biological explorations of deep-sea hydrothermalsystems in the northwestern Pacific began in 1988 with the dis-covery of active hydrothermal vents at Izena Hole (OkinawaTrough; TANAKA et al. 1990) by scientists using Shinkai 2000.Subsequent work discovered venting in yet another setting onthe arc volcanos of the Izu-Bonin seamount chain south ofJapan (KOJIMA 2002). Recent exploration of the Mariana Arcreveals variability in venting conditions and associated com-munities (EMBLEY et al. 2004). Together, the southwestern andnorthwestern Pacific hydrothermal fields represent a fertile re-gion for the study of population genetics and the biogeographyand evolution of vent taxa (DESBRUYÈRES et al. 2006).
Discovery of Seeps and OtherChemosynthetic Settings
The 1980s included other milestones: Seep communitiesdiscovered first in 1984 at the base of the Florida Escarpment inthe Gulf of Mexico (PAULL et al. 1984; HECKER 1985) and inSagami Bay (OKUTANI & EGAWA 1985; HASHIMOTO et al.1989), and then in association with subduction zone settings offOregon (SUESS et al. 1985), in the northwestern Pacific(LAUBIER et al. 1986; OHTA & LAUBIER 1987), and off Barbados(FAUGERES et al. 1987), taught us that vent-like taxa and troph-ically complex, chemosynthetically based ecosystems are notrestricted to hydrothermal settings. Discovery of chemosyn-thetic ecosystems and of vent- and seep-like taxa associatedwith whale skeletons on the seafloor (SMITH et al. 1989; re-viewed in SMITH & BACO 2003) underscored the need to adopta broad view of chemosynthetic faunas. Vent taxa were also dis-covered in diverse volcanic and hydrothermal settings, includ-ing alvinocarid shrimp at a mid-plate, hot-spot volcano (Loihiseamount; KARL et al. 1989; WILLIAMS & DOBBS 1995), alvinel-lids, mussels, crabs, and alvinocarid shrimp at Pito Seamount(NAAR et al. 2004), and relatively shallow (489 m) populationsof vesicomyid clams at Piips volcano (SELIVERSTOV et al. 1986;SAGALEVITCH et al. 1992). Thus we learn that vent and vent-like taxa and trophically complex, chemosynthetically basedecosystems are not restricted to mid-ocean ridge hydrothermalsettings, and that an understanding of the evolution and bio-geography of vent taxa can only be understood in the broadercontext of the evolution and biogeography of allied taxa at oth-er reducing environments (SIBUET & OLU 1998; VAN DOVER etal. 2002; TUNNICLIFFE et al. 2003a).
Other shallow hydrothermal settings were found to lack en-demic invertebrate species and do not support invertebrateswith endosymbiotic chemoautotrophic bacteria [e.g., subtidalvents off the coast of Southern California (KLEINSCHMIDT &TSCHAUDER 1985); vents of the volcanic arc in the Mediter-ranean (reviewed in DANDO et al. 1999); vents on the Kolbein-sy (Olafsson et al. 1989) and Jan Mayen ridges (FRICKE et al.1989) north of Iceland; vents in the crater of Ushishir volcanoof the Kamchatka region (TARASOV et al. 1990, TARASOV
2006)].
Hydrothermal vents also occur in fresh-water lakes. Fresh-water hydrothermal systems, similar to those in the ocean, arerelated either to rift zones, like lakes Baikal (CRANE et al. 1991)and Tanganyika (TIERCELIN et al. 1993), or volcanic activity insubduction zones, e.g. Crater Lake in Oregon (Anonymous1989). No examples of fresh-water, vent-specific metazoa areknown. Bacterial mats are common at all known fresh-watervents. Mineral structures analogous to Black smoker chimneys,reaching up to 10 m in height, can also develop (e.g., in CraterLake). The Russian-operated Pisces submersible studies in LakeBaikal documented aggregations of non-vent fauna in associa-tion with bacterial mats; carbon derived from bacterialmethane oxidation is incorporated into local food web of thebenthic fauna (GEBRUK et al. 1993).
Eruptions and the Hydrothermal Cycle
1991 was the year of the eruption at 9°50’N on the East Pa-cific Rise (HAYMON et al. 1993) and the beginning of a long-term study of hydrothermal cycles in this region (SHANK et al.1998). The bloom of bacteria that marked the commencementof the cycle (NELSON et al. 1991; HAYMON et al. 1993) and oth-er biological indications of recent volcanic activity were subse-quently observed at other locales, including the NE PacificRidges (e.g., TUNNICLIFFE et al. 1997; JUNIPER et al. 1998) and at17°S on the East Pacific Rise (EMBLEY et al. 1998). Repeated vis-its to developing vent communities documented the rapiditywith which colonization of new sites of venting takes place (e.g.,well-established colonies of tubeworms within one year; TUNNI-CLIFFE et al. 1997; SHANK et al. 1998) and confirmed earlier re-ports of rapid growth rates in vent species (LUTZ et al. 1994).Careful re-sampling of evolving vents allows insights into troph-ic strategies and the role of biological interactions such as com-petition (LEVESQUE et al. 2003).
The 9°50N vent field became a favored field site for eco-logical studies for several reasons: (1) the eruption took placethere, (2) its relatively simple topography has been thoroughlymapped, (3) its vents support a variety of foundation specieswith different habitat preferences (e.g., tubeworm clumps, mus-sel beds, alvinellid galleries, serpulid and anemone fields), (4)there is a long record of contemporaneous and complementarygeological and geochemical studies, and (5) the transit timefrom major ports in Mexico is relatively short. The 15+ years ofresearch at 9°50‘N since the eruption have resulted in impor-tant new insights into successional sequences (SHANK et al.1998), physiological adaptations to the vent environment (e.g.,GOFFREDI et al. 1997; SHILLITO et al. 2001), reproductive at-tributes of vent invertebrates (e.g., MARSH et al. 2000; PRADIL-LON et al. 2001), biological interactions among species (e.g.,MULLINEAUX et al. 2000; MICHELI et al. 2002; MULLINEAUX etal. 2003), animal-chemical relationships (e.g., LUTHER et al.2001; LE BRIS et al. 2006), population genetics (reviewed inVRIJENHOEK 1997), and community structure (e.g., VAN DOVER
2003; GOVENAR et al. 2004).
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One of the longest records of time-series studies at vents isthat for 13°N on the East Pacific Rise, where French and Amer-ican scientists have made intermittent observations since 1982(reviewed by DESBRUYÈRES 1995). At this site, a previously“dead” vent area with relic populations resumed activity in1987, and followed the successional pattern described for 9°N bySHANK et al. (1998). The presumption is that there was a sub-surface dyking event (i.e., an upward migration of molten rockthrough a fissure, but without overflow onto the seabed) in 1987that delivered heat to the system and restored the hydrothermalcirculation at this site.
More Atlantic Discoveries
Attention returned to the Mid-Atlantic Ridge in 1992 withthe chance collection of vent animals in a rock-dredge haul, andin 1993 during dives at the site that came to be known as LuckyStrike at 37°N (VAN DOVER et al. 1996; LANGMUIR et al. 1997).Although Lucky Strike shares a few species with the TAG andSnake Pit vent sites, most species at Lucky Strike were new toscience. This site and the nearby Menez Gwen vents explored in1993 (FOUQUET et al. 1995; DESBRUYÈRES et al. 2001) appear tobelong to a biogeographic province different from that of themore southerly TAG and Snake Pit faunas. Variables associatedwith the depth differential between the two provinces (<1700 mat Lucky Strike and Menez Gwen vs. >3000 m at TAG, SnakePit) have been implicated in the biogeographic differentiationof the two vent faunas (VAN DOVER et al. 1996; DESBRUYÈRES etal. 2000, 2001).
Continuing geological and geochemical exploration for hy-drothermal activity on the Mid-Atlantic Ridge between the Ve-ma transform fault at 11°N and the Azores hotspot (38°N) re-sulted in the first submersible dives to hydrothermal sites at Lo-gatchev in 1993 (BOGDANOV et al. 1995; GEBRUK et al. 2000),Broken Spur in 1993 (MURTON et al. 1995), and Rainbow in1997 (FOUQUET et al. 1997). The Logatchev site at 14°45’N wasthe first vent discovery where the host rock was ultramafic (i.e.,of mantle rather than crustal origin; BOGDANOV et al. 1997;FOUQUET et al. 2006). Logatchev represents the current south-ern limit of published information on vent faunas along theMid-Atlantic Ridge, although a new active vent field known asAshadze at 12°38’N has been reported (BELTENEV et al. 2004), ashave vent fields south of the equator (C. German, pers. comm).Logatchev is notable as the only known site on the Mid-At-lantic Ridge that supports large populations of vesicomyid clams;in other aspects, it has a species list similar to that found atSnake Pit (e.g., TURNIPSEED et al. 2004). Broken Spur (29°10’N)is a hybrid zone for mussels, where two species of mussels (Ba-thymodiolus puteoserpentis, dominant at southern locales, and B.azoricus, known from the northern vent sites) co-occur and in-ter-breed (WON et al. 2003a). Rainbow (FOUQUET et al. 1997),Saldanha (BARRIGA et al. 1998), and Lost City (KELLEY et al.2001) vent fields, like Logatchev, are hosted by ultramafic rocks.Their megafaunal and macrofaunal communities are generally
meager [except for dense shrimp Rimicaris exoculata populationsat Rainbow], possibly due to the distinctive fluid chemistries as-sociated with these sites (KELLEY et al. 2001).
The Indian Ocean
By 2000, a basic understanding of vent faunas of the north-ern Mid-Atlantic Ridge, the East Pacific Rise, the NortheastPacific, and back-arc basins in the western Pacific was estab-lished. One of the biggest missing pieces of the biogeographicjigsaw puzzle was the ridge system in the Indian Ocean. Cyanadives in the Gulf of Aden in 1984 (1400-1600 m) had docu-mented low-temperature flow dominated by shrimp, anemones,and galatheid squat lobsters, but specimens were not collected(JUNIPER et al. 1990). Japanese scientists brought attention tothe Indian Ocean in 2000, using the ROV Kaiko to explore theKairei hydrothermal field on the Central Indian Ridge, justnorth of the Rodriguez Triple Junction (HASHIMOTO et al.2001). Shrimp closely related to Rimicaris exoculata of Atlanticvents dominate the biomass of the Kairei field so that the siteresembles TAG and Snake Pit, but other taxa, including hairygastropods and mussels, are more closely allied with Pacific ventfaunas. These observations supported the hypothesis that theIndian Ocean ridges serve as a link between Atlantic and Pa-cific vent faunas (HASHIMOTO et al. 2001). Further explorationsof the Kairei Field (VAN DOVER et al. 2001) led to the discov-ery of a remarkable scaly-footed gastropod with no close al-liance to any known Pacific or Atlantic vent genera (Waren etal. 2003). Unlike other mollusks at vents that house autotroph-ic endosymbionts in epithelial bacteriocytes of their gills, thescaly-footed gastropod hosts its symbionts in a hypertrophiedesophageal gland (GOFFREDI et al. 2004).
Fossil Records
The first accounts of fossil vent invertebrates comes fromstudies of Cretaceous Oman and Cyprus ophiolites that yieldedwell-preserved tubes of worms, including what might be a ves-timentiferan (HAYMON et al. 1984; OUDIN & CONSTANTINOU
1984; HAYMON & KOSKI 1985). Invertebrates have occupiedvents (KUZNETSOV 1993; LITTLE et al. 1997) and seeps (BARBI-ERI et al. 2004) at least since the Silurian, but the ancient (old-er than the early Cretaceous) assemblages described so far weredominated by now-extinct families of brachiopods, monopla-cophorans, bivalves, and gastropods (reviewed in CAMPBELL
2006). There is little evidence to support the view that vents(or seeps) have served as a refuge during global extinctionevents for most taxa (reviewed in WARÉN & BOUCHET 2001;LITTLE & VRIJENHOEK 2003), although there may be exceptions(YAMAGUCHI et al. 2004). As in extant chemosynthetic com-munities, fossil records include indications of successional re-sponses to changing conditions at a given site (CAMPBELL et al.2002), and paleobiogeographic patterns that are likely to reflectpast plate tectonic configurations (CAMPBELL 2006).
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Prospects for Exploration and Discovery
There remain a number of oceanic regions where we cananticipate discovery of entire communities of new species at hy-drothermal vents (VAN DOVER et al. 2002; TYLER et al. 2003).Of these, the Arctic Ocean has long been of great interest: itsrelative youth and degree of bathymetric isolation from the restof the deep waters of the world’s oceans suggest that the ventfauna of the region might provide insight into the invasibilityof vent environments by metazoans. Hydrothermal vents on theGakkel Ridge in the Arctic Ocean have been localized (ED-MONDS et al. 2003), but their faunas remain to be photo-docu-mented and sampled. Shells of vesicomyid clams (Archivesicasp.), indicative of reducing environments, occur in dredge sam-ples from depths of ~2000 m near the Gakkel Ridge and seemlikely to have been associated with a hydrothermal site whenthe animals were alive (SIRENKO et al. 1995).
Other sites of considerable biogeographic and taxonomicinterest are isolated ridge systems, such as the Scotia Ridge inthe southwest Atlantic, the Cayman Rise in the Caribbean, andthe Andaman back-arc ridge in the northeast Indian Ocean.Even on the mid-ocean ridge system, there remain large stretch-es of unexplored ridge axis. One of the most compelling ridgesegments is the Mid-Atlantic Ridge north of the Azores – herethe ridge axis dives from shallowest depths back down to morethan 2500 m – has the Azores hot spot served as a faunal breakto north and south? The deep, greatly offset fracture zones ofthe Mid-Atlantic Ridge north of the Azores and south of theequator (i.e., south of the Romanche Fracture Zone) beg forstudies of gene flow and the role of these fracture zones as bar-riers or filters to species dispersal. The Chile Rise in the south-east Pacific and the Antarctic ridges are also certain to be of in-terest in comparative studies of taxonomy, diversity, and geneflow. Phylogeographic studies of the vent fauna of the tripleridge system of the Indian Ocean, in comparison to faunas ofthe Atlantic and western Pacific, should allow us to determinethe direction of dispersal of species between the Atlantic andPacific.
Even within extremely well-explored vent fields, the po-tential for discovery of previously unknown adaptations exists,as in the recent documentation of a blood-sucking polychaete(Galapagomystides aristata; JENKINS et al. 2002) that had beendescribed from some of the earliest collections at the GalapagosSpreading Center (BLAKE 1985). Little is known of the biologyof most vent species. There is great potential in careful studiesof populations, life history traits and behavior of individualspecies to learn more of the range of adaptations present inthese unusual habitats. Investigation of relationships of bio-mass-dominant metazoans to microbial communities and chem-ical settings will yield better insight into the complex processesthat control vent community structure and function (TUNNI-CLIFFE et al. 2003b).
The ChEss program (a component of the Census of Marine
Life) was conceived to promote international efforts to improveour understanding of the identity and biogeography of the fau-na of these reducing environments (TYLER et al. 2003). ChEssalso promotes tests of hypotheses regarding potential barriersand filters to dispersal of species whose adults are restricted tothese environments. Through ChEss and other ocean explo-ration activities, we look forward to the next decade of discov-ery that will undoubtedly add many new species, genera, andfamilies to the compendium of taxa so carefully compiled here.
Acknowledgements
We thank our many shipboard colleagues – Captains, pilots,ships’ crews, scientists, technicians, and students – who havecontributed to the discovery of deep-sea hot springs and of thestrange animals that live at them. There are too many individ-uals to cite by name; we honor their efforts with images of someof the many vehicles that have given us such remarkable accessto the seafloor (Fig. 1, 2).
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WILLIAMS A.G. & F.C. DOBBS (1995): A new genus and species of carideanshrimp (Crustacea: Decapoda: Bresiliidae) from hydrothermal ventson Loihi Seamount, Hawaii. — Proc. Biol. Soc. Wash. 108: 228-237.
WOLERY T.J. & N.H. SLEEP (1976): Hydrothermal circulation and geother-mal flux at mid-ocean ridges. — J. Geol. 84: 249-275.
WON Y., HALLAM S.J., O’MULLAN G.D. & R.C.VRIJENHOEK (2003a): Cytonucleardisequilibrium in a hybrid zone involving deep-sea hydrothermalvent mussels of the genus Bathymodiolus. — Mol. Ecol. 12: 3185-3190.
WON Y., YOUNG C.R., LUTZ R.A. & R.C. VRIJENHOEK (2003b): Dispersal barri-ers and isolation among deep-sea mussel populations (Mytilidae:Bathymodiolus) from eastern Pacific hydrothermal vents. — Mol.Ecol. 12: 169-184.
YAMAGUCHI T., NEWMAN W.A. & J. HASHIMOTO (2004): A cold seep barnacle(Cirripedia: Neolepadinae) from Japan and the age of the vent/seepfauna. — J. Mar. Biol. Ass. U.K. 84: 111-120.
23
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Fig. 1: Human-occupied vehicles (and maximum depth rating) in which many of the observations reported here were made. A: Alvin(4500 m), operated by the Woods Hole Oceanographic Institution, U.S.A.; B: Mir I and Mir II (6000 m), operated by the Shirshov Institute,Russia (by courtesy of Prof. A.M. Sagalevitch); C: Cyana (3000 m), operated by Ifremer, France; D: Nautile (6000 m), operated by Ifremer,France; E: Pisces IV (2000 m), formerly operated by the Department of Fisheries and Oceans, Canada, currently operated by the HawaiiUndersea Research Laboratory, U.S.A.; F: Shinkai 6500 (6500 m), operated by JAMSTEC, Japan.
A
C
B
D
E F
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Fig. 2: A selection of remotely-operated vehicles (andmaximum depth rating) used in deep-ocean exploration. A: ROPOS (5000 m), operated by the Canadian ScientificSubmersible Facility, Canada (by courtesy of V. Auger); B: Victor (6000 m), operated by Ifremer, France; C: Jason I(6000 m; retired); D: Jason II (6500 m), operated by theWoods Hole Oceanographic Institution, U.S.A.; E: Kaiko(11000 m), operated by JAMSTEC, Japan.
A
C
B
D
E F
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Meiofauna, small animals and protists, which pass througha net with 1 mm mesh size and are retained on a net with 63µm (or smaller) mesh size (GIERE 1993), is part of the hy-drothermal vent community. In general, very little is knownabout the diversity of vent meiofauna. So far, only a few phylasuch as the Arthropoda (Acariformes, Copepoda, Cumacea,Isopoda, Ostracoda, Tanaidacea), the foraminiferan Granu-loreticulosa, and the Nematoda have been described fromvents. In addition, Plathelminthes and three protist phyla in-cluding amoebae, colonial chrysophytes and ciliates are knownbut have yet to be described (SMALL & GROSS 1985; M. B.,pers. obs.). In contrast, other marine habitats, such as sedi-ments from shallow waters to the deep sea harbor far more di-verse meiofauna. For example gnathostomulids, gastrotriches,loriciferans, tardigrades, and small-sized representatives of hy-droids, bryozoans, nemertines, rotifers, or gastropods have notyet been discovered (HIGGINS & THIEL 1988; GIERE 1993).
All together, two foraminiferan, seven nematode, nine ost-racod, two acari, two cumacean, six tanaidacean, two isopod,and 78 copepod species are included in this edition. Only theCopepoda are known to a greater extent (IVANENKO & DEFAYE,this volume). Currently, meiofauna species contribute to about20% of the total diversity at vents. However, at large, the diver-sity of meiofauna is unknown. There are entire mid-ocean rid-ges or back-arc basins known for a relatively long time, in whichnot a single species of ostracod, tanaidacean, mite, or nematodeis described. Most known species are reported from a single site.The biogeographic range of meiofauna species is virtually un-
known. The few ostracod species described exclusively comefrom the eastern Pacific (DEEVEY 1969; KORNICKER 1969, 1991,KORNICKER & HARRISON-NELSON 2005; MADDOCKS 2006). Thefew tanaidaceans and the two isopod species were collected atthe site Lucky Strike at the Mid-Atlantic Ridge (CUNHA &WILSON, in press; LARSEN et al., in press); tanaidaceans were al-so observed at the site Rainbow (M. Segonzac, pers. comm.)Only one mite is described from Lau and North Fiji Back-ArcBasins (KRANTZ 1982) and the second species comes from theMid-Atlantic Ridge (BARTSCH 1990). Similarily, one cumaceanand one foraminiferan species each are decribed from the EastPacific Rise (BRÖNNIMANN et al. 1998; CORBERA this volume),the second species each come from the Mid-Atlantic Ridge(KAMENSKAYA et al. 2002; CORBERA this volume). The few des-cribed free-living nematode species exclusively were collected atsedimented vent sites from the East Pacific Rise, the GuaymasBasin, and the Lau and the North Fiji Back-Arc Basins (DE-CRAEMER & GOURBAULT 1998; VERSCHELDE et al. 1998).
To date there is not a single site for which the entire meio-fauna community structure is known. The picture emergingfrom the few meiofauna studies points to a vent meiofauna,which is low in abundance and diversity (DINET et al. 1988;SHIRAYAMA 1992; VANREUSEL et al. 1997; TSURUMI et al. 2003).Comprehensive studies are urgently needed not only to under-stand the community structure of meiofauna but also to provi-de insight in the biodiversity of the entire vent community andtheir trophic interactions.
Hydrothermal vent meiofauna
1, 2: Folliculinid ciliates from the East Pacific Rise: 9°N; by M. Bright.
M. BRIGHT Denisia 18 (2006): 27–28
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To our knowledge, vent meiofauna can be found in virtual-ly all vent habitats, from inactive bare basalt or sediments todiffuse flow areas up to Alvinella pompejana DESBRUYÈRES &LAUBIER, 1980 aggregations located on black smokers. Manyvent meiofauna animals such as copepods, nematodes, and ost-racods survive the transit from the vent environment at depthsof more than 2500 m to the water surface and can be maintai-ned without pressure at 4°C for at least a few days (M. B., pers.obs.).
Quantitative sampling of meiofauna is usually carried out insoft sediments with various types of corers. Hard substrate sam-pling and as well as sampling of megafauna aggregations such astubeworms clumps or mussel beds requires some sort of specialdesigned devices such as the “mussel pot” (VAN DOVER 2002)or the “Bushmaster Jr” (GOVENAR et al. 2005). Abundance andbiomass are standardized to 10 cm-2 surface area.
As there is no single technique available for extracting andfixing the entire meiofauna community quantitatively, the usu-ally applied compromise is either to sort through the entiresample including the sediment or to use various centrifugationtechniques in order to separate organic from inorganic materi-al. Fixation then is done by using 4% buffered formalin. Howe-ver, it has to be kept in mind that mostly the more robust so-called “hard” meiofauna such as copepods and nematodes isextracted and fixed and “soft” meiofauna such as plathelmin-thes or gastrotrichs, which is often not separated from the sedi-ment grains and/or is more delicate and sometimes requires ot-her fixation media, is lost. For storage, 70-80% ethanol is re-commended (see HIGGINS & THIEL 1988).
References:
BARTSCH I. (1990) Bull. Mus. Natl Hist. Nat., Paris, 4è sér., A. 12: 69-73.BRÖNNIMANN P., VAN DOVER C. & J.E. WHITTAKER (1989) Micropaleontology 35: 142-149.CUNHA M.R. & G.D.F. WILSON (in press) Zootaxa.DECRAEMER W. & N. GOURBAULT (1997) Zool. Scr. 26: 1-12.DEEVEY G.B. (1968) Proc. Biol. Soc. Wash. 81: 539-570.DINET A., GRASSLE F. & V. TUNNICLIFFE (1988) Oceanol. Acta 85: 7-14. GIERE O. (1993) Meiobenthology, the Microscopic Fauna in Aquatic Sediments. Springer Verlag: Berlin: 1-328.GOVENAR B., LE BRIS N., GOLLNER S., GLANVILLE J., APERGHIS A.B., HOURDEZ S. & C.R. FISHER (2005) Mar. Ecol. Prog. Ser. 305: 67-77.HIGGINS R.P. & H. THIEL (1988) Introduction to the Study of Meiofauna. Smithsonian Institution Press. Washington D.C., London: 1-488.KAMENSKAYA O.E., BAGIROV N.E. & T.G. SIMDIANOV (2002) Adaptation Aspects of Evolution of Marine Fauna. Collected Proc. Moscow VNIRO Publ. House:
144-152 [in Russian].KORNICKER L.S. & E. HARRISON-NELSON (2005) Zootaxa 1071: 19-38.KRANTZ G.W. (1982) Can. J. Zool. 6: 1728-1731.LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.MADDOCKS R.F. (2006) Micropaleontology 51: 345-372.SHIRAYAMA Y. (1992) Proc. JAMSTEC Symp. Deep Sea Res.: 287-290.SMALL E.B. & M.E. GROSS (1985) Bull. Biol. Soc. Wash. 6: 401-410.TSURUMI M., DE GRAAF R. C. & V.TUNNICLIFFE (2003) J. Mar. Biol. Ass. U. K. 83: 469-477.VAN DOVER C.L. (2002) Mar. Ecol. Prog. Ser. 230: 137-158.VANREUSEL A., VAN DE BORSCHES I. & F. THIERMANN (1997) Mar. Ecol. Prog. Ser. 157: 207-219.VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.
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Currently, very few macroparasites are known from hydro-thermal deep-sea vents. Only four copepods, Genesis vulcanoc-topusi (East Pacific Rise, LOPEZ-GONZALEZ et al. 2000), Cholidyapolypi (Endeavour Segment, HUMES & VOIGHT 1997), Ceuthoe-cetes aliger and Rimitantalus hirsutus (Galapagos Spreading Cen-ter, HUMES & DOJIRI 1980), one leech Bathybdella sawyeri (EastPacific Rise, BURRESON 1981; BURRESON & SEGONZAC, inpress), one acanthocephalan Hypoechinorhynchus thermaceri(East Pacific Rise, BURON 1988), and one nematode, Moravec-nema segonzaci (Mid-Atlantic Ridge, JUSTINE et al. 2002) havebeen reported from these sites. Significantly, each of these spe-cies was new to science. At least one monogenean and two di-geneans are currently being described by various specialistsfrom material collected on crabs and fishes collected from sou-thern East Pacific Rise sites and each of these also appear to bepreviously unreported species.
Because the concentration of potential hosts in a vent eco-system could possibly increase parasite transmission and diversi-ty, BURON & MORAND (2004) hypothesized that the low diversi-ty of macroparasites presently reported from these sites is likely areflection of these organisms having been overlooked by ventbiologists due to their inconspicuous nature. In support of thisidea, the use of molecular tools has shown the presence of a highdiversity of parasitic protists genetically linked to apicomplexans,perkinsozoans, syndiniales, and kinetoplastids (MOREIRA & LO-PEZ-GARCIA 2003). Additionally, Rickettsia-like inclusions havealso been found in numerous limpets from various vents (TERLIZ-ZI et al. 2004), and in mussels from Snake Pit and Logatchev si-tes (WARD et al. 2004) are further indications that parasite di-versity is vastly underreported from these important sites.
In brief, most often parasites must carefully looked for in or-der to be noticed. Hosts are various and diverse and range fromthe smallest of copepods to the largest of fish.
Since numerous parasites have a complex life cycle invol-ving more than one host, a better knowledge of the parasitevent fauna would help researchers to understand the trophicstructure present at each vent. Because trophically transmittedparasites are in many cases known to alter intermediate hostbehavior to increase capture by definitive hosts (LAFFERTY &MORRIS 1996) these organisms may play significant roles in in-creasing or directing the flow of energy into higher trophic le-vels at the vents. For example, acanthocephalans at the EastPacific Rise use fish as definitive hosts and appear to be trans-mitted via amphipod ingestion (I. B., pers. obs.): evidence fromother systems indicates that such infected amphipods exhibitaltered behavior (BAUER et al. 2005). Significantly, at the At-lantic sites these crustaceans were found to contain nematodelarvae of the ascarid genus Hysterothylacium although the adultswere not found (A. Petter, unpublished data).
Furthermore, non-trophically transmitted parasites presentat vents may also pose interesting problems for researchers. Forinstance, the leech Batybdella sawyeri, living among Riftia pachyp-tila and mussel Bathymodiolus thermophilus at the Galapagos Spre-ading Center, raises an interesting specificity problem: althougha parasite of fish, it has never been found on such hosts but wasobserved on vent crabs and bivalves from the southern East Pa-cific Rise (BURRESON & SEGONZAC, in press). Because of the wellknown specificity of many parasite-host relationships the recog-nition and identification of vent parasites may play an importantrole in deciphering vent colonization. In this regard it is impor-
Hydrothermal vent parasites
I. DE BURON & M. SEGONZAC Denisia 18 (2006): 29–30
1: Hirudinea Bathybdella sawyeri from East Pacific Rise: 17°S; cruiseBiospeedo (Ifremer, CNRS).
2: Nemertea Carcinonemertes sp., semi-parasite onbythograeid crabs from Lau Basin; cuise TUIMO6MV(MBARI); by G. Rouse.
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References:
BAUER A., HAINE E.R., PERROT-MINNOT J. & T. RIGAUD (2005) J. Zool. 267: 39-43.BURON I. DE & S. MORAND (2004) Parasitology 128: 1-6.BURON I. DE (1988) J. Parasitol. 74: 339-342.BURRESON E.M. & M. SEGONZAC (in press) Zootaxa.BURRESON E.M. (1981) Proc. Biol. Soc. Wash. 94: 483-491.HUMES A.G. & J.R. VOIGHT (1997) Ophelia 46: 65-81.HUMES A.G. & M. DOJIRI (1980) Proc. Biol. Soc.Wash. 93: 697-707.JUSTINE J.-L., CASSONE J. & A. PETTER (2002) Folia Parasitol. 49: 299-303.LAFFERTY K.D. & A.K. MORRIS (1996) Ecology 77(5): 1390-1397.LOPEZ-GONZALEZ P.J., BRESCIANI J. & R. HUYS (2000) Cah. Biol. Mar. 41: 241-253.MOREIRA D. & P. LOPEZ-GARCIA (2003) Trends Parasitol. 19(12): 556-558. PRITCHARD M.A. & G.O.W. KRUZE (1982) The Collection and Preservation of Animal Parasites. Univ. Nebraska, USA: 1-141.SHIELDS J.D. (2001) J. Crust. Biol. 21(1): 304-312.TERLIZZI C.M., WARD M.E. & C.L. VAN DOVER (2004) Diseases Aquat. Org. 62: 17-26.WARD M.E., SHIEDS J.D. & C.L. VAN DOVER (2004) Diseases Aquat. Org. 62: 1-16.
tant to note the presence of carcinonemertids (nemerteanworms) at vents. These worms are known to feed on the eggs ofshallow water decapods and were discovered on several vent crabspecies (SHIELDS 2001). Given the depth (1800-2400 m) and iso-lated nature of vent communities, the origin of vent colonizationby carcinonemertids has been raised and it has been speculatedthat infections were acquired from a more shallow-water host(see SHIELDS 2001 for review). Such egg predators were recentlycollected on several bythograeid crab species from western back-arc basins, and northern and southern East Pacific Rise (J.Shields & M. Segonzac, unpublished data). Moreover, because ofthe isolated nature of the vent ecosystems these sites will likelyprovide excellent sources of material for the study of the proces-ses of co-evolution in a relatively simple trophic system.
In summary, the study of parasitism in isolated vent com-munities should be emphasized not just because it would furt-her increase our understanding of the unique parasite faunapresent at these sites, but also because it would allow us to ad-dress and answer important questions regarding the coloniza-tion of vents and vent community structure as well as allow usto identify and recognize new strategies for parasite survival.
Parasites may be located almost anywhere on or in a host.Ideally, parasites should be isolated from their host while stillfresh and then prepared in a specific manner dependent on thetaxon collected (see, for example, PRITCHARD & KRUZE 1982for macroparasite fixation techniques). Importantly, parasitescollected from formaldehyde fixed hosts, such as museum spe-cimens, are most often of no use to taxonomic study.
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Granuloreticulosa, Foraminifera
Members of the order Foraminifera are widely distributed inmarine settings and can be found from polar regions to tropicallatitudes and from shallow water environments to the abyssalplains of the deep sea. More than 300 benthic foraminiferal ta-xa have been reported from deep-sea hydrothermal vent re-gions, but most of them are not restricted to these environ-
ments. Some foraminiferal species that have been collected invent regions also belong to planktonic species whose emptytests are accumulating on the sea floor. So far, only two end-emic foraminiferal species have been described from vent re-gions, which are listed in the present work.
References:
ARNOLD A.J., D’ESCRIVAN F. & W.C. PARKER (1985) J. Foraminiferal Res. 15: 38-42.MOLINA-CRUZ A. & A. AYALA-LOPEZ (1988) Geo-Mar. Lett. 8: 49-56.NIENSTEDT J.C. & A.J. ARNOLD (1988) J. Foraminiferal Res. 18: 237-249.QUINTERNO P.J. (1994) U.S. Geol. Surv. Bull. 2022 (Chpt.18): 337-359.AYALA-LOPEZ A. & A. MOLINA-CRUZ (1994) Journal of Micropaleontology 13: 133-146.JONASSON K.E, SCHRÖDER-ADAMS C.J. & R.T. PATTERSON (1995) Mar. Micropaleontol. 25: 151-167.JONASSON K.E. & C.J. SCHRÖDER-ADAMS (1996) J. Foraminiferal Res. 26: 137-149.
M. HOLZMANN Denisia 18 (2006): 31
1: Abyssotherma pacifica from East Pacific Rise: 9°N, Riftia Field; by M. Bright.
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32
Luffammina atlantica KAMENSKAYA, BAGIROV & SIMDIANOV, 2002
Size: Up to 2 cm in length.
Morphology: Agglutinated foraminifer with soft cylindricaltests are attached to the substrate by extended basal part. Up-per extended part of the test may be dichotomously divided.Test wall consists of fine detritus and agglutinated coccolithes.Surface of the test is covered by a thick layer of Methanotrix-likebacteria. Inner volume of the test with fine agglutinated parti-cles, diffuse cytoplasm and cavities. The genus Luffammina is
close to the genus Arborammina but differs from it by weakbranching of the upper test, absence of bead-like structures onthe test and absence of globogerins inside the test.
Biology: Found on the surface of relict hydrothermal chimneys.
Distribution: Mid-Atlantic Ridge, site Rainbow.
Reference:
KAMENSKAYA O.E., BAGIROV N.E. & T.G. SIMDIANOV (2002) in Adaptation Aspects of Evolution of Marine Fauna. Coll. Proc., Moscow VNIRO Publ. House:144-152 [in Russian].
M. HOLZMANN Denisia 18 (2006): 32
3: Specimens (arrows) on hydrothermal chimneys;by O. Kamenskaya.
4: Specimens (arrows) on hydrothermal chimneys; by O. Kamenskaya.
2: Test surface covered with Methanotrix-like bacteria; scalebar 100 µm; by O. Kamenskaya.
Granuloreticulosa, Foraminifera, Allogromida, Arboramminidae
1: Test; scale bar 1 mm; by O. Kamenskaya.
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Abyssotherma pacifica BRÖNNIMANN, VAN DOVER & WHITTAKER, 1989
Size: Maximum and minimum test diameter, 940 and 830 µm,respectively (holotype).
Morphology: Test free, a low watchglass-shaped trochospire.Adult chambers spirally elongate, umbilically asymmetric andmushroom-shaped; interior subdivided by secondary septaformed by infolding of inner organic sheet. Aperture double:primary opening interiomarginal, in strongly incurved anteriorflank of septum, anteriorly directed; secondary opening in axi-al-sutural position at tip of posterior flank of septum, posterior-ly directed, also interiomarginal. Wall imperforate, consistingof agglutinated layer between outer and inner organic sheets.
Biology: According to BRÖNNIMANN et al. (1989), A. pacificaoccurs in artificial recruitment arrays placed in the vicinity ofdeep-sea (2600 m) hydrothermal springs in the East Pacific.Water temperature immediately surrounding the arrays wasnear ambient. Abyssotherma pacifica occurs also associated withRiftia pachyptila and on bare basalt, suggesting a wide range oftemperature and sulfide tolerance (pers. comm. Bright).
Distribution: East Pacific Rise: 21°N (BRÖNNIMANN et al.1989) and 9°N (pers. comm. Bright).
References:
BRÖNNIMANN P., VAN DOVER C. & J.E. WHITTAKER (1989) Micropaleontol. 35: 142-149.LEE J.J., ANDERSON O.R., KARIM B. & J. BERI (1991) Micropaleontol. 37: 303-312.JONASSON K.E. & C.J. SCHRÖDER-ADAMS (1996) J. Foraminiferal Res. 26: 137-149.KAMINSKI M.E. (2000) Proc. 5th Intern. Workshop on agglutinated Foraminifera. Grzybowski Foundation Special Publication 7: 185-219.MICHELI F., PETTERSON C.H., MULLINEAUX L.S., FISHER C.R., MILLS S.W., SANCHO G., JOHNSON G.A. & H.S. LENIHAN (2002) Ecol. Monogr. 72(3): 365-382.MULLINEAUX L.S., PETTERSON C.H., MICHELI F.& S.W. MILLS (2003) Ecol. Monogr. 73(4): 523-542.
M. HOLZMANN Denisia 18 (2006): 33
1: Umbilical view x 80 © Micropaleontology.
2: Spiral view x 80 © Micropaleontology.
3: Close-up of agglutinated test, spi-ral view x 125 © Micropaleontology.
Granuloreticulosa, Foraminifera, Textulariida, Remaneicidae
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Porifera
Deep-sea representatives of the Porifera (sponges) belongmainly to the classes Hexactinellida and to a special family ofDemospongiae, class Poecilosclerida, the Cladorhizidae, al-though many other Demospongiae are able to live in bathyalenvironments. Hexactinellida and Cladorhizidae display twovery different life strategies allowing them to survive in deep-sea conditions where life is difficult for sponges, which are fil-ter-feeders living on tiny particles. The hexactinellids have ahighly developed filter-feeding system with large cavities linedby extremely thin living tissue to maximize the volume of fil-tered water and the ability for retention of particles. Converse-ly, the Cladorhizidae have lost their filter-feeding mode and re-ly on carnivory of relatively large prey, mostly crustaceans, avery unusual and unexpected mode of feeding in sponges. Thisstrategy allows cladorhizids to reach the hadal zone with adepth record of 8840 m, whereas the hexactinellids and “nor-mal” demosponges do not exceed 7000 m.
Both hexactinellids and cladorhizids are not members ofthe true vent communities. Up to now, they have never beenobserved in the immediate environment of active smokers.However, a few hexactinellids and numerous cladorhizids occurat few distance from the active vents in high diversity, and at ahigh proportion of undescribed species in the case ofcladorhizids (VACELET, in press). These carnivorous spongesbenefit, at least in part, from a general distant enrichmentaround the vents At the Lau sites, cladorhizid specimens caughttens of larval zoe of crab Austinograea alaysae (M. Segonzac,pers. obs.). It cannot be excluded, however, that they belong toa “non-vent” fauna taking advantage only of the presence ofhard substrata offered by the lava issued at rapidly spreading ar-eas and that their high diversity is due to a higher sampling ef-fort in these areas.
1: Abyssocladia sp. from Kilo Moana, Lau Back-Arc Basin, TUIM 05 cruise © M. Tivey.
J. VACELET Denisia 18 (2006): 35
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References:
HAJDU E. & J. VACELET (2002) in HOOPER J.N.A. & R.W.M VAN SOEST (Eds.) Systema Porifera: A Guide to the Classification of Sponges. Vol. 1: 636-641.VACELET J. (in press) Zool. J. Linn. Soc.
The cladorhizids are generally small sponges with a mor-phology rather unusual in sponges. They are erect, with a pin-nate or symmetrical shape, and their aquiferous system, longconsidered as a diagnostic characteristic of the phylum Porifera,has been discarded in most genera. A special mention, howev-er, should be made for the genus Chondrocladia, in which acanal system and choanocyte chambers are maintained, butserve to inflate large spheres on which the prey is captured. Theprey, mostly small crustaceans, are passively captured by meansof hook-like cheloid microscleres lining the surface of thesponge and catching the setae or appendages of various inver-tebrates, acting as a Velcro cover. In the absence of a digestivecavity, cells act individually to digest the prey.
The classification of the carnivorous sponges is presentlyrather problematic (HAJDU & VACELET 2002). They all belongto the order Poecilosclerida, and most are classified in the fam-ily Cladorhizidae, with several genera dinstinguished mainly bythe microsclere spicules. These microscleres, however, belongto different evolutionary lines of the Poecilosclerida. Further-more, it appears that a carnivorous feeding habit also exists insome other poecilosclerids displaying a similar morphology, butclassified in different families due to their different cheloid mi-croscleres. Molecular characters have not yet been used for thedistinction of the evolutionary lines of these sponges.
These deep-sea sponges are highly fragile. The hexactinel-lids, although supported by a highly developed siliceous skele-ton, have very thin, partly syncytial living tissue which is verydifficult to preserve properly. They must be preserved in goodhistological or cytological fixatives as soon as possible after col-lection. The carnivorous sponges have very fragile appendagesthat are generally only partially preserved during collection.Observation of their morphology from manned submersibles orROVs is important. For instance, a long time was needed to re-alize that a mysterious organism with pedunculate, translucentspheres actually corresponds to the genus Chondrocladia as itwas known from preserved specimens, in which the sphereswere contracted. Their histology and reproduction is poorlyknown and could be studied only from specimens preserved informalin or fixatives for electron microscopy. The present un-certainties in the distinction of evolutionary lines in carnivo-rous sponges require more information on their molecular char-acters, and the preservation of fragments in alcohol is highlydesirable.
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Abyssocladia dominalba VACELET, in press
Size: 30 mm high, with a body 10 mm in diameter.
Color: White in alcohol.
Morphology: A thin peduncle bearing an ovoid or subsphericalbody made of radiating fascicles. No aquiferous system. Skele-ton: Peduncle made of longitudinally arranged long fusiformstyles; radiating fascicles made of fusiform styles and smallerstyles with the tip outwardly directed. Spicules: fusiform styles,620-2500 x 7-35 µm; arcuate isochelae, 80-170 µm; abysso-chelae with the front alae long, nearly in contact with the op-posite ala, 40-45 µm; anisochelae, generally twisted, one endtridentate, the other end with fused alae, 9.5-11 µm; sigman-cistra in two classes, 30-40 µm and 9.5-12.5 µm.
Biology: On a dead smoker near an active site, water tempera-ture 2.6°C at the site of collection. A carnivorous feeding habithas been demonstrated in another species of the genus.
Distribution: North-Fiji Back Arc Basin: site White Lady.
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
J. VACELET & M. SEGONZAC Denisia 18 (2006): 37
1A: View of the holotype, scale bar 3.4 mm; B: Arcuate isochela 1, scale bar 12 µm; C: Arcuate isochela 1, scale bar 22 µm; D: Arcuate isochelae 2 (abyssochelae), scale bar 7.4 µm; E: Anisochelae, scale bar 2.5 µm; F: Sigmancistra 2, scale bar 2.2 µm; G: Sigmancistra 1, scale bar 4.4 µm; H: Sigmancistras 1 and 2, scale bar 4.4 µm; I: Style, scale bar 47 µm; from VACELET (in press).
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
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Abyssocladia naudur VACELET, in press
Size: Up to 40 mm high, with thin lateral filaments 6 mm long.
Color: White in situ, yellowish gray to clear brown in alcohol.
Morphology: Small erect sponge, forming a flattened axis withnumerous lateral filaments, frequently with a bud-like branch-ing process. Filaments regularly arranged in two lateral rows, al-ternating on each side. No visible aquiferous system. Skeleton:main axis of fusiform styles longitudinally arranged, lined bysubstrongyles at the base; axis of the processes conical at thebase, with the styles anchored by their head entirely crossingthe stem; stem and base of the filaments with a continuous lin-ing of isochelae. Spicules: Styles of the axis of the stem and fil-aments, fusiform, shorter in the filaments, 330-1600 x 5-37 µm;
substrongyles or strongyles of the basal coating, fusiform, bentor slightly flexuous, 30-825 x 8-30 µm; abyssochelae withfrontal alae roughly parallelepipedal, nearly in contact with theopposite frontal ala, 48-72 µm; sigmancistra in two classes, 15-19 µm and 5-8 µm.
Biology: Several specimens collected from a dead smoker, a fewmeters from active black smokers. Presumably with a carnivo-rous feeding habit.
Distribution: East Pacific Rise: 17°S.
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
J. VACELET & M. SEGONZAC Denisia 18 (2006): 38
1A: View of the holotype, scale bar 4.3 mm; B: Part of a paratype, scale bar 3.4 mm; C: Paratypes, scale bar 2 mm; D: Isochelae,scale bar 5.5 µm; E: Sigmancistra 1 and 2, scale bar 1.4 µm; F: Style of the axis, scale bar 64 µm; G: Styles of the lateral processes,scale bar 40 µm; H: Diverse sizes of substrongyles of the base, scale bar 33.4 µm; from VACELET (in press).
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
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Asbestopluma agglutinans VACELET, in press
Size: Axis 4 cm high and 0.8-1 mm in diameter, lateral fila-ments up to 5-6 mm long.
Color: Brown in alcohol.
Morphology: Small erect sponge, consisting of a flattened axissmooth on the base, then with biserially arranged filamentsarising perpendicularly to the axis in two opposite series. Fila-ments thick, cylindrical at the basis, then abruptly reduced to athin spicular axis. No visible aperture. Skeleton: axis of largefusiform styles longitudinally arranged, surrounded by a felt-work of acanthotylostrongyles including numerous skeletons ofradiolarians and foraminiferans; base with a cover of tangentialflexuous styles or strongyles. Spicules: fusiform styles of the ax-
is, 1550-2100 x 30-35 µm; mycalostyles of the filaments, 370-780 x 8.5-17 µm; styles or strongyles of the base, 220-535 x 15-42 µm; acanthotylostrongyles, 65-165 x 0.8-2.3 µm; aniso-chelae, 32-36 µm and 9.8-10.5 µm; sigmancistras, 23-28 µm.
Biology: Collected on a dead smoker and a basalt fragment.
Distribution: East Pacific Rise: collected at 18°S and 14°S.
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
J. VACELET & M. SEGONZAC Denisia 18 (2006): 39
1A: Holotype and paratype, scale bar 3.8 mm; B: Style of the axis, scale bar 83 µm; C: Head and tip of a style of the axis, scalebar 28 µm; D: Style of the filament axis, scale bar 35 µm; E: Substrongyle of the base, scale bar 37 µm; F: Acanthotylostrongyle,scale bar 7.5 µm; G: Head and tip of an acanthotylostrongyle, scale bar 2.3 µm; H: Anisochela 1, scale bar 4.1 µm; I: Anisochelae2, scale bar 2 µm; J: Anisochela 2, back view, scale bar 2 µm; K: Sigmancistra, scale bar 2.9 µm; from VACELET (in press).
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
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Asbestopluma formosa VACELET, in press
1: Fragment of the holotype, scale bar 10.2 mm;from VACELET (in press).
3A: Two fusiform styles of the axis, scale bar 65 µm; B:Style of the lateral processes, scale bar 21.5 µm; C:Substrongyle, scale bar 14.3 µm; D: Microstrongyleand detail of the head, scale bars 6.2 µm and 1.5 µm;E: Two anisochelae 1 and an immature one, scale bar10 µm; F: Two microtylostyles, scale bar 4.3 µm; G:Anisochelae 2, scale bar 2 µm; from VACELET (in press).
2: Collection of the holotype by the arm of “Nau-tile“ submersible, 1997 m; from VACELET (in press)© Ifremer/Starmer 1.
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
Size: Approximately 8 cm high and 16 cm wide, with stem andbranches up to 1 mm in diameter.
Color: White in situ and after preservation.
Morphology: Erect, fan shaped, consisting of a short stem fromwhich arise branches which divide dichotomously three or fourtime in a single plane, the last branches being long and paral-lel. Terminal branches flattened, bearing on both sides numer-ous, regularly spaced thin filaments. Flattened enlargementspresent at each dichotomy containing numerous reproductionbodies. No visible apertures or aquiferous system. Skeleton:Stem and branches with an axis of large fusiform styles with a
dense outer cover of microstrongyles, filaments with an axis ofsmaller styles; base of the stem with a cover of special microty-lostyles and short fusiform substrongyles. Spicules: Styles, 200-1025 x 20-45 µm and 180-350 x 7-9 µm; curved substrongyles,80-500 x 15-30 µm; microstrongyles minutely spinose, 25-60 x5-7 µm; microtylostyles, minutely spinose, 25-45 x 4-7 µm;anisochelae sometimes in rosettes, 72-90 µm and 10-15 µm.
Biology: On a fossil chimney ca. 50-100 m distant from an ac-tive site White Lady. Presumably with a carnivorous feedinghabit.
Distribution: North-Fiji Back-Arc Basin.
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
J. VACELET & M. SEGONZAC Denisia 18 (2006): 40
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Asbestopluma pennatula (SCHMIDT, 1875)
1A: Specimen from Lucky Strike, scale bar 5 mm; B: Style of the filament, scale bar 50 µm; C: Style of the filament andhead of style of the axis, scale bar 40 µm; D: Strongyle of the basis of the axis, scale bar 30 µm; E: Spinose mi-crostrongyle, scale bar 6 µm; F: Sigmancistra and anisochela II, scale bar 4 µm; G: Anisochela I, scale bar: 8 µm; H:Anisochela II, scale bar 2 µm; I: Basis of anisochela II and end of sigmancistras, scale bar 2 µm.
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
Size: 100 mm high. Lateral filaments up to 2.5 mm x 0.1-0.2mm, possibly longer but easily broken. Specimens up to 180mm with filaments 7 mm long have been recorded near Ice-land.
Color: Cream in situ and in alcohol.
Morphology: Erect axis with distinct groups of lateral filamentsarising perpendicularly, regularly spaced along the stalk. Noaquiferous system. Skeleton: spicular axis of styles in the stalkand in lateral filament, covered at the basis of the sponge by acoating of spinose microtylostrongyles. Spicules: Styles of theaxis of the stalk, fusiform, 700-920 x 17-22 µm, shorter andthicker at the basis of the axis where they gradually becomesubstrongyles 200-540 x 18-30 µm. Styles of the filaments,
slightly fusiform with an acerate end, straight, fusiform, 380-520 µm x 7-10 µm. Microtylostrongyles coating the surface ofthe axis near its basis, slightly flexuous, with a short spination,40-130 x 0.5-1 µm. Palmate anisochelae I, moderately abun-dant, longitudinally arranged with the teeth upwards on the fil-aments, with well marked lateral teeth in the small end, 33-40µm. Palmate anisochelae II, very abundant, perpendicular tothe surface of the filaments with the large tooth upwards, 9-10µm. Sigmancistras very abundant, 17-25 µm.
Biology: Attached to solid substrata in the bathyal zone.
Distribution: North Sea, Arctic, Bay of Biscay. Collected atMid-Atlantic Ridge: Lucky Strike and Menez Gwen.
References:
LUNDBECK W. (1905) The Danish Ingolf-Expedition 6(2): 1-219.
J. VACELET & M. SEGONZAC Denisia 18 (2006): 41
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Chondrocladia lampadiglobus VACELET, in press
Size: Up to approximately 50 cm high, with inflated spheres 3-5 cm in diameter.
Color: White in life, yellowish white to clear brown in alcohol.
Morphology: Large stalked sponge, composed of a rhizoid fixa-tion system, a cylindrical stalk ending in an enlarged, ovoidbody from which radiate in all directions secondary branches,each ending in translucent sphere in the living animal, in an ir-regular swelling including crustacean debris on the preservedspecimen. Aquiferous system present, with large choanocytechambers and canals ending in the inflatable spheres. Skeleton:stalk and branches made of large fusiform styles longitudinallyarranged, covered with a feltwork of rugose tylostyles; terminalswellings made of smaller styles with an outer cover of numer-ous microscleres. Spicules: fusiform styles, 700-4750 x 15-75 µmand 510-580 x 17-30 µm; rugose tylostyles, 300-535 x 5-6 µm;anchorate isochelae in two sizes, with six alae at each end, 123-
140 µm, alae 25 µm long, and 20-32 µm, alae 10-11 µm long;sigmas 45-120 x 2-3 µm.
Biology: The collected specimen was rooted in sediment be-tween pillow lava, near active hydrothermal sites, but in anarea still with low density of animal life. Sponges of similarmorphology have been often observed on various sites of theEast Pacific Rise, either rooted in sediment or attached to pil-low lava, always at some distance from the rich animal com-munities of the active hydrothermal sites. Their identificationto C. lampadiglobus cannot be ascertained from external mor-phology alone, and several species may be present in this largegeographic zone. Carnivorous mode of feeding.
Distribution: East Pacific Rise: collected at 17°S. Sponges ofsimilar morphology observed in various sites extending from23°S to 13°N, 2600-3000 m deep.
1: ROV Tiburon/2003, dive 556, 20°47.03’N, 109°08.98’W, 2555 m; by R.Vrijenhoek © MBARI.
2A: Holotype; scale bar 17 mm; B: Tylostyle of thecover of the stalk; scale bars: 42 µm and 4.2 µm; C:Style of the stalk; scale bar 30 µm; D: Style of thebody surface; scale bar 63 µm; E: Anchorate isochelae1; scale bar 14 µm; F: Anchorate isochelae 1; scale bar9.6 µm; G: Two anchorate isochelae 2; scale bar 7 µm;H: Sigma; scale bar 10 µm; from VACELET (in press).
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
J. VACELET & M. SEGONZAC Denisia 18 (2006): 42–43
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Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
3A: Holotype, East Pacific Rise: 17°S, 2714 m © Ifremer/Naudur; inset: Unidentified worm gliding on the lower left sphere; B: Collection of the holotype by the “Nautile“ submersible © Ifremer/Naudur; C: Presumed C. lampadiglobus, Geocyarise 3 (CY 30), 2622 m, 12°54’N, 103°58’W; cruise Geocyarise © Ifremer; D: Presumed C. lampadiglobus, Geocyarise 1 (CY 07), 2623 m;© Ifremer; modified from VACELET (in press).
43
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Cladorhiza abyssicola G.O. SARS, 1872
References:
LUNDBECK W. (1905) The Danish Ingolf-Expedition 6(2): 1-219.SARS G.O. (1872) in Kongelige Norske Universitet (Ed.) Spongiae. Volume I. Brøgger & Christie, Christiania, Norway: 1-82.
J. VACELET & M. SEGONZAC Denisia 18 (2006): 44
1A: Type specimen from Lofoten, 550 m, approximately 60 mm high, from SARS (1872); B: Specimen from MAR, Logatchevvent site, DiversExpedition, dive 3668, 3012 m, with oocytes and embryo; scale bar 3 mm; C: Style and anisochelae; scalebar 30 µm; D: Anisochela; scale bar 5 µm; E: Sigmas; scale bar 25 µm; F: Sigmancistra; scale bar 8 µm.
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
Size: Up to 75 mm high.
Color: Cream white in alcohol.
Morphology: Erect, with numerous lateral branches arising atnearly right angle from a central axis and bearing short second-ary processes, generally anchored in the sediment by richlybranched roots. No aquiferous system. Main skeleton: polyspic-ular fibres of styles in the axis of stem, branches, lateral process-es and roots. Spicules: Styles, fusiform, 390-730 µm x 14-22 µm;
unguiferate anisochelae with five teeth at each end, 21-25 µm;sigmas, 78-100 µm; sigmancistras, 40-42 µm.
Biology: A common species in the bathyal and abyssal zones,usually anchored in mud by the roots, with a carnivorous modeof feeding. A near relative, C. methanophila VACELET & BOURY-ESNAULT, 2002, is both carnivorous and symbiotic withmethanophilous bacteria.
Distribution: Atlantic N.E., Arctic, Mediterranean, Mid-At-lantic Ridge.
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Cladorhiza segonzaci VACELET, in press
Size: Axis up to 32 mm high and 0.4-0.5 mm in diameter, withlateral processes 4-9 mm long.
Color: In situ white; cream to clear brown in alcohol, with thebasal portion of the stem darker.
Morphology: Small erect sponge, forming an unbranched axiswith numerous lateral processes arranged in opposed series atright angle all around the axis. No aquiferous system. Skeleton:main axis of fusiform styles longitudinally arranged; axis of theprocesses conical at the base, with the styles anchored by theirhead reaching the centre of the stem and the point outwardlydirected. Spicules: Styles of the axis of the stem and lateral
processes, slightly fusiform, 380-990 x 14-23 µm; anchorate/un-guiferate anisochelae numerous, with five lanceolate alae andwell developed fimbriae at the large end, and three fang-likealae at the small end, 15-17.5 µm; sigmas 50-80 x 1-1.5 µm; sig-mancistras without notch, 20-25 x 2.0 µm.
Biology: 11 specimens collected from a dead smoker, a few me-ters from active black smokers. Presumably carnivorous mode offeeding.
Distribution: East Pacific Rise: 17°S.
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
J. VACELET & M. SEGONZAC Denisia 18 (2006): 45
1A: View of the holotype (left), two paratypes and a fragment of a paratype; scale bar 3 mm; B: Style; scale bar 90 µm; C: Developmental stage of anisochela; scale bar 3.1 µm; D: Anisochela; scale bar 3.7 µm; E: Anisochela, back view; scale bar 3.2 µm; F: Sigma; scale bar 12 µm; G: Sigmancistra; scale bar 5.3 µm; H: Sigmancistra; scale bar 5.3 µm; from VACELET (in press).
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
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Euchelipluma pristina TOPSENT, 1909
References:
TOPSENT E. (1909) Bull. Inst. Océanogr. Monaco 151: 1-23. TOPSENT E. (1928) Résult. Camp. Sci. Prince Albert Ier de Monaco 74: 1-376.
J. VACELET & M. SEGONZAC Denisia 18 (2006): 46
1A: Two specimensfrom Barbados; scalebar 4 mm; B: Head of a subty-lostyle, Barbados;scale bar 10 µm; C: Subtylostyles,placochelae I and II,isochela, sigmancistrasI and II, Barbados;scale bar 50 µm; D: Immatureplacochela, holotype;scale bar 10 µm; E: Placochela, holo-type; scale bar 10 µm;F: Placochelae,Barbados; scale bar 15 µm; G: Placochela,Barbados; scale bar 5 µm; H: Sigmancistra I,Barbados; scale bar 2 µm; I: Isochela, Barbados;scale bar 15 µm.
Porifera, Demospongiae, Poecilosclerida, Guitarridae
Size: 14-120 mm high, shaft 3-4 mm wide, lateral filaments upto at least 8 mm long and 70-100 µm in diameter.
Color: Clear brown in alcohol.
Morphology: Erect, pennaceous, composed of a short peduncleand a flattened shaft bearing symmetrically paired lateral fila-ments, broken and reduced to their basis in most specimens. Noaquiferous system. Skeleton a spicular axis of styles, condensedat the basis and divided in parallel fibres upward; lateral fila-ments with a twisted axis of subtylostyles. Microscleres most of-ten regularly arranged at the surface of the shaft and of the fil-aments, with the teeth outwardly directed. Spicules: Styles ofthe axis, fusiform with an obtuse point, 900-1550 x 15-40 µm;strongyles in the basis of the peduncle, slightly curved or flexu-ous, 210-490 x 14-22 µm; subtylostyles in the filaments, 330-
700 x 6-14 µm; palmate isochelae, 75-130 µm, in two size cate-gories in some specimens; placochelae I, 60-90 µm; placochelaeII, 100-130 µm, absent in the type specimen; sigmancistra I, 11-14 µm; sigmancistra II, 22-30 µm, sometimes absent.
Biology: Type specimens collected from 91 m. Fixed on solidsubstrata up to 4960 m deep. The presence of lateral filamentscovered by microscleres regularly arranged with the teeth up-wards, and the absence of canal system and apertures in the col-lected specimens suggest a carnivorous mode of feeding, similarto that developed in Cladorhizidae. This is supported by thepresence of debris of small crustaceans in the best preservedspecimens from Barbados (cruise Manon © Ifremer).
Distribution: Cape Verde Islands (91 m); Barbados (4960 m);Mid-Atlantic Ridge: Lucky Strike; Rainbow.
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Caulophacus cyanae BOURY-ESNAULT & DE VOS, 1988
Size: Up to 50 cm
Morphology: Mushroom-like body, white with a solid stalk anda convex discoid upper part; ectosomal skeleton of small pen-tactine, pinnular dermalia and of strong pentactine hypoder-malia. Choanosomal megascleres of hexactines, pentactinesand rhabdodiactines. Microscleres are discohexasters of threesizes. Large choanocyte chambers composed of anucleatechoanocytes.
Distribution: East Pacific Rise: 13°N. On vertical walls, inac-tive sulphide edifices and basaltic pillars in the graben.Caulophacus-like Porifera occur along the northern and south-ern East Pacific Rise and Pacific-Antarctic Ridge: 38°S, but infewer abundance.
Reference:
BOURY-ESNAULT N. & L. DE VOS (1988) Oceanol. Acta 8: 51-60.
N. BOURY-ESNAULT Denisia 18 (2006): 47
1: Some specimens in situ; cruise Hope @ Ifremer.
3: A preserved specimen; by P. Briand @ Ifremer.2: Spicules (SEM); after BOURY-ESNAULT & DE VOOS (1988).
Porifera, Hexactinella, Lyssacinosida, Caulophacidae
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Cnidaria, Hydrozoa, Leptolida (= Hydroida)
The leptolid fauna of hydrothermal vents of oceanic ridgesites is still poorly investigated and understood. Fragmentarycolonies of leptolids from oceanic ridges in the Pacific have sofar made it possible to identify Zygophylax cervicornis (NUTTING,1905), Zygophylax sp. and Halecium tenellum HINCKS, 1861.While from vents on the Mid-Atlantic Ridge only a single an-thoathecate leptolid, Candelabrum serpentarii SEGONZAC &VERVOORT, 1995 can be considered to be a true companion ofthe vent community. A second species, Candelabum phrygium(FABRICIUS, 1780) is a North Atlantic deep-sea inhabitant oc-casionally found associated with the hydroythermal vent fauna.
Another anthoathecate, Bouillonia sp. was found on theMid-Atlantic Ridge in the Gulf of Guinea area; it probably rep-resents the deep-water leptolid community without directaffinities to hydrothermal vents (SVOBODA, STEPANJANTS &LJUBENKOV, in press). Photographs taken during surveys of Mid-Atlantic vent areas show a third species of anthoathecate, oc-casionally in great numbers; it may represent a species of Tubu-laria or Ectopleura. There is presently no material available tosubstantiate that guess. A probably undescribed species of Hy-dractinia has also been observed.
Collecting activities at or near Mid-Atlantic vents areashave currently provided material for proper identification of 3anthoathecates and 23 leptothecates; 4 more leptothecatescould only be identified to the genus (CALDER & VERVOORT
1998). One more species (Hydrallmania falcata) was dead whencollected and accidentally introduced; all remaining leptothe-cates belong to the deep-water fauna and are only party re-stricted to the Atlantic.
Because of the technique used to collect animals at ridgesites, only rarely do complete animals or colonies become avail-able for scientific study. Anthoathecates in particular are hardto collect and they must be narcotized prior preservation. Sofar, only a single complete animal of Candelabrum serpentarii hasbeen obtained, in addition to many incomplete individuals,which have been studied. Anthoathecates are best preserved in6% formalin (after prior relaxation by adding MgSO4 solutionto the seawater), but Leptothecates get very brittle in formalinand are best preserved in 70% ethanol. These preservationtechniques only apply to material for routine taxonomic inves-tigations; more sophisticated methods are necessary for modernDNA techniques.
References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 1-65. SEGONZAC M. & W. VERVOORT (1995) Bull. Mus. Natl. Hist. Nat., Paris (4)17(1-2): 31-64. SVOBODA A., STEPANJANTS S. & J. LJUBENKOV (in press) Zool. Meded., Leiden.
W. VERVOORT Denisia 18 (2006): 48
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Candelabrum phrygium FABRICIUS, 1780
Size: 7-9 cm.
Color: White to brownish.
Morphology: Body in extended condition slender, composed offoot, blastostyle bearing region and trunk. Foot lobed and flat-tened part of body, attaching animal to rocks or bivalve mol-luscs. Blastostyle bearing region occupying one third to onequarter part of body, with 10-15 large, tubular blastostyles, bear-ing male or female gonophores in various stages of developmentand dispersed capitate tentacles; apex of each blastostyle withcircle of 4-5 such tentacles. Remainder of body forming elon-gated, tubular trunk, completely covered by capitate tentacles.Mouth at distal end of trunk. Nematocysts: Haplonemes 19.7-
20.5 x 8.2-9.9 µm; small desmonemes 8.2-9.0 x 6.4 x 6.6 µm;large desmonemes 12.5-13.0 x 9.0-9.8 µm; stenoteles 10.6-11.5x 8.2-9.8 µm.
Biology: Attached to rocks or other solid objects. Lucky Strikespecimens were attached to flange formations and occasionallyto organic support (shells of living bivalves). Carnivorous ani-mal, feeding on shrimps and other small crustaceans. Dioecious,development of larva unknown, but probably young polyp de-velops inside female gonophore.
Distribution: Circumarctic, occurring both in Atlantic and Pa-cific Oceans. Mid-Atlantic Ridge: Lucky Strike and Rainbow.
Reference:
SEGONZAC M. & W. VERVOORT (1995) Bull. Mus. Natl Hist. Nat., Paris, 4e sér. 17(1-2): 31-64.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 49
1: Several animals in situon substrate; cruise Diva 2© Ifremer.
2: Preserved specimen on small rock in Lucky Strike area, Mid-Atlantic Ridge; by P. Briand © Ifremer.
Cnidaria, Hydrozoa, Leptolida, Anthoathecata, Candelabridae
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Candelabrum serpentarii SEGONZAC & VERVOORT, 1995
Size: 10 cm.
Color: white, whitish or pale orange.
Morphology: Robust, gelatinous body composed of foot, blas-tostyle bearing region and trunk, attached to pillow lava or sul-fide rocks by means of (flattened) foot covered by thick, darkbrown perisarc. Blastostyles on proximal part of body large,bearing a number of male and female gonophores in variousstages of development; elongated trunk completely coveredwith capitate tentacles, in extended condition several mmlong. Mouth at distal end of trunk. Nematocysts: Haplonemes16.5-18 x 7.8-8.2 µm; desmonemes 13-14.5 x 9.8-10.5 µm;stenoteles 9.8-11.5 x 9.5-10.5 µm.
Biology: Attached to rocks (pillow lava or sulfide rocks) invicinity of active vents. Probably exclusively carnivorous, feed-ing on pericarideans and shrimps. Reproduction: Monoeciouswith hermaphroditic gonophores; development of larva un-known, but probably young polyp (actinula) developing in fe-male gonophore.
Distribution: Mid-Atlantic Ridge: Snake Pit.
Reference:
SEGONZAC M. & W. VERVOORT (1995) Bull. Mus. Natl Hist. Nat., Paris, 4e sér. 17(1-2): 31-64.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 50
1: In situ, Snake Pit area, Mid-Atlantic Ridge; cruise Hydrosnake © Ifremer.
2: In situ, Snake Pit area, Mid-Atlantic Ridge;cruise Hydrosnake © Ifremer.
Cnidaria, Hydrozoa, Hydroida, Anthomedusae, Candelabridae
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Eudendrium planum BONNEVIE, 1898
Cnidaria, Hydrozoa, Leptolida, Anthoathecata, Eudendriidae
1 right side: Part of hydrocaulus withpedicel and (damaged) hydranth; scalebar 0.5 mm; left side: Male gonophoreattached to base of hydranth; scale bar0.25 mm; from CALDER & VERVOORT (1998).
Reference:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 51
Size: Colonies up to 90 mm high, branching reduced.
Morphology: Colonies slender, stiff, sparsely branched, hy-drorhiza creeping, growth monopodial with terminal hydranth.Stem stiff, upright, basally polysiphonic but largely monosi-phonic; branching irregular, more or less in one plane, pedicelsalternate, long. Perisarc thick, yellowish, thinning out distally,terminating at bases of hydranths. Annulations present at baseof pedicels and branches, occasionally elsewhere but perisarcmostly smooth. Hydranths about 500 µm long, urn-shaped,with large, flared hypostome. Tentacles filiform, numberingabout 20. Gonophores fixed sporosacs. Male gonophores with
one or two chambers, attached to non-reduced hydranths. Fe-male gonophores undescribed. Nematocysts: macrobasic eury-teles 12.9-14.1 x 5.5-6.4 µm; heterotrichous microbasic euryte-les 7.5-8.4 x 3.9-4.6 µm.
Biology: Attached to fixed object like stones or rocks. AtLucky Strike site, attached to flanges where mussels are fixed.Probably a dioecious species feeding carnivorously.
Distribution: Deep water of the north-eastern Atlantic; Mid-Atlantic Ridge: Logatchev, Lucky Strike, Rainbow.
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Ectopleura larynx ELLIS & SOLANDER, 1786
Size: Height of individual polyps up to 50 mm.
Morphology: Colonies formed by branched hydrocauli or bydevelopment of actinulae on individual hydrocauli arising fromcreeping, twisted stolon with wrinkled, straw-coloured perisarcattached to fixed objects. Diameter of hydrocaulus about 400Ìm, perisarc firm basally, gradually thinning about distally andterminating just below bulbous base of terminal, vasiform hy-dranth. Height of hydranth about 1.7 mm, width 1.2 m; onewhorl of short oral tentacles, up to 750 µm long and one whorlof much longer aboral tentacles, up to 2.800 µm long.Gonophores fixed sporosacs, developing just above insertion ofaboral tentacles on short, slender dichotomously branches blas-tostyles. Both male and female gonophores are eumedusoidwithout remnants of radial canals but with three rudimentarytentacles. Nematocysts: Desmonemes 3.8-4.6 x 3.0 x 3.6 µm;
small stenoteles 6.7-7.3 x 5.4-5.7 mm; large stenoteles 10.6-21.1 x 9.5-10.1 µm; mastigophores 8.4-9.6 x 3.5-4.0 µm; O-isorhiras 9.2-9.4 x 8.4-8.7 µm.
Biology: Attached to fixed objects but frequently developinginto a free-living, detached colony as the result of fragmenta-tion of the original colony. Monoecious or dioecious colonies.The eggs develop inside the female gonophore and become freein the actinula stage. As the possibilities for dispersion of theactinulae are limited many settle on or in the vicinity of theelder.
Distribution: Mid-Atlantic Ridge: Menez Gwen. Sub-arctic,temperate and subtropical waters of the northern Atlantic,from the littoral zone down to considerable depth in the ba-thyal zone.
Reference:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 52
1: Hydranth with female gonophores, seen from above(oral view); from CALDER & VERVOORT (1998).
2: Tubularia or Ectopleura sp. in situ, from Menez Gwen ventfield at the boundary of active area; cruise Ataos © Ifremer.
Cnidaria, Hydrozoa, Leptolida, Anthoathecata, Tubulariidae
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Polyplumaria flabellata G.O. SARS, 1874
Size: Height of colony up to 350 mm, spread almost equal.
Morphology: Stolonal mass of fine tubules supporting a rigid,pinnately branched colony of very regular structure. Stem poly-siphonic and repeatedly branched, branches in opposite pairs;branches originating from secondary tubes; primary axis visiblein polysiphonic parts in front of bundle of secondary tubules,divided by oblique nodes into internodes of varied length, eachinternode with one to five apophyses, alternately directedobliquely upwards and to left and right. Primary and secondarytubules with nematothecae, each apophysis with a ‘mamelon’and three nematothecae; in addition two to four nematothecaebetween two successive apophyses. Hydrocladia about 3 mmlong, supported by apophyses, slightly curved, first internodeathecate, with one nematothecae; all following internodes the-cate, with one proximal, a pair of flanking, one distal and sev-eral additional and variously distributed nematothecae on dis-tal part of internode. All nematothecae bithalamic with deeplyscooped adcauline wall. Hydrotheca cylindrical with almostparallel walls, adcauline wall adnate for half its length, length
axis of hydrotheca at 45∞ to length axis of internode; hy-drothecal rim circular and smooth, perpendicular to hydrothe-cal length axis; lateral (flanking) nematothecae on shortapophyses near hydrothecal axil. Hydrocladia occasionallyforked, secondary branch springing from base of first hydrothe-ca. Depth of hydrotheca 140-190 µm, diameter at rim 135-160µm. Female and male gonothecae of same shape, curved ovoid,attached to apophyses by means of short pedicel; basal part ofgonothecae with 6-9 nematothecae. Aperture circular, laterallydisposed, turned towards hydrocladium, closed by circular lid.Length of gonothecae 610-750 µm; diameter 340-400 µm. Ne-matocysts: Not studied in detail.
Biology: Usually on soft bottoms (off vent areas). Serves as hostfor many epizoic leptolid species.
Distribution: Mid-Atlantic Ridge: near Menez Gwen, 997-696m. Moderately deep to deep waters of tropical, temperate, andnorthern parts of the eastern Atlantic.
1 bottom: Colony; from COR-NELIUS (1995); right: Central axisof branch with apophysis andbase of hydrocladium; scalebar 0.25 mm; left: Hydrocladialinternode with hydrotheca andnematothecae; scale bar 0.25mm; top: Gonothecae; scalebar 0.10 mm; from CALDER &VERVOORT (1998).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Halopterididae
References:
ANSÍN AGÍS J., RAMIL F. & W. VERVOORT (2001) Zool. Verh., Leiden 333: 3-268.CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.CORNELIUS P.F.S. (1995) North-West European Thecate Hydroids and their Medusae, 2. Synopses of the British Fauna (New Series) 50: vii, 1-386.RAMIL F. & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 53
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Halisiphonia arctica KRAMP, 1932
Size: Length of hydrotheca about 1 mm; length of pedicel up to3 mm.
Morphology: Stolonal colony with hydrothecae and gonothe-cae developing from branching and anastomosing stolons. Hy-drothecae pedicellate, pedicel up to 3 mm long, smooth, withsome proximal annulations, merging into almost tubular toslightly conical hydrotheca, separated from pedicel by quitethin diaphragm. Hydrothecal rim slightly everted, frequentlyrenovated. Hydranth unknown. Gonothecae big, flattened,about 2.5 mm long, spade-shaped, proximally narrowing into a
short, ringed pedicel, distally open, truncated, rim notched.Gonophores developing free medusae that are so far unde-scribed. Nematocysts: Unknown.
Biology: Attached to firm substrate. Although it seems clearthat the gonophore develops free medusae these are unde-scribed so far. The presense of free medusae in Halisiphoniabrings it close to such species of Hebella that also have a freemedusa in their life-cycle.
Distribution: Mid-Atlantic Ridge: Logatchev.
References:
KRAMP P.L. (1932) Medd. om Grønland 79: 1-86.SCHUCHERT P. (2001) Medd. Grønland Biosci. 53: 1-184.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 54
1: Colony composed of hydrothecae of varied lengths and a gonotheca on stem ofhydroid. Length of hydrotheca, including pedicel, 0.8- 5.0 mm; from KRAMP (1932).
2: Gonotheca with developing medusa;scale bar 0.5 mm; from SCHUCHERT (2001).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Hebellidae
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Grammaria abietina M. SARS, 1850
Size: Colonies from the Mid-Atlantic Ridge are a few cm high;in boreal and subarctic waters colonies may reach a size of 100mm.
Morphology: Irregularly branched, erect colonies with polysi-phonic stems and branches, only distal parts of branches mono-siphonic. In these parts hydrothecae placed in four, six or(rarely) eight longitudinal series with hydrothecae of adjacentrows alternate; in quadriseriate colonies hydrothecae in oppo-site pairs and decussate. Polysiphony caused by copious devel-opment of secondary tubules obscuring structure of polysiphon-ic stems and branches. Hydrothecae tubular, half of adcaulinewall adnate to stem or branch, curved outwards, rim circular,not flared, frequently renovated. No nematophores or nema-tothecae. Total length of hydrotheca 1.1-1.3 mm, diameter at
rim 270-306 µm. Gonothecae aggregated into ovoid coppiniaewith many strongly curved tubules projecting from betweengonothecae. Coppiniae hermaphroditic, female gonothecaeamphora-shaped, narrowing into short neck distally; malegonothecae globular and pedicellate. Eggs developing in anacrocyst projecting from female gonothecae. Nematocysts: Notdescribed in detail, “large and small nematocysts“ (BROCH
1918) being present.
Biology: Colonies developing on solid substrata. Eggs broodedin an acrycyst.
Distribution: Mid-Atlantic Ridge: Menez Gwen. Generallyrecognized as a circumpolar species, extending its distributionsouthwards in the deep water of the northern Atlantic.
References:
BROCH H. (1918) Danish Ingolf-Expedition 5(7): 1-205.CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.CORNELIUS P.F.S. (1995) North-West European Thecate Hydroids and their Medusae, 1. Synopses of the British Fauna (New Series) 50: vii, 1-347.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 55
1 left: Colony; high about 6 cm; by CORNELIUS (1995); middle: Part of stem with side-branch; scale bar 0.5 mm; from CORNELIUS
(1995); right: Arrangement of hydrothecae around stem; scale bar 0.25 mm; from CALDER & VERVOORT (1998).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Lafoeidae
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Lafoea dumosa FLEMING, 1820
Size: Erect colony up to 100 mm high; length of hydrothecavaried between 150 and 800 µm, diameter at rim 100-200 µm;greatest dimensions in specimens from cold water.
Morphology: Stolonal or erect colonies, often a combination ofboth; erect colonies very irregularly branched, often with anas-tomoses, mono- or polysiphonic, the latter being composed ofseveral adnate parallel tubes all bearing hydrothecae on allsides, occasionally in more or less opposite and decussatearrangement. Hydrotheca tubular to elongated conical, fre-quently asymmetrical, rim circular, usually renovated. Hy-drotheca narrowing basally into frequently almost impercepti-ble pedicel; other colonies may have a well developed, twistedhydrothecal pedicel. Hydranth extensile, with about 20 amph-icoronate tentacles around conical proboscis, attached deep in-side hydrothecae by means of ring of desmocytes; coenosarc yel-
lowish. Gonothecae aggregated into ovoid, several mm longcoppinia on stem or branches, hermaphroditic, with manystrongly curved accessory tubes protruding from betweengonothecae. Gonothecae amphora-shaped, strongly adnate.Presence of acrocyst unknown. Coppiniae rare, reproductionapparently frequently vegetative. Nematocysts: Isorhiza’s of twosize classes (about 21 and 16 µm length, not in same colony!)and a smaller unidentified capsule, 6 µm long (SCHUCHERT
2001).
Biology: On fixed objects, also epizoic. Although a well-dis-tributed species few details of its biology are available.
Distribution: Mid-Atlantic Ridge: Rainbow. Shallow to deepwaters of all oceans [concept of species as defined by CORNELIUS
(1975)].
References:
CORNELIUS P.F.S. (1975) Bull. Br. Mus. Nat. Hist., Zool. 28: 373-426.CORNELIUS P.F.S. (1995) North-West European Thecate Hydroids and their Medusae, 1. Synopses of the British Fauna (New Series) 50: vii, 1-347.RAMIL F. & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.SCHUCHERT P. (2001) Medd om Grønland. Bioscience 53: 1-184.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 56
Left: Erect colony, 6 cm height; middle: Part of erect colony with almost sessile hydrothecae; scale bar 0.5 mm; right: Coppinia;scale bar 0.5 mm; from CORNELIUS (1995).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Lafoeidae
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Zygophylax echinata CALDER & VERVOORT, 1998
Size: Height of erect colony 25 mm.
Morphology: Colony erect, stem strongly polysiphonic, mono-siphonic only in upper parts and there slightly geniculate. Hy-drocladia short, alternately arranged on either side of axis, bis-eriate, with axillar hydrotheca bearing two opposite nema-tothecae at base of pedicel. Hydrocladia may become polysi-phonic by apposition of secondary tubes and presence of nema-tothecae in axillar hydrothecae may become obscured. Hy-drothecae on axis and hydrocladia alternately arranged, placedon distinct, swollen apophysis with a pair of opposite nema-tothecae. Besides axillar hydrotheca there are two hydrothecaebetween two successive hydrocladia. Nematothecae small,elongated tumbler-shaped, narrowing proximally into shortpedicel, length 42-68 µm, diameter at rim 24-31 µm, renova-tions frequent. Hydrothecae small, slender vase-shaped, slight-
ly asymmetrical by bulging adcauline wall. Proximal portion ofhydrotheca narrowing into short, plump pedicel; diaphragmdistinct, oblique. Length of hydrotheca (without renovations)250-400 µm, rim moderately everted, circular, frequently reno-vated, diameter 94-117 µm. Gonothecae aggregated into char-acteristic, oval coppinia, 3.5 mm long, 2 mm high, surroundingaxis basally. Gonothecae flask-shaped, strongly adnate, taperingdistally into a sharply pointed structure with a lateral aperture.Nematophorous ramules protrude from between gonothecae,curving over coppinia, bearing both nematothecae and somehydrothecae. Nematocysts: Undescribed.
Biology: Attached to solid substrate like rocks or stones.
Distribution: Mid-Atlantic Ridge: near Menez Gwen.
1 left: Distal part of stem with hydrothecae, nematothecae and insertion of two hydrocladia, scale bar 0.25 mm; top row left:Axillary hydrotheca with nematotheca, scale bar 0.1 mm; middle: Hydrotheca and nematotheca, scale bar 0.1 mm; right: End ofnematophorous ramule with hydrotheca and two nematothecae, scale bar 0.1 mm; bottom row middle: Three gonothecae andnematophorous ramule from coppinia, scale bar 0.25 mm; right: Apophysis of hydrocladium with pedicel of hydrotheca andtwo nematothecae, scale bar 0.1 mm; after CALDER & VERVOORT (1998).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Lafoeidae
Reference:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 57
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Zygophylax leloupi RAMIL & VERVOORT, 1992
Size: Height of colony up to 140 mm.
Morphology: Structure of colony in principal as in Z. echinataCALDER & VERVOORT, 1998, but colony generally bigger, withstronger, repeatedly bifurcated stem, with longer hydrocladiaand distinctly pedicellate, big hydrothecae that are distinctlyfrontally directed. Length of hydrotheca (from diaphragm on-wards, without renovations) 500-650 µm, diameter at rim 280-340 µm; renovations frequent. Pedicel long, wrinkled or withsepta, 270-250 µm; diaphragm distinct but thin, occasionallyduplicated. Aperture of hydrotheca slightly tilted, rim everted,diameter 210-325 µm. Nematothecae scarce, frequently dis-
lodged, on apophyses of hydrothecae, cylindrical, proximallyrounded and shortly pedicellate. Gonothecae aggregated intoloose coppinia without nematophorous ramules, ovoid, distallywith two or three short tubular processes each with terminalaperture. Coppinia surrounding proximal part of stem. Nema-tocysts: Not described.
Biology: Attached to solid substrata, among mussel bed.
Distribution: Mid-Atlantic Ridge: Rainbow; seamount at Seg-ment 38°N, 919 m.
References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.RAMIL F. & VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 58
1 left: Monosiphonic part of stemwith three hydro-thecae; middle:Hydrotheca and nematotheca; right:Gonotheca isolated from coppinia.Length of hydrotheca about 0.8mm, length of gonotheca 1.0-1.4mm; from RAMIL & VERVOORT (1992).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Lafoeidae
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Hydrallmania falcata LINNAEUS, 1758
Size: Full grown colonies may reach a length of 650 mm; usual-ly much smaller.
Morphology: Colony lax, with characteristic, spirally twisted,monosiphonic stem, supporting branches of 20-40 mm lengthwith pinnately arranged hydrocladia. Each hydrocladium divid-ed into internodes, supporting closely packed groups of 3-10 hy-drothecae, alternately inclined left or right to a moderate de-gree. Hydrotheca more or less tubular, largely free, slightly nar-rowing distally, aperture circular, operculum two-flapped. Totallength of hydrotheca 300-400 µm; greatest diameter 130-190µm; diameter at rim 80-140 µm. Male and female gonothecaesimilar, elongated pear-shaped, narrowing proximally into shortpedicel attaching gonothecae to internode of hydrocladium;
aperture on short distal tube, rather wide. Length of gonothe-cae 1.2-1.7 mm; greatest diameter 550-700 µm. Nematocysts:Not described in detail.
Biology: Frequent on soft bottoms in shallow to moderatelydeep waters, particularly in temperate regions. One of the fewspecies of Leptolida was (and probably still is) commercially ex-ploited. Gonothecae produced in early spring. Colonies servethe attachment for mussel spat.
Distribution: Mid-Atlantic Ridge: Lucky Strike. Shallow tomoderately deep waters of the temperate and northern At-lantic; not a deep water species.
References:
CORNELIUS P.F.S. (1995) North-West European Thecate Hydroids and their Medusae, 2. Synopses of the British Fauna (New Series) 50: VII, 1-386.RAMIL F & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 59
1: Colony, height about 9 cm;from CORNELIUS (1995).
2 from left to right: Normally developed hydrocladium; hydrocladiumfrom young colony (similar arrangement of hydrothecae occasionallyalso in older colonies); hydrotheca; gonothecae; scale bars 0.5 mm;from CORNELIUS (1995).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Sertulariidae
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Sertularella tenella ALDER, 1856
Size: Erect colonies about 20 mm high or smaller.
Morphology: Monosiphonic, occasionally branched stem aris-ing from firm, tortuous stolon attached to solid substrate orhost; stem geniculate, composed of slender internodes with afew basal rings; nodes indistinct. Hydrotheca with almost feeabcauline wall, almost cylindrical to slightly swollen in proxi-mal half, with six annulations, best visible on abcauline side;depth 450-840 µm, greatest diameter 220-370 µm. Rim withfour low cusps; operculum four-flapped, flaps attached in em-bayments between marginal cusps. Female and male gonothe-
cae undistinguishable, ovoid, 900-2.000 µm long, greatest di-ameter 550-1.000 µm, with 8-10 transverse annulations; apexwith 3-4 low cusps surrounding a small, circular aperture. Ne-matocysts: Not studies in detail.
Biology: Frequently on foliate Bryozoa and epizoic on other hy-droids.
Distribution: Mid-Atlantic Ridge: Lucky Strike, Sintra. Cir-cumglobal in deeper parts of the littoral zone to greater depths(1000-2000 m).
References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.RAMIL F. & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 60
1: From left to right: three stems arising from stolonal tubes; part of stem; another part of stem from different colony to illus-trate variability in development of nodes and shape of hydrotheca; hydrotheca; gonotheca; scale bars 0.5 mm; by Cornelius.
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Sertulariidae
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Symplectoscyphus bathyalis VERVOORT, 1972
Size: Height of colony up to 80 mm.
Morphology: Erect colony, rising from creeping stolon. Proxi-mal parts of stem and branches polysiphonic; branching irregu-lar, basal part of stem ahydrothecate. Monosiphonic parts ofcolony composed of long, slender, geniculate internodes sepa-rated by nodes sloping in alternate directions. Hydrothecae950-1.200 µm deep, greatest diameter about 500 µm, placed atend of internode, one-fifth of abcauline wall adnate, curvedoutwards, slender, abcauline wall convex, abcauline wallbulging in proximal third. Hydrothecal rim not everted, occa-sionally renovated, with three prominent cusps, one abcauline,two lateral. Three triangular opercular plates attached in em-bayments between marginal cusps, no renovations of opercu-
lum observed. Gonotheca pyriform, 1.7 mm high; greatest di-ameter at two-thirds of distance from top, with about seventransverse ribs, petering our proximally. Orifice at end of a shorttube in middle of apical ‘field’ formed by distalmost rib. Pedicelquite short, attaching gonothecae to internode at hydrothecalbase. Nematocysts: Not studies in detail.
Biology: On solid substrates in deep ocean water.
Distribution: Mid-Atlantic Ridge: Lucky Strike, Sintra site.Deep water of Atlantic (Bay of Biscay) and Pacific (off Chileand SE of New Caledonia).
References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.VERVOORT W. (1972) Zool. Verh., Leiden 120: 3-247.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 61
1 from left to right: Monosiphonic part of stem with three hydrothecae; part of monosiphonicstem with hydrotheca and gonotheca (drawn from slide; gonotheca slightly compressed); scalebars 0.5 mm; from CALDER & VERVOORT (1998).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Sertulariidae
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Stegolaria geniculata ALLMAN, 1888
Size: Height of colonies up to 10 cm.
Morphology: Ramified, irregularly shaped, erect colonies withthick, geniculate, polysiphonic stems, resulting from geniculateaxial tube with hydrothecae becoming covered by many acces-sory tubules; ultimate ramifications monosiphonic. Stem basal-ly with stolonal tubes attaching colony to firm substrate. Hy-drothecae alternately arranged, in same plane as hydrocaulusand branches, adnate to axial tube and immersed to varying de-gree, curving outwards, gradually widening from base onwards;walls smooth, abcauline wall concave, free adcauline wall con-vex to nearly straight. Operculum a roof-shaped enclosure withtwo longitudinally pleated valves, suspended in U-shaped em-bayments of hydrothecal margin. Perisarc firm on stem andbranches, yellowish-brown, thinning out considerably alongwalls of hydrothecae and becoming almost hyaline distally; dis-tal part of many hydrothecae collapsed. No diaphragm; hy-dranths attached to internal wall of hydrotheca by means of cir-cular collar of tissue. Gonothecae elongated sack-shaped,
widest in mid-region, attached to axial tube amongst hydrothe-cae. Distal part narrowing fairly sharply, curving outwards as ashort tube with circular aperture. Condition of gonophore un-known; development of a free medusa phase is not unlikely. Ne-matocysts: Not described.
Biology: Attached to solid objects like stones or rocks. Re-silient colonies frequently without living tissue in deep waterhauls; abraded specimens difficult to recognize. Though this isby no means a rare species in deep oceanic waters the structureof polyp and gonophore still remain unknown because of thescarcity of well preserved material. Reproduction and mode offeeding unknown.
Distribution: Mid-Atlantic Ridge: Lucky Strike and Rainbow.Deep water of the Atlantic Ocean; probably also in deep waterof Indian and Pacific Oceans or replaced by closely alliedspecies [Stegolaria operculata (NUTTING, 1905)].
References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.RAMIL F. & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.
W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 62
1 from left to right: Monosiphonic top part of colony;terminal hydrotheca; hydrotheca on monosiphonic partof stem; length of hydrotheca 1.3-1.6 mm; from RAMIL &VERVOORT (1992).
2 from left to right: Renovated hydrotheca onpolysiphonic part of colony; scale bar 0.25 mm;branch with empty gonotheca and two damagedhydrothecae; scale bar 0.5 mm; from CALDER & VERVOORT (1998).
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Tiarannidae
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Thermopalia taraxaca PUGH, 1983 “Galapagos dandelion“
Size: Height of main body up to 40 mm.
Morphology: Ball-like structure, with an apical, smooth-walledfloat (pneumatophore), pinkish-orange in colour. Immediatelybelow this is a corona of flimsy, swimming bells; but these areonly apparent if the animal is actively swimming. The mainbody consists of a mass of orange-coloured feeding polyps (gas-trozooids), together with other structures. The specific charac-teristics are rather esoteric, and include a smooth-walled au-rophore; cormidia attached directly to the main body of thecorm; and the absence of gonopalpons.
Biology: The animals are very obvious structures as they float afew centimetres above the substrate, and are attached, liketethered hot-air balloons, by means of numerous lines (tenta-cles).
Distribution: Galapagos Spreading Center and East PacificRise. T. taraxaca or relatives have been observed on the Juan deFuca Ridge but never collected.
Reference:
PUGH P.R. (1983) Philos. Trans. R. Soc. Lond. 301(1105): 165-300.
P.R. PUGH Denisia 18 (2006): 63
1: In situ from East Pacific Rise: 13°N © Ifremer. 2: In situ from East Pacific Rise: 13°N © Ifremer.
3: In situ from Galapagos Spreading Center; bycourtesy of R.R. Hessler.
4: In situ from Galapagos Spreading Center; bycourtesy of R.R. Hessler.
Cnidaria, Hydrozoa, Siphonophora, Rhodaliidae
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Lucernaria janetae COLLINS & DALY, 2005 “stalked jellyfish”
1: Aggregation in situ (East Pacific Rise: 9°N, 2540 m); by R. Zierenberg.
2: Cross-section through monocameral peduncle,showing four septal cords (East Pacific Rise: 9°N,2540 m; FMNH 10328).
3: Close-up of perradial notch. Note secondary ten-tacles and absence of perradial anchor (East PacificRise: 9°N, 2540 m; FMNH 10327).
Cnidaria, Scyphozoa, Stauromedusae, Eleuthrocarpidae
Size: Calyx diameter to 100 mm, height to 60 mm; peduncleheight to 60 mm.
Color: Creamy greenish-white to orangish-white.
Morphology: Extremely large stauromedusan with goblet-shaped calyx and eight equa-distant clusters of capitate second-ary tentacles. Adults have no perradial anchors (primary tenta-cles); juveniles may have small, ovoid perradial anchors.
Biology: On basalt at vent periphery or areas of diffuse flow.Many individuals of varying sizes co-occur; may dominate localfauna. Co-occurs with tubeworm Tevnia, sea anemones(Cyananthea sp.), and vent-associated decapods (LUTZ et al.1998; HALANYCH et al. 1999). Aggregations comprised of spec-imens of varying sizes and age stages. Smallest juveniles may beattached to basal end of adults.
Distribution: East Pacific Rise: 7°S to 21°N. A similar formwas observed at Pacific-Antarctic Rise, 38°S, 2330 m (cruisePAR5, April 2005).
References:
COLLINS A.G. & M. DALY (2005) Biol. Bull. 208: 221-330.HALANYCH K.M., TIEGER M., O’MULLAN G.D., LUTZ R. & R.C. VRIJENHOEK (1999) InterRidge News 8: 23-27.LUTZ R.A., DESBRUYÈRES D., SHANK T.M. & R.C. VRIJENHOEK (1998) Deep-Sea Res. II 45: 329-334.
M. DALY Denisia 18 (2006): 64
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Cnidaria, Anthozoa, Actiniaria
According to the current knowledge, the sea anemones,with eight presently known species, constitute about 1.6% ofthe total number of identified vent species. Several other mor-photypes have been photographed in situ or are currently understudy. Thus the number of sea anemone species will probablyincrease slowly but continually when taxonomic studies con-tinue and new vent field are discovered.
Although the total number of sea anemone species report-ed for these environments is apparently not very significant –compared with other groups like gastropods or polychaetes, forexample – the number of species living in a specific vent fieldis not higher that two or three these few species are sometimesso abundant that they form characteristic fields completelycovered by individuals, sometimes equidistantly separated.
Most of the species reported belong to the family Actinos-tolidae (six species), while only one species each of Hormathi-idae and Boloceroididae are known to date. One of the acti-nostolid species (Actinostola sp.) is only known at the genericlevel at vents. However, this genus is well known from non-vent environments. The other five actinostolid species wereoriginally described from hydrothermal vents and were alsoused as types of their monotypic genera. The specific status ofthe single hormathiid species is currently under revision.
The term sea anemones it is widely used to refer to softhexacorallians of the orders Actiniaria (including Ptycodac-tiaria), Zoanthidea and Corallimorpharia. Members of the firsttwo orders have been collected from these chemoautotrophicenvironments although, for the moment, only identifications ofactiniarian specimens are available.
The current knowledge of sea anemone species associatedwith vents is far from completed, most of the characters used toidentify the different genera being anatomic (histology) or mi-croscopic (types and sizes of cnidocysts). In addition, severalspecies belonging to different genera can have a similar appear-ance in visual surveys. All these aspects often made the specif-ic identification of sea anemone from photographic or videorecords a difficult task. At present, molecular studies on ventsea anemone lack. Such studies as well as studies on reproduc-tion, feeding, and other biological aspects should be carried outwith the appropriate number of specimens per species in thenext decades.
Sea anemones are usually delicate animals. Also those withthick body walls and a cuticle are internally high delicate. As
the study of the internal anatomy and cnidocysts is essential fora correct identification, sea anemones should be carefully treat-ed. If specimens are obtained in good condition infrequent atbest they can be relaxed adding menthol’s crystals on the sur-face of the water. Fixation is best in a big volume of 10%buffered formalin for at least a week, then they can be trans-ferred to 70% ethanol or maintained in 10% buffered formalin10%. It is highly recommended to inject a solution of concen-trated 20-30% formalin into the gastrovascular cavity. This willfacilitate proper fixation of the internal structures used for his-tological studies.
For molecular studies, either parts of animals or if dissectionof some tentacles is possible in large animals, fixation in 100%alcohol and several changes of the fixation medium or freezingare recommended. It is very important that the rest of the spec-imen from which the tentacles or fragments have been removedis fixed in formalin as described above. Thus, molecular as wellas traditional taxonomic studies can be carried out using thesame voucher specimen. Material entirely frozen or fixed inethanol is seldom useful for the current taxonomic work.
P. LÓPEZ-GONZÁLEZ Denisia 18 (2006): 65
1: Boloceroides daphnae from East Pacific Rise: 17°S,Biospeedo cruise © Ifremer.
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Actinostola VERRILL, 1883
1: In situ specimen; cruise Hope 99 © Ifremer.
2: In situ specimen; cruiseHot 96 © Ifremer.
Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae
Reference:
DOUMENC D. & M. VAN PRAËT (1986) Oceanol. Acta 8: 61-68.
P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 66
Size: Diameter of the pedal disc, 90 mm; mouth, 70 mm; col-umn height, 60 mm; length of contracted tentacles, 11 mm.
Color: Column, tentacles, pharynx, and mesenteries yellowishwhite in situ.
Morphology: Column cylindrical, with conspicuous mesogloealtubercles arranged regularly in longitudinal rows; tentaclesarranged in four or five cycles, with an apical orifice, aboral sideof tentacles often thickened at the base; sphincter weak,mesogloeal, thus tentacles can not be completely retracted;longitudinal musculature of tentacles mesogloeal; mesenteriesarranged according to Actinostola-rule; oldest cycles of mesen-teries sterile.
Remarks: DOUMENC & VAN PRAËT (1986) discussed the possi-ble specific identity of the single specimen examined without a
definitive conclusion. Although in the figure 1 of that paper,the name A. callosa is included, it was probably a mistake dur-ing the editorial process of the manuscript. DOUMENC & VAN
PRAËT (1986: 63) also reported the presence of four cycles oftentacles, this number of cycles corresponding to 48 tentacles(6+6+12+24). However, the number of tentacles seen in the insitu photographs suggest five cycles. Additional material isneeded to study the specific pertinence and other anatomicaldetails.
Biology: As other sea anemones, probably feeds on caught prey,with a relative wide prey size-range. All other biological aspectsare unknown. Often observed within the gravel, attached atthe top of lava pillars.
Distribution: East Pacific Rise: 13°N.
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Cyananthea hydrothermala DOUMENC & VAN PRAËT, 1986
Size: Diameter of pedal disc, 10 mm, height, 35 mm; tentacles,ca. 25 mm.
Color: Whitish in situ, column and tentacles dark brownish,mesenteries pale beige.
Morphology: Column smooth, with distal margin distinctlymarked; tentacles occupying most oral disc, reaching near tothe mouth; more mesenteries distally than proximally; sphinc-ter mesogloeal, strong, on the endodermic face of themesogloeal layer; retractor musculature diffuse and weak; longi-tudinal musculature of tentacles ectodermic; radial musculatureof the oral disc ecto-mesogleal.
Remarks: This species was insufficiently described to be cor-rectly diagnosed or compared with other actinostolid generaand species. The familial pertinence was tentative after the sup-posed absence of acontia in the single fragment available tostudy by DOUMENC & VAN PRAËT (1986). Material collected inthe type locality should be re-described in order to precise thetaxonomic status of this genus and species.
Biology: As other sea anemones, probably feeds on caught prey,with a relative wide prey size-range. All other biological aspectsare unknown. The association with bacterial symbionts has notbeen demonstrated. Lives on the walls of the black smokers attemperatures of 12-20°C and on sulphide rocks.
Distribution: East Pacific Rise: 13°N.
Reference:
DOUMENC D. & M. VAN PRAËT (1986) Oceanol. Acta 8: 61-68.
P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 67
1: Two specimens taken onboard from the East Pacific Rise: 13°N, cruise Phare© Ifremer.
2: In situ specimen taken from the East PacificRise: 13°N, among siboglinids Riftia pachyptilaand limpets Lepetodrilus elevatus, cruise Phare© Ifremer.
Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae
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Maractis rimicarivora FAUTIN & BARBER, 1999
Size: Diameter of the pedal disc, up to 55 mm; mouth, up to 30mm; column height, up to 20 mm; length of contracted tenta-cles, up to 25 mm.
Color: Column, tentacles, pharynx and mesenteries yellowishwhite in situ.
Morphology: Column conical in contraction, with transversalfurrows due to contraction; sphincter mesogloeal, strong; tenta-cles about 100 in number, closely placed at the rim; longitudi-nal musculature of tentacles ectodermal; no mastigophore ne-matocysts in tentacles; mesenteries hexamerously arranged, butnot according Actinostola-rule; equal number of mesenteriesdistally and proximally, 4 cycles, only the first cycle perfect, allstronger mesenteries fertile; parietobasilar musculature only dif-ferentiate in the stronger cycles.
Remarks: SEGONZAC (1992) reported Parasicyonis ingolfi CAR-GREN, 1942 from Snake Pit, Mid-Atlantic Ridge. This specieswas described before the discovery of hydrothermal vents. De-
spite of that report, there is no formal morphological descrip-tion of this poorly known species from Mid-Atlantic Ridge ma-terial. At present, all other material subsequently studied fromthe Mid-Atlantic Ridge corresponds with the description ofMaractis rimicarivora. Although Parasicyonis ingolfi was includedin the first edition of the Handbook of deep-sea hydrothermalvent fauna (DESBRUYÈRES & SEGONZAC 1997), at present, wedecided not to include this species until new material will bestudied.
Biology: As other sea anemones, probably feeds on caught prey,with a relative wide prey size-range. This species was reportedpreying on shrimps Rimicaris exoculata. All other biological as-pects are unknown. Observed in peripheral regions of activeblack smokers, attached to the crumbly substratum of oxidizedsulfide.
Distribution: Mid-Atlantic Ridge: TAG, Snake Pit-Elan, andAshadze 1.
1: Specimens taken in situ by theRussian Polar Marine Expedition atthe site Ashadze, amongchaetopterid polychaeta tubeworms;by courtesy of A. Shagin © PMGE.
2: Specimens taken in situ by the American cruise DiversExpedition at the site Snake Pit-Elan, with zoarcid (Pachycara thermophilum), shrimps (Rimicaris exoculata), mytilidbivalves (Bathymodiolus puteoserpentis), buccinid gastropods (Phymorhynchus spp.);by courtesy of C. L. Van Dover.
Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae
References:
BEL’TENEV V., IVANOV V., SHAGIN A. et al. (2005) InterRidge News 14: 14-16FAUTIN D.G. & B.R. BARBER (1999) Proc. Biol. Soc. Wash. 112(3): 624-631.LÓPEZ-GONZÁLEZ P.J., RODRÍGUEZ E., GILI J.-M. & M. SEGONZAC (2003) Zool. Verh. Leiden 345: 215-243.SEGONZAC M. (1992) C. R. Acad. Sci. Paris, Sér. III 314: 593-600.SEGONZAC M. (1997) in DESBRUYÈRES D. & M. SEGONZAC (Eds.) Handbook of Deep-sea Hydrothermal Vent Fauna, Ifremer: 31.
P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 68
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Marianactis bythios FAUTIN & HESSLER, 1989
Size: In life, typical column length and oral disc diameter about125 mm in animal with 50 mm pedal disc diameter; length ofinner tentacles about equal to oral disc diameter. Dimensions ofcontracted specimens 25-30% of expanded ones.
Color: White in situ.
Morphology: Adherent pedal disc; smooth column; tentaclesabout 60 in number tapering, pointed, the outer ones of whichare much shorter than the inner. Mesenteries not arranged ac-cording to Actinostola-rule; first cycle perfect; stronger ones ga-metogenic. Tentacles without basal thickenings, no microbasicb-mastigophores in tentacles but with microbasic amastigo-phores; fewer than mesenteries. Marginal sphincter mesogloeal;longitudinal musculature of tentacles ectodermal; parietobasi-lar musculature distinct.
Remark: Another sea-anemone, morphologically similar to M.bythios and attributed to M. aff. bythios, dominates the periph-ery of black smokers at the Kairei vent field (Central IndianOcean).
Biology: High abundance in the immediate vicinity of low tem-perature vents. No morphological adaptations to this habitatdetected. Probably carnivorous, feeding on vent shrimps.
Distribution: Mariana Back-Arc Basin.
References:
FAUTIN D.G. & R.R. HESSLER (1989) Proc. Biol. Soc. Wash. 102: 815-825.HASHIMOTO J., OHTA S., GAMO T., CHIBA H., YAMAGUCHI T., TSUCHIDA S., OKUDAIRA T., WATABE H., YAMANAKA T. & M. KITAZAWA (2001) Zool. Sci. 18(5): 717-721.VAN DOVER C.L., POLZ M., ROBINSON J, CAVANAUGH D., KADKO D & J.P. HICKEY (1997) BRIDGE Newsletter 12: 33-34.VAN DOVER C.L. (2002) Mar. Biol. 141: 761-772.
P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 69
1: Specimens taken in situ in the Mariana Back-Arc Basin; by courtesy of Daphne Fautin.
2: Marianactis cf. bythios taken in situ at Central IndianOcean, Kairei vent field; in the center, mytilid bivalves(Bathymodiolus marisindicus); by courtesy of J. Hashimoto© JAMSTEC.
Cnidaria, Anthozoa, Actiniaria, Actinostolidae
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Size: Diameter of the pedal disc, 15 mm; column distally, 30mm; column height, 20 mm; length of contracted tentacles, 10mm. All measurements from preserved and contracted speci-mens.
Color: Preserved material of a whitish color. Living animals arewhitish to pinkish, with oral disc light brown.
Morphology: Column smooth, distal part including oral discwider than mid-column and pedal disc, not divisible into sca-pus and scapulus; sphincter distinctly marked distally as aprominent circumferential marginal ridge, mesogloeal, relative-ly weak; tentacles about 100 in number, of uniform thicknessalong entire length, with microbasic b-mastigophores; longitu-dinal tentacle and oral disc circular musculature ectodermal,that of tentacles equally well developed on all sides; mesenter-ies hexamerously arranged, but not according to Actinostola-
rule, first and second cycles of mesenteries perfect, all strongerones fertile; same number of mesenteries distally and proximal-ly; parietobasilar musculature not differentiated.
Biology: As other sea anemones, probably feeds on caught prey,with a relative wide prey size-range. All other biological aspectsare unknown. This species has been observed on living musselsor rocks, within 10 cm of a crack emitting a transparent fluid atca 40°C. Some anemones have been observed outside the areaof thermal influence, where isolated mytilid bivalves (Bathy-modiolus sp.) and siboglinid tubeworms (Arcovestia ivanoviSOUTHWARD & GALKIN, 1997) also occurred and where theseawater temperature was 2.8°C.
Distribution: Manus Back-Arc Basin: Fields D and E; probablyat Lau Back-Arc Basin.
Pacmanactis hashimotoi LÓPEZ-GONZÁLEZ, RODRÍGUEZ & SEGONZAC, 2005
References:
GALKIN S.V. (1997) Mar. Geol. 142:197-206.LÓPEZ-GONZÁLEZ P.J., RODRÍGUEZ E. & M. SEGONZAC (2005) Mar. Biol. Res. 1(5): 326-337.
P. LÓPEZ-GONZÁLEZ & E. RODRÍGUEZ Denisia 18 (2006): 70
1: Preserved specimen collected from Manus Basin(cruise BIOACCESS’98), specimen showing partiallyprotruded pharynx, oral disc about 20 mm in di-ameter; from LÓPEZ-GONZÁLEZ et al. (2005).
2: Preserved specimen collected from Manus Basin (cruise BIOACCESS’98), specimen cutlongitudinally, showing wide oral disc and mar-ginal ridge formed by the sphincter; from LÓPEZ-GONZÁLEZ et al. (2005).
Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae
3: Probably the same species on basalt covered by zoantharian, withgalatheid crabs Munidopsis spp.; in situ view from Lau Back-Arc Basin;Cruise TUIM7; by courtesy of C.R. Fisher.
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Paranthosactis denhartogi LÓPEZ-GONZÁLEZ, RODRÍGUEZ & SEGONZAC, 2003
Size: Diameter of the pedal disc, 35 mm; mouth, 16 mm; col-umn height, 15 mm; tentacles, 8 mm length. All measurementsfrom preserved and contracted specimens.
Color: Preserved material of a brownish color, the tentaclesdarker. Living animals show a brown crown of tentacles, whilethe column is dirty white.
Morphology: Column smooth, dome-shaped to cylindrical inpreserved state; tentacles about 70 in number, of uniform thick-ness along entire length; sphincter distinctly marked as aprominent marginal ring, mesogloeal; longitudinal tentaclemusculature ectodermal, equally well developed on all sides;
tentacles with mastigophore nematocysts; mesenteries hexam-erously arranged, but not according Actinostola-rule; same num-ber of mesenteries distally and proximally, four cycles, only thefist cycle perfect, all stronger mesenteries fertile, parietobasilarmusculature not differentiate as a separate lamella.
Biology: As other sea anemones, probably feeds on caught prey,with a relative wide prey size-range. All other biological aspectsare unknown. This species has been observed living among Rif-tia, on sediment where 10 cm depth a temperature of 98°C wasrecorded.
Distribution: East Pacific Guaymas Basin.
Reference:
LÓPEZ-GONZÁLEZ P.J., RODRÍGUEZ E., GILI J.-M. & M. SEGONZAC (2003) Zool. Verh. Leiden 345: 215-243.
P. LÓPEZ-GONZÁLEZ & E. RODRÍGUEZ Denisia 18 (2006): 71
1: Specimen collected from Guaymas Basin (Cruise Guaynaut, 1991); preserved specimen,about 35 mm along the larger axis at pedal disc; by courtesy of P. López-González.
2: In situ view of specimens amongtubeworms Riftia pachyptila; cruiseGuaynaut © Ifremer.
Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae
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Boloceroides daphneae DALY, in press “giant sea anemone”
References:
DALY M. (in press) Mar. Biol.DESBRUYÈRES D. & M. SEGONZAC (1997) Handbook of Deep Sea Hydrothermal Vent Fauna. Editions Ifremer, Brest: 1-279.
M. DALY Denisia 18 (2006): 72
1: Habitus, in situ, from East Pacific Rise: 17°S; cruise Biospeedo © Ifremer.
3: Spirocyst from the column ectoderm; scale bar 40 µm.
4: Cross section through the column wall, showinglongitudinal muscles of the ectoderm (Ec). The endo-derm (En) has weak circular muscles; scale bar 250 µm;by M. Daly.
2: A typical preserved specimen,with actinopharynx extrudedthrough the mouth and most ofthe tentacles missing; by M. Daly.
Cnidaria, Anthozoa, Actiniaria, Boloceroididae
Size: Diameter of column 0.1-0.5 m, diameter of tentacle crown0.5-2.0 m.
Morphology: Very large, pale pink to deep purple sea anemonewith crown of extremely long, strongly tapering tentacles. Col-umn with ectodermal longitudinal muscles and numerous large(~ 100 µm) spirocysts. Not more tentacles than mesenteries.Preserved specimens may lack tentacles, having only circularopenings on oral disc where tentacles had been attached.
Remark: Possibly misidentified as a species of Cerianthus inDESBRUYÈRES & SEGONZAC (1997): accompanying photographresembles this species but description does not.
Biology: Solitary specimens attached to boulders and cliff faces;long, tapering tentacles stream in current or contact sediment,shed when animal is disturbed. Prey unknown but cnidae ex-tremely large.
Distribution: East Pacific Rise: 9°N to 17°S; the specimens ob-served in southern areas belong probably to the same species.
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Chondrophellia cf. coronata (VERRILL, 1883)
Reference:
DOUMENC D. & M. VAN PRAËT (1986) Oceanol. Acta 8: 61-68.
P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 73
1: In situ view showing numerous specimens among stalked barnacles (Neolepas n. sp.), ophidiid fishand galatheid crabs; cruise Biospeedo © Ifremer.
Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Hormathiidae
Size: Diameter of the pedal disc, 27 mm; column distally, 30mm; column height, 30 mm; length of contracted tentacles, 4mm. All measurements from preserved and contracted speci-mens.
Color: Column brownish due to the presence of cuticle, tenta-cles orange in living specimens, to pale rose in preserved state.
Morphology: Pedal disc sharp, covered by cuticle; column di-visible into scapus and scapulus, the former with a cuticle, with12 rows of three high pointed tubercules distally, tubercles notclearly isolated from each other, with smaller tubercles betweenthe principal rows; cuticle thick, especially on the tubercles;sphincter mesogloeal, alveolar; tentacles up to 96 in number,the last circle can be incomplete; about similar number ofmesenteries distally and proximally, 6 pairs of perfect but ster-ile mesenteries; acontia present on oldest mesenteries; thirdand following cycles of mesenteries without filaments andacontia; without cinclidies.
Remarks: Despite of the report of this species at Pacific hy-drothermal vent areas, the correspondence of Atlantic and Pa-cific material attributed to this species is unclear. Chondrophel-lia coronata is known from the North Atlantic. Pacific materialassigned to this species shows similar cnidae biometry, but isdifferent in external morphology. This species was also report-ed – with doubts – from the coast of Chile. Specimens from hy-drothermal vents agree well with the drawings illustrated byDOUMENC & VAN PRAËT (1988: figure 2). A revision of thespecies described in this genus by López-González & Segonzacis in progress, and will give more information on the specificpertinence of the specimens collected at hydrothermal vents.
Biology: As other sea anemones, probably feeds on caught prey,with a relative wide prey size-range. All other biological aspectsare unknown. Very abundant mainly around the Southern EastPacific Rise sites, settled on sulphide deposits or basalts.
Distribution: East Pacific Rise: 13°N, 7°S to 32°S.
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Mollusca, Solenogastres
The aplacophoran molluscs are represented in the fauna as-sociated with the hydrothermal vent biotope by the Solenogas-tres (neomeniomorphs); members of the second class Caudo-foveata (chaetodermomorphs) have not yet been evidenced.Solenogastres show a narrowed body, glide upon their restrict-ed foot (pedal groove), and are carnivorous. They includeabout 245 known species. However, neither these nor thepresently three described species from the hot vent areas docu-mented here reflect the true biodiversity. Several new speciesunder description and additional representatives come from theEast Pacific Rise 13°N (Genesis, Parigo, Elsa), 9°N (Tica),18°S, and 21°S; and the Mid-Atlantic Ridge, Rainbow.
Recorded hot vent Solenogastres range between 1.5-6 mmlength and are – characteristically for all aplacophoran mol-luscs – covered by small aragonitic sclerites. Because these scle-rites are important specific as well as supraspecific characters,the collected animals should be preserved in 70% ethanol (orin formalin and soon transferred to ethanol) to save the scle-rites for determination. Furthermore, specimens should be pre-served as soon as possible after sampling, because internal or-gans (as important as the sclerites) rapidly undergo histolysis(as was the case in quite a number of specimens from the aboverecords).
L. SALVINI-PLAWEN Denisia 18 (2006): 75
1: Helicoradomenia sp. 2 from southern East Pacific Rise: 21°S; cruise Biospeedo © Ifremer.
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Helicoradomenia acredema SCHELTEMA, 2000
Size: Up to 3 mm.
Morphology: Solenogastres of cylindric body somewhat taper-ing at both ends, posterior end bluntly rounded and somewhatflattened dorsoventrally; appearance similar to H. juani, butshorter and fuzzier. Dominant sclerites slender and up to 190μm long with distal end rounded, in part with distal swelling;dorsofrontal sensory pit often obvious as transverse slit. Radulain sheath distinctly bipartite, paired ventral radula sack at endcurved; radula plates with 5-7 denticles, the lateral-most dis-tinctly longer and close to next one. Paired copulatory styletapparatus with two elements each.
Remarks: There is no information concerning the internal softorgans. Due to the similarities or even possible identities ofhard parts in different species (two new species are under de-scription), the conspecifity of specimens far off the type locali-ty (see distribution) needs to be confirmed according to inter-nal organisation. See also H. juani.
Distribution: East Pacific Rise: 21°N (type locality), 17°24’S;Galapagos Spreading Center.
References:
SCHELTEMA A.H. (2000) Argonauta XIV(2): 15-25.
L. SALVINI-PLAWEN Denisia 18 (2006): 76
1A, B: Holotype in lateral (A) and ventral (B) views (anterior end at left); arrow points to pedal pit; C: Mantle sclerites; maintype elongate and often with distal swelling; D: Two radula plates in dorso-frontal view, above from a right row, below from aleft row; after SCHELTEMA (2000).
Mollusca, Solenogastres, Cavibelonia, Simrothiellidae
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Helicoradomenia bisquama SCHELTEMA, 2000
Size: Up to 3.5 mm.
Morphology: Solenogastres of cylindric body slightly taperingat anterior end, posterior end truncate, appearance rough andbumpy. Sclerites up to 135 μm long, most of which broad andthickest at margins as if formed of two joined spicules, oftenwith bifurcate distal end. Dorsofrontal sensory pit not obvious.Radula biserial with paired ventral sack, at end spirally en-rolled; radula plates with nine denticles with increasing lengthtowards lateral. Paired copulatory stylet apparatus with seven(3+3+1) elements each.
Remarks: Though there is poor information concerning the in-ternal soft organs, the present species is within Helicoradomenia(see H. juani) well-defined by the characters of the hard parts.
Biology: From vent clam (Calyptogena) or vestimentiferan (Rif-tia) beds.
Distribution: East Pacific Rise: 21°N.
References:
SCHELTEMA A.H. (2000) Argonauta XIV(2): 15-25.
L. SALVINI-PLAWEN Denisia 18 (2006): 77
1A, B: Holotype in lateral (A) and ventral view (B, anterior end at left) with part of respiratory papillae protruding from mantlecavity; C: Mantle sclerites; D: Three radula plates of a right row: two plates (above) upon radular membrane (stippled) in dorso-frontal view,w one plate (below) in ventro-abfrontal view showing basal bar with serrations; after SCHELTEMALL (2000).
Mollusca, Solenogastres, Cavibelonia, Simrothiellidae
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Helicoradomenia juani SCHELTEMA & KUZIRIAN, 1991
Size: Up to 5 mm.
Morphology: Solenogastres of cylindric body somewhat enlarg-ing posteriorly, anterior end rounded, posterior end somewhatflattened ventroterminally; appearance fuzzy due to one layer ofsolid, elongate, slightly curved sclerites, distally with bluntpoint, and measuring posteriorly up to 200 μm in length. Dor-sofrontal sensory pit obvious as transverse slit. Radula biserialwith paired ventral sack, at end spirally enrolled; radula plateswith 5-6 denticles, the lateral-most distinctly longer and closeto next one. Mouth opening within posterior atrium/vestibu-lum; with multicellular dorsal foregut gland and with shortesophagus; one pair of shortly stalked seminal receptacle.Paired copulatory stylet apparatus with two elements each.
Remarks: Helicoradomenia is particularly defined by a biserialradula with paired ventral sack, by solid, elongate sclerites inone layer and by a dorsofrontal sensory pit. Specific charactersrefer to detailed shape of hard parts (sclerites, radula plates,copulatory stylets) as well as to soft internal organs (configura-tion of foregut, of accessory genital organs and of pallial cavi-ty). There are other hydrothermal vent representatives underdescription belonging to Helicoradomenia and to different gen-era (see Fig. 2-7).
Biology: Carnivorous, but probably not on Cnidaria.
Distribution: Juan de Fuca Ridge: Endeavour Segment; Explor-er Ridge; Gorda Ridge.
1A, B: Holotype in lateral (A) and ventral views (B, anterior end at left); line points to pedal pit; scale bar 2 mm; C: Mantle scle-rites of anterior (left) and posterior body (right); scale bar 0.1 mm; D: Three radula plates: two plates (above) of the right rowin dorso-frontal view, one plate (below) of the left row in ventro-abfrontal view; scale bar 0.5 mm; after SCHELTEMA & KUZIRIAN
(1991).
Mollusca, Solenogastres, Cavibelonia, Simrothiellidae
L. SALVINI-PLAWEN Denisia 18 (2006): 78–79
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Reference:
SCHELTEMALL A.H. & A.M. KUZIRIAN (1991) Veliger 34: 195-203.
2: H. cf. acredema, in vivo specimen from East Pacific Rise:9°N, site Tica; by M. Bright.
3: Helicoradomenia sp. 1, in vivo specimen from East Pacific Rise:9°N, site Tica; by M. Bright.
4: Helicoradomenia sp. 2, body length 2 mm, in ventralview,w anterior end at left showing slit-like dorsofrontal sen-sory pit and atrio-buccal area, pedal groove (foot) wide;from East Pacific Rise: 18°S; cruise Biospeedo © Ifremer.
5: Simrothiellidae gen. et sp. 1, body length 3 mm, lateral view,anterior end at left; mantle in part with dorsal incrustation of or-ange iron oxide; from East Pacific Rise: 13°N, site PP Hot 3 (Elsa);cruise Phare © Ifremer.
6: Helicoradomenia sp. 3, body length 2.3 mm, lateral view(anterior at left), mantle with incrustation by yellow-orangedeposits; from East Pacific Rise: 21°S; cruise Biospeedo© Ifremer.
7: Two specimens of Simrothiellidae sp. in vivo from North FijiBack-Arc Basin, site White Lady; cruise TUIM06MV (June 2005,MBARI) © G. Rouse.
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Thermochiton undocostatus SAITO & OKUTANI, 1990
Size: Length up to 13 mm.
Morphology: Animal elliptical, little elevated, with a sub-cari-nated back. Girdle rather broad, with imbricating obliquely set,convex, finely ribbed scales, which possess a granular dorsaledge and with slender and smooth, marginal spiculae. Tegmen-tum (dorsally visible part of the valves) sculptured with some-what irregular, concentrically arranged undulatin costae, strongtowards the outer margins and weaker and more irregular nearthe apices. Articulamentum (layer beneath tegmentum) at themiddle of the intermediate valves with a transverse callus, su-tural laminae roughly triangular in intermediate valves, trape-
zoid in the tail valves. Valves and girdle white, exposed surfacewith rusty brown deposits. Radula ca. 4 mm long in the 13 mmspecimen, with 170 densely packed teeth rows.
Biology: Found together with the other species Lepidochitontenuidontus among bivalves Bathymodiolus and Calyptogena, si-boglinid tubeworms and shrimps. It is not yet known whetherthe two species are endemic to vents or they also occur else-where.
Distribution: Okinawa Trough: Iheya Ridge.
Reference:
SAITO H. & T. OKUTANI (1990) Venus 49(3): 165-179.
R. VON COSEL Denisia 18 (2006): 80
1 left: Dorsal view; top right: Head valve, exterior and interior; bottom: Valve 4, anterior view; after SAITO & OKUTANI (1990).
Mollusca, Polyplacophora, Neoloricata, Ischnochitonidae
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Leptochiton tenuidontus SAITO & OKUTANI, 1990
Reference:
SAITO H. & T. OKUTANI (1990) Venus 49(3): 165-179.
R. VON COSEL Denisia 18 (2006): 81
1 left: Dorsal view; top right, head valve, exterior and interior; bottom: Valve IV, anterior and dorsalVVview; after SAITO & OKUTANITT (1990).
Mollusca, Polyplacophora, Neoloricata, Leptochitonidae
Size: Length up to 16 mm.
Color: Valves pale buff with some blackish deposits, girdle buffin preserved specimens.
Morphology: Animal oblong, moderately elevated, with anevenly rounded back. Girdle narrow, with elongate calcareousscales, each ornamented with 7-8 longitudinal ribs, and withlong, smooth, needle-like marginal spiculae. Tegmentum (dor-sally visible part of the valves) sculptured with elongated,close-set granules, arranged in longitudinal series. Articula-mentum (layer beneath tegmentum) with callus at the middle
portion of each valve, sutural laminae roughly sub-triangularand widely separated. Radula long, 7.3 mm in the 16 mm spec-imen, with 193 densely packed teeth rows in the holotype.
Biology: Found together with the other species Thermochitonundocostatus among bivalves Bathymodiolus and Calyptogena, si-boglinid tubeworms and shrimps. It is not yet known whetherthe two species are endemic to vents or they also occur else-where.
Distribution: Okinawa Trough: Iheya Ridge.
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Mollusca, Gastropoda
Currently about 60 genera and more than 100 species of gas-tropods have been recognized in hydrothermal communities inthe Indo-Pacific, Indian and Atlantic Oceans. That makes thegastropods the most species-rich group of vent animals thanks toall researchers who have sent specimens to taxonomists for iden-tification. The gastropods are probably also the group whereknowledge on distribution and zoogeography is the best.
Although the present handbook aims to present a fairlycomplete overview with diagnostic information and illustra-tion, it is important to stress several points:
(1) The identification of several genera of small gastropodsis difficult and is a matter of specialist work.ff
(2) The limpets can be assigned to genus based on shell andexternal morphology only, but many can be confusingly similaron superficial examination by an untrained eye. It is sometimeseven easier to identify limpets to species than to genus.
(3) For any work deemed to have results of more than tem-porary importance and where species identification is involved,
it is recommended that voucher specimens are deposited in arecognized scientific museum collection. Many natural historymuseums readily accept such deposits. For morphological workformalin-fixed specimens are preferred, although also ethanol-preserved specimens can be used. Frozen specimens are usuallytoo macerated to show details in tentacles and epithelia. As al-ways when calcareous shells are involved, do not forget tobuffer the formalin, for example with a tea spoon of borax perlitre 4% formalin.
(4) Beginning genetic work has revealed existence of isolat-ed genotypes or populations within some of the previously rec-ognized species. In some species this is accompanied by morpho-logical differences and/or geographic separation (e.g. Lepetodrilusfucensis JOHNSON et al., in press). In others (e.g. Alviniconcha hess-leri) there are no noticeable differences in morphology and theirgeographic distribution is not easily explicable. It should be re-membered that distribution, dispersal and speciation of deep-seaanimals still is “terra incognita“ and should be explored.
(5) This manual may give the impression that the fauna is ad-equately known. However, many vent inhabitants are still knownfrom a single or very few specimens, and almost every cruise tothe site at 13°N on the East Pacific Rise (probably the bestknown gastropod fauna) has resulted in one or a couple of unde-scribed species; new species are also likely to be discovered by newexpeditions at other “well-known“ and “well-collected“ sites.
Many gastropods and other hard-bodied animals from ventsites and other reducing environments may have thick crusts ofdeposited materials covering the surface. These may be black orreddish brown due to the presence of amorphic pyrite (iron sul-phide, FeS2) and rust (ferric oxide-hydroxide mixtures) respec-tively. SEM examination often reveals intense bacterial growthin protected cracks and crevices, whereas exposed areas seem tobe grazed with only a short stub of bacterial filaments in borderzones. Exposed areas have smooth and hard surface, but theprocess behind this transition seems unknown. The descriptionof colours of the gastropod shells and periostracum is based onshells free from deposits. Moreover, the shells of many gas-tropods are covered by pustules formed by bacteria (P. Dando,pers. comm.; cf. Figure of Depressigyra). These may to some ex-tent be host specific since their appearance differs between hostffspecies. They have only been observed in vent environments.
References:
JOHNSON P., VRIJENHOEK R.C. & A. WARÉNWW (in press) Bio. Bull.SASAKI T., OKUTANI T. & K. FUJIKURA (2005) Venus 64: 87-133.
A. WARÉNWW Denisia 18 (2006): 82
1: Ifremeria nautilei from Kilo Moana, Lau Back-Arc Basin, TU-IM 07 cruise © C.R. Fisher.
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Eulepetopsis vitrea MCLEAN, 1990 “translucent limpets”
References:
FRETTER V. (1990) J. Zool. Lond. 222: 529-555.MCLEAN J. (1990) J. Zool. Lond. 222: 485-528 [503].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [123].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 83
1: Exterior, interior and lateral views; by R. von Cosel, A. Le Goff & P. Briand.
Mollusca, Gastropoda, Patelligastropoda, Lepetopsoidea, Neolepetopsidae
Size: Shell length up to 17 mm.
Morphology: Shell oblong, very flat; apex one-quarter shelllength from anterior end; shell smooth on superficial examina-tion, with fine radial striae on closer inspection. Shells im-mersed in water or ethanol nearly transparent, dried shellswhite; interior with metallic sheen when viewed at angle. Nooperculum. No appendage along foot margin
Remarks: An unnamed species known from the Kairei VentField (Indian Ocean).
Biology: Specimens have been collected on the basalt substra-tum and on the mussel Bathymodiolus. Genus endemic to vents.Larval development lecithotrophic with planktonic dispersalstage.
Distribution: East Pacific Rise: 21°N to 17°S; GalapagosSpreading Center.
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Neolepetopsis MCLEAN, 1990 “symmetrical limpets”
References:
FRETTER V. (1990) J. Zool Lond. 222: 529-555.MCLEAN J. (1990) J. Zool. Lond. 222: 485-528 [492, 496].WARÉNWW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [122-123]
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 84
1: N. gordensis, exterior, interior and lateral views; by R. von Cosel & A. Le Goff.
Mollusca, Gastropoda, Patelligastropoda, Lepetopsoidea, Neolepetopsidae
Species DistributionN. densata MCLEAN, 1990 East Pacific Rise: 12-13°N, Galapagos Spreading CenterN. gordensis MCLEAN, 1990 Gorda Ridge: 41°N, S Gulf of California: 20°N; possibly also off Peru: 5°SN. occulta MCLEAN, 1990 East Pacific Rise: 21°NN. verruca MCLEAN, 1990 East Pacific Rise: 21°N
Size: Shell length up to 4-7.5 mm (varies with species).
Morphology: Shells oblong, depressed; apex 1/3 to 2/5 shelllength from anterior end; sculpture of strong beads produced atintersections of radial and concentric ribs. Species differ in de-tails of sculpture. No operculum. No appendage on sides of foot.
Remarks: Demarcation of genus against Paralepetopsis uncer-tain.
Biology: Genus endemic to vents and seeps; some species havebeen collected on inactive chimneys devoid of other megafau-nal species. Larval development lecitotrophic with planktonicdispersal stage. As in Paralepetopsis, species demarcation basedon morphology is close to impossible since the species are fea-ture-less, variable and often badly corroded (except N. densa-ta).
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Paralepetopsis ferrugivora WARÉNWW & BOUCHET, 2001 “rust-eating limpets”
Size: Shell length up to 17 mm.
Color: Shell semi-transparent, white or slightly brown; interiorwhitish.
Morphology: Shell oblong, sturdy, depressed, without trace ofcoiling; apex one-third shell length from anterior end, alwaysworn; shell with rough surface, numerous radial ribs. No oper-culum. No remarkable appendage or extension on head or footmargin. Eyes absent.
Remarks: Demarcation of genus against Neolepetopsis uncer-tain. Other species of Paralepetopsis are common in seep envi-
ronments in the Atlantic and eastern Pacific and on whaleskeletons off California. They are difficult to identify due totheir feature-less external morphology and often corrodedshells. Several species adjust the shell to the substrate with aslender outline and concave shell base, when living on tubes.
Biology: Specimens have been collected on and among Bathy-modiolus. Intestine filled with orange-brown matter of granularstructure with nematode and ciliate fragments. Larval develop-ment lecithotrophic with planktonic dispersal stage
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [123-125].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 85
1: Exterior, interior and lateral views; by R. von Cosel, A. Le Goff, P. Briand & A. Warén.
Mollusca, Gastropoda, Patelligastropoda, Lepetopsoidea, Neolepetopsidae
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Clypeosectus MCLEAN, 1989 “slit-limpets”
References:
HASZPRUNAR G. (1989) Nat. Hist. Mus. Los Angeles Cty, Contrib. Sci. 408: 1-17 [5].MCLEAN J. (1989) Nat. Hist. Mus. Los Angeles Cty, Contrib. Sci. 407: 1-29 [18, 21].WARÉNW A. & P. BOUCHET (2001) Veliger 44 (2): 116-231 [155].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 86
1: C. delectus; top: Exteri-or and interior view; bot-tom left, same specimen,lateral view; by A. LeGoff © MNHN; bottomright: Another, inclinedview; P. Briand © Ifremer.
2: Larvae; by courtesy of L. Mullineaux.
Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Lepetodrilidae
Species DistributionC.CC curvus MCLEAN, 1989 Explorer and Juan de Fuca RidgesC. delectus MCLEAN, 1989 East Pacific Rise: 13°N and 21°N, Galapagos Spreading Center
Size: Shell length up to 5-8 mm (varies with species).
Morphology: Shells of limpet form with oblique, elongate slit;sculptured by fine radiating ribs. Slit open at margin. No oper-culum. Eyes absent.
Biology: Found with siboglinid tubeworms, details of habitatnot known. Genus endemic to vents.
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Gorgoleptis MCLEAN, 1988 “dimorphic limpets”
References:
FRETTER V. (1988) Philos. Trans. R. Soc. Lond. B 319: 33-82 [58, 64-65].MCLEAN J. (1988) Philos. Trans. R. Soc. Lond. B 319: 1-32 [19-24].WARÉNW A. & P. BOUCHET Veliger 44(2): 116-231 [154-155].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 87
1 top:G. emarginatus;middle: G. patuluswith incrustations;bottom: G. spiralis.All with exterior,interior and lateralview; by R. von Cosel& A. Le Goff.
Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Lepetodrilidae
Species DistributionG. emarginatus MCLEAN, 1988 East Pacific Rise: 9-21°NG. patulus MCLEAN, 1988 East Pacific Rise: 13°N, Galapagos Spreading CenterG. spiralis MCLEAN, 1988 East Pacific Rise: 13°N
Size: Shell length up to 3-9 mm (varies with species).
Morphology: Shells depressed, ear-shaped, with a distinctlycoiled initial whorl; sculpture of beaded or imbricate radial ribs.Operculum small, not closing the shell. Male with penis formedby expansion of snout on left side. Five pairs of long epipodialtentacles.
Biology: Details of habitat unknown. Genus endemic to vents.Larval development lecithotrophic with planktonic dispersalstage
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Lepetodrilus MCLEAN, 1988 “dimorphic limpets”
1: L. atlanticus from Mid-Atlantic Ridge; left top to bottom: Exterior, interior and lateral view;by R. von Cosel & A. Le Goff; top right: In situ © Ifremer/Atos; bottom right: P. Briand © Ifremer.
Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Lepetodrilidae
Species DistributionL. atlanticus WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: 38-23°NL. corrugatus MCLEAN, 1993 Juan de Fuca RidgeL. cristatus MCLEAN, 1988 East Pacific Rise: 13°, 21°N, Galapagos Spreading CenterL. elevatus MCLEAN, 1988 East Pacific Rise: 21°N-17°S, Galapagos Spreading Center, North Fiji Basin,
Lau Basin, Mariana BasinL. fucensis MCLEAN, 1988 Juan de Fuca RidgeL. galriftensis MCLEAN, 1988 East Pacific Rise: 9°N, Galapagos Spreading CenterL. guaymasensis MCLEAN, 1988 Guaymas BasinL. japonicus OKUTANI, FUJIKURA & SASAKI, 1993 Okinawa BasinL. nux OKUTANI, FUJIKURA & SASAKI, 1993 Okinawa BasinL. ovalis MCLEAN, 1988 East Pacific Rise: 21°N-17°S, Galapagos Spreading CenterL. pustulosus MCLEAN, 1988 East Pacific Rise: 21°N-17°S, Galapagos Spreading CenterL. schrolli BECK, 1993 Manus BasinL. tevnianus MCLEAN, 1991 East Pacific Rise: 11°NFurther species Indian Ocean hydrothermal vents
Size: Shell length up to 6-20 mm (varies with species).
Morphology: Shells of limpet form; coiling distinct to indis-tinct. Apex posterior, lower than highest shell elevation, slight-ly to strongly projecting. Species differ by shell proportions andsculpture. No operculum. Male usually with penis near base ofright cephalic tentacle. Three pairs of short epipodial tentacles.
Biology: At East Pacific Rise sites, specimens of L. elevatushave been collected in such abundance from washings of si-
boglinid tubes that there is no doubt that a primary habitat ofthese limpets is directly on Riftia and TevniaTT . Association ofchemoautotrophic bacteria with the gill confirmed for Lepeto-drilus fucensis, which has been found in densities of 400000 m-2.The genus occurs in vents and seeps. Development with free-swimming lecithotrophic larvae with planktonic dispersalstage.
Remark: Differff ent genetic types, probably species, occur, espe-cially of the forms similar to L. elevatus and L. schrolli.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 88–90
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2: L. cristatus, on an active chimney, near an alvinellid colony; East Pacific Rise: 13°N; cruise Phare. P. Briand © Ifremer.
3: L. fucensis, among tubeworms Ridgeia picesae, Juan de Fuca Ridge; by courtesy of K. Juniper.
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References:
BATES A.E., TUNNICLIFFE V. & W.L. RAYMOND (2005) Mar. Ecol. Prog. Ser. 305: 1-15.BECK L. (1993) Ann. Nat.hist. Mus. Wien B 94-95: 167-179.DE BURGH M.E. & C.L. SINGLA (1984) Mar. Biol. 84 1-6.FRETTER V. (1988) Philos. Trans. R. Soc. Lond. B 319: 33-82.MCLEAN J. (1988) Philos. Trans. R. Soc. Lond. B 319: 1-32.MCLEAN J. (1993) Veliger 36: 27-35.OKUTANI T., FUJIKURA K. & T. SASAKI (1993) Bull. Natn. Sci. Mus.,Tokyo, Ser. A 19(4): 123-143.WARÉNWW A. & P.PP BOUCHET (2001) Veliger 44(2): 116-231 [143-154].
4 top: L. elevatus; middle: L. ovalis; bottom: L. pustulosus. All with exterior, interior and lateral view; by R. von Cosel & A. LeGoff; in situ views, limpets on tubes of Riftia pachyptila © Ifremer/Phare.
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1: P. midatlantica; top: Dorsal, lateral and ventral view, no precision of origin; by R. von Cosel & A. Le Goff; middle: Specimenfrom Logatchev (Mid-Atlantic Ridge), dorsal and ventral view; bottom: specimen from Snake Pit; by A. Warén.
Pseudorimula MCLEAN, 1989 “slit-limpets”
Size: Shell length up to 5-8 mm (varies with species).
Morphology: Shells of limpet form with oblique, elongate slit;sculptured by fine radiating ribs. Slit closed at margin. No op-erculum. Eyes absent.
Biology: On rocks. P. midatlantica also occurs on Bathymodiolusmussels. Genus endemic to vents. Development without plank-totrophic larvae.
Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Lepetodrilidae
Species DistributionP.PP marianae MCLEAN, 1989 Mariana Back-Arc BasinP. midatlantica MCLEAN, 1992 Mid-Atlantic Ridge: 38-15°N
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 91–92
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References:
MCLEAN J. (1989) Nat. Hist. Mus. Los Angeles Cty., Contrib. Sci. 407: 1-29 [22].MCLEAN J. (1992) Nautilus 106: 115-118 [116].WARÉNW A. & P.PP BOUCHET (2001) Veliger 44 (2): 116-231 [155-157].
2: P. midatlantica; by P. Briand.
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Pyropelta MCLEAN & HASZPRUNAR, 1987
Size: Shell length up to 5 mm.
Morphology: The family is defined by anatomical charactersand the shells are rather featureless, cap-shaped, with centralapex; sculpture unknown, exterior surface deteriorated. No op-erculum. Foot with a pair of posterior epipodial tentacles.
Biology: Specimens have been collected on sulphide crust inthe vents and on surrounding rocks, and they are absent in
washings of vestimentiferan tubes. Pyropelta corymba lives onshells of Provanna spp., grazing bacteria. The genus occurs onvents, seeps and also on whale bones. Larval developmentlecithotrophic with planktonic dispersal stage. The featurelessshell, always badly corroded, makes identification very uncer-tain.
References:
BECK L. (1996) Arch. Moll. 125: 87-103.MCLEAN J. & G. HASZPRUNAR (1987) Veliger 30: 196-205 [197-200].SASAKI T., OKUTANI T. & K. FUJIKURA (2003) Veliger 46(3): 189-210 [197].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [125-129].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 93
1: P.PP corymbar ; top left: Inclined and dorsal view of a specimen; right: The same specimen; SEM by A. Warén; bottom left: Lateralview of another specimen, from Guaymas; by P. Briand © Ifremer; bottom right: Another specimen; by J. McLean.
Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Pyropeltidae
Species DistributionP.PP bohlei BECK, 1996 Lihir Volcano, Edison Seamount, West PacificP. corymba MCLEAN & HASZPRUNAR, 1987 Oregon Margin: 45°N, Guaymas BasinP. musaica MCLEAN & HASZPRUNAR, 1987 Juan de Fuca Ridge: 45°56’N, off California: 36-33°N, Jalisco Block: 20°NP.PP yamato SASAKI, OKUTANI & FUJIKURA, 2003 Izu-Ogasawara Arc
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Bruceiella globulus WARÉNWW & BOUCHET, 1993
References:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 94
1 upper left: Specimen 1, 1.19 mm, apertural view, with periostracum; upper right: specimen 2, 1.19 mm, cleaned;both from cruise Biolau, Lau Back-Arc Basin; lower left: specimen 3, apertural view, 2.31 mm; lower right, specimen4, apical view,w 1.9 mm; both cruise Starmer II North Fiji Back-Arc Basin; after WARÉNWW & BOUCHET (1993).
Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Skeneidae
Size: Shell diameter up to 2.3 mm.
Morphology: Shells globular, almost as high as broad, with lowspire, rather robust, surface almost smooth. Umbilicus deep andwide. Whorls rounded, with a deep suture, aperture circular, al-most or completely detached from the following whorl. Proto-conch with spiral sculpture, about 0.65 whorls, with a diameterof about 340 μm. Operculum round, multispiral and thin.
Remark: Another species of this genus, B. athlia WAWW RÉN &BOUCHET, 2001, was described from cold seeps of the AleutianTrench, Shumagin Site, 54°18’N, 157°12’W, 4808 m.
Biology: At hydrothermal vents.
Distribution: North Fiji and Lau Back-Arc Basins.
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Fucaria WARÉNWW & BOUCHET, 1993
1: View ofdifferentsides ofseveralspecimens;by R. vonCosel.
Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Skeneidae
Species DistributionFFF mystax WARÉN & BOUCHET, 2001 S of Lihir, Edison Seamount, West PacificF. striata WARÉN & BOUCHET, 1993 Juan de Fuca Ridge
Size: Diameter up to 5.8 mm (F. mystax) and 10.6 mm (F. striata).
Color: White, greenish or brownish.
Morphology: Shells turbinate, about as high as broad, rathersturdy (only known with top of the shell eroded), almost closedumbilicus; operculum with central nucleus. Surface smooth (F.mystax) or sculptured by spiral lirae and groves (F. striata).Aperture rounded, completely closed by horny, multispiral op-
erculum. No eyes. Right neck-lobe fused with eye-lobe andequipped with marginal tentacles.
Biology: Species of Fucaria occur only in seeps and vents. Foodconsists of the detritus layer on the bottom (stomach contentsconsist of mineral particles, sponge spicules and radiolarianfragments in a matrix of mucus and detritus). Larval develop-ment lecithotrophic with planktonic dispersal stage.
References:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [15-17].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [136-137].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 95
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Leptogyra inflata WARÉNWW & BOUCHET, 1993
Size: Shell diameter up to 1.32 mm.
Morphology: Shell very small, skeneiform, broader than high,with evenly rounded whorls. Umbilicus deep and broad.Whorls with a deep suture. Protoconch with 0.6 whorls, regu-larly coiled, the initial part with an irregular net sculpture, theremaining half smooth; diameter 200 μm. Teleoconch with
2.25 whorls, with about nine low and indistinct spiral ribs andaxial growth lines. Periostracum thin and transparent. Opercu-lum thin, multispiral, brownish.
Biology: Only known from hydrothermal vent sites.
Distribution: Lau Back-Arc Basin.
Reference:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.
A. WARÉNWW & P. BOUCHET & R. VON COSEL Denisia 18 (2006): 96
1: Holotype, 1.32 mm; top: apical view; bottom left: apertural view wit periostracum and in-crustations; bottom right: same specimen, cleaned; cruise Biolau; after WARÉNWW & BOUCHET (1993).
3: Umbilicus; after WARÉNWW & BOUCHET (1993). 2: Protoconch; after WARÉNWW & BOUCHET (1993).
Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Skeneidae
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Protolira WARÉNWW & BOUCHET, 1993
References:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [22-26].WARÉNW A. & P. BOUCHET (2001) Veliger 44 (2): 116-231 [138-139].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 97
1 top: P. thorvaldssoni, apertural,abapertural and lateral view; by R. von Cosel & A. Le Goff;
right: P. valvatoides; by P. Briand.
Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Skeneidae
Species DistributionP.PP thorvaldssoni WARÉN, 1996 Mid-Atlantic Ridge: Snake Pit, to IcelandP. valvatoides WARÉN & BOUCHET, 1993 Mid-Atlantic Ridge: Menez Gwen and Lucky Strike
Size: Shell diameter up to 4.2 mm.
Morphology: Shells small, fragile, turbinate, about as high asbroad or broader than high, with spirally arranged micro-tuber-cles on the protoconch (spire almost always corroded) and analmost smooth, globular shell with open umbilicus and a deepsuture. Aperture circular, completely closed by horny opercu-lum. Shell whitish, greenish, brownish; surface covered bythick periostracum and mineral deposits. Right neck-lobe con-tinuous with eye-lobe. Right anterior corner of propodiumdrawn out into a spirally coiled tentacle.
Biology: Found among Bathymodiolus, on sediment and onrocks, Protolira thorvaldssoni also on whale bones, from which itwas originally described. Food consists of the detrital surfacelayer on the bottom (stomach contents consist of a mixture oforganic material and mineral particles, scattered spongespicules, polychaete bristles, diatoms and crustacean frag-ments). Larval development lecithotrophic with planktonicdispersal stage.
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Sutilizona MCLEAN, 1989 “slit-limpet”
Size: Shell length up to 2.4 mm.
Morphology: Shells fragile, limpet-like with long, moderatelyoblique slit; axial and spiral sculpture of various development.Protoconch with rough pit sculpture. Slit closed (S. theca, S.tunnicliffae) or open (S. pterodon) near the margin. Operculumsmall and thin, vestigial, multispiral with central nucleus.
Biology: Details of habitat not known. Genus endemic tovents. Larval development lecithotrophic with planktonic dis-persal stage.
References:
MCLEAN J. (1989) Nat. Hist. Mus. Los Angeles Cty., Contrib. Sci. 407: 1-29 [15].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [141-143].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 98
1: S. pterodon; two specimens, dorsal views and apertural view (SEM); by A. Warén.
Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Sutilizonidae
Species DistributionS. pterodon WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: Snake Pit,S. theca MCLEAN, 1989 East Pacific Rise: 12-13°NS. tunnicliffae WARÉN & BOUCHET, 2001 Juan de Fuca Ridge: Endeavour Segment
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Temnocinclis euripes MCLEAN, 1989 “slit-limpet”
Size: Shell length up to 4 mm.
Morphology: Coiled limpet with long, moderately oblique slit;reticulate sculpture formed by intersection of fine radiating ribsand concentric ridges. Operculum small, vestigial.
Biology: Details of habitat not known. Larval developmentlecithotrophic with planktonic dispersal stage.
Distribution: Juan de Fuca Ridge.
References:
HASZPRUNAR G. (1989) Nat. Hist. Mus. Los Angeles Cty., Contrib. Sci. 408: 1-17 [3].MCLEAN J. (1989) Nat. Hist. Mus. Los Angeles Cty., Contrib. Sci. 407: 1-29 [5-7].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [143].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 99
1: Different views; by R. von Cosel & P.PP Briand.
2: Specimen, dorsal view; by R. von Cosel & P. Briand.
Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Sutilizonidae
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Temnozaga parilis MCLEAN, 1989
Size: Shell length up to 4 mm.
Morphology: Symmetrical limpet with median, oblique, elon-gate slit; sculptured by strong radiating ribs bearing raisedscales. Operculum small, vestigial.
Biology: Details of habitat not known. Monotypic genus en-demic in vents. Larval development lecithotrophic with plank-tonic dispersal stage.
Distribution: East Pacific Rise: 21-13°N.
References:
HASZPRUNAR G. (1989) Nat. Hist. Mus. Los Angeles Cty., Contrib. Sci. 408: 1-17.MCLEAN J. (1989) Nat. Hist. Mus. Los Angeles Cty., Contrib. Sci. 407: 1-29.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 100
1: Habitus; by A. Warén & P.PP Briand.
Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Sutilizonidae
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Bathymargarites symplector WARÉNWW & BOUCHET, 1989
References:
WARÉNW A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [87-94].WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [11-13].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 101
1 top: Specimen 1, apertural, abapertural and lateral view by A. Le Goff © MNHN;bottom: Specimen 2, inclined dorsal view,ww apertural view,w inclined basal view; by P. Briand © Ifremer.
2: Larva; by courtesyof L. Mullineaux.
Mollusca, Gastropoda, Vetigastropoda, Trochoidea, Chilodontidae
Size: Shell up to 11 mm.
Color: Off-white.
Morphology: Shell turbinate, about as high as broad, almostsmooth, without umbilicus. Umbilical area and columella cov-ered by a solid callus in adult specimens. Interior with a well de-veloped nacreous layer. Eyes at base of cephalic tentacles, butpigment spot of variable shape, sometimes double or absent.Male with left neck lobe modified to function as a penis.
Biology: Specimens have been obtained from rubble samplesand washings of Riftia and Calyptogena. Monotypic genus en-demic to vents. Food consists of the detrital surface layer on thebottom. Stomach content consists of mineral particles, spongespicules, polychaete bristles, crustacean and diatom fragments,radiolarian tests in a matrix of undefined organic matter. Larvaldevelopment lecithotrophic with planktonic dispersal stage.
Distribution: East Pacific Rise: 13°N and 21°N.
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Helicrenion reticulatum WARÉNWW & BOUCHET, 1993
Size: Shell diameter up to 1.56 mm.
Morphology: Shell very small, skeneiform, like a vitrinellid, de-pressed, fragile, with an almost smooth surface. Umbilicuswidely open, without any basal area or spiral sculpture. Whorlswith a deep suture. Protoconch with 0.6 whorls, the initial partwith sharp spiral lines and irregularly spaced axial lines, form-ing a large-meshed net sculpture, later half smooth. Teleoconchwith about 1.5 rapidly increasing whorls, with very fine, close-
set granular incremental axial lines. Animal not known, radulawith unusual few marginal teeth for a neomphalid radula type.
Biology: Known only from hydrothermal vent sites.
Distribution: Only known from Lau Back-Arc Basin: Hine Hi-na vent field.
Reference:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 102
1 top: holotype apertural view,w 1.56 mm; bottom: paratype 1.10 mm, apical and basal view;Lau Basin; from WARÉNWW & BOUCHET (1993).
2: Protoconch showing reticulated sculpture; LauBasin; from WARÉNWW & BOUCHET (1993).
3: Protoconch showing umbilicus; Lau Basin;from WARÉNWW & BOUCHET (1993).
Mollusca, Gastropoda, Vetigastropoda, Trochoidea, Trochidae
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Vetulonia phalcata WARÉNWW & BOUCHET, 1993
Size: Shell diameter up to 1.24 mm.
Morphology: Shell trochiform, thin and fragile, with regularand sharp axial ribs parallel to the outer lip, ribs basally distin-cly flexuous. Deep, narrow umbilicus which has an indistinct,steeply ascending spiral rib. Whorls slightly depressed, with adeep suture. Surface between the axial ribs almost smooth. Pro-toconch with 0.65 whorls, strongly corroded, with a diameter ofat least 200 μm; teleoconch with 2.45 whorls; operculumround, multispiral, thin and transparent. Animal not known.
Biology: At vent sites; it is likely that the species feeds on su-perficial detritus and bacterial film on the bottom, like manyother deep water archaegastropods.
Distribution: North Fiji Back-Arc Basin.
Reference:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 103
1: Holotype, 1.06 mm; left apical view; right apertural view: collected during the cruise Starmer II; after WARÉNWW & BOUCHET (1993).
Mollusca, Gastropoda, Vetigastropoda, Trochoidea, Trochidae
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Cyathermia naticoides WARÉNWW & BOUCHET, 1989
1: Apertural, abapertural and lateral views; by R. von Cosel, A. Le Goff & P. Briand.
2: Larva; by courtesyof L. Mullineaux.
Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae
Size: Shell diameter up to 7 mm.
Color: White.
Morphology: Shell globular, regularly coiled, without sculpture.Adults with deep notch in the lower part of the outer lip. Verylarge bipectinate gill. Left cephalic tentacle modified to a penis.
Biology: Specimens have been found in abundance in washingsof Riftia tubes, more rarely with Alvinella tubes. Monotypicgenus endemic to vents. Probably a filter-feeder, possibly incombination with cleaning the worms’ tubes of bacterialgrowth. Larval development lecithotrophic with planktonicdispersal stage.
Distribution: East Pacific Rise: 9-21°N.
References:
WARÉNW A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [69-70].WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22 (1): 1-90 [33].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [158].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 104
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Lacunoides WARÉNWW & BOUCHET, 1989
Size: Shell diameter up to 2.5 mm.
Morphology: Shells small, thin and fragile, colourless andtransparent, globular, with low, depressed spire, large apertureand rapidly increasing diameter of the whorls. Surface withfine, dense, sometimes slightly irregularly spaced, sharp axiallines or low lamellae and still finer, dense spiral striations. Um-
bilicus not present or indistinct. Whorls round with a deep su-ture. Protoconch with diameter of 160 μm (L. exquisitus ) or180 μm (L. vitreus), about 0.5 whorls, the initial part with ir-regular net sculpture, later half smooth. Teleoconch with 2.25(L. exquisitus ) or about 2 (L. vitreus) round whorls. Operculummultispiral, thin and without color, slightly larger than theaperture.
Biology: Near hydrothermal vents on mussel beds or on otherhard substrate.
References:
WARÉNW A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102.WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 105
1: L. exquisitus, top specimen 1, aperturalview,ww 3.4 mm; bottom specimen 2, apicalview, 2.3 mm; from WARÉNWW & BOUCHET (1989).
2: L. exquisitus, close-up of protoconch ofthe same specimen; from WARÉNWW & BOUCHET
(1989).
3: L. vitreus, top holotype, apertural view,2.4 mm; bottom paratype, apical view,ww 2.2mm; from WARÉNWW & BOUCHET (1989).
4: L. vitreus: close-up of protoconch ofthe same paratype; from WARÉNW &BOUCHET (1989).
Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae
Species DistributionL. exquisitus Galapagos Spreading Center
L. vitreus Juan de Fuca Ridge: Axial Seamount – Ashes vent field
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Melanodrymia HICKMAN, 1984
References:
HASZPRUNAR G. (1989) Acta Zool. 70: 175-186.HICKMAN C.S. (1984) Zool. Scr. 13: 19-25 [19-20].WARÉNW A. & P. BOUCHET 1989) Zool. Scr. 18: 67-102 [75].WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22(1): 1-90 [41-44].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [158-161].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 106
1: Top: TT M. aurantiaca; by R. von Cosel; middle: M. aurantiaca;by P. Briand © Ifremer; bottom: M. galeronae; by A. Warén.
2: M. aurantiaca;by P. Briand © Ifremer.
Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae
Species DistributionM. aurantiaca HICKMAN, 1984 East Pacific Rise: 13°N, 21°N-17°SM. brightae WARÉN & BOUCHET, 1993 Juan de Fuca Ridge: Endeavour segmentM. galeronae WARÉN & BOUCHET, 2001 East Pacific Rise: 13°NMelanodrymia sp. “rust covered“ East Pacific Rise: 13°N
Size: Shell diameter up to 3.5 mm.
Color: Rusty orange (M. aurantiaca) or whitish (M. brightae).
Morphology: Shells depressed (M. aurantiaca, M. brightae) orhigher than broad (M. galeronae) with one or two strong pe-ripheral keels. Umbilicus open. Surface of shell above and be-low keel covered by raised riblets. Most specimens have a thickmineral crust.
Biology: Genus with three species endemic in vents. Thespecies are locally common in washings of Riftia, Ridgeia, Ca-lyptogena and Alvinella. Genus endemic to vents. Food consistsof the detrital surface layer of the bottom. Larval developmentlecithotrophic with planktonic dispersal stage.
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Neomphalus fretterae MCLEAN, 1981
Size: Shell diameter up to 30 mm.
Morphology: Shell cap-shaped with coiled apical whorl, apexsubcentral, sculptured with fine radiating ribs. No trace of op-erculum. Head with long neck, no snout, left cephalic tentacleenlarged.
Biology: Occurs locally in dense aggregations on the walls ofthe vents. The Neomphalidae are endemic to vents. Probablyfilter feeder. Larval development lecithotrophic with plankton-ic dispersal stage.
Distribution: Common at Galapagos Spreading Center, rare atthe East Pacific Rise: 9-21°N.
References:
FRETTER V., GRAHAM A. & J. MCLEAN (1981) Malacologia 21: 337-361.MCLEAN J. (1981) Malacologia 21: 291-336 [294].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [162].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 107
1: Habitus; by R. von Cosel & P. Briand.
2: Habitus; by R. von Cosel & P. Briand.
3: In situ specimens, from East PacificRise: 13°N © Ifremer.
Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae
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Pachydermia WARÉNWW & BOUCHET, 1989
References:
WARÉNW A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [75-80].WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [40-41].WARÉNW A. & P. BOUCHET (2001) Veliger 44 (2): 116-231 [161-162].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 108
1 top left to right: Apertural, abapertural and lateral view of P. laevis; by A. Le Goff © MNHN; bottom: anotherspecimen; by P.PP Briand © Ifremer.
Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae
Species DistributionP.PP laevis WARÉN & BOUCHET, 1989 East Pacific Rise: 21°N -17°SP. sculpta WARÉN & BOUCHET, 1993 North Fiji and Lau Back-Arc Basins
Size: Shell height up to 4.6 mm.
Morphology: Shell small and rather fragile, regularly coiled, upto 3.5 whorls, with part of the body whorl disjunct and with cir-cular aperture. Surface of protoconch net-sculptured, teleo-conch with fine incremental and indistinct spiral lines, other-wise smooth. Shell beige to greenish, interior not nacreous; pe-riostracum thick. Specimens are often encrusted with mineraldeposits. Operculum multispiral, closing the aperture com-pletely.
Biology: Many specimens found in washings of tubes of Alvinel-la and siboglinids but both species seems to occur also on othersubstrates. Genus endemic to vents. Stomach contents indicatedetritus feeding. Larval development lecithotrophic withplanktonic dispersal stage.
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Planorbidella WARÉNWW & BOUCHET, 1989
Size: Shell diameter up to 5.1 mm (P. planispira) and 1.56 mm(P. depressa).
Morphology: Shells flat-spired, regularly coiled, aperture al-most circular and strongly prosocline, umbilicus very wide anddeep. Surface smooth (P. planispira) or with strong, oblique ax-ial ribs and fine spiral cords (P. depressa), protoconch with fine-mesh net sculpture. Last part of the body whorl detached. Op-
erculum multispiral, completely closing the aperture. Speci-mens may be partly encrusted with mineral deposits.
Biology: Found in washings of tubes of Alvinella and siboglin-ids. Genus endemic to vents. Stomach contained only unde-fined organic material. Larval development lecithotrophic withplanktonic dispersal stage.
References:
WARÉNW A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [81-82].WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [35-39].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [162].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 109
1 top left, right and bottom: Dorsal, ventral and apertural view of P.PP planispira; by R. von Cosel & A. Le Goff.
Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae
Species DistributionP. depressa WARÉN & BOUCHET, 1993 Lau Back-Arc Basin: Hine HinaP.PP planispira WARÉN & BOUCHET, 1989 East Pacific Rise: 21°N-17°S
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Symmetromphalus MCLEAN, 1990
References:
BECK L. (1992) Ann. Nat.hist. Mus. Wien B 93: 243-257.MCLEAN J. (1990) Nautilus 104: 77-86.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 110
1: S. regularis; by R. von Cosel & P.PP Briand.
Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae
Size: Shell length up to 14-21 mm (varies with species).
Morphology: Shells of limpet form with coiled apical whorl,sculptured by finely beaded radial ribs. Operculum small, ves-tigial. Head with long neck, no snout, left cephalic tentacle of
male enormously distended, bearing a deep sperm groove con-nected to neck groove. Gill large, overlying head.
Biology: Symmetrophalus regularis occurs in dense aggregationson the walls of vents; the species at Lau and Fiji lives on mus-sels and on the subsutural ramp of Ifremeria nautilei. Endemic tovents. Sexually dimorphic, with males much smaller. Feedingbiology not known. Pallial furrow and especially epipodial ten-tacles with growth of large filiform bacteria. Larval develop-ment lecithotrophic with planktonic dispersal stage.
Species DistributionS. hageni BECK, 1992 Manus Back-Arc BasinS. regularis MCLEAN, 1990 Mariana Back-Arc BasinSymmetromphalus sp. North Fiji and Lau Back-Arc Basins
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Ctenopelta porifera WARÉNWW & BOUCHET, 1993
Size: Shell length up to 10 mm.
Morphology: Shell depressed, ear-shaped, sculptured with adozen spirally arranged rows of hollow conical tubercules con-nected via pore to the interior of the shell. Surrounding the tu-bercules are soft, tubular hollow appendages of organic materi-al. No eyes. Sides of foot and epipodium finely setose; posteriorpart of visceral mass carrying warts corresponding to the poresin the shell.
Biology: Genus with a single species endemic to vents. Speci-mens have been rarely collected in washings of tubes of Tevniaand Riftia or sulphide crusts. The setae of the foot and thestrange tubular processes on the back of the shell may be in-volved in some kind of symbiosis with chemosynthetic organ-isms (?). Sexes separate, females larger than males. Larval de-velopment lecithotrophic with planktonic dispersal stage.
Distribution: East Pacific Rise: 13°N.
References:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [33-35].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [170].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 111
1: Several specimens; by R. von Cosel.
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
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Depressigyra globulus WARÉNWW & BOUCHET, 1989
Size: Shell diameter up to 5.4 mm.
Color: Shell greenish to brownish, interior not nacreous. Pe-riostracum thick, brownish green.
Morphology: Shell broader than high with rather large bodywhorl, about three whorls; aperture subcircular with an indis-tinct shallow basal notch. Umbilicus reduced to a small chink,suture deep. Protoconch strongly ridged, teleoconch with irreg-ular incremental lines, otherwise smooth. Operculum multispi-ral, densely coiled, with central nucleus. Animal with tentaclesof even size in both sexes and a snout of approximately evenwidth.
Biology: On vestimentiferan tubes, extremely common. Occa-sionally, near acidic outflows, the calcareous layer may be dis-solved and the living animal is surrounded only by the strongperiostracum. Monotypic genus endemic to vents. Stomachcontains amorphous organic matter. Larval developmentlecithotrophic with planktonic dispersal stage.
Distribution: Juan de Fuca Ridge.
References:
BATESAA B. & V.VV TUNNICLIFFE (2006) Mar. Ecol. Prog. Ser. 305: 1-15.WARÉNW A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [80-81].WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [35].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [173].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 112
1: Several specimens; by R. von Cosel © MNHN.
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
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Echinopelta fistulosa MCLEAN, 1989 “tapersnout limpet”
Size: Shell length up to 9 mm.
Morphology: Shell of limpet form, apex close to posterior mar-gin but left of center. Sculpture with widely spaced tubularspines. Periostracum thick. Large specimens coated with rustcoloured iron deposits. No operculum. Mantle edge bearing nu-merous, crowded and elongate tentacles.
Biology: Common on outer face of active black smokers.Monotypic genus endemic to vents. Larval developmentlecithotrophic with planktonic dispersal stage.
Distribution: East Pacific Rise: 21°N, 13°N.
References:
FRETTER V. (1989) J. Zool. Lond. 218: 123-169.MCLEAN J. (1989) Zool. Scr. 18: 49-66 [58-60].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [170].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 113
1: Two specimens in various views; by R. von Cosel.TT
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
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Hirtopelta MCLEAN, 1989
References:
BECK L.A. (2002) Arch. Moll. 130 (1-2): 249-257.FRETTER V. (1989) J. Zool. Lond. 218: 123-169.MCLEAN L. (1989) Zool. Scr. 18: 49-66 [60-62].WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [35].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [169].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 114
1: All specimens H. hirta; D. Brabant © MNHN and P. Briand © Ifremer.
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
Species DistributionH. hirta MCLEAN, 1989 East Pacific Rise: 13-21°NH. tufari BECK, 2002 East Pacific Rise: 21°S
Size: Shell length up to 13 mm.
Color: Olive-brown.
Morphology: Shells more or less depressed, ear-shaped, sculp-tured by raised, scale-like projections arranged along growthlines. Shells usually covered by rust-like crusts. Gill huge in re-lation to body size. Stomach and intestine very narrow, intes-tine forming only a very short simple loop.
Biology: Genus endemic to vents. Gill and intestine morphol-ogy indicate that Hirtopelta uses the gill for chemosyntheticpurposes. Larval development lecithotrophic with planktonicdispersal stage.
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Lirapex WARÉNWW & BOUCHET, 1989
1 top: Apertural view of two specimens of L. costatus; by A. Warén; bottom: Same species;by P.PP Briand.
2: Larvae; by courtesyof L. Mullineaux.
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
Species DistributionL. costellata WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: Lucky Strike, Snake PitL. granularis WARÉN & BOUCHET, 1989 East Pacific Rise: 9-21°NL. humata WARÉN & BOUCHET, 1989 East Pacific Rise: 21°N
Size: Shell height up to 3.4 mm.
Morphology: Shells very small to small, valvatoid, regularlycoiled, about three whorls, rather sturdy, aperture almost circu-lar, umbilicus narrow to wide, suture deep, last part of bodywhorl often detached. Protoconch with sculpture of strong spi-ral ridges, teleoconch with axial ribs, variable from species tospecies. Shell whitish to brownish with more or less thick pe-riostracum; specimens are often encrusted with mineral de-
posits. Operculum multispiral with central nucleus, completelyclosing the aperture. Head with simple tentacles and a snout ofeven width.
Biology: On soft bottom and among Bathymodiolus. Genus en-demic to vents. Stomach contains amorphous organic materialbut rarely sponge spicules, crustacean remains. Larval develop-ment lecithotrophic with planktonic dispersal stage.
References:
WARÉNW A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [84-86].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [170-171].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 115
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Nodopelta MCLEAN, 1989 “tapersnout limpets“
1 top: N. heminoda; middle: N. subnoda; bottom: N. rigneae. All with exterior,rr interior and lateral view. Top andTTmiddle row by R. von Cosel & A. Le Goff; bottom row by A. Warén.
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
Species DistributionN. heminoda MCLEAN, 1989 East Pacific Rise: 13°N, 21°NN. rigneae WARÉN & BOUCHET, 2001 East Pacific Rise: 13°NN. subnoda MCLEAN, 1989 East Pacific Rise: 13°N
Size: Shell length up to 20 mm.
Color: Beige to olive-brown.
Morphology: Shells limpet-shaped, depressed, apex close toposterior margin but not overhanging it. Sculpture finelyclathrate with scattered imbricate nodes. Periostracum thick.No eyes, no operculum. Mantle margin with transverse ridgesaligned perpendicular to mantle edge.
Biology: Closely associated with black smokers, recovered fromaggregations of Alvinella. Genus endemic to East Pacific Risevents. Larval development lecithotrophic with planktonic dis-persal stage.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 116–117
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References:
FRETTER V. (1989) J. Zool. Lond. 218: 123-169.MCLEAN J. (1989) Zool. Scr. 18: 49-66 [53-56].WARÉNW A. & P.PP BOUCHET (2001) Veliger 44(2): 116-231 [168-169].
2: Nodopelta sp., exterior, interior and close up of interior view; by S. Hourdez © Roscoff.
3: N. subnoda top middle and right; bottom left and middle; bottom right is N. heminoda; by P. Briand © Ifremer.
4: Larva (SEM); by courtesy of L. Mullineaux. 5: In situ Nodopelta sp. among alvinellid worms; EastPacific Rise: 13°N, Phare cruise © Ifremer.
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Peltospira MCLEAN, 1989 “tapersnout limpets”
Size: Shell length up to 12 mm.
Color: Olive-brown.
Morphology: Shells depressed, ear-shaped, with a distinctlycoiled initial whorl; sculptured by numerous concentric lamel-lae (species differ in sculpture). Strong periostracum. Opercu-fflum small, not closing the shell (P. operculata) or even absent
(P. delicata). Eyes absent. Epipodium bearing club-shapedprocesses of irregular size, along opercular lobe.
Biology: Specimens have been collected from washings ofAlvinella and appear to be closely associated with active smok-ers. Genus endemic to vents. Larval development lecithotroph-ic with planktonic dispersal stage.
References:
FRETTER V. (1989) J. Zool. Lond. 218: 123-169.MCLEAN J. (1989) Zool. Scr. 18: 49-66 [51-53].WARÉNW A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [84-85].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [165-168].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 118
1 top: Four specimens of P.smaragdina; by A. Le Goff © MNHN; middle: Five specimensof P. operculata; from these theupper three specimens by A. LeGoff © MNHN and the lower twospecimens with lateral and exteri-or view; by P. Briand © Ifremer;bottom: Three specimens of P.delicata, interior,rr lateral andexterior; by A. Le Goff © MNHN.
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
Species DistributionP.PP delicata MCLEAN, 1989 East Pacific Rise: 9-13°NP. lamellifera WARÉN & BOUCHET, 1989 East Pacific Rise: 13°NP. operculata MCLEAN, 1989 East Pacific Rise: 9-21°N, 17°SP.PP smaragdina WARÉN & BOUCHET, 2001 Mid Atlantic Ridge: 15-38°N
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Rhynchopelta concentrica MCLEAN, 1989 “tapersnout limpet”
Size: Shell length up to 13 mm.
Color: Yellowish-brownish.
Morphology: Shell of limpet form, apex projecting close to theposterior margin. Sculpture of fine concentric ridges and radialstriae. Periostracum thin. No operculum. Epipodium and man-tle edge simple, without modification.
Biology: Associated with the tubes of Riftia. Genus endemic tovents. Larval development lecithotrophic with planktonic dis-persal stage.
Distribution: East Pacific Rise: 21-17°N.
References:
FRETTER V. (1989) J. Zool. Lond. 218: 123-169.MCLEAN J. (1989) Zool. Scr. 18: 49-66 [57-58].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [170].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 119
1 top: Exterior, lateral and interior view; by R. von Cosel & A. Le Goff; bottom: Exterior, lateral and interi-or view; by P. Briand.
2: Larva; by courtesyof L. Mullineaux.
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
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“scaly foot gastropod”
Size: Shell length up to 50 mm.
Morphology: Shell globular, ear-shaped, distinctly coiled;sculptured by numerous spiral ribs and growth lines. Perios-tracum thick, dark olive brown. Operculum absent in adult.Cephalic tentacles thick, conical without eyes. Epipodiumstrongly reduced, consisting of a series of inconspicuous tuber-cles on the side of the foot above the scales. Operculum modi-fied into several hundred horizontally aligned scales, arrangedin a roof tile fashion along the sides of the foot. Scales coveredby thick layers of quite pure pyrite and greigite (iron sulphides)deposited in a very uniform way, indicating active participationby the snail in the process.
Remark: This genus and species is still not formally named, butits conspicuous morphology and interesting symbiosis makes itdesirable to include this novelty.
Biology: The scaly-foot gastropod harbours thiotrophic γ-pro-γγteobacteria in an enormously enlarged oesophageal gland. It isa sedentary organism firmly attached to rocks at the base ofblack smoker chimneys. Genus endemic to vents. Sexes sepa-rate, sperm transfer by spermatophores. Developmentlecithotrophic, presumably with a planktonic dispersal stage.
Distribution: Indian Ocean: Rodriguez Triple Junction.
References:
WARÉNW A., BENGTSSON S., GOFFREDI S.K. & C. VANVV DOVER (2003) Science 302: 1007.GOFFREDI S.K., WARÉNWW A., ORPHAN V.J., VANVV DOVER C.L. & R.C. VRIJENHOEK (2004) Appl. Environ. Microbiol. 70(5): 3082-3090.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 120
1 top from left to right: Complete specimen, lateral, ventral view, and front view of head-foot (shell and mantle removed); bot-tom from left to right: Shell, front, apical, and basal view. Maximum diameter of complete specimen 50 mm; by A. Warén.
Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae
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Olgasolaris BECK, 1992
Size: Shell diameter up to 13 mm.
Color: Animal pinkish-reddish when alive.
Morphology: Shells almost circular, of limpet shape, depressed,with subcentral apex; sculptured by very fine, radiating, beadedribs; narrow shelf-like septum on the posterior inner side of
shell. Small, vestigial operculum. Animal with large oral lobe,penis near right cephalic tentacle. Eyes rudimentary; similarsize in veliger larva and adult.
Biology: Genus endemic to vents. Feeds by grazing bacterialmats from surfaces of sulphide chimneys and of shells of bi-valves and gastropods. The reddish colour of live animals isprobably caused by hemoglobin. Larval development withplanktotrophic larvae. Egg capsules, 1 mm diameter, depositedon shells of other molluscs, often in large numbers on Ifremeria.
Reference:
BECK L. (1992) Ann. Nat.hist. Mus. Wien B 93: 259-275.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 121
1: O. tollmanni; top left, bottom right, bottom left: exterior view, top right: Interior view; middle lateral view; by R. von Cosel.
Mollusca, Gastropoda, Neritimorpha, Neritoidea, Phenacolepadidae
Species DistributionO. tollmanni BECK, 1992 Manus Back-Arc BasinOlgasolaris sp. North Fiji and Lau Back-Arc Basins
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Shinkailepas OKUTANI, SAITO & HASHIMOTO, 1989
1: S. briandi: exterior,rr interior and lateral views; by R. von Cosel & A. Le Goff.
Mollusca, Gastropoda, Neritimorpha, Neritoidea, Phenacolepadidae
Species DistributionS. briandi WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: Lucky Strike, Menez Gwen, LogatchevS. kaikatensis OKUTANI, SAITO & HASHIMOTO, 1989 Kaikata SeamountS. mojinensis SASAKI, OKUTANI & FUJIKURA, 2003 Ogasawara RidgeS. tufari BECK, 1992 Manus Back-Arc BasinShinkailepas sp. North Fiji Back-Arc BasinFurther undescribed species Mariana Back-Arc Basin, Indian Ocean, East Pacific Rise:
Size: Shell length up to 11 mm.
Color: Animal bright red infreshly dead and living specimens.
Morphology: Shells symmetrical, of limpet shape, depressed,with posteriorly inclined apex; sculptured by radiating ribs;shelf-like septum on the posterior inner side of shell. Small,vestigial operculum. Animal with large oral lobe, penis near
right cephalic tentacle; very small eyes present in larvae and atleast some species.
Biology: Genus endemic to vents. The reddish colour of freshanimals is probably caused by hemoglobin. Development withplanktotrophic larvae. Egg capsules common on shells and oth-er hard surfaces.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 122–123
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2: S. briandi: exterior, interior view;by P. Briand.
4: S. briandi: in situ; by P. Briand.3: S. briandi: eggs; by P. Briand.
References:
BECK L.A. (1992) Ann. Nat.hist. Mus. Wien B 93: 259-275.OKUTANI T., SAITO H. & J. HASHIMOTO (1989) Venus 48: 223-230 [224].SASAKI T., OKUTANI T. & K. FUJIKURA (2003) Veliger 46(3): 189-210 [201].WARÉNW A. & P.PP BOUCHET (2001) Veliger 44(2): 116-231 [174-177].
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Eosipho THIELE, 1929
Size: Shell length up to 70 mm.
Color: Chestnut brown to black.
Morphology: Large whelks with solid, smooth shell. Spire al-ways truncated (early whorls dissolved) in subadults and adults.Thick corneous operculum smaller than aperture.
Biology: A small radiation of buccinids living at vents; provi-sionally placed in the genus Eosipho, a genus known fromsunken drift wood and normal bathyal environments. A relat-ed species lives at Caribbean seeps. Buccinidae are carnivorousor scavengers and E. desbruyeresi has been collected in quanti-ty in baited traps.
1: E. desbruyeresi; top left to right: Abapertural, lateral and apertural view; bottom: Lateral views; by R. von Cosel & P. Lozouet.
Mollusca, Gastropoda, Caenogastropoda, Muricoidea, Buccinidae
Species DistributionE. auzendei WARÉN & BOUCHET, 2001 East Pacific Rise: 17-23°S; Pacific-Antarctic Ridge: 31-38°SE. desbruyeresi OKUTANI & OHTA, 1993 Mariana, North Fiji and Lau BasinsE. desbruyeresi nipponensis OKUTANI & FUJIWARA, 2000 Okinawa Trough, Ogasawara
P. BOUCHET Denisia 18 (2006): 124–125
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References:
OKUTANI T. & S. OHTA (1993) Venus 52: 217-221.OKUTANI T. & Y. FUJIWARA (2000) Venus 59: 123-128 [125-126].WARÉNW A. & P.PP BOUCHET (2001) Veliger 44(2): 116-231 [190-191].
2: E. auzendei, in vivo specimens;s by R. von Cosel & P. Lozouet.
3: E. auzendei in situ, with bythograeid crab,chiridotid holothurian and serpulid wormLaminatubus alvini from southern East PacificRise, cruise Biospeedo © Ifremer.
4: E. desbruyeresi in situ, among mytilid Bathymodiolus bre-vior from Lau Back-Arc Basin, cruise TUIM07; by courtesy ofC.R. Fisher.
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Speculator cariosus WARÉNWW & BOUCHET, 2001
Size: Shell height up to 8.3 mm.
Color: Brownish yellow.
Morphology: Shell tall and very slender, rather fragile, withhigh spire and small, rounded aperture with obliquely drawn-out siphonal canal. Whorls distinctly convex. Surface with fourstrong spiral cords and two more on the body whorl and withsomewhat variable axial ribs, about 30 on the body whorl, re-sulting in a reticulate sculpture. Uppermost whorls eroded. Op-erculum thin, paucispiral, with indistinct coiling and stronglyexcentric nucleus.
Biology: The only known specimen was collected together withtubeworms Ridgeia piscesae, but nothing is known about its diet.
Distribution: Northern Pacific, known only from ExplorerRidge: Magic Mountain (Steve 4 vent).
Reference:
WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 126
1: Apertural view; after WARÉNWW &BOUCHET (2001).
2: Enlargement of aperture; after WARÉNWW & BOUCHET (2001).
Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Cerithiopsidae
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Laeviphitus desbruyeresi WARÉNWW & BOUCHET, 2001
Size: Shell height up to 1.8 mm.
Morphology: Shell very small, not especially fragile, about fourwhorls, with shallow suture, narrow umbilical crevice and an-teriorly bluntly rounded and posteriorly slightly pointed aper-ture, smooth teleoconch and distinctly demarcated and cancel-late protoconch. Shell surface usually with strong ferrugineousdeposits. Operculum thin, yellowish brownish, paucispiral.
Biology: Among Bathymodiolus and in sediment. Genus knownfrom vents and driftwood (unpubl.). Development with plank-totrophic larvae.
Distribution: Mid-Atlantic Ridge: Menez Gwen to Rainbow.Genus also known from Japan (Laeviphitus japonicus OKUTANI,FUJIKURA & SASAKI, 1993) and the Marianas (unpubl.). Larvaecommon at the East Pacific Rise: 13°N, but no juvenile or adultspecimens have been found.
References:
OKUTANI T. FUJIKURA K. & T. SASAKI (1993) Bull. Natn. Sci. Mus., Tokyo A 19(4): 123-143.WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [182].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 127
1: The same specimen with and without ferrugineous layer; scale bar 0.5mm; by A. Warén.
2: Different views of larvae; by L. Mullineaux.
Mollusca, Gastropoda, Caenogastropoda, Elachisinidae
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Alviniconcha hessleri OKUTANI & OHTA, 1988
Size: Shell height up to 85 mm.
Color: Yellowish.
Morphology: Shell globose, rather elastic, spectacularly orna-mented with regularly spirally arranged periostracal hairs.Aperture with a shallow sinus in front. Operculum horny andovo-quadrate.
Biology: Genus endemic in vents. Usually in stacks aroundvent openings where they are exposed to warm (up to 13°C),sulphide-rich (up to 750 μM) water. Scattered specimens liveon side of chimneys and attain a very large size. The gill is hy-
pertrophied and specialized gill cells contain endosymbioticchemoautotrophic bacteria of the sulphur cycle. Reduced di-gestive tract as compared to related species of the same family.
Distribution: A single morphospecies in the Western Pacific:Mariana, North-Fiji and Lau Back-Arc Basins; Indian Ocean:Rodriguez Triple Junction, Kairei hydrothermal field. Never-theless, A. hessleri from the type location is genetically differentfrom the three Alviniconcha populations collected at otherplaces.
A. WARÉNWW Denisia 18 (2006): 128
Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae
1: Apertural view; by P.PP Briand © Ifremer.
2: Dorsal and ventral view; by P.PP Briand © Ifremer.
4: In situ population at Lau Back-Arc Basin; Biolau cruise © Ifremer.3: In situ specimen in the Lau Back-ArcBasin; Biolau cruise © Ifremer.
References:
DENIS F., JOLLIVET D. & D. MORAGA (1993) Biochem. Syst. Ecol. 21: 431-440.ENDOW K. & S. OHTA (1989) Bull. Jap. Soc. Microb. Ecol. 3: 73-82.HEALY J. (1992) Bull. Mus. Natl. Hist. Nat. 14: 273-291.KOJIMA S., FUJIKURA K., OKUTANITT T. & J. HASHIMOTO (2003) Venus 63: 65-68.KOJIMA S., OHTA S., FUJIWARA Y. & J. HASHIMOTO (1999) JAMSTEC J. Deep-Sea
Res. 14: 501-506.
OKUTANI T., HASHIMOTO J. & T. SASAKI (2004) Venus 63: 1-11.OKUTANI T. & S. OHTA (1988) Venus 47: 1-9.STEIN J.L., CARY S.C.,HESSLER R.R., OHTA S., VETTER R.D., CHILDRESS J.J. & H. FEL-
BECK (1988) Biol. Bull. 174: 373-378.WARÉNWW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.
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Desbruyeresia WARÉNWW & BOUCHET, 1993
Size: Shell height up to 10-12 mm.
Morphology: Shells regularly coiled, distinctly slender, withhigh spire and small, rounded aperture. Sculpture consisting ofaxial ribs, spiral cords, knobs and occasionally short spines. Tipoften corroded, shell often encrusted. Species differ in charac-
ters of the sculpture. Eyes reduced or absent. Right pallial ten-tacle absent.
Biology: Genus endemic in vents; detritus feeders; develop-ment unknown.
References:
OKUTANI T. & K. FUJIKURA (1990) Venus 49: 83-91.OKUTANI T., HASHIMOTO J. & T. SASAKI (2004) Venus 63: 1-11.WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [71-73].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 129
1 top row left: D. spinosa;middle: D. melanoides; right:D. cancellata, holotype coated forSEM; by R. von Cosel & A. Le Goff;bottom row left: D. spinosa;middle and right: D. provanna;by P.PP Briand.
Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae
Species DistributionD. cancellata WARÉN & BOUCHET, 1993 North Fiji and Lau Back-Arc Basins D. marianaensis (OKUTANI & FUJIKURA, 1990) Mariana Back-Arc BasinD. marisindica OKUTANI, HASHIMOTO & SASAKI, 2004 Central Indian RidgeD. melanioides WARÉN & BOUCHET, 1993 Manus and Lau Back-Arc BasinsD. spinosa WARÉN & BOUCHET, 1993 North Fiji Back-Arc BasinD. sp. aff. spinosa Mariana Back-Arc Basin
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Ifremeria nautilei BOUCHET & WARÉNWW , 1991
Synonym: Olgaconcha tufari BECK, 1991.
Size: Up to 95 mm.
Color: Brown (juvenile) to black (adult).
Morphology: Shell with about two whorls, umbilicus, nearlyoval aperture, and a conspicuous subsutural ramp; periostracumthickened, faintly glossy, dissolved at the apex. While the shellis growing at the aperture, the apex dissolves. Front and sides offoot conspicuously light blue in living specimens.
Biology: Genus with a single species endemic in vents, usuallyin massive heaps at the edge of Alviniconcha stacks. As in
Alviniconcha, the gill and circulatory system are hypertrophiedand the alimentary system is unexpectedly small. Two types ofbacterial symbionts are present in the gills, one dominant sul-phide-oxidizing bacteria and one in lower abundance, likelymethane-oxidizing. Different commensal polychaetes werefound inside the pallial cavity (scale worms) and the umbilicus(Amphisamytha cf. galapagensis). The female broods the larvaein a brood chamber in the foot (A. Warén, unpublished). De-velopment probably lecithotrophic.
Distribution: North Fiji, Lau, Manus Back-Arc Basins.
1: Adult specimens, apertural and apical view; juvenile specimens, apertural view; by P. Maestrati.
Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 130–131
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References:
BECK L.A. (1991) Ann. Nat.hist. Mus. Wien 92B: 277-287.BOROWSKI C., GIERE O., KRIEGER J., AMANN R. & N. DUBILIER (2002) Cah. Biol. Mar. 43: 321-324.BOUCHET P. & A. WARÉNWW (1991) C. R. Acad. Sci. Paris, Sér. III 312: 495-501.DESBRUYÈRES D., ALAYSEAA -DANET A.-M. & S. OHTATT (1994) Mar. Geol. 116: 227-242.GALCHENKO V.F., PIMENOV N.V., LEIN A.Y., GALKIN S.V., MILLER Y.M. & M.V. IVANOV (1992) Dokl. Biol. Sci. 323: 125-129.WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [64-71].WINDOFFER R. & O. GIERE (1997) Biol. Bull. 193: 381-392.
2 top: Living specimens in situ, inhabited by limpets Olgasolaris sp.; bottom: Left by P. Briand © Ifremer; right byBiolau cruise © Ifremer.
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Provanna DALL, 1918
1 top left: P. laevis; top right: P. variabilis; bottom left: P. ios; bottom right: P. buccinoides; by P. Briand.
Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae
Species DistributionP. buccinoides WARÉN & BOUCHET,TT 1993 North Fiji and Lau Back-Arc BasinsP.PP glabra OKUTANI, TSUCHIDA & FUJIKURA, 1992 Sagami Bay, Okinawa BasinP. ios WARÉN & BOUCHET, 1986 Galapagos, East Pacific Rise: 13-21°N, 17°S P. laevis WARÉN & PONDER, 1991 Guaymas BasinP. muricata WARÉN & BOUCHET, 1986 North Fiji and Lau Back-Arc Basin, Galapagos Sprea-
ding Center, East Pacific Rise: 21°NP. nassariaeformis OKUTANI, 1990 Mariana and Manus Back-Arc BasinsP. segonzaci WARÉN & PONDER, 1991 Lau Back-Arc BasinP. variabilis WARÉN & BOUCHET, 1986 Juan de Fuca Ridge, Oregon Margin
Size: Shell height up to 10-12 mm.
Color: Olive-brown to dark brown or greenish, often coveredby thick mineral deposits.
Morphology: Shells regularly coiled, moderately slender, withrather high spire and rounded aperture. Surface smooth or withsculpture consisting of axial ribs and spiral cords. Species differff
in characters of the sculpture. Tip often corroded, shells oftenencrusted. Eyes reduced or absent. Right pallial tentacle present.
Biology: Genus known from vents, seeps and sunken drift wood(only the vent species are listed herein). Detritus feeders; devel-opment without planktotrophic larvae in species with knownprotoconchs. One species from seeps has adelphophagy andhatches in the crawling stage (A. Warén, unpublished).
References:
OKUTANITT T. & K. FUJIKURA (1990) Venus 49: 83-91.OKUTANI T., TSUCHIDA S. & K. FUJUKURA (1992) Venus 51: 137-148.WARÉNW A. & P. Bouchet (1986) Zool. Scr. 15: 157-164.WARÉNW A. & P.PP BOUCHET (1989) Zool. Scr. 18: 67-102 [94-95].WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [74-76].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [178-179].WARÉNW A. & W.F. PONDER (1991) Zool. Scr. 20: 27-56.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 132
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1 left: Specimen 1, 2.6 mm, apertural view; right: Specimen 2, 2.3 mm; Menez Gwen; after WARÉNWW & BOUCHET (2001).
Alvania cf. stenolopha BOUCHET & WARÉNWW , 1993
Size: Shell height up to 2.6 mm.
Morphology: Shell small, conical, thin and fragile, with bluntspire and large aperture, outer lip not thickened. Whorls dis-tinctly and evenly convex, suture well-marked. Surface withvery fine but distinct spiral lines, 4-8 stronger spiral cords atand below the periphery and distinct, evenly spaced sharp andnarrow axial ribs, 20 on the last whorl, which end abruptly atthe first to third spiral cord. Protoconch dome-shaped, with 1.4whorls, diameter about 460 μm, teleoconch with 2.4 whorls.
Remark: It is not sure whether or not this species is a memberof the vent fauna; the species was described from a localityabout 150 km northeast of Menez Gwen.
Biology: Development lecithotrophic; each egg capsule con-tains a single juvenile. At vent sites encountered on sulphiderocks or at the base of black smokers, partly with Hydrozoa.
Distribution: Mid-Atlantic Ridge: Lucky Strike, Menez Gwenand surroundings.
References:
BOUCHET P. & A. WARÉNWW (1993) Boll. Malacol. Suppl. 3: 579-840.WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231.
A. WARÉNWW & P. BOUCHET Denisia 18 (2006): 133
Mollusca, Gastropoda, Caenogastropoda, Rissoidae
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1 top: Apertural view; bottom left: Basal view; bottom right: Apical view (note the inclined protoconch); from Menez Gwen;after WARÉNWW & BOUCHET (2001).
Neusas marshalli SYKES, 1925
Size: Shell diameter up to 2.06 mm.
Morphology: Shell planispiral, resembling a planorbid, withrounded and almost smooth whorls with a deep suture. Proto-conch tall-spired and obliquely inserted, with slightly morethan two whorls, smooth. Teleoconch with about three slightlyirregularly coiled whorls. Operculum corneous, multispiral,round with central nucleus.
Remarks: The species was also collected from a non-hy-drothermal locality off Portugal, 39°42’N, 09°43’W, 1092-1993m (type locality). Two other species of the genus are known;they are confined to non-vent localities off New Caledonia andoff New Zealand.
Biology: No data. Apparently the species is not obligatorilyconfined to vents.
Distribution: Mid-Atlantic Ridge: Menez Gwen.
Reference:
WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231.
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 134
Mollusca, Gastropoda, Caenogastropoda, Vitrinellidae
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Hyalogyrina MARSHALL, 1988
1: H. globularis,apertural and dor-sal view of a speci-men; Juan de FucaRidge, EndeavourSegment, ClamBed; by A. Warén.
2: H. globularis,critical pointdried specimen;front view ofhead-foot, shelland pallial skirtremoved; Juande Fuca Ridge,Endeavour Seg-ment, ClamBed; by A.Warén.
3: H. umbellifera,early protoconch;A. Warén.
Mollusca, Gastropoda, Heterobranchia, Hyalogyrinidae
Species DistributionH. globularis WARÉN & BOUCHET, 2001 Juan de Fuca Ridge: Endeavour SegmentH. grasslei WARÉN & BOUCHET, 1993 Guaymas Basin
Size: Maximum height ca 3.4 mm.
Morphology: Shell small, rather fragile, depressed, globular ormore tall-spired, 2.5-4 whorls on the teleoconch, vitreous andfragile, with deep suture, deep umbilicus, aperture rounded. Ini-tial part of protoconch sculptured by small, crowded pits, latersmooth, teleoconch smooth. Operculum round, transparentand colorless to brown, multispiral with distinct growth linesand central nucleus. Animal with thick tentacles and almost
cylindrical snout. Additional tentacles behind the normalcephalic ones of variable development, possibly changing withage. Foot large, broad and flat, rounded or truncated posterior-ly and shallowly bilobed anteriorly, no propodium distinguish-able.
Biology: Epifaunal grazers. Genus known from vents, seeps,whalebone and driftwood. Development unknown.
References:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [49-52].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [200-207].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 135
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Lurifax vitreus WARÉNWW & BOUCHET, 2001
References:
SASAKI T. & T. OKUTANI (2005) Venus 63: 121-124. WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [207-208].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 136
1 top from left to right: Apertural, apical and ventral view of the same specimen;by R. von Cosel & A. Le Goff; bottom: Three specimens by A. Warén.
2 left and right: Specimens (SEM); by A. Warén; middle: Living specimen; by P.PP Briand @ Ifremer.
Mollusca, Gastropoda, Heterobranchia, Orbitestellidae
Size: Diameter up to 2.8 mm.
Morphology: Shell small, rather fragile, depressed conical, fivewhorls, vitreous, with shallow suture, broad and deep umbili-cus. Protoconch smooth, teleoconch with spiral striae andstronger ribs and radiating flexuous incremental lines. Surfaceoften covered by thick crusts of rust. Operculum stiff, almosttransparent, multispiral with six whorls and central nucleus.Animal with well developed eyes at the dorsal base of simple
cylindrical tentacles. Foot with well-demarcated propodium, noappendages except metapodial lobes.
Biology: Epifaunal. Genus known from vents and seeps. Devel-opment unknown.
Distribution: Mid-Atlantic Ridge: Menez Gwen and LuckyStrike. Genus with three species endemic to vents and seeps.One at Lau and Fiji Back-Arc Basins. Lurifax japonicus fromSumisu Caldera, Southern Japan.
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Xylodiscula analoga WARÉNWW & BOUCHET, 2001
Size: Diameter up to 3 mm.
Morphology: Shell small, fragile, subplanispiral, 2.5 whorls(teleconch), with rather deep suture, very wide and deep um-bilicus and subradial and slightly prosocline aperture. Proto-conch always corroded, teleoconch with incremental lines,otherwise smooth. Surface with a thick, yellowish-brownish pe-riostracum. Operculum thin and transparent, round, smooth,multispiral with central nucleus.
Biology: Found among Bathymodiolus and on sediment. Genusknown from vents, seeps and biogenic substrates. Developmentunknown.
Distribution: Mid-Atlantic Ridge: Menez Gwen and LuckyStrike. Another species, Xylodiscula major WARÉNWW & BOUCHET,1993 was collected from North Fiji Back-Arc Basin.
References:
WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [35].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [208, 210].
A. WARÉNWW , P. BOUCHET & R. VON COSEL Denisia 18 (2006): 137
1: Two specimens: left dorsal view, right ventral view; by A. Warén.
Mollusca, Gastropoda, Heterobranchia, Xylodisculidae
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Phymorhynchus DALL, 1908
1 top row, left to right: Twospecimens of P. carinatus; onespecimen of P. moskalevi; by A.Warén (SNHM) & P. Briand(Ifremer); bottom row: Threespecimens of P.PP ovatus, fromMid-Atlantic Ridge; by R. vonCosel & A. Le Goff (MNHN).
Mollusca, Gastropoda, Prosobranchia, Conoidea, Turridae
Species DistributionP.PP carinatus WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: 23-15°NP. hyfifluxi BECK, 1996 North Fiji BasinP. major WARÉN & BOUCHET, 2001 East Pacific Rise: 13-9°NP.PP moskalevi SYSOEV & KANTOR, 1995 Mid-Atlantic Ridge: 26-23°NP. ovatus WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: 37-15°NP. starmeri OKUTANI & OHTA, 1993 North Fiji and Manus Back-Arc Basins; may be
common in both vent and non-vent areas in up-per abyssal bottom
P.PP wareni SYSOEV & KANTOR, 1995 Edison Seamount, Lihir Is, West Pacific
Size: Shell height to 72 mm.
Morphology: Shells short, obese fusiform or bucciniform. Shellsurface white with strong spiral ribs (about 8-10 on penultimatewhorl and some 25-30 on body whorl including base) whichoverlie weak growth lines. Spire roundly conical, but bodywhorl well inflated, occupying about 50% of shell length. Aper-ture wide, lunate, with crenulated outer lip corresponding tospiral ribs. Columellar lip almost straight. Siphonal canal open,not twisted. No operculum present. Head with big rhyn-
chodaeum. Toxoglossate radula teeth hollow, needle-like inshape with basal expansion. Distal tip sharp, monocuspidatewith a slit.
Biology: Often observed at the periphery of dying vents, prob-ably predator on molluscs and scavenger. Genus known fromseeps and vents as well as free-living. Larval developmentplanktotrophic.
T. OKUTANI Denisia 18 (2006): 138–139
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2 left to right: One specimen of P. major from East Pacific Rise 13°N, and two specimens of P.starmeri from North Fiji Back-Arc Basin; by P. Briand (Ifremer).
References:
BECK L. (1996) Arch. Moll. 126(1/2): 109-115.BOUCHET P. & A. WARÉNWW (1986) Mem. Mus. Natl. Hist. Nat., Paris, Sér. A, Zool. 133: 457-571.OKUTANI T. & S. OHTA (1993) Venus 52(3): 217-221.SYSOEV A.V. & Y.I. KANTOR (1995) Ruthenica 5: 17-26.WARÉNW A. & P.PP BOUCHET (2001) Veliger 44(2): 116-231 [192-199].
3: Phymorhynchus sp. from East PacificRise: 18°S, cruise Biospeedo © Ifremer;middle: Phymorhynchus sp. from Juan deFuca Ridge; by K. Juniper; bottom: Larvafrom Juan de Fuca Ridge; by K. Juniper &L. Mullineaux.
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Dendronotus comteti VALDÈSVV & BOUCHET, 1998
Reference:
VALDÈSVV A. & P. BOUCHET (1998) Deep-Sea Res. II 45: 319-327.
P. BOUCHET & M. SEGONZAC Denisia 18 (2006): 140
1A: Dorsal view of the holotype (SEM); scale bar 1 mm; B: Details of the anterior region ofthe same specimen (SEM); scale bar 100 μm; C: Jaws of a paratype, arrow indicates masti-catory border; scale bar 100 μm; cruise Diva 2, Ifremer; from VALDÈSVV & BOUCHET (1998).
Mollusca, Gastropoda, Nudibranchia, Dendronotidae
Size: Up to 5 mm.
Color: Unknown in life. Preserved specimens: uniformly palecream background color, without traces of colored spots orlines.
Morphology: The frontal velum has four short papillae, two oneach side of the body midline. Medial pair longer and alwaysbranched, but outer two only branched in the larger specimens.2-4 pairs of cerata on each side of the dorsum. When branched,the cerata form three conical processes. Rhiniphores have 7-8
lamellae. The margin of the rhinophoral sheath has four un-branched tentacular papillae of even size.
Biology: First species of nudibranch recorded with certaintyfrom a vent site. Living among mussel bed of Bathymodiolusazoricus, but probably not restricted to this environment. Its oc-currence at vents can be explained on the one hand by thepresence of prey such as hydrozoa (Candelabrum phrygium), andon the other hand by the absence of predators.
Distribution: Mid-Atlantic Ridge: Lucky Strike, Eiffel Tower.
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Mollusca, Bivalvia
The Solemyidae are represented at hydrothermal vent orcold seep biotopes by the genus Acharax. Eight named speciesas well as several still unnamed species are known, but only asingle species has been collected at hydrothermal vents (LauBack-Arc Basin). Representatives of this genus can grow to rel-atively large sizes between 10 and 22 cm. They live deeplyburied in soft sediment. At least some species are characterizedby the absence of a digestive tract, and their nourishment reliesexclusively on their chemosynthetic symbiotic bacteria.
Of the large mussels living at hydrothermal vents or coldseeps, 20 species have been currently described, 18 of them inthe genus Bathymodiolus. Of these, 10 are known from hy-drothermal vents and are covered herein. Several other largemussels are still awaiting description. All known Bathymodiolushave a larval shell of about 0.5 mm or smaller with set-off pro-toconch 1; in general, the protoconch has a distinct rose color.
The Vesicomyidae are a rather diverse family in size, shapeand species number: at the moment 84 recently named speciesare known, but there are still numerous undescribed speciescurrently under study by several authors. The family compriseson one hand small species (genus Vesicomya sensu stricto),which occur in the deep sea but are not necessarily associatedwith cold seep and/or hot vent habitats and on the other hand,most of the numerous medium-sized or large to very largespecies are confined to reducing sediments, cold seeps, hydro-
carbon seeps or hydrothermal vents. In the past, species of Vesi-comyidae have been assigned to different genera or subgeneraand many of the large species are now commonly placed in thegenus Calyptogena (sensu lato). Recently however, some au-thors have revised tentatively all Vesicomyidae in the solegenus Vesicomya (in a broad sense) pending future supra-specif-ic revisions based mainly on molecular research. Anyway, shelland soft part morphology remain important and key charactersare among others hinge dentition, shell size and shape and pres-ence or absence of a well-developed pallial sinus. Herein, thegenus Calyptogena is maintained. The assignment of a vesi-comyid species to Calyptogena differs with authors, and thesupra-specific systematics of the Vesicomyidae is still far fromsettled. Most of the larger species have been described fromcold seep biotopes, but of the 28 named species currently treat-ed as Calyptogena, seven were found inhabiting also (or exclu-sively) hydrothermal vents. Not very much is known of the lar-val development of vesicomyid clams, but many are known tohave lecithotrophic development.
The Pectinidae are represented by two species, Bathypectenvulcani, present at the Galapagos Spreading Center, the north-ern East Pacific Rise 13°N and the southern East Pacific Rise38°S and Sinepecten segonzaci, from the Manus Back-arc Basin.More sampling need of these small bivalves, difficult to see insitu, to understand the processes of their distribution.
R. VON COSEL Denisia 18 (2006): 141
1: Bathymodiolus azoricus from Rainbow, Mid-Atlantic Ridge, Atos cruise © Ifremer.
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Calyptogena edisonensis OKUTANI, KOJIMA & KIM, 2003
Reference:
OKUTANI T.K., KOJIMA S. & D. KIM (2004) Venus 63: 29-32.
R. VON COSEL Denisia 18 (2006): 142
1: Several specimens; by R. von Cosel.
Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae
Size: Shell length up to 100 mm.
Morphology: Shell rather large, moderately thick, elongate-oval, dorsal and ventral margin almost parallel in their middlepart. Umbones not prominent, prosogyrous, situated anteriorlyat 1/4 to 1/5 of shell length. Surface with irregular commargin-al growth lines, periostracum thin, yellowish-straw-coloured,on earlier parts of the valves more or less eroded. Hinge platestrong but not very broad, typical for Calyptogena, with threecardinals in each valve.
Biology: At hydrothermal vents.
Distribution: Western Pacific: Edison Seamount.
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Calyptogena extenta KRYLOVA & MOSKALEV, 1996
References:
COAN E.V., SCOTT P.V. & F.R. BERNARD (2000) in COAN E.V., SCOTT P.V. & F.R. BERNARD (Eds.) Bivalve Seashells of Western North America: 336-343.KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406 [400]. KRYLOVA E. & L.I. MOSKALEV (1996) Ruthenica 6: 1-10.
R. VON COSEL Denisia 18 (2006): 143
1: Habitus; by R. von Cosel.
Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae
Size: Shell length up to 246 mm.
Morphology: Shell very large and very elongate, thick, slightlyinequivalve, somewhat irregular, bean-shaped, curved withmarkedly concave ventral margin and somewhat convex dorsalmargin, gaping anteriorly and posteriorly. Umbones not promi-nent, prosogyrous, situated anteriorly at 1/6 of shell length.Surface with low growth ridges, periostracum brownish-olive,persistent only posteriorly and near margins. Hinge plate rathernarrow but strong, with three cardinals in the left valve andtwo cardinals in the right valve.
Biology: At cold seeps and hot vents. The specimens liveburied in the sediment but with the posterior part (2/3 of shelllength) free, they are inclined at about 50° to the sediment sur-face and are capable of moving around by means of their largemuscular foot, leaving short tracks in the sediment.
Distribution: Gorda Ridge (hydrothermal vents); MontereyCanyon: 36°35’N, 122°30’5’’W, 3041 m (cold seeps).
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Calyptogena gigas DALL, 1896
Size: Shell length up to 125 mm.
Morphology: Shell large, regularly oval-oblong, rather thin-shelled, very inflated, with broadly rounded anterior and poste-rior margin. Ventral margin almost straight, dorsal marginslightly convex. Umbones not prominent, prosogyrous. Surfacewith fine irregular growth lines, sometimes more or less erodedin the umbonal region. Periostracum pale olive to dark brown,often eroded on the earlier parts of the valves. Inside with veryshort and broad pallial sinus. Hinge plate short and ratherstrong, with three cardinals in the left valve and three cardinalsin the right valve.
Biology: At hot vents and cold seeps, collected between 550and 2610 m.
Distribution: Gulf of California, 1567 m (type locality); Guay-mas Basin; Juan de Fuca Ridge.
References:
COAN E.V., SCOTT P.V. & F.R. BERNARD (2000) in COAN E.V., SCOTT P.V. & F.R. BERNARD (Eds.) Bivalve Seashells of Western North America: 336-343.DALL W.H. (1896) Proc. U.S. Natl. Mus. 18(1034): 7-20 [18-19].KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406 [400].
R. VON COSEL Denisia 18 (2006): 144
2: In situ; by courtesy of K. Juniper.
Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae
1: Habitus; by R. von Cosel.
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Calyptogena magnifica BOSS & TURNER, 1980
Size: Shell length up to 263 mm.
Morphology: Shell very large, variable, oval to elongate-oval inoutline, inequilateral and equivalve, usually without gape;valves white, entirely aragonitic; periostracum present, but fre-quently eroded away on older portions of shell; umbos prosogy-rous, sometimes partially enrolled; lunule and escutcheon vari-able, present or absent; ligament external, opisthodetic, andparivincular; foot strong and rugose, with byssal gland.
Biology: At hot vents only. The life habit of the adult is semi-epibiotic on bare basalt, living in cracks and crevices fromwhich hydrothermal fluids emanate; early life history stages are
often encountered living within basaltic rubble associated withsuch warm cracks and crevices. The presence of this species iscorrelated with elevated levels of hydrogen sulphide. The softtissue is coloured dark red when retrieved living due to thepresence of intracellular hemoglobin. The thick and large gillscontain sulfur oxidizing chemoautotrophic symbionts. Thelarge rugose foot is often seen protruding when the clams areviewed in life position.
Distribution: Entire Northern East Pacific Rise and SouthernEast Pacific Rise: 21°N to 22°S; Galapagos Spreading Center.
1: In situ, among barnacle bed ofVulcanolepas n. sp. and chiridotidholothurian; southern East Pacific Rise:17°S, Biospeedo cruise © Ifremer.
2: In situ; by courtesy of K. Smith Jr.
Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae
3: Exterior, interior and dorsal view of leftvalve, specimen from 21°N; by R. von Cosel & P. Lozouet.
ARP A. J., CHILDRESS J.J. & C.J. FISHER (1984) Physiol. l Zool. 57: 648-662.BOSS K. & R. TURNER (1980) Malacologia 20: 161-194.CHILDRESS J.J., FISHER C.R., FAVUZZI J.A. & N.K. SANDERS (1991) Physiol. Zool.
64: 1444-1470.FATTON E. & M. ROUX (1981) C. R. Acad. Sc. Paris, Série III 293: 63-68.FIALA-MÉDIONI A. & C. METIVIER (1986) Mar. Biol. 90: 215-222.FISHER C. et al. (1988) Deep-Sea Res. 35: 1811-1831.
HURTADO L.A., MATEOS M., LUTZ R.A. & R.C. VRIJENHOEK (2003) Appl. Envi-ronm. Microbiol. 69(4): 2058-2064.
KENNISH M. & R. LUTZ (1992) Rev. Aquat. Sci. 6: 29-66.KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406
[400].LUTZ R.A., FRITZ L.W. & R.M. CERRATO (1988) Deep-Sea Res. I 35: 1793-1810.RIO M. & M. ROUX (1984) C. R.Acad Sci. Paris, Série II 299: 167-172.VRIJENHOEK R. et al. (1994) Deep-Sea Res. 41: 1171-1189.
R. LUTZ & R.C. VRIJENHOEK Denisia 18 (2006): 145
References:
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Calyptogena nankaiensis OKUTANI, KOJIMA & ASHI, 1996
References:
KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406 [400].OKUTANI T., KOJIMA S. & J. ASHI (1996) Venus 55: 257-263.
R. VON COSEL Denisia 18 (2006): 146
1: Habitus of a specimen; by R. von Cosel.
Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae
Size: Shell length up to 191 mm.
Morphology: Shell very large, solid, elongate-oval, with slight-ly concave middle part of ventral margin and with posteriorpart slightly higher than anterior part. Anterior margin nar-rowly rounded, posterior margin broadly rounded. Umbonesprosogyrous, almost subterminal, situated anteriorly at about1/7 of shell length. Surface with fine, irregular, close-set growthlines. Periostracum thick, straw-coloured, sometimes more orless eroded. Hinge plate strong, with three cardinals in rightand left valve.
Biology: At cold seeps and hot vents, on seep sites sometimesco-occurring with C. soyoae OKUTANI, 1957.
Distribution: Nankai Trough (cold seeps); Okinawa Trough,North Iheya Knoll (hot vents).
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Calyptogena okutanii KOJIMA & OHTA, 1997
Size: Shell length up to 120 mm.
Morphology: Shell large, thick and solid, elongate-oval in out-line, with slightly concave middle part of ventral margin. Um-bones low, prosogyrous. Surface with irregular, close-set com-marginal growth ridges and lines. Periostracum thin and brown-ish, but often more or less erodedon the earlier parts of thevalves. Hinge plate strong and rather broad with two strongcardinals in both valves.
Remarks: Calyptogena okutanii and C. soyoae are sibling specieswith slight but constant morphological and molecular differ-ences
Biology: At cold seeps and hot vents, at some seep sites co-oc-curring with C. soyoae OKUTANI, 1957.
Distribution: Sagami Bay and Nankai Trough (cold seeps); Ok-inawa Trough, Iheya Ridge (hydrothermal vents).
References:
KOJIMA S. & S. OHTA (1997) Venus 56: 189-195.KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406 [400].
R. VON COSEL Denisia 18 (2006): 147
1: Habitus; by R. von Cosel.
Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae
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Calyptogena solidissima OKUTANI, HASHIMOTO & FUJIKURA, 1992
Size: Shell length up to 128.5 mm.
Morphology: Shell large, thick and robust, oblong-oval in out-line; with ventral margin slightly concave in its middle part.Umbones prosogyrous, not prominent. Surface with irregulargrowth lines and very fine and densely spaced radial threads,visible under a lens only. Periostracum extremely thin and dullyellowish, but generally eroded and persistent only near themargins. Hinge plate strong, with three cardinals in each valve,sub-umbonal pit present.
Biology: At hot vents and cold seeps. Living clams bury them-selves about one half to two-thirds of the shell length into thesediment. A temperature anomaly of 0.3°C was recorded 30 cmbelow the white-stained bottom surface. The thick and largegills contain sulphur oxidizing chemoautotrophic endosym-bionts.
Distribution: Mid-Okinawa Trough: Minami-Ensei Knoll.
References:
HASHIMOTO J., OTHA J., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. 42(4): 577-598.KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406 [400].OKUTANI T., HASHIMOTO J. & K. FUJIKURA (1992) Venus 51(4): 225-233.
J. HASHIMOTO & R. VON COSEL Denisia 18 (2006): 148
1: Habitus of two specimens; by R. von Cosel.
Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae
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Acharax alinae MÉTIVIER & COSEL, 1993
Size: Shell length up to 135 mm (including periostracum).
Morphology: Shell elongate-oval. Dorsal margin straight. An-terior and posterior margin rounded. Ventral margin nearlystraight or somewhat concave. Umbos at posterior third, broadand flattened, beaks eroded. Periostracum very strong, darkbrown, extending far beyond the calcified part of the valve.Animal with very large gills and a voluminous, roughly cylin-drical foot which distally ends in a pedal disk. Digestive tractabsent.
Biology: In reducing sediments; found buried 20-30 cm deep inpale-coloured coarse sediment at the base of an isolated large si-boglinid tube in a seep area at the edge of a hydrothermal vent.Known from vents only. Gills very likely harbouring chemosyn-thetic symbiosis. Lecitrotrophic development, most probablyno, or extremely short, free swimming larval phase. Larval shell(no separate protoconch I & II) 1.35 mm long and 0.68 mmhigh.
Distribution: Lau Basin: Valu Fa Ridge, Hine Hina.
Reference:
MÉTIVIER B. & R. VON COSEL (1993) C. R. Acad. Sci. Paris, Sér. III 316: 229-237.
D. DESBRUYÈRES Denisia 18 (2006): 149
1: Holotype, exterior and interior of both valves; by P. Maestrati © MNHN; bottom: Another specimen with soft parts; by P. Briand © Ifremer.
Mollusca, Bivalvia, Protobranchia, Solemyoida, Solemyidae
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Bathymodiolus aduloides HASHIMOTO & OKUTANI, 1994
Size: Shell length up to 96 mm.
Morphology: Shell large, rather thick and solid. Umbones sub-terminal, at about one-tenth of the shell length. Ligamentstrong. Extremely thick and large ctenidia with long demi-branchs. No extreme mantle fusion, valvular siphonal mem-brane short. Anterior adductor muscle scar located in front ofthe umbo. Posterior adductor muscle scar rounded trapezoid.Gut with a single clockwise loop.
Biology: This species is found at both hydrothermal vent andcold seeps. The specimens are found with their anterior endthrust into diffused vents and attached with byssus to outcropsclose to the vent openings. Periostracum colour is variable ac-cording to habitat. Development with planktotrophic larvae.
Distribution: Central Japan: Sagami Bay; Okinawa Trough:Minami-Ensei Knoll and Iheya Ridge, 710 m to 1389 m.
References:
HASHIMOTO J. & T. OKUTANI (1994) Venus 53 (2): 61-83.HASHIMOTO J., OHTA S., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. I 42(4): 577-598.
J. HASHIMOTO Denisia 18 (2006): 150
1: Paratype MNHN Paris, interior of right valve, exterior of both valves; by R. von Cosel & P. Lozouet © MNHN.
2: In situ © JAMSTEC.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
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Bathymodiolus azoricus COSEL & COMTET, 1998
Size: Shell length up to 121 mm.
Morphology: Shell variable, more or less elongate-modio-liform, beaks subterminal but close to the anterior margin. Ven-tral margin straight to more or less concave. Postero-dorsalmargin slightly to markedly convex, occasionally straight. Lig-ament plate slightly arched. Exterior with dense irregulargrowth lines and growth waves. Periostracum dull, warm chest-nut brown, in umbonal region and often also postero-dorsallylighter brown. Anterior byssus retractor scar in the umbonalcavity, under the beaks. Anterior part of posterior byssus re-tractor muscle scar under ligament’s end or slightly forward.Animal with large gills. Mantle lobes on anterior half of ven-tral side separate. Valvular siphonal membrane short, narrowand rather strong.
Biology: In dense clusters byssally attached to hard substratearound the hydrothermal vent, mostly on the walls and flangesof active edifices, at temperatures from 6° to up to 30°C. En-demic to vents. Development with long planktonic larvalphase; protoconch is 0.5 mm long.
Distribution: Mid-Atlantic Ridge: Menez Gwen, Lucky Strike,Rainbow (hybrids between B. azoricus and B. puteoserpentiswere observed at Broken Spur vent field).
1: Holotype MNHN, exterior and interior of both valves; by R. von Cosel & P. Lozouet © MNHN.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
R. VON COSEL Denisia 18 (2006): 151–152
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2: Post-larvae fixed on gastropod probably Protolyra thorvaldssoni (det. A. Warén); byP. Briand © Ifremer.
3: In situ at Menez-Gwen vent field, withlimpets Lepetodrilus atlanticus on theshells; Atos cruise © Ifremer.
4: In situ at Rainbow vent field with strongsulphide deposit; Atos cruise © Ifremer.
References:
COMTET T., LE PENNEC M. & D. DESBRUYÈRES (1999) J. Mar. Biol. Ass. U.K. 79: 1149-1150.COMTET T., JOLLIVET D., KHRIPOUNOFF A., SEGONZAC M. & D.R. DIXON (2000) Limnol. Oceanogr. 45: 1655-1661.COSEL R. VON, MÉTIVIER B. & J. HASHIMOTO (1994) Veliger 42(3): 218-248 [220-231].FIALA-MÉDIONI A., MCKINESS Z., DANDO P., BOULEGUE J., MARIOTTI A., ALAYSE-DANET J., ROBINSON J. & C. CAVANAUGH (2002) Mar. Biol. 141: 1035-1043.O’MULLAN G.D., MAAS P.A.Y., LUTZ R.A. & R.C. VRIJENHOEK (2001) Mol. Ecol. 10: 2819-2831.
152
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Bathymodiolus brevior COSEL, MÉTIVIER & HASHIMOTO, 1994
Reference:
COSEL R. VON, METIVIER B. & J. HASHIMOTO (1994) Veliger 37(4): 374-392 [375-380].
R. VON COSEL Denisia 18 (2006): 153
1: Holotype MNHN, exterior of both valves, interiorof left valve, dorsal view; by P. Maestrati © MNHN.
2: Other specimen, left valve; by P. Briand. 3: In situ view of mussel bed, Lau Back-Arc Basin; Biolaucruise © Ifremer.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
Size: Shell length up to 143 mm.
Morphology: Shell oval-wedge shaped, stout, beaks subtermi-nal. Ventral margin straight to more or less concave. Postero-dorsal margin slightly convex to almost straight. Ligament plateslightly arched. Exterior smooth, with irregular growth lines.Periostracum dull, dark brown, in umbonal region lighterbrown. Anterior retractor scar at the anterior part of the um-bonal cavity. Anterior part of posterior byssus retractor musclescar at 2/3 of the ligament. Animal with large gills. Mantlelobes on anterior half of ventral side separate. Valvular siphonalmembrane short, narrow and rather strong.
Biology: In dense clusters byssally attached to hard bottomaround the hydrothermal vents at temperatures up to 18°C. En-demic to vents. Development with long planktonic larvalphase; protoconch is 0.4 mm long.
Distribution: North Fiji and Lau Back-Arc Basins.
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Bathymodiolus elongatus COSEL, MÉTIVIER & HASHIMOTO, 1994
1: Several specimens, exterior, interior, dorsal view; bottom: Juveniles, exterior and interior of right valve; fromNorth Fiji Back-Arc Basin, Mussel Valley site; cruise Starmer 2; by P. Maestrati © MNHN & P. Briand © Ifremer.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
Size: Shell length up to 155 mm.
Morphology: Shell elongate-wedge shaped, slender, beaks wellsubterminal. Ventral margin straight or slightly convex. Liga-ment plate somewhat arched to straight. Exterior with irregulargrowth lines and narrow to broad irregular, concentric grooves,mostly on the ventral part. Periostracum glossy, light chestnutbrown, in umbonal region lighter brown. Anterior retractorscar at the anterior part of the umbonal cavity. Anterior part ofposterior byssus retractor muscle scar at 2/3 of the ligament.Animal with large gills. Mantle lobes on anterior half of ven-tral side separate. Valvular siphonal membrane short.
Biology: Byssally attached to lava around diffuse vents, withabsence of massive hydrothermal deposits and vent fluid tem-perature not exceeding 8.5°C. Endemic to vents. Developmentnot known but most probably with planktonic larval phase.
Distribution: North Fiji Back-Arc Basin.
Reference:
COSEL R. VON, METIVIER B. & J. HASHIMOTO (1994) Veliger 37(4): 374-392 [380-386].
R. VON COSEL Denisia 18 (2006): 154
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Bathymodiolus japonicus HASHIMOTO & OKUTANI, 1994
Size: Shell length up to 107 mm.
Morphology: Shell rather thick, stout, moderately tumid. Um-bones subterminal, at about 5% anterior of the shell length.Posterior end of ligament abrupt. Extremely thick and largectenidia with long demibranchs. No extreme mantle fusion,valvular siphonal membrane short. Anterior byssus retractormuscle scar located in the anterior part of the umbonal cavity.Pallial line smooth, concave in adult, but slightly concave injuvenile and sub-adult.
Biology: This species is found at both hydrothermal vents andcold seeps. It is the most abundant and conspicuous organism ofthe hydrothermal vent community on the Minami-Ensei Knoll.Development not known but most probably with planktoniclarval phase.
Distribution: Off Hatsushima site and Sagami Bay, centralHonshu, 1170 m. Mid-Okinawa Trough: Minami-Ensei Knolland Iheya Ridge.
References:
HASHIMOTO J. & T. OKUTANI (1994) Venus 53(2): 61-83.HASHIMOTO J. HASHIMOTO J., OHTA S., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. 42(4): 577-598.
J. HASHIMOTO Denisia 18 (2006): 155
2: In situ view of a mussel bed © JAMSTEC.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
1: Paratype MNHN Paris, exterior of both valves, interiorof right valve; by R. von Cosel & P. Lozouet © MNHN.
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Bathymodiolus marisindicus HASHIMOTO, 2001
Size: Shell length up to 86 mm.
Morphology: Shell oval-modioliform, stout, beaks subterminal.Ventral margin almost straight to slightly concave. Postero-dor-sal margin somewhat convex. Ligament plate slightly arched.Exterior with dense fine commarginal lines. Periostracumsmooth and glossy, strong, dark brown to blackish brown, inyoung specimens chestnut brown. Anterior retractor scar at an-terior extremity of the umbonal cavity. Anterior part of poste-rior byssus retractor muscle scar at 2/3 the ligament length. An-imal with large gills. Inner mantle folds separate along thewhole ventral margin. Valvular siphonal membrane short, nar-row and rather strong.
Biology: In dense beds byssally attached to hard bottom alongshimmering crevices near black smokers. Endemic to vents. De-velopment with long planktonic larval phase.
Distribution: Indian Ocean: Rodriguez Triple Junction, Kaireihydrothermal field.
References:
HASHIMOTO J. (2001) Venus 60(3): 141-149.HASHIMOTO J., OTHA S., GAMO T., CHIBA H., YAMAGUCHI T., TSUCHIDA S., OKUDAIRA T., WATABE H., YAMANAKA T. & M. KITAZAWA (2001) Zool. Sci. 18(5): 717-721.
R. VON COSEL Denisia 18 (2006): 156
1: Some specimens, exterior and interior of valves; by R. von Cosel & P. Lozouet © MNHN.
2: In situ view of a smallmussel bed, among numeroussea anemones Mariactis cf.bythios, alvinocaridid shrimpsRimicaris kairei and gastro-pod Phymorhynchus sp.© JAMSTEC.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
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Bathymodiolus platifrons HASHIMOTO & OKUTANI, 1994
Size: Shell length up to 115.6 mm.
Morphology: Shell rather thick and solid. Umbones often near-ly subterminal, but mostly on the same plane with anterior endof the shell. No extreme mantle fusion, valvular siphonal mem-brane short. Extremely thick and large ctenidia with long demi-branchs.
Biology: This species is found at both hydrothermal vents andcold seeps. In Sagami Bay, B. japonicus is occasionally sympatricin the same habitat. Periostracum colour variable according tohabitat. Development not known but most probably withplanktonic larval phase.
Distribution: Okinawa Bank and off Hatsushima site in Saga-mi Bay, central Honshu, 1180 m; Mid-Okinawa Trough: IheyaRidge and Izena Caldron.
Reference:
HASHIMOTO J. & T. OKUTANI (1994) Venus 53(2): 61-83.
J. HASHIMOTO Denisia 18 (2006): 157
2: In situ view of a mussel bed © JAMSTEC.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
1: Paratype MNHN Paris, exterior and interior of right valve; by R. von Cosel & P. Lozouet © MNHN.
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Bathymodiolus puteoserpentis COSEL, MÉTIVIER & HASHIMOTO, 1994
Size: Shell length up to 141 mm.
Morphology: Shell oval-wedge shaped, rather stout, beaks sub-terminal. Ventral margin straight to very weakly convex. Pos-tero-dorsal margin markedly convex. Ligament plate slightlyarched in anterior part, straighter in posterior part. Exteriorsmooth, with pronounced irregular growth lines. Periostracumdark brown and rather glossy. Anterior byssus retractor scar onanterior part of the umbonal cavity, in front of the beaks. An-terior part of posterior byssus retractor muscle scar under the
posterior third of the ligament, near the end. Animal with largegills. Mantle lobes on anterior half of ventral side separate.Valvular siphonal membrane short.
Biology: Bysally attached to sulfur blocks immediately arounddiffuse venting of water. Endemic to vents. Development un-known, most probably with long planktonic larval phase.
Distribution: Mid-Atlantic Ridge: TAG, Snake Pit and Lo-gatchev.
References:
CAVANAUGH C. M., WIRSEN C.O. & H.W. JANNASCH (1992) Appl. Environ. Microbiol. 58: 3799-3803.COSEL R. VON, METIVIER B. & J. HASHIMOTO (1994) Veliger 37(4): 374-392 [387-389].O’MULLAN G.D., MAAS P.A.Y., LUTZ R.A. & R.C. VRIJENHOEK (2001) Mol. Ecol. 10: 2819-2831.
R. VON COSEL Denisia 18 (2006): 158
1 top: Holotype MNHN, exterior of both valves; mid-dle: Other specimen, interior and exterior of left valve;bottom: Interior of left valve of holotype; by R. vonCosel & P. Lozouet © MNHN.
2: In situ view of a mussel bed, with seaanemones Maractis cf. rimicarivora andzoarcid fish Pachycara cf. thermophilum;cruise Microsmoke © Ifremer.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
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Bathymodiolus septemdierum HASHIMOTO & OKUTANI, 1994
Size: Shell length up to 124 mm.
Morphology: Shell large, rather thin but solid, inflated, ellipti-cal, more or less compressed. Umbones subterminal, less thanone-eighth anterior of the shell length. Ligament weak. Ex-tremely thick and large ctenidia with long demibranchs. No ex-treme mantle fusion, valvular siphonal membrane short. Ante-rior adductor muscle scar located below the umbo. Anteriorpart of the posterior byssus retractor muscle scar under posteri-or end of ligament, at about four-fifths of ligament length.
Biology: At hydrothermal vents emitting fluids over 310°Caround the living beds. Endemic to vents. The shell surfaces arecovered with numerous filamentous bacteria. Periostracum col-or changes according to growth stages. Development notknown but most probably with planktonic larval phase.
Distribution: Izu Ogasawara Arc: Suiyo Seamount and MokuyoSeamount.
Reference:
HASHIMOTO J. & T. OKUTANI (1994) Venus 53(2): 61-83.
J. HASHIMOTO Denisia 18 (2006): 159
2: In situ © JAMSTEC.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
1: Paratype MNHN, interior and exterior of right valve;by R. von Cosel & P. Lozouet © MNHN.
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Bathymodiolus thermophilus KENK & WILSON, 1985
1: In situ views; cruise Hope © Ifremer; top right, by P. Briand © Ifremer.
Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae
R. LUTZ & R. VRIJENHOEK Denisia 18 (2006): 160–161
Size: Shell length up to 18.4 cm.
Morphology: Shell smooth, modioliform, with subterminal um-bones; periostracum present and straw-yellow to brown incolour; external surface lacks sculpture and is dull white be-neath periostracum; hinge edentulous; ligament opisthodetic,paricincular, strong, extending most of the length of the dorsalmargin; well developed byssus. Valvular siphonal membranelong and stretching towards ventrally; on anterior part fusion ofinner mantle fold reaching ventrally to 1/3 shell length, to-gether with siphonal membrane leaving rather short byssusopening in the middle.
Biology: The life habit of the adult is epibiotic on bare basaltand other hard substrates (e.g. tubes of vestimentiferans) asso-ciated with deep-sea hydrothermal vents. The presence of thisspecies is correlated with elevated levels of hydrogen sulfide.Endemic to vents. Protoconch 0.4 mm in length, indicative ofa planktotrophic larval stage with a high dispersal capability.Paired ctenidia consist of inner and outer demibranchs, eachwith descending and ascending lamellae. The gills contain sul-phur-oxidizing chemoautotrophic symbionts. A commensalpolychaete (Branchipolynoe symmytilida) is frequently foundwithin the mantle cavity of the mussel.
Distribution: Galapagos Spreading Center, East Pacific Rise:13°N to 22°S. At the Pacific-Antarctic Ridge (31°S and 38°S)another species of Bathymodiolus was collected in 2005 (R. vonCosel & R. Vrijenhoek, unpublished data).
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References:
BELKIN S., NELSON D.C. & H.W. JANNASCH (1986) Biol. Bull. 170: 110-121.CRADDOCK C., HOEH W.R., GUSTAFSON R.G., LUTZ R.A., HASHIMOTO J. & R.J. VRIJENHOEK(1995) Mar. Biol. 121: 477-485.CRADDOCK C., HOEH W.R., LUTZ R.A. & R.C. VRIJENHOEK (1995) Mar. Biol. 124: 137-146.FISHER C.R., CHILDRESS J.J., ARP A.J., BROOKS J.M., DISTEL D., FAVUZZI J.A., FELBECK H., HESSLER R.R., JOHNSON K.S., KENNICUT II M.C., MACKO S.A., NEWTON A., POW-
ELL M.A., SOMERO G.N. & T. SOTO (1988) Deep-Sea Res. 35: 1769-1791.GRASSLE J.P. (1985) Bull. Biol. Soc. Wash. 6: 429-442.KENK V. & B. WILSON (1985) Malacologia 26: 253-271.LUTZ R., JABLONSKI D., RHOADS D.C. & D. TURNER (1980) Mar. Biol. 57: 127-133.NELSON D.C., HAGEN K.D. & D.B. EDWARDS (1995) Mar. Biol. 121: 487-495.RHOADS D., LUTZ R.A., REVELAS E.C. & R.M. CERRATO (1981) Science 214: 911-913.
2 top: Interior of right valve and exterior of left valve of specimen 1; bottom: Exterior of left valve and ventral view of speci-men 2 to show restricted byssal opening; all from East Pacific Rise: 17°S; by R. von Cosel.
161
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Gigantidas gladius COSEL & MARSHALL, 2003
References:
COSEL R. VON & B.R. MARSHALL (2003) Nautilus 117: 31-46.HASHIMOTO J. & M. HORIKOSHI (1989) Deep-Sea Newsletter 15: 32-34.HASHIMOTO J. & T. YAMANE (2005) Venus 64: 1-10.JONES W.J., WON Y.J., MAAS P.A.Y., SMITH P.J., LUTZ R.A. & R.C. VRIJENHOEK (2006) Mar. Biol. 148: 841-851.
R. VON COSEL Denisia 18 (2006): 162
1: Two paratypes (MNHN), exterior and interior of right valve, exterior of left valve, dorsal view; by D. Brabant © MNHN.
Mollusca, Bivalvia, Pteriomorphia, Mytiloida, Mytilidae
Size: Shell length up to 31.6 cm.
Morphology: Shells large to very large, aduliform, extremelylong, elongate and sword-shaped, rather thin for the size butsolid, with flattened and very broad umbones situated well sub-terminally in the anterior half of the valves. Ventral marginwell concave. Periostracum strong and dark brown; valves dullwhite beneath the periostracum. Inner mantle folds separatealong whole length of the ventral margin their edges frilledalong the posterior fifth of the shell length or less. Valvularsiphonal membrane almost absent, but a deep cleft between in-ner mantle fold of right and left valve. Attachment point of theanterior foot-byssus retractor muscle directly above the anteri-or adductor scar and united with it; posterior byssus retractorcomplex multibundle with two principle diverging muscle bun-dles and two additional thin bundles.
Biology: Protoconch II about 0.4 mm in length, which indi-cates a long planktonic larval stage with a high dispersal capa-bility. The extremely enlarged gills contain sulphide oxidizingchemoautotrophic symbiotic bacteria. A commensal poly-chaete of the genus Branchipolynoe was encountered within themantle cavity of most specimens. Mussels of the genus Giganti-das are found on warm seeps near active submarine volcanoesand therefore are included herein. The species lives in densepopulations at sulphur-rich hydrothermal seepings, they arepartly buried in sediment.
Distribution: Southwestern Pacific: Rumble III and Rumble Vsubmarine volcanoes, S-Kermadec Ridge, New Zealand; West-ern Pacific: Kaikata Seamount, SW of Ogasawara (Bonin) Is-land.
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Gigantidas horikoshii HASHIMOTO & YAMANE, 2005
Size: Shell length up to 19.5 cm.
Morphology: Shells large, aduliform, long and slender, ratherthick, solid. Ventral margin markedly concave in adults, givingthe shell a curved appearance. Umbonal cavity large andswollen. Periostracum strong chocolate or dark brown, um-bonal region light brown; valves dull white beneath the perios-tracum. Inner mantle folds entirely separate, terminating ante-riorly on anterior adductor. No valvular siphonal membrane.
Anterior retractor scar situated in front of umbonal cavity; pos-terior byssus retractor complex with two principle divergingmuscle bundles.
Biology: The mussels were observed on sandy bottom withwarm water seepage (18°C). The species live in dense popula-tions, partly buried in the sediment.
Distribution: Kaikata Seamount, SW of Ogasawara.
References:
HASHIMOTO J. & M. HORIKOSHI (1989) Deep-Sea Newsletter 15: 32-34.HASHIMOTO J. & T. YAMANE (2005) Venus 64: 1-10.
J. HASHIMOTO Denisia 18 (2006): 163
1: Muscular system; after HASHIMOTO & YAMANE (2005).
2 top: Dorsal view; bottom: Lateral view; after HASHIMOTO & YAMANE (2005).
3: Specimens in situ, from Kaikata Seamount; by J. Hashimoto © JAMSTEC.
Mollusca, Bivalvia, Pteriomorphia, Mytiloida, Mytilidae
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Bathypecten vulcani SCHEIN-FATTON, 1985
1: Specimen viewed from the right (lower) side; by P. Briand.
3: Lower (right) side of the antero-dorsal region showingthe anterior auricle above the byssal notch; by P. Briand.
2: Same specimen viewed from the left (upper) side; by P. Briand.
Mollusca, Bivalvia, Pteriomorphia, Eupteriomorphia, Pectinidae
4: In situ view of a population of B. vulcani on basalt,among polychaete serpulid tubeworms (Laminatubusalvini), and small sea anemones, probably Chon-drophellia cf. coronata; by courtesy of R. Vrijenhoek.
Size: Up to 17 mm.
Morphology: Small, thin, flattened, left (upper) valve moreconvex than right (lower); smooth, but early stage with con-centric undulations, more developed and regular on uppervalve; hinge line straight, posterior auricle not delimited, ante-rior auricle above a distinct byssal notch in lower valve.
Biology: Can be abundant at the periphery of vents (30-50 in-dividuals m-2) with mussels and gastropods. Byssally attached indiffuse venting areas with a low temperature anomaly. Usual bi-valve filter-feeding status. Non planktotrophic larval develop-ment.
Distribution: Galapagos Spreading Center; East Pacific Rise:9°N and 13°N; Pacific-Antarctic Ridge: 32°S, observed at 38°S.
References:
LE PENNEC G., BENINGER P.G., LE PENNEC M. & A. DONVAL (2003) J. Mar. Biol. Ass. U.K. 83: 479-482.ROUX M., RIO M., SCHEIN E., LUTZ R.A., FRITZ L.W. & L. RAGONE (1989) C. R. Acad. Sci., Paris, Sér. III 308: 121-127.SCHEIN-FATTON E. (1985) C. R. Acad. Sci., Paris, Sér. III 301: 491-496.SCHEIN-FATTON E. (1988) Oceanol. Acta 8: 83-98.
E. SCHEIN & P. BRIAND Denisia 18 (2006): 164
1-4: Specimens from East Pacific Rise: 13°N, cruise HOT 96.
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Sinepecten segonzaci SCHEIN, in press
Reference:
SCHEIN E. (in press) Zootaxa.
E. SCHEIN Denisia 18 (2006): 165
1: Outer view of a juvenile left valve (SEM).
1-3: Specimens collected by J. Hashimoto, cruise Bioaccess 98; from SCHEIN (in press).
3: Upper side (left) and lower side (right) of large adult holotype.
2: Lower side of a juvenile specimen.
Mollusca, Bivalvia, Pteriomorphia, Eupteriomorphia, Pectinidae
Size: Up to 28 mm.
Morphology: Shell thin, lower (right) valve flat; upper (left)valve convex. Commarginal ridges or lamellae covering thewhole of the upper valve, and the lower valve only after the ju-venile stage. Posterior auricle poorly delimited. Anterior auri-cle of the lower valve above a byssal notch which is widelyopen in the juvenile, but gradually closing and finally over-lapped by its lower edge in the adult.
Biology: Byssally attached on glassy basalt near vents where thetemperature ranged between 2.8°C and about 40°C.
Distribution: Manus Back-Arc Basin. A juvenile specimen ofthe same genus was collected at North-Fiji Back-Arc Basin,close to the vent field White Lady in June 2005 (cruise TU-IM06MV, R. Vrijenhoek © MBARI).
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Mollusca, Cephalopoda
The living cephalopods, commonly the finned and finlessoctopuses, squid and cuttlefishes with the vampire squid andNautilus, comprise a well-defined class of Mollusca. Althoughthese marine predators have undergone an extensive evolu-tionary radiation, only benthic incirrate octopuses of Granele-done and Benthoctopus and benthopelagic cirrate octopusspecies of Cirroteuthis and Grimpoteuthis have been reportednear vents, and only one octopod genus is recognized as en-demic to hydrothermal vents, Vulcanoctopus on the East Pacif-ic Rise. Octopods may form an ecologically important compo-nent of the vent ecosystem as opportunistic rather than en-demic predators, but they remain under-collected in the venthabitat. The largest single cause is the difficulty in capturingthese animals. Manipulators can grab large individuals of somespecies, especially cirrate and incirrate octopuses of Granele-done, and suction samplers and traps effectively collect Vul-canoctopus specimens, but individuals of Benthoctopus tend to bemuch more wary. Even when secured, specimens can be severe-ly damaged.
A second problem limiting our knowledge of these animalsis that, even with the specimen in the hand, species-level dif-ferences in these little-known groups are often subtle at bestand rely heavily on internal characters of sexually maturemales. As gravid females of some species are suggested to con-gregate in rocky areas, such as mid-ocean ridges, collectionstend to be female-biased, although in Vulcanoctopus hydrother-malis almost all known specimens are male. Based on isolateddeep-sea specimens, current species boundaries may be found tobe artificial, but only after careful reassessment of type speci-mens and newly collected specimens.
Videotape from research submarines and remotely operatedvehicles have documented a wide repertoire of cirrate octopusbehaviours in response to submersible-linked disturbances.They also suggest that incirrate octopus may nearly continual-ly feed as they move slowly across sediment. Despite these ad-vances, we remain largely unable to address questions as basicas whether the animals can change colour, a notable characterof shallow-water octopods. Bioluminescence has been observedin some cirrate species and may be a more widespread phenom-enon than suspected. Careful collection of live specimens and
their maintenance under suitable aquaria conditions can an-swer this and other intriguing questions on the behaviour ofthese deep-sea cephalopods.
After specimen collection, a tissue sample from arm muscu-lature should be taken by slitting the skin on the dorsal side ofthe arm to expose the muscle. Opening the skin avoids per-turbing the ventral suckers and minimizes contamination fromthe skin. The tissue sample should be frozen or preserved in95% ethanol for subsequent analysis. The specimen should bethen placed in a 6% solution of buffered formalin in seawater,if possible with the arms extended. The specimen can be keptin the formalin for several days before being shifted to ethanol.The specimens should be examined for any parasites and theirlocations should be noted. The presence of adaptations to hy-drothermal vent habitat, such as high concentrations of heavymetals in the tissues or presence of amoebocytes clots in the ve-nous system and in the renal sacs, should be carefully exam-ined.
References:
BOYLE P.R & P. RODHOUSE (2005) Cephalopods. Ecology and Fisheries. Blackwell Science Ltd, Oxford, UK: 1-452.NIXON M. & J.Z. YOUNG (2003) The Brains and Lives of Cephalopods. Oxford Univ. Press: 1-368.NORMAN M.D & F.G. HOCHBERG (2005). Phuket Mar. Bio, Center Res. Bull. 66: 127-154.VOSS G.L. (1988) Malacologia 29: 295-307.
A. GUERRA, J. VOIGHT & R. VILLANUEVA Denisia 18 (2006): 166
1, 2: Two in situ views ofan unknown cephalopodtaken at the Mid-AtlanticRidge, north of Rainbow;cruise Flores © Ifremer.
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Cirrothauma murrayi CHUN, 1911
Size: Total length up to 940 mm; mantle length 220 mm, fromsampled specimens; may be longer from observations at the EastPacific Rise: 13°N.
Color: In situ photographs show cirroteuthids with purple, redand/or brown color on both oral and dorsal surfaces.
Morphology: External features: body relatively elongate (i.e.,not compressed) and gelatinous. Eyes degenerate, cup-like,without lenses or iris, embedded within the jelly of the skin,look like small black balls. Fins large, wide, longer that headwidth. Mantle aperture closed around a long, slender funnel.Each arm bears a single longitudinal row of suckers alternatingwith paired, very long cirri. Suckers strongly modified in barrel-like form. Intermediate web present, linking arms to primaryweb. Internal features: butterfly-shaped shell.
Biology: The spermatophores of the males are small, rounded,simple in structure, and stored in the oviducal gland of the fe-male, indicating an internal fertilization. In gravid females, theovarian oocytes reach 200 in number and are present in differ-ent sizes and stages of development, from 0.4-9 mm long. One
or two eggs are stored in the distal oviduct, nearly to be re-leased, measuring to 14 x 8.9 mm. This indicates that once sex-ual maturity it is attained, eggs are probably released one or twoat a time, following a continuous spawning strategy of repro-duction. No data exist about the feeding habits of this speciesand no predators are reported. Despite their small size, the eyesof C. murrayi are probably sufficient to detect the biolumines-cence produced by other animals and the wide aperture of thecornea allows for detection of flashes over wide angles and forgreater sensitivity. Locomotion it is basically by the fins and theanimal becomes streamlined during fin swimming.
Distribution: Specimens of this species have been collected inAtlantic, Pacific and Arctic Oceans. The animals were seen atgreat depths (1500-5000 m), often near the bottom, but some-times 300-450 m above it. In the NE Atlantic COLLINS et al.(2001) collected 27 specimens, most of them below 3000 mdepth. Always one or two specimens observed at the East Pa-cific Rise: 13°N-17°S, around the vents. Probably the samespecies observed near other vents along the East Pacific Rise.
References:
ALDRED R.G., NIXON M. & J.Z. YOUNG (1983) Philos. Trans. R. Soc. Lond. B 301: 1-54.CHUN C. (1913) in MURRAY J. & J. HJORT (Eds) Report of the scientific results of the “Michael Sars“ North Atlantic deep-sea expedition 1910, Zoology.
Bergen Museum 3: 1-21.COLLINS M.A. & R. VILLANUEVA (in press) Oceanogr. Mar. Biol. Ann. Rev. 44.COLLINS M.A., YAU C., ALLCOCK & M.H. THURSTON (2001) J. Mar. Biol. Ass. U.K. 81: 105-117.
R. VILLANUEVA & M. SEGONZAC Denisia 18 (2006): 167
1: Specimen in situ, East Pacific Rise: 13°N; cruise Biocyarise © Ifremer.
2: Specimen in situ, East Pacific Rise: 17°S; cruise Biospeedo© Ifremer.
Mollusca, Cephalopoda, Octopoda, Cirroteuthidae
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Cirroteuthis magna HOYLE, 1885
Size: Up to 1300 mm total length. The largest known cirrateoctopod.
Morphology: Deep-sea animals of very great size (up to 1300mm TL). Butterfly-like shell. Shell Width Index: 26-31. Veryvoluminous eyes (Eye-ball Diameter Index: 39-43), with largelenses (Lens Diameter Index: 12-18). Arm length up to 940mm. Primary web inserting at different levels on the oral andaboral ends of the dorso-lateral and ventro-lateral arms on bothsides, and at the same levels on both ends of the dorsal and ven-tral arms. Very long non-retractile cirri (Cirrus Length Index:96-71); the first cirri commences between the fourth and fifthsuckers. Three types of suckers on all arms: cylindro-conicalform and those with the acetabulum highly deformable on thefirst 2/3 and barrel-shaped on the rest of the arm; no enlargedsuckers in male or female.
Biology: C. magna is, as other cirrate octopods, a cephalopod,typically adapted to the deep-sea environment in the abyssalecosystem. These gelatinous animals are neutrally buoyant.
They can be considered as abyssopelagic animals, although theycan also rest on the bottom, where they probably feed. Its vo-luminous and operative eyes indicate that these animals areable to detect light produced by themselves, other animals ofthe same species, or by potential predators and prey. Thisspecies, and other related ones, have been observed to swim at2.2 km/h and 0.46 km/h. It occurs near hydrothermal vents, butwas rarely observed at the Mid-Atlantic Ridge.
Distribution: South Indian (2557 m) and Pacific (1500 m) tosubtropical North Atlantic (1350 m). One specimen (1300 mmlength) was caught at the Mid-Atlantic Ridge, near Logatchev,3351 m. Another specimen was captured near a vent site in thesouthern East Pacific Rise: 17°S, 2574 m (M. Lilley & K. VanDamm, chief Scientists; J. Voight, det.).
References:
GUERRA A.R., VILLANUEVA R., NESIS K. & J. BEDOYA (1998) Bull. Mar. Sci. 63(1): 51-81.HOYLE W.E. (1885) Annals Magazine Natural History Series 5 15: 222-236.HOYLE W.E. (1886) Sciences Researches Voyage HMS “Challenger” 1873-76, Zoology 16: 1-245.HOYLE W.E. (1904) Bull. Mus. Comp. Zool., Harvard Coll. 43: 1-71.NESIS K.N. (1987) T.F.H. Publications, Inc., Neptune City, USA: 1-351. ROPER C.F.E. & W.L. BRUNDAGE (1972) Smithon. Contrib. Zool. 121: 1-46.VILLANUEVA R., SEGONZAC M. & A. GUERRA (1997) Mar. Biol. 129: 113-122.
A. GUERRA & M. SEGONZAC Denisia 18 (2006): 168
1: Ventral view of mature male(220 mm). Reconstructionbased on the specimencaptured and video images;from GUERRA et al. (1998).
2, 3: In situ views of the specimen caught north of Logatchev; cruise Faranaut © Ifremer. Thespecimen swans by moving its fins a few meters of the bottom, a locomotion mode namedfin-swimming; from VILLANUEVA et al. (1997).
Mollusca, Cephalopoda, Octopoda, Cirroteuthidae
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Grimpoteuthis ROBSON, 1932
Size: Up to 115 mm mantle length.
Color: Skin of red brown, orange or purple color in fresh spec-imens. Usually oral side more pigmented that dorsal side.
Morphology: External features: body gelatinous and bell-shaped. Fins medium to large, with distinct lobe near the ante-rior fin insertion. Thick primary web. Intermediate web absent.Each arm bears a single longitudinal row of suckers alternatingwith paired, medium-sized cirri. Internal features: optic lobespherical, optic nerves pass though white body as a single boun-dle of fibres. Shell U-shaped with other edges of lateral wallsparallel, not tapered to single fine points. The genus comprises14 species.
Biology: Stomach contents in G. wuelkeri and G. boylei showedpolychaetes, copepods, amphipods and isopods. Mature femaleshave few and large eggs in oviducts, measuring from 10-18 mmin length, suggesting a continuous spawning mode as observedin other cirrate octopods. Individuals of Grimpoteuthis sp. havebeen observed resting and crawling on the sea bottom, swim-ming basically by the use of fins and also by arm-web contrac-tions. As a response to disturbance, individuals have been ob-served in web inversion, with arms and web upturned, oral sur-face facing outward, completely covering mantle, head and fins.
Distribution: Specimens of this genus has been collected in theAtlantic and Pacific Oceans. In the NE Atlantic, abundancesof Grimpoteuthis sp. ranged from 1.3-25 individuals km-2 atdepths from 1500-4850 m (COLLINS et al. 2001).
References:
COLLINS M.A (2003) Zool. J. Linn. Soc. 139: 93-127.COLLINS M.A & R. VILLANUEVA (2006) Oceanogr. Mar. Biol. Annual Review 44: xx-xx.COLLINS M.A., YAU C., ALLCOCK & M.H. THURSTON (2001) J. Mar. Biol. Ass. U.K. 81: 105-117.O’SHEA S. (1999) NIWA Biodiversity Memoir Wellington 112: 1-280.VILLANUEVA R., SEGONZAC M. & A. GUERRA (1997) Mar. Biol. 129: 113-122.
R. VILLANUEVA & M. SEGONZAC Denisia 18 (2006): 169
1: In situ specimen taken at East Pacific Rise: 13°N,site Elsa; cruise HOPE 99 © Ifremer.
2: In situ specimen taken at Mid-Atlantic Ridge,north of Rainbow; cruise MARVEL © Ifremer.
Mollusca, Cephalopoda, Octopoda, Grimpoteuthidae
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Vulcanoctopus hydrothermalis GONZÁLEZ, GUERRA, PASCUAL & BRIAND, 1998
1: Specimen female (scale bar 5 cm) collected at East Pacif-ic Rise: 21°S, hydrothermal vent site Grommit (Biospeedo,17.04.2004); by P. Briand © Ifremer.
2: Specimen observedon tubes of Riftia atEast Pacific Rise:13°N; cruise HOPE’99© Ifremer.
3: Specimen observedat East Pacific Rise:23°S in April 2005;cruise PAR 5 © MBARI.
Mollusca, Cephalopoda, Octopoda, Octopodidae
Size: Up to 56 mm mantle length, 235 mm total length and 45g body weight.
Color: In situ, white, eyes black.
Morphology: Body semi-translucent with a muscular consisten-cy; mantle pear-shaped and posteriorly mitre-like; presence of alarge white body (which covers the eye, the optic nerves andthe optic lobe), an inflated black bulb (dark swelling), in prox-imal third of the intestine, a crop, and a multilobulate digestivegland; absence of an ink sac. Arms 1.5-4.3 times mantle length.Two rows of suckers on each arm. Arm formula typically 1.2.4.3or 2.1.4.3. No enlarged suckers. Maximum depth of the largestsector of the web about 22% of the longest arm. Gills with 7-8lamellae per demibranch. Right arm III hectocotylized (HA1.5-2.1 times mantle length) in males. Ligula short (8-10% ofHA), lance-shaped and without transverse ridges. Calamus rep-resents 30-50% of the ligula length in fully mature specimens.Spermatophore length 70-125% of mantle length. Only one fe-male of 35 mm mantle length was collected; external morphol-ogy similar to male. Finger-like oocytes (80) ranging from 0.15-4 mm maximum length.
Biology: This benthic species has characters that represent ei-ther adaptations to the deep-sea (absence of ink sac, loss of theanal flaps, eye without iris and optic chiasma) or to a hy-drothermal vent habitat (eyes are covered by a thin semi-translucent skin, high concentrations of metals and presence ofamoebocytes clots in the venous system and in the renal sacs).It inhabits an isolated extreme environment among aggrega-tions of tubeworms Riftia pachyptila, Alvinellidae polychaetes ormussels, very close to the chimneys (2-10°C); also observed onthe pillow lava at several meters from the active areas. No pre-dation over these species was observed. Octopuses forage on ba-thypelagic amphipods, apparently targeting their attacks basedon contact with the swarming amphipods (e.g. Halice hes-monectes). Some male specimens were parasitised by Genesisvulcanoctopusi, a species of cholidynid harpacticoid (copepod).
Distribution: East Pacific Rise: collected at 13°N (Fig. 2) and21°S (Fig. 1); observed at 23°S (Fig. 3).
References:
GONZÁLEZ A.F., GUERRA A., PASCUAL S. & P. BRIAND (1998) Cah. Biol. Mar. 39: 169-184.GONZÁLEZ A.F., GUERRA A., ROCHA F. & P. BRIAND (2002) Bull. Mar. Sci. 71(1): 289-298.LÓPEZ-GONZÁLEZ P.J., BRESCIANI J., HUYS R., GONZÁLEZ A.F., GUERRA A. & S. PASCUAL (2000) Cah. Biol. Mar. 41: 241-253.NIXON M. & J.Z. YOUNG (2003) The Brain and Lives of Cephalopods. Oxford Univ. Press: 1-368.NORMAN M.D & F.G. HOCHBERG (2005) Phuket marine biol. Center Res. Bull. 66: 127-154. ROCHA F., GONZÁLEZ A.F., SEGONZAC M. & A. GUERRA (2002) Cah. Biol.Mar. 43: 299-302.VOIGHT J.R. (2005) J. Mar. Biol. Ass. U.K. 85: 985-988.
A. GONZÁLEZ, A. GUERRA & M. SEGONZAC Denisia 18 (2006): 170
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Benthoctopus GRIMPE, 1921
Size: Total lengths can approach one meter, but typically 50 cmor less.
Color: Variable, from light to red to violet or deep purple. Re-verse counter-shading (the dorsal surface lighter than ventralsurface) is pronounced in some species, absent in others.
Morphology: The genus is poorly delineated and is likely notmonophyletic; it includes octopuses without an ink sac withtwo rows of arm suckers. Field identification of members of thisgenus relies on their smooth skin, double rows of suckers(which are distinct from zig-zag sucker rows of octopus ofGraneledone), the comparatively narrow heads and mantlesand, in some species, the mantle being lighter in dorsally thanventrally. Although internal examination of specimens is re-quired to identify species, external characters such as col-oration, eye size and, in a few species, dramatically enlargedsuckers contribute to species identification.
Biology: Members of Benthoctopus are typically more active andmore apt to jet away from submersibles and ROVs than areother deep-sea octopus, making submersible-collected speci-mens rare. These octopuses sometimes extend their dorsal armsvertically into the water column, perhaps to enhance chemore-ception. Egg-brooding females may aggregate on hard substratewhere they often sit with the suckered surfaces of their armsfacing away from the rock.
Distribution: Octopuses of the genus Benthoctopus occur world-wide, typically between about 400 m rarely to as deep as 2000m. Octopuses of Benthoctopus have been seen near vents at Juande Fuca Ridge including Endeavour segment, and Gorda Ridge(where they brood eggs), and cold seeps off South America.
References:
OLU K. SIBUET M., HARMEGNIES F., FOUCHER J.-P. & A. FIALA-MÉDIONI (1996) Mar. Ecol. Progr. Ser. 132: 109-125. VOIGHT J.R. & A.J. GREHAN (2000) Biol. Bull. 198: 94-100. VOSS G.L. (1988) Malacologia 29: 295-307.
J.R. VOIGHT Denisia 18 (2006): 171
1: Brooding females; with numerous ophiuroids, probably Ophiuridae © MBARI.
2: Individual on the bottom.
Mollusca, Cephalopoda, Octopoda, Octopodidae
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1: Graneledone sp. with tubeworms Ridgeia; by courtesy of V. Tunnicliffe, HiRise Expedition.
Graneledone JOUBIN, 1918
Size: Adults from shallow in the depth range of the genus (800-1000 m) can exceed 1 m in length; those from in deepest partof the range (2800 m) are only about half as long.
Morphology: Large octopuses with a very broad head and man-tle, suckers arranged in a single or at times zig-zag row on eacharm; skin texture warty (warts more prominent in specimensfrom the greater depths), with conspicuous papillae over theeyes which can make the eyes appear larger. Individuals tend tobe uniformly colored, but individuals vary in color from violetto orange to blue. These octopuses tend to move with theirarms curled dorsally in what appears to be a semi-protective po-sition.
Remark: Identifying species of Graneledone is difficult, but eggscollected in the Northeast Pacific at near 2660 m depth wereshorter than those from near 1500 m depth indicating two dis-tinct species are present.
Biology: Hydrothermal vent polychaetes and gastropods werefound in the gut of an octopus from Axial Volcano. Egg-brood-ing females sit with their ventral (suckered) surface toward thehard substrate to which their eggs are attached; groups of egg-brooding females can form locally dense aggregations. On open,sedimented seafloor the octopuses may probe the sediment withtheir arms to locate infaunal prey.
Distribution: Specimens of the genus Graneledone have beencollected from Juan de Fuca and Explorer Ridges. Photos docu-ment an individual of the genus near clams at the GalapagosSpreading Center, near cold seeps off Oregon and Californiaand on the Kermadec Arc at Monowai Caldera (http://ocean-explorer.noaa.gov/explorations/05fire/logs/april12/april12.html).Members of the genus, also known from near Antarctica andSouth Africa, may associate with chemosynthetic areas.
References:
CORLISS J.B. & R.D. BALLARD (1977) Natl. Geogra. Mag. 152: 441-453. DRAZEN J.C., GOFFREDI, S.K., SCHLINDING, B. & D.S. STAKES (2003) Biol. Bull. 205: 1-7.MOORE J.C., ORANGE D. & L.V.D. KULM (1990) J.Geophys. Res. 95(B6): 8795-8808. VOIGHT J.R. (2000) Malacologia 42: 63-74. VOIGHT J.R. (2000) J. Zool. Lond. 252: 335-341.VOIGHT J.R. & J.C. DRAZEN (2004) J. Molluscan Stud. 70: 400-402.VOIGHT J.R. & A.J. GREHAN (2000) Biol. Bull. 198: 94-100. VOSS G.L. (1988) Malacologia 29: 295-307.
J.R. VOIGHT Denisia 18 (2006): 172
Mollusca, Cephalopoda, Octopoda, Octopodidae
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Nematoda
1: Thalassomonhystera sp. from East Pacific Rise: 9°N, Tica; by M. Bright.
173
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Desmodora alberti VERSCHELDE, GOURBAULT & VINCX, 1998
Size: 750-1150 µm.
Morphology: Desmodorid with rounded head capsule and long,slender, conical tail. Fine somatic setae; absence of subcephalicsetae. Body cuticle annulated. Amphid cryptospiral. Buccalcavity with large dorsal tooth, ventral teeth not observed.Cylindrical pharynx with muscular endbulb, bipartite cuticularvalves in terminal bulb of pharynx. Large globular sperm cellsand short bent spicule in males. Females with vulva at approx-imately 56% of body length.
Biology: Living in heterogeneous sediment covered with bacte-ria. Specific structure of buccal cavity points to epistrate feeder.
Distribution: Guaymas Basin.
Reference:
VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.
D. VERSCHELDE, J. ZEKELY & M. BRIGHT Denisia 18 (2006): 174
2: Holotype male anteri-or end; scale bar 50 µm;by D. Verschelde.4: Allotype female total; scale bar
50 µm; by D. Verschelde.
5: Head capsule (SEM); scale bar 1 µm; by D. Verschelde.
Nematoda, Adenophorea, Desmodorida, Desmodoridae, Desmodorinae
1: Holotype male total;scale bar 50 µm; by D. Verschelde.
3: Holotype male posterior end;
scale bar 50 µm; by D. Verschelde.
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Desmodora marci VERSCHELDE, GOURBAULT & VINCX, 1998
Size: 2110-2760 µm.
Morphology: Desmodorid with long slender body. Head cap-sule. Slender conico-cylindrical tail. Fine somatic setae. Bodycuticle annulated. Amphid cryptospiral. Buccal cavity withlarge dorsal tooth and one or two ventral teeth hard to distin-guish. Cylindrical pharynx with muscular endbulb. Spicules ofmales with complex capitulum. Vulva of females located at 50-55% of body length.
Biology: Colleted with mussels and Solenidae. Maximal tem-perature below 20°C. Specific structure of buccal cavity pointsto epistrate feeder.
Distribution: Lau Back-arc Basin, Hine Hina site.
Reference:
VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.
D. VERSCHELDE, J. ZEKELY & M. BRIGHT Denisia 18 (2006): 175
Nematoda, Adenophorea, Desmodorida, Desmodoridae, Desmodorinae
1: Holotype maletotal;scale bar 50 µm;by D. Verschelde.
4: Allotype female total; scale bar 50 µm;by D. Verschelde.
2: Paratype maleanterior end; scale bar 50 µm; by D. Verschelde.
3: Holotype maleposterior end; scale bar 50 µm;by D. Verschelde.
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Desmodorella balteata VERSCHELDE, GOURBAULT & VINCX, 1998
Size: 870-1080 µm.
Morphology: Desmodorid with long head capsule and short,conical tail approximately 10% of total body length. Slenderbody annuli. Two pairs of lateral rows of distinct spines amongother longitudinal rows of spines. Absence of subcephalic setae.Amphid multispiral, buccal cavity with cheilorhabdia incheilostome, large dorsal tooth and two subventral teeth.Cylindrical pharynx with muscular endbulb. Males with strong-ly built precloacal setae. Vulva in females at 68% of bodylength.
Biology: Living in heterogeneous sediment covered with bacte-ria. Specific structure of buccal cavity points to epistrate feed-er.
Distribution: Guaymas Basin.
Reference:
VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.
D. VERSCHELDE, J. ZEKELY & M. BRIGHT Denisia 18 (2006): 176
2: Holotype maleanterior end;scale bar 50 µm;by D. Verschelde.
5: Total view female (SEM); scalebar 100 µm; by D. Verschelde.
6: Head capsule male (SEM); scale bar 1 µm; by D. Verschelde.
Nematoda, Adenophorea, Desmodorida, Desmodoridae, Desmodorinae
1: Holotype male to-tal, note epizoic suc-torians attached tobody cuticle;scale bar 50 µm; by D. Verschelde.
3: Holotype maleposterior end; scale bar 50 µm; by D. Verschelde.
4: Allotype female total; scale bar 50 µm; by D. Verschelde.
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Desmodorella spineacaudata VERSCHELDE, GOURBAULT & VINCX, 1998
Size: 745-956 µm.
Morphology: Desmodorid with smooth three-part roundedhead capsule and short, conical tail. Slender body annuli. Twopairs of lateral rows of distinct spines among other longitudinalrows of spines. Extreme anterior position of multispiral amphid.Subcephalic setae at posterior edge of head capsule. Buccal cav-ity with large dorsal tooth and two subventral teeth. Cylindri-cal pharynx with muscular endbulb. Males with long filiformspicules and postcloacal thorns on the tail. Vulva in females atapproximately 62% of body length.
Biology: Living in heterogeneous sediment covered with bacte-ria. Specific structure of buccal cavity points to epistrate feed-er.
Distribution: Guaymas Basin.
Reference:
VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.
D. VERSCHELDE, J. ZEKELY & M. BRIGHT Denisia 18 (2006): 177
Nematoda, Adenophorea, Desmodorida, Desmodoridae, Desmodorinae
1: Holotype maletotal; scale bar 50 µm;by D. Verschelde.
5: SEM micrograph head capsule male;scale bar 1 µm; by D. Verschelde.
4: Allotypefemale total;
scale bar 50 µm; by D. Verschelde.
2: Holotype male anterior end; scale bar 50 µm; by D. Verschelde.
3: Holotype maleposterior end;
scale bar 50 µm;by D. Verschelde.
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Cephalochaetosoma pacificum notium DECRAEMER & GOURBAULT, 1997
Size: 575-850 µm.
Morphology: Draconematid with coarsely striated cuticle ex-cept in head region. Helmet present. Amphid spiral. Buccalcavity well developed, with a dorsal tooth and two small sub-ventral teeth. Pharynx with terminal bulb. Numerous cephalicadhesion tubes with non-swollen base and open tip located pos-terior head region and extending to about two head diamtersalong the cervical region; posterior adhesion tubes with bell-shaped end, all located anterior to cloacal opening/anus. Tailcylindro-conoid with numerous annules. Four pairs of cloacalsetae. Females with vulva at about 48% of body length. Maleswith spicule length 40-50 µm.
Biology: Hard substrate with oxide deposits and bacterial mats.Temperature 5-17°C. Specific structure of buccal cavity pointsto epistrate feeder.
Distribution: Lau Back-Arc Basin.
Reference:
DECRAEMER W. & N. GOURBAULT (1997) Zool. Scr. 26: 1-12.
J. ZEKELY & M. BRIGHT Denisia 18 (2006): 178
Nematoda, Adenophorea, Chromadorida, Draconematidae, Prochaetosomatinae
1: Holotype male total; scale bar 20 µm;by W. Decraemer.
2: Paratype female total; scale bar 20 µm; by W. Decraemer.
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Dinetia nycterobia DECRAEMER & GOURBAULT, 1997
Size: 320-490 µm.
Morphology: Draconematid with short, stout and sigmoid toepsilonematoid habitus. Body annulation reaching the lip re-gion and surrounding the small, spiral amphid. Cuticle notthickend in head region. Buccal cavity narrow and unarmed.Pharynx with terminal bulb with thickend cuticle. Cephalicadhesion tubes without expanded base located in cervical re-gion; posterior tubes with bell-shaped end. In females vulva at45% of body length.
Biology: Sedimented vent site. Specific structure of buccal cav-ity points to deposit feeder.
Distribution: East Pacific Rise: 21°N.
Reference:
DECRAEMER W. & N. GOURBAULT (1997) Zool. Scr. 26: 1-12.
J. ZEKELY & M. BRIGHT Denisia 18 (2006): 179
Nematoda, Adenophorea, Chromadorida, Draconematidae, Prochaetosomatinae
3: Paratype 4th
stage juvenilefemale; scale bar 20 µm:by W. Decraemer.
2: Paratype femaletotal; scale bar 20 µm;
by W. Decraemer.
1: Holotype maletotal; scale bar 20 µm; by W. Decraemer.
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Moravecnema segonzaci JUSTINE, CASSONE & PETTER, 2002
Reference:
JUSTINE J.-L., CASSONE J. & A. PETTER (2002) Folia Parasitol. 49: 299-303.
J.-L. JUSTINE Denisia 18 (2006): 180
1: Holotype male (A, B, D), paratype (C); A: Anterior end, lateral view; B: General body shape; C: Posterior end, ventral view(protruding spicules not drawn); D: Distal end of long spicule; from JUSTINE et al. (2002).
Nematoda, Secernentea, Spirurida, Habronematoidea, Cystidicolidae
Size: Male, 5 mm long, female, 4-10 mm long.
Morphology: Body filiform. Oral opening dorsoventrally elon-gated, hexagonal in shape. Pseudolabia rudimentary. Four sub-median buccal processes on margin of oral opening. Four sub-median papillae and two lateral amphids. Buccal cavity long,dilated dorsoventrally at anterior extremity. Oesophagus divid-ed into short anterior muscular and long posterior glandularparts. Deirids tiny, spine-like, located just anterior to posteriorend of buccal cavity. Nerve ring and excretory pore located atanterior and posterior thirds, respectively, of muscular oesoph-agus. Tail rounded in both sexes. Male: Caudal alae present.Area rugosa absent. Spicules unequal: short spicule arcuate,rounded at distal extremity, 80-100 µm long; long spicule 260-
360 µm long, distal extremity pointed with a cuticular spur at23 µm from end. Gubernaculum absent. Four pairs of precloa-cal pedunculate papillae arranged in two groups, Six pairs ofpostcloacal pedunculate papillae. Female: amphidelphic. Vulvaat about two thirds from cephalic apex. Eggs larvated in vagina,37-42 x 25 µm with 1-5 thin filaments, 40-140 µm long, aris-ing from a small plug at each pole. Tail 65-95 µm long.
Biology: Parasitic in the intestine of the zoarcid fish Pachycarathermophilum GEISTDOERFER, 1994.
Distribution: Mid-Atlantic Ridge: Logatchev; Snake Pit-Moose.
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Hypoechinorhynchus thermaceri BURON, 1988
References:
BURON I. DE (1988) J. Parasitol. 74: 339-342.BURON I. DE, HUNDLEY J.L. & M. SEGONZAC (2000) InterRidge News 9(2): 14-15.
I. DE BURON Denisia 18 (2006): 181
1: Female (left) and male (right);by I. de Buron.
4: In situ specimens in the gut of a zoarcidfish Thermarces cerberus; by I. de Buron.
5: In situ specimen of a digenean in the in-testine of a zoarcid fish Thermarces cerberus,among specimens of Hypoechinorhynchusthermaceri; by I. de Buron.
2: Proboscis, face view (SEM); by I. de Buron. 3: Proboscis, lateral view (SEM); by I. de Buron.
Acanthocephala, Palaeacanthocephala, Echinorhynchida, Hypoechinorhynchidae
Size: 3.5 to 5 mm long by 0.4 to 0.7 mm wide.
Color: White, or sometimes yellowish.
Morphology: Globular proboscis armed with ~2 rows of 2-3hooks each. Trunk unarmed. Testis arranged in tandem. Six ce-ment glands. Female genital pore subterminal.
Remarks: Needs to be carefully isolated from the mucosa bydissecting the intestine around the proboscis. For morphologi-cal studies, worms should be placed in distilled water for aboutone hour in order to ensure optimal evagination of the pro-boscis and fixed in AFA. For molecular studies, fix the wormsdirectly in 70% alcohol.
Biology: Intestinal parasite of zoarcid fish Thermarces cerberus.No intermediate host was identified but other species from thesame family have amphipods for intermediate hosts. Gono-choric with a slight sexual dimorphism. Attaches to intestinalmucosa using the proboscis.
Distribution: East Pacific Rise: 13°N.
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Thermonemertes valens ROGERS, GIBSON & TUNNICLIFFE, 1996
Reference:
ROGERS A.D., GIBSON R. & V. TUNNICLIFFE (1996) Deep-Sea Res. I 43(10): 1581-1599.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 183
1: T. valens (preserved specimen); left anterior end, right total;from ROGERS et al. (1996).
2: Thermonemertes sp.; live specimen from East Pacific Rise:9°N; by M. Bright.
Nemertini, Hoplonemertini, Monostilifera
3: Thermonemertes sp.; live specimen anterior end with par-tially extruded proboscis; from East Pacific Rise: 13°N; by M.Bright
Size: 5 cm.
Color: Bright red; after fixation uniformly flesh colored.
Morphology: The head is spatuloate and dorsoventrally flat-tened. A single cephalic furrow on the ventral surface towardsthe rear of the head. Mouth and rhynchodeum open into a sin-gle opening behind the tip of the head. The body is slender anddorsoventrally compressed. In the posterior half the ventral sur-face appears distinctly concave.
Remarks: Another yet undescribed species of Thermonemerteswas found at East Pacific Rise: 9°N from inactive sulfide chim-neys and at 13°N from Riftia pachyptila aggregations.
Biology: Lives on rocks and bacterial mats; not associated withvestimentiferan tubeworms. The diet remains unknown. Mightbe able to swim. It has been suggested that this species is an ear-ly colonizer of vent communities, but is displaced in aging com-munities.
Distribution: Juan de Fuca Ridge.
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Annelida, Polychaeta1
According to different countings, the polychaete worms, with111 presently described species, constitute 18-20% of the totalnumber of species fully identified from vent samples. 30% of thesebristle worms are belonging to scale worms (Polynoidae), 10% toalvinellids, 9% to siboglinids, 6% to spionids, 5% to hesionids anddorvilleids. Other families account for less than 5% of describedannelid species. The present recorded species are largely (94%)considered as ‘endemic’ of the deep-sea hydrothermal vent envi-ronment (or according to Andrey Gebruk, vent ‘obligate’), lessthan 4% are shared with other deep-sea reduced environments andca. 2% are ‘regular’ deep-sea species. However this high ratio of en-demism will probably be revised downward when the sampling ofthe peripheral areas will make progress accounting for opportunis-tic groups as dorvilleids or spionids which could be thriving in ad-jacent disturbed habitats.
Because of their odd biology and ecology, two species of an-nelids Riftia pachyptila (‘the giant tube worm’) and Alvinella pompe-jana (‘the Pompeii worm’) became emblematic of the vent researchand are still considered as ‘biological models’. Numerous researchprojects dealing with the functioning of endosymbiosis in Riftiapachyptila and the physiological adaptation of Alvinella pompejanato its extreme habitat are presently underway.
Nevertheless, the basic taxonomy of vent annelids is far to becompleted. Recent cruises which occurred in the Western Pacific(Lau Basin, Kermadec and Mariana Arcs) and on southern East Pa-cific Rise brought back a lot of new interesting specimens present-ly under study. For example, five new species of scale-worms fromLau Basin are being described belonging to Levensteiniella,Branchinotogluma, Lepidonotopodium and Harmothoe as well as onespecies of ampharetid, one sigalionid, one alvinellid (Paralvinella n.sp.), a new species of dorvilleid belonging to Parougia and one tere-bellid (cf. Polycirrus). From southern East Pacific Rise, severalspecies of scale worms, one species of alvinellid and one surprisingnew genus of spionid are currently under description. New familiesfor this environment were found and the description of new speciesof flabelligerid and sphaerodorid are under wording. Former collec-tions of worms from Mid-Atlantic Ridge and East Pacific Rise arestill not exhaustively studied and several families (e.g. dorvilleid,spionid, capitellid, cirratulid) need further taxonomic work.
The molecular identification of sibling species among wide-spread morphotypes (e.g. Amphisamytha galapagensis or Archinomerosacea) lead ‘classical’ taxonomists to dig back throughout formercollections looking for new diagnostic characters. For example, ajoint work between morphologists and molecular taxonomists al-
lows recognizing three different new species of Archinome and onenew genus of Amphinomidae among specimens previously identi-fied as A. rosacea (J. Kudenov, pers. comm.) Conversely, newrecords of previously described species from new locations (e.g. Hes-iolyra bergi on Mid-Atlantic Ridge vents or Hesiospina vestimentiferafrom Lau Basin) question the dispersal and allopatric speciation inresponse to major vicariance events. Because of the influence ofjoint occurrences in biogeographic analyses, the solution of thesetaxonomic riddle must be seriously considered and we urge scien-tists working at sea to focus on these widespread morphotypes andto build up parallel samples for classical and molecular taxonomy.
Polychaetes are delicate and fragile animals, and special careshould be taken when handling them. To be described, the speci-mens must be unfragmented and as intact as possible and damagedspecimens may be misidentified. Thus a gentle sorting is desirable assoon as possible after recovery of the samples, using soft pliers formacroscopic individuals and gentle sieving for others (avoid stackof sieves and split the samples before sieving). Specimens may be re-laxed prior to preservation (7.5% of magnesium chloride) and tubesof tubicolous annelids must be opened; fixation is best in bufferedformalin 5-10%. After a suitable time of fixation, depending of thesize (in general < 24 hours), the worms may be transferred to 80%ethyl or isopropyl alcohol (formaldehyde even buffered, is a verypoor preservative).
1 Even if the question is still strongly debated and the molecular information isnot consistent, we chose herein, following ROUSE & FAUCHALD (1997) to in-clude the Pogonophora (Perviata and Vestimentifera) within the family Si-boglinidae (Polychaeta: Sabellida). Contrarily SOUTHWARD et al. (2005) chosea more conservative standpoint and retain the class ‘Pogonophora’ within An-nelida, waiting for more conclusive information (HALANYCH 2005).
1: Laminatubus alvini; by courtesy of R. A. Lutz.
References:
HALANYCH K.M. (2005) Hydrobiologia 535/536: 297-307.ROUSE G.W. & K. FAUCHALD (1997) Zool. Scr. 26: 139-204.SOUTHWARD E.C, SCHULZE A. & S.L. GARDINER (2005) Hydrobiologia 535/536: 225-249.
D. DESBRUYÈRES DENISIA 18 (2006): 185
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Archinome rosacea (BLAKE, 1985)
Size: Up to 13 mm long, 5.8 mm wide.
Morphology: Body short, fusiform, roughly trapezoidal in cross-section, with 18 segments. Prostomium with paired dorsal andinconspicuous median antenna. Caruncle fused throughchaetiger 2, cantilevered to posterior margin of chaetiger 3 andanterior margin of chaetiger 4 (sometimes to chaetiger 5).Paired palps similar in shape, just slightly longer than pairedprostomial antennae. Peristomium glandular, ciliated; chordateshape divided into two lobes by a long, deep midventral groove,extending through anterior lip of mouth into pharynx. Mouthventral, opening at junction of chaetigers 1-2. Chaetae simple,calcareous and bifurcate. Long prongs of bifurcate chaetae near-ly straight, smooth to distally serrated with around three in-
conspicuous denticles confined to tips. Largest notochaetaeslightly less robust than largest neurochaetae. Ratio of long toshort prongs of bifurcate notochaetae ranging from 5.3-10.7: 1;neurochaetae from 2.1-4.7: 1; notoaciculae from 20.5-22: 5: 1;and neuroaciculae around 36: 1. Branchiae present fromchaetiger 3, palmate. Pygidium opening on dorsum of last twochaetigers; cirrus digitiform, recurved anterodorsally.
Biology: Ubiquitous; associated with tube worms and in musselbeds. Carnivore and scavenger with dietary preferences forpolychaetes, crustaceans; also some molluscs.
Distribution: Galapagos Spreading Center, Rose Garden,Northern East Pacific Rise.
References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 6: 67-101.KUDENOV J.D. (1991) Ophelia Supplement 5: 111-120.KUDENOV J.D. (1993) Antarc. Res. Ser. 58: 93-50.WARD M.E., JENKINS C.D. & C.L. VAN DOVER (2003) Can. J. Zool. 81(4): 582-590.
J.D. KUDENOV Denisia 18 (2006): 186
1: Specimens from East Pacific Rise: 13°N;by P. Briand © Ifremer. 4: Branchiae (SEM) © Ifremer.
5: Bifurcate chaetae withshort prongs; by Kudenov.
Annelida, Polychaeta, Eunicida, Amphinomidae
3: Prostomium and caruncle,dorsal view; by Kudenov.
2: Branchiae; by Kudenov.
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Exallopus jumarsi BLAKE, 1985
References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.PETRECCA R.F & J.F. GRASSLE (1989) in G.R. MC MURRAY (Ed.) Proc. Gorda Ridge Symp. May 11-13, 1987, Springer: 279-283.
J.A. BLAKE Denisia 18 (2006): 187
1A: Anterior end, dorsal view; B: Same, ventral view; C: Heavymodified chaetae from chaetiger 1; E: Group of composite fal-cigers from anterior chaetiger; F: Normal simple chaetae from anterior chaetigers; from BLAKE (1985).
2: Maxillary apparatus, showing maxillary carriers and indi-vidual plates. Labels on maxillary apparatus indicate maxil-lary carriers (MC) and maxillae (1-7); from BLAKE (1985).
Annelida, Polychaeta, Eunicida, Dorvilleidae
Size: Small, about 3 mm long, 0.6 mm wide for 26 segments.
Morphology: Prostomium wider than long, rounded along an-terior margin; with two annulated dorsal antennae and twoshort, smooth lateral palps. Chaetiger 1 modified, without dor-sal and ventral cirri, but bearing two types of heavy modifiedsimple chaetae: (1) upper group of 4-5 narrow, finely denticu-late chaetae; (2) lower group of four large spines with lateral ac-cessory tooth. Following segments with thick dorsal cirri, noventral cirri, with simple serrated chaetae and compound fal-cigers. Jaw apparatus with seven pairs of maxillae, all with larg-er terminal or subterminal tooth and numerous small denticles.
Biology: From fine sediments among hydrothermal mounds.Densities of 2667 individuals m-2 were recorded from mats ofBeggiatoa sp.
Distribution: Guaymas Basin.
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Ophryotrocha akessoni BLAKE, 1985
Size: Up to 8.5 mm long, 1.1 mm wide, for 55 segments.
Morphology: A small, compact species. Color in alcohol lighttan. Prostomium broadly rounded anteriorly with two smoothantennae and two similar palps. Segments 1-2 achaetous, fol-lowed by similar appearing chaetigers. Each segment with sin-gle ciliary band encircling body. Pygidium with two anal cirri.Each chaetiger with long dorsal cirrus and short, stubby ventralcirrus. Chaetae include dorsal group of simple serrated chaetae,ventral group of composite falcigers, and 1-2 ventralmost sim-ple chaetae. Maxillary apparatus with seven pairs of maxillae,each with numerous short teeth; maxillary carriers large, for-cep-like in adults.
Biology: An abundant, opportunistic epifaunal species occur-ring with siboglinids, mussels, and clams. In the GuaymasBasin, O. akessoni dominated in sediments saturated with pe-troleum hydrocarbons. Densities of 3,222 individuals per m2
were recorded from mats of Beggiatoa sp. in the Guaymas Basin.
Distribution: Widespread at the East Pacific Rise: 21°N, 13°N,9°N, Galapagos Spreading Center and Guaymas Basin.
References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 8: 103-116.GRASSLE J.F. (1988) Adv. Mar. Biol. 23: 301-366.PETRECCA R.F & J.F. GRASSLE. (1989) Proc. Gorda Ridge Symp.: 279-283.
J.A. BLAKE Denisia 18 (2006): 188
1: Specimen in vivo, apicalview Ifremer
3: Anterior part, lateral view (SEM) Ifremer 4: Composite falciger(SEM) Ifremer
2: Maxillary apparatus: left juvenile, rightadult; by courtesy of J. Blake.
Annelida, Polychaeta, Eunicida, Dorvilleidae
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Ophryotrocha globopalpata BLAKE & HILBIG, 1990
Size: Up to 7 mm long, 1.0 mm wide, with 40+ segments.
Color: In alcohol opaque white.
Morphology: Body wide, dorsoventrally compressed, blunt an-teriorly, tapering towards pygidium; with long, prominent para-podia. Prostomium rounded, twice as wide as long; with twolong, antennae; with two biarticulate palps bearing wide, glob-ular palpophores and slender palpostyles; eyes absent. Peris-tomium with two achaetous rings about as long as followingchaetigers. Parapodia uniramous, with triangular acicular lobe,low prechaetal lobe and retractable ventral chaetal lobe; dorsaland ventral cirri absent. Chaetae of three kinds: (1) supraacic-ular fascicle of long, flat capillaries with fine subdistal serration;(2) subacicular fascicle of heterogomph falcigers with distallyserrated shaft and smooth, narrow, distally blunt blades; (3) in-feriormost smooth capillary. Pygidium rounded with two long
dorsolateral anal cirri. Mandibles elongate, rodlike, with short,rounded, serrated cutting edge and large, unsclerotized lateralwings giving mandible triangular outline. Maxillae with P-typeforceps and eight pairs of free denticles with numerous smallteeth along cutting edge.
Biology: An epifaunal species associated with smokers and ac-tive vents. Mature females with about 6-8 eggs in each para-podium and sometimes 3-4 additional ones in large ventralpouches formed from ventral body wall; egg diameters from 64-120 µm, those in pouches larger (mean = 102 µm) than thosein the parapodium (mean = 81 µm). Mature males with spermfrom chaetiger 3 or 4 to end of body, visible in parapodia andunder dorsal body wall.
Distribution: Juan de Fuca Ridge.
1A: Anterior end dorsal view; B: Posterior end, dorsal view; C: Middle seg-ments of ovigerous female with ventral pouches, ventral view; D: Parapodiumanterior view; E: Supraacicular chaetae; F: Subacicular falcigers; G: Ventralmostcapillary; from BLAKE & HILBIG (1990).
1: Dorsolateral view of the anterior part (SEM)© Ifremer.
Annelida, Polychaeta, Eunicida, Dorvilleidae
References:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.TUNNICLIFFE V. (1988) Proc. R. Soc. Lond. B 233: 347-366.
J.A. BLAKE Denisia 18 (2006): 189
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Ophryotrocha platykephale BLAKE, 1985
2: Prostomium, dorso-lateral view (SEM) © Ifremer.
Annelida, Polychaeta, Eunicida, Dorvilleidae
1A: Anterior end, dorsalview; B: Same, ventral view;C: Left middle parapodium;D-G: chaetae; adapted fromBLAKE (1985).
Size: Up to 40 mm long, 2.5 mm wide, with more than 150 seg-ments.
Morphology: Body dorsoventrally compressed throughout,thickest anteriorly; posterior segments appearing ragged fromprojecting parapodia. Color in life and alcohol opaque white.Prostomium flattened, wider than long, with cirriform, distallytapering antennae and palps, both of similar length; eyes ab-sent. Peristomium with single achaetous ring. Uniramous para-podia projecting far away from body, with pre- and postchaetallobes short and rounded anteriorly, becoming elongate by aboutchaetiger 15. Dorsal cirri from chaetiger 17-19, becoming long,digitiform, bifid in posterior chaetigers. Ventral cirri short, re-tractile from chaetiger 3 to end of body. Dorsal branchiae fromchaetiger 14; ventral branchiae from chaetiger 2; ventralbranchiae longer than dorsal ones. Large parapodial glandspresent between dorsal cirrus and dorsal parapodial base.Chaetae include dorsal fascicle of simple spines and ventral fas-cicle of compound spinigers. Maxillae with large P-type forcepsand seven pairs of free denticles.
Remarks: (1) The illustration of the parapodium in the origi-nal description of O. platykephale (BLAKE 1985: Fig. 14C) wasshown upside down and misinterpreted. This error was laternoted by BLAKE & HILBIG (1990) and corrected by SOLÍS WEISS
& HILBIG (1992). The original figure has been corrected here.(2) A closely related morphotype of O. platykephale is presentin the same samples from the Guaymas basin hydrothermalmounds. It differs externally from O. platykephale having a peri-stomium with two achaetous rings and a globose dorsal cirrus(see Fig. 2-6). Its mandibles differ also from O. platykephale be-ing more spindly and curved. Its specific status remains to bediscussed (likely morphological variants or different stages ofO. platykephale).
Biology: Living specimens of O. platykephale were observedswimming among a colony of the ampharetid Amphisamythagalapagensis. The general habitat was of a seep site warmed bypercolating hot water, but away from chimneys.
Distribution: Guaymas Basin hydrothermal mounds.
J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 190–191
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References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 6: 67-101.BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.SOLÍS WEISS V. & B. HILBIG (1992) Bull. South Calif. Acad. Sci. 91: 92-96.
3: Medial parapodia, dorso-lateral view showing branchia andglobular dorsal cirri (SEM) © Ifremer.
4: Medial parapodia, dorso-lateral view showing the twochaetal fascicles (SEM) © Ifremer
5: Anterior parapodia, dorso-lateral view showing the twochaetal fascicles (SEM) © Ifremer.
6: Composite blunt-tipped chaetae (SEM) © Ifremer.
191
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Parougia wolfi BLAKE & HILBIG, 1990
Size: Up to 11.5 mm long, 1.0 mm wide, with about 55 seg-ments.
Color: In alcohol opaque yellowish-white.
Morphology: Body long, cylindrical, with projecting parapodia,tapering within last 5-10 segments towards pygidium. Segmen-tal ciliation absent (present on specimens observed by DD).Prostomium rounded anteriorly, triangular to pear-shaped, withlarge, flattened palps, longer than greatest prostomial width;antennae unknown. Peristomium with two rings, as long as fol-lowing chaetigers. Parapodia in chaetiger 1 uniramous, follow-ing chaetigers subbiramous; neuropodium with low, squareprechaetal lobe and longer, conical postchaetal lobe; notopodi-um with conspicuous acicula in long basal part and short, con-ical distal article; ventral cirri short, cirriform. Chaetae of
supraacicular fascicle simple, with subdistal serrations and fur-cate tip (seldom observed by D. DESBRUYERE); capillaries ab-sent. Subacicular fascicle with 10-15 compound falcigers withsubdistally serrated shaft and bifid, distally hooded blade. Py-gidium small, with terminal anus surrounded by heavily ciliat-ed, rounded papillae. Mandibles triangular, with heavily sclero-tized handle and transparent lateral wings; cutting edge withouter hook-shaped tooth and about five large, rounded and fivesmaller, irregular teeth. Maxillae with superior basal plates and30-35 superior and inferior free denticles; maxillary carriers andinferior basal plates absent.
Biology: Collected from active vents.
Distribution: Juan de Fuca Ridge.
1: Prostomium, dorso-lateral view (SEM) © Ifremer.
2: Anterior part, ventral view (SEM) © Ifremer.
4: Middle parapodia, dorsal view (SEM)© Ifremer.
3: Three compound falcigers of the sub-acicular fascicle with subdistally serratedshaft and bifid, distally hooked blade(SEM) © Ifremer.
6: Simple chaetae ofthe supraacicular fasci-cle with subdistal serra-tions (SEM) © Ifremer.
5: Simple chaetae ofthe supraacicular fas-cicle with a furcatetip (SEM) © Ifremer.
Annelida, Polychaeta, Eunicida, Dorvilleidae
Reference:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.
J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 192
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Eunice pulvinopalpata FAUCHALD, 1982
Size: A large species, 450 mm long, 12 mm wide, for more than350 chaetigers.
Morphology: A large, robust eunicid. Prostomium short, trans-verse, with pair of large palps along anterior edge; with fiveweakly articulated antennae. Peristomium with two achaetousrings, with separation between the two indistinct; a pair of peri-stomial cirri on second ring. Branchiae from chaetiger 3-4 toend of body; branchiae mostly pectinate, with multiple fila-ments. Parapodia all similar, with rounded acicular lobe bearing
three protruding dark brown or black aciculae; with long dorsalcirrus and short, thick ventral cirrus. Chaetae include bidentatecompound hooded hooks, limbate capillaries, pectinatechaetae, bidentate subacicular hooks, and acicula.
Biology: Found on smoker walls in rusted areas; probably a car-nivore.
Distribution: East Pacific Rise: 21°N and 13°N.
References:
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 8: 103-116.FAUCHALD K. (1982) Proc. Biol. Soc. Wash. 95: 871-877.
J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 193
1: Anterior part, dorsal view;by P. Briand © Ifremer.
3: Anterior part, ventral view;by P. Briand © Ifremer.
5: Anterior part of the proboscis (SEM) © Ifremer.
2: Midpart, dorsal view;by P. Briand © Ifremer.
4: Parapodium, SEM viewshowing one of the darkbrown acicula © Ifremer.
6: Bidentate compound hooded hooks(SEM) © Ifremer.
7: Pectinate chaeta (SEM) © Ifremer.
Annelida, Polychaeta, Eunicida, Eunicidae
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Thraumastos dieteri WATSON, 2001
Size: > 10 mm maximum size (type truncated).
Morphology: Anterior mid-body rectangular, long tapered an-teriorly, numerous segments. Pale gold coloured palae fans(brown areas present). Prostomium small quadrate compressedbetween anterior segments. Subulate median antennae, withshort ceratophore, inserted medially on anterior edge of theprostomium, dorsally to two similar sized lateral antennae.Prostomial cirri with filiform tips. Two ovoid palps inserting on
the ventral edge of the prostomium. Mid-body notopodia withlateral and main paleal fascicles. Neuropodia with subacicularfascicle of heterogomph falcigerous neurochaetae. Glandularpads present.
Biology: Common in washings of Bathymodiolus brevior.
Distribution: North Fiji and Lau Basins and cold seep south ofLihir Papua New Guinea.
1: Habitus in vivo Ifremer
2: Prostomium, dorsal view (SEM) Ifremer.
4: Prostomium, ventral view (SEM) Ifremer.
3: Notopodial fan, middle body (SEM) Ifremer.6: Detail of two median paleae (SEM) Ifremer.
5: Neuropodial spinigerous falciger (SEM) Ifremer.
Annelida, Polychaeta, Phyllodocida, Chrysopetalidae
References:
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.WATSON C. (2001) The Beagle, Records of the Museums and Art Galleries of the Northern Territory 17: 57-66.
D. DESBRUYÈRES Denisia 18 (2006): 194
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Glycera branchiopoda MOORE, 1911
References:
BLAKE J.A. (1985) Bull. Biol. Soci. Wash. 8: 67-101.BOGGEMANN M. (2002) Abh. Senckenb. Naturforsch. Ges. 555: 1-249.FAUCHALD K. (1972) Allan Hancock Monography in Mar. Biol. 7: 1-573.
M. BOGGEMANN Denisia 18 (2006): 195
1A-D: Proboscidial papillae, main and additional types (A-C posterior; D anterior view, USNM 17304; C SMF 9391);from BOGGEMANN (2002, page 97, fig. 26) with the editor’s kind permission.
Annelida, Polychaeta, Phyllodocida, Glyceridae
Synonym: Glycera profundi CHAMBERLIN, 1919.
Size: Body up to 90 mm with up to 137 segments.
Morphology: Conical prostomium long with 8-9 rings. Pro-boscis with two types of papillae: (1) numerous tall thin, withstraight, median longitudinal ridge and very small papillae sur-rounding core; (2) isolated shorter broader oval to globularpapillae without ridge. Aileron with pointed triangular base.First two parapodia uniramous; following parapodia biramous.Two slender triangular to digitiform praechaetal lobe. Oneshorter, rounded, sometimes blunt triangular postchaetal lobe.Dorsal cirri distinctly visible from 3rd parapodium, oval to glob-ular. Posterior parapodial lobes prolonged. Branchiae absent.
Biology: Burrows in soft sediments; not endemic. Glycerids,sometimes called bloodworms, are long-bodied active burrow-ing predatory worms, which lives in soft sediments.
Distribution: Pacific coast of North America, Central Ameri-can Trench; off central Oregon and Gulf of California deepbasins including Guaymas Basin hydrothermal mounds. Depthrange 10-2562 m.
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Glycera tesselata GRUBE, 1863
Size: Up to 76 mm long with 181 segments.
Morphology: Proboscidial papillae mainly digitiform withstraight, median, longitudinal ridge; aileron with deeply incisedbases; parapodia of mid-body with two short roundedpostchaetal lobes; branchiae absent. Mid-body segments bian-nulate. Conical prostomium consisting of about 8-9 rings. Pro-boscis with two types of papillae: (1) numerous digitiform papil-lae with straight, median, longitudinal ridge; (2) isolated,shorter and broader conical papillae with more or less distinct-ly straight, median longitudinal ridges. First two parapodia uni-ramous; following parapodia biramous. Two slender triangular
to digitiform prechaetal lobes; neuropodial lobe usually slightlylonger and wider than notopodial lobe. Dorsal cirrus from thirdparapodium, conical to oval; inserted on body wall far aboveparapodial base. Branchiae absent.
Biology: Glycerids are considered to be carnivorous worms.
Distribution: Northwestern and northeastern Atlantic, Gulf of Mexico, Caribbean Sea, Mediterranean Sea, Red Sea, south coast of Africa, Indo-Pacific, northwestern Pacific; 2000-4066 m; Mid-Atlantic Ridge: Lucky Strike.
1: Top left, proboscis and anteriorpart in a living specimen; by P. Briand © Ifremer.
2: Same part, lateral view (SEM) ©Ifremer.
Annelida, Polychaeta, Phyllodocida, Glyceridae
3: Proboscidial papillae (SEM) © Ifremer.
D. DESBRUYÈRES Denisia 18 (2006): 196–197
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Reference:
BÖGGEMANN M. (2002) Abhandl. Senkenberg. Naturforsch. Ges. Frankfurt 555: 1-249.
4: Proboscidial papillae (SEM) © Ifremer.
5: Median parapodia, dorsal view (SEM) © Ifremer.
6: Distal part of main papillae (SEM) © Ifremer.
7: Main and additional type of proboscidial papillae (SEM) © Ifremer.
8: Notochaetae, median parapodia (SEM) © Ifremer.
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Amphiduropsis axialensis (BLAKE & HILBIG, 1990)
Size: All known specimens incomplete, anterior end fragmentsup to 8 mm long for 21 segments.
Color: Preserved animals yellow-white, pigmentation absent orwith dark brown-black pigmentation on lips, laterally on pros-tomium, palpophores and nuchal organs, distally on proboscis,and on venter.
Morphology: Prostomium rounded rectangular, wider thanlong, posteriorly with small median incision. Palpophores veryshort, palpostyles of equal width or with slightly enlargedrounded tips. Paired antennae as long as palps, tapered. Bothpalpostyles and paired antennae very long and slender. Medianantenna much shorter than paired antennae, with tapering tip,inserted medio-dorsally on prostomium. Eyes absent. Proboscissmooth and unarmed, ending anteriorly in ring of cilia butwithout papillae. Dorsal cirri and cirrophores on segment 1–5larger than on following segments, ventral cirri on segment 1–4larger and longer than on following segments, with distinct cir-
rophores. Neuropodial lobes and neurochaetae present fromsegment 5, notopodial lobes and notochaetae from segment 6.Notopodial lobes conical, pointed. Notochaetae include manyfine capillaries with two rows of small teeth, three to four ven-trally situated curved chaetae with distal serration on ventralside, and two (rarely one) medially situated stout emerging aci-cular chaetae. Neuropodial lobes conical, much larger than no-topodial lobes. Neurochaetae all compound. Ventral cirri dis-tally inserted on neuropodium.
Remarks: The new record from East Pacific Rise: 9°N is basedon a single specimen from the expedition Hope’99 (Nautiledive 1372, 2500 m, 09°46’17’’N, 104°21’70’’W); although notin good condition it agrees well with A. axialensis.
Distribution: Juan de Fuca Ridge: Axial Seamount and MiddleValley; East Pacific Rise: 9°N (new record; see Remarks).
References:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.PLEIJEL F. (2001) Ophelia 54: 15-27.
F. PLEIJEL Denisia 18 (2006): 198
1A: Anterior end, dorsal view, pharynx extended; B: Parapodium from middle segment, anterior view; by courtesy of J. Blake.
Annelida, Polychaeta, Phyllodocida, Hesionidae
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Hesiodeira glabra BLAKE & HILBIG, 1990
Size: No complete specimens known. Anterior end of 21 seg-ments 5.6 mm long.
Morphology: Body shape cylindrical with flattened venter.Prostomium rounded rectangular, wider than long, with anteri-or median incision (Fig. 1A). Palpophores prominent, annulat-ed, palpostyles tapered with pointed tips. Paired antennaecylindrical with short pointed tips, median antenna small,pointed, inserted dorsally on prostomium (not visible on Fig.1A). Eyes absent. Proboscis unknown. Dorsal cirri and cir-rophores on segment 1–5 and ventral cirri and cirrophores on
segment 1–3 larger than on following segments. Dorsal cirriconnected by several muscle bands. Ventral cirri on segment 4very small. Neuropodial lobes and neurochaetae from segment5, notopodial lobes and notochaetae from segment 6. No-tochaetae include capillaries with two rows of teeth and a fewfurcate chaetae. Neurochaetae all compunds, with indentatetips. Ventral cirri inserted distally on neuropodium.
Distribution: Juan de Fuca Ridge: Axial Seamount.
References:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.PLEIJEL F. (1998) Zool. Scr. 27: 89–163.
F. PLEIJEL Denisia 18 (2006): 199
1A: Anterior end, dorsal view; B: Right parapodium from middle segment, anterior view; by courtesy of J. Blake.
Annelida, Polychaeta, Phyllodocida, Hesionidae
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Hesiolyra bergi BLAKE, 1985 “caterpillar worm“
Size: Up to 80 mm for 100 segments.
Morphology: Body anteriorly truncate and posteriorly tapered,dorso-ventrally flattened. Prostomium rounded rectangular,slightly wider than long. Palpophores short, stout; palpostylesmuch longer and thinner than palpophores, cylindrical withrounded tips. Paired antenna similar to palpostyles; median an-tenna absent. Eyes absent. Outside of everted proboscis withpointed papillae, increasing in size distally. Terminal ring ofproboscis with large number of pointed papillae. Inside of pro-boscis with large number of denticles in band on ventral side;dorsal and ventral teeth present, dorsal much larger. Dorsal cir-ri and cirrophores on segment 1–5 and ventral cirri and cir-rophores on segment 1–4 larger and longer than on followingsegments. Neuropodial lobes and neurochaetae from segment 5,notopodial lobes and notochaetae from segment 6. No-tochaetae surrounded by large pre- and postchaetal lamellae,smaller lamellae also surrounding neurochaetae. Notochaetaeof four kinds: one-two serrated flattened (difficult to detect),
large number of lyriform, tapered spinulose, and subdistally en-larged (and often twisted) spinulose. Neurochaetae all com-pound, dark to black in color (possibly due to some inclu-sions?), with unidentate tips. Ventral cirri inserted medially onneuropodium, with distinct cirrophores.
Biology: On smoker walls, often visiting tubes of Alvinella pom-pejana. Can occur in large numbers. Epibiotic filamentous bac-teria on parapodia and chaetae. Egg diameter about 125 µm.
Distribution: East Pacific Rise: 9°N and 21°N, and a closely re-lated species on Mid-Atlantic Ridge: at the Azores TripleJunction (expeditions Diva 1 and 2, Marvel) and 22°N(Microsmoke). Newly collected specimens from Mid-AtlanticRidge agree well with H. bergi. Whether they should be referredto a new species or not cannot be stated at present; they arecurrently best referred to as “Hesiolyra sp.”.
2: Population on a smoker wall, visit-ing Alvinella’s tubes (East Pacific Rise13°N, 2630 m). Phare cruise Ifremer.
3: Proboscis, frontal view (SEM) Ifremer.
1: Habitus in vivo; by P. Briand Ifremer.
Annelida, Polychaeta, Phyllodocida, Hesionidae
F. PLEIJEL Denisia 18 (2006): 200–201
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References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 6: 67–101.DIXON D.R., DIXON, L. R. J., SHILLITO, B. & J.P. GWYNN (2002) Cah. Biol. mar. 43: 333–336.PLEIJEL F. (1998) Zool. Scr. 27: 89–163.SHILLITO B., JOLLIVET D., SARRADIN P.M., RODIER P., LALLIER F., DESBRUYÈRES D. & F. GAILL (2001) Mar. Ecol. Progr. Ser. 216: 141–149.
4A: Habitus; by V. Martin © Ifremer; B-D: Denticles of the proboscis; E: Left parapodium ofsegment 13; by courtesy of J. Blake; F: Lyriform notochaeta; G: Tapered spinulose notochaeta;H: Compound neurochaeta (B-D & F-H © F. Pleijel).
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Hesiospina vestimentifera BLAKE, 1985
Size: Up to 36 mm long for 45 segments.
Color: Live specimens yellowish transparent, with yellow-or-ange gut, dorsal and parapodial blood vessels red. Preservedspecimens yellowish.
Morphology: Body anteriorly truncate and posteriorly tapered.Prostomium rounded trapezoid, anteriorly wider (Fig. 5A).Palpophores long; palpostyles much shorter than palpophores,varying in shape from ovoid to elongated, tips rounded. Pairedantenna thin, evenly tapering to rounded tips, slightly shorteror of same length as palps. Median antenna absent. Eyes absent.Everted proboscis basally with papillae appearing in c. 10–15poorly defined rows, with three to five conical, pointed papillaein each row. Papillae increasing in size distally. Anterior end ofproboscis with 10 conical papillae. Dorsal and ventral pair ofjaw plates inside terminal ring of papillae, both pairs with twobasally jointed lamellae, dorsal pair anteriorly rounded, ventralpair pointed (PLEIJEL, 2004, fig. 10A, B). Dorsal cirri and cir-
rophores on segment 1–5 larger than on following segments,ventral cirri on segment 1–4 larger and longer than on follow-ing segments, with distinct cirrophores. Neuropodial lobes andneuropodia from segment 5, notopodial lobes and notochaetae(except hooks; see below) absent from all segments. Notopodiafrom segment 8–9 to last segments with single stout, straight toslightly curved protruding hook, emerging on frontal side of cir-rophore. Neuropodial lobes elongated triangular. Neurochaetaeof two kinds: one to two stout, bent acicular chaetae, usuallyshorter than shafts of compounds, and c. 50 compounds withunidentate tips. Ventral cirri inserted subdistally on neuropodi-um, without distinct cirrophores.
Biology: Commonly associated with the siboglinid Riftia. Eggsize 100–120 µm in diameter.
Distribution: Explorer Ridge, Juan de Fuca Ridge, GalapagosSpreading Center, and East Pacific Rise.
Annelida, Polychaeta, Phyllodocida, Hesionidae
1A: Anterior end, dorsal view with proboscis everted; B: Middle left parapodium, ante-rior view; C: Notopodium, showing aciculum and protruding acicular spine; by courtesyof J. Blake.
F. PLEIJEL Denisia 18 (2006): 202–203
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3: Prostomium, frontal view Ifremer.4: Parapodia, dorso-lateral view Ifremer.
2: Habitus, dorsal view; by P. Briand Ifremer. 3: Anterior part in vivo; by P. Briand Ifremer.
References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.DESBRUYÈRES D., GAILL F., LALLIER L. & Y. FAUQUET (1985) Bull. Biol. Soc. Wash. 8: 103-116.PLEIJEL F. (2004) J. nat. Hist. 38: 2547–2566.TUNNICLIFFE V. (1988) Proc. R. Soc. London B 233: 347-366.
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Nereimyra alvinae BLAKE, 1985
Size: Length of largest observed anterior fragment 3 mm for 20segments.
Morphology: Prostomium wider than long. Palpophores cylin-drical, palpostyles thinner, basally inflated with tapering tips.Paired antennae thinner and longer than palpostyles, medianantenna absent. Eyes absent. Everted proboscis smooth, anteri-or end with ring of papillae, jaws absent. Dorsal cirri and cir-rophores on segment 1–3 (possible 1–4) longer than on follow-ing segments, ventral cirri (and cirrophores?) on segment 1–3larger than on following segments. Noto- and neuropodial lobesand noto- and neurochaetae from segment 4. Notochaetae allcapillaries, neurochaetae unidentate capillaries and com-pounds. Ventral cirri inserted subdistally on neuropodium.
Biology: Associated with mussels at the Galapagos Rift andsorted from sediment in box cores from the Guaymas Basin.
Remarks: Poorly known and requires redescription (the typematerial is in bad condition and the above description is basedon the original one); the affinities with other members ofNereimyra is uncertain.
Distribution: Galapagos Spreading Center, Guaymas Basin.
Reference:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.
F. PLEIJEL Denisia 18 (2006): 204
1A: Anterior end, dorsal view with proboscis everted; B: Middle parapodium, posterior view; by courtesy of J. Blake.
Annelida, Polychaeta, Phyllodocida, Hesionidae
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Sirsoe grasslei (BLAKE, 1985)
Synonym: Orseis grasslei BLAKE, 1985.
Size: Length 8–9 mm for 32 segments (holotype).
Morphology: Body shape anteriorly truncate and posteriorly ta-pered. Prostomium rounded rectangluar, slightly wider thanlong. Palpophores short and large, palpostyles much longer,evenly tapered and pointed. Paired antennae similar to pal-postyles but slightly smaller, median antenna similar to pairedones but much smaller, inserted on dorsal, posterior part ofprostomium. Eyes absent. Everted proboscis smooth, unarmed,anterior end with ring of 10 small pointed papillae. Dorsal cir-ri of segment 1–2 or 1–3 (uncertain) larger than on followingsegments, ventral cirri on segment 1–2 larger than on following
segments and with developed cirrophores (also on segment 3).Notopodial lobes and notochaetae absent from all segments.Neuropodial lobes and neurochaetae present from segment 2.Neurochaetae all unidentate compounds. Ventral cirri subdis-tally inserted on neuropodium.
Remarks: Poorly known, requires redescription.
Biology: Densities of 2,844 individuals per m2 have beenrecorded from mats of Beggiatoa (PETRECCA & GRASSLE 1989).Eggs large, about 250 µm in diameter.
Distribution: Guaymas Basin.
References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.PETRECCA R.F & J.F. GRASSLE (1989) Proc. Gorda Ridge Symp.: 279-283.PLEIJEL F. (1998) Zool. Scr. 27: 89-163.
F. PLEIJEL Denisia 18 (2006): 205
1A: Anterior end, dorsal view, proboscis everted; B: Right parapodium from chaetiger 6, anterior view; by courtesy of J. Blake.
Annelida, Polychaeta, Phyllodocida, Hesionidae
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Iheyomytilidicola tridentatus MIURA & HASHIMOTO, 1996
Size: Up to 20 mm in length. Up to 2.3 mm in width.
Color: Preserved specimens pale.
Morphology: Body flattened ventrally and arched dorsally, with106 segments. Prostomium short with a pair of short cirriformantennae, without eyes. Second or tentacular segment com-pletely fused to prostomium, lacking dorsal cirri and neu-rochaetae, defined by ventral cirri. Parapodia subbiramousthroughout the body, with very short dorsal and ventral cirri.Notopodia slightly inflated, supported by single stout acicula.Neuropodia supported by a single stout neuroacicula, project-ing from the parapodial base, slightly elongated on middle andposterior segments. Ventral cirri on ventro-posterior side ofneuropodia, decreasing in size on middle and posterior parapo-dia. Neuropodial hooks on dorsal side of chaetal lobes, stout,slightly inflated subdistally, with minute projection on cutting
edge of the main fang; occurring up to five on each parapodi-um. Additional neurochaetae two kinds; tridentate chaetae onouter base of chaetal lobes, simple slender than neuropodialhooks, shorter than other chaetae, with blunt main tooth onone side of distal tip and two shorter blunt teeth on oppositeside, occurring 10-20 per parapodium; minute slender chaetaeon inner side of chaetal lobe, enlarged subdistally, with singledistal mucronate spin, longer and more numerous than triden-tate chaetae. Pygidium rounded, without anal cirri.
Biology: Living in the mantle cavity of the deep-sea mussel Ba-thymodiolus aduloides. Foregut with well muscularized part; sto-modeum may be eversible. The species is thought to be para-sitic.
Distribution: Okinawa Trough.
Reference:
MIURA T. & J. HASHIMOTO (1996) Publ. Seto Mar. Biol. Lab. 37(3/6): 257-274.
T. MIURAS Denisia 18 (2006): 206
1A: Anterior end, dorsal view; B: Posterior end, dorsal view; C:Anterior end, ventral view; D: Same enlarged; E: Parapodiumof the 12th segment (or parapodium 11); F: Parapodium 40,anterior view; from MIURA & HASHIMOTO (1996).
2A: Neuropodial hooks; B: Tridentate chaetae; C: Minute mu-cronate chaetae; D: Anterior end of a juvenile; from MIURA &HASHIMOTO (1996).
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae
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Mytilidiphila enseiensis MIURA & HASHIMOTO, 1993
Size: Up to 28 mm in length. Up to 1.5 mm in width.
Color: Preserved specimens pale.
Morphology: Body long, vermiform, flattened ventrally andarched dorsally, with up to 164 segments; first segment tentac-ular, chaetigerous. Prostomium short with a pair of short cirri-form antennae, without eyes. Second segment well fused withprostomium, defined by embedded acicula and ventral cirri aswell as projected simple hooks. Notopodia very short, with dor-sal cirri and stout acicula. Neuropodia cylindrical, with very
short ventral cirri and stout acicula; ventral hooks simple, thin,occurring more than 35 as maximal number of posterior para-podia. Pygidium simple, without anal cirri.
Biology: Living in the mantle cavity of a burrowing deep-seamussel (Adula?). Foregut with well muscularized part; sto-modeum may be eversible. The species is thought to be para-sitic.
Distribution: Okinawa Trough.
References:
HASHIMOTO J., OHTA S., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. 42: 577-598.MIURA T. & J. HASHIMOTO (1993) Zool. Soc. 10: 169-174.
T. MIURA Denisia 18 (2006): 207
1A: Anterior end, dorsal view; B: Posterior end, dorsal view; C: Anteriorend enlarged, dorsal view; D: Same, ventral view; E: Parapodium of the 11th
segment, anterior view; F: Parapodium of the 27th segment, anterior view;G: Hooks; from MIURA & HASHIMOTO (1993).
2: Anterior part in dorsal view; by T. Miura.
3: Details of the prostomium; by T. Miura.
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae
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Mytilidiphila okinawaensis MIURA & HASHIMOTO, 1993
1A: anterior end, dorsal view; B: Same enlarged, dorsal view; C: Same, ventral view; D: Posterior end, dorsalview; E: Parapodium of the 11th segment (or parapodium 10) anterior view; F: Parapodium of the 36th segment(or parapodium 35), anterior view; G: Hooks; from MIURA & HASHIMOTO (1993).
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae
Size: Up to 15 mm in length. Up to 1.4 mm in width.
Color: Preserved specimens pale.
Morphology: Body long, vermiform, flattened ventrally andarched dorsally, with about 100 segments; first segment tentac-ular, achaetous. Prostomium short with ventrally a pair of cirri-form antennae, without eyes. Second or tentacular segmentwell fused with prostomium, achaetous, defined by embeddedacicula and ventral cirri. Notopodia very short, with dorsal cir-ri and stout acicula. Neuropodia short cylindrical, with very
short ventral cirri and stout acicula; ventral hooks simple, thin,ocuring more than 30 as maximal number on posterior parapo-dia. Pygidium rounded, without anal cirri.
Biology: Living in the mantle cavity of a deep-sea mussel Ba-thymodiolus japonicus. Foregut with well muscularized part; sto-modeum may be eversible. The species is thought to be para-sitic.
Distribution: Okinawa Trough.
References:
HASHIMOTO J., OHTA S., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. 42: 577-598.MIURA T. & J. HASHIMOTO (1993) Zool. Soc. 10: 169-174.
T. MIURA Denisia 18 (2006): 208
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Shinkai longipedata MIURA & OHTA, 1991
Size: Up to 200 mm in length. Up to 6.0 mm in width.
Color: In alcohol light greenish brown; parapodia darker thantrunk (small individuals pale or colorless).
Morphology: Body long, vermiform, flattened ventrally andarched dorsally, with about 242 segments (excluding preanalachaetous segments); first segment tentacular, chaetigerous.Prostomium with a pair of antennae. Notopodia elongated,with short dorsal cirri and thin acicula. Neuropodia globular,with very short ventral cirri and stout acicula; ventral hooks
simple, stout, 2-10 as maximal number on anterior parapodia oroccuring singly on successing parapodia.
Biology: Living in the mantle of a giant clam species of thegenus Calyptogena. Foregut with well muscularized part; sto-modaeum may be eversible. The species is thought to be para-sitic.
Distribution: Okinawa Trough.
References:
MIURA T. & J. HASHIMOTO (1996) Publ. Seto Mar. Biol. Lab. 37(3/6): 257-274.MIURA T. & S. OHTA (1991) Zool. Soc. 8: 383-387.
T. MIURA Denisia 18 (2006): 209
1: Ventral view of the anterior part.
2: Variations in parapodia and chaetae.
3: Female as found in the clams’ mantle cavity; by courtesyof T. Miura.
4: Male as found in the clams’ mantle cavity; by courtesy ofT. Miura.
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae
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Shinkai semilonga MIURA & HASHIMOTO, 1996
Size: Up to 100 mm in length. Up to 3.9 mm in width.
Color: The female is deep purplish brown on parapodia and themale whitish.
Morphology: Body long, vermiform, flattened ventrally andarched dorsally, with about 267 segments (excluding preanalachaetous segments). Prostomium short, with a pair of short an-tennae, without eyes. Tentacular segment well fused with pros-tomium, with dorsal cirri and ventral cirri, neuroacicula, andnumerous neuropodial hooks. Foregut with strongly muscular-ized part. Pygidium rounded, without anal cirri. Parapodia subbiramous, with dorsal and cirri. Notopodia slight-ly elongated, supported by single, very thin notoacicula; dorsalcirri short, on distal end of notopodia. Neuropodia well devel-oped, supported by single stout neuroacicula; ventral cirri onventro-posterior side of neuropodia, shorter than dorsal cirri.
Neuropodial hooks simple, stout, slightly curved on distal end;number of hooks per parapodium, about 15 on parapodia 1-3,more than 25 on parapodia 4-6 as maximum, then decreasingto reach about ten on parapodium 10, 5-8 on parapodia 50-200.
Remark: A closely related species has been sampled at ManusBack-Arc Basin: Desmos Cauldron (Shinkai 2K # 916). It dif-fers from S. semilonga by the shape of the neuropodial hooks,which have a tridentate apical end.
Biology: Living in the valve cavity of a giant clam Calyptogenasolidissima. The species is thought to be parasitic.
Distribution: Okinawa Trough: Minami-Ensei Knoll.
1A: Anterior end, dorsal view; B: Same, ventral view; C: Same, lateral view; D: Posterior end, dorsal view; E: Neuropodial hooksfrom parapodia 51 (right) and 200 (left); F: Parapodium; from MIURA & HASHIMOTO (1996).
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae
T. MIURA Denisia 18 (2006): 210–211
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2: Shinkaia sp., middle segments dorsal view (SEM); ManusBasin © Ifremer.
3: Shinkaia sp., distal view of posterior hooks (SEM); ManusBasin © Ifremer.
4: Shinkaia sp., tridentate hooks from anterior parapodia(SEM); Manus Basin © Ifremer.
5: Shinkaia sp., tridentate hooks from anterior parapodia(SEM); Manus Basin © Ifremer.
Reference:
MIURA T. & J. HASHIMOTO (1996) Publ. Seto Mar. Biol. Lab. 37(3/6): 257-274.
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Nereis piscesae BLAKE & HILBIG, 1990
Size: A large, robust species, some fragments up to 5 cm long;largest complete specimens 75 mm long, 7 mm wide, for 100segments.
Morphology: Prostomium triangular, narrowing anteriorly torounded tip bearing two short antennae; eyes absent, but twopairs of sunken depressions representing positions of missingeyes present; palps thickened, fleshy, each bearing two finger-like palpostyles. Peristomium enlarged, inflated, surroundingprostomial structures; four pairs of tentacular cirri. Jaws golden,pointed, each with 78 teeth. Paragnaths all conical, arranged asfollows: I – two large cones in vertical row; II – 12 large conesin nearly circular arc; III – 21-22 small and large cones in clus-ter; IV – 19-20 small and large in three distinct clustersarranged in three transverse rows; V – 0; VI – 46 large pointedcones; VII – two small cones; VIII – two groups of two smallcones. Parapodia of middle and posterior segments with uppernotopodial ligule becoming enlarged and flattened, with dis-tinct dorsal curvature and tapering triangular tip, with thick-ened glandular border, bearing thin dorsal cirrus; lower notopo-dial ligule tapering, not enlarged; neuropodial ligule thickened
basally, tapering distally; ventral cirrus narrow, cirriform. No-topodial acicular lobes small in both anterior and posterior seg-ments. Neuropodial acicular lobes conical throughout. Fourtypes of chaetae present. Anterior notochaetae all homogomphspinigers replaced from segments 30-35 by 1-2 heavy ho-mogomph falcigers with elongated blades bearing 6-8 bluntdenticles [not seen in specimens from type locality studied byD. Desbruyères, probably worn falcigers]; upper neurochaetaeincluding heterogomph spinigers and heterogomph falcigers;lower neurosetae include homogomph spinigers and hetero-gomph falcigers; blades of all spinigers with thin denticles oncutting edge; falcigers with thin, pointed denticles; shafts of allchaetae smooth; internally ornamented with dark brown toblack camerations.
Biology: An epifaunal species associated with siboglinids. Thespecies is probably an omnivore, feeding on filamentous bacte-ria.
Distribution: Juan de Fuca Ridge.
1: Anterior end, ventral view (SEM) © Ifremer. 2: Anterior end, lateral view (SEM) © Ifremer.
Annelida, Polychaeta, Phyllodocida, Nereididae
J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 212–213
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Reference:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.
3: Enlarged notopodial ligule of middle segments (SEM) © Ifremer.
4: Anterior notochaetae homogomph spinigers (SEM) © Ifremer.
5: Heterogomph falcigers ofupper neurochaetae (SEM) © Ifremer.
6: Heavy homogomph falciger of poste-rior notopods (SEM) © Ifremer.
7: Lower neurochaetae of posterior segments in-cluding heterogomph falcigers and homogomphspinigers (SEM) © Ifremer.
213
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Nereis sandersi BLAKE, 1985
Size: Largest specimen 95 mm.
Morphology: Parapodia elongated in posterior segments, pro-viding ragged appearance to worm. Body terminated in twolong anal cirri, each with basal swelling. Jaws black to golden,with 10-12 blunt teeth. Paragnaths all conical, arranged as fol-lows: I – 4-6 in vertical cluster; II – about 30 small cones in anarc; III, IV – 85-100 small cones in dense field; V – 0; VI 4-5dark large, curved cones in cluster; VII – two irregular rows ofabout 10 cones each; VIII – 1-2 large colored cones and 3-4cones. Parapodia of segments 1-2 uniramous, following parapo-dia biramous with two black acicula. Four types of chaetae pres-
ent: anterior chaetae homogomph spinigers replaced by 4-6heavily homogomph falcigers. Lower neurochaetae include ho-mogomph spinigers and heterogomph falcigers; blades of allspinigers with thin denticles on cutting edge.
Biology: On smoker’s walls, dwelling inside active/inactive de-posits. Some nereids are carnivorous, others are grazers of bac-terial mats.
Distribution: Galapagos Spreading Center, Northern East Pa-cific Rise, Guaymas Basin.
Reference:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 6: 67-101.
D. DESBRUYÈRES Denisia 18 (2006): 214
Annelida, Polychaeta, Phyllodocida, Nereididae
4: Specimen on a smo-ker wall (EPR 13°N);cruise Phare Ifremer.
1: Anterior part; byV. Martin Ifremer.
2: Anterior part invivo Ifremer.
3: Prostomium, frontal view (SEM) Ifremer.
6: Posterior parapodia elon-gated in vivo Ifremer.
7: Homogomph falcigers(SEM) Ifremer.
5: Everted proboscis showing jawsand paragnaths (SEM) Ifremer.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Eulalia papillosa (BLAKE, 1985)
Reference:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.
J.A. BLAKE Denisia 18 (2006): 215
1A: Anterior end, dorsal view with pharynx everted; B: Left parapodium from posterior region, anterior view; C-D: Compositechaetae; from BLAKE (1985).
Annelida, Polychaeta, Phyllodocida, Phyllodocidae
Size: Up 13 mm long, 0.8 mm wide, for 75 segments.
Color: In alcohol opaque white with brown pigment granuleson antennae, cirri, and body surface.
Morphology: Prostomium pear-shaped, slightly wider than longwith four frontal antennae; with short, clavate median antennaon posterior one-fourth. With four pairs of tentacular cirri onthree segments with following formula: 1 +S1/1 + S1/n; ventraltentacular cirrus of segment 2 short, broad, pointed; other threetentacular cirri long, tapering, cirriform. Dorsal cirri from seg-ment 4 longer than wide, oval-shaped; ventral cirri asymmetri-cal, not pointed, shorter than podial lobe. Parapodia uniramouswith single internal aciculum and 10-12 compound spinigers,
each with shaft bearing single medial tooth and fine lateraldenticles, blade with finely serrated cutting edge. Pygidiumwith two long broad flattened cirri.
Remarks: E. papillosa was originally described as a species ofProtomystides. However, the median antenna was misinterpret-ed as a papilla and when correctly interpreted, the speciesagrees with the definition of Eulalia.
Biology: An epifaunal species associated with Riftia and Calyp-togena. The species probably feeds on small epifaunal inverte-brates.
Distribution: East Pacific Rise at 21°N.
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Galapagomystides aristata BLAKE, 1985
Size: Up to 20 mm long, 1.2 mm wide, with about 90 segments.
Morphology: Color dark brown to light tan, with dark brownspots on prostomium, antennae, tentacular cirri, and dorsalventral cirri. Prostomium wider than long bearing four long,cirriform frontal antennae shifted ventrolaterally. Proboscisnarrow, smooth. Segment 1 dorsally fused to prostomium, freeventrally; segments 2-3 distinct dorsally and ventrally. Threepairs of long, cirriform tentacular cirri with following formula:1 + S1/1 +S0/n. Segment 3 without dorsal cirrus. Dorsal cirri fromsegment 4 long, thickened, fleshy, tapering distally. Ventral cir-ri small, oval subequal to podial lobe. Parapodia uniramous
bearing 4-8 heavy compound chaetae with heavy shaft andthin, aristate-like narrow blade. Pygidium with pair of long,thick cirri.
Biology: An epifaunal species associated with siboglinid tubesand mussels. The species is probably a scavenger, or a possiblepredator on small invertebrates. This worm was proposed to bean hematophagous.
Distribution: Galapagos Spreading Center; East Pacific Rise:13°N, 9°N; common species.
References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.DESBRUYÈRES D., GAILL F., LALLIER L. & Y. FAUQUET (1985) Bull. Biol. Soc. Wash. 8: 103-116.JENKINS C.D., WARD M.E., TURNIPSEED M., OSTERBERG J. & C. VAN DOVER (2002) Invertebr. Biol. 121: 243-254.
J.A. BLAKE Denisia 18 (2006): 216
1: Habitus in vivo (dorsal); by P. Briand Ifremer.
2: Ventral view (SEM) Ifremer.
3: Middle parapodia, right side(SEM) Ifremer.
5: Aristate-like composite chaetae (SEM) Ifremer.
4: Shaft of a composite chaetae (SEM) Ifremer.
Annelida, Polychaeta, Phyllodocida, Phyllodocidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Protomystides verenae BLAKE & HILBIG, 1990
References:
BLAKE J.A. & B. HILBIG, (1990) Pac. Sci. 44: 219-253.TUNNICLIFFE V. (1988) Proc. R. Soc. Lond. B 233: 347-366.
J.A. BLAKE Denisia 18 (2006): 217
1A: Anterior end in dorsal view; B: Right middle parapodium; C-F: Composite chaetae;from BLAKE & HILBIG (1988).
2: Habitus; scale bar 1 mm; by courtesy of V. Tunnicliffe.
Annelida, Polychaeta, Phyllodocida, Phyllodocidae
Size: Up to 27 mm long, 2.1 mm wide, for 75 segments.
Color: Body dark brown to light tan with dark spots on pros-tomium, all cirri, and body segments.
Morphology: Prostomium trapezoidal, broadly rounded anteri-orly, as wide as long, V-shaped posteriorly; with four long, cirri-form antennae, ventral pair shifted ventrolaterally. Anteriorsegments free from one another and prostomium. With threepairs of long, tapering, cirriform tentacular cirri with followingformula: 1 + S1/1 + S1/v. Dorsal cirri from segment 4, each short,oval, thickened, about 2-3 times as long as wide. Dorsal cirri ofanterior segments long, extending well beyond podial lobe;those of middle segments smaller, shifted to higher location onbody wall; ventral cirri shorter than ventral cirri throughout;
both dorsal and ventral cirri with dense concentrations ofbrown pigment spots. Parapodia uniramous bearing single acic-ula and fascicle of 8-12 compound spinigers of two types: (1)large, heavy, brown in color, (2) small, thin, non-colored; bothwith shaft bearing large, pointed terminal tooth and subapicalnotch from which thin blade emerges; tip of shaft with fineteeth along one edge. Pygidium bearing two long, cirriform analcirri.
Biology: The species is likely an epifaunal associate of vesti-mentiferans and other larger invertebrates found at vent sites.The species is probably predatory on small invertebrates.
Distribution: Explorer Ridge and Juan de Fuca Ridge.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Bathykurila guaymasensis PETTIBONE, 1989
Reference:
PETTIBONE M. (1989) Proc. Biol. Soc. Wash. 102(1): 154-168.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 218
1: Dorsal view in vivo; by courtesy of A. Glover.
4: Elytra (SEM); by courtesy of A. Glover.
5: Distal part of sexual male ventralpapillae; (SEM); by courtesy of A.Glover.
2: Ventral view in vivo; by courtesy of A. Glover.
3: Prostomium, dorsal view (SEM); by courtesy of A. Glover.
6: Anterior end, ventral view (SEM); by courtesy of A. Glover.
Annelida, Polychaeta, Phyllodocida, Polynoidae
Size: Up to 8 mm in length.
Color: Pinkish red.
Morphology: Body fusiform, flattened, with 15 segments (firstachaetous). Elytra seven pairs on segments 2, 4, 5, 7, 9, 11, and13, large, oval, covering dorsum, delicate, with larger conical torounded tubercles variable in size on posterior and lateral bor-ders with smaller ones scattered on surface. Prostomium oval,deeply bilobed with triangular anterior continuations bearingsmall frontal filaments. First or tentacular fused to prostomium,not visible dorsally; tentaculophores lateral to prostomium,each with prominent acicular lobe on inner side, withoutchaetae, and pair of subulate tentacular cirri, shorter thanpalps; upper lip large bilobed. Notopodia rounded basally with
projecting acicular lobes on lower side. Neuropodia with coni-cal projecting prechaetal acicular lobe and shorter, roundedpostchaetal lobe. Notochaetae numerous, short to longer form-ing radiating bundles stouter than neurochaetae, with row ofwidely spaced teeth along one side and blunt tips. Neu-rochaetae very numerous forming fan-shaped bundles; neu-rochaetae with two rows of long spines. Pair of long ventralpapillae present in males on segment 11, sometimes extendingfar beyong posterior end. Posterior two segments smaller, en-closed in elytragerous segment 13. Pygidium long with pair ofslender short anal cirri.
Biology: Found also on whale carcasses.
Distribution: Guaymas Basin.
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Branchinotogluma hessleri PETTIBONE, 1985
Synonym: Opisthotrochopodus alvinus PETTIBONE, 1985.
Size: Up to 16 mm.
Morphology: Segments 21. Body flattened, tapering slightlyanteriorly and posteriorly, with parapodia longer than bodywidth. Dorsum with transverse ciliated bands. Elytra 10 pairs.Elytra covering dorsum, round to oval with branched “veins“.Dorsal cirri with short cylindrical cirrophores and long taperedsmooth styles extending beyong tips of chaetae. Groups of ar-borescent branchiae attached on the lateral sides of elytro-phores and dorsal tubercles. Branchiae begining on segment 3as two small groups and becoming larger posteriorly. Prostomi-um bilobed, oval anterior lobes triangular with delicate frontalfilaments. Median antenna with bulbous ceratophore. Palpstout, tapering, smooth. First segment with two tentacular cir-ri. Notopodium with hood or bract encircling small bundle ofchaetae. Everted pharynx with three dorsal and two ventralpapillae, two pairs of prominent jaws with minute denticles onthe inner border. Notopodia with short conical acicular lobesand on elytrigerous segments, with prominent bracts enclosingnotochaetae dorsally and posteriorly. Notochaetae acicularmuch stouter than neurochaetae, smooth or with 1-2 rows ofshort spines. Neurochaetae slender, very numerous, formingfan-shaped bundles. Few upper neurochaetae with more promi-nent spines, with tip flattened and finely spinous. Tips slightlyhooked, with finely spinous hood.In females pygidium consisting of small squarish, to roundedlobes wedged between bases of posteriorly-directed parapodia of
posterior two segments, with pair of anal cirri. Six pairs of smallrounded segmental lamellae near ventral bases of neuropodia ofsegments 11-16.In males (described as Opisthotrochopodus alvinus), segment 12with a greatly extended papilla, parapodia of posterior four seg-ments modified and directed posteriorly differing from one an-other. Parapodia of chaetigerous segment 18 smaller than pre-ceding and following parapodia. Notopodium represented as asmall acicular lobe with a bundle of short smooth acicularchaetae and delicate rounded flattened lamella. Neuropodiumwithout neurochaetae. Modified parapodia of segment 19 with-out chaetae with small elytrophore and elongate-oval elytron.Parapodium 20 greatly modified. Notopodium represented byachaetous elongated cylindrical acicular lobe with distal style ofdorsal cirrus. Neuropodium forming large cyclindrical acicularlobe with projecting conical acicular process and enclosing cir-cle of chaetae of several types. Segment 21 with right and leftparapodia closely approximated medially and directed posteri-orly. Notopodium consisting of expanded thin lamella andthickened acicular lobes without notochaetae. Neuropodiumalso with expanded thin lamella and thickened acicular lobeswith a small bundle of needle-like neurochaetae. Pygidium con-sisting of small ovale lobe without anal cirri.
Biology: Found in Riftia and Calyptogena washes.
Distribution: East Pacific: from Guaymas Basin to East PacificRise 9°N, Galapagos Spreading Center.
1: Habitus in vivo; by P. Briand © Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
2: In vivo, ventral view of a male specimen;by P. Briand © Ifremer.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 219–220
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Reference:
PETTIBONE M.H. (1985) Proc. Biol. Soc. Wash. 98: 447-469.
8: Upper neurochaetae (SEM) © Ifremer.
5: Notochaetae (SEM) © Ifremer.
6: Ventral view of the posterior part of amale specimen showing the elongated papil-la of the segment 12 (SEM) © Ifremer.
7: Modified segments 18-21, ventral view(SEM) © Ifremer.
3: Prostomium, apical view (SEM) © Ifremer.
4: Jaws and proboscis papillae, apical view(SEM) © Ifremer.
9: Parapodium (SEM) © Ifremer.
10: Hooked neurochaetae from chaetiger 20(SEM) © Ifremer.
11: Bract second segment (SEM) © Ifremer.
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Branchinotogluma marianus (PETTIBONE, 1989)
Synonym: Opisthotrochopodus marianus PETTIBONE, 1989.
Size: Up to 6 mm in length. Up to 3 mm in width.
Morphology: Body flattened, with 21 segments (firstachaetous). Dorsum with transverse ciliated bands. Elytra 10pairs on segments 2, 4, 5, 7, 9, 11, 13, 15, 17 and 19. Elytra cov-ering the dorsum, except for posterior modified segments (ely-tra very small on segments 17 and 19). Branchiae delicate, ar-borescent, on lateral bases of elytrophores and dorsal tuberclesand dorsal bases of notopodia, beginning on segment 3 as twosmall groups, becoming larger and continuing to segment 15 asa small group. Muscular pharynx encircled distally by four pairsof papillae, dorsal with two medial ones larger and lateral onessmaller, ventral with four small papillae; two pairs of curvedreddish jaws occupying most of opening, without denticles oninner border. Parapodia 2-16 biramous with notopodia shorterthan neuropodia. Notopodia long, conical with notochaetaeshort smooth, or longer with faint spinuous rows. Larger neu-ropodia with prechaetal lobe long, subconical, with projectingacicular process, postchaetal lobe shorter rounded. Neu-rochaetae with slightly hooked tips. Elongated ventral papillaeon segment 12 extending posteriorly to segment 15. six pairs ofshort, flat ventral lamellae on segments 13-18. Parapodia from
segment 15 more or less modified. Parapodia 16 and followingparapodia without branchiae. Parapodium 17 with a neu-ropodium shorter than ventral cirrus. Parapodium 18 withsmall notopodial acicular lobe fused to cirrophore, conical neu-ropodium shorter than ventral cirrus. Parapodium 19 with asmall elytron, inflated oval notopodium; neuropodium withsmall bundle of long capillary neurochaetae. Parapodium 20 di-rected posteriorly, with small notopodial acicular lobe fused tocirrophore of dorsal cirrus. Notochaetae long, wider subdistallyspinuous, tapering to capillary tips. Parapodium 21 directedposteriorly, with notopodial acicular lobe fused to cirrophore ofthe dorsal cirrus, without notochaetae; neuropodial conical aci-cular lobe with slender capillary neurochaetae and a large ovallateral lamella.
Remark: A closely related species was sampled in North FijiBack-Arc Basin. It differs from PETTIBONE’s (1989) original de-scription by the shape of the muscular pharynx which is encir-cled by four double large papillae instead of four pairs of papil-lae, with two dorsomedial ones larger.
Biology: Female unknown.
Distribution: Mariana Back-Arc Basin.
Annelida, Polychaeta, Phyllodocida, Polynoidae
D. DESBRUYÈRES Denisia 18 (2006): 221–222
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Reference:
PETTIBONE M.H. (1989) Proc. Biol. Soc. Wash. 102(1): 137-153.
1: Muscular pharynx in distal view (SEM) © Ifremer.
2: Posterior chaetigers (ventral view) showing the transformed laterallamella of the 21st segment (SEM) © Ifremer.
3: Ventral papillae of the 12th segment (SEM) © Ifre-mer.
4: Long notochaetae of the 20th segment, wider subdistally spinuous andtapering to capillary tips © Ifremer.
1-4: Branchinotogluma aff. marianus from North Fiji Back-Arc Basin; cruise Starmer.
222
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Branchinotogluma sandersi PETTIBONE, 1985
Synonym: B. grasslei PETTIBONE, 1985.
Size: Up to 60 mm in length.
Color: Pink to light red in living specimens. Golden-coloredchaetae.
Morphology: Body rectangular flattened, tapering slightly an-teriorly and posteriorly. Dorsum with transverse ciliated bands.Elytra 10 pairs, large covering dorsum, round to oval, stiff,opaque. Each segment from 4rd chaetiger with two groups of del-icate arborescent compact branchiae, with terminal filaments.A single group of branchiae on 3rd segment. Prostomiumbilobed, anterior lobes cylindrical with a terminal filament.Median antenna with a short style. Palps stout tapered smooth.Tentaculophores with two pairs of tentacular cirri, dorsal onesas long as palps. Muscular pharynx showing five papillae aroundopening (3 dorsal and two ventral). Jaws denticled in their in-ner border.In males, parapodia of segments 2-18 biramous. Notopodia con-ical with projecting acicular processes, without bracts except onsegment 2. Neuropodia with long conical prechaetal acicularlobes, postchaetal lobes rounded and shorter. Notochaetae
short and acicular, superior neurochaetae with two rows ofprominent spines and a slightly hooked tip, lower neurochaetaewith flattened spinous hooked hood. Parapodia of the posteriorthree segments modified directed posteriorly and enclosed inparapodia of segment 18, all with ventral cirri but branchiaelacking. Parapodia of elytrigerous segment 19 biramous with ra-mi similar in length; notopodia with thickened dorsal bract.Parapodia of segment 20 much smaller and enclosed in parapo-dia of segment 19. Notopodial acicular lobe fused to cirrophoreof dorsal cirrus. Notochaetae few, short, stout, curved with tworows of distal spines. Upper neurochaetae with scattered longspines and rounded tips. Segment 21 similar to 20th but lackingnotochaetae. Four pairs of segmental ventral papillae on seg-ment 12-15. Three pairs of rounded ventral lamellae on seg-ments 16-18.In females (described as B. grasslei) segments 19-21 not trans-formed. With or without five pairs of small squarish segmentalventral papillae near ventral bases of neuropodia 11-15.
Distribution: Galapagos Spreading Center, East Pacific Rise:21°N to 17°S, Guaymas Basin.
Reference:
PETTIBONE M.H. (1985) Proc. Biol. Soc. Wash. 98: 447-469.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 223–224
1: Dorsal view; by P. Briand © Ifremer. 2: Ventral view of the posterior segments, female;by P. Briand © Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
3: Frontal view of the prostomial part (SEM) © Ifremer.
6: Upper neurochaetae of the 20th segment,male (SEM) © Ifremer.
7: Notopodial bract on segment 2 (SEM) © Ifremer.
8: Subacicular neurochaeta (SEM) © Ifremer.
4: Ventral view of the posterior segments, male (SEM) © Ifremer.
5: Ventral view of the segmental papillae, male (SEM) © Ifremer.
224
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Branchinotogluma segonzaci (MIURA & DESBRUYÈRES, 1995)
Size: Up to 49 mm in length. Up to 19 mm in width.
Morphology: Body flattened, with 21 segments (firstachaetous). Elytra 10 pairs on segments 2, 4, 5, 7, 9, 11, 13, 15,17 and 19. Branchiae arborescent, separated into two groups onsegments 3-19, or on segments 3-21 on half specimens. Muscu-lar pharynx encircled distally by four pairs of small papillae,subequal in size, bearing lateral lamellar expansions with fourdistal papillae, and numerous small papillae forming proximalband. Two pairs of jaws without denticulation on the inner bor-der. Dorsal cirri on non-elytrigerous segments with short cylin-drical cirrophores, and short styles with rounded tips, extendingto tips of neurochaetae. Notopodia subconical, with projectingacicular lobe. Neuropodia subconical, deeply notched on upperpart. On males, pair of segmental papillae at the most extend-ing to the base of ventral cirrus on segment 12; five pairs of
squarish ventral lamellae on following segments 13-17; lackingboth papillae on female specimens. On males, parapodium ofsegment 20 modified with reduced notopodium and elongatedneuropodium, lacking notochaetae. Notopodial acicular lobefused to cirrophore with ventral lamellar expansion and em-bedded notoacicula. Neuropodium with lamellar expansionfolding small bundle of chaetae. Parapodium of segment 21strongly modified on males, lacking chaetae. Notopodial acicu-lar lobe fused to cirrophore of the short conical dorsal cirrus,ventral lamellar expansion enlarged.
Biology: Free living polynoids, found on whitish parts of activechimney walls. Active predators of shrimps and bacterial matgrazers.
Distribution: North Fiji, Lau and Manus Back-Arc Basins.
T. MIURA Denisia 18 (2006): 225–226
1: Specimen on active smoker walls; cruise TUIM07, LauBasin; by courtesy of C.R. Fisher.
2: Prostomium in dorsal view (SEM). Notice the proximal band ofpapillae on the pharynx © Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Reference:
MIURA T. & D. DESBRUYÈRES (1995) Proc. Biol. Soc. Wash. 108: 583-595.
3: Second left parapodia in dorsal view, with the typical dor-sal cirrus (SEM) © Ifremer.
4: Ventral view of the left side of the 12th and 13th segmentsin male showing the ventral papilla and the first flat semiovalventral lamella (SEM) © Ifremer.
5: Upper part of the pharynx with two dorsal rounded papil-lae (SEM) © Ifremer.
6: Lower part of the pharynx showing two small papillae(SEM) © Ifremer.
7: Ventral view of the posterior end in male (SEM) © Ifremer. 8: Ventral view of the posterior end in female (SEM) © Ifremer.
226
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Branchinotogluma trifurcus (MIURA & DESBRUYÈRES, 1995)
1: Living specimens dorsal and ventral view; by courtesy of F.Pleijel.
4: Anterior part in dorso-anterior view(SEM) © Ifremer.
5: Ventral view of the posterior part, fe-male (SEM) © Ifremer.
6: Dorsal view of the posterior part, male(SEM) showing the transformed segments© Ifremer.
2: Ventral papillae on segment11, female (SEM) © Ifremer.
3: Trifurcate neurochaetaefrom segment 20, male (SEM)© Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
Reference:
MIURA T. & D. DESBRUYÈRES (1995) Proc. Biol. Soc. Wash. 108: 583-595.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 227
Size: Small species up to 15 mm in length and 7 mm in width.
Color: Bright red in vivo. Preserved specimens pale.
Morphology: Body short with 21 segments, including the ten-tacular segment. Elytra 10 pairs, large imbricated, oval smooth.Dorsal cirri with very long cirriform styles and tapered tips.Branchiae arborescent in two groups, present on segments 3-21(females) or 3-20 (males). Prostomium bilobed. Anterior lobesprominent, without frontal filaments. Tentacular segmentachaetous not distinct dorsally. Muscular pharynx encircled byfour large papillae, subequal in size; two pairs of jaws with den-ticulations on inner border. Notopodia long, conical, with sev-eral notochaetae; neuropodia long, conical, deeply notched onupper part. In males, ventral segmental papillae on segment 12, as long assegment; five pairs of flat semioval ventral segmental lamellae
present on segments 13-17. Segments 18-21 modified with re-duced parapodia. Segment 18 with very long dorsal cirri. Seg-ment 19 with achaetous notopodia bearing broad dorsal lamel-lar expansion. Segment 20 modified strongly, with longachaetous notopodia bearing bifurcate distal end, dorsal digiti-form expansion and ventral fringed lamella, club-shaped neu-ropodia fringed by trifurcate neurochaetae. Segment 21 re-duced in size achaetous, with uniramous parapodia, notopodiadigitiform with very short cirri and basal lamellar expansion.Anal cirri short.Females lack posterior modified parapodia and ventral lamel-lae, but bear ventral papillae on segment 11.
Biology: Very often found in washings of Ifremeria nautilei; noevidence of co mmensalisms.
Distribution: North Fiji and Lau Back-Arc Basins.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Branchinotogluma tunnicliffae (PETTIBONE, 1988)
Synonym: Opisthotrochopodus tunnicliffae PETTIBONE, 1988,Branchinotogluma grasslei sensu PETTIBONE, 1988 (specimensfrom Northeast Pacific).
Size: Up to 31 mm.
Morphology: Segment 21, elytra 10 pairs. Body flattened ven-trally, arched dorsally, with parapodia longer than body width.Elytra large, overlapping covering dorsum. Elytra smooth, stiff,somewhat vaulted around place of attachment to elytrophores.Elytrophores large, projecting posteriorly, with arborescentbranchiae attached on lateral sides. Dorsal tubercules elongate,projecting posteriorly with arborescent branchiae on distalpart. Prostomium bilobed, anterior lobes subtriangular, withlong frontal filaments; median antenna with bulbous cer-atophore in anterior notch, subulate style with long slender tip,shorter than tentacular cirri; palp stout, tapered, smooth. Ten-taculophores lateral to prostomium, without chaetae, each withpair of tentacular cirri, dorsal tentacular cirrus longer than palp,ventral one shorter. Second segment forming posterior lip ofthe mouth, bearing biramous parapodia and first pair ofelytrophores. Notopodia with hood or bract on anterodorsalside encircling small bundle of notochaetae and conical acicu-lar lobe; ventral buccal cirri similar to tentacular cirri longerthan following ventral cirri. Muscular pharynk showing fivepapillae around opening (3 dorsal and 2 ventral); two paires ofjaws minutely denticulated.In females (described as Branchinotogluma grasslei): Biramousparapodia on segments 3-21 similar with smaller notopodium
on anterodorsal side of a large neuropodium. Pygidium consist-ing of small squarish lobe with pair of anal cirri.In males (described as Opisthotrochopodus): Biramous parapodiaon segments 3-17 similar with smaller notopodium on an-terodorsal side of a large neuropodium. Notochaetae stouterthan neurochaetae acicular with two rows of spines. Neu-rochaetae long, slender with slightly hooked tips. Cirrigerousparapodium of segment 18 much smaller, with long dorsal cir-rus projecting posteriorly and single arborescent branchiae.Segment 19 with smaller elytrophores approaching medially,with small parapodia hidden from view dorsally, notopodia withdelicate lamella on dorsal side, notochaetae stout acicular withtwo rows of spines. Segment 20 with parapodia greatly modi-fied. Neuropodia enlarged, inflated with projecting conical aci-cular lobe with neurochaetae projecting from wide opening andventral cirrus attached near the base. Neurochaetae consistingof four stout acicular harpoon chaetae and long slenderchaetae. Neuropodia inflated and extended according to thespecimens (even according to the side of the same specimen).Segment 21 with elongated cirrophores fused medially and tothe notopodial lobe. Ventral segmental papillae long taperingon segment 12, shorter and thicker with slender distal part onsegments 13-15 reduced to a rounded lamellae on 16 and 17.
Distribution: Explorer Ridge; Juan de Fuca Ridge: Endeavoursegment; Axial Seamount and Southern Juan de Fuca Ridge.
Annelida, Polychaeta, Phyllodocida, Polynoidae
1: In situ; by courtesy of S.K. Juniper.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 228–229
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Reference:
PETTIBONE M.H. (1988) Proc. Biol. Soc. Wash. 101: 192-208.
5: Neurochaetae(SEM) © Ifremer.
6: Stout acicular harpoon chaetae ofthe 20th segment (SEM) © Ifremer.
2: Proboscis, left anterior view showing the jaws(SEM) © Ifremer.
3: Posterior part, male (SEM) © Ifremer.
4: Parapodium (SEM) © Ifremer.
229
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Branchiplicatus cupreus PETTIBONE, 1985
Size: 50 mm in length, 15 mm in width.
Morphology: Segments 31-35. Prostomium truncate. No eyes,no lateral antenna. Elytra 12 pairs on segments 2, 4, 5, 7, 9, 11,13, 15, 17, 19, 21 and 23. Up to 12 posterior segments withoutelytra. Elytra large covering dorsum excepted for posterior seg-ments. Elytra stiff, smooth, opaque, with « veins » Posteriorborder of elytra with variable number of small rounded tuber-cules. Elytra surface with minute sensory papillae. Branchiaebegin on segment 3 of unique type, formed of flattened elon-gated sacs, deeply folded and convoluted attached by broadbases to lateral sides of elytrophores and dorsal tubercules and
to dorsal sides of notopodia and dorsal cirrophores. Parapodiabiramous. Notopodia shorter than neuropodia. Notochaetaenumerous forming radiating bundles short to long. Notochaetaemuch stouter than neurochaetae, acicular, smooth with doublerows of spines near distal bare tips. Neurochaetae very numer-ous, slender, forming fan-shaped bundles. Subacicular neu-rochaetae more slender, with shorter spines and slightly hookedbare tips.
Biology: From Riftia and Calyptogena washings.
Distribution: East Pacific Rise: 9°N to 21°N; Guaymas Basin.
Reference:
PETTIBONE M.H. (1985) Proc. Biol. Soc. Wash. 98: 150-157.
D. DESBRUYÈRES Denisia 18 (2006): 230
1: Dorsal view; by P. Briand Ifremer.
3: Elytron (SEM) Ifremer.
2: Prostomium (SEM) Ifremer.
4: Close up of the surface ofthe elytron (SEM) Ifremer.
5: Folded branchiae (SEM) Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
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Branchipolynoe pettibonae MIURA & HASHIMOTO, 1991
Size: Up to 43 mm in length. Up to 19 mm in width.
Morphology: Body flattened, with 21 segments (firstachaetous). Elytra 10 pairs on segments 2, 4, 5, 7, 9, 11, 13, 15,17 and 19. Branchiae well developed, arborescent on segment3-21; separated into upper larger and lower smaller groups. No-topodia very small, digitiform, with a few notochaetae. Neu-ropodia large, long, with slightly bilobed acicular lobe androunded postchaetal lobe. Ventral segmental papillae long; twopairs on segments 11 and 12.
Biology: Living in the mantle cavity of the deep-sea vent mus-sels (Bathymodiolus platyfrons, B. japonicus, B. brevior, B. elonga-tus).
Distribution: Izu Ogasawara Arc, Okinawa Trough, North Fijiand Lau Back Arc Basins.
References:
COSEL R. & B. MÉTIVIER (1994) Veliger 37: 374-392.DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.HASHIMOTO J., OHTA S., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. I 42: 577-598.MIURA T. & J. HASHIMOTO (1991) Proc. Biol. Soc. Wash. 104: 166-174.
T. MIURA Denisia 18 (2006): 231
1: Habitus © Greg Rouse (TUIM06).
2: Anterior part, ventral view froma specimen of Hine Hina, LauBassin Briand/Ifremer.
3: Parapodium, 11th segment,right side, anterior view Briand/Ifremer.
4: 11th and 12th segments,ventral view Briand/Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
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Branchipolynoe seepensis PETTIBONE, 1986
Size: Up to 31 mm in length and 13 mm in width.
Morphology: Body short, spindle-shaped, tapered anteriorlyand posteriorly, flattened ventrally and arched dorsally. 21 seg-ments with 10 pairs of elytra attached on segments 2, 4, 5, 7, 9,11, 13, 15, 17 and 19. Elytra moderatly large, oval, covering lat-eral thirds of body on larger specimens and nearly coveringbody of juveniles. Branchiae well developped, arborescent.Prostomium oval, bilobed, with rounded anterior lobes lackingfrontal filaments. Median antenna lacking distinct cer-atophore. Palp rather short, thick, smooth and tapered. Thickmuscular pharynx, five pairs of small delicate oval papillaearound opening and two pairs of small jaws lacking denticulatebase. Lower neurochaetae slender and having hooked tips withsome longer lateral denticles.
Remark: According to CHEVALDONNÉ et al. (1998) B. seepensisfrom the Gulf of Mexico and B. seepensis from Mid-AtlanticRidge appear to have recently diverged and must be consideredas two isolated phylogenetic species, although no morphologi-cal differences have been found on large series of specimens.
Biology: Commensal within the mantle cavity of mussels. Twoindividuals (male and female) are frequently observed within asingle mussel. Sexual dimorphism with females larger thanmales; females having two pairs of ventral papillae on segments11 and 12. Sex ratio 0.5-0.7: 1. Females contain mature spermand there is evidence of internal fertilization. Inferred mode ofdevelopment lecitotrophic or direct. Asynchronous gametoge-nesis, rapid oogenesis.
Distribution: Hypersaline seeps at the base of Florida Escarp-ment, 26°02’24’’N and 84°54’48’’W; Mid-Atlantic Ridge:Lucky Strike.
Annelida, Polychaeta, Phyllodocida, Polynoidae
2: Tip of upper neurochaetae(SEM) © Ifremer.
1: Proboscis, frontal view (SEM) © Ifremer.
D. DESBRUYÈRES Denisia 18 (2006): 232–233
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
3: Habitus, dorsal view; by A. Fifis © Ifremer.
4: Habitus, ventral view; by A. Fifis © Ifremer.
References:
CHEVALDONNÉ P., JOLLIVET D., FELDMAN R. A., DESBRUYÈRES, d. & LUTZ R. A. & R. C. VRIEJENHOEK (1998) Cah. Biol. Mar. 39: 347-350.DESBRUYÈRES D., ALAYSE A.M., ANTOINE E., BARBIER G., BARRIGA F., BISCOITO M., BRIAND P., BRULPORT J.P., COMTET T., CORNEC L., CRASSOUS P., DANDO P., FABRI M.C.,
FELBECK H., LALLIER F., FIALA-MÈDIONI A., CONÇALVES J., MÈNARD F., KERDONCUFF J., PATCHING J., SALDANHA L. & P.M. SARRADIN (1994) InterRidge News 3(2):18-19.
HOURDEZ S. & C. JOUIN-TOULMOND (1998) Zoomorphology 118: 225-233.HOURDEZ S., LALLIER F.H., GREEN B.N. & A. TOULMOND (1999) Proteins 34: 427-434.JOLLIVET D., COMTET T., CHEVALDONNÉ P., HOURDEZ S. DESBRUYÈRES D. & D. DIXON (1998) Cah. Biol. Mar. 39: 359-362.PETTIBONE M.H. (1986) Proc. Biol. Soc. Wash. 99(3): 444-451.VAN DOVER C.L., TRASK J., GROSS J. & A. KNOWLTON (1999) Mar. Ecol. Progr. Ser. 181: 201-214.
233
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Branchipolynoe symmytilida PETTIBONE, 1984
Size: Up to 50 mm.
Morphology: Scale worm dwelling commensal of Bathymodiolusthermophilus; body flattened, segments 21 first achaetous. Elytraand elytrophores 10 paires on segments 2, 4, 5, 7, 9, 11, 13, 15,17 and 19. Branchiae well developped, arborescent on all para-podia from segment 2. Notopodia small, with few short acicu-lar notochaetae. Neuropodia short truncate, without projectingacicular lobes; with very numerous neurochaetae including up-per stout, slightly hooked ones and very slender ones. Adultswith very small elytra. 0/2 pairs of ventral papillae on segments11 and 12.
Biology: All worms were collected from mussels and werefound occupying the anterior and ventral parts of the mantlecavity. A study of the digestive contents led to hypothesize anutrition based on mussel gills and pseudofaeces.
Distribution: Galapagos Spreading Center, East Pacific Rise(not collected at 21°N and Guaymas Basin).
References:
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.HOURDEZ S., LALLIER F., MARTIN-JEZEQUEL V. WEBER R.E. & A. TOULMOND (1999) Proteins 34: 427-434.PETTIBONE M.H. (1984) Proc. Biol. Soc. Wash. 97: 226-239.
D. DESBRUYÈRES Denisia 18 (2006): 234
1: Habitus; by V. Martin Ifremer.
2: Gill in vivo; by P. Briand Ifremer.
3: Ventral view showing the two pairs ofpapillae; by P. Briand Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Iphionella risensis PETTIBONE, 1986
Size: Up to 11 mm in length, width 6 mm including chaetae.
Color: Reddish.
Morphology: Body ovate greatly flattened. Thirteen pairs ofelytra, 28 segments. The elytra are reniform, their surface cov-ered by polygonal areas enclosing small areolae. Prostomiumbilobed forming to separate rounded lobes, with anterolateralbulbous extensions fused to the facial tubercle. Antennae andeyes lacking. Long cylindrical tentaculophores emerge lateralto the prostomium and palps each with a stout aciculum. Shortpapillate dorsal and ventral tentacular cirri. Small oval medialnodules are found on segments 4 and 5, variable in number andposition among studied specimens. Very numerous notochaetae
are straw-colored and bipinnate feathered, with a slender axisand lateral spines. The upper group of neurochaetae are similarto notochaetae. The rest of neurochaetae are stouter, withclose-set spinous rows on the basal enlarged part.
Biology: Uncommon, in mussels and clams washings. Morecommon among stalked cirripeds.
Distribution: East Pacific Rise: 21°N, 9°N. A posteriorly in-complete, unique specimen was sampled at Mid-AtlanticRidge: Lucky Strike and could correspond to I. risensis or aclosely related species.
References:
PETTIBONE M.H. (1986) Smithson. Contr. Zool. 428: 1-43.VAN DOVER C.L. (2002) Mar. Ecol. Progr. Ser. 230: 137-158.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 235
1: Habitus in vivo Ifremer.
2: Surface of elytron in vivo Ifremer. 3: Prostomium, dorsal view (SEM) Ifremer.
5: Notochaetae (SEM) Ifremer.4: Surface of elytron (SEM) Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Lepidonotopodium atalantae DESBRUYÈRES & HOURDEZ, 2000
Size: Up to 9 mm in length for 23 segments.
Color: Red to pinkish, notopodial chaetae are straw colored,neuropodial chaetae are light brown.
Morphology: The body is short, suboval in outline, flatteneddorsoventrally, slightly tapered and rounded anteriorly and pos-teriorly. The 11 pairs of elytra are located on segments 2, 4, 5,7, 9, 11, 13, 15, 17, 19 and 21. They cover the dorsum. The ely-tra are subreniform, overlapping with 6-12 macrotuberculesraised on the posterior border. The elytral surface appearssmooth but is covered with numerous globular or clavate mi-cropapillae. The surface of both elytrophores and dorsal tuber-cles have bands or tufts of cilia. The prostomium is bilobed, theanterior lobes subtriangular with a frontal filament, lateral an-tennae are absent. The tentacular segment is not visible dorsal-ly. The tentaculophores of this segment are lateral to the pros-tomium and lack chaetae. The facial tubercle is lacking. Seven
pairs of unequal bulbous papillae encircle the opening of theextended pharynx. The three dorsal papillae are pear-shaped,longer than the other ones. The two pairs of hooked jaws areminutely serrated with numerous teeth. The notopodium issubconical with acicular lobes and is enclosed anterodorsally byflaring bracts. The neuropodium is deeply cleft in the upper andlower part. The notochaetae are numerous forming thick radi-ating bundles. The notochaetae have two rows of spines andbare tips. The neurochaetae are numerous; they have two rowsof numerous spines and bare tips. On one third of the speci-mens (males), there are four ventral pairs of elongated papillaeon segments 11-14, extending to the tip of neurochaetae.There is a pair of very long ventral anal cirri.
Distribution: East Pacific Rise: 13°N to 17°S.
1: Habitus in vivo © Ifremer.
2: Ventral view in vivo of thesegments 11-14 showing thesexual elongated papillae © Ifremer.
3: Anterior part in dorsal view (SEM) © Ifremer.
5: Elytron from the middle part of thebody (SEM) © Ifremer.
4: Frontal view of the distal part of the proboscis(SEM) © Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
Reference:
DESBRUYÈRES D. & S. HOURDEZ (2000) Cah. Biol. Mar. 41: 47-54.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 236
6: Distal part of a neuropodial chaeta(SEM) © Ifremer.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Lepidonotopodium fimbriatum PETTIBONE, 1983
Size: Maximum observed 35 mm in length, 18 mm in width.
Color: Living specimens pinkish or rustish to whitish in color,neurochaetae brown to black.
Morphology: Body stout, short, rectangular in outline, flat-tened ventrally, strongly arched dorsally, and slightly taperedand rounded anteriorly and posteriorly. The elytra are thick;leathery, imbricated, covering the dorsum. There are 11 pairesof elytra attached on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19and 21. Except for the first and the last pairs, each elytron hastwo raised smooth macrotubercules on the posterior one-third.The elytra appears smooth but is covered with numerous round
microtubercules. The prostomium is bilobed, the anterior lobessubtriangular, each with a small frontal filament. The biramousparapodia have a shorter notopodia located on the anterodor-sal sides of the longer neuropodia. The distal margins of the no-topodial acicular lobes, notopodial bracts, and neuropodiallobes are fimbriated with slender papillae.
Biology: Mainly observed abundant on smoker walls. Likely acarnivorous species (amphipods observed crushed by the jaws).
Distribution: East Pacific Rise: 21°N to 9°N.
Reference:
PETTIBONE M.H. (1983) Proc. Biol. Soc. Wash. 96(3): 392-399.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 237
3: Specimen on a smoker wall; East Pacific Rise: 13°N, 2630 m. Cruise Phare Ifremer.
2: Prostomium, dorsal view Ifremer.
1: Habitus; by V. Martin Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Lepidonotopodium jouinae DESBRUYÈRES & HOURDEZ, 2000
Size: Maximum observed 14 mm in length and width 7 mm.
Color: Light brown after preservation.
Morphology: Body short, suboval in outline, flatteneddorsoventrally, slightly tapered and rounded anteriorly and pos-teriorly. The elytra are opaque, oval to subreniform, imbricate,covering the dorsum. There are 11 pairs of elytra attached onsegments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19 and 21. The elytra ap-pears smooth to the naked eye. Under SEM, the elytral surfacein the non-overlapping region appears covered with numerousglobular or clavate micropapillae. The prostomium is bilobed,the anterior lobes subtriangular, each with a small frontal fila-ment. Six pairs of unequal pear-shaped papillae and two medi-an papillae encircle the opening of the extended pharynx. Thetwo pairs of dorsal and ventral jaws are minutely serrated with
numerous teeth. The biramous parapodia have a shorter no-topodia located on the anterodorsal sides of the longer neu-ropodia. The notopodium is enclosed by a flaring bract. Theneuropodium is deeply cleft on the upper and lower parts. Thenotochaetae are stouter than neurochaetae. They have tworows of alternating teeth. The dorsal neurochaetae have onerow of spines and straight tips. The ventral ones have two rowsof numerous spines. 0/5 pairs of elongated papillae on segments11-15.
Biology: Mainly observed in washings of mussels. Likely a car-nivorous/omnivorous species.
Distribution: Mid-Atlantic Ridge: Azores Triple Junction,Menez Gwen, and Lucky Strike.
1: Habitus (preserved specimen) Ifremer.
6: Anterior elytron (SEM) Ifremer.
3: Prostomium, fronto-dorsal view(SEM) Ifremer.
7: Clavate micropapilla at the surface ofelytron (SEM) Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
4: Proboscis everted showing fourpairs of jaws with numerous teeth(SEM) © Ifremer.
5: Upper neurochaetae (SEM) Ifremer. 8: Notochaetae (SEM) Ifremer.2: Segments 11-15, ventral view © Ifremer.
Reference:
DESBRUYÈRES D. & S. HOURDEZ (2000) Cah. Biol. Mar. 41: 399-405.
D. DESBRUYÈRES Denisia 18 (2006): 238
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Lepidonotopodium piscesae PETTIBONE, 1988
Size: Maximum observed 29 mm in length, 13 mm in width.
Color: Living specimens brownish to tan in color, elytra cov-ered with bacterial filaments giving them a white color.
Morphology: Body sturdy, elongated oval, rounded anteriorlyand posteriorly, flattened ventrally, arched dorsally. The elytraare large, overlapping, covering the dorsum, thick, stiff, opaque.There are 11 pairs of elytra attached on segments 2, 4, 5, 7, 9,11, 13, 15, 17, 19 and 21. Elytral surface thickly covered withopaque rounded to conical microtubercles, especially thick onsomewhat transverse areas in the middle of elytra and alongborders with scattered globular and filiform micropapillae. Theprostomium is bilobed, the anterior lobes prominent, each witha long frontal filament. The biramous parapodia have a shorternotopodia located on the anterodorsal sides of the longer neu-ropodia. Neuropodium diagonally truncate, with shallow notch
on posterior lower side and deep notch on posterior upper side.Notochaetae with two rows of spines and blunt tapered and“hairy“ tips. Very numerous neurochaetae with slightly hookedtips and fine spiny rows on cutting edge. On some specimens,neuropodia of segment 13 differing by the presence of only 1-2stout, reddish sabre like chaetae. 0/5 elongate papillae on seg-ments 11-15.
Biology: Widely dispersed species, but low in abundance. Ob-served at the periphery of vents, crawling on the rocks, activesulphide and vestimentiferan tubes, grazing on the bacterialmats and protozoans. Trophic generalist or/and carnivorous.
Distribution: Explorer Ridge; Juan de Fuca Ridge: Endeavoursegment, Axial Seamount and Southern Juan de Fuca Ridge;Gorda Ridge.
1: Habitus in vivo Ifremer.
2: Prostomium, dorsal (SEM) Ifremer.
3: Elytron (SEM) Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
D. DESBRUYÈRES Denisia 18 (2006): 239–240
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
References:
PETTIBONE M.H. (1988) Proc. Biol. Soc. Wash. 101: 192-208.SARRAZIN J. & S.K. JUNIPER (1991) Mar. Ecol. Progr. Ser. 185: 1-19.SARRAZIN J., JUNIPER S.K., MASSOTH G. & P. LEGENDRE (1999) Mar. Ecol. Progr. Ser. 190: 89-112.TSURUMI M. & V. TUNNICLIFFE (2003) Deep-Sea Res. I 50: 611-629.TSURUMI M., DE GRAAF R.C. & V. TUNNICLIFFE (2003) J. Mar. Biol. Assoc. U.K. 83: 469-477.
4: Proboscis and jaws (SEM) Ifremer.
5: Elongated ventral papillae on segments 11-15 (SEM) Ifremer.
6: Sabre-like supra acicular neurochaetae on segment 13(SEM) Ifremer.
7: Elytron surface (SEM) Ifremer.
240
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Lepidonotopodium riftense PETTIBONE, 1984
Size: Maximum observed 13 mm in length and 7 mm in width.
Color: No color in preservative.
Morphology: Body short, suboval in outline, flatteneddorsoventrally, slightly tapered and rounded anteriorly and pos-teriorly. The elytra are oval to subreniform, imbricate, coveringthe dorsum. There are 11 pairs of elytra attached on segments2, 4, 5, 7, 9, 11, 13, 15, 17, 19 and 21. The elytra appear smoothwith branching “veins“ emanating from the place of attach-ment to the elytrophores. Near the posterior and lateral bordersof the elytra, scattered micropapillae with short tapered orcylindrical bases. On some specimens from the EPR, the elytra
have a variable number of oval projections near the posteriorborder. The prostomium is bilobed, the anterior lobes subtrian-gular, each with a small frontal filament. The biramous parapo-dia have a shorter notopodia located on the anterodorsal sidesof the longer neuropodia. The neuropodium is diagonally trun-cated and deeply cleft on the upper part. 0/2 pairs of elongatedpapillae on segments 11-12.
Biology: Mainly observed in washings of mussels, clams andtube worms. Likely a carnivorous/omnivorous species.
Distribution: East Pacific Rise: 21°N to 9°N and GalapagosSpreading Center.
1: Habitus (preserved specimen) © Ifremer.
6: Prostomium, dorsal view (SEM) © Ifremer.
7: Two pairs of dorsal and ventral jaws minutelyserrated with numerous teeth (SEM) © Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
References:
PETTIBONE M.H. (1984) Proc. Biol. Soc. Wash. 97: 849-863.VAN DOVER C.L. (2002) Mar. Ecol. Progr. Ser. 230: 137-158.
D. DESBRUYÈRES Denisia 18 (2006): 241
2: Ventral view (preserved speci-men) showing the two pairs ofelongated papillae on segments 11-12 attached basally on theventroposterior sides of theneuropodia. © Briand/Ifremer.
3: Opening of the pharynxencircled by nine pairs ofbulbous papillae © Ifremer.
4: Oval projections of theposterior border of one elytron(SEM) © Ifremer.
5: Globular micropapillae at the surface of elytron (SEM) © Ifremer.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Lepidonotopodium williamsae PETTIBONE, 1984
Size: 26 mm in length, 19 mm in width. Largest up to 36 mmin length.
Color: Brownish to tan.
Morphology: Body short, stout, rectangular in outline, 24-26segments. 11 pairs of elytra attached eccentrically. The elytraare large, imbricate, covering the dorsum. They are thick, stiffopaque; their dorsal surface is nearly covered with rounded toconical microtubercules (inner anterior part) and rounded ele-vations at the distal posterior part. The prostomium is bilobed,the anterior lobes prominent, each with a frontal filament; lat-eral antennae are absent. The palps are cylindrical, smooth.The eyes are lacking. The biramous parapodia have shorter no-
topodia located on the anterodorsal sides of longer neuropodia.The notopodia are subconical with projecting acicular lobeshidden by the numerous notochaetae and enclosed dorsally bywell developed large flaring bracts. The distal margins of thenotopodial acicular lobes, notopodial bracts and neuropodiallobes are fimbriate with slender papillae. 0/2 elongated papillaeventral on segments 11-12.
Biology: Ubiquitous at vents.
Distribution: Galapagos Spreading Center; East Pacific Rise:21°N to 9°N.
References:
DESBRUYÈRES D. & S. HOURDEZ (2000) Cah. Biol. Mar. 41: 47-54.PETTIBONE M.H. (1984) Proc. Biol. Soc. Wash. 97: 849-863.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 242
1: Habitus; by V. Martin Ifremer.
2: Habitus, in vivo Ifremer.
5: Prostomium, dorsal view (SEM) Ifremer.
3: Proboscis, frontal view (SEM) Ifremer.
4: Elytron, posterior border(SEM) Ifremer
Annelida, Polychaeta, Phyllodocida, Polynoidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Levensteiniella iris HOURDEZ & DESBRUYÈRES, 2003
Size: Maximum observed 21 mm in length for 25 segments.
Color: Living specimens transluscent or greenish with gold col-ored chaetae. Elytra and chaetae often coated with mineral de-posits.
Morphology: Body short, tapering anteriorly and posteriorly.There are 11 pairs of elytra (they are deciduous). Elytra overlapanteriorly and posteriorly, and cover the mid-dorsal line. Shortdorsal cirri on the non-elytra bearing segments. Elytra aretranslucent, smooth, oval, and bear posterior, raised, irregular-ly-spaced macrotubercles. The elytral surface appears smooth orslightly covered of small scattered bumps. The cirri are as longas the parapodia, tapering to tips. The prostomium is bilobed,with cephalic peaks and frontal filaments as long as 1/3 of theprostomium length. Lateral antennae are absent and the medi-an antenna is approximately as long as the prostomium, tapersclose the tip and bears a terminal filament. The median anten-na is inserted in the anterior notch of the prostomium and hasa short, spherical ceratophore. Palps are short and thick, ap-proximately 1.5 times the length of the prostomium, and bearterminal filaments. When dissected, the pharynx opening re-vealed seven pairs of papillae, all similar in size. There are twopairs of jaws which do not bear any teeth but the tip is slightly
curved and appears as a parrot beak. The parapodia are bira-mous. Notopodia are shorter in length to the neuropodia andlocated on their anterior-dorsal side. Notopodia are cylindricalin shape, with the bundle of notochaetae radiating on the dor-sal side, close to the tip. The acicular lobe forms a pointed,scoop-like ligule located underneath the notochaetae. Theneuropodia are not cleft in the upper and lower part, but theacicular lobe forms a pointed, scoop-like ligule on the dorsal-anterior side. Notochaetae are stouter than the neurochaetae.Notochaetae are short, curved on one side, with rows of teethappearing as scales on the curved side. They taper to blunt tips.The neurochaetae are numerous, forming a fan-shaped bundleand bear two lateral rows of spines on one side. They areunidentate and and slightly hooked. Ventral cirri are short andtapered, located at the base of the neuropodia. The pygidium issmall, rounded, bearing a pair of anal cirri. All the ventralpapillae are inconspicuous, except of segment 11 where theyare very developed in some of the specimens.
Biology: Among Bathymodiolus azoricus beds.
Distribution: Mid-Atlantic Ridge: Rainbow, Lucky Strike (un-common species).
Reference:
HOURDEZ S. & D. DESBRUYÈRES (2003) Cah. Biol. Mar. 44: 13-21.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 243
1: Habitus, dorsal view Ifremer. 2: Prostomium, dorsal view (SEM) Ifremer.
3: Posterior part of elytron(SEM) Ifremer.
4: Acicular lobe of notochaetae(SEM) Ifremer.
5: Ventral pair papillae11th segment Ifremer.
Annelida, Polychaeta, Phyllodocidae, Polynoidae
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Levensteiniella plicata HOURDEZ & DESBRUYÈRES, 2000
Size: Maximum size observed 57 mm in length, width 21 mmincluding chaetae.
Color: Pinkish in vivo, light brown when preserved. Chaetaeare straw colored.
Morphology: Body long suboval, flattened dorso-ventrally. 11pairs of elytra. The elytra are smooth, thick and with a promi-nent longitudinal fold. They leave uncovered the middle partof the body and the posterior end. The tapering tip of the dor-sal cirrus does not extend beyong the neurochaetae. The pros-tomium is trapezoid, deeply bilobed with frontal terminal fila-ments on the cephalic peaks. The ceratophore of the medianantennae is short, cylindrical, and inserted in the anteriornotch. The style is shorter than the palps, which are stout,shorts and with a terminal articulated filament. The proboscis
has two pairs of straight jaws with up to 20 teeth. Both rami ofthe parapodia well developed; the notopodia shorter than theneuropodia are rounded and the acicula project from acicularlobes. The notochaetae are numerous, stouter than neu-rochaetae. They are straight, flattened, tapering to blunt tipsand with scales on one side. The neurochaetae are slender, nu-merous, forming a fan-shaped bundle. They have two longitu-dinal rows of spines on one side and their tip is bare andstraight. 0/2 elongated papillae on 11 and 12.
Biology: Living among mussel beds of Bathymodiolus ther-mophilus.
Distribution: East Pacific Rise: 9°N.
1: Habitus Ifremer.
2: Posterior part Ifremer.
3: Parapodia, dorsal view Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 244–245
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4: Right parapodium, anterodorsal view (SEM) Ifremer.
5: Prostomium, frontal view; median antenna removed (SEM) Ifremer.
6: Elongated ventralpapillae 11 and 12segments Ifremer.
7: Notochaetae (SEM) Ifremer. 8: Neurochaetae (SEM) Ifremer.
References:
HOURDEZ S. & D. DESBRUYÈRES (2000) Cah. Biol. Mar. 41: 97-102.VAN DOVER C.L. (2002) Mar. Ecol. Progr. Ser. 230: 137-158.
245
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Thermiphione fijiensis MIURA, 1994
Size: Up to 20 mm in length and 10 mm in width.
Morphology: Body flattened, with 30 or 31 segments. Elytra 14pairs on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26 and27; elytra on segment 2 oval, with fringes of short papillae cov-ered with filamentous bacteria; other elytra elongated subreni-form, lateral borders with fringes of short papillae; elytral sur-face covered with hexagonal or polygonal areas with secondaryareolae. Prostomium partially fused to tentacular segment, andwithdrawn in anterior segments. Prostomium bilobed formingseparate rounded lobes with anterolateral extensions. Tentacu-lar segment with long cylindrical tentaculophores lateral toprostomium; each with a single aciculum. Nodular papillae aris-ing from dorsum of anterior segments; more than 20 papillae onanterior segments; arranged in two or more rows before segment
14, thereafter in single row, absent posterior to segment 20.Elytrophores bulbous, transversaly elongated; places of attach-ment with latero-posterior extensions. Dorsal tubercles on cir-rigerous segments bulbous, prominent, transversally elongated,continuous with enlarged cirrophores of dorsal cirri. Dorsal andventral cirri with short clavate papillae. Branchiae absent. No-topodia subconical, with projecting acicular lobes hidden bynumerous notochaetae. Neuropodia large truncate. Hookedneurochaetae occurring on segment 3.
Biology: Free living scale-worm. Epibiotic filamentous bacteriaon parapodia.
Distribution: North Fiji and Lau Back-Arc Basins.
Reference:
MIURA T. (1994) Proc. Biol. Soc. Wash. 107: 532-543.
T. MIURA Denisia 18 (2006): 246
1: Dorsal view; by P. Briand © Ifremer.
3: Anterior (segment 2) andmedian (segment5) elytra with de-tails of the hexag-onal areas andmarginal papillae;from MIURA (1994).
Annelida, Polychaeta, Phyllodocida, Polynoidae
2: Anterior end, dorsal viewelytra removed; from MIURA (1994).
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Thermiphione tufari HARTMANN-SCHRÖDER, 1992
Size: Up to 30 mm in length, width 7.5 mm including chaetae.
Color: Reddish.
Morphology: Body ovate greatly flattened. Fourteen pairs ofelytra (…23, 26, 27), 29-32 segments. The elytra are reniform,their surface are covered by polygonal areas with secondary are-olae; elytrae ciliated at the outer and posterior borders. Pros-tomium bilobed forming two separate rounded lobes, with an-terolateral bulbous extensions fused to the facial tubercle. An-tennae and eyes lacking. Long cylindrical tentaculophoresemerge lateral to the prostomium and palps each with an aci-culum as well as one or two chaetae. Short papillae on dorsaland ventral tentacular cirri. Small oval medial nodules are
found on segments 4 (1), 5 (2) and two following segments (2)but variable in number according to specimens. Segment 3 notvisible dorsally. Elytrophores bulbous, transversally elongated.Dorsal tubercles on cirrigerous segments transversally elongat-ed and striated. Dorsal and ventral cirri with short clavatepapillae. Notochaetae forming radiating bundles of dense tufts,shorter than neurochaetae. Hooked neurochaetae first presentfrom chaetiger 4.
Biology: Rare, found at the periphery of active vents, withstalked barnacles.
Distribution: East Pacific Rise: 7°S to 21°S.
1: Anterior part, dorsal view © Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
2: Proboscis, frontal view (preserved specimen) © Ifremer.
D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 247–248
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
8: Ventral cirrus (SEM) © Ifremer.
4: Elytra withciliated border (SEM) © Ifremer.3: Median elytron © Ifremer.
6: Medial parapodia, dorsal view (SEM) © Ifremer.5: Polygonal area with a central areola (SEM) © Ifremer.
7: Notochaetae (SEM) © Ifremer.
Reference:
HARTMANN-SCHRÖDER G. (1992) Helgol. Meeresunters. 46: 389-403.
248
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Thermopolynoe branchiata MIURA, 1994
1: Habitus in vivo; by courtesy of C. Rouse (cruise TUIM06).4: Prostomium and buccal region, frontal view (SEM) © Ifremer.
2: Elytron surface (SEM) © Ifremer. 5: Inferior neurochaetae (SEM) © Ifremer.
3: Elytron surface detail (SEM) © Ifremer. 6: Superior notochaetae (SEM) © Ifremer.
Annelida, Polychaeta, Phyllodocida, Polynoidae
T. MIURA & D. DESBRUYÈRES Denisia 18 (2006): 249–250
Size: Up to 53 mm in length. Up to 20 mm in width.
Morphology: Body flattened, with 27 segments (firstachaetous). Elytra 11 pairs on segments 2, 4, 5, 7, 9, 11, 13, 15,17, 19 and 21, large, covering dorsum, stiff, rough with numer-ous brownish papillae. Branchiae well developed, arborescenton segment 3-26; separated into two groups on anterior andposterior sides of notopodia on segment 3 and on some less de-veloped posterior segments; forming single large branchial areasencircling central parts of notopodia on other fully developedparapodia. Prostomium bilobed. Anterior lobes prominent,cylindrical, with small frontal filaments. Mouth opening situat-ed between segments 1 and 2. Muscular pharynx encircled by
seven pairs of bulbous papillae. Dorsal and ventral pairs of jawsfused medially each with up to 15 teeth on basal sides. No-topodia subconical with projecting acicular lobes and large flar-ing bracts. Neuropodia diagonally truncate, deeply notched onupper part; distal margins fimbriated with slender papillae cov-ered with numerous filamentous bacteria. Ventral segmentalpapillae long; three pairs present on segments 12, 13 and 14 andlacking in half of the specimens.
Biology: Free living. Found in washings of Ifremeria and mus-sels. Epibiotic bacteria on parapodia.
Distribution: North Fiji, Lau Basin, and Manus Back-ArcBasins.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
7: Middle notochaetae (SEM) © Ifremer.
8: Flaring bracts first chaetigers, frontal view (SEM) © Ifremer.
10: Jaw (dissection) (SEM) © Ifremer.
11: Median elytrophorous chaetiger (SEM) © Ifremer.
12: Median cirriphorous chaetiger (SEM) © Ifremer.
Reference:
MIURA T. (1994) Proc. Biol. Soc. Wash. 107: 532-543.
9: Arborescent branchiae forming large branchial areas (SEM) © Ifremer.
250
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Vampiropolynoe embleyi MARCUS & HOURDEZ, 2002
Reference:
MARCUS J. & S. HOURDEZ (2002) Proc. Biol. Soc. Wash. 111(2): 341-349.
S. HOURDEZ Denisia 18 (2006): 251
4: Ventral view of a partially everted pharynx showing up-per, lateral and ventral lips with their papillae and ridges(SEM); by courtesy of J. Marcus and S. Hourdez.
5: Dorsale view of the anterior part of the worm, showingthe strong aciculae protrudring from the first segment(SEM); by courtesy of J. Marcus & S. Hourdez.
2: Keratinized teeth at the junction between the mouth openingand the pharynx (SEM); by courtesy of J. Marcus and S. Hourdez.
3: Dorsal view of the segments 11-14 (SEM); by courtesy of J. Mar-cus and S. Hourdez.
Annelida, Polychaeta, Phyllodocida, Polynoidae
1: Paratype, dorsalview of the whole
specimen; from MARCUS &HOURDEZ (2002).
Size: Up to 52 mm.
Morphology: Scale worm found on bacterial mats; body flat-tened, segments 43-45 first achaetous. Elytra (smooth) andelytrophores 10 paires on segments 2, 4, 5, 7, 9, 11, 13, 15, 17and 19. Dorsal tubercles developed as gills on cirrigerous seg-ments starting on segment 6. Prostomium bilobed, strong sharpacicular lobes on segment 1. Notochaetae stouter than neu-rochaetae. No jaws, keratinized teeth inside the pharynx.Mouth opening with papillae. No elongated ventral papillae.
Biology: All worms were collected from bare basalt on vent pe-ripheries, after the 1998 eruption of Axial Volcano. The speciesis thought to feed on bacterial mats and may be a pioneerspecies.
Distribution: Juan de Fuca Ridge: Axial Volcano.
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Sphaerosyllis ridgiensis BLAKE & HILBIG, 1990
Size: Small species, holotype 3.1 mm long for 23 chaetigeroussegments, 1 mm wide.
Morphology: Prostomium weakly notched on anterior margin,about as long as wide, clearly fused with tentacular segment;medial antenna inserted at point of folded merger of tentacularsegment with prostomium: lateral antennae inserted nearly inline with medial antenna; antennae glandular, with bulbousbases and narrow tapering tips. Eyes lacking. Tentacular cirripresent laterally on tentacular segment similar to antennae;parapodia similar throughout body, conical with single imbed-ded aciculum lacking papillae; dorsal cirri glandular, with bul-bous bases and narrow tapering tips; without dorsal cirri on
chaetiger 2; ventral cirri long, cirriform. Chaetae including sin-gle, long dorsalmost simple chaetae with bidentate tips and 8-10 compound falcigers with bidentate tips and conspicuous ser-rations on blade. Pharynx red in color, occupying tentacularsegment, and bearing single middorsal tooth sometimes emerg-ing from oral opening; proventriculus occupying chaetigers 1-4,tan colored, with about 20 rows of muscle cells.
Biology: Unknown. The syllids are known to be suctorial-feed-ers.
Distribution: Explorer Ridge, Juan de Fuca Ridge.
References:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.TUNNICLIFFE V. (1988) Proc. R. Soc. London B 233: 347-366.
J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 252
1: Habitus, dorsal view; by courtesy of J. Blake.
3: Median parapodium, dorsal view(SEM) © Ifremer.
2: Anterior part, ventral view(SEM) © Ifremer.
4: Proventricle (SEM)© Ifremer.
5: Compound falcigers with bidentate tips (SEM) © Ifremer.
Annelida, Polychaeta, Phyllodocida, Syllidae
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Laminatubus alvini TEN HOVE & ZIBROWIUS, 1986
6: Population in situ from East Pacific Rise13°N; cruise Phare Ifremer.
7: Two special collarchaetae and onecapillary chaeta below;from TENHOVE & ZIBROVIUS
(1986).
3: Tubes of specimens from East Pacific Rise13°N Ifremer.
Annelida, Polychaeta, Sabellida, Serpulidae
1: Habitus opercu-lum missing; by V. Martin Ifremer.
Size: Tube, until 100 mm long, 5.3 mm wide.
Morphology: The tube is thick-walled, smooth, subtriangularin cross-section. It has a wide flattened area of attachment; thelateral parts are solids, not with the alveolar structure as in. Acrest-like, sometimes undulating medial keel is the only orna-mentation present. The wall of the tube comprises two distinctlayers. The operculum is radially or slightly bilaterally symmet-rical. It consist of a bulbous proximal part, which may be some-what conical, and a rounded to inverted saucer-like distal cap.
Branchial crown about 1/2 of the animal. Branchial radioles ina short spirale. Collar with capilaries and special chaetae, fivepairs of subsequent bundles. Abdomen with up to 109 seg-ments.
Biology: At the border of active vent fields. Filter feeding or-ganisms (mixotrophic?).
Distribution: Galapagos Spreading Center; East Pacific Rise:21°N to 23°S.
4: Geniculated chaetafrom anterior abdomen(SEM) Ifremer.
5: Thoracic uncini(SEM) Ifremer.
2: Habitus in vivo; by P. Briand Ifremer.
References:
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Biol. Soc. Wash. Bull. 6: 103-116.TEN HOVE H.A. & H. ZIBROWIUS (1986) Zool. Scripta 15(1): 21-31.
D. DESBRUYÈRES & M. SEGONZAC Denisia 18 (2006): 253
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Protis hydrothermica TEN HOVE & ZIBROWIUS, 1986
Size: Tube, up to 70 mm long, 3.8 mm wide, the thickness ofthe wall is ca 0.2 mm; length of the body 47 mm, width of thethorax up to 2 mm.
Morphology: The tube is semicircular in cross-section, adher-ing relatively flatly to the substrate. If a flattened area of at-tachment is present, it is very small. The tube is rather indis-tinctive, somewhat rugged. Very careful illumination revealsthat, in addition to the faint medial ridge, there may be a pairof shallow grooves or ridges laterally, making the cross-sectionof the tube slightly trapezoidal. Free anterior tube parts have
not been observed. The wall of the tube is homogenous, anddoes not consist of two distinct layers. The surface is lusterless.Older tube parts are frequently encrusted by yellowish-browndeposits. No operculum.
Distribution: Galapagos Spreading Center; East Pacific Rise:21°N, 13°N, 17°S; Pacific-Antarctic Ridge: 38°S.
Reference:
TEN HOVE H.A. & H. ZIBROWIUS (1986) Zool. Scr. 15(1): 21-3.
H. ZIBROWIUS & M. SEGONZAC Denisia 18 (2006): 254
1 left to right: Specialcollar chaeta, capillarycollar chaeta and capil-lary chaeta; from TEN
HOVE & ZIBROWIUS (1986).
2: Anterior part of tube; from TEN
HOVE & ZIBROWIUS (1986).
4: Specimen removed from tube; East Pacific Rise: 13°N; by D. Desbruyères © Ifremer.
5: Colony of tubes on an artificial substrate left severalmonths on the vent site Parigo, East Pacific Rise: 13°N; cruiseHydronaut; by P. Briand © Ifremer.
3: Tube of specimen onthe capitulum of
stalked banacle Vul-canolepas sp., from EastPacific Rise: 17°S; cruiseBiospeedo; by P. Briand
© Ifremer.
Annelida, Polychaeta, Sabellida, Serpulidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Alaysia spiralis SOUTHWARD, 1991 “small spiral tube-worm“
References:
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.HALANYCH K.M. (2005) Hydrobiologia 535/536: 297-307.KOJIMA S., OHTA S., YAMAMOTO T., YAMAGUCHI T., MIURA T., FUJIWARA T.& J. HASHIMOTO (2003) Mar. Biol. 142: 625-635.SOUTHWARD E.C. (1991) J. Nat. Hist. 25: 859-881.
E.C. SOUTHWARD Denisia 18 (2006): 255
1 left: Anterior region of larger tube; right: Outer lamellae, inner filaments and obturaculum, dorsal view; from SOUTHWARD (1991).
Annelida, Polychaeta, Sabellida, Siboglinidae
Size: Tube length max. 300 mm; diameter 0.7-1.0 mm.
Color: White/grey, semi-transparent.
Morphology: Anterior part of the tube coiled like a corkscrew,rising from a sinuous basal region. Surface smooth, with narrowflanges at irregular intervals. The top of the obturaculum is cup-shaped, with a thin colorless lining, on a stalk which is trian-gular in cross section, having a dorsal groove and lateral flaps.Branchial filaments are parallel to the obturaculum, forminglamellae, composed of two types: one pair of sheath lamellae,composed of filaments without pinnules, surrounds the innerlamellae made of pinnulate filaments.
Biology: Tubes are fixed to rocks close to mussel beds in areasof diffuse venting; temperature about 7°C. Endosymbiotic bac-teria present, probably sulphur-oxidisers.
Distribution: Valu Fa Ridge in the Lau Back-Arc Basin.
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Arcovestia ivanovi SOUTHWARD & GALKIN, 1997
1: Co-dominant species at the DESMOS site (Manus Back-Arc Basin) aregalatheids (Munidopsis lauensis), vestimentiferans (Arcovestia ivanovi) andzoarcids (Pyrolycus manusanus); by courtesy of J. Hashimoto © JAMSTEC.
3: Habitus of two animals in their tube. Specimens from Lau Basin; cruise TUIM 06; by courtesy of Greg Rouse.
2: Anterior end of the tube, obturaculum andplume; specimen from Lau Back-Arc Basin;cruise TUIM 06; by courtesy of Greg Rouse.
Annelida, Polychaeta, Sabellida, Siboglinidae
Size: The sinuous tubes are up to 200 mm long, the anteriorpart is up to 3 mm in diameter.
Color: The semi transparent tube wall is light-tan to olive-green (grey in preserved specimens).
Morphology: The tube has an anterior funnel followed by a se-ries of collars (flanges) diminishing in size toward the posteriorend. The tube surface is slightly ridged, but its general appear-ance is shiny and smooth. The plume is made of slender fila-ments that surround a slender obturaculum with a spoon-shaped top. The obturaculum stalk has a sharp ventral ridge,extending from the base to the apex and a dorsal groove sepa-
rates prominent lateral ridges. The branchial filaments aregrouped in numerous paired lamellae, parallel to the obturacu-lum. The collar at the anterior end of the vestimental region islong, covering the base of the branchial lamellae. The lateralvestimental folds meet at the dorsal mid line, enclosing the dor-sal vestimental space, and join posteriorly to form an entireventral fold. The branchial plume including the obturaculum isslightly longer than the vestimental region. A pair of genitalgrooves runs from the posterior end of the vestimental region tothe base of the anterior collar in mature males.
Distribution: Manus and Lau Back-Arc Basins.
References:
SOUTHWARD E.C. & S.V. GALKIN (1997) J. Nat. Hist. 31: 43-45.KOJIMA S., OHTA S., YAMAMOTO T., MIURA T., FUJIWARA Y., FUJIKURA K. & J. HASHIMOTO (2002) Mar. Biol. 141: 57-64.
D. DESBRUYÈRES Denisia 18 (2006): 256
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Lamellibrachia barhami WEBB, 1969
Size: Tube length max. ca. 1500 mm; anterior diameter 7-12mm.
Color: Grey/white, opaque.
Morphology: Tube rigid, thick walled, sinuous, tapering to <1mm at posterior end; short external flanges anteriorly, erodedand indistinct over much of the tube. Top of obteraculum cup-shaped, smooth interior surface, stalk elliptical in section. Ob-turaculum short (max. 12 mm), about 1/3 length of vestimen-tal region. Branchial filaments are parallel to the obturaculum,forming lamellae of two types: pale outer sheath lamellae (2-5pairs) made of adherent filaments without pinnules and red in-ner lamellae of pinnulate filaments.
Biology: Sprawling tangles of tubes lie on muddy surface, withposterior ends burried in sediment or under rocks. Cold seepanimals live at ambient sea temperature; Middle Valley animalsat fringes of diffuse venting regions with sulphide seepage butno apparent temperature anomaly. Use internal symbiotic sul-phide-oxidizing bacteria.
Distribution: Subduction zone cold seeps on the North Amer-ica continental margin and a sedimented hydrothermal regionat Middle Valley on the Juan de Fuca Ridge.
References:
JONES M.L. (1985) Bull. Biol. Soc. Wash. 6:117-158.JUNIPER S.K., TUNNICLIFFE V. & E.C. SOUTHWARD (1992) Can. J. Zool. 70: 1792-1809.SOUTHWARD E.C., TUNNICLIFFE V., BLACK M.B., DIXON D.R.& L.R.J. DIXON (1996) J. Geol. Soc., Spec. Publ. 118: 211-224.SUESS E., CARSON B., RITGER S.D., MOORE J.C., JONES M.L., KULM L.D.& G.R. COCHRANE (1985) Bull. Biol. Soc. Wash. 6: 475-484.WEBB M. (1969) Bull. Mar. Sci. 19: 18-47.
E.C. SOUTHWARD Denisia 18 (2006): 257
1: Population on the bottom at Middle valley onthe Juan de Fuca Ridge; by courtesy of V. Tunni-cliffe.
4: Anterior region in vivo; bycourtesy of V. Tunnicliffe.
3: Population on the bottom at Middle val-ley on the Juan de Fuca Ridge; by courtesyof V. Tunnicliffe.
Annelida, Polychaeta, Sabellida, Siboglinidae
2: Branchial regionin lateral view; byE. Southward.
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Lamellibrachia columna SOUTHWARD, 1991
1: Left, anterior end of thetube and anterior end of theanimal, dorsal view; fromSOUTHWARD (1991).
2: Anterior end of the animal showing obturaculum andbranchial filaments; cruise TUIM 06; by courtesy of G. Rouse.
3: A tube projecting vertically,Valu Fa Ridge; cruise Biolau ©Ifremer.
Annelida, Polychaeta, Sabellida, Siboglinidae
References:
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.SOUTHWARD E.C. (1991) J. Nat. Hist. 25: 859-881.
E.C. SOUTHWARD Denisia 18 (2006): 258
Size: Tube length max. 820 mm; anterior diameter 14-20 mm.
Color: Tube white/grey, opaque.
Morphology: Tubes smooth surfaced, straight, hick walled,rigid, tapering to sinuous posterior region 10 to <5 mm diame-ter. The white obturaculum (15-42 mm long) has a funnel-shaped top, with thin anterior rim and smooth inner surface, ona slender stalk elliptical in section. Branchial filaments are par-allel to the obturaculum forming lamellae of two types: paleouter sheath lamellae (8-16 pairs) made of adherent filamentswithout pinnules and red inner lamellae composed of pinnulatefilaments.
Biology: The tubes are attached within rock crevices, and proj-ect vertically, near sources of diffuse venting, but probablythere is no temperature anomaly in the surrounding water; in-ternal bacteria present probably sulphide oxidizers.
Distribution: Valu Fa Ridge in the Lau Back-Arc Basin.
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Lamellibrachia satsuma MIURA, TSUKAHARA & HASHIMOTO, 1997
1: Anterior end, ventral view, body fixedafter removal from the tube; by T. Miura.
3: Living specimens; by T. Miura.
4: A clump of vestimentiferan tube worms, 82 m deep,Kagoshima Bay; by T. Miura © Jamstec.
2: Uncini of the opisthosome (SEM);by T. Miura.
Annelida, Polychaeta, Sabellida, Siboglinidae
Size: Tube length max. 1400 mm; anterior diameter max. 9mm.
Color: Tube grey/white.
Morphology: Tube rigid, thick walled, tapering to less than 1mm at posterior end; short external flanges anteriorly, erodedand indistinct in posterior half; sinuous but anterior parts morestraight than posterior: Obturaculum cup- or funnel- shape,with smooth interior surface; stalk elliptical in transverse sec-tion, with up to 19 pairs of branchial lamellae hidden by up tofour pairs of peripheral lamellar sheets. Vestimentum of alewith paired ciliated grooves, diverging at anterior ends. Vesti-mental cuticular plaques 35-63 mm in diameter, trophosomalones 51-82 mm.
Biology: This tubeworm species lives in shallow water at depthsof 82-110 m in Kagoshima Bay and of about 300 m in NankaiTrough. The worms make a cluster on sediment bottom. Mostof clusters are less than 5 m in length with about 1 m height,but reaches more than 10 m in length and 3 m in height. Am-bient temperature is constant at about 16°C throughout theyear. Internal bacteria present, sulfide oxidizers, may chiefly usebiogenic hydrogen sulfide rather than volcanic gas.
Distribution: West Pacific: Kagoshima Bay; Nankai Trough.
References:
MIURA T., TSUKAHARA J. & J. HASHIMOTO (1997) Proc. Biol. Soc. Wash. 110: 447-456.MIURA T., NEDACHI M. & J. HASHIMOTO (2002) J. Mar. Biol. Ass. U. K. 82: 537-540.
T. MIURA Denisia 18 (2006): 259
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Oasisia alvinae JONES, 1985
References:
HUNT H.L., METAXAS A., JENNINGS R.M., HALANYCH K.M. & L.S. MULLINEAUX (2004) Deep-Sea Res. I 51: 225-234.JONES, M.L. (1985) Bull. Biol. Soc. Wash. 6: 117-158.JONES M.L. (1988) Oceanol. Acta, Spec. 8: 69-82.
D. DESBRUYÈRES & E.C. SOUTHWARD Denisia 18 (2006): 260
1: Right lateral view of the anteriorregion; by E. Southward.
2 left: Tube; right: Habitus; from East Pacific Rise: 13°N; cruise Biocyarise;by P. Briand © Ifremer.
Annelida, Polychaeta, Sabellida, Siboglinidae
Size: Tube length max. 123 mm; anterior diameter 1.5-2.5 mm.
Morphology: Tube transparent, tapering, flexible, with widelyspaced flanges. Top of obturaculum with conspicuous centralaxial rod carrying 2-3 translucent white saucer-like structures(small specimens found at East Pacific Rise 9°N having 5-6saucers); broad dorsal groove. Branchial filaments in concentriclamellae, most filaments pinnulate, a few without pinnules ondorsal end of basal lamellae. Paired excretory pores openingdorsally at the base of obturaculum. Anterodorsal wings of ves-timentum become sheath at base of obturaculum, continuingventrally, free from obturaculum and vestimentum proper.Opisthosome with as many as 37 segments.
Biology: Found on sites of diffuse venting; occurs with Tevniajerichonana; endosymbiotic sulfide oxidizing bacteria.
Distribution: East Pacific Rise: 21°N to 9°N.
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Ridgeia piscesae JONES, 1985
1: Population in situ (Endeavour segment); by courtesy of S.K. Juniper.
Annelida, Polychaeta, Sabellida, Siboglinidae
Size: Tube length max. 1900 mm; anterior diameter 2-13 mm.
Morphology: Tube very variable, straight or sinuous, stiff orsoft, tapering to 1mm posterior end; anterior funnel and exter-nal flanges present. Color white, grey, gold, brown; commonlytranslucent. Top of obturaculum carries 1-15 brown saucers ona central axial rod; if these are shed, traces of the axial structureremain. Red branchial filaments in concentric lamellae parallelto the obturaculum; most bear pinnules, but there are a fewthin, smooth filaments at the ends of each row.
Remark: R. piscesae was amended by SOUTHWARD et al. (1995)to include R. phaeophiale JONES 1985
Biology: Grows gregariously in clusters at sites of diffuse orwarm venting, temperature range 5-60°C (?). Uses internalsymbiotic sulphide-oxidizing bacteria. This species can evengrow and survive in areas of low diffuse vent flow with very lowplume level exposure to sulfide.
Distribution: Explorer Ridge, Juan de Fuca Ridge, GordaRidge.
References:
BLACK M. B. (1991) M.Sc., University of Victoria.JONES M. L. & S.L. GARDINER (1989) Biol. Bull. 177: 254-276.SARRAZIN J., ROBIGOU V., JUNIPER S.K. & J.R. DELANEY (1997) Mar. Ecol. Progr. Ser. 153: 5-24.SOUTHWARD E.C., TUNNICLIFFE V. & M. BLAKE (1995) Can. J. Zool. 73: 282-285.TUNNICLIFFE V. & S.K. JUNIPER (1990) Progr. Oceanogr. 24: 1-13.TUNNICLIFFE V. (1988) Proc. R. Soc. London B 233: 347-366.URCUYO I., MASSOTH G., JULIAN D. & C.R. FISHER (2003) Deep Sea Res. I 50: 763-780.
E.C. SOUTHWARD Denisia 18 (2006): 261
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Riftia pachyptila JONES, 1981 “giant tube worm“
Size: Individual tubes up to 1.5 m.
Morphology: Four body regions (1) anterior tentacular plumeon obturaculum; winged vestimentum; trunk; segmented poste-rior opisthosome. Plume red, with tentacular lamellae perpen-dicular to axis; paired halves of apical split end forming oper-culum upon withdrawal into tube. Vestimentum with dorsolat-eral flaps overlapping one another ventrally. Opisthosome withvariable number of segments, ending in rounded posterior tip.Anterior segments completely encircled with paired single rowsof chaetae, becoming incomplete posteriorly. Tube white,smooth, flexible, extremely sturdy, essentially cylindrical, basal-ly blind ending. Tube made of a chitin-protein system.
Biology: Forms clusters on rocks in zone of diffuse venting.Feeds only on internal symbiotic sulfide oxidizing bacteria.
Distribution: Galapagos Spreading Center, East Pacific Rise(including Guaymas Basin).
1: Population in situ from East Pacific Rise: 13°N; cruise Phare Ifremer.
Annelida, Polychaeta, Sabellida, Siboglinidae
D. DESBRUYÈRES Denisia 18 (2006): 262–263
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References:
ARP A.J., CHILDRESS J.J. & R.D. VETTER (1987) J. Exp. Biol. 128: 139-158.ARP A.J., DOYLE M.L., DI CERA E. & S.J. GILL (1990) Resp. Physiol. 80: 323-
334.BELKIN S., NELSON D.C. & H.W. JANNASCH (1986) Biol. Bull. 170: 110-121.BLACK M.B., LUTZ R.A. & R.C. VRIJENHOEK (1994) Mar. Biol. 120: 33-39.BRIGHT M., KECKEIS H. & C R. FISHER (2000) Mar. Biol. 136: 621-632.CHILDRESS J.J., ARP A.J. & C.R. FISHER (1984) Mar. Biol. 83: 109-124.CHILDRESS J.J., FISHER C.R., FAVUZZI J.A., KOCHEVAR R.E., SANDERS N.K. & A.M.
ALAYSE (1991) Bio. Bull. 180: 135-153.HAAS F. de, ZAL F., LALLIER F., TOULMOND A. & J.N. LAMY (1996) Proteins 26:
241-256.FELBECK H. (1985) Physiol. Zool. 53: 272-281. GAILL F., HERBAGE D. & L. LEPESCHEUX (1988) Oceanologica Acta Numéro Spé-
cial 8: 155-160.GAILL F., SHILLITO B., MÉNARD F., GOFFINET G. & J.J. CHILDRESS (1997) Mar. Ecol.
Progr. Ser. 148: 135-143.
GARDINER S.L. & M.L. JONES (1985) Trans. Amer. Microsc. Soc. 104: 19-44.GIRGUIS P.R., LEE R.W., DESAULNIERS N., CHILDRESS J.J., POSPESEL M., FELBECK H. &
F. ZAL (2000) Appl. Environ. Microbiol. 66: 2783-2790.GOFFREDI S. K., CHILDRESS J.J., DESAULNIERS N.T. & F.H. Lallier (1997) J. Exp. Bi-
ol. 200: 2609-2616.JONES M.L. (1981) Proc. Biol. Soc. Wash. 93: 1295-1313.MARSH A.G., MULLINEAUX L.S., YOUNG C.M. & D.T. MANAHAN (2001) Nature
411: 77-80.POWELL M.A. & G.N. SOMERO (1983) Science 219: 297-299.SCOTT K.M., FISHER C.R., VODENICHAR J.S., NIX E.R. & E. MINNICH (1994) Physi-
ol. Zool. 67: 617-638.THIEBAUT E., HUTHER X., SHILLITO B., JOLLIVET D. & F. GAILL (2002) Mar. Ecol.
Progr. Ser. 234: 147-157.ZAL F., LALLIER F.H., WALL J.S. & S.N. VINOGRADOV (1996) J. Biol. Chem. 271:
8869-8874.
2: Specimen, total view (left) and detail of anterior body regions, dorsal view (middle), ventral view (right) at EastPacific Rise: 9°N; by M. Bright.
263
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Siphonobrachia lauensis SOUTHWARD, 1991
References:
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.SOUTHWARD E.C. (1991) J. Nat. Hist. 25: 859-881.
E.C. SOUTHWARD Denisia 18 (2006): 264
1 left: Anterior end of thetube; right: Anterior end ofanimal; by E. Southward.
2: Tubes in hydrothermal sediment, close to bacterial mats. Stalked barnacles andgalatheids in background. Biolau cruise © Ifremer.
Annelida, Polychaeta, Sabellida, Siboglinidae
Size: Tube length max. 290 mm; anterior diameter 0.8-1.2 mm.
Color: Tube, transparent yellow anteriorly, then orange browndarkening to opaque brown posteriorly.
Morphology: The tube is straight and stiff, with a funnel at thetop and external collars at irregular intervals, tapering to < 0.7mm posteriorly. No obturaculum. Tentacles (10-21) stuck to-gether side by side to form a hollow cylinder 5-18 mm long.Dark V-shaped bridle on dorsal side of forepart of body, fol-
lowed by two rows of dark crescentic plaques on the anteriortrunk papillae.
Biology: The tubes are partly burried in sediment, projectingvertically, between rocks covered with oxides and bacterialmats. Temperature “not elevated“. Internal bacteria present,probably sulphur-oxidizers.
Distribution: Lau Back-Arc Basin: Valu Fa Ridge.
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Tevnia jerichonana JONES, 1985 ”Jericho worm“
Size: Tube length max. 350 mm; anterior diameter 3-8 mm.
Morphology: Tube sinuous, rigid, tapering to <1 mm posterior-ly; closely spaced external flanges. Color white or gold/brown,opaque. Top of obturaculum has yellow/brown crust and smallaxial lamina; obturaculum is short (length < 2 x diameter);conspicuous dorsal groove. Short red branchial filaments inconcentric lamellae parallel to opteraculum, most bear pin-nules, but a few without pinnules at dorsal ends of rows.
Biology: Uses internal symbiotic sulphide-oxidizing bacteria.Forms cluster on rocks in zone of diffuse venting, temperature5-30°C; an early colonizer of new vent sites.
Distribution: East Pacific Rise: 13°N to 21°S.
References:
EDWARDS D.B. & D.C. NELSON (1991) Appl. Environ. Microbiol. 57: 1082-1088.FUSTEC A., DESBRUYÈRES D. & S.K. JUNIPER (1987) Biol. Oceanogr. 4: 121-164.JOLLIVET D. (1993) Thèse d’Université, Université de Bretagne Occidentale.JONES M.L. (1985) Bull. Biol. Soc. Wash. 6: 117-158.LAUBIER L. & D. DESBRUYÈRES (1984) La Recherche 161: 1506-1517.MCMULLIN E.R., HOURDEZ S., SCHAEFFER S.W. & C.R. FISHER (2003) Symbiosis 34: 1-41.SHANK T.M., FORNARI D.J., VON DAMM K.L., LILLEY M.D., HAYMON R.M. & R.A. LUTZ (1998) Deep-Sea Res. II 45: 465-515.SHILLITO B., LECHAIRE J.P., GOFFINET G. & F. GAILL (1995) in PARSON L., WALKER C.L. & D.R. Dixon (Eds.) Hydrothermal Vents and Processes. Geological Society
Special Publications 87: 295-402.
E.C. SOUTHWARD Denisia 18 (2006): 265
1: Population in situ from East Pacific Rise: 13°N; cruise Hydronaut Ifremer.Upper left: Anterior region; drawing by E. Southward. Box: Tubes.
2: Plume; by courtesy of P. Baston.
Annelida, Polychaeta, Sabellida, Siboglinidae
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Nicomache arwidssoni BLAKE, 1985 “bamboo worm“
Size: A large species, up to 84 mm long, 3 mm wide for 21chaetigers.
Morphology: Color in alcohol light to dark brown, with goldpigment on anterior end and dark, rusty pigment posteriorly;some EPR specimens with glandular areas appearing dark greenor black. Prostomium broadly rounded anteriorly, merging pos-teriorly with arched cephalic keel; cephalic plate lacking,nuchal grooves curved. Segments longer than wide, flaring an-teriorly, but not forming collars. Notochaetae capillaries; longfilamentous capillary notochaetae absent. Neurochaetae of
chaetigers 1-3 with 1-2 blunt-tipped spines; rostrate chaetaefrom chaetiger 4 increasing to 9-10 per fascicle middle seg-ments; hooks with four apical teeth and tuft of bristles belowmain fang. Anus surrounded by funnel bearing about 25 papil-lae.
Biology: An epifaunal species, found in association with mus-sels, clams, siboglinids, and other associations. Maldanids areknown as bulk-ingestors of particles.
Distribution: Galapagos Spreading Center; East Pacific Rise:13°N, 21°N; Mariana Back-Arc Basin (questionable).
References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 6: 67-101.BLAKE J.A. (1991) Proc. Biol. Soc. Wash. 104: 175-180.DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.VAN DOVER C.L. (2002) Mar. Ecol. Prog. Ser. 230: 137-158.
J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 266
1: Habitus, dosalview; by V. Martin Ifremer.
2: Prostomium, lateral view(preserved specimen) Ifremer.
3: Funnel with papillae sur-rounding anus Ifremer.
4: Tube Ifremer.
5: Hooks with tuffs of bristles(SEM) Ifremer.
Annelida, Polychaeta, Scolecida, Maldanidae
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Nicomache venticola BLAKE & HILBIG, 1990
Size: A large species, up to 130 mm long, 2.5 mm wide, for 27-32 segments.
Color: In alcohol greenish gray or light brown, with dark pig-mented areas sometimes apparent on neuropodia.
Morphology: Prostomium rounded on anterior margin, contin-uing posteriorly as arched cephalic keel; cephalic plate lacking;nuchal grooves curved. Anterior segments except first one or 2,longer than wide, flared anteriorly, overlapping preceding seg-ments but not forming collars. Few posterior segments crowded,with swollen parapodial lobes. Parapodia biramous. No-tochaetae simple capillaries with smooth shafts and clear nar-row sheaths; long filamentous capillary notochaetae absent.Neuropodia of chaetigers 1-3 with 4-6 heavy straight acicular
spines. Following neuropodia with 6-8 rostrate hooks in singlerows, increasing to 9-10 in middle segments; hoods with mainfang surmounted by 2-3 teeth and with numerous fibrils belowmain fang; unworn hooks with distinct tuft of fibrils emergingbelow main fang; worn spines lacking tuft of fibrils; rostratehooks with prominent manubrium on shaft. Anus terminal,surrounded by irregular funnel bearing about 22 papillae.
Biology: An epifaunal species, collected from washings of larg-er biota.
Distribution: Juan de Fuca Ridge: Axial Seamount, SouthernJuan de Fuca Ridge; Explorer Ridge; Guaymas Basin.
Reference:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.
J.A. BLAKE Denisia 18 (2006): 267
1: Specimen in vivo; by courtesy of V. Tunnicliffe. 2: Prostomium, lateral view showing arched cephalickeels (SEM) © Ifremer.
3: Neuropodium of chaetiger III with four acicular spines(SEM) © Ifremer.
4: Rostrate hooks of middle segments, with main fangsurmounted by 2-3 teeth and with numerous fibrils belowmain fang (SEM) © Ifremer.
Annelida, Polychaeta, Scolecida, Maldanidae
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Leitoscoloplos pachybranchiatus BLAKE & HILBIG, 1990
1: Prostomium, lateral view (SEM) © Ifremer.
2: Nucchal organ (SEM) © Ifremer.
Annelida, Polychaeta, Scolecida, Orbiniidae
3: Camerated capillaries (SEM) © Ifremer.
4: Lateral view showing transitionbetween thoracic and abdominalregions (SEM) © Ifremer.
5: Lateral view showing abdominalparamodia with thick glandularbranchiae (SEM) © Ifremer.
6: Far abdominal segments, lateralview (SEM) © Ifremer.
Reference:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.
J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 268
Size: Small species, up to 9.0 mm long and 0.64 mm wide for 54chaetigers.
Color: In alcohol tan.
Morphology: Prostomium conical, smoothly rounded along an-terior margin, with small nuchal organs on lateral boundary ofprostomium and peristomium. Peristomium with one or twoachaetous rings depending on size and state of preservation;both rings apparent in smaller specimens, vaguely apparent inlarger specimens. Thorax with nine similar chaetigers. Notopo-dia with thin, cirriform postchaetal lobes; neuropodia, withshort triangular shaped postchaetal lobes, developing thick-ened base in abdominal region; neuropodial lobe of abdominalsegments becoming thicker, blunted, then developing weakbilobed appearance in far abdominal segments. Notochaetaecamerated capillaries; furcate chaetae absent. Thoracic neu-
rochaetae camerated capillaries; abdominal neurochaetae in-cluding 2-3 capillaries and 1-2 thin acicular spines. Branchiaefrom chaetiger 13-15; each thick in cross section, appearingglandular; anterior and posterior abdominal branchiae short,stubby in appearance, with broad base and narrow apex;branchiae from middle abdominal segments, longer, but stillwith broad base. Pygidial segment broadly rounded, with twolateral cirri.
Biology: Exact habitat unknown, collected near an active vent.The species probably burrows into mud and is likely a depositfeeder. Two large eggs measuring approximately 100 mm in di-ameter were observed in one specimen. They were enclosed ina chamber that was located on the dorsum between chaetigers6 and 7.
Distribution: Juan de Fuca Ridge.
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Orbiniella aciculata BLAKE, 1985
1A: Dorsal view; B: Anterior end, dorsal view; C: Acicular spine from notopodium; D: Basal partof capillary notochaetae; from BLAKE (1985).
Annelida, Polychaeta, Scolecida, Orbiniidae
Size: Small, probably no more than 4 mm long, 0.6 mm wide,for 25 segments.
Color: In alcohol opaque white.
Morphology: Body small, wide, not divided into thorax and ab-domen. Prostomium broadly rounded along anterior margin;eyes and nuchal organs absent. Peristomium consisting of twoachaetous rings with first narrower than second. Noto- and
neurochaetae include 1-4 acicular spines and 2-10 long barbedcapillaries; number of chaetae size dependent, smaller speci-mens with fewest chaetae. Pygidium with two short cirri.
Biology: Infaunal, in mud; exact habitat unknown, may not bea true vent species; a deposit feeder.
Distribution: Galapagos Spreading Center hydrothermal ventfields.
Reference:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.
J.A. BLAKE Denisia 18 (2006): 269
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Orbiniella hobsonae BLAKE & HILBIG, 1990
Size: Small, threadlike, up to 4.5 mm long, 0.2 mm wide, for 25chaetigers.
Color: In alcohol opaque white.
Morphology: Prostomium nearly circular in outline, roundedalong anterior margin, with pair of lateral nuchal organs. Firstachaetous peristomial ring reduced, compressed between pros-tomium and larger second ring. All chaetigers similar, withoutapparent abdominal or thoracic regions. Chaetigers 1-4 gener-ally narrower, shorter, more compressed; middle chaetigers larg-er, more elongate; posterior chaetigers becoming compressed
again. Pygidium simple, without lobes or cirri. Parapodia re-duced, without distinct podial or postchaetal lobes; branchiaelacking. Chaetae include 1-2 barbed acicular spines and 2-5bristled capillaries in both noto- and neuropodia; bristles ofcapillaries arranged in uniform transverse rows.
Biology: Unknown, but probably living in crevices containingpockets of soft sediments; a deposit feeder.
Distribution: Juan de Fuca Ridge.
1: Anterior end, dorsolateral view (SEM) © Ifremer. 2: Barbed acicular spines (SEM) © Ifremer.
Annelida, Polychaeta, Scolecida, Orbiniidae
References:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.TUNNICLIFFE V. (1988) Proc. R. Soc. Lond. B 233: 347-366.
J.A. BLAKE Denisia 18 (2006): 270
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Scoloplos ehlersi BLAKE, 1985
1A: Anterior end dorsal view; B: Chaetigers 10-13, dorsal view; C: Left parapodium from abdominal segment, posteriorview; D: Thoracic capillary notochaetae; E: Abdominal flail chaetae; F: Thoracic acicular neurochaetae; from BLAKE (1985).
Annelida, Polychaeta, Scolecida, Orbiniidae
Size: Up to 10 mm long, 0.5 mm wide, for 50+ segments.
Color: In alcohol opaque white, with golden flocculent materi-al observed in gut.
Morphology: Body elongate, cylindrical in cross section; ante-rior nine chaetigers swollen. Thorax consisting of 11chaetigers; transition to abdomen abrupt, denoted by appear-ance of elongate neuropodial cirrus and disappearance of tho-racic neuropodial uncini. Prostomium elongate, pointed anteri-orly; eyes and nuchal organs absent. Parapodia of chaetigers 1-3 without postchaetal lamellae; from chaetiger 4, notopodiawith fingerlike postchaetal lamellae, becoming longer over fol-
lowing segments; abdominal neuropodia elongate, flattened,with ventral lobe in middle chaetigers. Branchiae fromchaetiger 21, continuing to posterior end [late occurrence ofbranchiae unusual]. Thoracic notopodia with fascicles contain-ing numerous camerated capillaries; neuropodia with dense fas-cicles of capillaries and short, ribbed spines.
Biology: Infaunal, in mud; may not be a true vent species; a de-posit feeder.
Distribution: Galapagos Spreading Center, hydrothermal ventfields.
References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.VAN DOVER C.L. (2002) Mar. Ecol. Progr. Ser. 230: 137-158.
J.A. BLAKE Denisia 18 (2006): 271
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Spiochaetopterus SARS, 1853
Size: Unknown, only fragmented specimens.
Color: Dark tan in alcohol with ventral white glandular shieldextending from segment 5-8.
Morphology: Body long, slender, soft; tubes ringed, horny. Pros-tomium small, and enfolded by the truncate buccal segmentwhich bears long grooved palps but no tentacular cirri. Fourthchaetiger with a fan of lanceolate chaetae, a large and longspine with a truncated top, slightly cordate and a small aristatespine. Middle region (very damaged) with numerous segments(>15), with two rami, all similar with a bilobed Y-shaped innerpart and a small unilobed foliaceous outer part. The neuropodiaare unilobed on B1 and bilobed on following segments. The no-
topodia of the posterior region are erected, slightly inflated pos-teriorly.
Remarks: Likely two different species of Spiochaetopterus arepresent at Mid-Atlantic Ridge vent fields. The poor conserva-tion state of the collections does not allow a specific identifica-tion; however, the shape of the specialized chaetae is unique.
Biology: All chaetopterids have highly modified body regionsthat enable them to strain water through mucous bags. Allchaetopterids secrete the tube from glands located on the ven-tral side of the anterior body region.
Distribution: Mid-Atlantic Ridge: TAG and Rainbow.
1: Chaetopterid on the bottom atTAG; cruise Exomar © Ifremer.
4: Dorsal view of the posterior part of a specimencollected at the Rainbow vent field; by P. Briand ©Ifremer.
5: Short and thick aristate chaeta from segmentfour (SEM) © Ifremer.
4: Long and thickslightly cordate spinefrom chaetiger four(SEM) © Ifremer.
2: View of the extremity and themiddle part of the tube, by P.Briand © Ifremer.
3: Lateral view, anterior part of a specimen collected atthe Rainbow vent field; by P. Briand © Ifremer.
Annelida, Polychaeta, Spionida, Chaetopteridae
References:
BHAUD M. (1998) Sarsia 83: 243-263.BHAUD M. (2001) J. Mar. Biol. Ass. U. K. 81: 225-234.BHAUD M. (2003) Sci. Mar. 67(1): 99-105.
D. DESBRUYÈRES Denisia 18 (2006): 272
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Laonice athecata SIGVALDADÓTTIR & DESBRUYÈRES, 2003
Size: Up to 51 mm long.
Color: Greenish in vivo, white when preserved in ethanol.
Morphology: Prostomium broadly triangular with straight an-terior margin, laterally tapering backwards into a carunclereaching to posterior margin of chaetiger four. Small occipitalantennae on level of chaetiger one. Eyes absent. Nuchal organarranged in two rows enclosing caruncle. Peristomium sur-rounding prostomium almost completely, laterally formingprominent lateral wings, partly fused with chaetiger one.Postchaetal notopodial lamellae of chaetiger one reduced, tri-angular, with dorsal projection. Lamellae on followingchaetigers large, foliate, with a dorsal protrusion, extending al-most over dorsum. Lamellae successively becoming smaller, onposteriormost chaetigers ventrally round with dorsal tip.Prechaetal lamellae small. Neuropodial lamellae on chaetiger
one reduced, round ventrally, with dorsal projection. On subse-quent chaetigers lamella larger, dorsally and ventrally evenlyprojected. On posterior chaetigers lamella becoming smallerand less projecting. Interparapodial pouches missing. Branchi-ae from chaetiger two, present on about 30 chaetigers. Branchi-ae smooth, little longer than notopodial lamellae, heavily cili-ated laterally. Chaetae of anterior chaetigers arranged in simplerow, not particularly long, becoming longer and reduced innumber on posterior chaetigers. Neuropodial hooded hooksfrom chaetiger 30-32. Hooks with one pair of teeth over mainfang which is slightly grooved longitudinally. Notopodial hooksabsent. Sabre chaeta from chaetiger 20-23. Chaetae long, thin,with a curved delicate tip. Pygidium with 6-8 short cirri.
Distribution: Mid-Atlantic Ridge: Lucky Strike and Loga-tchev.
1: Anterior part (preserved specimen) Briand/Ifremer.
2: Anterior part (SEM) Ifremer.
3: Neuropodial lamella with hooded hooks, and sabre chaeta(SEM) Ifremer.
4: Hooded hook showing pair of teeth over main fang (SEM) Ifremer.
Annelida, Polychaeta, Spionida, Spionidae
Reference:
SIGVALDADOTTIR E. & D. DESBRUYÈRES (2003) Cah. Biol. Mar. 44: 219-225.
D. DESBRUYÈRES Denisia 18 (2006): 273
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Laubieriellus grasslei MACIOLEK, 1981
1A: Anterior end, dorsolateral view; B: Anterior end, lateral view; C: Ante-rior end, dorsal view; D, E: Pygidium, ventral view; from MACIOLEK (1981).
Annelida, Polychaeta, Spionida, Spionidae
Size: Small, up to 11.5 mm long, 0.75 mm wide, with about 45segments.
Morphology: Prostomium rounded anteriorly, continuing pos-teriorly as caruncle to chaetiger 2-3 (A-C). Peristomium sepa-rated from chaetiger 1, without lateral wings. Branchiaesmooth, apinnate, four pairs, from chaetiger 2. Neuropodia ofanterior segments connected by ventral crests from chaetiger 2;dorsal crests present on several postbranchial chaetigers (A).Notochaetae all capillaries; neurochaetae include capillariesand multidentate hooded hooks from chaetiger 10; ventral
sabre chaetae from chaetiger 10-11. Pygidium with two short,rounded lobes and one long cirrus (D: E).
Biology: A cryptic species, found among mussels, from experi-mental fouling panels, and present with other invertebrateswashed from various collections. The species is likely a surfacedeposit feeder that forms temporary tubes.
Distribution: Galapagos Spreading Center, hydrothermal ventfields.
Reference:
MACIOLEK N.J. (1981) Proc. Biol. Soc. Wash. 94: 826-837.
J.A. BLAKE Denisia 18 (2006): 274
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Lindaspio dibranchiata BLAKE & MACIOLEK, 1992
1A: Anterior end, dorsal view; B: Posterior parapodium, anterior view; C: Modified neuropo-dial spines from anterior chaetiger; D: Neuropodial hooded hooks; E: Notopodial hoodedhooks; from BLAKE & MACIOLEK (1992).
Annelida, Polychaeta, Spionida, Spionidae
Size: A large species, holotype 31 mm long, 4 mm wide, with165 segments.
Color: Brown in alcohol.
Morphology: Anterior end broad, widest between chaetigers12-20, dorsoventrally flattened throughout. Body terminatingin conical pygidial cone, lacking appendages. Prostomium bifidanteriorly, with two thick rounded lobes, extending posteriorlyas short, mounded caruncle to chaetiger 1. Palps short, thick.Chaetiger 1 reduced, lacking notochaetae; following segmentswith well-developed noto- and neuropodia bearing elaboratepre- and postchaetal lamellae encompassing chaetal fascicles.Notopodia of middle and posterior chaetigers with anterior andposterior lamellae, but these shorter, more triangular and onlypartly encompassing chaetae. Dorsal branchiae present fromchaetiger 2, each anterior branchia thickened, extending acrossdorsal midline; after chaetiger 20, becoming very thin, contin-uing to posterior end. Ventral branchiae from about chaetiger20 as swollen protuberance of neuropodium, becoming well de-
veloped by chaetiger 30 and fully developed by chaetiger 40;ventral branchiae broader than dorsal branchiae, not meetingat ventral midline, continuing to posterior end. Notochaetae ofchaetigers 2-4 modified into unique rosette of 9-10 heavyspines; subsequent notochaetae consisting of capillaries untilabout chaetiger 40 where 3-5 unidentate hooded hooks begin;hooks sharply curved with closely adhering hood. Neu-rochaetae of chaetigers 1-5 capillaries; with distinct fascicles of15-20 heavy spines and thin capillaries from chaetigers 6-28;spines replaced by multidentate hooded hooks by aboutchaetiger 45; neuropodial spines distinctly tapered distally,when worn, appearing acicular; neuropodial hooks thinner,more delicate than notopodial hooks.
Biology: A large benthic infaunal species, probably a surfacedeposit feeder.
Distribution: Guaymas Basin, Southern Trough, in sedimentsof hydrothermal mounds.
Reference:
BLAKE J.A. & N.J. MACIOLEK (1992) Proc. Biol. Soc. Wash. 105 (4): 723-732.
J.A. BLAKE Denisia 18 (2006): 275
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Lindaspio southwardorum BLAKE & MACIOLEK, 1992
Annelida, Polychaeta, Spionida, Spionidae
Size: A large species, recorded up to about 160 mm long, 6-7mm wide, for more than 340 segments.
Color: In life pinkish white with pale violet streak down dor-sum; in alcohol light brown.
Morphology: Anterior half of body dorsoventrally flattened,with dorsum becoming rounded posteriorly. Pygidium not ob-served. Prostomium narrow, pear-shaped, flaring anteriorly,forming two broadly swollen lobes, continuing posteriorly asnarrow, folded caruncle to anterior margin of chaetiger 2. Palpsshort, thick. Chaetiger 1 reduced, lacking notochaetae. No-topodia of chaetigers 2-4 modified, dorsally elevated, with pre-and postchaetal lamellae forming cup enclosing cluster of mod-ified spines; notopodia from chaetiger 5 and neuropodia fromchaetiger 2 with well-developed, elaborate pre- and postchaetallamellae enclosing chaetal fascicles; notopodial lamellae even-tually becoming more elongate, somewhat triangular; neuropo-dial lamellae remaining broadly rounded throughout. Dorsalbranchiae first present from chaetiger 2, each relatively shortfor first 45-50 chaetigers, extending only half of distance tomidline, thereafter branchiae becoming thinner, longer, ex-tending to midline. Ventral branchiae from about chaetiger 55as extensions of postsetal lamellae, remaining relatively short,until about chaetiger 100-125, then becoming longer, more
cylindrical; in far posterior segments, nearly reaching ventralmidline, but never as long as dorsal branchiae. Notochaetae ofchaetigers 2-4 modified into cluster of about 20 heavy spines;subsequent notochaetae numerous, thin capillaries; 5-8 hoodedhooks from chaetiger 75; capillaries become heavier and morelimbate in far posterior segments; individual notopodial hooksstrongly curved, with pointed main fang surmounted by severalminute teeth. Anterior neurochaetae include row of 25-30heavy spines, thin companion capillaries, and ventral bundle ofthin capillaries; neuropodial spines each with smooth shaft thattapers abruptly, then continues as fine, pointed tip with fineserrations or bristles sometimes visible along edge; spines pres-ent until about chaetiger 40, then replaced by thin capillaries;neuropodial hooded hooks from about chaetiger 75; each hooksmaller, more delicate than notopodial hook; each hook withseveral minute teeth above main fang.
Biology: A benthic infaunal species living in sediments nearhydrothermal vents; the species is likely a surface deposit feed-er.
Distribution: Juan de Fuca Ridge: Middle Valley Segment, insediment from high heat areas.
J.A. BLAKE Denisia 18 (2006): 276–277
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1A: Anterior end, dorsal view; B: Anterior parapodium anterior view; C: Middle parapodium, anteriorview; D: Posterior parapodium, anterior view; E: Modified anterior neurochetae and accompanyingcapillaries; F-G: Notopoal hooked hooks; H: Neuropodial hooked hooks; from BLAKE & MACIOLEK (1992).
Reference:
BLAKE J.A. & N.J. MACIOLEK (1992) Proc. Biol. Soc. Wash. 105(4): 723-732.
277
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Prionospio sandersi MACIOLEK, 1981
1: Drawings from MACIOLEK (1981). A: Anterior end, dorsolateral view; B:Prostomium, dorsal view; C: Branchia, dorsal view, showing details ofthe wringled surface; D: Ventral sabre chaetae; E, F: Hooded hook; G:Pygidium, lateral view; H. Pygidium, ventral view.
2: Living specimen from East Pacific Rise: 13°N; by P.Briand © Ifremer.
3: Two neuropodia of the middle part of the body inlateral view showing sabre chaetae and multidentatehooks (SEM); East Pacific Rise: 13°N © Ifremer.
Annelida, Polychaeta, Spionida, Spionidae
Size: Small, up to 3.2 mm long, 0.5 mm wide, with about 35segments.
Morphology: Prostomium broadly rounded anteriorly, endingposteriorly at chaetiger 1. Peristomium encompassing prostomi-um laterally and ventrally, not developing lateral wings; fuseddorsally to chaetiger 1. Branchiae from chaetiger 2, nine pairs;each gill broad, robust, appearing wrinkled, but lacking pin-nules; branchiae longest anteriorly, becoming shorter, stubbyposteriorly. Dorsal crests absent. Anterior chaetae capillaries;multidentate hooded hooks from chaetiger 15 in neuropodia,chaetiger 29 in notopodia; ventral sabre chaetae from chaetiger14. Pygidium cup-shaped, with deeply rounded ventral edgeand small dorsal elongation.
Remark: A closely related form of Prionospio sp. has been com-monly sampled on the Northern East Pacific Rise at 13°N and9°N. It differs from Prionospio sandersi by the number ofbranchiae (12-14 pairs instead nine pairs) by the occurrence ofthe sabre chaetae on neuropodia from chaetiger 15 instead of14 and the occurrence of the multidentate hooks on neuropo-dia from chaetiger 17 instead 15 and in notopodia fromchaetiger 23 instead of 29.
Biology: The known specimens were collected from washingsof Riftia pachyptila and might no be a fully developed. Thespecies is likely a surface deposit feeder, scavenging particleswith its short palps.
Distribution: Galapagos Spreading Center, hydrothermal ventfields.
J.A. BLAKE Denisia 18 (2006): 278–279
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4: A microphotograph mosaic, lateralview (SEM); East Pacific Rise: 13°N © Ifremer.
5: A microphotograph mosaic, dorsalview (SEM); East Pacific Rise: 13°N © Ifremer.
Reference:
MACIOLEK N.J. (1981) Proc. Biol. Soc. Wash. 94: 826-837.
279
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Prionospio unilamellata SIGVALDADÓTTIR & DESBRUYÈRES, 2003
Size: Up to 17 mm long and 0.9 mm width for 114 chaetigers.
Morphology: Prostomium broadly triangular, widest at anteriormargin, narrowing into a raising caruncle, reaching to posteri-or delineation of first chaetiger. Anterior margin sometimeswith 3-4 ciliated tubercles. Peristomium partly surroundingprostomium, forming a low collar, not fused to chaetiger one.Pair of indistinct ciliated nuchal organs situated on each side ofcaruncle. Postchaetal notopodial lamella of chaetiger one re-duced, dorsally pointed, free from prechaetal lamella. Onchaetigers 2 to 16-17, a single large notopodial lamella is en-closing chaetae. Postchaetal lamellae on postbranchial seg-ments becoming smaller, separated from prechaetal lamellae,round in form. On anterior chaetigers, postchaetal lamellaewith dorsal tip. On posteriormost chaetigers lamellae dorsallypointed. Prechaetal lamellae small on postbranchial chaetigers.Dorsal crests absent. Neuropodial postchaetal lamellae reducedon first chaetiger, elliptical in shape. Lamellae largest onbranchial chaetigers, successively becoming smaller. Onchaetigers 2-3 lamellae ventrally round, dorsally with a tip, onsubsequent chaetigers becoming more round. On posteriormost
chaetigers lamellae leaflike, dorsally pointed. Prechaetal lamel-lae separated from postchaetal lamellae. Branchiae fromchaetiger two, 15-16 in number. Anterior branchiae slightlylonger than notopodial lamellae, on subsequent chaetigers suc-cessively longer and more slender. Branchiae apinate withdense lateral ciliation. Interparapodial pouches missing. Small,papilla-like, structures can sometimes be observed laterally be-tween neuropodia. These structures start on 2-3 posteriormost,or posterior to, branchial segments and occur on 7-8 segments.Chaetae on anterior chaetigers slender, arranged in dense dou-ble rows. Neuropodial hooded hooks from chaetiger 19-27. In-troduction of hooks moving backwards with increasing size.Hooks with 4-6 secondary teeth over main fang. Notopodialhooks from chaetiger 25-46. Notopodial hooks with longershaft than neuropodial hooks. Sabre chaeta from chaetiger 12-20, one or two per rami. Sabre chaeta distally granulated. Py-gidium with long median cirrus and two rounded lateral flaps.
Distribution: Mid-Atlantic Ridge: Lucky Strike, Rainbow andSnake Pit.
Reference:
SIGVALDADOTTIR E. & D. DESBRUYÈRES (2003) Cah. Biol. Mar. 44: 219-225.
D. DESBRUYÈRES Denisia 18 (2006): 280
1: Habitus (preserved specimen) Ifremer. Box: Distalpart of notopodial hooded hook (SEM) Ifremer.
2: Anterior part (SEM) Ifremer.
Annelida, Polychaeta, Spionida, Spionidae
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Xandaros acanthodes MACIOLEK, 1981
Reference:
MACIOLEK N.J. (1981) Proc. Biol. Soc. Wash. 94: 826-837.
J.A. BLAKE Denisia 18 (2006): 281
1A: Anterior end, dorsal view, palps and several branchiae removed; B: Anterior end, lateral view; C: Distal tip of branchia inlateral view; D: Entire branchia dorsal view; E: Pygidium and last three chaetigers, lateral view; from MACIOLEK (1981).
Annelida, Polychaeta, Spionida, Spionidae
Size: A small species, 6 mm long, 0.3 mm wide, with about 55segments.
Morphology: A unique spionid. Prostomium rounded anterior-ly, without posterior keel or caruncle. Peristomium well devel-oped, but not producing lateral wings. Notochaetae absent onchaetiger 1. Branchiae from chaetiger 4, continuing tochaetiger 10-13; each gill elongate, cylindrical, wrinkled, lack-ing distinct pinnules. Notochaetae capillaries; neurochaetae in-clude capillaries on chaetigers 1-2, replaced with unhooded aci-cular spines on chaetigers 3-10, these in turn becoming thinner,
straighter, then replaced by bidentate hooks with half hoodsfrom chaetiger 16 continuing to posterior end; spines and hood-ed hooks accompanied by 4-5 large and 2-3 thin capillariesthroughout; ventral sabre chaetae absent. Pygidium with twodorsal and two larger ventral lobes.
Biology: Epifaunal, collected among siboglinids and mussels.The species is likely a surface deposit feeder.
Distribution: Galapagos Spreading Center, hydrothermal ventfields.
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Alvinella caudata DESBRUYÈRES & LAUBIER, 1986
Size: Up to 140 mm.
Morphology: Four pairs of lamellate gills; two transformedchaetigerous segments bearing stout spines; body in two parts,the posterior one with enlarged notopods bearing distal digita-tions; bacterial epibiosis.
Biology: Dwelling inside organic tubes in active chimney walls.Temperature ranging from 10-50°C. Seldom observed in situ.Feeding on free bacteria, associated with filamentous bacteriaepibiosis. Gut functional; retractile buccal tentacles; gonochor-ic, sexual dimorphism.
Distribution: East Pacific Rise: 21°N to 17°S (not observed inGuaymas Basin and Galapagos Spreading Center).
References:
ALAYSE-DANET A.M., DESBRUYÈRES D. & F. GAILL (1987) Symbiosis 4: 51-62.CHEVALDONNÉ P. & D. JOLLIVET (1993) Mar. Ecol. Prog. Ser. 95: 251-262.DESBRUYÈRES D. & L. LAUBIER (1980) Oceanol. Acta 3: 267-274.DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.DESBRUYÈRES D. & L. LAUBIER (1986) Can. J. Zool. 64: 2227-2245.GAILL F. & S. HUNT (1991) Rev. Aquat. Sci. 4: 107-137.JOLLIVET D. (1993) Thèse de doctorat de l’Université de Bretagne Occidentale.
D. DESBRUYÈRES Denisia 18 (2006): 282
1: Population in situ (East Pacific Rise: 13°N, 2630 m); Phare cruise Ifremer.
3: “Caudal“ part in dorsal view;by P. Briand Ifremer.
4: “Caudal“ part in ventral view; by P. Briand Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
2: Habitus, ventral view;by V. Martin Ifremer.
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Alvinella pompejana DESBRUYÈRES & LAUBIER, 1980 “Pompei worm“
Size: Up to 150 mm.
Morphology: Tube dwelling polychaetes with four pairs oflamellate branchiae; two transformed chaetigerous segments;body tappering posteriorly, no obvious abdominal part; bacteri-al epibiosis on the dorsum. Retractile buccal tentacles and twospecialized sexual ones.
Remark: HURTADO et al. (2004) found a very significant mito-chondrial divergence between north and south EPR popula-tions, leading to hypothesize the presence of a southern cryptic
species, although no significant morphological differences havebeen detected.
Biology: Dwelling inside organic tubes in active chimney walls.Temperature ranging from 10-80°C (debated?). Feeding on freebacteria, associated with filamentous bacteria epibiosis. Func-tional gut, retractile buccal tentacles, gonochoric, sexual di-morphism.
Distribution: East Pacific Rise: 21°N to 23°S (not observed inGuaymas Basin and Galapagos Spreading Center).
1: Population in situ; East Pacific Rise: 13°N; Cruise Phare Ifremer.
2: Cluster offilamentous bacteriafrom dorsal part ofthe worm (SEM) Ifremer.
3: Transformednotosetae on twoanterior segments Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
References:
CARY C.S., SHANK T. & J. STEIN (1998) Nature 391: 545-546.CHEVALDONNÉ P., JOLLIVET D., VANGRIESHEIM A. & D. DESBRUYÈRES (1997) Limnol. Oceanogr. 42: 67-80.CHEVALDONNÉ P., FISHER C.R., CHILDRESS J.J., DESBRUYÈRES D., JOLLIVET D., ZAL F & A. TOULMOND (2000) Mar. Ecol. Prog. Ser. 208: 293-295.DESBRUYÈRES D. & L. LAUBIER (1980) Oceanol. Acta 3: 267-274.DESBRUYÈRES D., GAILL F., LAUBIER L., PRIEUR D. & G. RAU (1983) Mar. Biol. 75: 201-205.DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.DESBRUYÈRES D. & L. LAUBIER (1986) Can. J. Zool. 64: 2227-2245.DESBRUYÈRES D. & L. LAUBIER (1991) Ophelia Suppl. 5: 31-45.DESBRUYÈRES D., CHEVALDONNÉ P., ALAYSE A.M., JOLLIVET D., LALLIER F.H., JOUIN-TOULMOND C., ZAL F., SARADIN P.M., COSSON R., CAPRAIS J.C., ARNDT C., O‘BRIEN J.,
GUEZENNEC J., HOURDES S., RISO R., GAILL F., LAUBIER L. & A. TOULMOND (1998) Deep-Sea Res. II 45: 383-422.GAILL F., DESBRUYÈRES D. & D. PRIEUR (1987) Microb. Ecol. 13: 129-139.HURTADO L.A., LUTZ R.A. & R.C. VRIJENHOEK (2004) Mol. Ecol. 13: 2603-2615.JOLLIVET D., DIXON L.R.J., DESBRUYÈRES D. & D.R. DIXON (1998) J. Mar. Biol. Ass. U. K. 78: 113-130.LUTHER III G. W., ROZAN T.F., TAILLEFERT M., NUZZIO D.B., DI MEO C., SHANK T.M., LUTZ R.A. & S.C. CARY (2001) Nature 410: 813-816.
D. DESBRUYÈRES Denisia 18 (2006): 283–284
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4: Specimen in vivo; by Dugornay © Ifremer.
284
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Paralvinella (Miralvinella) bactericola DESBRUYÈRES & LAUBIER, 1991
Size: Up to 42 mm for 130 segments.
Morphology: Alvinellid worm with four pairs of pinnatebranchiae. Animals brownish in colour after preservation. Pros-tomium reduced, with a dorsal lobe, having a dorsal incisionand two well developed lateral expansions. Dorsal part of thedorsal cavity with several rows of smooth oral tentacles with aciliated groove, surrounding dorsal organ dorsally. Dorsal organwith a deep median hollow ending ventrally in two stout lobesfrom which arise two large, long and strong pointed tentacles(male), each with a deep longitudinal groove without any cili-ation. Tentacles are often rolled in a spiral. Notopodia of 7th
chaetigerous segment modified with 4-6 stout and slightlycurved acicular hooks instead of capillary chaetae. Notopodiaof chaetigers 8-32 with digitiform lobes. Capillary notochaetae
covered with minute spines. Uncinigerous neuropodial toripresent from chaetiger 35-37 to the end of the body. Unciniger-ous tori increasing slightly in length posteriorly. Uncini bre-viacicular, stout as in other alvinellids, arranged in a single ver-tical row and oriented with the teeth pointing backwards. Py-gidium rounded without appendages.
Biology: The animal seems to live freely in the sediment with-out conspicuous tubes. On living animals, the tentacles extendfar over the bacterial mats and move gently in the moiré water,while the tips of the branchiae reach the surface of the sedi-ment.
Distribution: Guaymas basin.
2: Anterior end, ventro-lateral view (pre-served specimen) © Ifremer.
3: Buccal part of a male specimen showingthe sexual grooved tentacles and dorsalview (SEM) © Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
Reference:
DESBRUYÈRES D. & L. LAUBIER (1991) Ophelia Suppl. 5: 31-45.
D. DESBRUYÈRES Denisia 18 (2006): 285
1: Habitus, lateral view of a male specimen by V. Mar-tin © Ifremer. Upper box, prostomium, dorsal view;center, buccal segment ventral view; bottom,chaetigers 7 and 8, showing digitiform lobes (SEM) © Ifremer.
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Paralvinella (Miralvinella) dela DETINOVA, 1988
Size: Up to 110 mm (170 chaetigerous segments).
Color: Dark red when preserved.
Morphology: Alvinellid worm with four pairs of pinnatebranchiae. Prostomium very reduced, similar to that of P. bac-tericola with a median lobe having a median incision and twovery well developed and bilobed lateral expansions. Numerousgrooved buccal tentacles inserted on two lateral pads. As in P.grasslei, a median hemispherical body is observed ventrally inthe buccal cavity, bordered by two lateral pads. Two long point-ed and grooved peribuccal tentacles are present in males, eachwith a deep longitudinal groove devoid of ciliation. Four stout
and slightly curved acicular spines in the 7th chaetiger.Uncinigerous neuropodial tori from chaetiger 48-52 to the endof the body. No notopodial or anal cirrus.
Remarks: The species is a sister species of P. bactericola fromGuaymas basin and P. hessleri from Western Pacific Back-ArcBasins which were grouped in the subgenus Miralvinella.
Biology: This worm is usually found in tube worm clusters.
Distribution: Juan de Fuca Ridge: Axial Seamount, sites Ashesand Casm.
References:
DESBRUYÈRES D. & L. LAUBIER (1989) Proc. Biol. Soc. Wash. 102: 761-767.DESBRUYÈRES D. & L. LAUBIER (1993) Proc. Biol. Soc. Wash 106: 225-236.DETINOVA N.N. (1988) Zool. Zh. 67: 858-864.DETINOVA N.N. (1989) Trans. Shirshov Instit. Oceanol. 123: 71-80.
D. DESBRUYÈRES Denisia 18 (2006): 286
1: Female specimen from Axial Seamount, frontal view © Ifremer.
2: Living specimens in: “A Bestiary ofthe Endeavour Hot Vents”; by cour-tesy of V. Tunnicliffe.
Annelida, Polychaeta, Terebellida, Alvinellidae
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Paralvinella (Miralvinella) hessleri DESBRUYÈRES & LAUBIER, 1989
Size: Up to 22 mm.
Morphology: Alvinellid worm with four pairs of pinnatebranchiae. Up to 61 segments. Prostomium medially reducedwith a median incision and two well developed lateral lobesventrally enclosing peristomium. Male buccal apparatus com-prising a ventral globular bulky organ, two lateral strong andpointed tentacles bearing a deep groove without ciliation, andmany grooved and ciliated smaller tentacles inserted in twogroups on a quadrilobed upper lip. First two segments (II andIII) achaetous and fused to the first three chaetigerous. The first15-20 chaetigerous segments with notopodia only. First threenotopodia smaller than others and dorsally elevated. Chaetiger4 with a median dorsal expansion which protrude forward. No-topodia from 4 to 13-17 (7 excepted) cylindrical with a dorsal
digitiform lobe bearing two groups of capillary chaetae, onewith short and the other with long. Chaetigerous segment 7strongly modifies lacking cylindrical notopodia but bearing oneach side 4-5 strongly modified acicular hooks directed posteri-orly. Segment 8 with cylindrical notopodium and very strongdigitiform lobe directed forward. Uncinigerous neuropodial torion each segment from segment 15-20 to the end of the body.
Biology: All specimens found in tubes on rocks directly ex-posed to venting water whose temperature was recorded up to25°C. Tubes whitish and corneous in aspect, amoeba like inshape with long anchor filaments. Tube walls thick and multi-layered. Inner surface bearing important bacterial mats.
Distribution: Mariana and Manus Back-Arc Basins.
Reference:
DESBRUYÈRES D. & L. LAUBIER (1989) Proc. Biol. Soc. Wash. 102: 761-767.
D. DESBRUYÈRES Denisia 18 (2006): 287
2: Male buccal apparatus (SEM) Ifremer.
3: Cross cut of a tube wall, show-ing epibiotic bacteria on the innerpart (SEM) Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
1: Habitus;by V. Martin.
Box: Anterior part,dorsal view (SEM)
Ifremer.
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Paralvinella (Nautalvinella) pandorae DESBRUYÈRES & LAUBIER, 1986
Size: Up to 20 mm.
Color: Light brown to pinkish in ethanol.
Morphology: Body tapering posteriorly comprising about 60chaetigerous segments. Seventh chaetigerous segment with no-topodial chaetae deeply modified forming stout hooks directedbackward. Prostomium reduced. Buccal apparatus retractilemade of a median groove with a pointed end. Numerous buccaltentacles grooved and smooth inserted in two groups. The firstthree chaetigerous segments bearing only notopodia withchaetae. The branchial region comprises the first achaetoussegment plus the three following chaetigerous segments. Fourpairs of pinnate gills with lanceolated secondary filaments in
two adjacent lines. Uncinigerous tori from fifth chaetigeroussegment (P. p. p.) or sixth chaetigerous segment (P.p.i.). Nodigitiform lobes.
Biology: Associated with early colonizer siboglinids (e.g. Oa-sisia, Tevnia). Continuous or semi-continuous recruitment (sin-gle mode size-frequency distribution). MCHUGH (1989) hy-pothesized a brooding of larvae by adults.
Distribution: P. p. pandorae is found on North East Pacific:Juan de Fuca Ridge; P. p. irlandei is found at the East PacificRise: 21°N to 20°S.
4: P. p. irlandei (East Pacific Rise: 13°N), ventral view; Box:Median groove (SEM) Ifremer.
2: P. p. irlandei (East Pacific Rise: 13°N), dorsolateral view;Box: Gill (SEM) Briand/Ifremer.
3: P. p. irlandei (East Pacific Rise: 13°N), latero-ventral view;the arrow is pointing at the first uncinigerous torus in thesixth chaetigerous segment Briand/Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
References:
DESBRUYÈRES D. & L. LAUBIER (1986) Can. J. Zool. 64: 2227-2245.LEVESQUE C., JUNIPER K. & J. MARCUS (2003) Mar. Ecol. Progr. Ser. 246: 173-182.MCHUGH D. (1987) M. Sc., University of Victoria.MCHUGH D. (1995) Invertebr. Biol. 114: 161-168.MCHUGH D. (1989) Mar. Biol. 103: 95-106.
D. DESBRUYÈRES Denisia 18 (2006): 288
1: Drawing of P. p. pandorae;by V. Martin Ifremer.
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Paralvinella (Nautalvinella) unidentata DESBRUYÈRES & LAUBIER, 1993
Size: 4.8-11 mm in length for 77-88 chaetigerous segments.
Color: Pale grey-pinkish in ethanol.
Morphology: Prostomium well developed, with oviform shieldshape, clearly separated from buccal segment, with anterior me-dian incision on two third of the length. Buccal apparatus withmany grooved buccal tentacles. Paired sexual tentacles or ven-tral organ not observed in the type series. First 25-30 chaetiger-ous segments with notopodia only. Branchiae four pairs all sim-ilar arranged as funnel-like structure, with strong basal stembearing small secondary filaments and a thin terminal tip de-void of secondary filaments as long as the basal stem. Secondaryfilaments inserted along stem on two opposite areas; each leaf-shaped, strongly flattened with median ciliated area and point-ed tip. Notopodia (from chaetiger 1 to the end of the body - ex-
cepted 7th) cylindrical bearing two groups of capillary chaetae.Notopodia without digitiform lobes. Chaetiger 7 strongly mod-ified bearing two to three straight short acicular notopodialchaetae on each side. Uncinigerous neuropodia from 26-29chaetiger; uncini numerous (20-50 per torus) in single rows,with teeth directed anteriorly. Uncini with only a single maintooth, lacking a secondary tooth. Pygidium rounded with fiveconspicuous rounded papillae.
Biology: In anhydrite close to vent opening and on rocks underAlviniconcha beds in active areas. Tubes horny with apical digi-tations.
Distribution: North Fiji Back-Arc Basin; Lau Back-Arc Basin:Vaï Lili vent field.
References:
DESBRUYÈRES D. & L. LAUBIER (1993) Proc. Biol. Soc. Wash. 106: 225-236.DESBRUYÈRES D., ALAYSE-DANET A. M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.
D. DESBRUYÈRES Denisia 18 (2006): 289–290
1: Habitus; by V. Martin © Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
2: Anterior portion (SEM); by V. Martin © Ifremer.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
3: in vivo; by courtesy of Greg Rouse.
4: Anterior part of a preserved speci-men showing the funnel-like structureof gills. Photo by P. Briand © Ifremer.
6: Funnel like structure of gills (SEM) © Ifremer.
8: Prostomium (SEM) © Ifremer. 9: Uncinigerous torus (SEM) © Ifremer.
7: 7th and 8th parapods, left side of the body (SEM) © Ifremer.
5: Tubes of P. unidentata on a sulphide spire © Ifremer.
290
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Paralvinella (Paralvinella) fijiensis DESBRUYÈRES & LAUBIER, 1993
Size: Up to 25 mm long.
Color: In vivo reddish and yellow gills.
Morphology: Body maggot-shaped in large specimens (50-65 cs). Prostomium reduced medially, with two anterior lobes.Buccal apparatus comprising numerous exertile grooved tenta-cles and two paired sexual tentacles in males ending with threeunequally developed rounded lobes. Notopodia of chaetigeroussegments one and two reduced. Branchiae four pairs, all similar,with a strong basal stem and secondary filaments abundant in-serted on two opposite areas. Notopodia from about 9 cs to 30 cs
bearing dorsal and ventral rounded lobes. Chaetigerous seg-ment 7 bearing three to four stout transformed chaetae.Uncinigerous neuropodial tori present from 12-19 to the end ofthe body. Each uncinus with a main tooth surmounted bysmaller secondary tooth. Pygidium blunt.
Biology: Most specimens were collected on active edifice walls.Tubes are cylindrical and isolated.
Distribution: North Fiji and Lau Back-Arc Basins.
2: Habitus in vivo; Lau Basin, cruise TUIM06; by courtesy of F. Pleijel.
3: Notopodialchaeta (SEM) © Ifremer.
4: Posterioruncinigerous tori(SEM) © Ifremer.
5: Sexual trilobed tenta-cles (SEM) © Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
1: Habitus; by V. Martin;box preserved specimen© Ifremer.
Reference:
DESBRUYÈRES D. & L. LAUBIER (1993) Proc. Biol. Soc. Wash. 106: 225-236.
D. DESBRUYÈRES Denisia 18 (2006): 291
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Paralvinella (Paralvinella) grasslei DESBRUYÈRES & LAUBIER, 1982
Size: Up to 80 mm.
Color: Brownish when preserved, reddish in vivo.
Morphology: Alvinellid worm with four pairs of pinnatebranchiae. One transformed notopod wearing stout acicularspines. Uncinigerous segment from 13-17th chaetigerous seg-ment to the posterior part. Chaetigerous segments up to 110.Gonochoric species; males with two robust ventral peribuccaltentacles ending in three rounded lobes. No bacterial epibiosis.Mucous tubes unconspicuous.
Biology: Worm dwelling both in warm part of active chimneyswhere mineral load is low and on Riftia tubes. Deposit feederfeeding mainly on bacterial mats.
Distribution: East Pacific Rise, Guaymas Basin, GalapagosSpreading Center.
References:
DESBRUYÈRES D. & L. LAUBIER (1982) Proc. Biol. Soc. Wash. 95: 484-494.DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.DESBRUYÈRES D. & L. LAUBIER (1986) Can. J. Zool. 64: 2227-2245.ZAL F., DESBRUYÈRES D. & JOUIN-TOULMOND C. (1994) C R. Acad Sci. Paris, Sciences de la Vie / Life Sciences 317: 42-48.ZAL F. JOLLIVET D., CHEVALDONNÉ P. & D. DESBRUYÈRES (1995) Mar. Biol. 122: 637-648.
D. DESBRUYÈRES Denisia 18 (2006): 292
2: Population in situ on a whitesmoker (East Pacific Rise: 13°N);cruise Phare Ifremer.
3: Anterior part Ifremer.1: Habitus; by V. Martin Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
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Paralvinella (Paralvinella) palmiformis DESBRUYÈRES & LAUBIER, 1986
1: Endeavour segment, P. palmiformis associatedwith Ridgeia piscesae; bycourtesy of S.K. Juniper.Bottom right: Habitus; by V. Martin Ifremer. Topleft: Comparison betweenthe edges of peribuccaltentacles (SEM) in P.grasslei (upper) and P.palmiformis (lower) Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
References:
ALAIN K. OLAGNON M., DESBRUYÈRES D., PAGÉ A., BARBIER G., JUNIPER S.K., QUÉRELLOU J. & M.A. CAMBON-BONAVITA (2002) FEMS Microbiol. Ecol. 42: 463-476.DESBRUYÈRES D. & L. LAUBIER (1986) Can. J. Zool. 64: 2227-2245.JOLLIVET D., DESBRUYÈRES D, LADRAT C. & L. LAUBIER (1995) Mar. Ecol. Progr. Ser. 123: 125-136.JUNIPER S.K. (1988) Oceanol. Acta N° SP. 8: 167-172.MCHUGH D. (1989) Mar. Biol. 103: 95-106.SARRAZIN J. & S.K. JUNIPER (1999) Mar. Ecol. Progr. Ser. 185: 1-19.SARRAZIN J., JUNIPER S.K., MASSOTH G. & P. LEGENDRE (1999) Mar. Ecol. Progr. Ser. 190: 89-112.TAGHON G.L. (1988) Comp. Biochem. Physiol. 91B: 593-596.TUNNICLIFFE V.& A.R. FONTAINE (1987) J. Geophys. Res., B 92: 11303-11314.TUNNICLIFFE V. & S.K. JUNIPER (1990) Progr. Oceanogr. 24: 1-13.
D. DESBRUYÈRES Denisia 18 (2006): 293
Size: Up to 80 mm.
Color: Red or pinkish after preservation in ethanol.
Morphology: Body gradually tapering. Holotype with 118 seg-ments, paratypes 100-118. Prostomium medially reduced. Ven-trally on the peristomium of males, two blind cavities. Buccalapparatus comprising large number of smooth and grooved buc-cal tentacles and in males two robust peribuccal tentacles end-ing in three rounded lobes bordered by composite papillae. Thefirst 20-31 chaetigerous segments with notopodia only.Chaetigerous segment 7 strongly modified.
Biology: Most commonly found with its caudal end coiledaround the distal portion of siboglinid tubes or on the surface ofsulfide mineral deposits covered in a sheath of inorganic par-ticulates accumulated by mucus secretion. Deposit feeder.
Distribution: Gorda Ridge, Explorer Ridge, Juan de FucaRidge.
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Paralvinella (Paralvinella) sulfincola DESBRUYÈRES & LAUBIER, 1993 “sulfide worm“
Size: Up to 80 mm.
Color: Bright red in life and chocolate brown with brownchaetae after preservation in formalin.
Morphology: 54-68 chaetigerous segments. Body regular, taper-ing abruptly in terminal 10 segments. Buccal apparatus com-prising small grooved and ciliated tentacles, one pair of largebasal tubercles and in males two large lateral tentacles and ter-minating in a curved margin. First 24-30 chaetigerous segmentswith notopodia only. Four pairs of branchiae present, all similarstrong, abruptly attenuated. Branchial stem relatively short,with a large number of slender filaments arising in two oppositeranks and bearing ciliated longitudinal line. First 22 notopodia
(excepted 7) bearing a dorsal digitiform lobe. Chaetigerous 7strongly modified with 4-5 very large stout notopodial hooks di-rected posteriorly. Tubes cylindrical translucent and thin.
Biology: Temperatures of 20-80°C have been measured on sur-faces colonized by sulfide worms and its most likely that theworms regularly experience temperatures within the lower partof this range. All the collections containing this species arecoming from active sulfide edifices where the animal occupiesa distinct microhabitat.
Distribution: Explorer Ridge, Juan de Fuca Ridge, GordaRidge.
References:
JUNIPER S.K. (1994) Paper presented at the 4th International Polychaete Conference, Angers, France.SARRAZIN J. & S.K. JUNIPER (1998) Cah. Biol. Mar. 39: 255-258.SARRAZIN J. & S.K. JUNIPER (1999) Mar. Ecol. Progr. Ser. 185: 1-19.TUNNICLIFFE V. & S.K. JUNIPER (1990) Progr. Oceanog. 24: 1-13.TUNNICLIFFE V., DESBRUYÈRES D., JOLLIVET D. & L. LAUBIER (1993) Can. J. Zool. 71: 286-297.
D. DESBRUYÈRES Denisia 18 (2006): 294
3: Population in situ (Endeavour segment); by courtesy of S.K. Juniper.
1: Anterior part (preserved specimen) Ifremer.
Annelida, Polychaeta, Terebellida, Alvinellidae
2: Population in situ (Endeavour seg-ment); by courtesy of S.K. Juniper.
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Amathys lutzi DESBRUYÈRES & LAUBIER, 1996
Size: 1.25-15.7 mm in length, 2.1-6 mm in width.
Color: No color pattern observed on preserved material.
Morphology: 38 chaetigerous segments of which 20 thoracicand 18 abdominal; 17 thoracic uncinigerous segments. Pros-tomium lacking glandular ridge or eyes; no pallae. Buccal ten-tacles smooth, grooved and inserted on a buccal membrane.Four pairs of branchiae, all smooth and similar, regularly atten-uated. Thoracic parapodia without cirri. Each thoracic uncinuswith a single row of four teeth, abdominal uncini with the same
shape as thoracic ones. Abdominal notopodia absent. Pygidiumterminal without circle of papillae or anal cirri. Mucus linedtubes covered with rusty colored mineral particles, mussel pe-riostracum pieces and fragments of byssal threats; sometimes,tube simple covered with gray mud.
Biology: Sorted from mussel washings, or found in tubes at-tached to mussel shell hinges and to sulfide or basaltic rocks.
Distribution: Mid-Atlantic Ridge: Lucky-Strike, Broken Spur,Snake Pit.
References:
COLAÇO A., DEHAIRS F. & D. DESBRUYÈRES (2002) Deep-Sea Res. 49: 395-412.DESBRUYÈRES D. & L. LAUBIER (1996) Proc. Biol. Soc. Wash. 109: 248-255.
D. DESBRUYÈRES Denisia 18 (2006): 295
2: Anterior part, lateral view;by P. Briand Ifremer.
3: Tube from Mid-Atlantic Ridge:Rainbow; by P. Briand Ifremer.
4: Prostomium, frontal view;by P. Briand Ifremer.
5: Thoracic uncini (SEM) Ifremer.
Annelida, Polychaeta, Terebellida, Ampharetidae
1: Habitus; by V. Martin Ifremer.
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Amphisamytha galapagensis ZOTTOLI, 1983
Size: Maximum size in Guaymas individuals 18 mm, smallerelsewhere.
Color: Whitish or grey when preserved. Pinkish to green in vi-vo.
Morphology: Ampharetinae with four pairs of smooth gills, nopallae, first three chaetigerous segments reduced, uncini fromfourth chaetigerous segment continuing posteriorly on 14 tho-racic segments. Uncini avicular with one row of teeth. Buccaltentacles smooth, inserted on a “buccal membrane”, 12-15
abdominal segment. Tube mucus-lined covered with smallchips of volcanic glass or mud.
Biology: Ubiquitous, often found in other invertebrate shells orcarapaces. Deposit feeder.
Distribution: Galapagos Spreading Center, East Pacific Rise,Guaymas Basin, North East Pacific, Lau Back-Arc Basin, NorthFiji Back-Arc Basin, Manus Back-Arc Basin, Okinawa Trough,Mariana Back-Arc Basin. Likely a group of cryptic species (seeCHEVALDONNÉ et al. 2002).
References:
CHEVALDONNÉ P., JOLLIVET D., DESBRUYÈRES D., LUTZ R.A. & R.C. VRIJENHOEK (2002) Cah. Biol. Mar. 43: 367-370.MCHUGH D. & V. TUNNICLIFFE (1994) Mar. Ecol. Prog. Ser. 106: 111-120.ZOTTOLI R. (1983) Proc. Biol. Soc. Wash. 96: 379-391.
D. DESBRUYÈRES Denisia 18 (2006): 296
3: Anterior part, lateral view(SEM) Ifremer.
4: Posterior end (SEM) Ifremer.
2: Specimen in vivo Briand/Ifremer.
5: Buccal membrane and tentacles (SEM) Ifremer.
6: Abdominal segments, lateral view (SEM) Ifremer.
7: Thoracic unici (SEM) Ifremer.
Annelida, Polychaeta, Terebellida, Ampharetidae, Ampharetinae
1: Habitus; by V. Martin Ifremer.
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Bathybdella sawyeri BURRESON, 1981
Annelida, Oligochaeta, Hirudinida, Piscicolidae, Platybdellinae
1: Habitus,reconstruc-tion of diges-tive system,reproductivesystem, bodyshape andsucker shapefrom dorsalaspect; A – anus; C – cropcaecum; I – intestine; LI – lateralinvagination; M – mouth-pore; P – proboscis; PC – post-caeca; R – rectum; T – testisac;from BURRESON
(1981).
3: Leech cocoons of Johanssonia arc-tica on the walking leg of the tan-ner crab, Chionoecetes bairdi, offthe coast of Oregon (USA); scale bar6 mm; by E. Burreson © VIMS.
4: Leech cocoons of Myzobdellalugubris on the carapace of the bluecrab, Callinectes sapidus, in Chesa-peake Bay, Virginia (USA); scale bar6 mm; by E. Burreson © VIMS.
2: Specimen taken onboard, after collectionfrom Southern East Pacific Rise: 17°S, Oasis site;cruise Biospeedo; by S. Hourdez © CNRS.
Size: Maximal 13 mm.
Morphology: The morphology of most specimens correspondswith the original described morphology of B. sawyeri, withsuckers of approximately equal size, but all specimens collectedfrom Cyanagraea praedator have unusually large oral suckersthat were distinctly larger than the caudal sucker. However,based on the internal morphology using serial sections, also thespecimens from C. praedator could be identified as Bathybdellasawyeri. The leech is unpigmented and lacks eyespots and ocel-li, but often appears red in color because of the blood in the gut.The body is smooth and lacks papillae, tubercles or lateral pul-satile vesicles. Internally B. sawyeri is characterized by six pairsof testisacs and a very unusual reproductive system that in-cludes paired lateral invaginations in segment 12 connected viavector tissue to a large bilobed spermatheca.
Biology: Found free-living in washings of bivalves vesicomyidCalyptogena magnifica and mytilid Bathymodiolus thermophilus,bythograeid crabs Bythograea thermydron and Cyanagraeapraedator, tubeworm Riftia pachyptila, and bythitid fish. Little isknown on the biology of Bathybdella sawyeri. No specimenswere collected from fish hosts, but the presence of nucleatedred blood cells in the gut suggest that this leech feeds on theblood of fishes, as do all other members of the family Piscicoli-dae. The high abundance of leeches found among vent inver-tebrates suggests that the leech leaves the fish host after eachblood meal and seeks refuge among various invertebrates untildigestion is complete and the leech is ready to feed again. Al-though cocoons (egg cases) have not been found, B. sawyeri un-doubtedly deposits cocoons on available hard substrate includ-ing shells or carapaces of invertebrates.
Distribution: Galapagos Spreading Center; Southern East Pa-cific Rise: 14°S, 17°S.
References:
BURRESON E.M. (1981) Proc. Biol. Soc. Wash. 94(2): 483-491.BURRESON E.M. & M. SEGONZAC (submitted) Zootaxa.
E.M. BURRESON & M. SEGONZAC Denisia 18 (2006): 297
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Copidognathus papillatus KRANTZ, 1982
Arthropoda, Chelicerata, Arachnida, Acariformes, Halacaridae
1: Habitus; by I. Bartsch.
Size: 500-570 µm (from tip of rostrum to end of anal papillae).
Morphology: Marine mite with ovate idiosoma, slendergnathosoma with almost parallel-sided rostrum, two pairs oflegs directed forward and two pairs backward. Dorsal plates ofidiosoma with areolate-reticulate sculpturing. Anal papillae en-larged, extending beyond anal aperture. Legs with aciculatereticulation; they lack long spine-like setae.
Biology: Present within detritus among colonies of vestimen-tiferan tubeworms, barnacles, and mussels.
Distribution: Galapagos Spreading Center, East Pacific Rise:13°N, Juan de Fuca Ridge: Main Endeavour; North Fiji and LauBack-Arc Basins.
References:
BARTSCH I. (1991) Zool. Sci. 8: 789-792.KRANTZ G.W. (1982) Can. J. Zool. 6: 1728-1731.
I. BARTSCH Denisia 18 (2006): 299
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Halacarellus auzendei (BARTSCH, 1990)
References:
BARTSCH I. (1990) Bull. Mus. Natl. Hist. Nat., Paris, 4è sér., A 12: 69-73.BARTSCH I. (1994) Cah. Biol. Mar. 35: 479-490.SEGONZAC M. (1992) C. R. Acad. Sci. Paris, Sér. III 314: 593-600.
I. BARTSCH Denisia 18 (2006): 300
Arthropoda, Chelicerata, Arachnida, Acariformes, Halacaridae
Size: 650-700 µm (from tip to rostrum to end of anal papilla).
Morphology: Marine mite with ovate idiosoma and two pairs oflegs directed forward and two pairs backward. Gnathosomaabout 1/3 as long as the idiosoma; rostrum slender. Dorsal idio-somatic plates reticulate, membranous integument betweenplates with minute denticles. First pair of legs with long, blunt-ly ending ventral spines; in adults, third, fourth and fifth seg-ment with five, two, and five spines respectively.
Biology: Several mites have suctorians (Ciliophora) fixed onthe idiosoma, gnathosoma or legs.
Distribution: Mid-Atlantic Ridge: Snake Pit and Broken Spur.
1: Habitus; by I. Bartsch.
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Ammothea verenae CHILD, 1987
Synonym: Scipiolus thermophilus TURPAEVA, 1988.
Size: Male leg span about 43 mm. Female slightly larger.
Morphology: Trunk slender with segment posteriors flared intocowling. Ocular tubercle a low, blind, truncate cone at neckmid length. Lateral processes separated by less than half theirdiameters. Proboscis long, with median and distal constrictions.Abdomen long, down curved. Chelifores tiny, short, chelae at-rophied bumps. Palps nine-segmented, 5 distal segments veryshort, heavily setose ventrally. Oviger segment 4 and 5 subequal in length, proximally setose, strigilis segments 6, 7 and 8with many long lateral setae, segments 8, 9 and 10 with 1-2small endal denticulate spines. Eggs tiny, about 0.25 diameter ofmain oviger segments, carried in large round clusters. Legs long,slender, extremely setose. Third coxae with many long ventralsetae. Distal leg segments alike, tarsus very short, propodusslender, well curved, sole spines alike. Main claws with long
auxiliary claws. Male cement gland opening a tiny dorsodistalpore. Female oviger strigilis with segments lacking dense setae,with 1-2 denticulate spines on distal segments. Female legs withfar fewer setae, without large fields of setae.
Remarks: This species differs in basic morphology from anyother known member of the genus. It is blind, lacks the con-spicuous dorsomedian tubercles common on the trunk cowls ofmost species, lacks differentiated heel spines, and does not havedifferences in propodi shape between the anterior and posteri-or legs.
Biology: Found commonly in close proximity to hydrothermalvents and sometimes encrusted with polymetallic sulfides.
Distribution: Juan de Fuca Ridge: Axial Seamount, EndeavourSegment; Explorer Ridge.
References:
CHILD C.A. (1987) Proc. Biol. Soc. Wash. 100(4): 892-896.TURPAEVA E.P. (1988) Zool. Zh. 67(6): 950-953.
R. BAMBER Denisia 18 (2006): 301
1: Male holotype. A: Trunk, dorsal view; B: Trunk, lateralview; C: Oviger; female paratype: D: Third leg with ova;from CHILD (1987).
2: In situ view showing numerous specimens and a scaleworm (Branchinotogluma?) © NOAA Ocean Exploration.
Arthropoda, Pycnogonida, Ammotheidae
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Sericosura cochleifovea CHILD, 1989
Synonym: Sericosura bifurcata STOCK, 1991.
Size: Small for the genus, male leg span about 13 mm.
Morphology: Trunk slightly ovoid, lateral processes separatedby about half their diameters, armed with one or more dor-sodistal and laterodistal spines. Trunk segments with swollenposterior cowls. Ocular tubercle only as tall as its base, blind.Proboscis ovoid, originating on slender stalk, greatly constrict-ed distally. Abdomen long, downcurved, with two pairs of longdorsal spines, pair of short distal spines. Chelifores typical;short, with atrophied chelae. Palps nine-segmented, secondsegment longest, five distal segments little longer than wide,heavily setose. Oviger fourth segment longest, fifth slightlyshorter. Strigilis sixth segment with field of long lateral setae,with 1-2 tiny denticulate spines on distal three segments. Legsmoderately slender, very setose. Male leg major segments withrows of many setae, some longer than segment diameter. Femaleleg major segments with few very long femoral setae, lateralrows of extremely long setae and few long dorsal setae on tibi-
ae and propodus. No thick fields of setae. Tarsus short, propo-dus slender, almost straight, claws moderately long. Eggs large,almost as wide as oviger segment diameters around which theyare wound. Cement gland at extreme proximal end of femurwith laterally pointing tube slightly longer than segment diam-eter.
Remarks: One of the major differences between this speciesand the others in this genus is the sexual dimorphism displayedin leg setae. The very long lateral leg setae of females differen-tiate them conspicuously from males. The shorter leg setae andvery long cement gland tube of the males sets them off from fe-males of this species and males of other known species whichhave much shorter tubes.
Biology: Taken in the proximity of a vent site with other fau-na: snails, crabs, shrimps and anemones.
Distribution: Only known from the Mariana Islands Back-ArcBasin hydrothermal vent fields.
References:
CHILD C.A. (1989) Proc. Biol. Soc. Wash. 102(3): 732-737.STOCK J.H. (1991) Résultats Campagnes Musorstom, 8. Mém. Mus. Natl. Hist. Nat., Paris (A) 151: 158-160.
R. BAMBER Denisia 18 (2006): 302
1: Holotype male. A: Trunk, dorsal view; B: Trunk, lateral view; C: Palp; D: Third leg, with cement gland tube enlarged; E: Ovigerwith several eggs attached; from CHILD (1989).
Arthropoda, Pycnogonida, Ammotheidae
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Sericosura cyrtoma CHILD & SEGONZAC, 1996
Size: Leg span 17 mm.
Morphology: Trunk robust, very broad anteriorly, armed withmany short spines on sockets on trunk anterior, distally on lat-eral processes and chelifores, and on palp second segment. Oc-ular tubercle broad, not as tall as basal width, blind. Proboscistapering from broad base, downcurved distally, lips flat. Ab-domen broad, long, with dorsal spines. Chelifore scapes twice aslong as wide, chelae atrophied to rounded nubs. Palp seven-seg-mented, fourth segment slightly longer than second, fifthlonger than sixth and seventh combined, few setae. Oviger (fe-male) small, plain, strigilis with 1-2 tiny denticulate spines per
segment. Legs with dorsal, ventral and lateral rows of shortspines in sockets, tibiae with 1-2 longer dorsal setae. Tarsus veryshort, propodus typical, with few sole spines. Claw long, auxil-iaries half main claw length. Male features unknown.
Remark: The tapered and bent proboscis is a unique characteramong the known species of Sericosura.
Biology: Found associated with siboglinid worms in a hy-drothermal vent area.
Distribution: East Pacific Rise: 13°N.
Reference:
CHILD C.A. & M. SEGONZAC (1996) Proc. Biol. Soc. Wash. 109(4): 664-676.
R. BAMBER Denisia 18 (2006): 303
1: Holotype female. A: Trunk, dorsal view; B: Trunk, lateral view; C: Palp; D: Oviger; E: Third leg; from CHILD & SEGONZAC (1989).
Arthropoda, Pycnogonida, Ammotheidae
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Sericosura heteroscela CHILD & SEGONZAC, 1996
1: Holotype male. A: Trunk, dorsal view; B: Trunk, lateral view; C:Palp; D: Oviger; E: Third leg; from CHILD & SEGONZAC (1996).
3: Specimen collected at Rainbowsite; cruise Atos; note the strongsulphide deposit on the entirebody; by P. Briand © Ifremer.
2: Specimen collected at Lucky Strike site(cruise Atos); by P. Briand © Ifremer.
4: Specimen with eggs, and colonized by fila-mentous bacteria; by P. Briand © Ifremer.
Arthropoda, Pycnogonida, Ammotheidae
References:
CHILD C.A. & M. SEGONZAC (1996) Proc. Biol. Soc. Wash. 109(4): 664-676.TURPAEVA E.P. (1998) in KUZNETSOV A.P. & O.N. ZEZINA (Eds.) Benthos of the High Latitude Regions. Collected Proc.: 1-138 [in Russian].
R. BAMBER Denisia 18 (2006): 304
Size: Leg span 26.7 mm.
Morphology: Trunk and lateral processes typical. Lateralprocesses with many short dorso- and latero-distal spines. Ocu-lar tubercle slender, twice as long as diameter, blind, bifurcateat tip. Proboscis and abdomen typical. Chelifore scapes hardlylonger than wide, chelae rounded atrophied nubs. Palps seven-segmented, only slightly longer than proboscis, moderately se-tose. Oviger typical, field of long lateral setae on sixth and sev-enth segments, distal three segments with tiny denticulatespines. Male legs of two kinds. Anterior two pairs with normal-ly proportioned distal segments, posterior two pairs with gross-ly enlarged and inflated propodi twice diameter and slightlylonger than anterior propodi. Claw of inflated propodi largerand longer than normal propodi and auxiliary claws shorter inrelation to main claw. Cement gland of all male legs a smallbulbous dorsolateral inflation on proximal femur with a trun-
cate conical tube directed anterolaterally. All female legs withnormal sized propodi and fewer short setae than those of male.Neither sex with fields of leg setae.
Remarks: The grossly inflated posterior four propodi of malesserve to set this species off from any other known in the genus.TURPAEVA (1998) allocates this species to a new genusAnisopes: A. heterocella (sic). Nevertheless, we suggest that weneed more material to check the probable morphological vari-ations.
Biology: Specimens of this species were found in associationwith hydrothermal vents in several different collecting locali-ties.
Distribution: Mid-Atlantic Ridge: Snake Pit, Logatchev, Rain-bow, Lucky Strike, and Menez Gwen.
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Sericosura mitrata (GORDON, 1944)
References:
CHILD C.A. (1982) Smithon. Contrib. Zool. 349: 19-21, Fig. 6.FRY W.G. & J.W. HEDGPETH (1969) Mem. New. Zeal. Oceano. Inst. 49: 112-113.GORDON I. (1944) Brit., Austr., New Zeal. Ant. Res. Exped. ser. B 5(1): 54-57.SEGONZAC M. (1992) C. R. Acad. Sci., Paris 314(III): 593-600.
R. BAMBER Denisia 18 (2006): 305
1: Male. A: Trunk, dorsal view; B: Third leg, with enlargement of cement gland; C: Palp; D: Oviger; from GORDON (1944).
Arthropoda, Pycnogonida, Ammotheidae
Synonym: Achelia mitrata GORDON, 1944.
Morphology: Trunk typical with lateral processes spaced at dis-tances of half their diameters, armed with 2-3 short dorsodistaland laterodistal setae. Ocular tubercle more than twice as longthan its basal diameter, blind, bifurcate at tip with prominentsensory papillae. Proboscis ovoid with basal and distal constric-tions. Abdomen long with 2-3 short distal setae. Cheliforesvery short, scapes broad, chelae tiny, atrophied. Palps seven-segmented, segments 2 and 4 subequal in length, seventh seg-ment as long as fifth and sixth coined lengths, moderately se-tose. Oviger second segment longest, fifth slightly longer thanfourth. Sixth segment with field of lateral setae longer than seg-ment diameter. Strigilis segments 8, 9, 10 with pairs of tiny den-ticulate spines. Male legs dimorphic in setae lengths andarrangement. Tibiae with sparse setae longer than segment di-ameters, with 1-2 ventral spines. Propodus slender, slightlycurved, with few long and short setae, few short sole spines.Claws moderately long. Cement gland forming a small proxi-mal bulge on femur, tube slender, about as long as femur diam-
eter. Female legs with dorsal, lateral and ventral rows of shortsharp spines pointing distally, lateral spines slightly longer.Some specimens with very long lateral setae. Propodus withrow of many short sole spines.
Remarks: The very long leg setae of Gordon’s female specimenmay be an infestation of something growing on the integumentand not setae. One of the easily identified characters in thisgenus is the number of palp segments. The genus has specieswith either seven or nine segments with two of the above hav-ing seven and two having nine.
Biology: This species was described before the discovery of hy-drothermal vent areas. It is an opportunistic species in the At-lantic vent communities where it has been collected since.
Distribution: Eastern margin of Antarctica at 219 m, in theRoss Sea at 106 m, on the Walvis Ridge off South Africa at2100 m; Mid-Atlantic Ridge: Snake Pit.
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Sericosura venticola CHILD, 1987
Reference:
CHILD C.A. (1987) Proc. Biol. Soc. Wash. 100(4): 896-899.
R. BAMBER Denisia 18 (2006): 306
1: Male holotype. A: Trunk, dorsal view; B: Trunk, lateral view; C: Palp; D: Third leg, with enlargement of cement gland; E:Oviger; from CHILD (1987).
Arthropoda, Pycnogonida, Ammotheidae
Size: Male leg span about 28 mm. Female size unknown.
Morphology: Trunk moderately slender, first three segmentswith flaring posterior cowls. Lateral processes separated by lessthan half their diameters, armed with 3-6 lateral spines. Oculartubercle taller than basal diameter, blind. Proboscis massive,with proximal and distal constrictions. Abdomen long, down-curved. Chelifores small, short, chelae atrophied to nubs. Palpsseven-segmented, segments 2 and 4 subequal in length, distalthree short segments heavily setose. Oviger second segmentlongest, fourth and fifth subequal in length. Strigilis segments6, 7 and 8 with many long lateral setae, segments 8, 9, 10 with1-2 small denticulate spines. Eggs very tiny, about one fifth aswide as the diameter of segment they are on. Legs slender, heav-ily setose with many ventral setae on coxa 3 and proximal fe-mur. Distal leg segments of uniform size, tarsus short, propodusslender, well curved, claws of moderate length. Male cementgland a swelling at proximal end of femur, with a short tubepointing laterally. Female characters unknown.
Remarks: The main difference between specimens of thisgenus and those in the genus Ammothea is the placement of themale cement gland and its orifice. They are otherwise difficultto separate.
Biology: The only known specimens were found associatedwith siboglinids at a hydrothermal vent on the Endeavour Seg-ment.
Distribution: NE-Pacific: Endeavour Segment, Juan de FucaRidge; a juvenile belonging probably to the same species wascollected at Southern East Pacific Rise: 17°S, Rehu-Marka site.
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Archiconchoecia (Archiconchoecia) chavturi KORNICKER & HARRISON-NELSON, 2005
Size: Adult female carapace length 0.60 mm, height 0.40 mm.
Morphology of adult female (male unknown): Carapace: ovalin lateral view with greatest height just posterior to midlength;anterior rostrum and incisure small; posterior edge of eachvalve with small glandular process dorsal to midheight; withoutposterodorsal spines. First antenna with proximal black spots.Second antenna: exopod with short medial setae on article 1;endopod article 1 without processus mamillaris; ventral marginwith minute digitations and spines. Basal endite of mandiblewith row of triangular teeth. Bellonci Organ elongate, reaching
bend at tip of first antenna, with rounded tip bearing twominute spines.
Biology: Unique female with 15 eggs in marsupium. Gut withappendage fragments, bristles, and bristle-like claws indicatingthat the species is a scavenger or predator on small inverte-brates. The collecting site suggests that the species, which is anactive swimmer, dwells in deep-water close to the substrate;collected within Riftia pachyptila aggregations.
Distribution: East Pacific Rise: 9°N, site Tica.
Reference:
KORNICKER L.S. & E. HARRISON-NELSON (2005) Zootaxa 1071: 19-38.
L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 307
1: Complete specimen from left side.
2: Complete specimen, dorsal view, anterior to right (black spotsare on first antenna; BO – tip of Bellonci Organ).
Arthropoda, Crustacea, Ostracoda, Myodocopa, Halocyprida, Halocypridae
3: Left first antenna andBellonci Organ.
4: Rightmandible,distal coxa andbasis.
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Bathyconchoecia deeveyae KORNICKER, 1969
Size (length and height excluding spines): Adult or A-1 femalecollected off Surinam: length 1.49 mm, height 1.10 mm. Juve-nile collected off Peru: length 1.12 mm, height 0.89 mm. A-4 instar collected in Guaymas Basin: length 0.66 mm, height0.46 mm.
Morphology: Carapace with seven spines (each valve with an-terior spine, dorsal spine near midlength, and lateral spine be-low middle; right valve with posterior spine). Surface with orwithout punctae and arcuate ridges between punctae. Pos-terodorsal corner of each valve with glandular process.
Biology: Collections indicate that the species, a swimmer, livesclose to the bottom at bathyal and abyssal depths.
Distribution: Guaymas Basin, Southern Trough (2000 m), col-lected in plankton net 3-4 m above bottom. Also collected offPeru in the Peru-Chile Trench system (520 m), and in westernAtlantic off Surinam (508–523 m).
References:
KORNICKER L.S. (1969) Proc. Biol. Soc. Wash. 82: 403–408.KORNICKER L.S. (1981) Proc. Biol. Soc. Wash. 89(4): 1237–1243.KORNICKER L.S. (1991) Smithson. Contr. Sci. 516: 1–46.
L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 308
1: Juvenile carapace, dorsal, lateral, and ventral views(length 0.87 mm); by Kornicker.
2: Juvenile carapace, lateral view (length 0.87 mm); by Kornicker.
3: Endopod of second antenna(adult or A-1 instar); by Kornicker.
Arthropoda, Crustacea, Ostracoda, Myodocopa, Halocyprida, Halocypridae
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Bathyconchoecia paulula DEEVEY, 1968
Size: Adult male length 0.95 mm, height 0.60–0.61 mm (Guay-mas Basin); length 0.95 mm, height 0.70 mm (Gulf of Mexico).
Morphology: Adult male carapace rostrum with prolongeddownward pointed tip; surface with crescent-like reticulationscontaining minute pits; height at least 70% length; posterodor-sal corner of each valve with glandular opening; list along pos-terior infold with four transparent lamella with four triangularflagella-like cusps. First antenna terminal segments with morethan 200 sensory filaments. Second antenna exopod with un-usually long first article (71–72% length of protopod); endopodfirst article with two bristles (distal bristle about twice length ofproximal bristle). Fifth limb terminal article of endopod withtwo claws and one shorter ringed bristle. Sixth limb terminalarticle of endopod with three bristles (one very long, twoshort). Seventh limb with two bristles. Furca with eight claws
on each lamella and long unpaired bristle; small oval area prox-imal to claw 2. Copulatory organ inner rod with pointed re-curved tip. Adult female carapace and appendages, except en-dopod of second antenna, fairly similar to those of adult male.
Biology: Collections indicate that the species, a swimmer, livesclose to the bottom at bathyal and abyssal depths. Specimensfrom the Gulf of Mexico were collected in the intestines of bot-tom fish.
Distribution: Guaymas Basin, Southern Trough (collected inplankton net 3-4 m above bottom). Also collected in Gulf ofMexico (from intestines of bottom fish), 1000 m and southwestof Sao Miquel, Azores (620–800 m) in plankton net open be-tween 680 and 780 m.
References:
DEEVEY G.B. (1968) Proc. Biol. Soc. Wash. 81: 539–570.KORNICKER L.S. (1991) Smithson. Contr. Sci. 516: 1–46.POULSEN E.M. (1972) Tethys 4(2): 445–456.
L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 309
Arthropoda, Crustacea, Ostracoda, Myodocopa, Halocyprida, Halocypridae
2: Adult maleendopod of secondantenna; by Kornicker.
1: Adult male carapace from right side; by Kornicker.
3: Adult male sixth limb; by Kornicker.
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Euphilomedes climax KORNICKER, 1991
References:
KORNICKER L.S. (1991) Smithson. Contr. Zool. 516: 1–46.TSURUMI M. & V. TUNNICLIFFE (2003) Deep-Sea Res. I 50: 611–629.TUNNICLIFFE V. (1988) Proc. R. Soc. Lond. B 233: 347–366.
L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 310
1: Complete carapace showing four eggs; by Kornicker.
Arthropoda, Crustacea, Ostracoda, Myodocopa, Myodocopida, Philomedidae
2: Right lamella of furca; by Kornicker.
3: Seventh limb; by Kornicker.
4: Anterior of bodyfrom right side show-ing Bellonci Organ;by Kornicker.
Size: Adult female carapace length 2.84 mm, height 2.02 mm.
Morphology of adult female (adult male unknown): Carapaceoval in lateral view with broad rostrum and deep incisure; sur-face with many short pointed bristles, and long widely separat-ed bristles more numerous along valve edge. Mandible exopodabout half length of dorsal margin of article 1 of endopod. Sev-enth limb with 17–19 bristles; comb with 13 alate teeth withtwo small teeth on each side of base, side opposite comb withtwo or three pegs. Furca with 11–14 claws on each lamella; claw3 small, secondary, remaining claws decreasing in length alonglamella. Bellonci Organ elongate with suture at midlength andpointed or narrowly rounded tip. Lateral eyes absent in femalesat all stages, present in males at all stages.
Biology: Benthic swimmer and burrower. One female with foureggs in marsupium. Detritus feeder; guts generally with uniden-tified particles, but one specimen with two copepods in gut.The collecting site suggests that the species may be restricted todeep water.
Distribution: Explorer Ridge: Pogo Peaks Vent, Gulati GusherVent, Lunch Hour Vent, Crab Vent, Upper Magic MountainVent, Busted Thruster Vent; Juan de Fuca Ridge: Long TermObservatory Vent, Axial Seamount, Hamond’s Hell Vent.
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Polycopetta pax KORNICKER & HARRISON-NELSON, 2005
Size: Adult female carapace length 0.54 mm, height 0.47 mm.
Morphology of female (male unknown): Carapace: anteriormargin with concavity, but without rostrum or anterodorsaltooth; ventral margin with about 25 small teeth and minutespines (or serrations) between teeth. First antenna dorsal mar-gin without processes, but with long proximal bristle.Mandible: coxa endite bifurcate distally; exopod interpreted tohave two articles, but without suture separating articles: article1 broad with terminal bare bristle; article 2 with proximalspines and funnel-like tip. Maxilla: proximal dorsal half of ba-sis with long backward projection; distal end of exopod reach-
ing distal end of endopod, and with eight long terminal bristles.Fifth limb exopod well-developed, and with four terminal spin-ous bristles (three long, one short). Bellonci Organ consists ofspinous bristle divided at midlength.
Biology: Unique specimen with several eggs in marsupium, andfilled internal sperm sac near base of furca. Gut with brown par-ticles, some round or disc-like, and with few fragments of tubu-lar structures. Species is a bottom dweller; found within Riftiapachyptila aggregations, maximal water temperature 23°C.
Distribution: East Pacific Rise: 9°N, site Riftia Field.
Reference:
KORNICKER L.S. & E. HARRISON-NELSON (2005) Zootaxa 1071: 19-38.
L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 311
1: Complete specimen from right side.
2: First antennae and Bellonci Organ (not all bristles shownon first antennae).
3: Right mandible.
Arthropoda, Crustacea, Ostracoda, Myodocopa, Myodocopida, Polycopidae
4: Right maxilla (notall bristles shown;precoxa endite notshown).
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Prionotoleberis styx KORNICKER, 1991
Reference:
KORNICKER L.S. (1991) Smithson. Contr. Zool. 516: 1–46.
L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 312
2: Sixth limb; by Kornicker.
4: Mandible; by Kornicker.
Arthropoda, Crustacea, Ostracoda, Myodocopa, Myodocopida, Cylindroleberidae
1: Complete carapace showing central adductormuscles; by Kornicker.
Size: Adult female carapace length 1.89 mm, height 1.09 mm.
Morphology of adult female (adult male unknown): Carapaceelongate with deep incisure. First antenna sensory bristle withfairly long proximal bristle and six long terminal filaments.Sixth limb anterior margin with 18–20 short bristles. Seventhlimb with 24 or 25 bristles. Furca with 10 claws on each lamel-la. Lateral eyes absent.
Biology: Benthic swimmer and burrower. Filter feeder; gut slen-der, containing fine-grained material. The collecting site sug-gests that the species may be restricted to deep water.
Distribution: East Pacific Rise: National Geographic Site,Clam Acres.
3: Seventh limb;by Kornicker.
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Thomontocypris brightae MADDOCKS, 2006
References:
MADDOCKS R.F. (1991) Zool. J. Linn. Soc. 103: 309-333.MADDOCKS R.F. (2006) Micropaleontology 51: 345-372.MADDOCKS R.F. & P.L. STEINECK (1987) Micropaleontology 33: 318-355.
R.F. MADDOCKS Denisia 18 (2006): 313
1: Left side of entire animal.
2: Exterior of right valve, male specimen.
4: Left side of entire female animal.
5: Dorsal view of entire male animal.
3: Exterior of left valve, male specimen.
6: Right side of anterior body, showing anteri-oventral horn on upper lip and palps of male fifthlimb. 1-6; from MADDOCKS (2006).
Arthropoda, Crustacea, Ostracoda, Podocopida, Pontocyprididae
Size: Carapace length 0.53-0.57 mm, height 0.30-0.33 mm, nocarapace dimorphism.
Morphology: Carapace compressed, lateral outline rounded-subtriangular with very broadly rounded anterior margin andthree-segmented dorsal margin, distinct ventral indentation;conspicuous anteroventral hornlike process on upper lip; palpof male fifth limb broad, subtriangular, with thumb-like ven-trodistal projection; terminal setae of seventh limb smooth;Zenker’s organ with three bulbous swellings.
Remarks: Other species of Thomontocypris have been reportedliving in anchialine pools and caverns (Bermuda), sublittoral
and reefal sand and plants (Madagascar, Australia), and exper-imental wood falls in the deep sea (MADDOCKS & STEINECK
1987; MADDOCKS 1991).
Biology: Living specimens were collected in washings of Riftiapachyptila. Appendages and genitalia are normal for the genus,and gut contents include no recognizable objects. Well-devel-oped swimming setae suggest active demersal swimming, whilethe rather weak mouthparts of this family indicate a soft detri-tus or bacterial diet. The function of the hornlike process is un-known.
Distribution: East Pacific Rise: 9°N, site Riftia Field and Tica.
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Thomontocypris gollnerae MADDOCKS, 2006
References:
MADDOCKS R.F. (1991) Zool. J. Linn. Soc. 103: 309-333.MADDOCKS R.F. (2006) Micropaleontology 51: 345-372.MADDOCKS R.F. & P.L. STEINECK (1987) Micropaleontology 33: 318-355.
R.F. MADDOCKS Denisia 18 (2006): 314
1: Ventral view of entire female.
2: Right valve exterior with ovaries.
4: Furca.
5: Detail of left valve exterior, showing adductormuscle scars (asterisk) and normal pore canals. 1-6; from MADDOCKS (2006).
3: Left valve exterior with ovaries.
Arthropoda, Crustacea, Ostracoda, Podocopa, Podocopida, Pontocyprididae
Size: Carapace length 0.65 mm, height 0.32 mm, no carapacedimorphism.
Morphology: Carapace egg-shaped, slightly flexible, swollenventrolaterally, with rounded-subtrapezoidal lateral outline,weakly convex ventral margin, and slightly produced anteriorend; Y-aesthetasc of antenna small, elongate; terminal setae ofseventh limb smooth; furcal claws with opposing setules form-ing terminal thorn.
Remarks: Other species of Thomontocypris have been reportedliving in anchialine pools and caverns (Bermuda), sublittoraland reefal sand and plants (Madagascar, Australia), and exper-
imental wood falls in the deep sea (MADDOCKS & STEINECK
1987; MADDOCKS 1991).
Biology: Living specimens were collected in washings of Riftiapachyptila. Appendages and genitalia are normal for the genus,and gut contents include no recognizable objects. Well-devel-oped swimming setae suggest active demersal swimming, whilethe rather weak mouthparts of this family indicate a soft detri-tus or bacterial diet. The unusually weak calcification of thecarapace may be either a response or a preadaptation to low pH.
Distribution: East Pacific Rise: 9°N, site Riftia Field.
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Xylocythere vanharteni MADDOCKS, 2006
Syn.: Xylocythere sp. A in VAN HARTEN (1993).
Size: Carapace length 0.56-0.66 mm, height 0.27-0.32 mm,males smaller than females.
Morphology: Lateral outline elongate-subquadrate to subreni-form, with broad ventral indentation; reticular network of low,narrow muri and flat, polygonal sola uniformly developed overentire lateral surface, except for nearly smooth submarginalfields; 20-45 pores in pore clusters; prominent ventrolateralspine and single anterior and posterior marginal spines on eachvalve.
Remarks: Other living and fossil species of Xylocythere are re-ported from experimental wood falls and deep-sea sediments
(Atlantic and Pacific, Upper Oligocene to Holocene; MAD-DOCKS & STEINECK 1987; STEINECK et al. 1990).
Biology: Living specimens were collected in washings of Riftiapachyptila. The mouthparts and legs are normal for the family,and the gut contents contains no recognizable objects. A crawl-ing habit and bacterial or detrital diet are probable. The func-tion of the pore clusters is unknown but might be related to res-piration (VAN HARTEN, 1992, 1993).
Distribution: East Pacific Rise: 13°N and 9°N, sites Tica andRiftia Field.
References:
MADDOCKS R.F. (2006) Micropaleontology 51: 345-372.MADDOCKS R.F., & P.L. STEINECK (1987) Micropaleontology 33: 318-355.STEINECK P.L., MADDOCKS R.F., COLES G. & R.C. WHATLEY (1990) in WHATLEY R. & C. MAYBURY (Eds.) Ostracoda and Global Events,
Chapman & Hall: 307-319.VAN HARTEN D. (1992) Deep-Sea Res. 39: 1067-1070. VAN HARTEN D. (1993) in MCKENZIE K.G. & P.J. JONES (Eds.) Ostracoda in the Earth and Life Sciences. A.A. Balkema: 571-580.
R.F. MADDOCKS Denisia 18 (2006): 315
1: Entire animal, right side.
2: Right valve exterior.
4: Left valve exterior (SEM).
5: Right valve interior showing hinge and pore clusters (SEM). 1-5: from MADDOCKS (2006).3: Left valve exterior.
Arthropoda, Crustacea, Ostracoda, Podocopida, Cytheruridae
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Arthropoda, Crustacea, Copepoda
Almost 80 species of copepods are described from hydro-thermal vents. More than half of the copepods species recordedrepresent copepods of the family Dirivultidae (order Siphonos-tomatoida) which were exclusively found at different vent siteswith exception for the type species of Dirivultus dentaneus (HU-MES & DOJIRI 1980; IVANENKO & FERRARI 2003). Additionally,three dirivultid species descriptions from animals of the EastPacific Rise, the West Pacific, and the Mid-Atlantic Ridge arein preparation; and at least two dozens of new species, whichare in our disposition, are waiting for thorough descriptions.These are mostly representatives of calanoids, cyclopoids andharpacticoids from different localities of the East Pacific Rise
and the Mid-Atlantic Ridge, which came mainly from the sam-ples obtained during in situ colonization experiments and sedi-ment traps. Some of these copepods represent common genera(such as the harpacticoid genus Tisbe and the cyclopoid genusHeptnerina close to Cyclopina) and even common families (theharpacticoid family Tegastidae) known till now only from shal-low waters (IVANENKO & DEFAYE 2004a, b; V. Ivanenko & D.Defaye, unpublished).
Certainly new methods of meiofauna sampling and explo-ration of new sites will reveal many new copepods representingdifferent taxonomical and ecological groups. Further ecologicalstudies of copepods from different microhabitats, distinct loca-lities, environments surrounding deep-sea hydrothermal ventsand cold seeps (see HEPTNER & IVANENKO 2002; V. Ivanenko,D. Defaye & Cuoc, unpublished), as well as meiofauna associa-ted with whale remnants (to date unknown) will let us betterunderstand the role of the remarkably diverse copepods instructuring and functioning of deep-sea chemosynthetic com-munities.
For more information about taxonomic composition, distri-bution, morphological, and biological traits, and the presumedhabitat specificity of the copepods from deep-sea chemosynthe-tic environments we refer to the paper of HEPTNER & IVANEN-KO (2002) as an attempt to summarize all data available for thatmoment.
1: Oncaea preclara from East Pacific Rise: 9°N, Tica;by M. Bright.
V.N. IVANENKO Denisia 18 (2006): 316–317
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References:
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta 11(2): 117-134.HUMES A.G. & M. DOJIRI (1980) Pac. Sci. 34: 143-151.IVANENKO V.N. & D. DEFAYE (2004a) Cah. Biol. Mar. 45(3): 255-268.IVANENKO V.N. & D. DEFAYE (2004b) Zoosystema 26(1): 49-64.IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11(3): 177-185.
2: Aphotopontius sp. from East Pacific Rise: 13°N; by M. Bright.
317
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Andromastax CONROY-DALTON & HUYS, 1999
Arthropoda, Crustacea, Copepoda, Harpacticoida, Aegisthidae
1: A. cephaloceratus; A: Male habitus, dorsal; B: Female cephalosome, dorsal;from LEE & HUYS (2000).
Species Distribution Body length in mmA. cephaloceratus LEE & HUYS, 2000 Okinawa Trough Female 3.05, male 2.85
A. muricatus CONROY-DALTON & HUYS, 1999 Galapagos Spreding Center Female 3.49, male 3.24
Morphology: Body with reticulated integument. Cephalicshield, coxae of legs 2-4 with spinous processes. Caudal ramislightly longer than rest of body. Antennule of female seven-segmented, second segment with two spine-like processes. An-tennule of male nine-segmented, segment 8 elongate. Feedingappendages of male reduced.
Biology: Large benthopelagic copepods with non-feedingmales. A. muricatus has been found in “washing of mussel sam-ple”. A. cephaloceratus has been collected 0.5-3 m above thebottom.
References:
CONROY-DALTON S. & R. HUYS (1999) J. Crustac. Biol. 19(2): 408-431.LEE W. & R. HUYS (2000) Zool. J. Linn. Soc. 129: 1-71.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 318
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Uptionyx verenae CONROY-DALTON & HUYS, 2000
Reference:
CONROY-DALTON S. & R. HUYS (2000) Cah. Biol. Mar. 41(4): 343-397.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 319
Arthropoda, Crustacea, Copepoda, Harpacticoida, Ancorabolidae
1: Female; A: Habitus, dorsal; B: Habitus, lateral; from CONROY-DALTON & HUYS (2000).
Size: Body length of female 1.11 mm. Male unknown.
Morphology: Body with dorsal and lateral bulbous processes onall somites except last two somites. Rostrum subtriangularprominence. Antennule three-segmented, with aesthetasc onsegments 2 and 3. Antenna three-segmented, without exopod.Legs 1-4 biramous; endopods of legs 1-4 and exopod of leg 1two-segmented, exopod of legs 2-4 three-segmented. Leg 5:long endopod fused with protopod, with four setae; exopodone-segmented, with five setae.
Biology: Two females were found associated with macroinver-tebrates. Most ancorabolids inhabit mud sediment.
Distribution: Juan de Fuca Ridge: Middle Valley Segment,Peanut Vent.
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Argestoides prehensilis HUYS & CONROY-DALTON, 1997
Arthropoda, Crustacea, Copepoda, Harpacticoida, Argestidae
Size: Body length of male 0.34 mm. Female unknown.
Morphology: Prosome four-segmented; urosome six-segmented.Antennule eleven-segmented, geniculation between segments8 and 9. Antennary exopod two-segmented, with three setae.Maxilliped subchelate, three-segmented. Legs 1-4 biramous,with three-segmented rami. Endopod of leg 1 prehensile, firstsegment elongate. Leg 5 two-segmented, baseoendopod withendopodal lobe bearing two spines, exopod with six setae.
Biology: One male has been found in a “fraction of bucket mus-sel washings”, bearing, attached to its shield of cephalothoraxand anal somite, two stages of a tantulocarid, an ectoparasiticcrustacean.
Distribution: Galapagos Spreading Center.
1: Male; A: Habitus, dorsal; B: Urosome, ventral; from HUYS & CONROY-DALTON (1997).
Reference:
HUYS R. & CONROY-DALTON S. (1997) Cah. Biol. Mar. 38: 235-249.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 320
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Jamstecia terazakii LEE & HUYS, 2000
Arthropoda, Crustacea, Copepoda, Harpacticoida, Aegisthidae
1: Female habitus, lateral;from LEE & HUYS (2000).
Size: Body length of female 3.38 mm. Male unknown.
Morphology: Body with reticulated integument and pairedspinous processes. Caudal rami 1.5 longer than rest of body.Antennule seven-segmented, first segment elongate. Antennawith two-segmented exopod, one-segmented endopod shorterthan allobasis. Leg 5 large, one-segmented.
Biology: Benthopelagic copepod sampled 0.5-3 m above thebottom.
Distribution: Okinawa Trough.
References:
LEE W. & R. HUYS (2000) Zool. J. Linn. Soc. 129: 1-71.LEE W. & R. HUYS (2001) Zool. J. Linn. Soc. 131: 249.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 321
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Scabrantenna yooi LEE & HUYS, 2000
Arthropoda, Crustacea, Copepoda, Harpacticoida, Aegisthidae
1A: Female habitus, dorsal; B: Male antenna; C: Antenna; from LEE & HUYS (2000).
Size: Body length of female 3.53 mm; male 3.32 mm.
Morphology: Body with reticulated integument. Caudal ramislightly longer than rest of body. Rostrum spiniform, small in fe-male and strongly developed in male. Antennule seven-seg-mented in female, nine-segmented in male. Antenna withthree-segmented exopod, allobasis and endopod sexually di-morphic. Mandible and maxillule reduced in male. Leg 1-4 bi-ramous, with three-segmented rami, excepting one-segmentedendopod of leg 1. Leg 5 almost reaching posterior edge of analsomite, one-segmented in female and three-segmented in male.
Biology: Benthopelagic copepod sampled 0.5-3 m above thebottom. Male non-feeding.
Distribution: Okinawa Trough.
Reference:
LEE W. & R. HUYS (2000) Zool. J. Linn. Soc. 129: 1-71.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 322
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Cheramomyzon abyssale HUMES, 1989
Size: Body length of female 2.09-2.27 mm. Male unknown.
Morphology: Prosome four-segmented, tergites of last twosomites pointed posteriorly. Urosome five-segmented; firstsomite with leg 5. Genital double-somite and caudal ramuselongate. Oral cone elongate. Antennule nineteen-segmented;segment 17 with aesthetasc. Antenna with short one-segment-ed exopod. Mandible with stylet-like gnathobase and two-seg-mented palp bearing two terminal setae. Maxilla two-segment-ed, distal segment claw-like. Legs 1-4 biramous, with three-seg-mented rami. Leg 5 two-segmented, distal segment with four se-tae.
Remark: Has been found in three samples.
Distribution: East Pacific Rise: 13°N.
Reference:
HUMES A.G. (1989) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 11, section A (4): 829-849.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 323
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Asterocheridae
1: Female; A: Habitus, dorsal; B: Habitus, lateral; C: Leg 5 and genital double-somite, ventral; D: Antennule; E: Antenna; F: Mandible. ae – aesthetasc; gn – gnathobase of mandible; l5 – leg 5; pl – palp of mandible; from HUMES (1989).
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Collocherides brychius HUMES, 1999
Size: Body length of female 0.55-0.58 mm; male 0.50-0.51 mm.
Morphology: Prosome four-segmented. Urosome five-segment-ed in female and six-segmented in male; first somite with leg 5.Longest seta of caudal ramus slightly longer (1.18:1) than ra-mus. Oral cone short. Antennule of female twenty-segmented,segment 18 with aesthetasc. Antenna with exopod representedby one short seta. Mandible with stylet-like gnathobase andone-segmented palp with one terminal seta. Maxilla three-seg-mented. Legs 1-4 biramous, with three-segmented rami. Second
segment of endopod of legs 1-3 with two inner setae. Leg 5 two-segmented, first segment with outer seta, second segment withfour setae in female and five setae in male.
Biology: Females and males have been found in washings oftube worms. All congeners are endosymbionts living in thestomach of shallow-water ophiuroids (HUMES 1993).
Distribution: Juan de Fuca Ridge: Cleft Segment, Marker M ofNorth Field, 2253 m.
References:
HUMES A.G.(1993) Bijdr. Dierkd. 63: 121-127.HUMES A.G. (1999) Proc. Biol. Soc. Wash. 112(1): 181-188.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 324
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Asterocheridae
1: Female; A: Antennule; B: Antenna; C: Mandible; D: Maxilla; E: Maxilliped; F: Leg 5; G: Male leg 5. ex – exopod of antenna represented by seta; gn – gnathobase of mandible; pl – palp of mandible; from HUMES (1999).
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Hyphalion captans HUMES, 1987
Size: Body length of female 1.94-2.17 mm; male 1.71-1.82 mm.
Morphology: Body elongate, flattened. Prosome four-segment-ed. Urosome six-segmented in female, five-segmented in male;first somite with leg 5. Rostrum broad, projecting anterior. An-tennule six-segmented in both sexes; male antennules non-geniculate. Antenna three-segmented, without exopod; distalsegment with three stout recurved claws. Mandible flexed,gnathobase with three spines and one seta; palp absent. Lobe ofmaxillule with five setae. Maxilla two-segmented; first segmentwith one seta; second segment with one seta and three spines.Maxilliped sexually dimorphic; maxilliped of female two-seg-
mented, second segment with long terminal seta; maxilliped ofmale subchelate, four-segmented, second segment triangularwith two rows of spinules, third segment not well defined,fourth segment claw-like. Legs 1-4 biramous with three-seg-mented rami. Leg 5 two-segmented, first segment with one se-ta; second segment with three spines and one seta.
Biology: Symbiont of bivalves; type of feeding unclear, mayfeed on tissues, mucus or utilize host’s food.
Distribution: Guaymas Basin.
References:
DEFAYE D. & T. TODA (1994) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 16, section A (1): 87-94.HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 325
Arthropoda, Crustacea, Copepoda, Cyclopoida, Clausidiidae
1: Female; A: Habitus, dorsal; B: Antennule; C: Antenna; D: Maxilliped; E: Male habitus, dorsal; F: Maxilliped; from HUMES (1987).
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Barathricola rimensis HUMES, 1999
Size: Body length of female 0.67-0.79 mm; male 0.47-0.57 mm.
Morphology: Prosome slender, five-segmented. Urosome five-segmented in female, six-segmented in male, first somite withleg 5. Caudal ramus long, of length/width ratio 11:1. Anten-nule thirteen-segmented; geniculation of male antennules be-tween segments 11 and 12. Antenna four-segmented, withoutexopod. Mandible with biramous palp: endopod two-segment-ed; exopod small, one-segmented, with two terminal setae. Legs
1-4 biramous, with three-segmented rami. Leg 1 with one setaon inner segment. Leg 5 three-segmented with intercoxal scle-rite: armature 0-0; 1-0; II, 1, I in female and 0-0; 1-0; II, 1, I, 1in male.
Distribution: Juan de Fuca Ridge: Coaxial Segment, Flow ventsite (Vent HDV).
Reference:
HUMES A.G. (1999) J. Nat. Hist. 33: 961-978.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 326
Arthropoda, Crustacea, Copepoda, Cyclopoida, Cyclopinidae
1: Female; A: Habitus, dorsal; B: Urosome, dorsal; C: Antenna; D: Mandible; E: Maxilliped; F: Leg 5; from HUMES (1999).
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Heptnerina confusa IVANENKO & DEFAYE, 2004
Size: Body length of female 0.82 mm; male 0.65 mm.
Morphology: Prosome five-segmented; tergite of somite bearingleg 1 covered laterally by shield of cephalosome. Urosome five-segmented in female, six-segmented in male; first somite withleg 5. Antennule ten-segmented in female and seventeen-seg-mented in male. Antenna four-segmented; exopod representedby two setae on first segment. Mandible with biramous palp.Legs 1-4 biramous, with three-segmented rami. Endopod of leg4 with three robust setae. Leg 5 two-segmented with armature1-0; I, 1, I in female and 1-0; I, 1, I, 2 in male. First segment offemale leg 5 can have stout endopodal element.
Biology: Many adult and subadult copepodid stages of H. con-fusa have been collected during in situ colonization experi-ments at the base of the active chimney Eiffel Tower covered bya layer of Bathymodiolus azoricus (Bivalvia, Mytilidae).
Distribution: Mid-Atlantic Ridge: Lucky Strike, Chimney Eif-fel Tower.
Reference:
IVANENKO V.N. & D. DEFAYE (2004) Zoosystema 26 (1): 49-64.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 327
Arthropoda, Crustacea, Copepoda, Cyclopoida, Cyclopinidae
1: Female; A: Habitus, dorsal; B: Antennule; C: Antenna; D: Leg 5, endopodal element arrowed; E: Male habitus, dorsal;from IVANENKO & DEFAYE (2004).
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Aphotopontius HUMES, 1987
Size: Body length 0.6-1.2 mm.
Morphology: Prosome four-segmented. Urosome five-segment-ed in female, six-segmented in male; first somite with leg 5.Oral cone short. Antenna with one-segmented exopod.Mandible with stylet-like gnathobase, without palp. First seg-ment of maxilla with inner sinuous seta. Exopods of legs 1-4and endopods of legs 1-3 three-segmented. Endopod of leg 4two-segmented; first segment with one inner seta; second seg-ment with two (terminal and inner) setae.
Remarks: A. rapunculus HUMES & SEGONZAC, 1998 transferredto Rhogobius HUMES, 1987; A. temperatus HUMES, 1997 was syn-onymized with A. atlanteus. Only females are known for A. flex-ispina and A. hydronauticus.
Biology: All species of Aphotopontius have been found free-liv-ing or associated with invertebrates. A. arcuatus, A. atlanteus,A. flexispina and A. probolus found in association with bivalves;A. arcuatus, A. flexispina and A. mammillatus with vestimen-tiferans; A. acanthinus and A. forcipatus with crustaceans. Eco-logical studies at the Juan de Fuca Ridge showed that A. forci-patus is more abundant at new vents appearing after eruptionand has female to male ratio 10.6:1 (TSURUMI et al. 2003).Functional analysis suggests that copepods of the genus Aphoto-pontius are swimming and moving over substratum, they canfeed on bacterial films and flakes on substratum and in near-bottom water column (HEPTNER & IVANENKO 2002).
Distribution: See the table after Ivanenko & Ferrari, 2003: 1 –Galapagos Rift; 2 – East Pacific Rise at 10°N; 3 – East PacificRise at 13°N; 4 – East Pacific Rise at 21°N; 5 – Guaymas Basin,27°N; 6 – Gorda Ridge, 41°N; 7 – Juan de Fuca Ridge, 46°N; 8– Explorer Ridge, 49°N; 9 – Mid-Atlantic Ridge at 37°N(Lucky Strike); 10 – Mid-Atlantic Ridge at 37°N (MenezGwen); 11 – Mid-Atlantic Ridge at 23°N (Snake Pit).
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
Species 1 2 3 4 5 6 7 8 9 10 11
A. acanthinus HUMES & LUTZ, 1994 +
A. arcuatus HUMES, 1987 + + +
A. atlanteus HUMES, 1996 + +
A. baculigerus HUMES, 1987 + +
A. flexispina HUMES, 1987 +
A. forcipatus HUMES, 1987 + + + +
A. hydronauticus HUMES, 1989 +
A. limatulus HUMES, 1987 + + +
A. mammillatus HUMES, 1987 + + + +
A. probolus HUMES, 1990 +
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 328-329
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References:
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta 11(2): 117-134.HUMES A.G. (1987) Bull. Mar. Sci. 41(3): 645-788.HUMES A.G.(1990) Sci. Mar. 54(2): 145-154.HUMES A.G. & R.A. LUTZ (1994) Jour. Crustac. Biol. 14(2): 337-345.HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11(3): 177-185. IVANENKO V.N & M.V. HEPTNER (1998) J. Mar. Syst. 15: 243-254.TSURUMI M., DE GRAAF R.C. & V. TUNNICLIFFE (2003) J. Mar. Biol. Ass. U.K. 83(3): 469-478.
1: Female of A. mammillatus; A: Habitus, dorsal; B: Habitus, lateral; C: Antenna; D: Mandible; E: Maxilla; F: Leg 4. b – basis; c – coxa; en1-en2 – endopodalsegments; ex1-ex3 – exopodal segments 1-3; gn – gnathobase of mandible; p1-p4 – segments of prosome; ss – sinuous seta of maxilla; u1-u5 – segments of urosome;from IVANENKO & HEPTNER (1998).
2: Female of A. atlanteus (SEM); A: Habitus, dorsal; B: Habitus, ventral;
C: Distal part of oral cone; D: Urosome, dorsal. es – egg sac; lb – distal part of labrum
of oral cone, oral cone, distal part of labrum; lm – distal part of labium of oral cone
ornamented with setiform elements;md – mandible, tip of gnathobase; original.
329
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Benthoxynus HUMES, 1984
References:
HOURDEZ S., LAMONTAGNE J., PETERSON P., WEBER R.E. & C.R. FISHER (2000) Biol. Bull. 199: 95-99.HUMES A.G. (1984) Can. J. Zool. 62: 2594-2599.HUMES A.G. (1989) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 11, section A, no. 4: 829-849.TSURUMI M., DE GRAAF R.C. & V. TUNNICLIFFE (2003) J. Mar. Biol. Ass. U.K. 83(3): 469-478.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 330
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1: Female of B. tumidiseta; A: Habitus, dorsal; B: Anal somite and elongate caudal ramus with reduced innermost terminal seta,dorsal; C: Antennule; D: Maxilla; female of B. spiculifer; E: Antennule; F: Leg 4. is – innermost terminal seta of caudal ramus;from HUMES (1989) (A-D) and HUMES (1984) (E, F).
Species Distribution Body length in mmB. spiculifer HUMES, 1984 Juan de Fuca Ridge: Axial Seamount, Heineken Hollow, Female 1.61-1.79; male 0.97-1.09
Megaplume Site; Gorda Ridge; Explorer Ridge
B. tumidiseta HUMES, 1989 East Pacific Rise: 13°N, sites Genesis, Elsa, Totem Female 1.53-1.76; male unknown
Morphology: Prosome four-segmented. Urosome five-segment-ed in female, six-segmented in male; first somite with leg 5.Caudal ramus elongate; innermost terminal seta minute. Oralcone short. Antenna with one-segmented exopod. Mandiblewith stylet-like gnathobase, without palp. First segment of max-illa with inner sinuous seta. Exopods of legs 1-4 and endopodsof legs 1-3 three-segmented. First exopodal segment of leg 3lacks inner seta. Leg 4 with two-segmented endopod; first seg-ment unarmed; second segment armed with terminal spine. Leg5: small lobe with 3 setae in female and two setae in male.
Biology: B. spiculifer has been found in debris from vestimen-tiferan tubes. Ecological studies at the Juan de Fuca Ridge re-vealed that it is more common at older vents (TSURUMI et al.2003). The blood of B. spiculifer has haemoglobin supportingaerobic respiration in low-oxygen conditions (HOURDEZ et al.2000).
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Ceuthoecetes HUMES & DOJIRI, 1980
Type species: C. aliger HUMES & DOJIRI, 1980. Other includedspecies: C. acanthothrix HUMES, 1987; C. cristatus HUMES, 1987;C. introversus HUMES, 1987.
Size: Body length 0.91-1.50 mm.
Morphology: Prosome four-segmented. Urosome five-segment-ed in female, six-segmented in male; first somite with leg 5.Oral cone short, with distal disk formed by labium. Antennawith small one-segmented exopod; endopod with curved termi-nal claw. First segment of maxilla without inner sinuous seta;second segment elongate, with distal setiform part. Exopods oflegs 1-4 and endopods of legs 1-3 three-segmented. Leg 4 with
two-segmented endopod; first segment unarmed; second seg-ment armed with terminal spine and inner seta. Maxilliped ofmale with stout second segment.
Remarks: Only female was described for C. aliger; only malesare known for three other species.
Biology: Abundant in washings of vestimentiferans and Calyp-togena. Type of feeding unclear, may feed on tissues or associat-ed mucus of their hosts. Ceuthoecetes aliger was found in wash-ing of tentacular crowns of vestimentiferans.
Distribution: East Pacific Rise, Galapagos Spreading Center.
2: Male of C. introversus (SEM). A: Habitus, dorsal;B: Habitus, ventral; C: Oral cone, ventral. a1 – antennule; cr – caudal ramus; dd – distal disk oforal cone formed by labium; l1-l3 – swimming legs1-3. lb – labium; m1 – maxillule; m2 – maxilla; mp – maxilliped; p1-p4 – segments of prosome; u1-u6 – somites of urosome; original.
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1: Female of C. aliger; A: Habitus, dorsal; B: Antenna; C: Maxilla; D: Leg 4; E: Male of C. cristatus, habitus, dorsal;F: Male of C. introversus, maxilla. ex – exopod; tc – terminal claw of endopod; from HUMES & DOJIRI (1980) (A-D) and HUMES (1987) (F).
References:
HUMES A.G. (1987) Bull. Mar. Sci. 41(3): 645-788.HUMES A.G. & M. DOJIRI (1980) Proc. Biol. Soc. Wash. 93(3): 697-707.HUMES A.G. & M. SEGONZAC (1998) Cah.Biol. Mar. 39: 51-62.IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11(3): 177-185.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 331
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Chasmatopontius thescalus HUMES, 1990
Size: Body length of female 1.14-1.39 mm; male 1.06-1.28 mm.
Morphology: Prosome four-segmented. Urosome four-segment-ed in female, five-segmented in male; first somite with leg 5.Copulatory pores near posterior border of female genital dou-ble-somite. Oral cone short. Antenna with one-segmented ex-opod. Mandible with stylet-like gnathobase, without palp. Firstsegment of maxilla with inner sinuous seta. Exopods of legs 1-4and endopods of legs 1-3 three-segmented. Leg 4 with two-seg-mented endopod; first segment unarmed; second segment withthree inner setae. Second segment of male maxilliped expand-ed, with knob on inner edge.
Biology: Abundant. Found in washings of tubes of Paralvinellahessleri DESBRUYÈRES & LAUBIER, 1989 in the Mariana Back-Arc Basin.
Distribution: Mariana Back-Arc Basin: Alice Spring, Illium;Lau Back-Arc Basin: Vailili.
References:
HUMES A.G. (1990) J. Nat. Hist. 24: 289-304.HUMES A.G. (1991) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 13, section A, nos. 1-2: 121-134.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 332
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1: Female; A: Habitus, dorsal; B: Urosome, dorsal; C: Maxilla; D: Male leg 4, anterior; E: Maxilliped, posterior. cp – copulatory pore; kb – knob on inner edge of male maxilliped; from HUMES (1990).
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Dirivultus HUMES & DOJIRI, 1980
References:
HUMES A.G. (1999) J. Mar. Biol. Ass. U.K. 79: 1053-1060.HUMES A.G. & M. DOJIRI (1980) Pac. Sci. 34: 143-151.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 333
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1A: Female habitus, dorsal; B: Habitus lateral; C: Antenna, posterior; D: Maxilla; E: Leg 4; F: Male habitus, dorsal; G: Femalehabitus, dorsal; H: Egg sac. ec – egg sac; ex – exopod; s2 – second segment of maxilla ornamented with setules; tc – terminalclaw of endopod; from HUMES & DOJIRI (1980) (A-F D. dentaneus) and HUMES (1999) (G-H D. spinigulatus, female).
Type species: D. dentaneus HUMES & DOJIRI, 1980; one furtherspecies D. spinigulatus HUMES, 1999.
Size: Body length of female 1.00-1.16 mm; male 0.92-1.09 mm.
Morphology: Prosome four-segmented. Urosome five-segment-ed in female, six-segmented in male; first somite with leg 5.Rostral area projecting anteriorly. Oral cone with distal diskformed by labium; labrum with dentiform spines directed pos-teriorly. Antenna with one-segmented exopod; endopod withcurved terminal claw. Mandible with stylet-like gnathobase,without palp. First segment of maxilla without inner sinuous se-ta. Second segment of maxilla armed with long slender clawand ornamented with long setules. Exopods of legs 1-4 and en-
dopods of legs 2-3 three-segmented; endopods of legs 1 two- orthree-segmented. Leg 4 with two-segmented endopod; first seg-ment unarmed; second segment armed with terminal spine andinner seta.
Biology: May “feed on tentacular lamellae or associated mucus”of siboglinids (HUMES & DOJIRI 1980). The type species D. den-taneus HUMES & DOJIRI, 1980 has been found in tentacularcrown of Lamellibrachia barhami WEBB, 1969 off Southern Cali-fornia.
Distribution: D. dentaneus from Southern California(32°19’6’’N, 117°19’08’’W; 1125 m); D. spinigulatus fromTabar-Feni Volcanic Fore-Arc: Edison Seamount.
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Exrima HUMES, 1987
References:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.HUMES A.G. (1989) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 11, section A, no. 4: 829-849.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 334
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1: Female of E. singula; A: Habitus, dorsal; B: Antenna; C: Maxilla; D: Leg 4; E: Female of E. dolichopus, habitus, dorsal; from HUMES (1987).
Species Distribution Body length in mmE. dolichopus HUMES, 1987 East Pacific Rise: 13°N, Parigo Female 1.12-1.15; male unknown
E. singular HUMES, 1987 East Pacific Rise: 21°N, Clam Acres Female 0.97-1.07; male unknown
Morphology: Prosome four-segmented. Urosome five-segment-ed; first somite with leg 5. Oral cone short. Antenna with one-segmented exopod; distal segment of endopod with two longterminal setae. Mandible with stylet-like gnathobase, withoutpalp. First segment of maxilla with inner sinuous seta. Exopodsof legs 1-4 and endopods of legs 1-3 three-segmented. Leg 4with two-segmented endopod; first segment unarmed; secondsegment armed with terminal spine. Leg 5 two-segmented.
Biology: E. singula has been found in washing of vestimentifer-ans and Calyptogena.
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Fissuricola caritus HUMES, 1987
References:
HUMES A.G. (1987) Bull. Mar. Sc. 41: 645-788.IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11 (3): 177-185.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 335
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
Size: Body length of female 1.19-1.34 mm. Male unknown.
Morphology: Prosome four-segmented. Urosome five-segment-ed; first somite with leg 5. Oral cone short. Antenna with one-segmented exopod. Mandible with stylet-like gnathobase, with-out palp. Maxilla reduced to small vestige. Exopods of legs 1-4and endopods of legs 1-3 three-segmented. Leg 4 with two-seg-mented endopod; first segment with inner seta; second segmentwith one terminal spine and three inner setae.
Remark: The only dirivultid with maxilla reduced to vestige.
Distribution: East Pacific Rise: 21°N, Clam Acres.
1: Female; A: Habitus, dorsal; B: Antenna; C: Maxilla; D: Leg 4; E: Leg 5; from HUMES (1987).
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Humesipontius arthuri IVANENKO & FERRARI, 2003
Reference:
IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11(3): 177-185.
V.N. IVANENKO & D. DEFAYE Denisia 18: 336
1: Female; A: Habitus, dorsal; B: Antenna; C: Maxilla; D: Maxilliped, posterior; E: Leg 4, anterior; after IVANENKO & FERRARI (2003).
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
Size: Body length of female 1.8 mm. Male unknown.
Morphology: Body elongate. Prosome four-segmented. Uro-some five-segmented; first somite with leg 5. Oral cone short.Antenna with one-segmented exopod; second segment of en-dopod with elongate terminal seta. Mandible with stylet-likegnathobase, without palp. First segment of maxilla with innersinuous seta. First segment of maxillipedal endopod with elon-gate seta. Exopods of legs 1-4 and endopods of legs 1-3 three-segmented. Leg 4 without endopod.
Remark: The only dirivultid missing endopod of leg 4.
Distribution: Juan de Fuca Ridge.
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Nilva torifera HUMES, 1987
Size: Body length of female 0.67-0.88 mm, male 0.79-0.96 mm.
Morphology: Prosome four-segmented. Urosome five-segment-ed in female, six-segmented in male; first somite with leg 5; lasttwo somites indistinctly separated. Genital double-somite of fe-male and genital somite of male with dorsal hump. Oral coneshort, with distal disk formed by labium. Antenna with smallone-segmented exopod; endopod with curved terminal claw.First segment of maxilla without inner sinuous seta; second seg-ment elongate with setiform distal part. Exopods of legs 1-4 andendopods of legs 1-3 three-segmented. Leg 4 with two-seg-mented endopod; first segment unarmed; second segmentarmed with inner seta and terminal spine.
Biology: Common in washings of vestimentiferans and Calyp-togena; type of feeding unclear, may feed on tissues or mucus ofhosts.
Distribution: Galapagos Spreading Center; East Pacific Rise:21°N, Clam Acres: 13°N, Parigo, Totem, Genesis.
Reference:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 337
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1: Female; A: Habitus, dorsal; B: Urosome, lateral; C: Antenna; D: Maxilla; E: Male habitus, dorsal; F: Urosome, lateral. dh – dorsal hump; tc – terminal claw of antennal endopod; from HUMES (1987).
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Rhogobius HUMES, 1987
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
Species Distribution Body length in mmR. contractus HUMES, 1987 Galapagos Spreading Center; East Pacific Rise: 13°N, 21° Female 0.7-0.94, male 0.6-0.71
R. pressulus HUMES, 1989 Galapagos Spreading Center Female 0.67-0.78; male unknown
R. rapunculus (HUMES & SEGONZAC, 1998) East Pacific Rise: 9°N Female 0.89-0.91; male unknown
References:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.HUMES A.G. (1989) Pac. Sci. 43: 27-31.HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 338
Morphology: Body with broad prosome. Prosome four-segment-ed. Urosome five-segmented in female, six-segmented in male;first somite with leg 5. Last abdominal somite with terminalpaired lobes. Oral cone short. Antenna with one-segmentedexopod; second segment of endopod elongate and slender.Mandible with stylet-like gnathobase, without palp. First seg-ment of maxilla without inner sinuous seta. Exopods of legs 1-4 and endopods of legs 1-3 three-segmented. Leg 4 with two-
segmented endopod; first segment with inner seta; second seg-ment with terminal spine and inner seta.
Remark: R. rapunculus (HUMES & SEGONZAC, 1998) transferredfrom Aphotopontius.
Biology: R. contractus has been found in number of samples(HUMES & SEGONZAC 1998). Only two females of R. rapunculusand 10 females of R. pressulus have been found until now.
1: Female of R. contractus.; A: Habitus, dorsal; B: Genital double-somite, abdominal somites and caudal rami; C: Antenna; D: Maxilla; E: Leg 5, male; F: Habitus,dorsal; G: R. rapunculus. Female leg 5; H: Habitus, dorsal; I: Leg 4; J: Female of R. pressulus, leg 5; K: Genital double-somite andabdominal somites. s2 – second segment of endopod; tl – terminal lobe on anal somite;from HUMES (1987) (A-G), HUMES &SEGONZAC (1998) (H-J) and HUMES
(1989) (K).
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Rimipontius mediospinifer HUMES, 1996
References:
HUMES A.G. (1996) Bull. Mar. Sci. 58: 609-653.HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.IVANENKO V.N. (1998) Zool. Zh. 77(11): 1249-1256.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 339
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1: Female; A: Habitus, dorsal; B: Habitus, lateral; C: Anal somite and caudal ramus, dorsal; D: Maxilla; E: Leg 4; F: Male habitus, dorsal. cr – caudal ramus; ds – dorsal spiniform process of genital double somite; en – endopod of leg 4; ss – sinuous seta of maxilla; from HUMES (1996).
Size: Body length of female 0.84-1.01 mm; male 0.57-0.69 mm.
Morphology: Prosome four-segmented. Urosome five-segment-ed in female, six-segmented in male; first somite with leg 5.Genital double-somite of female with dorsal longitudinal crest.Caudal rami with 5 setae: innermost terminal seta reduced.Oral cone short. Antenna with one-segmented exopod.Mandible with stylet-like gnathobase, without palp. First seg-ment of maxilla with inner sinuous seta. Exopods of legs 1-4and endopods of legs 1-3 three-segmented. Leg 4 with two-seg-mented endopod, first segment unarmed; second segment withterminal spine.
Biology: Found in washing of the decapod Rimicaris (HUMES
1996) and in plankton 80-300 m over the hydrothermal fieldamong shrimps and other copepods (IVANENKO 1998). It can“feed on the bacteria on the mouthparts of the shrimps, but al-so, when they are free, perhaps on the bacteria free in the wa-ter” (HUMES 1996).
Distribution: Mid-Atlantic Ridge: Logatchev, Irina-2; SnakePit, Broken Spur (80-300 m over Saracen’s Head).
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Scotoecetes introrsus HUMES, 1987
References:
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta, 11(2): 117-134.HUMES A.G.(1987) Bull. Mar. Sci. 41: 645-788.HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.SELL A.F. (2000) Proc. R. Soc. London 267: 2323-2326.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 340
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1: Female; A: Habitus, dorsal; B: Maxilla; C: Leg 1; D: Leg 4; E: Male habitus, dorsal. lb – labrum of oral cone; a2 – antenna; m1 – maxillule; m2 – maxilla; from HUMES (1987).
Size: Body length of female 1.65-1.79 mm; length of male 1.30-1.42 mm.
Morphology: Prosome four-segmented. Urosome five-segment-ed in female, six-segmented in male; first somite with leg 5.Oral cone short. Antenna with one-segmented exopod.Mandible with stylet-like gnathobase, without palp. First seg-ment of maxilla with inner sinuous seta. Exopods of legs 1-4and endopods of legs 2-3 three-segmented. Endopod of leg 1two-segmented in female and three-segmented in male; distalsegment of endopod with outer terminal seta directed inward.Leg 4 with two-segmented endopod; first segment unarmed,second segment with terminal spine and inner seta.
Biology: Abundant, associated with siboglinids, feeding onbacterial flakes; different copepodid stages have been found inwashings of vestimentiferans collected at 9°N (unpublished da-ta). The blood of S. introrsus as well as that of Benthoxynus spi-culifer has haemoglobin supporting aerobic respiration in low-oxygen conditions (SELL 2000).
Distribution: East Pacific Rise: 9°N, 13°N.
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Stygiopontius HUMES, 1987
Size: Body length 0.6-1.6 mm.
Morphology: Prosome four-segmented. Urosome five-segment-ed in female, six-segmented in male; first somite with leg 5.Oral cone short. Antenna with one-segmented exopod.Mandible with stylet-like gnathobase, without palp. First seg-ment of maxilla with inner sinuous seta. Exopods of legs 1-4and endopods of legs 1-3 three-segmented. Leg 4 with two-seg-mented endopod; first segment unarmed; second segmentarmed with terminal spine and inner seta.
Remarks: S. lumiger HUMES, 1989 and S. bulbisetiger HUMES,1996 were synonymized with S. sentifer and S. pectinatus, re-spectively. Only females are known for eight species (S.cinctiger, S. flexus, S. hispidulus, S. mucroniferus, S. pectinatus, S.sentifer, S. stabilitus, and S. teres), and only males are known forsix species (S. appositus, S. latulus, S. mirus, S. paxillifer, S. rimi-vagus, and S. verruculatus).
Biology: All species of the genus Stygiopontius have been foundfree-living or associated with invertebrates. S. hispidulus and S.rimivagus have been found in association with bivalves; S.hispidulus, S. sentifer, and S. stabilitus with polychaetes; S. flexuswith siboglinids; S. cladarus and S. pectinatus with crustaceans.S. sentifer lives in tubes of polychaete Alvinella pompejana DES-BRUYÈRES & LAUBIER, 1980. S. pectinatus has been found in gillcavity of the shrimps Rimicaris exoculata WILLIAMS & RONA,1986 and Chorocaris chacei (WILLIAMS & RONA, 1986) and free-living in plankton over hydrothermal field (IVANENKO 1998). S.quadrispinosus is more abundant at high temperature at the Juande Fuca Ridge and has the ratio of females to males 7.6:1 (TSU-RUMI et al. 2003).
Distribution: See the table after Ivanenko & Ferrari (2003): 1– East Pacific Rise at 10°N; 2 – East Pacific Rise at 13°N; 3 –East Pacific Rise at 21°N; 4 – Guaymas Basin, 27°N; 5 – Gor-da Ridge, 41°N; 6 – Juan de Fuca Ridge, 46°N; 7 – ExplorerRidge, 49°N. WP – West Pacific: 8 – Mariana Back-Arc Basin,18°N; 9 – Lau Back-Arc Basin, 23°S; 10 – Mid-Atlantic Ridgeat 37°N (Lucky Strike); 11 – Mid-Atlantic Ridge at 29°N (Bro-ken Spur); 12 – Mid-Atlantic Ridge at 26°N (TAG); 13 – Mid-Atlantic Ridge at 23°N (Snake Pit).
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae
1 2 3 4 5 6 7 8 9 10 11 12 13S. appositus + +S. brevispina +S. cinctiger + + +S. cladarus + +S. flexus + +S. hispidulus + + +S. latulus +S. lauensis +S. mirus + +S. mucroniferus + +S. paxillifer + +S. pectinatus + + + +S. quadrispinosus + + +S. regius +S. rimivagus +S. sentifer + + +S. serratus +S. stabilitus + +S. teres +S. verruculatus + +
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 341–342
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References:
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta 11(2):117-134.HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.IVANENKO V.N. (1998) Zool. Zh. 77 (11): 1249-1256.IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11 (3): 177-185.TSURUMI M., DE GRAAF R.C. & V. TUNNICLIFFE (2003) J. Mar. Biol. Assoc. U.K. 83(3): 469-478
1: Female of S. quadrispinosus;A: Habitus, dorsal; B: Maxilla; C: Leg 4; D: Male habitus,dorsal; from HUMES (1987).
2: S. pectinatus ventral view (left)and dorsal view (right); original.
342
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Ecbathyrion prolixicauda HUMES, 1987
Size: Body length of female 1.01-1.11 mm; male 0.77-0.84 mm.
Morphology: Prosome four-segmented. Urosome four-segment-ed in female and five-segmented in male; first somite with leg5. Caudal ramus elongate. Oral cone short. Antennule eight-een-segmented in female and seventeen-segmented in male;aesthetasc on segments 15 and 17 in female and male, respec-tively. Antenna with one-segmented exopod having pointedprojection. Mandible with stylet-like gnathobase, without palp.First segment of maxilla with inner sinuous seta. Legs 1-4 bira-
mous, with three-segmented rami. Basis of leg 1 without innerspine or seta. Leg 5: first segment fused with somite and repre-sented with one outer seta; second segment bearing three setaein female, and five setae in male.
Remarks: Found in many samples.
Distribution: Galapagos Rift; East Pacific Rise: 9°N, 13°N,21°N.
References:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 343
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Ecbathyriontidae
1: Female; A: Habitus, dorsal; B: Urosome, dorsal; C: Antennule; D: Antenna; E: Maxilla; F: Leg 4; G: Male habitus, dorsal. ae – aesthetasc; cr – caudal ramus; ds – genital double-somite; en – endopod; ex – exopod; ss – sinuous seta; from HUMES (1987).
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Ambilimbus IVANENKO, DEFAYE & HUYS, 2005
Arthropoda, Crustacea, Copepoda, Cyclopoida, Erebonasteridae
1: A. arcuscelestis, female; A: Habitus, dorsal; B: Mandible; C: Maxilla; D: Maxilliped; E: Male habitus, dorsal; F: Maxilliped; from IVANENKO et al. (2005).
Species Distribution Body length in mmA. altalis (HUMES & HUYS, 1992) Juan de Fuca Ridge Female 2.41-2.46; male 1.98-2.13
A. arcuscelestis IVANENKO, DEFAYE & HUYS, 2005 Mid-Atlantic Ridge: Rainbow Female 2.35; male 1.65
A. tuerkayi (MARTINEZ-ARBIZU, 1999) Fiji Basin Female 1.51; male 1.2
Synonym: Amphicrossus HUYS, 1991
Morphology: Body elongate and flattened, ornamented withspinules. Prosome five-segmented, epimera of somites bearinglegs 1-4 rounded. Urosome five-segmented in both sexes; firstsomite with leg 5; copulatory pore on ventral side of genitaldouble-somite. Rostrum prominent with row of dentiform spin-ules and long setules. Antennule six-segmented in both sexes.Antenna four-segmented, without exopod. Mandibulargnathobase with four spines, mandibular palp one-segmentedwith four setae. Maxillule represented by lobe with five setae.
Maxilla two-segmented, distal segment claw-like, with innerprocess. Maxilliped four-segmented, sexually dimorphic, distalsegment claw-like, with inner process. Legs 1-4 biramous withthree-segmented rami. Leg 5 two-segmented.
Biology: Deep-sea poecilostomatoid cyclopoids found in sedi-ment. Most likely associated with bivalves. Type of feeding un-clear; can feed on tissues, mucus or utilize hosts food. Ere-bonaster protentipes HUMES, 1987 has been found in mantle cav-ity and in washings of a Nuculana-like protobranch bivalvefrom cold seeps at Guaymas Basin.
References:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.HUMES A.G. & R. HUYS (1992) Can. J. Zool. 70: 1369-1380.IVANENKO V.N., DEFAYE D. & R. HUYS (2005) Marine Biology Research 19: 93-105.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 344
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Bathylaophonte LEE & HUYS, 1999
Morphology: Body elongate, sub-cylindrical, with reticulatedintegument and spinules. Prosome four-segmented; urosomefive-segmented in female and six-segmented in male; genitaldouble-somite of female subdivided dorsally. Rostrum bell-shaped. Anal operculum well-developed. Antennule seven-seg-mented in female, eight-segmented in male. Antenna withone-segmented exopod bearing four setae. Maxilliped sub-chelate, three-segmented. Legs 1-4 biramous, with three-seg-mented exopods and two-segmented endopods, except malewith three-segmented endopod of leg 3. Endopod of leg 1 pre-
hensile, first segment longer than exopod, first segment un-armed, second segment with claw and one seta. Leg 5 two-seg-mented; endopodal lobe of female well developed, with four se-tae; endopodal lobe of male reduced, with one seta; exopodwith five setae in female and four setae in male.
Biology: Both species were “extracted from the invertebratesliving in the hydrothermal fluids”.
Distribution: East Pacific Rise: 17°S north of Easter Island.
Reference:
LEE W. & R. HUYS (1999) Cah. Biol. Mar. 40: 293-328.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 345
Arthropoda, Crustacea, Copepoda, Harpacticoida, Laophontidae
1: B. azorica; A: Female habitus, lateral; B: Leg 5, anterior; C: Male leg 5, anterior; D: Female of B. pacifica, leg 1,anterior; en1 – first endopodal segment; from LEE & HUYS (1999).
Species Distribution Body length in mmB. azorica LEE & HUYS, 1999 Mid-Atlantic Ridge: Menez Gwen, Lucky Strike Female 0.69; male 0.66
B. pacifica LEE & HUYS, 1999 East Pacific Rise:17°S north of Easter Island Female 0.81; male 0.74
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Laitmatobius crinitus HUMES, 1987
Size: Body length of male 1.29-1.38 mm. Female unknown.
Morphology: Body slender. Prosome five-segmented. Urosomesix-segmented, first somite with leg 5, second somite elongate.Rostrum rounded lobe in lateral view. Caudal ramus elongate,length/width ratio 2.7:1. Antennule five-segmented. Antennafour-segmented with four terminal setiform claws and two sub-terminal setae. Mandible elongate, without palp; basal partwith two spines and one seta, distal part forming long lash.Maxillule bilobed. Maxilla two-segmented; second segmentwith one seta and two spines. Maxilliped subchelate, four-seg-mented. Legs 1-4 biramous, with three-segmented rami; basis ofleg 1 with inner spine. Leg 5 represented by small segment bear-ing two setae and adjacent seta.
Biology: More than 60 males of L. crinitus have been found intwo samples. Other family members occur in plankton at dif-ferent depths and probably associated with pelagic inverte-brates (HUYS & BÖTTGER-SCHNACK 1997 cited in BOXSHALL &HALSEY 2004).
Distribution: Guaymas Basin.
References:
BOXSHALL G.A & S.H. HALSEY (2004) An Introduction to Copepod Diversity. Ray Society, London: i-xv, 1-966.HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 346
Arthropoda, Crustacea, Copepoda, Cyclopoida, Lubbockiidae
1: Male; A: Habitus, dorsal; B: Rostrum, lateral; C: Antenna; D: Mandible; E: Maxilla; F: Maxilliped; from HUMES (1987).
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Hyalopontius boxshalli HUMES, 1988
Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Megapontiidae
Size: Body length of male 4.75-4.86 mm. Female unknown.
Morphology: Body elongate, 5.7 times as long as wide. Prosomefour-segmented, epimera pointed posteriorly. Urosome six-seg-mented, first somite with leg 5. Oral cone elongate. Antennuleeleven-segmented. Antenna with short one-segmented exopodbearing three long setae and two-segmented endopod bearingtwo unequal terminal setae. Mandible with stylet-likegnathobase, without palp. Maxillule with lobe bearing three se-tae and one adjusted to lobe seta. Maxilla two-segmented.
Maxilliped subchelate, three-segmented with long terminalclaw. Legs 1-4 biramous, with three-segmented rami. Leg 5 one-segmented, armed with three setae; one outer seta near seg-ment.
Biology: Two males have been found only. Functional analysissuggests that megapontiids suck out blood or soft tissues of un-known hosts (HEPTNER 1968).
Distribution: Galapagos Spreading Center.
References:
HEPTNER M.V. (1968) Zool. Zh. 47(11):1628-1638.HUMES A.G. (1988) Proc. Biol. Soc. Wash. 101(4): 825-831.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 347
1: Male; A: Habitus, dorsal; B: Oral cone, lateral;C: Antenna; from HUMES (1988).
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Stenhelia gundulae WILLEN, 2003
Size: Body length of female 0.89 mm; male 0.81 mm.
Morphology: Body with distinct separation between prosomeand urosome. Prosome four-segmented; urosome five-segment-ed in female and six-segmented in male. Rostrum separatedfrom cephalothorax, triangular, with bifid tip. Anal operculumlarge and prominent. Caudal rami 2.6 times as long as wide.Antennule short, eight-segmented in female and ten-segment-ed in male. Antennary exopod three-segmented, with one, one,four setae. Maxilliped three-segmented. Legs 1-4 biramous,with three-segmented rami, except two-segmented endopod ofleg 1 in female and of legs 1 and 2 in male. Leg 5 with exopodarticulated with baseoendopod in female and fused in male;
baseoendopod with one outer and four or two inner setae in fe-male and male, respectively; exopod with five setae in femaleand four setae in male.
Biology: Found only in “the North and South Clam fields, witha relatively high abundance”. The samples with the copepods“contained basaltic pepperite with numerous small xenoliths aswell as a great number of vesicomyiid clams and a rich associ-ated fauna indicating hydrothermal activity” (WILLEN 2003).
Distribution: Tabar-Feni Volcanic Fore-Arc: Edison Seamount,Southern Clam field, North Clam field.
Reference:
WILLEN E. (2003) J. Nat. Hist. 37(14): 1691-1711.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 348
Arthropoda, Crustacea, Copepoda, Harpacticoida, Miraciidae
1A: Female habitus, dorsal; B: Maxilliped; C: Leg 5; D: Male habitus, dorsal; E: Leg 5; from WILLEN (2003).
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Misophriopsis longicaudata HUMES, 1999
References:
BOXSHALL G.A. (1982) Philos. Trans. R. Soc. Lond. B 297: 125-181.BOXSHALL G.A. & S.H. HALSEY (2004) An Introduction to Copepod Diversity. Ray Society, London: i-xv, 1-966.HUMES A.G. (1999) J. Nat. Hist. 33: 961-978.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 349
Arthropoda, Crustacea, Copepoda, Misophrioida, Misophriidae
1A: Female habitus, dorsal; B: Habitus, lateral; C: Antenna; D: Leg 5; E: Male habitus, dorsal; F: Urosome, dorsal; G: Leg 5; from HUMES (1999).
Size: Body length of female 1.80-1.94 mm; male 1.28-1.45 mm.
Morphology: Prosome slender, five-segmented; tergite ofsomite bearing leg 1 covered by shield of cephalosome. Uro-some ornamented with lamellae and setules, five-segmented infemale, six-segmented in male; first somite with leg 5, genitaldouble-somite of female subdivided dorsally. Rostrum beak-likein lateral view. Caudal rami longer than wide. Antennuleeighteen-segmented in female, thirteen-segmented in male.Antenna with three-segmented endopod and six-segmentedexopod. Legs 1-4 biramous, with three-segmented rami. Secondendopodal segment of leg 2 with two inner setae. Leg 5 withone-segmented endopod and two- or three-segmented exopodin female and male, respectively.
Remarks: Misophriid copepods inhabit shallow waters, deep-sea hyperbenthos, deep-sea plankton, anchialine caves, and la-va tubes. Bathypelagic misophriids of the genus BenthomisophriaSARS, 1909 “are opportunistic gorgers, feeding on variety ofother animals, including copepods and cnidarians” (BOXSHALL
1982). Cave misophriids of the genus Speleophriopsis JAUME &BOXSHALL, 1996 (Speleophriidae) have been found in baitedtraps and can be scavengers (BOXSHALL & HALSEY 2004).
Distribution: Juan de Fuca Ridge: Coaxial Segment, Flow site(Vent HDV).
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Oncaea praeclara HUMES, 1988
Arthropoda, Crustacea, Copepoda, Cyclopoida, Oncaeidae
1A: Female habitus, lateral; B: Urosome, dorsal; C: Antenna; D: Maxilliped; E: Male habitus, dorsal; F: Maxilliped; from HUMES (1987).
Size: Body length of female 1.01-1.30 mm; male 0.91-0.98 mm.
Morphology: Body elongate. Prosome five-segmented. Uro-some five-segmented in female and six-segmented in male, firstsomite with leg 5. Genital double-somite of female and genitalsomite of male, anteriorly dilated and elongate. Caudal ramuslong, length/width ratio 6.7:1 in female and 3.2:1 in male. An-tennule six-segmented in both sexes. Antenna three-segment-ed, without exopod; distal segment with two groups of setae.Mandible with three setae and two broad spines; palp absent.Maxillule lobe with seven setae. Maxilla two-segmented, sec-ond segment with one spine and one seta. Maxilliped sexuallydimorphic, four-segmented in female and three-segmented inmale. Legs 1-4 biramous, with three-segmented rami. Femaleleg 5 as a small segment bearing two setae and adjacent dorsalseta; male leg 5 represented by three setae.
Biology: Common in the “samples taken in the vicinity of thevents by means of box corers and slurp guns” (HUMES 1988).Oncaea sp. has been found in plankton over the Mid-AtlanticRidge among dirivultids and subadult calanoids (IVANENKO
1998). Most of more than 70 species of Oncaea occur in theepipelagic zone, several species have been found in the deep ba-thypelagic zone. GO et al. (1998) recently reported that Oncaeafeeds on the integuments of various planktonic animals and in-flicts especially heavy injuries to chaetognaths; the copepods ofthis group “must gnaw or cut out pieces of host integument withtheir mandibles. To do this, they adhere tightly to the host bymeans of their antennae and maxillipeds” (HEPTNER & IVA-NENKO 2002).
Distribution: Galapagos Rift; East Pacific Rise: 9°N, 13°N,21°N; Guaymas Basin.
References:
GO Y.-B., OH B.-C. & M.J. TERAZAKI (1998) J. Mar. Sys. 15: 474-482.HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta 11(2): 117-134.HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.HUMES A.G. (1988) J. Plankton Res. 10: 475-485.HUMES A.G. & M. SEGONZAC (1998) Zool. Zh. 77(11): 1249-1256.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 350
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Grievella shanki FERRARI & MARKHASEVA, 2000
Reference:
FERRARI F.D. & E.L. MARKHASEVA (2000) Proc. Biol. Soc. Wash. 113 (4): 1079-1088.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 351
Arthropoda, Crustacea, Copepoda, Calanoida, Scolecitrichidae
1: Female; A: Habitus, lateral; B: Rostrum; C: Genital double-somite, lateral; D: Distal segment of maxilla; E: Leg 5;from FERRARI & MARKHASEVA (2000).
Size: Body length of female 2.1-2.2 mm. Male unknown.
Morphology: Prosome four-segmented; last segment of prosomewith leg 5. Urosome three-segmented; first somite is genitaldouble-somite with small integumental bumps. Rostrum bear-ing two filaments, each on robust base. Antennule twenty four-segmented; segment 22 with ear-like extension. Distal segmentof maxilla with nine sensory setae. Exopods of legs 1-4 and en-dopods of legs 3-4 three-segmented; endopods of leg 1 and leg2 one- and two-segmented, respectively. Distal exopodal seg-ment of leg 2 with two outer spines. Leg 5 two-segmented, dis-tal segment with three setae.
Biology: One female has been collected in “less than a meterabove the surface on the side of a small diffusing vent chim-ney”. The copepods of the family Scolecitrichidae “usually arecollected from pelagic and benthopelagic habitats of marinewaters below 200 m” (FERRARI & MARKHASEVA 2000).
Distribution: East Pacific Rise: 21°S, Droopy Vent.
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Isaacsicalanus paucisetus FLEMINGER, 1983
Size: Body length of female 2.44-2.56 mm. Male unknown.
Morphology: Prosome five-segmented; tergites of two proximalsomites fused laterally. Urosome four-segmented; first somite isgenital double-somite. Rostrum present, not bifurcate. Caudalrami slightly asymmetrical: left inner seta projecting ventrally,right inner seta directed dorsally. Antennule twenty three-seg-mented. Exopod of antenna longer than endopod. Exopods oflegs 1-4 and endopods of legs 3-4 three-segmented; endopods oflegs 1-2, 1- and two-segmented, respectively. Leg 1: first seg-ment of exopod without inner seta. Leg 5 absent.
Biology: The copepods have been sampled with the aid of aslurp gun within 1 m of the sea floor. They “were swimming ina tightly clustered swarm of several thousand individuals justabout a small depression… formed in collapsed pillow lava”(FLEMINGER 1983). Water temperatures in the depression rangefrom 5-15°C. The genus belongs to Spinocalanidae, “a com-mon group of small- to medium-sized calanoid copepods foundat meso- and bathypelagic depths of all oceans” (SCHULZ 1996).
Distribution: East Pacific Rise: 21°N.
References:
FLEMINGER A. (1983) Proc. Biol. Soc. Wash. 96 (4): 605-622.SCHULZ K. (1989) Mitt. Hamb. Zool. Mus. Inst. 86: 185-208.SCHULZ K. (1996) Polar Biol.16: 595-600.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 352
Arthropoda, Crustacea, Copepoda, Calanoida, Spinocalanidae
1: Female; A: Habitus, dorsal; B: Rostrum; C: Caudal rami; D: Antenna; from FLEMINGER (1983) (A, B, D) and SCHULZ (1989) (C).
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Smacigastes micheli IVANENKO & DEFAYE, 2004
Reference:
IVANENKO V.N. & D. DEFAYE (2004) Cah. Biol. Mar. 45: 255-268.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 353
Arthropoda, Crustacea, Copepoda, Harpacticoida, Tegastidae
1A: Female habitus, lateral; B: Male habitus, lateral; from IVANENKO & DEFAYE (2004).
Size: Body length of female 0.63 mm; male 0.57 mm.
Morphology: Body strongly compressed laterally, shield ofcephalothorax produced ventrolaterally, and male genital com-plex produced ventrally. Legs 1-4 biramous, with three-seg-mented rami. Elongate caudal rami and ten-segmented anten-nule of male are the distinctive features of Smacigastes.
Biology: Numerous adult and subadult copepodid stages of S.micheli have been collected during in situ colonization experi-ments at the base of the active chimney Eiffel Tower covered bya layer of Bathymodiolus azoricus (Bivalvia, Mytilidae).
Distribution: Mid-Atlantic Ridge: Lucky Strike.
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Cholidya polypi FARRAN, 1914
Size: Body length of female 0.74-0.83 mm; male 0.57-0.62 mm.
Morphology: Body of female with flattened shield-likecephalothorax and swollen metasome, urosome with slight in-dication of segmentation. Body of male with distinct somitesand six-segmented urosome. Rostrum produced anteriorly. An-tennule six-segmented in female and seven-segmented in male.Antenna: exopod one-segmented, endopod two-segmented.Mandibular palp three-segmented. Maxillule bilobed, withthree and five setae. Maxilla three-segmented, last two seg-ments forming claw. Maxilliped four-segmented, distal segmentclaw-shaped. Leg 1 biramous with three segmented rami; distalsegment of endopod with three claws; distal segment of exopodarmed with one spine and four setae ornamented with setules.Leg 2 of female biramous, with two-segmented rami. Legs 3-4 offemale represented by small segment with three and one setae,respectively. Legs 2-4 of male with three-segmented exopodsand two-segmented endopods. Leg 5 of female one-segmented,
with four setae and blunt terminal teeth. Leg 5 of male repre-sented with lateral expansion bearing four setae.
Biology: Widespread ectoparasite living on arms and web ofdeep-sea octopuses in the Atlantic and Pacific Oceans. One ofthese octopuses, Graneledone sp., sampled at the sediment-laden vent field was described by JUNIPER et al. (1992). Theparasitic tisbids are found on arms, head, and gills of the deep-sea octopuses associated to soft substrates. The redescription ofCholidya polypi and the list of species parasitizing octopuses isgiven in HUMES & VOIGHT (1997). Number of free-living tis-bids (including Tisbe sp. nov.) have been found in the Mid-At-lantic Ridge (Lucky Strike) during in situ colonization experi-ments (Ivanenko et al., in preparation).
Distribution: Juan de Fuca Ridge: Middle Valley; none vent lo-calities, Pacific and Atlantic Oceans (HUMES & VOIGHT 1997).
References:
HUMES A.G. & J.R. VOIGHT (1997) Ophelia 46(1): 65-81.JUNIPER S.K., TUNNICLIFFE V. & E.C. SOUTHWARD (1992) Can. J. Zool. 70: 1792-1809.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 354
Arthropoda, Crustacea, Copepoda, Harpacticoida, Tisbidae
1A: Female habitus, dorsal; B: Habitus ventral; C: Leg 1; D: Male habitus, dorsal. en – endopod; l5 – leg 5; from HUMES & VOIGHT (1997).
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Genesis vulcanoctopusi LÓPEZ-GONZÁLEZ, BRESCIANI & HUYS, 2000
Arthropoda, Crustacea, Copepoda, Harpacticoida, Tisbidae
1: Female; A: Habitus, dorsal;B: Habitus ventral; fromLÓPEZ-GONZÁLEZ et al. (2000).
Size: Body length of female 2.5-2.8 mm. Male unknown.
Morphology: Body indistinctly segmented; prosome laterallyexpanded with protruding margins. Rostrum produced beyondanterior margin of cephalothorax. Antennule four-segmented.Antenna with one-segmented exopod and two-segmented en-dopod. Mandibular palp two-segmented. Maxillule bilobed,with eight setae. Maxilla two-segmented, second segment claw-like. Maxilliped three-segmented, distal segment claw-shaped.Leg 1-4 biramous, exopods of legs 1-4 and endopod of leg 2three-segmented; endopods of legs 1, 3, and 4 two-segmented.Leg 5 represented by lobe fused to body and bearing six setae.
Biology: Females and subadults have been found embedded andencapsulated in the integument of the head and of the mantleof the deep-sea octopus Vulcanoctopus hydrothermalis GONZÁLEZ
& GUERRA, 1998. It was suggested that the life cycle of Genesisvulcanoctopusi would consist of both endoparasitic and ectopar-asitic phases.
Distribution: East Pacific Rise: 13°N, near Genesis.
Reference:
LÓPEZ-GONZÁLEZ P.J., BRESCIANI J., HUYS R., GONZÁLEZ A.F., GUERRA A. & S. PASCUAL (2000) Cah. Biol. Mar. 41: 241-253.
V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 355
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Arthropoda, Crustacea, Cirripedia
The cirripedes or barnacles are crustaceans found perma-nently attached as adults. They occur in brackish as well as ma-rine conditions, ranging from rigorous habitats such as estuariesand the highest reaches of the tides to the relatively benigndepths of the oceans. There are three superorders, the Acrotho-racica, Rhizocephala and Thoracica, and it is in the last thatthe greatest diversity is found. While the barnacle suborders as-sociated with hydrothermal vents and seeps are, like their deep-sea counterparts, predominantly pedunculates belonging to theScalpellomorpha and the asymmetrical sessile Verrucomorpha,their representatives, such as Ashinkailepas and Neoverruca, areremarkably generalized. Surprisingly, vents also include repre-sentatives of the two symmetrical sessile suborders, theBrachylepadomorpha and Balanomorpha. The former werethought to have gone extinct in the Miocene, until Neo-brachylepas was discovered at Lau. The latter, represented byEochionelasmus, first appear in the Paleocene and include the
dominant shallow-water barnacles today. All four of these gen-era are judged to be the most primitive surviving members oftheir suborders; e.g., relics of bygone ages.
In addition to being unusual in composition and primitivecharacter, as a group the vent and seep barnacles differ from allother thoracicans in having their mandibles uniquely modifiedfor feeding on very fine particles such as bacteria and fine de-bris. Some of the species have filamentous chemoautotrophicbacteria growing profusely on modified cirral setae, which theynurture by holding in prevailing vent currents and upon whichthey presumably feed. Furthermore, while vent barnacle larvaeare non-feeding, they are relatively large and yolky and studiesto date indicate they can remain in the plankton for threemonths or so before having to settle. This is five to six timeslonger than any known shallow-water barnacle which, as onewould expect, indicates they are very good dispersers.
W. NEWMAN, T. YAMAGUCHI, A.J. SOUTHWARD & M. SEGONZAC Denisia 18 (2006): 356–357
1: Barnacle Vulcanolepas “Lau A” sp. from Lau Back-Arc Basin; cruise TUIM07; by courtesy of C.R. Fisher.
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The relative abundance of vent barnacles appears to differgreatly from place to place as well as between species. WhileNeolepas zevinae is common along the Central East Pacific Rise,dense stands are rarely encountered. However, an undescribedneolepadine from the Central Indian Ocean Ridge, Vul-canolepas osheai from the Kermadecs, Leucolepas longa from offNew Guinea, Neoverruca brachylepadoformis from the MarianaBack-Arc Basin, and Eochionelasmus ohtai from Lau, may occurin extremely dense populations around vents or seeps. But cu-riously, the remains of vent barnacles have yet to be reportedfrom vent or seep deposits of any age. On the other hand, nospecimens of Neobrachylepas relica or Imbricaverruca yamaguchiihave been taken since first discovered, despite several subse-quent expeditions to Lau, and to date Neoverruca spp. areknown only by a chance photograph from Lau and in a samplefrom Manus, and both were among crowded Eochionelasmus. Itthus seems likely many new and exciting forms remain to bediscovered.
Vent-seep barnacles are now known to range pretty muchthroughout the Indo-Pacific vent-seep systems, except in theNE Pacific and the Galapagos where they are conspicuously ab-sence. They are also absent from the Atlantic, but so are nu-
merous other ancient groups of marine invertebrates. The cen-ter of distribution, where representatives of most genera arepresently found, is in the SW Pacific, specifically the Lau Basin,Tonga. It is noteworthy that the centers of diversity at high tax-onomic levels in numerous other relic groups of marine inverte-brates are also found in the SW Pacific. Taxonomic diversity at-tenuates to north and south, as well as to the east, and appar-ently also to the relatively unexplored vents of the far west. Bar-riers, such as disjunctions in active venting, the sill between theMarianas vents and the vents of Japan, or the Eastern Pacificand the Galapagos, have been proposed to explain these atten-uations. However, considering the ephemerality and patchinessof vent and seep fields, extinction rates associated with themmust be relatively high. It follows that the center of distributionis where availability of the vent-seep habitat has been the leastinterrupted in space and time, for otherwise such a high taxo-nomic endemicity could not have accumulated. Like the hightaxonomic endemicity of the SW Pacific in general, that of thevent barnacles appears to be the result of reliction due to habi-tat restriction rather than to barriers to dispersal per se.
357
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Eochionelasmus ohtai YAMAGUCHI, 1990
Size: This species, measuring up to 25 mm in height, is sub-stantially larger than its bathyal and more highly derived coun-terpart, Chionelasmus darwini, which is about 15 mm in height.
Morphology: Chionelasmatines are balanomorphs having awall comprising two pairs of dedicated latera, in addition to therostrum, carina and basal imbricating plates seen inbrachylepadomorphs. Eochionelasmus is distinguished fromChionelasmus by the basal whorls of imbricating plates of sever-al to numerous distinct whorls rather than being condensed in-to what appears to be a single whorl, and by its trophi beingmuch like those of other vent species. Two subspecies of Eo-chionelasmus were proposed by YAMAGUCH & NEWMAN (1997),E. ohtai ohtai and E. ohtai manusensis from Lau, Fiji and ManusBack-Arc Basins, respectively. They are distinguished fromeach other by differences in the frequency of appearance of therl1 and cl1 plates of the imbricating whorls (see E. paquensis).
Remarks: Eochionelasmus represents the most primitive livingbalanomorph. While the order in which the whorls of imbri-cating plates are added during ontogeny is the reverse of that in
brachylepadomorphs, the most parsimonious explanation wasthat balanomorphs evolved from them rather than independ-ently from scalpellomorphs. However, forthcoming genetic ev-idence indicates the balanomorphs evolved from very differentscalpellomorph stocks than the neoverrucids whereby the no-table similarities between neoverrucids and verrucids are con-vergent.
Biology: Eochionelasmus apparently occupies the same micro-habitat as Neoverruca, but when co-occurring it tends to bephysically closer to the mussels then are sympatric ne-olepadines. Equipped for feeding on the same material as allvent barnacles studied to date. Eggs are relatively large, butrarely recovered brooding whereby the nature of the larvalstages is unknown.
Distribution: Lau, North Fiji and Manus Back-Arc Basins.Morphological differences between the Lau-Fiji and Manuspopulations are considered sub-specific (see above) while dif-ferences between these three and a population from near East-er Island were interpreted as being specific (E. paquensis).
References:
BUCKERIDGE J.S. & W.A. NEWMAN (1992) J. Paleontol. 66: 341-345.GALKIN S.V. (1992) Zl. zh. Rss. Akad. Nauk. 71(11): 139-134.KLEPAL W, NEWMAN W. A. & W. TUFAR (2006) in SCHRAM F.R. & J.C. VON VAUPEL KLEIN (Eds.) Crustaceans and Biodiversity. Brill, Leiden: 195-199.PERÉZ-LOSADA M., JARA C.G., BOND-BUCKUP G., PORTER M.L. & K.A. CRANDALL (2002) J. Crust. Biol. 22: 661-669.TUFAR W. & H. JULLMANN (1991) Spiegel der Forschung 8(1): 39-44.YAMAGUCHI T. & W.A. NEWMAN (1990) Pacific Sc. 44(2): 135-155.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 358
1: E. o. othai, collected at Lau Back-Arc Basin; bycourtesy of T. Yamaguchi.
2: Group of E. o. manusensis from N-Fiji Back-Arc Basin; cruise StarmerII; P. Briand © Ifremer.
Arthropoda, Crustacea, Cirripedia, Balanomorpha, Chionelasmatoidae
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Eochionelasmus paquensis YAMAGUCHI & NEWMAN 1997
Size: 5 mm in diameter.
Morphology: The shell is low conic, its orifice large and rhom-boidal. Differs externally from E. ohtai in the alae of primaryplates being very indistinct and by the rostrum rising obliquelyrather than nearly vertically to the orifice of the wall. Oftencoated with ferromanganese. The scutum is very distinct inlacking an articular ridge and adductor muscle pit, a wider ar-ticular furrow, and a tergal magin that is indented rather thanstraight. The tergum has a produced articular ridge. The num-ber of basal imbricating plates ist greatly reduced compared toE. othai.
Biology: Living with mussels (Bathymodiolus thermophilus).
Distribution: East Pacific Rise: 17°S, site Rehu. Undescribedspecies of this genus are also known from Izu Ogasawara Arc,the Pacific-Antarctic Ridge: 38°S and the Central IndianOcean.
Reference:
YAMAHUCHI T. & W.A. NEWMAN (1997) J. Crust. Biol. 17(3): 488-496.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 359
1A: Right side of the holotype; B: Above side of the same specimen; scale bar 1 mm;from YAMAGUCHI & NEWMAN (1997). Capital letters indicate the principal wall plates: R – rostrum; RL – rostrolateral; CL – carinolateral; C – carina; small letters indicate theimbricating plates
2: Specimen, view from above; scalebar 1 mm; from East Pacific Rise:17°S, cruise Naudur © Ifremer; fromYAMAGUCHI & NEWMAN (1997).
Arthropoda, Crustacea, Cirripedia, Balanomorpha, Chionelasmatoidae
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Ashinkailepas seepiophilia YAMAGUCHI, NEWMAN & HASHIMOTO, 2004
Size: Up to 36 mm in height.
Morphology: Capitular plates ornamented by conspicuous lon-gitudinal ribs or ridges, peduncular to capitular ratio of 1:1 orless and, if unaltered by bending over to one side, with whorlsof six large scales which are considerably wider than high.
Remarks: Ashinkailepas is considered as the most primitive ofthe Neolepadinae. The subfamily Neolepadinae is consideredas the most primitive of the living Scalpellomorpha as well asclosest relative of the ancestor of the Brachylepadomorpha ofwhich the only living representative is Neobrachylepas relicaknown from Lau.
Biology: This species was collected from a cold deep. Associ-ates included a seep clam (Calyptogena sp.), and shrimps(Alvinocaris sp. and Lebbeus sp.). An ordinary deep-sea crab(Paralomis multispina) inhabited crevices between the boulders.Equipped with long cirri for setose feeding in gentle currentsand mouth parts uniquely modified for handling extremely fine
particles, but no significant differences noted compared toothose of ordinary vent barnacles (with the exception of thosewith cirri having exceptionally long setae generally festoonedwith filamentous bacteria). Peduncles not elongate, as in someneolepadines, but rather relatively short and bilaterally asym-metrical peduncular plates when individuals are bent over tooone side or the other. Function of bending over unknown, butit may be an advantage too extend the cirri closer too the sub-stratum, or perhaps too present a lower profile too maraudingpredators. Hermaphroditic; eggs large (~300 x 500 µm) and rel-atively few in number, as in N. zevinae NEWMAN, 1979, freenauplii likely present and lecithotrophic (cf. JONES 1993; TUN-NICLIFFE & SOUTHWARD 2004; WATANABE et al. 2004).
Distribution: Off Hatsushima Island, Sagami Bay (hydrother-mally driven cold seep). Undescribed species of this genus areknown from Japan: 32°N and 27°N, Lihir and the KermadecRidge.
1: Right side; from YAMAGUCHI et al. (2004).
2: Group of Ashinkailepas sp. from Japan;by courtesy of T. Yamaguchi.
Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae
References:
TUNNICLIFFE V. & A.J. SOUTHWARD (2004) J. Mar. Biol. Assoc. U.K. 84: 121-132.WATANABE H., KADO R., TSUCHIDA S., MIYAKE H., KYO M. & S. KOJIMA (2004) J. Mar. Biol. Assoc. U.K. 84: 743-745.YAMAGUCHI T., NEWMAN W.A. & J. HASHIMOTO (2004) Mar. Biol. Assoc. U.K. 84: 797-812.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 360
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Leucolepas longa SOUTHWARD & JONES, 2003
Size: Peduncular length commonly five but up to 12 timesheight of the capitulum, total length reaching 40 cm or more.This is, proportionately as well as literally, the longest peduncleof any known staked barnacle.
Morphology: Capitulum relatively narrow, tergal apex moreacute than in other neolepadines; e.g., <65° rather than some-what <75° as in Neolepas and >80° as in Vulcanolepas; capitularplates of mature individuals separated by a band of arthrodialmembrane; basal angle of tergum elevated well above ratherthan close to capitulo-peduncular junction; ratio of peduncularlength to capitular height generally >5:1 to as much as 12:1.Leucolepas longa can be distinguished from other neolepadinesin the basal angle of the tergum being well separated from thecapitulo-peduncular junction. This is true even in early juve-nile stages in which the relatively contiguous plates and themore convex occludent margin of the capitulum, as well asshort, blunt peduncular scales, suggest affinities with Vul-canolepas. In adult L. longa the capitular plates are well separat-ed rather than contiguous with each other.
Biology: It occurs in densities of over 1000 m-2, mostly associat-ed with vesicomyid clams, but occasionally on rocks and vesti-mentiferans over sulfide-rich sediments. The function of the
long, often gently curved peduncle appears to be to keep the ca-pitulum above those of its peers rather than adjusting to changesin prevailing currents, as seems to be the case in Vulcanolepas.Growth appears to be rapid with reproductive activity and re-cruitment continuous. Lecithotrophic nauplii, released upon re-trieval were cultivated in vitro for 45 days, metamorphosed intostage IV swimming nauplii containing lipid reserves deemed suf-ficient to insure wide dispersal. In a similar in vitro rearing of lar-vae of Neoverruca sp., it tooks 96 days to reach the cyprid stage.
Remarks: Leucolepas longa differs considerably in general fromNeolepas and Vulcanolepas. While what is apparently a newgenus from the Izu Ogasawara Arc has the same general ap-pearance, the basal angle of the tergum reaches the capitulo-junction, the plates are contiguous, the median latus has aheight to width ratio is two rather than 1.5:1 or less, and the p-c ratio is in the order of eigth rather than 12:1.
Distribution: Tabar-Feni Volcanic Fore-Arc: Edison Seamount.While presently monotypic, undescribed populations range asfar East as the Izu Ogasawara Arc (see remarks). An unde-scribed species is known from North Fiji Back-Arc Basin, an-other undescribed species from Sunda Trench, and perhaps a re-lated genus from Izu Ogasawara Arc.
1: Capitulum after removal of theprosoma and cirri which empha-sizes the uncalfified interspacesbetween the shell plates; fromSOUTHWARD & JONES (2003).
2: Group of specimens; from SOUTHWARD &JONES (2003).
3: In situ view of clumps of barnacles, somewith cirral net open, at the edge of a bed oflive vesicomyids; also visible, associated faunaof Phymorhynchus gastropods and squat lob-ster; scale bar 15 cm; from TUNNICLIFFE & SOUTH-WARD (2004); by courtesy of P. Hertzig.
Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae
References:
SOUTHWARD A.J. & D.S. JONES (2003) Senckenberg. maritima 32 (1/2): 77-93.TUNNICLIFFE V. & A.J. SOUTHWARD (2004) J. Mar. Biol. Ass. U.K. 84: 121-132.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 361
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Neolepas rapanuii JONES, 1993
Size: Up to approximately 8.0 cm in total length.
Morphology: Distinguished from N. zevinae by the apex of ros-trum generally being equal to rather than higher than the me-dian latus, a peduncle often as much as fives times the height ofthe capitulum and clothed with less robust scales, and amandible without a strong spine near the superior margin of thesecond tooth and denticles of the second and third teeth notrolled over toward the inside.
Biology: While little known, its biology is likely much the sameas that of N. zevinae, except for the development of a longerand often curved peduncle, presumably to keep the cirral net inprevailing currents, and the comb of denticles along the supe-rior margin of the second and third mandibular teeth and theupper part of the inferior angle, that stand nearly erect ratherthan being rolled over toward the inner surface of themandible. This is curious as they are rolled over in N. zevinae
and all subsequently described vents and seep species, but thefunctional basis for the difference is unknown.
Remarks: Peduncles elongated as in most other neolepadinegenera, evidently to keep the capitulum and cirral net in cur-rents rather than to host chemoautotrophic endosymbionts.This is a different strategy than practiced by Ashinkailepas, N.zevinae, and of course the sessile vent barnacles, Neoverruca,Imbricaverruca, Neobrachylepas and Eochionelasmus.
Distribution: East Pacific Rise: 23°S. An undescribed species ofthis genus is known from Pacific-Antarctic Ridge: 32°S, anoth-er species from Izu Ogasawara Arc and another morphological-ly similar from is known from Toto Caldera Marianos Arc.Three other populations from the Indian Ocean Rigde (25°S,17°S, 41°S) while genetically close to Neolepas, appear inter-mediate between it and Leucolepas.
References:
JONES D.S. (1993) Bull. Mar. Sci. 52(3): 937-948 SOUTHWARD A.J. & D.S. JONES (2003) Senckenberg. maritima 32(1/2): 77-93
D. JONES, W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 362
1: In vivo specimen; scale bar 1 cm; from JONES (1993).
Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae
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Neolepas zevinae NEWMAN, 1979
1: Specimen in vivo collected fromEast Pacific Rise: 21°N; by A. Southward.
2: Specimen in vivo collected fromEast Pacific Rise: 21°N; by A. Southward.
Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae
Size: This species is known to be up to 5.8 cm in total length,whereas alone the peduncle of N. rapanuii can exceed thatlength, and some neolepadines in the western Pacific have pe-duncles more than a decimeter in length.
Morphology: Capitulum with relatively smooth, unornament-ed capitular plates and an adult peduncle with whorls of 16 ormore scales which are about as wide as high. Distinguished fromN. rapanuii by the apex of rostrum generally being higher thanthat of the median latus, a peduncle clothed with proportion-ately more robust scales and of no more than three rather thanas much as five times the height of the capitulum, and amandible having a strong spine near the superior margin of thesecond tooth and the denticles of the second and third teethrolled over toward the inside.
Remarks: Neolepadines are considered the most primitive ofthe living scalpellomorphs known, and are evidently earlyMesozoic in age. They apparently arose in the Triassic (BUCK-ERIDGE & GRANT-MACKIE 1985) and are near the base of theclade that gave rise to the first sessile barnacles, thebrachylepadomorphs.
Biology: Neolepadines are hermaphroditic, like all vent barna-cles studied to date. They are gregarious but, like the cyprid lar-vae of Ashinkailepas, those of Neolepas zevinae apparently preferto settle on the substratum rather than on established individ-uals, such as on the long peduncles of Neolepadines from else-where. Individuals tend to align along ridges and contours fac-ing the prevailing current. Neolepadines and Eochionelasmusoccur sympatric with mussels, the latter tend to be closer to thevent. Equipped with long cirri for setose feeding in gentle cur-rents and equipped with mouth parts uniquely modified forhandling extremely fine particles, as in all vent barnacles stud-ied to date. Peduncles become elongate in some forms, evi-dently to keep the capitulum and cirral net in currents ratherthan to host chemoautotrophic endosymbionts. Eggs large insize, egg nauplius present in Neolepas rapanuii, but there arelikely free lecithotrophic stages, as in other vent barnacles.
Distribution: East Pacific Rise: 9-21°N.
References:
BUCKERIDGE J.S. & J.A. GRANT-MACKIE (1985) Géologie de la France 1: 77-80. JONES D.S. (1993) Bull. Mar. Sci. 52: 937-948.NEWMAN W.A. (1979) Trans. San Diego Soc. Nat. Hist. 19(11): 153-167.NEWMAN W.A. (1985) Bull. Biol. Sc. Wash. 6: 231-242.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 363
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Vulcanolepas osheai (BUCKERIDGE, 2000)
Size: Capitulum up to approximately 20 mm, overall length toat least 120 mm, so pedunclar to capitular 5:1 or 6.8:1.
Morphology: Capitulum broad, tergal apex blunter than in oth-er Neolepadines (>80 versus <75°); capitular plates approxi-mate, not separated by a band of arthrodial membrane; basalangle of tergum close to rather than elevated well above capit-ulo-peduncular junction; maximal observed ratio of pedunclarlength to capitular height 7:1 but usually less.
Biology: Found living on volcanic debris in the vicinity of anew dome forming within the caldera. While no other mac-robenthic animals were collected, the presence of shrimp wasobserved. Cirri apparently equipped with exceptionally long,relatively soft setae (BUCKERIDGE 2000, Fig. 4C, E), which sug-gests bacterial ectosymbiosis (cf. SOUTHWARD & NEWMAN
1998), an hypothesis compatible with the molecular phyloge-netic and isotopic evidence for ectosymbiosis in V. osheai(Suzuki et al., in prep.).
Remarks: This species is very similar in external appearance to“Lau A“ of SOUTHWARD & NEWMAN (1998). In most individu-als of Lau A, these setae were festooned with filamentous bac-teria. Since the cirral rami as well as the exterior surfaces of thebarnacle in general were not covered with such bacteria,SOUTHWARD & NEWMAN (1998) concluded those on the setaewere being farmed for consumption.
Distribution: Kermadec Arc: Brothers Seamount.
References:
BUCKERIDGE J.S. (2000) New Zealand J. Mar. Freshwater Res. 34: 409-418.SOUTHWARD A.J. (2005) Senckenberg. maritima 35(2): 147-156.SOUTHWARD A.J. & W.A. NEWMAN (1998) Cah. Biol. Mar. 39: 259-262.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 364
2: Group of 16 paratypes; from BUCKERIDGE (2000).
Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae
1: Habitus; scale bar 1 cm© T. Yamaguchi.
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Vulcanolepas parensis SOUTHWARD, 2005
Size: Capitulum up to 15 mm in height and peduncular lengthof 103 mm; peduncular to capitular ratio 6.86:1, while perhapsa smaller species than V. osheai, peduncular to capitular ratio es-sentially the same.
Morphology: Tergum shorter (basal angle well above capitulo-peduncular margin) and median latus is taller than in V. osheai,median latus forming an approximately equilateral trianglewhereas in V. osheai and Leucolepas longa it is curved along thetergal margin.
Remarks: Morphologically this species appears to be a Vul-canolepas, and if so, it is relatively isolated since the genus isotherwise only known from Lau, East to the Kermadecs. How-ever, there could be a vent fauna along the Pacific-Antarcticand Indian-Antarctic Ridges to be discovered that would helpexplain the situation.
Biology: Posterior cirri with ctenopod rather than lasiopod cir-ri as in Vulcanolepas sp.
Distribution: Pacific-Antarctic Ridge: 37°S to 38°S. A bacte-ria-farming form from Lau Back-Arc Basin is representative ofthis genus.
Reference:
SOUTHWARD A.J. (2005) Senckenberg. maritima 35(2): 147-156.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 365
1: Capitulum of a specimen with the capitularplates; from SOUTHWARD (2005).
2: Cluster of specimens on a small fragmentof the substratum, peduncles of mediumlength; from SOUTHWARD (2005).
Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae
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Neobrachylepas relica NEWMAN & YAMAGUCHI, 1995
References:
NEWMAN W.A. & T. YAMAGUCHI (1995) Bull. Mus. natl. Hist. nat., Paris. 4e sér 17A (3-4): 211-243.NEWMAN W.A. (1993) in TRUSEDALE J. (Ed.) The History of Carcinology. Crustacean Issues 8: 349-434.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 366
1: Viewed from above (A) and from the right (B). Me-dian latera (L), scuta (S) and terga (T) comprise the op-erculum; rostrum (R) and carina (C) are in contact attheir lateral margins; the other letters indicate the im-bricating plates; from NEWMAN & YAMAGUCHI (1995).
2: Specimen viewed from the left side, almost com-pletely buried beneath sulfide and oxide depositson a small block of basalt; cruise Biolau © Ifremer;from NEWMAN & YAMAGUCHI (1995).
Arthropoda, Crustacea, Cirripedia, Brachylepadomorpha, Neobrachylepadidae
Size: Only a few possibly juvenile or protandric specimens havebeen collected, suggesting they may have been waifs; that is,the microhabitat of the adult populations likely has yet to besampled. The largest specimen was mature as a male, but therewere no eggs and it was only 6 mm high. It is nonetheless pos-sible this is a small species compared to other vent barnacles.
Morphology: A living representative of the Brachylepadomor-pha (U. Jurassic-Miocene), symmetrical sessile barnacles withbasal whorls of imbricating plates surrounding a wall consistingof but two principal plates, the rostrum and carina. Neo-brachylepas differs from †Brachylepas in the rl-l-cl tiers of imbri-cating plates covering the gap between the principal wall platesstanding three rather than four plates high (compare to Imbri-caverruca in this regard). Distinguished from neolepadines inbeing sessile and operculate, from neoverrucids in being sym-metrical, and from balanomorphs in the operculum including apair of median latera and the oldest whorl of imbricating platesbeing basal rather than situated between the younger imbricat-ing whorls and the wall.
Biology: Unfortunately a reproductive population of thisspecies has yet to be discovered, but one would expect its mem-bers to occupy much the same situations occupied by Neoverru-ca brachylepadoformis and Eochionelasmus ohtai because, whileperhaps a smaller species, it has essentially the same feedingmechanism and is hermaphroditic. The first and often the sec-ond pair of cirri in vent barnacles are antenniform and likelyserve in part in orienting the cirral net to currents. However, inNeobrachylepas, the second pair is not only shorter than the firstbut it apparently functions as maxillipeds. Another unique fea-ture in Neobrachylepas is a median-dorsal appendage on the pro-soma that may serve to hold the egg mass in place while brood-ing, but ovigerous individuals have not been observed.
Remarks: Neobrachylepas relica is the only known survivingmember of the Brachylepadomorpha, fossil representatives ofwhich are known from the Jurassic to the Miocene.
Distribution: Lau Back-Arc Basin.
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Imbricaverruca yamaguchii NEWMAN, 2000
Size: Apparently a small species, largest of several specimenshaving a rostro-carinal diameter of 7 mm and standing about5.5 mm high.
Morphology: Differs from Neoverruca in having 1) a more ver-rucid-like operculum albeit including a well developed medianlatus, 2) a fixed scutum and tergum much wider than highrather than at least as high as wide, and 3) in retaining a com-plete set of well-developed imbricating plates on the movableside of the wall essentially for life whereas the imbricating sys-tem in Neoverruca is incomplete (at least plates r4 -l4-c4 fail todevelop) and the older (marginal) imbricating plates tend tofall off.
Biology: No data.
Distribution: Known only from the type locality; Lau Back-ArcBasin, site Hine Hina.
Reference:
NEWMAN W.A. (2000) Zoosystema 22(1): 71-84.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 367
1A: Oblique view of movable side; B: Viewed from scutal end; C – carina, FS,MS – scutum, FT, MT – tergum, R – rostrum, cl – carinolaterale, l – laterale, rlrostrolaterale; from NEWMAN (2000).
Arthropoda, Crustacea, Cirripedia, Verrucomorpha, Neoverrucidae
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Neoverruca brachylepadoformis NEWMAN, 1989
Size: Moderately large sessile barnacle measuring up to 25 mmin height.
Morphology: Neoverrucids are bilaterally asymmetrical sessilebarnacles built the same general plan as the brachylepado-morphs. They are distinguished from verrucids in having basalwhorls of imbricating plates surrounding the wall and an oper-culum including a median latus, which in N. brachylepadoformisbecomes vestigial with age and may even be lost, but in Imbri-caverruca it remains relatively large and significant part of theoperculum. Imbricaverruca is further distinguished from Neover-ruca in having the rl and cl tiers of imbricating plates as in†Brachylepas; e.g., standing four rather then three or less plateshigh, and in having the basal-most ones well attached ratherthan easily shed.
Remarks: Neoverrucids appear not only to be the most primi-tive living verrucomorphs but to represent the “missing link“between the Brachylepadomorpha and the remainder of theVerrucomorpha.
Biology: Neoverruca brachylepadoformis is hermaphroditic. It isgregarious, and the cyprid larvae settle on established individu-als as well as on the substratum. Settled juveniles first passthrough several pedunculate stages before undergoing meta-morphosis into the first sessile stage. Cyprids metamorphosis in-to juveniles with their cirral nets facing the prevailing current,and those that settled on the right side of an established indi-
vidual develop their operculum on the right side, while thosesettling on the left develop it on the left side. Thus right andleft-sidedness is ecotypically determined, as is likely the case inother verrucomorphans where right and left-sidedness occurs inthe same species. It appears that some species of the genus areoccasionally sympatric with Eochionelasmus, at Lau and ManusBasins for example.Equipped to feed on the same material as all vent barnaclesstudied to date. Eggs are relatively large and held in place byovigerous frenae. Lecithotrophic nauplii in vitro pass throughsix stages before metamorphosis after to the cyprid stage, andthis took 96 days in the deep-water form from the OkinawaTrough reared in vitro. Unusual among sessile barnacles in pass-ing through several pedunculate stages resembling neolepadidsduring ontogeny, as likely do the brachylepadomorphs. Whilethe earliest pedunculate stages are essentially symmetrical,asymmetry becomes apparent before an abrupt metamorphosisinto the first sessile juvenile, during which the peduncle be-comes buried beneath an expanded membranous basis, and thebasal whorl of capitular plates becomes the oldest or basal-mostwhorl of imbricating plates of the sessile form. Additionalwhorls are added between it and the wall plates until at leastthe median latera of the movable side stand four plates high.
Distribution: Mariana Back-Arc Basin. Undescribed speciesare found in Marianas, Japan, Manus and Lau.
References:
NEWMAN W.A. (1989) Bull. Mar. Sci. 45(2): 467-477.NEWMAN W.A. & R.R. HESSLER (1989) Trans. San Diego Soc. Nat. Hist. 21(16): 259-273.WATANABE H., KADO R., TSUCHIDA S., MIYAKE H., KYO M. & S.J. KOJIMA (2004) Mar. Biol. Assoc. U.K. 84(4): 743- 745.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 368
1: Large specimen(2.1 cm high), rightside, with two juve-niles lacking ferro-
manganese deposit;Mariana Back-Arc
Basin; from NEWMAN
& HESSLER (1989)
2: Closelypacked aggregation;Mariana Back-Arc Basin; by courtesy ofR.R. Hessler.
Arthropoda, Crustacea, Cirripedia, Verrucomorpha, Neoverrucidae
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Dahlella caldariensis HESSLER, 1984
Size: Up to 10 mm.
Morphology: Carapace ovoid, posterior edge emarginate, ex-posing at least part of pleonite one dorsally. Rostrum 0.4 timesof carapace length, 3 time longer than wide. First thoracomereseen from carapace even dorsally, postero-lateral angles ofpleonites 1-3 rounded, that of pleonite 4 pointed. Anal plateslike equilateral triangles. Eyestalk boomerang shaped (see up-per centre figure) and denticulate articulated to head without
visual elements. Scale of the first antenna with the anteriormargin densely clothed with setae ranging from stout and den-tate to long and slender (see upper right figure).
Biology: In washings of mussels and vestimentiferan worms.Observed swimming above clumps of animals at vents.
Distribution: Galapagos Spreading Center and East PacificRise: 21°N to 18°S.
Arthropoda, Crustacea, Malacostraca, Leptostraca, Nebaliidae
References:
HESSLER R.R. (1984) J. Crustac. Biol. 4(4): 655-664.LEDOYER M. (1995) Mar. Life 4(1): 25-29.
T. HANEY Denisia 18 (2006): 369
1: Habitus (critical point dried specimen) Ifremer.
2: Eyestalk (carapace removed) SEM Ifremer.
3: Distal penduncular article ofthe first antenna (SEM) Ifremer.
4: Lateral side of pleonites(SEM) Ifremer.
6: Caudal ramus (SEM) Ifremer.5: Pleopods (SEM) Ifremer.
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Atlantocuma bidentatum LEDOYER, 1988
Size: 3.9 mm.
Morphology: Subfamily: Mandibles not broad at base. Exopodsonly on maxilliped 3 and pereopod 1 in female. Males usuallywith five pairs of pleopods (sometimes two or three) but theyare absent in Atlantocuma that has been considered as an aber-rant genus (JONES , 1984). Without free telson. Uropod endo-pod one- or two-articulated.
Female: Carapace smooth, about twice as long as high andslightly shorter than 1/3 of total length; pseudorostrum not verylong, pointed; antennal notch shallow, anterolateral angle notprominent, marked by two teeth; eyelobe pointed at front, with-out lenses. Five thoracic segments visible. Pleon slightly longerthan carapace and thoracic segments combined. Antenna 1 pe-duncle three-articulated; main flagellum two-articulated withtwo aesthetascs terminally, accessory flagellum rudimentary. Ex-opods on maxilliped 3 and pereopod 1. Maxilliped 3 basis slen-der, 1.5 as long as the rest of appendage. Pereopod 1 basis slight-ly longer than rest of appendage with a small simple seta on dis-tal corner; carpus longer than propodus, which is 1.5 times as
long as dactylus. Pereopod 2 basis shorter than the remaining ar-ticles together, ischium very short, carpus longer than dactylus.Pereopod 3 basis longer than rest of appendage, carpus about 1/3of basis. Pereopod 4-5 basis shorter than the rest of appendage,carpus longer than half length of basis. Uropod peduncle longerthan pleonite 5 and twice length of endopod; exopod article 2with a simple seta on distal third of inner margin and a long sim-ple seta terminally; endopod one-articulated, with 10 setae oninner margin and a longer one terminally.
Remark: Although the specimen described above agrees withthe LEDOYER’S (1988) description in most of the features, it dif-fers by a greater number of setae on the endopod of uropod.
Biology: Collected in a sediment trap (12 m above the bottom)located at East Pacific Rise: 13°N, 2 m north of the vent siteParigo.
Distribution: East Pacific Rise: 13°N, site Parigo. Previouslyknown only from South-East of Glorieuses Islands, Mozam-bique Channel, at 3716 m depth (LEDOYER 1988).
References:
JONES N.S. (1984) Bull. Br. Mus. Nat. Hist. (Zool.) 46(3): 207-289.LEDOYER M. (1988) Mésogée 48: 131-172.
J. CORBERA Denisia 18 (2006): 370
1: Preadult female; A: Habitus; B: Anterolatereal angle of carapace; C: Maxilliped 3; D: Pereopod 1; E: Pereo-pod 2; F: Pereopod 3; G: Pereopod 4; H: Pereopod 5; I: Uropod; by J. Corbera.
Arthropoda, Crustacea, Cumacea, Bodotriidae
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Bathycuma brevirostre (NORMAN, 1879)
Size: 15.5 mm.
Morphology: Subfamily (Vaunthompsoniinae): Mandibles notbroad at base. Exopods on at least first three pairs of pereopods.Males usually with five pairs of pleopods (sometimes three).Without free telson. Uropod endopod one- or two-articulated.Species, adult male: Carapace with a mid-dorsal paired row ofteeth on anterior half; anterolateral angle acute and lateralmargin serrated. Antenna 1 peduncle three-articulated, article1 slightly shorter than article 2 and 3 combined lengths; mainflagellum two-articulated with 2 aesthetascs terminally, acces-sory flagellum rudimentary. Well developed exopods on maxil-liped 3 and pereopods 1-4. Maxilliped 3 basis longer than restof appendage, distal outer corner produced reaching merus.Pereopod 1 basis shorter than rest of appendage, with cuspidatesetae on ventral face and pappose setae on distal half of outerand inner margins; carpus and dactylus of the same length.Pereopod 2 basis as long as rest of appendage, ischium very
short. Pereopod 3 and 4 basis longer than rest of appendage,with pappose setae on margin. Pereopod 5 basis shorter thanthe three following articles combined lengths. Uropod pedun-cle longer than rami, with more than 20 setae on inner margin,being largest the distal one; exopod article 2 with simple setaeon outer margin, plumose setae on the inner margin and twolong simple setae terminally; endopod two-articulated, article 1twice long as article 2 with more than 20 setae on inner mar-gin; article 2 with eight setae on inner margin and 2 terminal-ly.
Biology: Collected in a sediment trap (2.5 m above the bot-tom) located between two active vents at 1630 m depth.
Distribution: Previously known from south of Ireland (CAL-MAN 1905), Bay of Biscay (JONES 1985) and Mediterranean Sea(REYSS 1973) up to 5000 m depth. Mid-Atlantic Ridge: LuckyStrike.
1: Adult male; A: Habitus; B: Antenna 1; C: Maxilliped 3; D: First pereopod; E: Second pereopod; F: Third pereo-pod; G: Fourth pereopod; H: Fifth pereopod; I: Uropod; by J. Corbera.
Arthropoda, Crustacea, Cumacea, Bodotriidae
References:
CALMAN W.T. (1905) Fish. Ireland, Sci. Invest.1904(1): 1-7.JONES N.S. (1985) in LAUBIER L. & C. MONNIOT (Eds.) Peuplements Profonds du Golfe de Gascogne: 429-433.NORMAN A.M. (1879) Ann. Mag. Nat. Hist. 3(5): 54-73.REYSS D. (1972) Deep-Sea Res. 20: 1119-1123.
J. CORBERA Denisia 18 (2006): 371
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Arthropoda, Crustacaea, Tanaidacea
In some deep-sea environments tanaids are the most di-verse and abundant fauna but because of their small size (mostare 2-5mm long) they are frequently overlooked. Their distri-bution in hydrothermal environments is virtually unknown. Upto now only two species (Leptognathia ventralis and Typhlotanaissp.) are known from Western Pacific Back-Arc Basins, sevenspecies from the Mid-Atlantic Ridge, Lucky Strike (LARSEN etal., in press) and one (Typhlotanais sp.) from Rainbow (M.Segonzac, pers. obs.).
At Lucky Strike, the highest diversity and abundance oftanaids occurred in peripheral mussel clumps. Besides the twospecies, Agathotanais ingolfi and Pseudotanais vulsella, knownfrom other deep-sea environments, Mesotanais styxis was foundboth inside and outside the vent field.
All species known from hydrothermal vents belong to pre-viously known genera, even Gordotanais was found before al-though not yet described indicating that the fauna at LuckyStrike is not dramatically different from the surrounding deep-sea habitat. A similar observation was made in the study oftanaids from the Juan de Fuca vent system (LARSEN, in press).
Although the present handbook aims to present diagnosticinformation and illustration it is important to stress that theidentification of tanaid species is often difficult and usually re-quires the dissection of the specimens.
References:
LARSEN K. (in press) Zootaxa.LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.
M.R. CUNHA Denisia 18 (2006): 372
1: Armaturatanais atlanticus; by K. Larsen, M. Blazewicz-Praszkowycz & M. R. Cuhna.
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Armaturatanais atlanticus LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press
Size: 2.0 mm.
Morphology: (Genus, female) Body slightly dorso-ventrallyflattened. Cuticle heavily calcified. Pleotelson acorn-shaped inlateral view. Antennula with four articles. Antenna with six ar-ticles and fusion line on article 4. Mandibles small; molar fair-ly broad with flat distal crushing area surrounded by small den-ticles. Maxilliped endites without denticles, processes or flat se-tae. Pereopods I-III merus and carpus without spiniform setae.Pereopods and pleopods attached on the inner side of the lat-eral shield. Pereopods with coxa; dactylus and unguis not fusedto a claw; pereopods I-III unguis longer than dactylus; pere-opods IV-VI dactylus longer than unguis. Pleopods present infemale, with simple or plumose setae. Uropods biramous, endo-pod with two articles; exopod with one article. (Genus, male)
Pleon marginally longer than in female. Antennular article 3shorter than in female and without fusion line. Functionalmouthparts retained. All pleonites bearing pleopods withplumose setae, pleopods larger than in female. (Species, fe-male) Antennula article 3 more than half as long as article 2.Right mandibular incisors smooth; left mandibular/lacinia mo-bilis with few dorsal denticles. Chelipedal dactylus naked. Pere-opods longer than pleon. Pereopods IV-VI dactylus naked; un-guis of same with simple apex.
Biology: Collected from samples of volcanic rocks (hyaloclas-tic) and hydrothermal slab in the proximity of active vents.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.
K. LARSEN & M.R. CUNHA Denisia 18 (2006): 373
1: Female; A: Habitus, lateral view; B: Antennula; C: Antenna; D: Cheliped; E: Maxilliped; scale bars1 mm (A), 0.1 mm (others); from LARSEN et al. (in press).
Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, fam. indet.
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Leptognathiella fragilis LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press
Size: 1.3 mm.
Morphology: (Family and Genus) Body cylindrical, relativelysmall (rarely over 2 mm in length). Pleon and pleotelson short(never longer than combined length of three last pereonites).Antennule with four or five articles; article 2 frequently withdorsal projection overlapping basal part of article 3. Antennawith five or six articles. Mouthparts well developed and func-tional in both sexes. Molar process thin with few terminalspines. Maxillule with seven or eight terminal spiniform setae.Maxilliped endite not fused, distal edge often with medial, fre-quently triangular, process. Chelipeds attached via sclerite.Pereopods I-III often stout, with or without coxa. Pereopods IV-VI without coxa and not stouter than pereopods I-III; dactylusand unguis not fused. Pleopods absent or present with simple
setae only. Uropods biramous; rami with one or two articles.(Species, female) Body fairly elongated (length/width ca. 9.5).Cephalothorax as long as combined length of first two pere-onites. Pereopods I-III carpus and propodus without small ven-tral spines. Uropods twice as long as pleotelson but shorter thancombined pleon; basal article shorter than pleotelson and biar-iculated exopod half as long as first endopod article.
Biology: Collected from samples of hydrothermal slabs in thevicinity of active venting.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.
K. LARSEN & M.R. CUNHA Denisia 18 (2006): 374
1: Female; A: Habitus lateral view; B: Antennula; C: Antenna; D: Maxilliped; E: Cheliped;scale bars 1 mm (A), 0.1 mm (others); from LARSEN et al. (in press).
Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Colleteidae
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Mesotanais styxis LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press
Size: 1.9 mm.
Morphology: (Family and Genus) Body cylindrical. Eye lobespresent but without visual pigment. Antennula with three arti-cles. Antenna with six articles; article 2 and 3 with spiniformdorsal setae. Mandibles well developed with broad molar. Labi-um with two pairs of lobes. Maxillule with nine spiniform ter-minal setae. Maxilliped basis and endites not fused; enditeswith two or three short, flat, unequal-sized setae and none orone simple seta; basis with one or two long simple setae nearpalp insertion. Mouthparts reduced in males. Chelipeds at-tached via sclerite. Pereopod I almost twice as long as followingpereopods; dactylus/unguis longer than propodus. On otherpereopods dactylus/unguis shorter than propodus. Pereopods I-III with coxa; dactylus/unguis not fused. Pereopods IV-VI with-
out coxa; basis thicker than pereopods I-III dactylus/unguis in-completely fused to an elongated claw. Pleopods present andwell developed, with plumose setae. Uropods biramous; exopodbiarticulated; endopod with three or more articles. (Species, fe-male) Antennula without long (almost as long as antennule)setae. Maxilliped basis with only one distal setae. Uropodal en-dopod with four articles.
Biology: Collected from samples of hydrothermal slabs in theproximity of active chimneys but also on volcanic rocks outsidethe vent field (Lucky Strike segment).
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.
K. LARSEN & M.R. CUNHA Denisia 18 (2006): 375
1: Female; A: Habitus, dorsal view; B: Habitus, lateral view; C: Antennula; D: Antenna; E: Cheliped;F: Pleopod; scale bars 1 mm (A, B), 0.1 mm (others); from LARSEN et al. (in press).
Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Leptocheliidae
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Obesutanais sigridi LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press
Size: 1.2 mm.
Morphology: (Family and Genus, female) Body habitus shortand stout (almost pseudotanais-like) with a wide head. Body ta-pering off in posterior direction. Antennula with long setae onarticle 1. Mandibular molar with few distal spines. Maxillipedendites with one distal tubercule. Mouthparts reduced in males.Cheliped merus and carpus with long seta (longer than fixedfinger). Pereopods II and III merus with one seta as long as orlonger than carpus; carpus with a long seta (almost as long aspropodus). Pereopod IV-VI without process with microspines(clinging apparatus). Pleopods setation, except for endopodproximal seta, restricted to the distal end; large gap betweenendopod proximal seta and other setae, long (more than half
the length of endopod). Uropod endopod is incompletely fusedto one article; endo- and exopod with thick specialized termi-nal setae; exopod unarticulated, almost as long as endopod.(Species, female) Pereopods long (longer than pleon). Pere-opods IV-VI dactylus and unguis partly fused but not into aclaw. Uropods longer than pleotelson; endopod with pseudoar-ticulation; exopod with one article, almost as long as endopod.
Biology: Collected from samples of volcanic rocks (hyaloclas-tic) and sulphide chimneys active and inactive. Often found in-side tubes built with small particles of sediment.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.
K. LARSEN & M.R. CUNHA Denisia 18 (2006): 376
1: Female; A: Habitus, dorsal view; B: Habitus, lateral view;C: Antennula; D: Antenna; E: Cheliped; scale bars 0.1 mm;from LARSEN et al. (in press).
2: Tubes on a piece of sulphide(Lucky Strike), cruise Seahma 1 © FCIJL; by P. Briand © Ifremer.
3: Specimen removedfrom tube; by M. Cunha.
Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Nototanaidae
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Typhlotanais incognitus LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press
Size: 1.5 mm.
Morphology: (Family and Genus, female) Body almost com-pletely cylindrical. Cephalothorax wider than pereon. Anten-nula with three articles. Antenna with five or six articles. Mo-lar process broad with several terminal denticles. Maxillulawith seven or eight terminal spiniform setae. Maxilliped basispartially fused, endites not fused; distal edge with lateralprocess. Chelipeds ventrally attached via sclerite. Pereopods I-III with coxae; merus and carpus with simple setae only. Pere-opods IV-VI with or without coxae; basis stouter or not stouterthan pereopods I-III; merus and carpus with at least one spini-form seta, usually with clinging apparatus; dactylus and unguis
fused. Pleopods present, with simple or plumose setae. Uropodsbiramous; endopod with two articles; exopod with one or twoarticles. (Species, female) Pereopods long (longer than pleon).Pereopods IV-VI not stouter than Pereopods I-III; dactylus andunguis fused but not into a claw. Uropods longer than pleotel-son; endopod with two articles; exopod with one article, almostas long as endopod.
Biology: Collected from samples of hydrothermal slab in theproximity of active vents.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.C.R CUNHA (in press) Zootaxa.
K. LARSEN & M.R. CUNHA Denisia 18 (2006): 377
1: Female; A: Habitus, lateral view; B: Habitus, dorsal view; C: Antennula; D: Antenna; E: Cheliped; scale bars 0.1 mm; fromLARSEN et al. (in press).
Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Nototanaidae
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Pseudotanais vulsella BIRD & HOLDICH, 1989
Size: 2.3 mm.
Morphology: (Family and Genus) Body habitus short and ro-bust. Eyes absent. Pereonites 1-2 reduced. Antennula withthree articles, distal article bearing setae with complex tips.Maxilliped endites fused. Cheliped attachment via sidepiece.Pereopods I-III coxae absent. Pereopod I different from II-III.Pereopods II-VI carpus antero-inferior spine flattened, “blade-like”. Uropod endopod with two articles; exopod with two arti-cles. One pair of oostegites only on pereopods IV. (Species)pereonites 1-3 combined shorter than pereonite 4. Cheliped offorcipate type. Pereopod I slender, basis about 12 times longerthan broad. Pereopods II-III similar, basis 6 times longer than
broad. Pereopods II-VI carpal blade-like spine long. PereopodsIV-VI similar dactylus and unguis fused. Pleopods well devel-oped.
Biology: On Lucky Strike area, this species was frequently col-lected from samples of volcanic rocks (hyaloclastic and lava)and sulphide rubble but also in areas of diffuse venting and nearactive chimneys active.
Distribution: North Atlantic: North Feni Ridge, RockallTrough, Hebridian slope, Porcupine Seabight and Celtic Slope,1028-1640 m; Mid-Atlantic Ridge: Lucky Strike.
References:
BIRD G.J. & D.M. HOLDICH (1989) Zool. J. Linnean Soc. 97: 233-298.LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.
M.R. CUNHA & M. SEGONZAC Denisia 18 (2006): 378
1: A: Female habitus, dorsal view; B: Male habitus, dorsal view; C: Right cheliped, female; D: Antenulla, male; scale bar 1 mm;from BIRD & HOLDICH (1989).
Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Pseudotanaidae
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Arthropoda, Crustacaea, Isopoda
Recent expeditions to the Lucky Strike vent field at theMid-Atlantic Ridge provided an uncommon collection of per-acarid crustaceans (CUNHA et al. 2001, CUNHA & WILSON
2003, in press) that accounted for about 50% of the speciesrichness in the samples and included about 20 isopod species.Among these four asellotes are new species: the two Heterome-sus presented herein, one undescribed Katianiridae and one un-described Munnopsidae. These species were collected in areasof diffuse low venting, in peripheral mussel (Bathymodiolusazoricus) aggregations and in a zone of filter feeding organisms.Nevertheless, we cannot infer that these isopods are vent en-demic because they do not show special adaptations to the en-vironment and our knowledge of their distribution is minimal.
The Lucky Strike isopod fauna also includes a number ofknown deep-sea asellotes (mostly Haplomunnidae, Haplonisci-
dae, and Munnopsidae) and a few species of other isopods (An-thurida, Cirolanidae, and Gnathiidae, unpublished data). Someof these were already cited in the previous version of the Hand-book (DESBRUYÈRES & SEGONZAC 1997) together with the epi-carid Thermaloniscus cotylophorus from East Pacific Rise: 13ºN.
Isopods are among the most abundant and diverse taxa inthe deep sea (HESSLER & SANDERS 1967, HESSLER et al. 1979;RAUPACH et al. 2004), but there are only very few records ofthese crustaceans in hydrothermal communities. The reasonsfor this are unclear because isopods, especially asellotes, areknown for their adaptative potential (HESSLER & THISTLE
1975) and at least in relatively shallow (<2000 m) hydrother-mal vent fields, such as in Lucky Strike, the less toxic environ-ment likely facilitates the immigration of background species(VAN DOVER 1995).
References:
CUNHA M.R. & G.D.F. WILSON (2003) Zootaxa 323:1-16.CUNHA M.R. & G.D.F. WILSON (in press) Zootaxa.CUNHA M.R., HILARIO A.M. & I.G. TEIXEIRA (2001) IOC Workshop Report 175: 73-74.DESBRUYÈRES D. & M. SEGONZAC (1997) Handbook of Deep-Sea Hydrothermal Vent Fauna. Ifremer Ed.: 1-279.HESSLER R.R. & T.D. THISTLE (1975) Mar. Biol. 32: 155-165.HESSLER R.R. & H.L. SANDERS (1967) Deep-Sea Research 14: 65-78.HESSLER R.R., WILSON G.& D. THISTLE (1979) Sarsia 64: 67-76.LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.RAUPACH M.J., HELD C. & J.-W. WÄGELE (2004) Deep-Sea Research II 51: 1787-1795.VAN DOVER C.L. (1995) in PARSON L.M., WALKER C.L. & D.R. DIXON (Eds.) Hydrothermal vents and processes. Geol. Soc. Spec. Publ. 87: 257-294.
M.R. CUNHA Denisia 18 (2006): 379
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Heteromesus calcar CUNHA & WILSON, in press
Size: 4 mm.
Morphology: (Family) Body elongate, subcylindrical and nar-row. Pereonites 4-5 elongate, pereonite 5 longest, pereonite 4widest anteriorly, 5 widest posteriorly. Head fused and embed-ded in pereonite 1. Pereonites 1 (posterior margin) to 4 free andarticulating. Anus separated from branchial chamber. Eyes ab-sent. Antennula terminating with simple seta, article 1 squatand globular, article 2 elongate, at least twice as long as article1. Antenna length more than half body length, without squa-ma. Maxilla inferior margin with two medial pectinate setae.Pereopod I robust strongly subchelate; pereopods II-VII ambu-latory. Uropod uniramous terminal. (Genus) Pereonite 5 freelyarticulated with pereonite 6, articulations not expressed be-tween pereonite 6, pereonite 7, pleonite 1 and pleotelson. Pere-onite 4 produced posteriorly but always broader than long. An-tennula article 1 globular, article 2 inserting dorsally; article 2strongly curved anteriorly at proximal insertion; articles distalto article 2 reduced to 1-3 articles, distal articles altogether
tiny, length less than 0.3 times of article 2 length. Pereopod Icarpus distally expanded, widest point distal to midpoint of car-pus, with one elongate robust seta and a proximal shorter robustseta. Pleopod II female operculum with narrow proximal neck,almost circular posteriorly, with plumose setae. Pleopod III ex-opod with plumose setae and fringe of fine setae. Pleopod Vabsent. Uropod with single article, conical, tapering distally;extending beyond posterior margin of pleotelson. (Species)Pereonites 1-3 with anterolateral spines. Pereonite 2 in femalewith no paired dorsal spines or tubercles. Pereonite 5 in femalelength 2.0 times the width. Antennula with 3 articles altogeth-er. Pereopods IV-V bases with pedestal spines; ischia with elon-gate pedestal spines.
Biology: Collected in the vicinity of active venting sites on sul-phide deposits, sulphide rubble and volcanic rocks.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
CUNHA M.R. & G.D.F. WILSON (in press) Zootaxa.
M.R. CUNHA & G.D.F. WILSON Denisia 18 (2006): 380
1: Female (left) head, lateral view and habitus, dorsal and lat-eral views; scale bar 1 mm; by M. Cunha.
2: Male (right) habitus, dorsal view, and pleotelson, dorsal andventral views; scale bar 1 mm; by M. Cunha.
Arthropoda, Crustacea, Isopoda, Asellota, Ischnomesidae
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Heteromesus ctenobasius CUNHA & WILSON, in press
Size: 4 mm.
Morphology: (Family) Body elongate, sub-cylindrical and nar-row. Pereonites 4-5 elongate, pereonite 5 longest, pereonite 4widest anteriorly, 5 widest posteriorly. Head fused and embed-ded in pereonite 1. Pereonites 1 (posterior margin) to 4 free andarticulating. Anus separated from branchial chamber. Eyes ab-sent. Antennula terminating with simple seta, article 1 squatand globular, article 2 elongate, at least twice as long as article1. Antenna length more than half body length, without squa-ma. Maxilla inferior margin with two medial pectinate setae.Pereopod I robust strongly subchelate; pereopods II-VII ambu-latory. Uropod uniramous terminal. (Genus) Pereonite 5 freelyarticulated with pereonite 6, articulations not expressed be-tween pereonite 6, pereonite 7, pleonite 1 and pleotelson. Pere-onite 4 produced posteriorly but always broader than long. An-tennula article 1 globular, article 2 inserting dorsally; article 2strongly curved anteriorly at proximal insertion; articles distal
to article 2 reduced to 1-3 articles, distal articles altogethertiny, length less than 0.3 times of article 2 length. Pereopod Icarpus distally expanded, widest point distal to midpoint of car-pus, with one elongate robust seta and a proximal shorter robustseta. Pleopod II female operculum with narrow proximal neck,almost circular posteriorly, with plumose setae. Pleopod III ex-opod with plumose setae and fringe of fine setae. Pleopod Vabsent. Uropod with single article, conical, tapering distally;extending beyond posterior margin of pleotelson. (Species)Pereonite 1-3 median pedestal spines. Pereonites 4-5 in femalewith lateral pedestal spines. Pleotelson terminal margin withpedestal spines. Antennula with five articles altogether. Pereo-pod II-VII bases and ischia with pedestal spines.
Biology: Collected in the vicinity of active venting sites onvolcanic rocks.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
CUNHA M.R. & G.D.F. WILSON (in press) Zootaxa.
M.R. CUNHA & G.D.F. WILSON Denisia 18 (2006): 381
1: Female habitus, dorsal view; the color picture is a false color composite of three different SEM images,by G.D.F. Wilson & S. Lindsay © Australian Museum.
Arthropoda, Crustacea, Isopoda, Asellota, Ischnomesidae
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Arthropoda, Crustacea, Amphipoda
Currently, 24 species of Amphipoda are described fromhydrothermal vent samples. In addition, some other species we-re collected in the surroundings of vents or at stations notstrictly linked with hydrothermal vent communities. A certainnumber of new species are currently being described.
The following preliminary conclusions can be drawn fromthe species so far identified (BELLAN-SANTINI 1998):
(1) The subfamily of Lysianassoidea (including Lysianassi-dae sensu lato and Uristidae) is exclusively found in Pacificsamples with 10 species.
(2) Twelve other families are represented by one to threespecies.
(3) The number of endemic species is very high. However,the large repartition of species most likely is due to insufficientidentification and appears doubtful.
(4) Four species are abundant and build swarms: Ventiellasulfuris and Halice hesmonectes in Pacific Ocean and Bouvierellacurtirama and Luckia striki in Atlantic Ocean (VAN DOVER et al.1992; MARTIN et al. 1993; SHEADER et al. 2003). The other spe-cies are known to occur only in small numbers per sample.
(5) Different techniques have been used in the past to sam-ple amphipods. Trawls are not recommended because large sur-faces are sampled without distinction of communities (in thecase of the sampling of the Guaymas area). Also, only large an-imals are collected and the delicate amphipods are often bro-ken. Slurp guns or grabs manipulated by the submarine are thebest methods for collecting fragile animals but for rare speciesthey are not very efficient. Washing of mytilids, tubeworms andother species is a good method if rinsing of samples is conduc-ted carefully. Sediment traps deployed for a certain period of ti-me are also a good, but probably selective method.
Amphipods are delicate and fragile animals in particularthe representatives of the deep-sea families. They frequentlyloose appendices and damaged individuals are difficult to de-termine. Generally there are only one or two individuals of aspecies per sample. Consequently, if they are damaged identifi-cations and descriptions are impossible. Alcohol is the best fi-xative, but colors will vanish after preservation.
References:
BELLAN-SANTINI D. (1998) Cah. Biol. Mar. 39: 143-152.BELLAN-SANTINI D. (2005) J. Nat. Hist. 39(39): 3435-3452.MARTIN J.W., FRANCE S.C. & C.L.VAN DOVER (1993) Can. J. Zool. 71: 1724-1732.SEGONZAC M. (1992). C.R. Acad. Sci. Paris. 314: 593-600.SHEADER M. VAN DOVER C.L. & M.H. THURSTON (2004) Mar. Biol. 144: 503-514.VAN DOVER C.L., KAARTVEDT S., BOLLENS S.M., WIEBE P.H., MARTIN J.W. & S.C. FRANCE (1992) Nature 358: 25-26.VINOGRADOV G.M. (1993) Zool. Zh. 72(2): 40-53.
D. BELLAN-SANTINI Denisia 18 (2006): 382
1: Ventiella sulfuris from East Pacific Rise: 9°N; by M. Bright.
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Ampelisca romigi BARNARD, 1954
References:
BARNARD J.L. (1954) Allan Hancock Pac. Exped. 18(1): 1-137.BARNARD J.L. (1966) Allan Hancock Pac. Expec. 27(5):1-166.VINOGRADOV G.M. (1993) Zool. Zh. 72(2): 40-53 [in Russian].
D. BELLAN-SANTINI Denisia 18 (2006): 383
Arthropoda, Crustacea, Amphipoda, Gammaridea, Ampeliscidae
1: Female 9.5 mm;after BARNARD (1966).
Synonyms: A. isocornea: BARNARD, 1954, A. romigi ciegoBARNARD, 1966.
Size: Up to 16 mm.
Morphology: (Family) Mouthparts basic. Eyes composed ofcorneal lenses or absent. Body without process except uro-somite 1. Urosomites 2 and 3 coalesced. Antenna 1 without ac-cessory flagellum. Gnathopods 1 and 2 slender subchelate ornearly simples. Pereopods 3-4 slender with merus elongate.(Genus) Pereopods 5-6 with basisvery broad. Pereopod 7 basisdilated with posterior lobe greatly expanded distally, bearingmarginal plumose setae. Telson longer than broad and deeplycleft. (Species) Eyes present. Pereiopod 7 lower edge of basisreaching joint between merus and carpus, merus with a posteri-or setose lobe, carpus with distal anterior edge notched.
Epimeral plate posterior edge convex, lower posterior cornerquadrate. Urosomite 1, dorsal surface with a weakly seddle-shaped process. Telson rather broad. (Subspecies A. romigiciego) Eyes absent. Pereiopod 7 with propodus more stronglynarrowed in the proximal part while the propodus is wider.
Biology: Unknown. Dredged mainly from soft-bottom areas.Trawl in periphery of hydrothermal vents.
Distribution: East Pacific Rise. Along the coast of southernCalifornia and in the Channel Islands; Gulf of California atAngel de la Guardia Island, Isla Partida; Angeles Bay, San Mar-cos Island, Tortuga Island and Tiburon Island; Isabel Island,Mexico; Salinas Bay, Costa Rica; Secas Islands, Panama; SantaElena Bay, Ecuador, 3-500 m. The subspecies A. romigi ciego wasfound also at Guaymas vent site.
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Gitanopsis alvina BELLAN-SANTINI & THURSTON, 1996
Size: 2.5 mm in adult female.
Morphology: (Family) Coxa 4 immensely broadened, coxa 1very small and hidden by coxa 2. Peduncle of uropode 3 elon-gate. Telson entire elongate. (Genus) Mandibular molar large,triturative. Dactylus of gnathopods simple. (Species) Head withrostrum. Accessory flagellum one-articulated. Antenna 2slightly longer than antenna 1. Gnathopods small and feeble.Epimeral plate 3 posterodistal margin rounded. Uropod 3, pe-duncle 1.5x longer than inner ramus, outer ramus shorter thaninner. Telson elongate, triangular, apex tridentate.
Biology: Collected in hydrothermal field.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
BELLAN-SANTINI D. & M. THURSTON (1996) J. Nat.Hist. 30: 685-702.
D. BELLAN-SANTINI Denisia 18 (2006): 384
1A: Head incomplete of female; B: Head of juvenile, C: Antenna 1, D: Gnathopod 1, E: Epimeral plates, F: Uropod 3, G: Telson;from BELLAN-SANTINI & THURSTON (1996).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Amphilochidae
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Autonoe longicornis CHEVREUX, 1909
Size: 8 mm.
Morphology: (Family) Head anteroventral margin weakly re-cessed, moderately excavate. Pereopods 3-4 basis glandular,pereopod 7 very elongate. Gnathopod 1 enlarged in males andfemales. Uropod 3 with slender and robust setae. Telsondorsoventrally thickened. Species, male: Antennae 1 and 2subequal in length, less than half body length. Antenna 1 pe-duncle article 3, one quarter length of article 1; flagellum short-er than peduncle; accessory flagellum composed of one rudi-mentary article. Mandible palp article 3 longer than article 2,terminally falcate. Maxilla 1 inner plate with a single, long,pectinate seta. Labium mandibular processes acute. Gnathopod1 coxa produced anterodistally, subacute; basis robust; carpusenlarged, longer than propodus, the posterodistal corner pro-duced into a spine. Gnathopod 2 carpus and propodus elongate,subequal in length. Pereopods 5-7 basis only weakly expanded.Epimera 1-3 rounded. Uropod 1 peduncle with inter-ramal
spine, about one third length of peduncle; inner ramus longerthan outer and subequal in length with peduncle. Uropod 2 pe-duncle with short inter-ramal spine about one sixth length ofpeduncle; inner ramus longer than outer and subequal with pe-duncle. Uropod 3 peduncle longer than broad, rami subequaland only a little longer than peduncle, inner ramus with smallsecond article. Telson with each dorsal crest bearing a fine seta.
Biology: Collected from samples of volcanic rocks (pillow lavaand hyaloclastic rocks). The proximity of active venting wasinferred from the occurrence of Bathymodiolus azoricus (livingspecimens and/or shell debris) in the samples. Previouslyknown only by the type specimens (only females) collected byCHEVREUX (1909) from similar depths (1360 m) also in theAzores region.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Arthropoda, Crustacea, Amphipoda, Corophiidea, Aoridae
1: Male; by A.A. Myers. G1 – gnathopod 1; G2 – gnathopod 2; Lab – labium; Md – mandible; Mx1 – maxilla; Mxp –maxilliped; T – telson; U1, U2, U3 – uropods 1-3.
References:
CHEVREUX E. (1909) Bull. Inst. Océanogr. Monaco 150: 1-7.MYERS A.A. & M.R. CUNHA (2004) J. Mar. Biol. Ass. U.K. 84: 1019-1025.
A.A. MYERS & M.R. CUNHA Denisia 18 (2006): 385
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Oradarea longimana (BOECK, 1871)
References:
BOECK A. (1871) Forhandl. Vidensk.-Selskabet i Christiania 1870: 83-280.SHAW P. (1989) Can. J. Zool. 67(8): 1882-1890.SHOEMAKER C.R. (1930) Contr. Canad. Biol. Fisheries, new series 5(10): 221-359.
D. BELLAN-SANTINI & D. JAUME Denisia 18 (2006): 386
1A: Posterior part of the body; B: Head; C: Gnathopod 1; D: Gnathopod 2; E: Telson; from SHOEMAKER (1930).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Calliopiidae
Size: Up to 11 mm.
Morphology: (Family) Lateral cephalic lobes small. Coxae ofmedium length or short. Antennae long; accessory flagellumvestigial or absent. Mouthparts basic. Gnathopods powerful orfeeble, usually subchelate, occasionnally simple. Telson ofmedium size, entire, acuminate, emarginate or notched.(Genus) Rostrum large. Accessory flagellum one-articulated.Antenna 2 longer than 1. Gnathopods diverse. Pereopods 3-7ordinary. Epimeron 3 smooth. (Species) Accessory flagellumvery short. Both gnathopods very slender; gnathopod 2 muchlonger than 1. Pereopods 3-4 alike in structure. Urosomite 1-2dorsally produced backward. Epimeron 3 with lower posteriorangle very obtuse and postero-lateral margin evenly convex.Uropod 3 peduncle one quarter of inner ramus and half of out-er. Telson a little longer than wide, apex rounded with a veryshallow cental notch.
Biology: Frequently associated to other invertebrates. SHOE-MAKER (1930) describes a collection from whelk egg cases, per-haps as a scavenger or predator of eggs. One of samples from theJuan de Fuca vent sites was taken from the decapod Macrorego-nia macrochira SAKAI. Oradarea is a genus attracted to baitedtraps, and it seems probable that these amphipods are scav-engers that cue on exudates of egg capsules and possibly pre-moult decapods.
Distribution: Recorded from the North Atlantic (54 m depth)and the N Pacific coast of North America, and probably broad-ly extended along the northern Hemisphere. Present also at hy-drothermal vent sites of the Juan de Fuca Ridge (EndeavourSegment, Lieutenant Obo vent, Dual Smoker; Explorer Seg-ment, Upper Magic Mountain, Magic Mountain.
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Caprella bathytatos MARTIN & PETTIT, 1998
Size: Largest male 14.9 mm; smallest juvenile 2.2 mm.
Morphology: (Family) Body slender, cylindrical. Eyes lateral.Gnathopods often different. Number of pereopods variable.(Genus) Mandible lacking palp. Antenna 2, flagellum two-ar-ticulate. Pereopods 3-4 lacking. Pereopods 5-7 normal.(Species) Head rounded, lacking spines or projectings. Eyes re-duced. Pereon lacking any spines or projections. Gnathopod 2,propodus bears one blunt, rectangular tooth, separated by a U-shaped notch from a slightly more acute tooth, an acute proxi-mal lobe, dactylus 2/3 length of propodus; gnathopod attachedto body at midlength of pereionite 2. Each pereopod with grasp-ing spines on propodus forming with the dactylus a mechanismfor attachment to host setae.
Biology: It is the first caprellid found in association with thecrab Macroregonia macrochira, or in the vicinity of marine hy-drothermal vent. 30 individuals were sampled on a single crab.
Distribution: Juan de Fuca Ridge.
Reference:
MARTIN J.W. & G. PETTIT (1998) Bull. Mar. Sci. 63: 189-198.
D. BELLAN-SANTINI Denisia 18 (2006): 387
1A: Habitus of male, attached to maxilliped 3 of the majid crab; B: Gnathopod 2; C: Pereopod 7; D: Abdomen, male; E: Abdominal appendage; from MARTIN & PETTIT (1998).
Arthropoda, Crustacea, Amphipoda, Caprellidea, Caprellidae
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Bouvierella curtirama BELLAN-SANTINI & THURSTON, 1996
Size: 5 mm in adult female.
Morphology: (Family) Accessory flagellum 0-2-articulated.Gnathopods variable. (Genus) Body not toothed. Eyes absent.Coxal plates 1-4 ordinary. Epimeron 3 smooth. (Species) Head,rostrum moderate, lateral cephalic lobe rounded. Antenna 1longer than antenna 2, flagellum with a number of articles vari-able with the size of the body, 8-46, 22 in the holotype. An-tenna 2 flagellum variable about 7-37, 12 in holotype.Gnathopods weakly subchelate, dissimilar, second longer thanfirst. Pereopods 3-7 ordinary, dactylus simple not prehensile.Epimeral plate 3 smooth. Telson rounded, without armament.
Biology: B. curtirama is abundant. This species lives in Bathy-modiolus azoricus beds. Females maturing at a minimum bodylength of 3.5 mm and males at 2.4 mm.
Distribution: Mid-Atlantic Ridge: Lucky Strike, sites Sintraand Tour Eiffel sites.
References:
BELLAN-SANTINI D. & M.H. THURSTON (1996) J. Mar. Biol. Ass. U.K. 30: 685-702.SHEADER M., VAN DOVER C.L. & M.H. THURSTON (2004) Mar. Biol. 144: 503-514.
D. BELLAN-SANTINI Denisia 18 (2006): 388
Arthropoda, Crustacea, Amphipoda, Gammaridea, Eusiridae
1: A: Habitus; B: Gnathopod 1; C: Epimeral plates; D: Uropod 1; E: Uropod 2; F: Uropods 3 and telson; from BELLAN-SANTINI &THURSTON (1996).
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Luckia striki BELLAN-SANTINI & THURSTON, 1996
Size: 7 mm in adult female.
Morphology: (Family) Accessory flagellum 0-2-articulated.Gnathopods variable. (Genus) Body normally compressed.Rostrum short. Antenna 1 longer than antenna 2. Accessoryflagellum one-articulated. Labrum entire, molar triturative.Gnathopods different in size. Epimeral plate 3 smooth. Outerrami of uropods 1-3 shorter than inner rami. Telson cleft with-out armament. (Species) Body smooth. Eyes absent. Rostrumshort. Antenna 1 long as body length. Antenna 2 slightlylonger than half of antenna 1. Gnathopods slender, linear. Tel-son triangular, cleft.
Biology: Collected in vent community among shrimps, gas-tropods, crabs, and limpets.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
BELLAN-SANTINI D. & M. THURSTON (1996) J. Nat.Hist. 30: 685-702.
D. BELLAN-SANTINI Denisia 18 (2006): 389
1: A: Habitus; B: Accessory flagellum; C: Gnathopod 2; D: Epimeral plates; E: Uropod 1; F: Uropod 2;G: Uropods 3 and telson; from BELLAN-SANTINI & THURSTON (1996).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Eusiridae
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Bonnierella compar MYERS & CUNHA, 2004
Size: 3 mm.
Morphology: Family: Head anteroventral margin moderately tostrongly recessed and moderately excavate. Gnathopod 2 en-larged in males and females. Pereopods 3-4 basis glandular.Uropod 3 peduncle broad proximally, narrow distally, rami withtiny apical setae. Telson dorsoventrally thickened. Species:Head anteroventral margin strongly regressed. Eyes absent. An-tenna 1 and 2 subequal in length, setose; accessory flagellumwith one long and one rudimentary article. Maxilliped raptori-al, with reduced plates and strong palp terminating in a dactyli-form palp article 4. Gnathopod 1 small, unornamented.Gnathopod 2 similar in both sexes, massive, with palmar exca-vation and with defining spine on posterior margin. Uropod 3
peduncle elongate, broad proximally, narrow distally, ramimuch less than half length of peduncle. Telson without cusps orspines.
Biology: Collected in the vicinity of active venting sites to-gether with filter-feeding organisms (small sponges, hydrozoansand cirripeds). Some of the specimens of B. compar were foundinside their tubes of fine pelagic sediments cemented with mu-cus, in some cases built around hydrozoan stalks. A previousrecord of the genus in hydrothermal vents is given by SHAW
(1989) who collected a single specimen of B. linearis from a sta-tion at the Juan de Fuca Ridge.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
References:
MYERS A.A. & M.R. CUNHA (2004) J. Mar. Biol. Ass. U.K. 84: 1019-1025. SHAW P. (1989) Can. J. Zool. 67: 1882-1890.
A.A. MYERS & M.R. CUNHA Denisia 18 (2006): 390
Arthropoda, Crustacea, Amphipoda, Corophiidea, Ischyroceridae
1: By A. Myers. FG1, FG2 – female gnathopods 1 and 2; MG2 – male gnatophod 2; Lbr – labrum; Md – mandible; Mx1 –maxilla; Mxp – maxilliped; T – telson; U1, U2, U3 – uropods 1-3.
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Bonnierella cf. linearis BARNARD, 1964
Size: Up to 4 mm.
Morphology: (Family) Head anteroventral margin moderatelyto strongly recessed and moderately excavate. Gnathopod 2enlarged in males and females. Pereopods 3-4 basis glandular.Uropod 3 peduncle broad proximally, narrow distally, rami withtiny apical setae. Telson dorsoventrally thickened. (Species)Eyes absent. Lateral lobes of head produced and acute. Anten-nae nearly as long as body. Gnathopod 1, palm not ornamed.Gnathopod 2, armed with three sharp cusps in male; with twoblunt process in female defining cusps obsolete. Uropod 1, in-ner ramus lacking a marginal spine. Uropod 3 with outer ramusbearing 5 to 6 minute fringe-like spinules and a distal setule.Telson triangular , the narrow apex blunt. The single specimenknown from hydrothermal vents, a female, differs slightly fromthe original description (BARNARD 1964) in the tuberosities ofthe palmar margin of gnathopod 2, minor differences in telson
setation, and number of spines in the mandibular spine row. Butvirtually all characters used to distinguish the present Bon-nieriella species are only present on the males, which makes thespecific assignment of this specimen dubious.
Biology: Unknown. At hydrothermal vents, collected fromwashings of associated vent fauna (vestimentiferans and ar-chaeogastropods).
Distribution: Off Peru, 10°13’S 80°05’W, 6324 m. Reported al-so from hydrothermal vents at Explorer Ridge, Juan de FucaRidge: site Gulati Gusher.
References:
BARNARD J.L. (1964) Bull. Am. Mus. Nat. Hist. 127(1): 1-46.SHAW P. (1989) Can. J. Zool. 67(8): 1882-1890.
D. BELLAN-SANTINI & D. JAUME Denisia 18 (2006): 391
Arthropoda, Crustacea, Amphipoda, Gammaridea, Ischyroceridae
1: Female 3.4 mm, with both antennules and antennae lost; after SHAW (1989).
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Apotectonia heterostegos BARNARD & INGRAM, 1990
Reference:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contrib. Zool. 499: 1-80.
D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 392
1: Female 18.3 mm. A: Head; B: Pleosome and urosome with attached uropods and telson; C: First gnathopod; D: Secondgnathopod; E: Proximal portion of pereopods 5-7; from BARNARD & INGRAM (1990).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
Size: Up to 18.3 mm.
Morphology: (Family) Body compact, robust. Coxae generallylarge. Antenna 1, peduncle article 1 large, inflated, articles 2-3much shorter, often telescoped. Mouthparts very variable.Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-acteristic for the family, slender and microchelate. Pereopods 5-7 basis usually broadly expanded. Telson variable. (Genus) Dor-sal process of urosomite 1 complexly toothed. Gnathopod 1simple. (Species) Antenna 1, primary flagellum with basalspines. Urosomite 1 with 2 sharp dorsal teeth. Mandibular rightlacinia mobilis tiny, bifid; left lacinia mobilis scarcely larger and
multitoothed; molar of both mandibles simple, pubescent. In-ner plate of either maxilla 1-2 fully setose medially. Inner plateof maxilliped with strongly oblique apex. Gnathopod 1 simpleand with reduced coxa. Basis of pereopods 5-7 rounded-attenu-ate. Oostegite present on coxa 1 of brooding females.
Biology: Largely unknown. Readily attracted to baited traps;some specimens sorted also from siboglinid washings.
Distribution: Galapagos Spreading Center, apparently endem-ic to the sites Garden of Eden and Rose Garden.
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Cyclocaris tahitensis STEBBING, 1888
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
1: Habitus; after STEBBING (1888).
Size: Up to 15 mm.
Morphology: (Family) Body compact, robust. Coxae generallylarge. Antenna 1 peduncle article 1 large, inflated, articles 2-3much shorter, often telescoped. Mouthparts very variable.Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-acteristic for the family, slender and microchelate. Pereopods 5-7 basis usually broadly expanded. Telson variable. (Genus)Head tall, horizontally short, weakly grotesque. Coxae 1-2small, strongly shortened and partly covered by coxa3. Coxa 4large excavate. Gnathopod 1 long, simple. Uropod 3 elongate.Telson elongate, deeply cleft. (Species) Base of primary flagel-lum of antenna 1 with callinophore. Body capable of coiling in-to a circle, bending its head round to the protection of the cox-ae of the third and fourth pereopods. Head extremely short,
lacking rostrum. Mandibular palp present. Coxa 1much widerat distal margin than at insertion. Epimeron 3 with posteriorangle rounded. Rami of uropod 1 about equal in length. Pe-duncle of uropod 2 shorter than rami.
Biology: Unknown. Originally described from volcanic mudbottoms around Tahiti and trapped also around Cape Verde onsandy bottoms, the only report of the species from hydrother-mal vents could be accidental since the accompanying faunawas also not typically hydrothermal.
Distribution: Off Tahiti, 17°30’26’’S, 149°33’45’’W (STEBBING
1888). Off Cape Verde Islands, 1477 m. East Pacific Rise,Guaymas Basin.
References:
CHEVREUX E. (1935) Résult. Camp. Sci. Prince Albert Ier de Monaco 90: 1-214. STEBBING T.R.R. (1888) Rep. Sci. Results Voyage H. M. S. Challenger Years 1873-1876, Zoology 29: 1-1737.VINOGRADOV G.M. (1993) Zool. Zh. 72(2): 40-53 [in Russian].
D. BELLAN-SANTINI & D. JAUME Denisia 18 (2006): 393
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Euonyx mytilus BARNARD & INGRAM, 1990
Reference:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contr. Zool. 499: 1-80.
D. BELLAN-SANTINI & D. JAUME Denisia 18 (2006): 394
1: Female 20.04 mm; A: anterior portion of body, lateral; B: pleosome and urosome with uropods and telson attached, lateral;C: First gnathopod; D: second gnathopod; E: fifth to seventh pereopods; from BARNARD & INGRAM (1990).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
Size: Up to 20 mm.
Morphology: (Family) Body compact, robust. Coxae generallylarge. Antenna 1 peduncle article 1 large, inflated, articles 2-3much shorter, often telescoped. Mouthparts very variable.Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-acteristic for the family, slender and microchelate. Pereopods 5-7 basis usually broadly expanded. Telson variable. (Genus)Mandible with palp attached strongly distal. Inner plate ofmaxilla 1 weakly setose. Coxa 1 strongly shortened. Gnathopod1 elongate. Inner ramus of uropod 2 without notch. Uropod 3ordinary. Telson elongate, deeply cleft. (Species) Body lackingany prominent dorsal tooth. Lateral cephalic lobes rounded.
Gnathopod 1 with propodus about 1.2 times as long as carpus.Palm of gnathopod 2 long. Epimeron 2 with a strong tooth.Epimeron 3 with tiny posteroventral tooth.
Biology: Members of this genus come readily to baited trapsand some appear occasionally associated with echinoderms anddeep-sea corals. Euonyx mytilus was sorted from washings ofvent clams, trapped also with baited traps, and also caught di-rectly with a slurp gun around vent fauna.
Distribution: East Pacific Rise, 13°N; Galapagos SpreadingCenter: Garden of Eden and Rose Garden.
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Hirondellea glutonis BARNARD & INGRAM, 1990
Reference:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contrib. Zool. 499: 1-80.
D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 395
1: Female 12.45 mm; A: head; B: Detail of coxal plates of anterior pereopods; C: Pleosome and urosome with attached uropodsand telson; D: First gnathopod; E: Second gnathopod; F: Proximal portion of pereopods 5-7; from BARNARD & INGRAM (1990).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
Size: Up to 12.6 mm.
Morphology: (Family) Body compact, robust. Coxae generallylarge. Antenna 1 peduncle article 1 large, inflated, articles 2-3much shorter, often telescoped. Mouthparts very variable.Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-acteristic for the family, slender and microchelate. Pereopods 5-7 basis usually broadly expanded. Telson variable. (Genus)Mandible with incisor ordinary, molar simple, palp attachedopposite molar. Coxa 1 strongly shortened. Gnathopod 1 short,subchelate, palm transverse. Gnathopod 2, propodus greatly
shorter than carpus, propodus minutely chelate. Telson elon-gate, cleft. (Species) Dactylus of gnathopod 1 scarcely overlap-ping palm. Epimera 3 rounded behind. Inner ramus of uropod 2constricted. Telson weakly cleft.
Biology: Attracted by baited traps, but caught also directly withslurp gun on and around, or sorted from vent mussel and clamwashings. Most specimens had the midgut densely packed withbait food, and with the sternites ventrally extended.
Distribution: Galapagos Spreading Center, East Pacific Rise.
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Orchomene (Abyssorchomene) abyssorum STEBBING, 1888
Size: 6.1 mm in adult male.
Morphology: (Family) Body compact. Peduncle of antenna 1short and stout, articles 2-3 much shorter than 1 and partlytelescoped. Article 3 of gnathopod 2 elongate. (Genus) Mouth-parts forming quadrate bundle. Coxa 1 large and visible not ta-pering. Gnathopod 1 subchelate. Inner ramus of uropod 2 with-out notch. (Species) Lateral cephalic lobes broadly rounded.Eyes long, lunate, colorless in alcohol, glandular, ommatidianot evident. Peduncle of antenna 1 barely keeled. Carpus ofgnathopod 1 short, posterior lobe narrow, well exceeding pos-terior tangential line between merus and propodus. Propodus ofgnathopod 2 with palm short, chelate, not lunate, dactylus
short and covering 100% of palmar edge, palm defined thinspinule and bearing weak inner setal basket. Article 2 of pere-opods 5-7 with sparse, weak posterior setule-notches. Epimeralplate 3 weakly sinuous posteriorly, produced into weak blunttooth posteroventrally, smooth. Dorsal process of urosomite 1hood-shaped, weakly over-vaulting urosomite 2. Telson cleftabout 50% of its length.
Biology: Sampled in mussel and clam beds.
Distribution: Galapagos Spreading Center; worldwide abyssaldistribution to 9000 m.
References:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contrib. Zool. 499: 1-80. THURSTON M.H. (1990) Prog. Oceanogr. 24: 257-274.
D. BELLAN-SANTINI Denisia 18 (2006): 396
1A: Head; B: Pleon; C: Gnathopod 1; D: Gnathopod 2; E: Pereopod 7; F: Uropod 3; G: Telson; from BARNARD & INGRAM (1990).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
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Orchomene (Abyssorchomene) distinctus BIRSTEIN & VINOGRADOV, 1960
Size: 12 mm in adult male.
Morphology: (Family) Body compact. Peduncle of antenna 1short and stout, articles 2-3 much shorter than 1 and partlytelescoped. Article 3 of gnathopod 2 elongate. (Genus) Mouth-parts forming quadrate bundle. Coxa 1 large and visible not ta-pering. Gnathopod 1 subchelate. Inner ramus of uropod 2 with-out notch. (Species) Lateral cephalic lobes broadly rounded.Eyes long, flask-shaped, very pale pink. Carpus of gnathopod 1short, posterior lobe narrow, scarcely exceeding posterior tan-gential line between merus and propodus. Propodus of gnatho-pod 2 with palm elongate, transverse, sinuate, dactylus long but
covering only 60% of palmar edge, palm defined by cusp. Arti-cle 2 of pereopods 5-7 with sparse, very weak posterior setule-notches. Epimeral plate 3 broadly rounded posteroventrally,smooth. Dorsal process of urosomite 1 hood-shaped, weaklyover-vaulting urosomite 2. Telson cleft about 53% of its length.
Biology: Sampled in sediments by slurp gun and in washings ofsiboglinid tubeworms.
Distribution: East Pacific Rise: 13°N; worldwide abyssal distri-bution to 4850 m.
References:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contrib. Zool. 499: 1-80. THURSTON M.H. (1990) Prog. Oceanogr. 24: 257-274.
D. BELLAN-SANTINI Denisia 18 (2006): 397
1A: Head; B: Pleon; C: Gnathopod 1; D: Gnathopod 2; E: Pereopod 7; F: Uropod 3; G: Telson; from BARNARD & INGRAM (1990).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
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Tectovalopsis diabolus BARNARD & INGRAM, 1990
Reference:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contrib. Zool. 499: 1-80.
D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 398
1: Female 19.1 mm; A: Head; B: Pleosome and urosome with uropods and telson attached; C: Antennule; D: Second gnathopod;from BARNARD & INGRAM (1990).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
Size: Up to 23.4 mm.
Morphology: (Family) Body compact, robust. Coxae generallylarge. Antenna 1 peduncle article 1 large, inflated, articles 2-3much shorter, often telescoped. Mouthparts very variable.Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-acteristic for the family, slender and microchelate. Pereopods 5-7 basis usually broadly expanded. Telson variable. (Genus)Mandible with molar large, conical, setulose; palp attachedstrongly distal of molar. Coxa 1 strongly shortened. Gnathopodstrongly subchelate. Urosomite 1 carinate. Telson elongatedeeply cleft. (Species) Mandibular incisor with two teeth; rightlacinia mobilis very small, flake-like, much broader than long;left lacinia also broader than long, evenly serrate; molar coni-
cal, densely setulose, tapering to tiny apical plaque with weaktriturative surface. Inner plate of maxillae with medial setae.Gnathopod 1 not elongate and subchelate, with reduced coxa.Palm of gnathopod 2 long, strongly oblique. Teeth on epimera2-3 weak. Epimeron 2 lacking facial spine. Keel of urosomite 1simple. Urosomite 3 with erect dorsal keel.
Biology: Unknown. Collected with slurp gun or baited traps.
Distribution: East Pacific Rise: 13°N. Apparently endemic tothis vent site.
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Tectovalopsis wegeneri BARNARD & INGRAM, 1990
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
Size: Up to 33.9 mm.
Morphology: (Family) Body compact, robust. Coxae generallylarge. Antenna 1 peduncle article 1 large, inflated, articles 2-3much shorter, often telescoped. Mouthparts very variable.Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-acteristic for the family, slender and microchelate. Pereopods 5-7 basis usually broadly expanded. Telson variable. (Genus)Mandible with molar large, conical, setulose; palp attachedstrongly distal of molar. Coxa 1 strongly shortened. Gnathopod1 strongly subchelate. Urosomite 1 carinate. Telson elongatedeeply cleft. (Species) Similar to the foregoing T. diabolusBARNARD & INGRAM, 1990 except for: (1) the wider peduncleof antenna 1; (2) the slightly less tapering coxa 1; (3) the less
adz-shaped coxa 2; (4) the slightly stouter lobe of coxa 4; (5)the slightly weaker gnathopods with shorter carpus and propo-dus on gnathopod 2; (6) the less oblique palm of gnathopod 2;(7) the more numerous and more widely spread spines onepimera 2-3; (8) the less stronger dorsal projection of urosomite2 and dorsal keel of urosomite 3; and (9) the outer ramus of uro-pod 2 a bit longer.
Biology: Largely unknown. Collected with slurp gun aroundAlvinella polychaetes.
Distribution: East Pacific Rise: 13°N. Apparently endemic tothis vent site.
1: Female 33.9 mm long; from BARNARD & INGRAM (1990).
Reference:
BARNARD J.L. & C. INGRAM (1990) Smithon. Contrib. Zool. 499: 1-80.
D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 399
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Transtectonia torrentis BARNARD & INGRAM, 1990
Reference:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contrib. Zool. 499: 1-80.
D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 400
1: Female 10.6 mm; A: Head; B: Pleosome and urosome with uropods and telson attached; C: Coxal plates of first and secondgnathopods; D: First gnathopod; from BARNARD & INGRAM (1990).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae
Size: Up to 21.2 mm.
Morphology: (Family) Body compact, robust. Coxae generallylarge. Antenna 1 peduncle article 1 large, inflated, articles 2-3much shorter, often telescoped. Mouthparts very variable.Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-acteristic for the family, slender and microchelate. Pereopods 5-7 basis usually broadly expanded. Telson variable. (Genus)Mandible with incisor strongly toothed, laciniae mobilis longerthan broad, molar large, conical. Coxa 1 not shortened and nostrongly covered by coxa 2. Pereopods 5-7 elongate, basis
strongly tapering. Urosomite 1 with a sharp dorsal tooth.Telosn elongate, deeply cleft. (Species) Antennae especiallyshort. Inner plate of maxillae setose medially only along distalhalf. Gnathopod 1, palm strongly oblique. Epimeron 3 roundedbehind with six spines narrowly spread.
Biology: Caught associated with worms and attracted to baitedtraps.
Distribution: East Pacific Rise: 13°N. This monotypic genus isapparently endemic to this vent site.
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Halice hesmonectes MARTIN, FRANCE & VAN DOVER, 1993
Size: 5.3 mm in adult male.
Morphology: (Family) Body laterally compressed, coxae short.Accessory flagellum well developed, molar absent. (Genus)Rostrum well developed. Eyes absent. Gnathopods simple, slen-der. Pereopods simple. (Species) Teeth of urosomites 1-2strong. Accessory flagellum biarticulate. Dactylus of pereopods3-4 minute, constricted at mid length. Pereopods 5-7 extreme-ly long. Telson cleft along 2/3 and terminating in acute tip.
Biology: In large monospecific swarms in the immediate vicin-ity of low temperature vent openings. 1000 ind. l-1 estimated.
Distribution: East Pacific Rise: 9°N.
Reference:
MARTIN J., FRANCE S.C. & C.L. VAN DOVER (1993) Pacific Can. J. Zool. 71: 1724-1732.
D. BELLAN-SANTINI Denisia 18 (2006): 401
1A: Habitus; B: Accessory flagellum; C: Lateral view of urosome; D: Gnathopod 1; E: Dorsal view of urosome and telson; fromMARTIN et al. (1993).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Pardaliscidae
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Pardalisca endeavouri SHAW, 1989
Reference:
SHAW P. (1989) Can. J. Zool. 67(8): 1882-1890.
D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 402
Arthropoda, Crustacea, Amphipoda, Gammaridea, Pardaliscidae
1: Male 11.5 mm; after SHAW (1989).
Size: 11.5 mm.
Morphology: (Family) Body laterally compressed, coxae short.Accessory flagellum well developed. Molar absent. Inner plateof maxilliped short to evanescent. (Genus) Rostrum small. Eyesabsent. Mandible asymetrical, incisor on left smooth, weaklytoothed, on right strongly toothed. Palp of maxilla 1 apicallyexpanded. Coxae 1-4 subquadrate. Gnathopods simple. Telsonscarcely elongate, partly cleft. (Species) Body slender smooth.Antenna 2 with peduncle segments 3 and 4 elongate. Epimeron
3, posteroventral corner acute. Telson cleft to 80%, lobes wide-ly separated, each apex tridentate, bearing paired marginalspines, in addition to paired penicillate setae on the upper face.
Biology: Unknown. Collected with slurp gun from the vicinityof siboglinids and archaeogastropods.
Distribution: Explorer Ridge: site Gulati Gusher.
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Seba profundus SHAW, 1989
Size: Up to 3.5 mm.
Morphology: (Family) Urosomites 2-3 free or coalescent. An-tenna 1, peduncle elongate, accessory flagellum two-articulate.Gnathopods 1-2 chelate or gnathopod 1 strongly subchelate.Gnathopod 1 larger than gnathopod 2. Uropod 3 uniramous.Telson entire. (Genus) Body slender. Urosomites 2-3 coales-cent. Gnathopods diverse. Gnathopod 1 much the larger, sub-chelate or chelate. Gnathopod 2 strongly chelate. Telson lin-guiform. (Species) Body stout. Antennae subequal. Epimeron 3lacking large posterodistal tooth. Gnathopod 1, subchelate,with posterodistal corner of propodus extending out from lon-
gitudinal axis of limb. Gnathopod 2 elongate, propodus slender.Basis of pereopods 5-7 evenly expanded, with convex anteriorand posterior margins. Telson triangular, armed with two pairsof marginal setae.
Biology: Sorted from samples of associated vent fauna (the si-boglinid Ridgeia piscesiae and archaeogastropods). Other mem-bers of the genus are typically associated either as a commensalor as inquiline with invertebrate hosts.
Distribution: Explorer Ridge: Crab Vent.
Reference:
SHAW P. (1989) Can. J. Zool. 67(8): 1882-1890.
D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 403
1: Male 3.4 mm; after SHAW (1989).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Sebidae
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Steleuthera ecoprophycea BELLAN-SANTINI & THURSTON, 1996
Size: 3 mm in adult female.
Morphology: (Family) Body appears stout and globular. Coxae1-4 forms lateral shield. Antennae very short. Mandible with-out molar and palp. Gnathopods feeble. (Genus) Accessory fla-gellum biarticulate. Mandibular incisor toothed. Pereopods 3-4simple. Article 2 of pereopod 6 unexpanded. Pereopod 7 with 7articles. (Species) Epimeron 3 with postero-distal cornerprominent and without seta. Gnathopods 1-2 with propodus ta-pered distally. Basis of pereopod 7 serrulate. Telson broaderthan long, scarcely cleft.
Biology: Collected with shrimps Rimicaris exoculata and Choro-caris chacei, and in mussel washing.
Distribution: Mid Atlantic Ridge: Snake Pit.
Reference:
BELLAN-SANTINI D. & M. THURSTON (1996) J. Nat.Hist. 30: 685-702.
D. BELLAN-SANTINI Denisia 18 (2006): 404
1: A: Habitus; B: Antenna 1; C: Mandible; D: Gnathopod 1; E: Pereopod 7; F: Epimeral plates; G: Uropod 3 and telson; from BELLAN-SANTINI & THURSTON (1996).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Stegocephalidae
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Metopa (Prometopa) samsiluna BARNARD, 1966
Size: 4.5 mm.
Morphology: (Family) Coxa 1 very small and partially coveredby following coxae. Coxa 4 enlarged, shield-like, not posteri-odorsally excavate. Uropod 3 uniramous. Telson entire.(Genus) Antenna 1 lacking nasiform process. Accessory flagel-lum absent or vestigial. Palp of mandible two- to three-articu-late. Gnathopods 1-2 subchelate, different from each other insize and shape. Pereopod 5 with basis rectolinear. Pereopod 6-7with expanded lobate basis. (Species) Eyes absent Antennaevery long. Accessory flaggellum vestigial. Coxa 2 very broad.Gnathopod 1 short. Gnathopod 2 with a large medial tooth,defining corner with large tooth. Telson spinose.
Biology: Unknown.
Distribution: Originally described from the submarine canyonsof off southern California (San Clemente Rift Valley, 32°44’N,118°12’W, 1096-1620 m). Found also in the Gulf of California,in the Guaymas Basin.
References:
BARNARD J.L. (1966) Allan Hancock Pac. Exped. 27(5): 1-166.VINOGRADOV G.M. (1993) Zool. Zh. 72(2): 40-53 [in Russian].
D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 405
Arthropoda, Crustacea, Amphipoda, Gammaridea, Stenothoidae
1: Habitus; from BARNARD (1966).
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Stenothoe menezgweni BELLAN-SANTINI, 2005
Size: 5 mm in adult female.
Morphology: (Family) Coxa 1 very small and partially coveredby following coxae, coxa 4 enlarged, shield-like, not posteri-odorsally excavate. Uropod 3 uniramous. Telson entire.(Genus) Antenna 1 lacking nasiform process on article 1, ac-cessory flagellum absent or 1-articulated. Gnathopods 1-2 sub-chelate, very different from each other in size and shape.Gnathopod 1 small. Gnathopod 2 large. Pereopod 5 with rec-tolinear basis. Pereopods 6-7 with expanded and lobate basis.
Telson ordinary, flat. (Species) Antenna 1 with accessory fla-gellum reduced at a small scale. Eyes present. Gnathopod 1small. Gnathopod 2 greatly enlarged, with the propodus longand narrow, palm excavate at the distal part dactylus as long ashalf propodus. Pereopods 6-7 slightly lobate. Telson entire.
Biology: Collected on hydrothermal site
Distribution: Mid-Atlantic Ridge: SW of Azores, MenezGwen.
Reference:
BELLAN-SANTINI D. (2005) J. Nat. Hist. 39(15): 1101-1110.
D. BELLAN-SANTINI Denisia 18 (2006): 406
1A: Habitus; B: Gnathopod 1; C: Gnathopod 2; D: Epimeral plates; E: Uropod 1; F: Uropod 2; G: Uropods 3; H: Telson; from BEL-LAN-SANTINI (2005).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Stenothoidae
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Torometopa saldanhae BELLAN-SANTINI, 2005
Size: 4 mm in adult male.
Morphology: (Family) Coxa 1 very small and partially coveredby following coxae, coxa 4 enlarged, shield-like, not posteri-odorsally excavate. Uropod 3 uniramous. Telson entire.(Genus) Antenna 1 lacking nasiform process on article 1, ac-cessory flagellum zero- to two-articulated. Gnathopods 1-2 dif-ferent from each other in size and shape. Gnathopod 1 small,almost simple or weakly subchelate. Gnathopod 2 large. Pereo-pod 5, basis not lobate in the middle of the article but with aposter-distal lobe expanded to the end of ischium. Pereopods 6-7 with expanded and lobate article 2. Telson ordinary, flat.(Species) Antenna 1 without accessory flagellum. Eyes presentof moderate size. Gnathopod 1 subchelate, feeble, propoduswith a short and oblique palm, defined by a spine. Gnathopod2 large, propodus distally expanded with palm transverse,
strongly indented, limited at the distal corner by a tooth, pos-terior margin fringed with six small spines, dactylus as long asthe palm, margin smooth. Pereopods 6-7 with expanded and lo-bate basis, more expanded in pereopod 7. Telson entire, longerthan wide, distal triangular.
Biology: Collected a few meters away from the fluid emissionamong sponges or dead mussels, and out of the active hy-drothermal areas amongst gorgonians. This species is presentwithin hydrothermal sites and in the bathyal environment, canbe regarded as an opportunist amongst hydrothermal communi-ties.
Distribution: Mid-Atlantic Ridge: Menez Gwen, Lucky Strike,Rainbow.
Reference:
BELLAN-SANTINI D. (2005) J. Nat. Hist. 39(15): 1101-1110.
D. BELLAN-SANTINI Denisia 18 (2006): 407
1A: Habitus; B: Gnathopod 1; C: Gnathopod 2; D: Uropod 1; E: Uropod 2; F: Uropod 3; G: Telson; from BELLAN-SANTINI (2005).
Arthropoda, Crustacea, Amphipoda, Gammaridea, Stenothoidae
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Ventiella sulfuris BARNARD & INGRAM, 1990
Size: 6.4 mm in adult female.
Morphology: (Family) Body compact. Peduncle of antenna 1short and stout, articles 2-3 much shorter than 1 and partlytelescoped. Article 3 of gnathopod 2 elongate. (Genus) Mouth-parts forming quadrate bundle, labrum dominant, molar tritu-rative. Eyes absent. Coxa 1 strongly shortened partly coveredby coxa 2. Gnathopod 1 short, poorly subchelate. Inner ramusof uropod 2 without notch. Telson short weakly cleft. (Species)Monotypic.
Biology: Collected from worms and clams washings as well ascrab trap wash. Very abundant. Ventiella sulfuris is the most fre-quent species in Eastern Pacific and represents up to 98% of thecatches.
Distribution: Galapagos Spreading Center; East Pacific Rise.
References:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contr. Zool. 499: 1-80. VINOGRADOV G.M. 1993. Zool. Zh. 72(2): 40-53 [in Russian].
D. BELLAN-SANTINI Denisia 18 (2006): 408
1A: Habitus; B: Gnathopod 1; C: Uropod 3; D: Telson; from BARNARD & INGRAM (1990).
Arthropoda, Crustacea, Amphipoda, Lysianassoidea, Uristidae
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Thysanoessa parva HANSEN, 1905
Size: Max. length 9-10.5 mm.
Morphology: Rostrum acute, ending beyond the eyes. Eyeslarge, with a transverse constriction between two lobes, the up-per one being smaller than the lower. Carapace with a lateralsmall denticle on its inferior margin, posterior to its mid-point.Thoracopods almost uniform in structure, although the secondpair is elongated; their setae are provided with setules visibleonly at high magnifications. The eighth pair is rudimentary.The seventh pair has well developed exopodites; endopoditesare absent in males and short in females. Sixth abdominal seg-ment shorter than the sum of lengths of the two preceding seg-ments. Photophores are present on the eye stalks, at the basesof the thoracopods 3 and 7 and on the mid ventral part of ab-dominal segments 1-4.
Biology: It is interesting to note the great abundance of speci-mens in almost all samples from the hydrothermal vent fieldRainbow (Mid-Atlantic Ridge). Indeed, a total of 700 speci-mens were caught in the different traps (KHRIPOUNOFF et al.2001). They were regularly present from September to Januaryand more markedly in October and November at different dis-tances from the vents. They were always absent in traps fromFebruary to June, except for traps laid far off vent sites (2 km).
Curiously, all the specimens were juveniles. Chemical analysisof the composition of particles from the stomach content hasbeen done by EDS X-ray microanalysis. A comparison betweenT. parva specimens and specimens of the related species T. gre-garia indicate that probably the T. parva population is more orless linked with the particle flux. Indeed, a peak of the Fe ele-ment is obvious and might be related with the high concentra-tion of iron (7%) in the vent particles (KHRIPOUNOFF et al.2001) that does not exist in T. gregaria (unpublished data).However, studies of T. parva specimens caught far from thevents have yet to verify this hypothesis. Curiously, there wereno euphausiaceans in traps laid in the South Atlantic (Gulf ofGuinea) and in the Pacific (East Pacific Rise: 13°N) (M.Segonzac, pers. comm.).
Distribution: It is a rare meso-bathypelagic species. In the At-lantic, it has been recorded off Portugal and Africa, from thelatitude of Gibraltar to near Cap Town; on the western side, itis mentioned near the Bermuda Islands. In the Pacific, it isknown in scattered areas: east of south Japan, off California andeast of New Zealand. In the Indian Ocean, it exists west ofSumatra. Mid-Atlantic Ridge: Rainbow.
References:
BAKER A. DE C., BODEN B.P. & E. BRINTON (1990) A Practical Guide of the Euphausiids of the World. Nat. Hist. Mus. Publ., London: 1-96.BRINTON E. (1962) Bull. Scripps Inst. Oceanogr. 8: 51-270.KHRIPOUNOFF A., VANGRIESHEIM A., CRASSOUS P., SEGONZAC M., COLACO A., DESBRUYÈRES D. & R. BARTHELEMY (2001) J. Mar. Res. 59: 633-656.MAUCHLINE J. & L.R. FISHER (1969) Adv. Mar. Biol. 7: 1-454.
R. BARTHELEMY & B. CASANOVA Denisia 18 (2006): 409
Arthropoda, Crustacea, Euphausiacea, Euphausiidae
1: Adult male; from BAKER et al. (1990).
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Decapoda, Caridea
MARTIN & HANEY (2005) listed 52 species of nine familiesof caridean shrimp from the vicinity of hydrothermal vents andcold seeps, although several of them have not been identifiedto species level. Of them species of the family Alvinocarididaeare endemic to such reduced environments, whereas the occur-rence of most of other families is considered to be rather op-portunistic. It is highly likely that pelagic or demersal species(particularly species of Oplophoridae) will be further recordedfrom waters around vent sites by future studies. Periclimenesthermohydrophilus of the Palaemonidae is associated with the tu-beworm Lamellibrachia satsuma, and such commensalism is rat-her unique among the shrimp species known from the chemo-synthetic communities.
This book may give an impression that inventoryof the caridean fauna of the hydrothermal vents or cold seeps isnearly completed. Indeed, in particular, thanks to the recentstudies, progress on the taxonomy of the Alvinocarididae hasbeen made, but there are still some species to be described(MARTIN & HANEY, 2005). For example, in spite of the revisionby KOMAI & SEGONZAC (2005) and subsequent work by KOMAI
et al. (2005), the taxonomic identity of some species of Alvino-caris remain unclear. There is a tendency that oceanographicresearchers publish lists of vent or seep communities based onidentification made in a rather preliminary manner and thatthe voucher material is stored at universities or personal free-zers or other non-recognized repositories. This makes access tothe specimens, as well as accurate assessment of specific identi-ties, difficult. It is advisable to send specimens for accurateidentifications to relevant specialists and to deposit voucherspecimens in museums or other appropriate institutions.
Below, some important points are given:
1. Juveniles of congeneric species of Alvinocarididae arevery similar in morphology and sometimes it is impossible toidentify them without locality data. In case of the occurrence ofmore than one congeneric species sympatrically, identificationshould only be done to genus level.
2. Adults of alvinocaridids often exhibit a high degree ofpolymorphism in the morphology of the chela and ambulatorylegs. Therefore careful observations of various characters is ne-cessary for accurate identification.
3. Some species of alvinocaridids, for example, species of Ri-micaris, shows dramatic ontogenetic change after postlarval sta-ges. Therefore, it is strongly recommended that a formal des-cription of new species is based on adult specimen(s).
4. For any work including results of identification of spe-cies, it is recommended that voucher specimens are depositedin a recognized museum collection. For morphological studies,specimens preserved in 70–75% ethanol is recommended, alt-hough formalin fixed specimens are also useful. However, 70–75% ethanol or formalin may be not adequate for DNA extrac-tion. Therefore, it is advisable to preserve specimens in two dif-ferent concentration of ethanol according to further treatment.
5. Color in life is sometimes very useful in discriminatingcaridean species, particularly hippolytids. It is advisable to takecolor photographs just after capture of specimens.
6. There is little doubt that new species are likely to be dis-covered by new expeditions even at “well-known“ or “well-col-lected“ sites.
References:
KOMAI T. & M. SEGONZAC (2005) Journal of Natural History 39(15): 1111-1175.KOMAI T. SHANK T.M. & C.L. VAN DOVER (2005) Zootaxa 1019: 27-42.MARTIN J.W. & T. HANEY (2005) Zool. J. Linn. Soc. 145: 445-522.
T. KOMAI Denisia 18 (2006): 410
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Nematocarcinus burukovskyi KOMAI & SEGONZAC 2005
Size: Carapace length 25.2 mm, total length ca. 74 mm.
Color: In life, body light orange, yellowish hepatopancreas vis-ible through integument. Cornea of eye brown, but reflective;eye-stalk dark orange. Pereopods nearly colorless.
Morphology: Body surface smooth. Rostrum slender, slightlyfalling short of anterior margin of antennal scale, about 0.6 ofcarapace length; dorsal margin armed with three fixed teeth;ventral margin with four fixed teeth. Carapace relatively wellsculptured with distinct postorbital, cervical, post-cervical, he-patic, branchiocardiac grooves; postrostral median ridge withsix teeth clearly separated from three teeth on rostrum proper,at least some of them with basal suture. Abdomen dorsallysmooth; fifth somite with one or two acute posteroventralteeth; inner pleural ridge on fifth somite low; sixth somite withtwo rows of setae on ventral surface, but devoid of paired pos-teroventral spots. Telson with six pairs of small dorsolateralspines; all dorsolateral spines aligned; terminal margin withthree pairs of spines, second pair longest. Eye with well devel-oped, distinctly faceted cornea. Second pereopod longest
among pereopod. Third to fifth pereopods markedly elongate,but still relatively short and stout for nematocarcinids; propodivery short, subcylindrical. Exopods on third and fourth pere-opods very short, particularly that on fourth pereopod rudi-mentary.
Remarks: This species can be confused in situ with Nemato-carcinus ovalis, whose a female ovigerous carapace length 27.5mm, collected a few tens of meters above the hydrothermalsites East Pacific Rise: 13°N, 2600 m, presents a similar color.This species is observed on the active sites, but also in theabyssal milieu at several hundred meters of sites.
Biology: Live around the active chimneys diffusing milky fluidat low temperature between 2 and 7°C, above beds of themytilid Bathymodiolus spp., close to siboglinid tubeworm Riftiapachyptila, crabs Bythograea spp., and galatheid crabs Munidopsisspp. Necrophagous feeder.
Distribution: Southern East Pacific Rise: 17°S, 23°S and 31°S.
1, 3-5: Holotype, subadult male carapace length 18.6 mm, from Wormwood vent site, Southern East Pacific Rise: 17°S © J.-F. Dejouannet/IRD.
2: Individual in situ, taken at Fred’s Fortressvent site, Southern East Pacific Rise: 31°S; cruisePAR 5; on mussel bed of Bathymodiolus sp.,with bythograeid crabs (Bythograeavrijenhoeki); by R. Vrijenhoek © MBARI.
Arthropoda, Crustacea, Decapoda, Caridea, Nematocarcinidae
1: Habitus; scale bar 1 cm.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 411-412
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Reference:
KOMAI T. & M. SEGONZAC (2005) Zoosystema 27(2): 343-364.
3: Rostrum, anterior part of carapace and cephalic appendages, lateral view;scale bar 5 mm.
4: Posterior part of third abdominal somite to telson and left uropod, lateralview; scale bar 5 mm.
6: Holotype taken on board, just after collection; cruiseBiospeedo; P. Briand © Ifremer.
7: Paratype juvenile carapace length 13.8mm, rostrum, anterior part of carapace andcephalic appendages, taken on board, justafter collection; Southern East Pacific Rise:31°S; cruise PAR 5; G. Rouse © Australia.
5: Telson, dorsal view; scalebar 2 mm.
412
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Alvinocaris brevitelsonis KIKUCHI & HASHIMOTO, 2000
Size: Only the ovigerous female holotype is known: carapacelength 13.8 mm, total length ca. 57 mm.
Color: Ivory to dull ivory white.
Morphology: Rostrum directed forward, slightly curved dorsal-ly, 0.55% of carapace length, reaching distal margin of secondsegment of antennular peduncle; dorsal margin armed with 13teeth, including eight teeth on rostrum proper and five rela-tively large teeth on carapace posterior to orbital margin, pos-teriormost tooth arising from 0.38% of carapace length; ventralmargin armed with seven small teeth on anterior 0.60%. Cara-pace 0.7 times as wide as long; postrostral ridge relatively high,dorsal angle about 150°; branchial region not notably inflated,slightly convex; pterygostomian tooth relatively weak. Fourthabdominal pleuron with three tiny teeth on posteroventralmargin. Fifth abdominal pleuron with strong posteroventraltooth and one additional tooth on posterior margin. Telsonfalling short of posterior margin of uropodal endopod, armedwith six or seven dorsolateral spines; posterior margin moder-ately convex, armed with six pairs of spines (second pair longestand third pair shortest) and two pairs of plumose setae subequalin length to longest second spine. Eye with small spiniform tu-
bercle on anterior surface. Antennular peduncle with secondsegment 1.8 times longer than wide. Antennal scale half lengthof carapace, 1.87 times longer than wide. Third to fifth pere-opods moderately slender; dactyli each with single row of ac-cessory spinules; meri armed with spines on ventrolateral sur-faces; ischia also with spines at least in third and fourth pere-opods. No epipods on third maxilliped to fourth pereopod. Sec-ond to fourth pereopods each with slender, cincinnulate ap-pendix interna.
Remarks: A. brevitelsonis is represented only by the holotype.The possession of more than two pairs of spines on the posteri-or margin of the telson links A. brevitelsonis to A. lusca and A.stactophila. As KIKUCHI & HASHIMOTO (2000) correctly point-ed out, the shorter telson, which falls short of the posterior endof the uropod, distinguishes A. brevitelsonis from A. lusca. A.dissimilis and Shinkaicaris leurokolos occurred sympatrically.
Biology: This species occurs at a hydrothermally influencedarea.
Distribution: Okinawa Trough: Minami-Ensei Knoll.
References:
KIKUCHI T. & J. HASHIMOTO (2000) Species Diversity 5: 135-148.KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.
T. KOMAI & M. SEGONZAC Denisia 18 (2006) 413
2: Anterior part of carapaceand cephalic appendages,dorsal.
3: Telson andleft uropod,dorsal.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Body, lateral.
1-3: Holotypeovigerous female(carapace length13.8 mm) from Mi-nami Ensei Knoll,Mid-OkinawaTrough;scale bars 1 mm; by T. Kikuchi.
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Alvinocaris dissimilis KOMAI & SEGONZAC, 2005
Size: Largest male carapace length 8.4 mm, largest female cara-pace length 9.8 mm, ovigerous females carapace length 7.3-9.8mm. Maximal total length 41 mm.
Morphology: Rostrum directed forward, straight or very slight-ly curved dorsally, usually reaching to second segment of an-tennular peduncle; dorsal margin armed with 13-17 teeth, in-cluding 8-10 teeth on rostrum proper and 5-8 relatively smallteeth on carapace posterior to orbital margin; ventral marginarmed with one or two small subdistal teeth. Carapace width0.65-0.80 of length; dorsal angle about 155°. Fourth abdominalpleuron with small posteroventral tooth and additional one ortwo teeth on posterior margin. Fifth abdominal somite similar-ly armed with strong posteroventral tooth and additional 1-5tiny teeth on posterior margin or ventral margin. Telson notreaching posterior margin of uropodal endopod; armed with 5-8 dorsolateral spines; posterior margin convex (rarely shallowlynotched medially), armed with two pairs of spines at lateral cor-
ners and 12-14 plumose setae all longer than mesial pair of lat-eral spines.
Remarks: Alvinocaris dissimilis, A. markensis, A. muricola and A.longirostris are morphologically very similar to one another.These four species share the rostrum reaching at least to thesecond segment of the antennular peduncle and armed usuallywith one or more ventral teeth, relatively high postrostral me-dian ridge on the carapace, and the posterior margin of the tel-son with long plumose setae and two lateral pairs of spines.These characters should be used with caution, as most of themare useful only for adult specimens.
Biology: This recently described species occurs in a hydrother-mally active area. Alvinocaris brevitelsonis and Shinkaicaris leu-rokolos occurred sympatrically with this species.
Distribution: Known only from Minami-Ensei Knoll, Mid-Ok-inawa Trough, 705 m.
References:
KIKUCHI T. & J. HASHIMOTO (2000) Species Diversity 5: 135-148.KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 414
1: Carapace and cephalic appendages, lateral.
2: Third to sixth abdominal somites, lateral.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
3: Anterior part of carapace andcephalic appendages, dorsal.
4: Telson and left uropod, dorsal (marginal setae onuropod omitted).
1-4: Holotypeovigerous female(carapace length11.9 mm) fromMinami-EnseiKnoll, Mid-Okinawa Trough; by T. Komai.
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Alvinocaris longirostris KIKUCHI & OHTA, 1995
Size: Largest female 16.9 mm. Maximal total length ca. 85 mm.
Color: Ivory to dull ivory white.
Morphology: Rostrum slightly descending or directed forward,straight or weakly curved dorsally, 0.64-1.26 of carapace lengthin females, usually overreaching distal margin of antennular pe-duncle, armed with 9-15 teeth including 5-10 relatively largeteeth on carapace posterior to orbital margin, posteriormosttooth arising 0.38-0.48 of carapace length; ventral marginarmed with 4-9 small teeth on anterior 0.30-0.50. Carapacewidth 0.65-0.72 of length; postrostral median ridge relativelyhigh, dorsal angle about 150°; branchial region not notably in-flated, slightly convex; pterytostomial tooth relatively strong.Fourth abdominal pleuron with 1-4 posterolateral teeth. Fifthabdominal pleuron similarly armed with strong posteroventraltooth and additional 2-5 small teeth. Telson not reaching toposterior margin of uropodal endopod, distinctly narrowed pos-teriorly, armed with 5-7 dorsolateral spines; posterior marginconvex, armed with two pairs of spines at lateral angle and 12-
14 plumose setae all longer than mesial pair of lateral spines.Eye with small spiniform tubercle on anterior surface. Anten-nular peduncle with second segment 1.58-1.69 times longerthan wide. Antennal scale about half lengh of carapace and1.72-1.90 times longer than wide. Third to fifth pereopodsmoderately slender; dactyli each with single row of accessoryspinules; meri armed with spines on ventrolateral surfaces; is-chia also with spines at least in third and fourth pereopods. Noepipods on third maxilliped to fourth pereopod. Second tofourth pereopods each with slender, cincinnulate appendix in-terna.
Biology: Usually associated with hydrothermal vents. FUJIKURA
ET AL. (1995) reported the occurrence of A. longirostris at coldseeps on off Hatsushima site, Sagami Bay.
Distribution: Okinawa Trough: Iheya Ridge and HatomaKnoll; Japan: Sagami Bay, off Hatsushima site; Kermadec-Arc:Brothers Seamount Caldera.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Adult female, lateral.
3: Anterior part ofcarapace and
cephalic ap-pendages, dorsal.
2: Posterior part of abdomen and uropods in ven-tral view.
4: Telson anduropod, dorsal.
1-4: from KIKUCHI &OTHA (1995).
References:
FUJIKURA K., HASHIMOTO J., FUJIWARA Y. & T. OKUTANI (1995) JAMSTEC J. Deep-Sea Res. 11: 227-241.KIKUCHI T. & S. OHTA (1995) J. Crustac. Biol. 15: 219-240.WATABE H. & H. MIYAKE. (2000) JAMSTEC J. Deep-Sea Res. 17: 29-34.WEBBER W.R. (2004) Zootaxa 444: 1-26.
T. KOMAI Denisia 18 (2006): 415
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Alvinocaris lusca WILLIAMS & CHACE, 1982
Size: Carapace length largest male 7.3 mm; largest female 13.5mm. Maximal total length ca. 85 mm.
Morphology: Rostrum slightly descending or directed forward,straight, 0.37-0.44 times of carapace length, usually reachingsecond segment of antennular peduncle, armed with 10-14teeth including 4-6 moderately large teeth on carapace posteri-or to orbital margin, posterior tooth arising 0.32-0.39 of cara-pace length; ventral margin armed with three or four smallteeth on anterior 0.20-0.40. Carapace width 0.55-0.67 oflength; postrostral median ridge relatively low, dorsal angleabout 170°; branchial region weakly convex; pterytostomialtooth relatively weak. Fourth abdominal pleuron with 1-5(most frequently four) posterolateral teeth. Fifth abdominalpleuron similarly armed with strong posteroventral tooth andadditional 1-5 small teeth. Telson not reaching to slightly over-reaching posterior margin of uropodal endopod, very slightlynarrowed posteriorly, armed with 7-9 dorsolateral spines; poste-rior margin shallowly notched medially or slightly convex,armed with 5-7 spines (mesial 3-5 spines unequal in length, but
third spine from lateral shorter than others) and one or twoshort plumose setae on either side of median notch. Eyes withsmall spiniform tubercle on anterior surface. Antennular pe-duncle with second segment 2.00-2.20 times longer than wide.Antennal scale about half lengh of carapace and 2.20-2.60times longer than wide. Third to fifth pereopods moderatelyslender; dactyli each with single row of accessory spinules; meriarmed with spines on ventrolateral surfaces; ischia also withspines at least in third and fourth pereopods. No epipods onthird maxilliped to fourth pereopod. Second to fourth pere-opods each with slender, cincinnulate appendix interna.
Biology: This species is associated with the vestimentiferanworm Riftia pachyptila, and could be nourished from its biologi-cal production. VAN DOVER et al. (1985) described plank-totrophic larval develoment.
Distribution: Known with certainty only from the GalapagosSpreading Center: Rose Garden area and East Pacific Rise:9°N.
References:
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.VAN DOVER C.L., FACTOR J.R., WILLIAMS A.B. & C.J. BERG (1995) Bull. Biol. Soc. Wash. 6: 223-227.WILLIAMS A.B. & F.A. CHACE (1982) J. Crustac. Biol. 2(1): 136-147.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 416
1: Body, lateral; enlargement x5.
3: Telson anduropods, dorsal;
enlargement x10.1-3: from WILLIAMS
& CHACE (1982).
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
2: Anterior part of carapaceand cephalic appendages; en-largement x10.
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Alvinocaris markensis WILLIAMS, 1988
2: Anterior part of carapace andcephalic appendages, dorsal; scale bar 2 mm.
3: Telson and right uropod, dorsal; scalebar 5 mm.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Female (carapace length 16.7 mm) from site Les Ruches, Snake Pit, MAR; carapaceand cephalic appendages, lateral; scale bar 5 mm.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 417–418
Size: Carapace length largest male 12.9 mm; largest female 22.3mm, ovigerous female 13.0 mm. Maximal total length ca. 82mm.
Color: In situ red-orange, abdomen paler.
Morphology: Rostrum directed forward or slightly descending,nearly straight or slightly curved dorsally, armed with 14-19teeth, including 8-12 on rostrum proper and 5-7 moderatelylarge teeth on carapace posterior to level of orbital margin, pos-teriormost tooth arising from 0.24-0.31 of carapace length; ven-tral margin armed with 6-9 small teeth. Carapace 0.52-0.65times as wide as long; postrostral median ridge relatively high,dorsal angle about 150°; pterygostomial tooth relatively strong;branchial region not particularly inflated, slightly convex.Fourth abdominal pleuron with 1-4 (most frequently two or 3)teeth posteriorly. Fifth abdominal pleuron similarly armed withone strong posterolateral tooth and 1-3 additional teeth on pos-terior margin. Telson narrowed posteriorly, not reaching orreaching posterior margin of uropodal endopod, armed with 6-8 dorsolateral spines; posterior margin always convex, armedwith two pairs of lateral spines and 12-14 plumose setae all
longer than mesial pair of lateral spines. Antennular pedunclemoderately stout, second segment 1.79-2.05 times longer thanwide. Eye with small spiniform tubercle on anterior surface.Antennal scale 1.91-2.04 times longer than wide. Third to fifthpereopods moderately slender; dactyli each with single row ofaccessory spinules; meri armed with spines on ventrolateral sur-faces; ischia also with spines at least in third and fourth pere-opods. No epipods on third maxilliped to fourth pereopod. Sec-ond to fourth pereopods each with slender, cincinnulate ap-pendix interna.
Biology: This species always lives more or less solitary at thebase and on walls of active chimneys, close to aggregates ofRimicaris exoculata, or on mussel beds. The trophic mode isnecrophagous, as captures of the shrimps by baited traps indi-cate. However, the gut contents examined generally containedmainly mineral particles. Ovigerous females collected in Au-gust.
Distribution: The occurrence of this species is confirmed atseveral locations at Mid-Atlantic Ridge: TAG, Snake Pit, Bro-ken Spur, Logatchev, Rainbow, and Lucky Strike.
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References:
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.SEGONZAC M., SAINT LAURENT M. DE & B. CASANOVA (1993) Cah. Biol. Mar. 34: 535-571.SHANK T., BLACK M.B., HALANYCH K.M., LUTZ R.A. & R.C. VRIJENHOEK (1999) Mol. Phylogenet. Evol. 13: 244-255.VAN DOVER C.L. (1995) in PARSON L., WALKER C.L. & D.R. DIXON (Eds) Hydrothermal Vents and Processes: 257-294.WILLIAMS A.B. (1988) J. Crustac. Biol. 2(1): 136-147.WILLIAMS A.B. (1988) Fish. Bull. 86(2): 263-287.
4: Fresh specimen taken on board; from Briand © Ifremer.
5: In situ at the base of hot vent on the site Les Ruches, Snake Pit. The shrimp is seen in pro-file; one notes the camber cephalothorax/abdomen, characteristic of this species. Surround-ed by Chorocaris chacei shrimps. Cruise Hydrosnake © Ifremer. 1-5: by T. Komai (from KOMAI
& SEGONZAC 2005).
418
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Alvinocaris niwa WEBBER, 2004
Size: Male holotype carapace length 15.38 mm, total length ca.57 mm.
Morphology: Rostrum short, not reaching distal margin of firstsegment of antennular peduncle, directed forward, weakly com-pressed laterally, terminating acutely, dorsal margin sharplycarinae, armed with 5-17 teeth including 4-12 on rostrum prop-er and 0-6 on anterior part of carapace posterior to level of or-bital margin; ventral margin usually unarmed or rarely with onetiny subterminal tooth. Carapace somewhat compressed later-ally, with sharp postrostral ridge reaching anterior 0.2 of cara-pace length; antennal spine acuminate; no conspicuous lobemesial to antennal spine; pterygostomial angle weakly to some-what produced in adults, reaching or distinctly overreachingantennal spine, terminating in sharp spine. Abdomen smoothdorsally; pleuron of third somite usually smooth, those of fourthand fifth somites at least with posterolateral tooth and fre-quently with additional small teeth ventrally and/or posterior-ly. Telson with six or seven dorsolateral spines arranged in sin-uous row on either side; posterior margin convex, with 1-3small spines at each lateral angle and row of numerous longplumose setae. Eyestalks degenerated, broadly fused mesially,cornea unfacetted; anterior surface unarmed. Chela of first
pereopod without fine row of long submarginal setae on outersurface along cutting edges of fingers. Third to fifth pereopodsmoderately slender; dactyli armed with two or more rows of ac-cessory spinules on ventral surface; meri usually unarmed; is-chia of third and fourth pereopods armed usually with two lat-eral spines. No strap-like epipods on third maxilliped and pere-opods. Appendices internae on second to fourth pereopodsslender, without coupling hooks.
Remarks: The reduced dorsal rostral series, the reduced arma-ture of the meri of the third to fifth pereopods and the posses-sion of two or more rows of accessory spinules on the ventralsurface of each dactylus of the third to fifth pereopods set thisspecies apart from Alvinocaris, and link it to a group composedof Shinkaicaris, Opaepele, Chorocaris and Rimicaris.
Biology: This species is associated with hydrothermal springson active submarine volcanoes, among mytilid bivalve Giganti-das gladius, but also abundant bathyal fauna (coral, barnacles,asteroid echinoderms).
Distribution: Kermadec-Arc: Brothers Seamount Caldera andRumble V Seamount.
3: Front, dorsal view.
2: Abdomen, ventrolateralview; scale bar 5 mm.
4: Telson and uropods, dorsal view. 1-4: Holotype, carapacelength 15.38 mm; from WEBBER (2004).
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Lateral view; scale bar 5 mm.
References:
WEBBER W.R. (2004) Zootaxa 444: 1-26.WEBBER W.R. & N.L. BRUCE (2002) Biodiversity Update 5: 6-7.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 419
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Alvinocaris williamsi SHANK & MARTIN, 2003
Size: Largest male carapace length 6.1 mm; largest female 10.3mm, ovigerous female 5.8 mm.
Morphology: Rostrum directed forward, straight or slightlydownward, not reaching or reaching distal margin of first seg-ment of antennular peduncle, 0.27-0.37 of carapace length;dorsal margin armed with 10-15 teeth, including 5-9 teeth onrostrum proper and 4-8 small to moderately large teeth on cara-pace posterior to orbital margin, posteriormost tooth arisingfrom 0.13-0.20 of carapace length; ventral margin usually un-armed. Carapace about 0.70 times as wide as long, postrostralmedian ridge relatively low, with dorsal angle about 150°;branchial region not notably inflated, slightly convex; bran-chiostegal tooth moderately strong. Fourth abdominal pleuronrounded or with small posteroventral tooth. Fifth abdominalpleuron armed with strong posteroventral tooth and 2-4 addi-tional tiny teeth on posterior margin. Telson not reaching pos-terior margin of uropodal endopod; armed with six or sevendorsolateral spines arranged in straight or slightly convex line;
posterior margin broadly rounded, armed with two pairs ofspines at posterolateral angles and 14-16 plumose setae alllonger than mesial pair of lateral spines. Eye with small tuber-cle on anterior surface. Antennular peduncle very stout, withsecond segment 1.05-1.11 times longer than wide. Antennalscale about 0.4 of carapace length, 1.40-1.50 times longer thanwide. Third to fifth pereopods moderately slender; dactyli eachwith single row of accessory spinules; meri usually armed withspines on ventrolateral surfaces in third and fourth; ischia alsowith spines at least in third and fourth pereopods. No epipodson third maxilliped to fourth pereopod. Second to fourth pere-opods each with slender, cincinnulate appendix interna.
Biology: The specimens were collected from the mussel beds ofthe mytilid Bathymodiolus azoricus.
Distribution: Known only from Mid-Atlantic Ridge: MenezGwen.
References:
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.SHANK T.M. & J.W. MARTIN (2003) Proc. Biol. Soc. Wash. 116: 158-167.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 420
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Body, lateral.
2: Anterior part ofcarapace and cephalicappendages, dorsal.
3: Telson and lefturopod, dorsal. 1-3: Holotype,ovigerous female;scale bar 1 mm;from SHANK & MAR-TIN (2003).
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Chorocaris chacei (WILLIAMS & RONA, 1986)
Size: Female, max. carapace length 17.4 mm; total length 55mm.
Color: Cephalothorax red orange, abdomen paler.
Morphology: Rostrum broadly rounded, flattened dorsoventral-ly, reaching or overreaching antennal spines; no dorsal or ven-tral teeth; ventral surface convex. Carapace somewhat com-pressed laterally, rounded dorsally; antennal spine blunt; ptery-gostomial angle strongly produced, terminating bluntly. Fourthabdominal pleuron blunt angular posteroventrally; fifth pleu-ron with subacutely pointed posterolateral angle. Telson nar-rowed posteriorly, bearing row of 6-8 dorsolateral spinesarranged in sinuous row on either side; posterior margin broad-ly convex, with 1-3 spines at each lateral angle and with row ofnumerous plumose setae. Eye-stalks degenerated, broadly fusedmesially without trace of median separation; no distinct spineor tubercle on anterior surface. Chela of first pereopod withoutfine row of long submarginal setae on outer surface along cut-ting edges of fingers. Third to fifth pereopods moderately slen-der to stout; dactyli armed with two or more rows of accessory
spinules on ventral surface; meri and ischia unarmed. Thirdmaxilliped and first to fourth pereopods without strap-likeepipods. Appendices internae on second to fourth pereopodssimple, slender, without cincinnuli. Protopod of endopod ta-pering distally, terminating subcautely or acutely.
Remarks: A single female specimen collected at the site EiffelTower on Lucky Strike differs from typical series of C. chacei inhaving an acuminate rostrum and acuminate antennal andpterygostomial spines on the carapace. We regard that the spec-imen is an abnormal individual of C. chacei, because it was col-lected along with individuals of typical C. chacei and we havenot encountered such a form in other collections from theLucky Strike.
Biology: This species is much less abundant (2-3 ind./dm2, byplaces) than Rimicaris exoculata. It is present close to chimneysand on sulfide blocks, mostly where fluid emissions are weak.
Distribution: Mid-Atlantic Ridge: TAG, Snake Pit, Logatchev,Rainbow, and Lucky Strike.
2: Specimen taken on board; cruise Exomar © Ifremer.
4: Anterior part of carapaceand cephalic appendages, dor-sal; cruise Exomar © Ifremer.
5: Telson and uropods,dorsal; cruise Exomar © Ifremer.
3: Cephalothoraxof a single aber-rant specimencollected at LuckyStrike in July2001 (Americancruise DiversExpe-dition) by P.Briand © Ifremer.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Holotype, lateral view; enlargement x5.2; from WILLIAMS & RONA (1986).
References:
RAMIREZ LLODRA E.,TYLER P.A. & J.T.P. COPLEY (2000) J. Mar. Biol. Ass. U.K. 80: 473-484.SEGONZAC M., SAINT LAURENT M. DE & B. CASANOVA (1993) Cah. Biol. Mar. 39(15): 1111-1175.WILLIAMS A.B. & P.A. RONA (1986) J. Crustac. Biol. 6: 446-462.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 421
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Chorocaris paulexa MARTIN & SHANK, 2005
Size: Female, carapace length 9.6 mm
Color: Live shrimp mostly pale translucent white to slightlypink on posterior and dorsal surfaces of carapace and abdomen.
Morphology: Rostrum broadly triangular, flattened dorsoven-trally, overreaching antennal spines; no dorsal or ventral teeth;ventral surface convex. Carapace somewhat compressed later-ally, rounded dorsally; spawning female with very shallow de-pression on either side of midline in anterior half; antennalspine blunt; pterygostomial angle strongly produced as relative-ly narrow triangular projection, terminating subacutely. Fourthabdominal pleuron bluntly angular posteroventrally; fifth pleu-ron with subacutely pointed posterolateral angle. Telson slight-ly narrowed posteriorly, bearing row of 6-8 dorsolateral spinesarranged in sinuous row on either side; posterior margin broad-ly convex, with 1-3 spines at each lateral angle and with row of
numerous plumose setae. Eye-stalks degenerated, broadly fusedmesially without trace of median separation; no distinct spineor tubercle on anterior surface. Chela of first pereopod withoutfine row of long submarginal setae on outer surface along cut-ting edges of fingers. Third to fifth pereopods moderately slen-der to stout; dactyli armed with two or more rows of accessoryspinules on ventral surface; meri and ischia unarmed. Thirdmaxilliped and first to fourth pereopods without strap-likeepipods. Appendices internae on second to fourth pereopodssimple, slender, without couppling hooks. Protopod of endopodtapering distally, terminating in blunt tooth.
Biology: Living directly on the black smokers in numbers up to47 individuals per half meter square.
Distribution: Southern East Pacific Rise: 17°S.
Reference:
MARTIN J. & T. SHANK (2005) Proc. Biol. Soc. Wash. 118(1): 183-198.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 422
2: Anterior part of carapace and cephalic appendages, dor-sal; scale bar 5 mm.
3: Telson and left uropod, dorsal;scale bar 2 mm. 1-3: from MARTIN & SHANK (2005).
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Body, lateral; scale bar 10 mm.
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Chorocaris vandoverae (MARTIN & HESSLER, 1990)
Size: Female carapace length 13.4 mm.
Morphology: Rostrum broadly rounded, flattened dorsoventral-ly, overreaching antennal spines; no dorsal or ventral teeth;ventral surface convex. Carapace somewhat compressed later-ally, rounded dorsally; antennal spine blunt; pterygostomial an-gle strongly produced, terminating bluntly. Fourth abdominalpleuron bluntly angular posteroventrally; fifth pleuron withsubacutely pointed posterolateral angle. Telson slightly nar-rowed posteriorly, bearing row of 6-8 dorsolateral spinesarranged in sinuous row on either side; posterior margin broad-ly convex, with 1-3 spines at each lateral angle and with row ofnumerous plumose setae. Eye-stalks degenerated, broadly fusedmesially without trace of median separation; no distinct spine
or tubercle on anterior surface. Chela of first pereopod withoutfine row of long submarginal setae on outer surface along cut-ting edges of fingers. Third to fifth pereopods moderately slen-der to stout; dactyli armed with two or more rows of accessoryspinules on ventral surface; meri and ischia unarmed. Thirdmaxilliped and first to fourth pereopods without strap-likeepipods. Appendices internae on second to fourth pereopodssimple, slender, without cincinnuli. Protopod of endopodslightly tapering and rounded distally.
Biology: Living in profusion at vents or adjacent rocks.
Distribution: Mariana, Manus, North Fiji and Lau Back-ArcBasins.
References:
MARTIN J.W. & R.R. HESSLER (1990) Nat. Hist. Mus. Los Angeles Cty 417: 1-11.Williams A.B. & P.A. Rona (1986) J. Crustac. Biol. 6: 446-462.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 423
2: Dorsal view of carapace and basesof antennae; by J. Martin.
3: Telson and uropods; by J. Martin.
1: Habitus, lateral view; by J. Martin.
4: In situ view of two specimens among gastropods(Alviniconcha hessleri); Mariana Back-Arc Basin; bycourtesy of R.R. Hessler.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
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Mirocaris fortunata (MARTIN & CHRISTIANSEN, 1995)
Size: Females carapace length 3.3-8.7 mm, males carapacelength 3.8-7.2 mm.
Morphology: Rostrum triangular, terminating bluntly or suba-cutely in dorsal view, flattened dorsoventrally, reaching toslightly overreaching antennal spine; no dorsal or ventral teeth.Carapace somewhat compressed laterally, with short transverse(vertical) rows of minute to short setae on lateral parts; dorsalsurface rounded in males and non-ovigerous females, broadlycarinate in ovigerous females, general outline in lateral viewfaintly sinuous to weakly convex; in ovigerous females, subme-dian areas very shallowly depressed and ornamented with nu-merous longitudinal striae; antennal spine acute; pterygostomi-al angle weakly produced, terminating subacutely. Fourth ab-dominal pleuron rounded posteriorly; fifth pleuron with suba-cutely pointed posterolateral angle. Telson barely narrowedposteriorly, bearing row of 7-9 dorsolateral spines arranged insinuous row; posterior margin broadly convex, occasionallywith shallow median emargination, bearing 12-19 spines in to-tal; 1-3 spines at posterolateral angle shorter than mesial spines,simple, while remaining mesial spines elongate, bearing minutemarginal setules. Eye-stalks rather large but degenerated, broad-
ly fused mesially without trace of median separation; no distinctspine or tubercle on anterior surface. Chela of first pereopodwith fine row of long submarginal setae on outer surface alongcutting edges of fingers. Third to fifth pereopods moderatelyslender to stout; dactyli armed with single row of accessoryspinules on ventral margin; meri unarmed; ischia with spines inthird, usually unarmed in fourth and fifth. Third maxilliped andfirst to fourth pereopods with strap-like epipods. Appendicesinternae on second to fourth pereopods rudimentary or absent.
Biology: Lives in close proximity to vent openings or in shim-mering water from crevices with diffuse flow, among numerousBathymodiolus bivalves. Numerous ovigerous females in theJune-July samples. This species was found in the enteron ofCandelabrum phrygium (Cnidaria, Hydrozoa) which lives in thesame area.
Distribution: Known from hydrothermal vent sites along theMid-Atlantic Ridge between 38°N and 14°N: Menez Gwen,Lucky Strike, Rainbow, Broken Spur, TAG, Snake Pit, Lo-gatchev.
1: Body, lateral; by J.W. Martin & J.C. Christiansen.
2: Anterior part of carapace andcephalic appendages, dorsal(setae partially omitted; rightantenna removed); scale bar 2 mm; by T. Komai.
3: Telson and left uro-pod, dorsal (setae omit-ted), scale bar 1 mm; by T. Komai.
4: In situ aggregation on crevice of suphide edifice, Lucky Strikesite during Exomar cruise © Ifremer.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
References:
KOMAI T. & M. SEGONZAC (2003) Cah. Biol. Mar. 44: 199-215.MARTIN J.W. & J.C. CHRISTIANSEN (1995) Proc. Biol. Soc. Wash. 108(2): 220-227.SEGONZAC M. & W. VERVOORT (1995) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 17: 31-64.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 424
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Mirocaris indica KOMAI, MARTIN, ZALA, TSUCHIDA & HASHIMOTO, in press
References:
KOMAI. T., MARTIN J.W., ZALA K., TSUCHIDA S. & J. HASHIMOTO (in press) Scientia Marina 70(1).
T. KOMAI Denisia 18 (2006): 425
2: Front, dorsal view.
3: Telson and right uropod,dorsal view (setae omitted).
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
Size: Carapace length 9.2-13.8 mm (females), 4.6-10.2 mm(males).
Color: In life, body and appendages generally white, but dorsalsurface of carapace and anterior two abdominal somites reddish;eye yellowish.
Morphology: Rostrum triangular, terminating bluntly or suba-cutely in dorsal view, flattened dorsoventrally, reaching toslightly overreaching antennal spine; no dorsal or ventral teeth.Carapace somewhat compressed laterally, with short transverse(vertical) rows of minute to short setae on lateral parts; dorsalsurface rounded, general outline in lateral view slightly convex;antennal spine acuminate; pterygostomial angle weakly pro-duced, terminating subacutely. Fourth abdominal pleuronrounded posteriorly; fifth pleuron with subacutely pointed pos-terolateral angle. Telson barely narrowed posteriorly, bearingrow of 7-9 dorsolateral spines arranged in sinuous row; posteri-or margin broadly convex, occasionally with shallow medianemargination, bearing 12-19 spines in total; 1-3 spines at pos-
terolateral angle shorter than mesial spines, simple, while re-maining mesial spines elongate, bearing minute marginal se-tules. Eye-stalks rather large but degenerated, broadly fused; nodistinct spine or tubercle on anterior surface. Chela of firstpereopod without submarginal row of setae on outer surfacealong cutting edges of fingers. Third to fifth pereopods moder-ately slender to stout; dactyli armed with single row of accesso-ry spinules on ventral margin; meri unarmed; ischia with spinesin third, usually unarmed in fourth and fifth. Third maxillipedand first to fourth pereopods with strap-like epipods. Appen-dices internae on second to fourth pereopods rudimentary orabsent.
Biology: The collection sites were an active vent field that in-cluded black smoker complexes with dense patches of vent as-sociated animals, including Rimicaris kairei, Bathymodiolus mus-sel, Alviniconcha snail, Austinograea crab, Marianactis anemone.
Distribution: Central Indian Ridge, Kairei Field, Edmond VentField.
1: Habitus holotype, lateral view; scale bar 5 mm.1-3 from KOMAI et al. (2006) © Scientia Marina.
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Nautilocaris saintlaurentae KOMAI & SEGONZAC, 2005
Size: Largest male carapace length 5.6 mm, largest female cara-pace length 7.8 mm. Maximal total length ca. 35 mm.
Color: Generally whitish; matured female gonads reddish, visi-ble throughout cuticle.
Morphology: Rostrum carinate and dentate dorsally, reachingdistal margin of basal segment of antennular peduncle; ventralsurface unarmed. Carapace somewhat compressed laterally;postrostral median carina low, blunt, restricted to anterior 0.15of carapace; antennal tooth acuminate; pterygostomial angleweakly produced anteriorly, extending as far as antennal spine,terminating in sharp tooth. Third to fifth pleonal pleura den-tate posteroventrally. Telson with 7-9 dorsolateral spinesarranged in slightly sinuous row; posterior margin convex, bear-ing 12-19 spines in total, 1-3 spines at each posterolateral cor-ner shorter than mesial spines, simple, while remaining mesialspines elongate, bearing minute marginal setules. Eyes ratherlarge but degenerate, broadly fused mesially; anterior surfacesmooth; no trace of pigment. Antennal scale broadly oval, with
distinct dorsolateral tooth. Chela of first pereopod with finerow of long submarginal setae on outer surface along cuttingedges of fingers. Third to fifth pereopods moderately slender tostout; each dactylus armed with single row of accessory spinuleson ventral margin; meri unarmed; ischia with spines in third,usually unarmed in fourth and fifth. Third maxilliped to fourthpereopods with strap-like, terminally hooked epipods, corre-sponding to setobranchs above first to fifth pereopods; appen-dices internae on second to fourth pereopods rudimentary.
Biology: This species lives around black smokers where the flu-ids are percolating at temperatures between 5 and 33°C, amongmolluscan beds. The similarity in the mouthparts and chelamorphology suggests that the shrimps ingest free bacteria, mol-lusc pseudo-faeces, or other detritus, as most other species ofthe family. Some ovigerous females were collected in May 2005(American cruise PAR 5).
Distribution: North Fiji and Lau Back-Arc Basins.
References:
DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA (1994) Mar. Geol. 116: 227-242.KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 426
3: Telson, dorsal view(setae omitted); scale bar 1 mm; by T. Komai.
2: Anterior part of carapaceand cephalic appendages,dorsal view; scale bar 0.5 mm;by T. Komai. 4: Several individuals in situ on sulphide wall; cruise TUIM07.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Holotype female (carapace length 7.3mm) from the North Fiji Basin; habitus,lateral view (fourth and fifth pereopodsand second to fourth pleopods missing);scale bar 2 mm; by T. Komai.
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Opaepele loihi WILLIAMS & DOBBS, 1995
Size: Male carapace length 13.6 mm.
Color: When collected, intensely orange, with orange to yel-lowish deposit on abdomen (iron oxyhydroxide particles).
Morphology: Rostrum short, not reaching distal margin of firstsegment of antennular peduncle, directed forward or slightlydownward, somewhat depressed dorsoventrally, terminatingacutely, triangular in dorsal view; dorsal surface bluntly carinaein midline, unarmed or armed with minute denticles in adults,armed with up to six very small teeth in juveniles; ventral sur-face flat or slightly elevated in midline, unarmed. Carapacesomewhat compressd laterally, rounded dorsally; antennal spineacute; no conspicuous lobe mesial to antennal tooth; pterygos-tomial angle somewhat produced in adults, distinctly exceedingantennal tooth, terminating in sharp tooth. Abdomen smoothdorsally; pleura of third and fourth somites smooth or faintlydenticulate posteriorly, posterolateral angle of fourth pleuronrounded or sharply pointed; fifth pleuron with marginal denti-cles posteriorly and sharply pointed posteroventral angle. Tel-son with 6-8 dorsolateral spines arranged in sinuous row on ei-
ther side; posterior margin convex, with 1-3 small spines ateach lateral angle and row of long plumose setae. Eyestalks de-generated, broadly fused mesially, cornea unfacetted, poorly or-ganized retinal pigment evident; anterior surface unarmed.Chela of first pereopod without fine row of long submarginal se-tae on outer surface along cutting edges of fingers. Third to fifthpereopods moderately slender; dactyli armed with two or morerows of accessory spinules on ventral surface; meri and ischiaunarmed. No strap-like epipods on third maxilliped and pere-opods. Appendices internae on second to fourth pereopodsslender, without coupling hooks.
Biology: Lives on the flank of small volcano, among numerousvents which discharge fluids at 37°C (ambiant water, 4°C).The fluid is clear. At Loihi Seamount, no other animal livethere.
Distribution: So far known only from Hawaii, Loihi Seamount.
Reference:
WILLIAMS A.B. & F.C. DOBBS (1995) Proc. Biol. Soc. Wash. 108(2): 228-237.
T. KOMAI Denisia 18 (2006): 427
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Female holotype, lateral view; scale bar 5 mm.
2: Female paratype, anterior re-gion, dorsal view.
3: Female paratype,telson and uropods,
dorsal view. 1-3: from WILLIAMS &
DOBBS (1995).
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Rimicaris exoculata WILLIAMS & RONA, 1986
Size: Carapace length 22.0 mm; total length up to 55 mm.
Morphology: Rostrum strongly reduced to very low, broadlyrounded lobe. Carapace with branchial region very strongly in-flated, distinctly wider than abdomen; dorsal surface rounded;antennal spine modified as broadly rounded lobe; pterygosto-mial angle somewhat produced anteriorly, broadly rounded; sur-face with scattered puntae each bearing tuft of short setae.Third abdominal pleuron with posterolateral margin forminglight angle. Fourth and fifth pleura with posterolateral anglessubacutely or blunly pointed. Telson barely narrowed posterior-ly, bearing row of 6-8 dorsolateral spines arranged in sinuousrow on either side; posterior margin broadly convex, with 1-3spines at each lateral angle and with row of numerous plumosesetae. Eye-stalks greatly reduced and modified into broad ocu-lar plate. Chela of first pereopod always slender, without finerow of long submarginal setae on outer surface along cuttingedges of fingers. Third to fifth pereopods stout, becoming
markedly shorter toward posterior; dactyli armed with severalrows of accessory spinules on ventral surface; propodi of thirdand fourth pereopods with covering of numerous spinules onventral surfaces; meri and ischia unarmed. No strap-likeepipods on third maxilliped to fourth pereopods. Appendicesinternae on second and fourth pereopods simple, slender, with-out coupling hooks, those of fourth and fifth pleopods normal-ly developed, with coupling hooks.
Biology: Shrimp highly mobile occurring in swarms reachingdensities of about 2500 ind./m2 on active chimney walls wheretemperatures range from 10-30°C. Primary consumer, eatingbacteria “cultivated” on the mouthparts, and on the mineralparticles ingested. Very few ovigerous females collected, be-tween August and November.
Distribution: Mid-Atlantic Ridge: TAG, Snake Pit, Rainbow,very few at Lucky Strike.
References:
MARTIN J.W. & R.R. HESSLER (1990) Nat. Hist. Mus. Los Angeles C. 417: 1-11.SEGONZAC M., SAINT LAURENT M. DE & B. CASANOVA (1993) Cah. Biol. Mar. 34: 535-571.WILLIAMS A.B. & P.A. RONA (1986) J. Crustac. Biol. 6: 446-462.ZBINDEN M. & M.-A. CAMBON-BONAVITA (2003) FEMS Microbiol. Ecol. 46: 23-30.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 428
2: In situ aggregation of adults and juveniles or subadults (red cephalothorax)specimens; TAG; cruise Exomar © Ifremer.
1: Specimen taken onboard, dorsal view;cruise Exomar by P. Briand © Ifremer.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
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Rimicaris kairei WATABE & HASHIMOTO, 2002
Size: Up to 21.6 mm in carapace length.
Morphology: Integument completely spineless, almost mem-branous on branchiostegite. Carapace longitudinally oval indorsal view, sparsely pitted by shallow punctuations withoutany setae. Rostrum and eyestalks completely absent in adult.Antennal and antennular flagella strongly developed. Anten-nal scale enlarged, completely filling anterior margin of cara-pace. First and second pereopods chelate, usually hidden be-neath branchiostegite. Third to fifth pereopods well developed,stout.
Remark: Comparing the sequence of gene coding for mito-chondrial cytochrome oxydase subunit one, VAN DOVER et al.(2001) found a small divergence (0.9%) from R. exoculata.
Biology: Swarming with extremely high density on the surfaceof active chimneys and diffuse vents. Like R. exoculata, symbio-sis with chemoautotrophic bacteria.
Distribution: Central Indian Ridge: Kairei and Edmond VentFields, near the Rodriguez Triple Junction.
References:
HASHIMOTO J., OHTA S., GAMO T., CHIBA H., YAMAGUCHI T., TSUCHIDA S., OKUDAIRA T., WATABE H., YAMANAKA T. & M. KITAZAWA (2001) Zool. Sci. 18(5): 717-721.VAN DOVER C.L., HUMPHRIS S.E., FORNARI D., CAVANAUGH C.M., COLLIER R., GOFFREDI S.K., HASHIMOTO J., LILLEY M.D., REYSENBACH A.L., SHANK T.M., VON DAMM
K.L., BANTA A., GALLANT R.M., GÖTZ D., GREEN D., HALL J., HARMER T.L., HURTADO L.A., JOHNSON P., MCKINESS Z.P., MEREDITH C., OLSON E., PAN I.L.,TURNIPSEED M., WON Y., YOUNG III C.R. & R.C. VRIJENHOEK (2001) Science 294: 818-823.
WATABE H. & J. HASHIMOTO (2002) Zool. Sci. 19(10): 1167-1174.
J. HASHIMOTO Denisia 18 (2006): 429
1: Two specimens taken onboard; scale bar 5 mm; by courtesy of J. Hashimoto.
2: In situ aggregation of adult specimens at the KaireiVent Field; below right, sea anemone Marianactis cf.bythios, and mytilid bivalves Bathymodiolus sp.; cruiseROV Kaiko © JAMSTEC.
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
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Shinkaicaris leurokolos (KIKUCHI & HASHIMOTO, 2000)
Size: An ovigerous female carapace length 12.5 mm.
Color: Ivory to dull ivory white.
Morphology: Integument of body thin, but not membranous,surface shining. Rostrum compressed laterally, reaching distalmargin of first segment to midlength of second segment of an-tennular peduncle; dorsal margin armed with 7-10 teeth slight-ly diminishing in size anteriorly, including 6-9 on rostrum prop-er and 1-3 on carapace. Carapace somewhat compressed later-ally; postrostral median ridge relatively low, rather blunt; an-tennal tooth sharp, distinctly buttressed, margin inferior to an-tennal tooth forming small rounded lobe. Strong median ster-nal spine between coxae of fifth pereopods. Abdomen smoothdorsally; pleural margin of anterior two somites broadly round-ed, that of third somite broadly rounded or with one small pos-teroventral tooth; pleuron of fourth somite with acute postero-lateral tooth and occasionally with additional 1-3 acute teethon posterior margin; pleuron of fifth somite with 2-4 posterior
teeth including posteroventral tooth. Telson elongate subrec-tangular, with 5-7 dorsolateral spines arranged in a sinuous row;posterior margin convex, with two pairs of lateral spines (mesialpair longer than lateral) and 22-24 long plumose setae. Eyes onbasally separated, movable stalks broadly fused mesially; divi-sion of corneal region and stalk unclear; corneal region, un-faceted, with irregular, scattered pigment-like masses withinstalk; anteromedial face flat, sloping to anterolateral face, with-out spiniform tubercle on anterior surface. Dorsolateral flagel-lum longer than carapace; ventromesial flagellum somewhatlonger. Antennal flagellum longer than body. Uropod with ra-mi subequal in length, exopod with small movable spine mesialto smaller distolateral tooth and sinuous diaeresis.
Biology: This species occurs in a thermally influenced area.
Distribution: Known only Okinawa Trough: Minami-EnseiKnoll, 705 m.
References:
KIKUCHI T. & J. HASHIMOTO (2000) Species Diversity 5: 135-148.KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.
T. KOMAI & M. SEGONZAC Denisia 18 (2006): 430
3: Telson and uropods,dorsal view. 1-3: Scale bar 1.0 mm;from KIKUCHI &HASHIMOTO (2000).
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
1: Holotype female in lateral view.
2: Anterior region,dorsal view.
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Lebbeus carinatus ZARENKOV, 1976
Size: Maximum carapace length 17 mm.
Morphology: Rostrum short, not reaching distal margin of firstsegment of antennular peduncle; dorsal margin armed with 4-6teeth including 2-4 on rostrum proper and two on carapace pos-terior to orbital margin, posteriormost tooth arising about 0.15length of carapace; ventral blade poorly developed; ventralmargin with 1-3 small teeth subdistally. Carapace with smallsupraorbital spine; broad V-shaped notch just below supraor-bital spine; orbital margin depressed, forming shallow concavi-ty accommodating eye-stalk; suborbital lobe acutely triangular,somewhat upturned, slightly exceeding antennal spine; antero-lateral margin between antennal spine and pterygostomial an-gle strongly sinuous; pterygostomial spine small; dorsal surfacestrongly convex in spawning females, not inflated in males. Ab-domen rounded dorsally; fourth pleuron broadly rounded, fifthpleuron with strong posteroventral tooth. Telson with 4-6 dor-solateral spines on either side; posterior margin slightly convexor truncate, with two pairs of spines at lateral angle and one ortwo pairs of plumose setae. Eye small, corneal diameter0.09–1.00 of carapace length; cornea semispherical, darkly pig-mented and clearly faceted. Antennule with stylocerite notreaching distal margin of first segment of peduncle; first pedun-cular segment armed with two or three spines on dorsodistalmargin laterally; second and third segments each with one dor-sodistal spine; flagella sexually dimorphic, those of males elon-gate, particularly ventral flagellum of male exceeding 1.5
length of carapace. Antennal scale moderately broad; distolat-eral tooth falling short of or reaching rounded distal lamella.Third maxilliped overreaching antennal scale by half length ofultimate segment. First pereopod chelate. Second pereopod al-so chelate, carpus divided into seven articles. Third to fifthpereopods slender, third overreaching antennal scale by fulllength of propodus; merus of third and fourth pereopods withfour or five spines on lateral surface; that of fifth unarmed orarmed with one spine; dactyli short, 0.10-0.13 length of propo-di, each with four or five accessory spinules on ventral margin.Third maxilliped and first and second pereopods each withstrap-like epipod.
Remark: This species is easily distinguished from L. washingto-nianus of KIKUCHI & OHTA (1995), the other congenericspecies occurring at hydrothermal vents, by the absence of astrap-like epipod on the third pereopod and the more slender,elongate third to fifth pereopods. Another different species wasdescribed by SAINT LAURENT (1984) under the same name. Itdiffers mainly from the other caridean species associated withhydrothermal vents by the well developed eyes and the carpusof the second pereopod divided in seven articles.
Biology: This species live among the tubeworm Riftia pachypti-la and mussel beds of Bathymodiolus spp.
Distribution: Known with certainty only from the East PacificRise: 13°N.
Arthropoda, Crustacea, Decapoda, Caridea, Hippolytidae
T. KOMAI Denisia 18 (2006): 431–432
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References:
KIKUCHI T. & S. OHTA (1995) J. Crustac. Biol. 15: 219-240.SAINT LAURENT M. DE (1984) C. R. Acad. Sc., Paris 299(9), sér. III: 355-360.ZARENKOV N.A. (1976) J. Mar. Biol. 5: 277-285.
1: Body, lateral view; from N. ZARENKOV (1976).
2: Cephalothorax, lateral view; fromEast Pacific Rise 13°N, cruise Hope 99; by P. Briand © Ifremer.
3: Specimen in situ, among tubeworms(Riftia pachyptila), from East Pacific Rise13°N, cruise Hope 99 © Ifremer.
432
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Thymopides laurentae SEGONZAC & MACPHERSON, 2004
2: Carapace, anterior part andcephalic appendages, lateral view.
3: Fifth and sixth abdominal somitesand tail fan with telson, dorsal view.
4: Carapace dorsalview.
5: Photograph taken on board, after sampling © Ifremer
6: In situ of one lobster (probably T. laurentae)taken from the submersible at Snake Pit field;cruise Hydrosnake © Ifremer
Arthropoda, Crustacea, Decapoda, Astacidea, Nephropidae
1–4: Scale bar 1 cm; drawings by M. de Saint Laurent.
1: Body, lateral view.
References:
MÈVEL C., AUZENDE J.-M., CANNAT M., DONVAL J.-P., DUBOIS J., FOUQUET Y., GENTE P., GRIMAUD D., KARSON J.A., SEGONZAC M. & M. STIEVENARD (1989) C. R. Acad.Sci., Paris 308(II): 545-552.
SEGONZAC M. (1992) C. R. Acad. Sci., Paris 314(III): 593-600.SEGONZAC M. & E. MACPHERSON (2004) Cah. Biol. Mar. 44: 361-367.
M. SEGONZAC & E. MACPHERSON Denisia 18 (2006): 433
Size: Carapace length 45 mm; total length 97 mm.
Morphology: Carapace smooth, rostrum slender, nearly 0.3 ofcarapace length, overreaching antennular and antennal pedun-cles. Abdomen without longitudinal median carina and palm offirst chela distinctly longer than wide. Telson 1.7 times longerthan wide and longer than abdominal somite 6. Eyes small, un-pigmented, cornea as wide as ocular peduncle.
Biology: Lives on the slope of the Snake Pit vent area, near theblack smoker edifice Beehive, among galatheid crab Munidopsisexuta, and chaetopterid polychaetes.
Distribution: Mid-Atlantic Ridge: near the Snake Pit hy-drothermal area.
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Arthropoda: Decapoda, Anomura
Worldwide, there are over 2500 species of anomourancrabs, which comprise ca 5% of all crustacean species. The In-fraorder Anomura represents a paraphyletic group that includesthe superfamilies Lomisoidea, Hippoidea, and the much morediverse Galatheoidea and Paguroidea. Species of these taxa arecommonly found living from the intertidal zone to the abyssalplain >2000 m, including one terrestrial representative. Mor-phologically they have little in common, some are like crabs(e.g. Lithodidae) and others are like hermit crabs (e.g. Paguri-dae). They only share one character: the small fifth pereiopod.Molecular studies have shown that Galatheoidea and Paguroi-dea are more related to each other than to Hippoidea, althoughmore work is needed to completely resolve these relationships.
The vent fauna contains representatives of the superfami-lies Galatheoidea and Paguroidea, including species of four fa-milies and a recent new family (Kiwaidae). Despite their ecolo-gical importance and high diversity, many aspects of their sys-tematics and distribution are still poorly known.
The anomurans exhibit a considerable diversity of repro-duction modes, life cycles and capacities for dispersal. The vastmajority of species have relatively small pelagic eggs, with theexception of some representatives of the families Galatheidaeand Chirostylidae, and a pelagic larval phase, which enhancestheir capacity for dispersal. There is evidence for prolongedbrooding periods. Usually they produce only a few large eggs,probably related to an abbreviated or direct larval develop-ment. Most species are not restricted to hydrothermal ventsand cold seeps. They usually occur in low densities, althoughthere are some interesting exceptions (e.g. Munidopsis lentigo,Shinkaia crosnieri). The abundance of anomurans in vent andseep environments probably reflects the benefit derived fromthe organic matter produced by the chemosynthetic communi-ty inhabiting these zones. In general, their abundance increasedin the vicinity of active hydrothermal sites. In the center ofhydrothermal activity, however, abundance decreases.
Information on the geographic distribution and taxonomyof the different species is still limited and incomplete. Most spe-cies have a broad geographic distribution, as has been shown fora number of deep-sea taxa. There appears to be some inter-change among the different communities separated by long dis-tances, associated with high dispersal capabilities, although ahigh degree of local endemism can also exist. Unfortunately,mechanisms for colonization by vent organisms remain largelyunknown for most of the mid-ocean ridge systems. Therefore,additional studies on molecular data and larval dispersal pro-cesses are desirable.
Munidopsis sp. (probably M. lauensis); cruise TUIM 07; by C.R. Fisher.
References:
MARTIN J.W. & G.E. DAVIS (2001) Nat. Hist. Mus. Los Angeles Cty, Sci. Ser. 39: 1-124.MARTIN J.W. & T.A. HANEY (2005) Zool. J. Linn. Soc. 145: 445-522.MORRISON C.L., HARVEY A.W., LAVERY S., TIEU K., HUANG Y. & C.W. CUNNINGHAM (2002) Proc. R. Soc. Lond., B 269: 345-350.PERÉZ-LOSADA M., JARA C.G., BOND-BUCKUP G., PORTER M.L. & K.A. CRANDALL (2002) J. Crustac. Biol. 22: 661-669.
E. MACPHERSON & K. BABA Denisia 18 (2006): 434
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Paragiopagurus ventilatus LEMAITRE, 2004
Size: Shield length 5.8 mm max. (male); 4.9 mm max. (oviger-ous female).
Color: In life, shield and cephalic appendages cream with somefaded pink areas. Chelipeds and ambulatory legs red or pink.
Morphology: Gills biserial. Ocular acicles subtriangular, termi-nating in strong, simple spine (rarely bifid) with transverse stri-ae. Outer faces of basis and ischium of third maxillipeds, che-lipeds, and second and third pereopods, with plumose, bacte-riophore setae. Chelipeds markedly dissimilar; proportions andarmature of right strongly affected by size and sex, with that ofmales considerably longer and more elongate than in females.Dactyls of second and third pereopods with irregular rows ofspines on ventromesial margins. Propodal rasp of fourth pereo-pod with one row of scales. Anterior lobe of sternite XII (third
pereopods) with bifid spine. Telson asymmetrical; posteriormargin separated by broad, shallow cleft, into rounded projec-tions armed with few short, corneous spines often ventrallycurved. Males lacking first pleopods; with unpaired rudimenta-ry left pleopod.
Biology: Specimens collected by beam-trawl among otherspecies newly described as mytilid and lucinid bivalves(BOUCHET & COSEL 2004), and numerous grapsid crabs (NG etal. 2000). At this area, diffuse gas seepage from the bottom.The specimens live in gastropod shells of the buccinid Siphona-lia. Probably not part of the “obligate” vent fauna. Ovigerous fe-males found in May 2001.
Distribution: Okinawa Arc: Northeastern Taiwan, Tashi fish-ing grounds.
References:
BOUCHET P. & R. VON COSEL (2004) Zool. Stud. 43: 704-711.LEMAITRE R. (2004) Cah. Biol. Mar. 45: 325-334.NG N.K., HUANG J.F. & P.H. HO (2000) Nat. Taiwan Mus., Spec. Publ. Ser. 10: 191-199.
R. LEMAITRE Denisia 18 (2006): 435
1: Male in vivo; by courtesy of T. Yam Chan. 2: Female in vivo; by courtesy of T. Yam Chan.
Arthropoda, Crustacea, Decapoda, Anomura, Parapaguridae
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Paralomis hirtella DE SAINT LAURENT & MACPHERSON, 1997
Size: Up to 59 mm carapace length.
Morphology: Carapace more or less hexagonal. Dorsal surfacesmooth, with few scattered small granules. Numerous tuft of se-tae of different length scattered on carapace surface. Rostrumwith basal spine and two small dorsal spines. Chelipeds bearingthick spines and acute granules. Walking legs moderately long,with some thick spines on dorsal border of merus, carpus andpropodus, dactylus slightly curved, with corneous spinelets
along ventral margin. Scattered tuft of setae on posterior sur-face, dorsal and ventral margins of articles.
Biology: Lives in few numbers around active chimneys and onmussel beds.
Distribution: North Fiji Back-Arc Basin: White Lady ventfield; Lau Back-Arc Basin: Hine Hina vent field.
1: Holotype male, 32 x 35 mm, habi-tus, dorsal view; scale 10 mm; by M.de Saint Laurent & E. Macpherson.
2: Holotype, abdomen; scale bar 10 mm; byM. de Saint Laurent & E. Macpherson.
3: Other specimen 44 x 45 mm, abdomenof ovigerous female; scale bar 10 mm; byM. de Saint Laurent & E. Macpherson.
4: Specimen collected at Lau Back-Arc Basin: Hine Hina site; by P. Briand © Ifremer.
5: Two specimens in situ on gastropod bed of Ifremerianautilei at North Fiji Back-Arc Basin: White Lady site;cruise Starmer 2 © Ifremer.
Arthropoda, Crustacea, Decapoda, Anomura, Lithodidae
References:
DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA (1994) Mar. Geol. 116: 227-242.SAINT LAURENT M. DE & E. MACPHERSON (1997) Zoosystema 19: 721-727.
E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 436
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Paralomis verrilli (BENEDICT, 1895)
Size: Up to 110 mm carapace length.
Morphology: Carapace more or less hexagonal, as long as wide.Dorsal carapace surface covered with small granules of varioussize. Granules usually with several setae. Regions well-defined.Gastric region rather more prominent than other regions, witha thick spine on apex. Cardiac region clearly less prominentthan branchial regions, with four thick granules in square pat-tern. Each branchial region with one median spiniform granuleand two smaller thick granules near intestinal region. Basalspine of rostrum more or less horizontal, slightly overreachingend of cornea, with spiniform tubercles on ventral side; two di-
vergent dorsal spines, upwardly directed, extending well pastend of cornea. External orbital spine slightly shorter than eyes.Anterolateral spine slightly shorter than external orbital. Che-lipeds bearing thick spines on mesial border of merus, carpusand hand. Walking legs moderately long, with well-developedspines along dorsal and ventral margins of meri and propodi,dactyli as long or longer than propodi.
Distribution: North Pacific, from Japan, Bering Sea to Califor-nia, 850-2379 m. Collected on vent sites of Juan de Fuca Ridge:Axial Seamount.
References:
DAWSON E.W. (1989) N.Z.O.I Misc. Publ. 101: 1-338.IKEDA H. (1998) The Deep-sea Crabs of Sagami Bay, Imamoto Printing: 1-180.MACPHERSON E. (2001) Zoosystema 23: 797-805.ZAKLAN S.D. (2002) Alsk. Sea Grant Coll. Progr.: 751-845.
E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 437
1: Specimen among siboglinid tubeworms and microbial mats on side of sulphide edifice; from R/V Thomas G. Thompson, cruiseTN149 (Zooarium vent field, Explorer Ridge; 31 July 2002) © NOAA.
Arthropoda, Crustacea, Decapoda, Anomura, Lithodidae
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Munida magniantennulata BABA & TÜRKAY, 1992
Size: Up to 12.8 mm carapace length.
Morphology: Dorsal carapace surface smooth, with epigastricspines. Lateral branchial margins with five small spines. Firstlateral spine of carapace near anterolateral angle, second spineclearly smaller than first. Second abdominal segment withtransverse row of small spines, other segments unarmed.Corneae small, width equal to or less than distance betweenthan distance between sinus formed by supraocular spines androstrum. Basal segment of antennular peduncle unusually large.
Distomesial spine of first antennal article very small, neverreaching midlength of second article. Merus of third maxillipedwith only one spine on flexor margin. Chelipeds moderatelymassive, about three times carapace length. Walking legs slen-der.
Biology: No data. Collected by grab TV from an area close toactive sites.
Distribution: Lau Back-Arc Basin: Hine Hina area.
References:
BABA K. (1994) Mem. Qld. Mus. 35: 1-21.BABA K. (2005) Galathea Report 20: 1-317.BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.BABA K. & M. TÜRKAY (1992) Senckenb. Marit. 22: 203-210.
E. MACPHERSON & K. BABA Denisia 18 (2006): 438
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
1: Holotype male, carapacelength 7.2 mm; dorsal view;scale bar 1 mm; from BABA & TÜRKAY (1992).
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Munidopsis acutispina BENEDICT, 1902
4: Specimen from Lost City; cruise Exomar; by P. Briand © Ifremer.
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
References:
FROGLIA C., MURA, M. & A. BONFITTO (2002) Crustac. Int. J. Crustac. Res. 75: 375-382.GEBRUK A.V., GALKIN S.V., KRYLOVA E.M., VERESHCHAKA A.L. & G.M. VINOGRADOV (2002) InterRidge News 11(2): 18-19.MACPHERSON E. & M. SEGONZAC (2005) Zootaxa 1095: 1-60.OLU-LE ROY K., SIBUET M., FIALA-MÉDIONI A., GOFAS S., SALAS C., MARIOTTI A., FOUCHER J.-P. & J. WOODSIDE (2004) Deep-Sea Res. I 51: 1915-1936.
E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 439
1: Carapace, dorsal view; male, carapacelength 8.0 mm; from FROGLIA et al. (2002).
2: Carapace and abdomen, lateral view; scale bar 5 mm;male; from FROGLIA et al. (2002).
Size: Up to 8.0 mm carapace length.
Morphology: Carapace with numerous pointed tubercles. Ros-trum narrow triangular, horizontal and dorsally carinated. Frontmargin convex, antennal spine small. Abdomen unarmed.Cornea well exposed, visible in dorsal view, eye spine mesialand small. Fixed finger of chelipeds without denticulate carinaon distolateral margin. Walking legs not reaching end of che-lipeds. Propodi of walking legs of uniform width, with distinctspines. Epipods absent from pereopods.
Biology: Occurs in both hydrothermal and cold seep areas.
Distribution: Mid-Atlantic Ridge: Lost City vent field, andEastern Mediterranean Sea, Kazan mud volcano, cold seep,2030 m. Known as well off Western North Africa and theAzores Islands, between 698 and 845 m, and from the Mediter-ranean Sea (W Tyrhenian Sea, Sardinia) between 374 and1036 m.
3: Telson; scale bar 5 mm; from FROGLIA etal. (2002).
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Munidopsis alvisca WILLIAMS, 1988
2: Carapace, lateral view; scale bar 5 mm; by A.B. Williams.
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
Size: Up to 27.9 mm carapace length.
Morphology: Dorsal carapace surface without spines, coveredby scale-like rugosities. Rostrum unarmed, broad at base, distal-ly narrowed and upturned, weakly carinated dorsally. Frontmargin oblique, small antennal spine present. Abdomen un-armed. Eyes not movable, eyespine well developed, cornea rel-atively large, clearly wider than eyespine. Chelipeds with sev-eral spines on merus and carpus, palm spineless, fixed finger ofchelipeds without denticulate carina on distolateral margin.
First walking leg reaching nearly tip of cheliped, dorsal crest ofmerus and carpus with row of spines along dorsal border; dacty-lus smoothly narrowed distally, flexor margin slightly curving,bearing low spines. Epipods absent from pereopods.
Biology: Living around hydrothermal vent sites.
Distribution: Explorer Ridge: Magic Mountain; Juan de FucaRidge; Guaymas Basin.
References:
BABA K. (2005) Galathea Report 20: 1-317.KHODKINA I.V. (1991) Zool. Zh. 70: 71-76.WILLIAMS A.B. (1988) Fish. Bull. 86: 263-287.
E. MACPHERSON & K. BABA Denisia 18 (2006): 440
1: Carapace and abdomen, dorsalview; holotype carapace length23.7 mm (including rostrum);scale bar 5 mm; by A.B. Williams.
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Munidopsis diomedeae (FAXON, 1893)
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
Size: Up to 30 mm (with rostrum).
Morphology: Carapace covered with tubercles, dorsally armedwith two well-developed epigastric, one extremely strong later-ally compressed mesogastric, and one moderately large cardiacspine, with two lateral spines. Dorsal carapace surface smooth,without spines. Rostral spine upturned distally. Second tofourth abdominal tergites each with prominent median spine.Eyes movable and spineless, corneae subglobular. Chelipedscovered with granules, nearly spineless. Walking legs long,
slender, with numerous granules, subcylindrical. First walkingleg overreaching chelipeds. Dactyli compressed laterally, flexorborder moderately curving, bearing proximally diminishing lowspines. Epipods present on chelipeds and first and second walk-ing legs.
Distribution: East Pacific Rise. The species has been sampledin Eastern Pacific, from the Gulf of California to Chile, be-tween 768 and 2026 m.
References:
BABA K. (2005) Galathea Report 20: 1-317.FAXON W. (1895) Mem. Mus. Comp. Zool. Harvard Coll. 18: 1-292.HAIG J. & M. WICKSTEN (1975) Bull. South Calif. Acad. Sci. 74: 100-105.KHODKINA I.V. (1991) Zool. Zh. 70: 71-76.
E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 441
1: Male, carapace length ca. 30 mm(with rostrum), habitus, dorsal view;from FAXON (1895).
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Munidopsis exuta MACPHERSON & SEGONZAC, 2005
3: Specimen from Snake Pit; cruise Hydrosnake; by P. Briand© Ifremer.
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
References:
MACPHERSON E. & M. SEGONZAC (2005) Zootaxa 1095: 1-60.SAINT LAURENT M. DE (1985) in Ifremer Brest (Ed.) Peuplements Profonds du Golfe de Gascogne: 469-478.SEGONZAC M. (1992) C. R. Acad. Sci. Paris III 314: 593-600.
E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 442
1: Carapace, dorsal view; holotype, carapace length34.7 mm; scale bar 5 mm; from MACPHERSON & SEGON-ZAC (2005).
Size: Up to 39.5 mm carapace length.
Morphology: Carapace with pair of epigastric spines, some-times 2-3 additional small spines, covered by scale-like rugosi-ties. Rostrum broad at base, distally narrowed and upturned,weakly carinated dorsally. Front margin oblique, antennal spineabsent. Abdomen unarmed. Eyes not movable, cornea relative-ly large, clearly wider than eyespine. Chelipeds with severalspines on merus and carpus, palm spineless, fixed finger moder-ately ridged along distal third of lateral margin. First walkingleg slightly exceeding cheliped, dorsal crest of merus and carpuswith row of spines along dorsal border; dactylus smoothly nar-rowed distally, flexor margin slightly curving, bearing lowspines. Epipods on chelipeds, not on walking legs. The specieswas identified as M. subsquamosa by SAINT LAURENT (1985)and M. crassa by SEGONZAC (1992).
Biology: This species occurs in both abyssal (Gulf of Biscay,2860 m) and hydrothermal vent environments (Mid-AtlanticRidge: Snake Pit, Elan). It has been collected by baited trap to-gether with a nephropid lobster Thymopides laurentae SEGON-ZAC & MACPHERSON, and a macrurid fish Coryphaenoides arma-tus (HECTOR), at the base of the active edifice, on sulphiderocks, among scattered sea anemones, chaetopterid tubewormsand alvinocaridid shrimps. The species has also been collectedat the base of inactive chimney complex, among dead mytilidbivalves. An additional specimen (unfortunately lost) was col-lected during the cruise BRIDGE 1993 at the Mid-AtlanticRidge hydrothermal vent site Broken Spur site (dive Alvin2625, 27.06.1993, E. Southward, personal communication).
Distribution: Mid-Atlantic Ridge: Snake Pit, TAG, probablyBroken Spur, and Logatchev; Gulf of Biscay.
2: Carapace, lateral view; holo-type, carapace length 34.7mm; scale bar 5 mm; fromMACPHERSON & SEGONZAC (2005).
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Munidopsis lauensis BABA & DE SAINT LAURENT, 1992
3: Munidopsis sp. (probably M. lauensis); cruise TUIM 07; by courtesy of C.R. Fisher.
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
References:
BABA K. (2005) Galathea Report 20: 1-317.BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA (1994) Mar. Geol. 116: 227-242.
E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 443
1: Carapace and abdomen, dorsal view; holo-type male, carapace length 10.8 mm; scalebar 1 mm; by K. Baba & M. de Saint Laurent.
Size: Up to 12.4 mm carapace length.
Color: Generally white, but some rare specimens are blackish,due to a manganese deposit.
Morphology: Carapace without dorsal spines or tubercles. Ros-trum spiniform, horizontal and unarmed. Eye with small mesialspine, cornea large. Abdomen unarmed. Sixth abdominal seg-ment having posteromedian margin transverse, exceeded bylateral lobes. Epipods absent from pereopods. First walking legs
overreaching chelipeds. Fixed finger of chelipeds without den-ticulate carina on distolateral margin. Chelipeds and walkinglegs, with few small spines. Dactylus of walking legs with spinesalong entire flexor border.
Biology: On active hydrothermal sites, often in high density.
Distribution: North Fiji Back-Arc Basin: White Lady ventfield; Lau Back-Arc Basin: Valu Fa Ridge, Hine Hina ventfields.
2: Carapace, lateral view; holotype male, carapace length 10.8 mm; scale bar 1mm; by K. Baba & M. de Saint Laurent.
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Munidopsis lentigo WILLIAMS & VAN DOVER, 1983
2: Carapace, lateral view; femaleparatype, carapace length 11.9 mm;from WILLIAMS & VAN DOVER (1983).
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
Size: Up to 12.8 mm carapace length.
Morphology: Dorsal carapace surface smooth, without spines.Rostrum unarmed, broad triangular and horizontal, not cari-nated dorsally. Front margin transverse, antennal spine present.Abdomen unarmed. Flattened eyes, depressed cornea, large me-dian eyespine present. Chelipeds with several spines on merusand carpus, and palm, fixed finger of chelipeds without dentic-ulate carina on distolateral margin. Lenticular-shapped, flat-
tened light brown spot on ventral surface of chela. First walk-ing leg reaching nearly tip of cheliped, dorsal crest of meruswith row of small spines; dactylus short, flexor margin bearinglow spines. Epipods absent from pereopods.
Biology: Lives around the hydrothermal vents.
Distribution: East Pacific Rise: 21°N; south of Baja California.
1: Habitus, dorsal view; scale bar 1 mm; female paratype, carapace length 11.9 mm; from WILLIAMS & VAN DOVER (1983).
References:
BABA K. (2005) Galathea Report 20: 1-317.WILLIAMS A.B. & C. VAN DOVER (1983) Proc. Biol. Soc. Wash. 96: 481-488.
E. MACPHERSON & K. BABA Denisia 18 (2006): 444
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Munidopsis marianica WILLIAMS & BABA, 1989
2: Rostrum and anterior part ofcarapace, dorsal view.
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
References:
BABA K. (2005) Galathea Report 20: 1-317.WILLIAMS A.B. & K. BABA (1989) Fish. Bull. 87: 899-910.
E. MACPHERSON & K. BABA Denisia 18 (2006): 445
1: Carapace and abdomen, dorsal view.
1-4: Female holotype, carapace length 38.0 mm; scale bar 10 mm; by A.B. Williams & K. Baba.
Size: Up to 38.0 mm carapace length.
Morphology: Dorsal carapace surface without spines, coveredby scale-like rugosities. Rostrum unarmed, broad triangular andhorizontal, weakly carinated dorsally. Front margin oblique,small antennal spine present. Abdomen unarmed. Eyes mov-able, large mesial eyespine small lateral eyespine, cornea rela-tively large, wider than mesial eyespine. Chelipeds with sever-al spines on merus and carpus, palm spineless, fixed finger of
chelipeds without denticulate carina on distolateral margin.First walking leg reaching nearly tip of cheliped, dorsal crest ofmerus and carpus with row of spines along dorsal border; dacty-lus smoothly narrowed distally, flexor margin nearly stright,bearing low spines. Epipods present on chelipeds.
Biology: Lives at both vent and non vent areas.
Distribution: Western Pacific Ocean, Mariana Back-Arc Basin.
3: Anterior part of carapace, lateral view.
4: Telson, no scale.
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Munidopsis sonne BABA, 1995
3: Telson.
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
1: Carapace and abdomen, dorsal view.
Size: Up to 12.8 mm carapace length.
Morphology: Dorsal carapace surface covered with numeroustubercle-like setiferous processes, gastric region with pair of epi-gastric processes. Rostrum nearly triangular, nearly horizontal,lateral margins serrate. Front margin with antennal acuteprocess. Anterolateral angle produced. Abdomen unarmed;sixth abdominal segment with weak posterolateral lobe, poste-rior margin not produced. Eyes immovable, cornea narroweddistally. Antennular basal segment cristate dorsolaterally, withrow of several spines. Chelipeds subcylindrical, covered with
tubercular processes and blunt short spines, longer than secondwalking legs; fixed finger without denticulate carina on disto-lateral margin. Walking legs diminishing in length posteriorly;dactyli rather straight in first and second legs, more curving onthird; setae along distal two-thirds of flexor border. Epipodspresent on chelipeds, first and second walking legs.
Biology: Lives around hydrothermal vents.
Distribution: North Fiji Back-Arc Basin.
2: Carapace and abdomen, lateral view.
1-3: Ovigerous female holotype, carapace length 10.5 mm; scale bar 1 mm; by K. Baba.
References:
BABA K. (1995) Crustac. Res. 24: 188-193.BABA K. (2005) Galathea Report 20: 1-317.
E. MACPHERSON & K. BABA Denisia 18 (2006): 446
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Munidopsis starmer BABA & DE SAINT LAURENT, 1992
3: Telson; scale bar 1 mm.
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
References:
BABA K. (2005) Galathea Report 20: 1-317.BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.
E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 447
1: Carapace and abdomen, dorsal view; scale bar 10 mm.
1-3: Female holotype, carapace length 31.4 mm; by K. Baba & M. de Saint Laurent.
Size: Up to 33.4 mm carapace length.
Morphology: Carapace with two epigastric spines, dorsal sur-face smooth. Rostrum spiniform, upwards directed, unarmed.Eye with small mesial spine, cornea large. Abdomen unarmed.Sixth abdominal segment having posteromedian margin trans-verse, exceeded by lateral lobes. Epipods absent from pere-opods. First walking legs overreaching chelipeds. Fixed finger of
chelipeds without denticulate carina on distolateral margin.Chelipeds and walking legs, with few small spines. Dactylus ofwalking legs with spines along entire flexor border.
Biology: Lives on active hydrothermal vents.
Distribution: North Fiji Back-Arc Basin: site White Lady.
2: Carapace and abdomen, lateral view; scale bar 10 mm.
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Munidopsis WHITEAVES, 1874
Size: Up to 45 mm carapace length.
Morphology: Carapace with gastric region having group ofspines including two epigastric spines and some additionalsmall spines or tubercular spines on scale-like ridges. Lateralmargin having first spine (anterolateral) distinctly larger thanantennal spine, directed anterolateral, second spine strongerthan first. Rostrum subtriangular in proximal half, distally nar-rowed, slightly upcurved, dorsally carinated. Front marginoblique, antennal spine small. Abdominal segments unarmed,sixth segment having posteromedian margin slightly convex,not produced. Telson composed of 10 plates, midlateral plateproduced on anterolateral margin. Ocular peduncles broad atbase, distomesially with eye-spine distinctly longer thancornea; cornea relatively small, as broad as eye-spine. Fixed fin-ger of chelipeds without denticulate carina on distolateral mar-gin. First walking leg exceeding chelipeds; palm shorter than
fixed finger, bearing a few spines along mesial margin; fingersspooned at tip. Dactyli of walking legs smoothly narrowed dis-tally, flexor margin bearing ultimate denticle equidistant be-tween penultimate denticle and end of article. Epipods presenton chelipeds.
Remarks. This species has been reported from the hydrother-mal vents of the Galapagos Spreading Center under the nameof M. subsquamosa HENDERSON, 1885, but it belongs to a differ-ent species, probably undescribed. A more careful examination,including molecular analyses, of the specimens from differentlocalities will clarify the systematic status of this species.
Biology: Occurrences around the hydrothermal vents may berevised.
Distribution: Galapagos Spreading Center, East Pacific Rise.
Reference:
BABA K. (2005) Galathea Report 20: 1-317.
E. MACPHERSON & K. BABA Denisia 18 (2006): 448
1: Carapace, dorsal view; scale bar 5 mm;from BABA (2005).
2: Right, carapace, lateral view; bar 5 mm; from BABA (2005).
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
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Shinkaia crosnieri BABA & WILLIAMS, 1998
Size: Up to 43.7 mm carapace length (Taiwanese specimen 45mm, 58.9 mm with rostrum).
Morphology: Dorsal carapace surface smooth, lateral marginsbearing many small spines. Rostrum prominent, flattened dor-sally, triangular, not carinated dorsally. Orbital margin concave,antennal spine present. Pterygostomian flap anteriorly pro-duced, covering greater part of antennal peduncle. Dense longplumose setae on sternum, pterygostomian flap and ventral sur-face of pereopods. Abdomen unarmed. Flattened eyes, with
dorsally flattened projection. Chelipeds strong, broad, de-pressed, with numerous short spines. Walking legs stout andmoderately flattened. Epipods present on chelipeds, first andsecond walking legs.
Distribution: Western Pacific: Edison Seamount; OkinawaTrough, Iheya Ridge, North Iheya Knoll; off NE Taiwan, 1200-1500 m.
1: Male carapacelength 45 mm,dorsal view; by T.-Y. Chan.
Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae
References:
BABA K. (2005) Galathea Report 20: 1-317.BABA K. & A.W. WILLIAMS (1998) Zoosystema 20: 143-156.CHAN T.Y., LEE D.-A. & C.S. LEE (2000) Bull. Mar. Sci. 67(2): 799-804.FUJIKURA K. & T. OKUTANI (2002) Benthos Research 57: 21-30.WATABE H. (2000) JAMSTEC J. Deep-Sea Res. 17: 29-34.
E. MACPHERSON & K. BABA Denisia 18 (2006): 449
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Kiwa hirsuta MACPHERSON, JONES & SEGONZAC, 2005 “yeti crab”
1: Carapace and abdomen, lateral view; scale bar 10 mm; by J.-F. Dejouannet © IRD.
2: Sixth segment of abdomen, telson and uropods, dorsal view; scale bar 5 mm; by J.-F. Dejouannet © IRD.
3: Two specimens on mussel bed, amongbythograeid crabs on vent site Annie’s Anthill(Southern East Pacific Rise); cruise PAR 5; by courtesyof R. Vrijenhoek © MBARI.
Arthropoda, Crustacea, Decapoda, Anomura, Kiwaidae
M. SEGONZAC Denisia 18 (2006): 450–451
Size: Carapace length 51.5 mm (58.6 mm with rostrum), totallength 88.4 mm.
Color: Uniformly white, with yellow corneous tip at the ex-tremity of the chelipeds 1.
Morphology: Carapace, excluding rostrum, 1.3 times longerthan broad, dorsal surface smooth. Front margin slightlyoblique, with small tooth near rostrum; anterolateral anglerounded. Rostrum broadly triangular, horizontal, slightly con-cave dorsally, lateral borders granulated, with long uniramoussetae; ventral side slightly carinated. Insertion of fifth pereopodnot visible and situated below sternal plastron. Eyes strongly re-duced, membranous remains, without pigment. Antennal scaleabsent. Antennal peduncle without scaphocerite. Flagellum aslong as carapace without rostrum. Chelipeds and walking legswith dense mat of setae. Telson as wide as long, median trans-verse suture separating anterior and posterior portions. Che-lipeds and walking legs with numerous rows of spines, eachspine with yellow corneal tip and tuft of long and denseplumose setae, only absent in cheliped fingers, setae denser andlonger in mesial and ventral sides than in lateral and dorsalsides. Chelipeds nearly symmetric, slightly more than twice aslong as carapace including rostrum. Fingers somewhat triangu-lar, without setae, having numerous spines decreasing in sizedistally, distal areas of fingers unarmed; slightly gapping, and
distally spoon-shaped; movable finger with proximal large den-ticulate tooth followed by cutting margin bordered withsmooth, low, corneous scales, ending in acute corneous point;fixed finger with some proximal small teeth, followed by cut-ting edge similar to movable finger and ending in acute cor-neous point, additional row of mesial granules ending in acutecorneous point; fingers distally crossing, corneous tip of mov-able finger crossing between two corneous tips of fixed finger.Paired pleopods present.
Setae: the pereopods, and in particular the chelipeds, are dense-ly covered with flexible setae (ca. 15 mm) having clusters of fil-amentous bacteria, mainly at distal part. Other rigid chitinoussetae (ca. 13 mm) are barbed in the distal part, ending in a rigidspine, and they are regularly inserted in pairs mainly on themerus of the chelipeds. They are deprived of bacteria.
Biology: Occurs at densities of one to two individuals per 10m2, more or less regularly spaced on the zone of pillow basaltsurrounding active hydrothermal vents, and at the base ofchimneys among vent mussels Bathymodiolus sp., crabsbythograeid spp. and ophidiid fish. Omnivorous.
Distribution: Pacific-Antarctic Rise: German Flats, 38°S. Thisanimal was first observed (and noted as “type Shinkaiinae”, butnot collected), in 2001 by the German cruise Sonne SO-157(STECHER et al. 2002).
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References:
MACPHERSON E., JONES W.J. & M. SEGONZAC (2005) Zoosystema 27(4): 709-723.STECHER J., TÜRKAY M. & C. BOROWSKI (2002) Cah. Biol. Mar. 43: 271-274.
4: Male holotype, dorsal view; by A. Fifis © Ifremer.
5: One specimen on pillow lava, taken on the site Pâle Etoile (Southern East Pacific Rise); cruise, PAR 5; by courtesy of R. Vrijenhoek © MBARI.
451
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Uroptychus bicavus BABA & DE SAINT LAURENT, 1992
1: Habitus, dorsal view; holotype male, carapace length 7.5 mm; scale bar 1 mm; by K. Baba & M. de Saint Laurent.
2: Habitus, lateral view; holotype male, carapace length 7.5 mm; scale bar 1 mm; by K. Baba & M. de Saint Laurent.
Arthropoda, Crustacea, Decapoda, Anomura, Chirostylidae
Size: Up to 7.5 mm carapace length.
Morphology: Carapace smooth, lateral margin without distinctspine other than anterolateral spine. Epigastric spines on dorsalsurface. Rostrum moderately broad triangular, slightly upturnedapically. Fourth sternite rounded on anterolateral corner. An-tennal scale barely reaching midlength of fifth antennal article.Propodus of first walking legs slightly longer than carpus.Dactylus of walking legs with flexor marginal spines not con-tiguous to flexor margin.
Biology: Living in low number among vent community.
Distribution: North Fiji Back-Arc Basin: site Mussel Valley.
References:
BABA K. (2005) Galathea Report 20: 1-317.BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA (1994) Mar. Geol. 116: 227-242.
E. MACPHERSON & K. BABA Denisia 18 (2006): 452
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Uroptychus edisonicus BABA & WILLIAMS, 1998
1: Ovigerous female holotype, carapace length6.2 mm; dorsal view; scale bar 3 mm; by K. Baba& A.B. Willians.
2: Ovigerous female holotype, carapace length 6.2 mm; lateral view; scalebar 3 mm; by K. Baba & A.B. Willians.
Arthropoda, Crustacea, Decapoda, Anomura, Chirostylidae
Size: Up to 8.4 mm carapace length.
Morphology: Carapace smooth, lateral margin without distinctspine other than anterolateral spine, not laterally serrate. Epi-gastric spines absent. Rostrum narrow triangular, weakly curv-ing dorsad distally. Fourth sternite relatively narrow on antero-lateral corner. Antennal scale overreaching end of penultimatesegment. Propodus of walking legs unarmed on dorsal crest,with spines on distal portion of flexor margin, distal-most re-motely separated from distal second, and situated near juncturewith dactylus; propodus with convex flexor distal margin.Dactylus of second walking leg with two distal spines remotelyseparated from proximal group of spines.
Distribution: Western Pacific: Bismarck Archipelago, EdisonSeamount, near Lihi Island.
References:
BABA K. (2005) Galathea Report 20: 1-317.BABA K. & A.B. WILLIAMS (1998) Zoosystema 20: 143-156.
E. MACPHERSON & K. BABA Denisia 18 (2006): 453
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Uroptychus thermalis BABA & DE SAINT LAURENT, 1992
1: Habitus, dorsal view; male holotype, carapacelength 8.4 mm; scale bar 1 mm; from BABA &SAINT LAURENT (1992).
2: Habitus, lateral view; male holotype, carapace length 8.4 mm; scale bar 1 mm; from BABA & SAINT LAURENT (1992).
Arthropoda, Crustacea, Decapoda, Anomura, Chirostylidae
Size: Up to 8.4 mm carapace length.
Morphology: Carapace smooth, lateral margin without distinctspine other than anterolateral spine. Epigastric spines absent.Rostrum narrow triangular, weakly curving dorsad distally.Fourth sternite relatively narrow on anterolateral corner. An-tennal scale overreaching end of penultimate segment. Propo-dus of walking legs with spines on distal portion of flexor mar-gin, distal-most remotely separated from distal second, and sit-uated near juncture with dactylus; propodus without convexflexor distal margin. Dactylus of second walking leg with twodistal spines remotely separated from proximal group of spines.
Biology: Living in few number among vent community.
Distribution: North Fiji Back-Arc Basin: White Lady. Knownalso from Queensland, at 1497 m.
References:
AHYONG S. & G.C.B. POORE (2004) Zootaxa 436: 1-88.BABA K. (2005) Galathea Report 20: 1-317.BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.
E. MACPHERSON & K. BABA Denisia 18 (2006): 454
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Chordata, Chondrichthyes & Osteichthyes
Since the discovery of animal communities in oceanic hy-drothermal vents in 1977, fishes have been regularly observedin association with these chemosynthetically driven communi-ties, but in most cases they are difficult to catch and thereforespecies identification can often only rely on images captured bythe diving vehicles.
Ichthyologic information pertaining to species inhabitingthe deep-sea hydrothermal vents is mentioned in over 30 pa-pers and even in the more detailed and updated lists (BISCOITO
et al. 2002; GEISTDOERFER 1996, 1998; TUNNICLIFFE 1991) thereare species missing. The situation for bathyal species inhabitingthe periphery of active vent fields is even less clear since thevast majority of identifications have been based on videorecords or photographs.
Species living inside active fieldsVent fishes were found at only 20 of some 50 active vent
fields discovered to date. The specific diversity found is low andthe degree of endemism is high, which seems to be an overallcharacteristic of the hydrothermal vent fauna (TUNNICLIFFE
1991).
The family Zoarcidae dominates in terms of the number ofspecies and biomass. Eelpouts form a highly diverse family withover 220 known, mostly benthic species (WEITZMAN 1997) andare one of the more successful fish families to occupy continen-tal slopes down to 5000 m (ANDERSON 1994; WEITZMAN 1997).Nonetheless, it is remarkable that they have been able to adapt(and evolve) to the vent environments. As pointed out byGEISTDOERFER (1996) these adaptations are neither anatomical,nor trophic. In order to cope with the chemical conditions ofvents and seeps, these species must have biochemical adapta-tions. These adaptations, as well as the factors conditioning thedistribution of the species need further investigation. At thepresent level of our knowledge on the taxonomy and distribu-tion of the species, zoogeographical considerations cannot beproduced.
Bathyal species living in the vicinity of the vents and seeps
As for the peripheral bathyal fish fauna, data available arefar from being satisfactory. The five best-represented familiessensu lato (Centrophoridae, Somniosidae and Etmopteridae),
Hydrolagus pallidus from Lucky Strike, Mid-Atlantic Ridge, Atos cruise © Ifremer.
M. BISCOITO Denisia 18 (2006): 489-490
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References:
ANDERSON M.E. (1994) Ichthyol. Bull. J.L.B. Smith. Inst. Ichtyol. 60: 1-120.BISCOITO M., SEGONZAC M., ALMEIDA A.J., DESBRUYÈRES D., GEISTDOERFER P., TURNIPSEED M. & C. VAN DOVER (2002) Cah. Biol. Mar. 43: 359-362.DESBRUYÈRES D., BISCOITO M., CAPRAIS J.-C., COLAÇO A., COMTET T., CRASSOUS P., FOUQUET Y., KHRIPOUNOFF A., LE BRIS N., OLU K., RISO R., SARRADIN P.-M., SEGONZAC
M. & A. VANGRIESHEIM (2001) Deep-Sea Res. Part I 48: 1325-1346.GEISTDOERFER P. (1996) Oceanol. Acta 19(5): 539-548.GEISTDOERFER P. (1998) Ann. Inst. Océanogr. 74(2): 201-215.MERRETT N.R. & R.L. HAEDRICH (1997) Deep-sea Demersal Fish and Fisheries. Chapman & Hall, London: 1-282.TUNNICLIFFE V. (1991) Oceanogr. Mar. Biol. Annu. Rev. 29: 319-407.WEITZMAN S.H. (1997) in RANDALL J.E. & A.P. FARRELL (Eds.) Deep-Sea Fishes: 43-78.
Macrouridae, Ophidiidae, Squalidae, Moridae and Synapho-branchidae correspond to the most common families in thedeep sea (WEITZMAN 1997). However, the variations in num-bers of recorded species from field to field are enormous anddefinitely reflect insufficient research.
Although there are twice as many known active sites in thePacific as in the Atlantic, the number of species recorded in thelatter is higher (38 versus 43). Depth may well be the main rea-son for this, as vent fields in the Atlantic range from 850 mdown to 3650 m and the ones in the East Pacific are all around2500 m of depth. As pointed out by MERRETT & HAEDRICH
(1997) for the Porcupine Seabight, demersal fish abundancetends to increase with depth attaining a maximum around 1000m and then decreases sharply. This is in accordance with avail-able data, wherein the highest number of records is from MenezGwen (850 m) and Lucky Strike (1700 m) on the Mid-AtlanticRidge (DESBRUYÈRES et al. 2001).
The fact that the fields near the Azores Triple Junction(Mid-Atlantic Ridge) have been more intensively studied byprofessional ichthyologists can also have an influence in the re-sults obtained. This bias precludes us from drawing conclusionson abundance, diversity and zoogeography of this fauna. To
overcome it, scientific teams working on hydrothermal vent bi-ology should be reinforced with fish biologists as well as makingmore ship-time available for this kind of research.
The species treated herein are not only those who live in-side the active fields, which are commonly considered as “ventendemic”, but also some of the more commonly seen in the pe-riphery of the vents, in some cases feeding even on vent inver-tebrates and thus contributing for the export of energy from thechemosynthetically-driven environment to the photosyntheti-cally-dependent bathyal environment.
New speciesAmongst the so-called “vent endemic” species, at least six
new to science are being described while this contribution isbeing published: three Zoarcidae from 9°N East Pacific Rise,the Kermadec Arc and the Rodriguez Triple Junction in the In-dian Ocean respectively, one Ophidiidae from 17°S southernPacific Pacific Rise, one Myxinidae from 38°S Pacific-Antarc-tic Ridge and one Cynoglossidae from seamounts at the Mari-ana Arc and the Kermadec Arc (E. Anderson, J. Hashimoto, J.Nielsen & C. Roberts, pers. comm.).
490
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Hydrolagus affinis (CAPELLO, 1868) “small-eyed rabbitfish”
Size: To about 1300 mm total length.
Color: Uniform dark violet/brown.
Morphology: Body greatly tapering from a massive head andtrunk to a pointed caudal fin with a short filament at its tip.Snout short, somewhat conical, overhanging mouth. First dor-sal fin short-based, triangular and high, with a strong spine infront. Second dorsal fin long and low. Pectoral fins not reach-ing to pelvic fin base when laid back. Anal fin continuous withcaudal fin.
Biology: Benthopelagic on continental slopes and down toabyssal plains. At Lucky Strike, specimens were observed swim-ming gently above the bottom and off the hydrothermal ventfield. Carnivorous. At Lucky Strike, preys included the ventendemic mussel, Bathymodiolus azoricus. Oviparous.
Distribution: General: Western Atlantic: from Davis Straitand off Newfoundland to Cape Cod. Eastern Atlantic: Den-mark Strait, Rockall Trough, northern Bay of Biscay and off thecoast of Portugal. Possibly a much wider Distribution thanrecords show. Mid-Atlantic Ridge vents: Lucky Strike, MountSaldanha, Famous Segment, Rainbow. Depth range: 300 to2400 m.
References:
HARDY G.S. & M. STEHMANN (1990) Arch. Fischereiwiss. 40(3): 229-248.MARQUES A. & F. PORTEIRO (2000) Copeia 2000(3): 806-807.MØLLER P.R., KULLBERG T. & O.A. JØRGENSEN (2004) Cybium 28(1): 55-60.STEHMANN M. & D.L. BÜRKEL (1984) in WHITEHEAD P.J.P., BAUCHOT M.-L., HUREAU J.-C., NIELSEN J. & E. TORTONESE (Eds.) Fishes of the North-eastern Atlantic
and the Mediterranean 1: 212-215.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 491
1: Habitus; by H. Encarnação © MMF, 2000.
Chordata, Vertebrata, Chondrichthyes, Chimaeriformes, Chimaeridae
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Hydrolagus pallidus HARDY & STEHMANN, 1990 “pallid ghost shark”
References:
HARDY G.S. & M. STEHMANN (1990) Arch. Fischereiwiss. 40(3): 229-248.MØLLER P.R., KULLBERG T. & O.A. JØRGENSEN (2004) Cybium 28(1): 55-60.SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 492
1: Habitus; by H. Encarnação © MMF, 2002.
2: At Rainbow, Mid-Atlantic Ridge; cruise Atos © Ifremer.
Chordata, Vertebrata, Chondrichthyes, Chimaeriformes, Chimaeridae
Size: To about 1300 mm total length.
Color: Uniform creamy to light greyish.
Morphology: Body greatly tapering from a massive head andtrunk to a pointed caudal fin with a short filament at its tip.Snout short, somewhat conical, overhanging mouth. First dor-sal fin short-based, triangular and high, with a strong spine infront. Second dorsal fin long and low. Pectoral fins not reach-ing to pelvic fin base when laid back. Anal fin continuous withcaudal fin.
Biology: Benthopelagic on continental slopes and down toabyssal plains. At Lucky Strike, specimens were observed swim-ming gently above the bottom and off the hydrothermal ventfield. Carnivorous, possibly feeding on small fishes and inverte-brates. Oviparous.
Distribution: General: Western Atlantic: Davis Strait, Cana-da, Bear Seamount, New England; Eastern Atlantic: Green-land, off Iceland, from the southern Bay of Biscay to off west-ern Scotland and Mid-Atlantic Ridge: Lucky Strike, MenezHom, Mount Saldanha, Rainbow. Depth range: 1200-2075 m.
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Dysommina rugosa GINSBURG, 1951
Size: Up to 352 mm total length.
Morphology: Naked ilyophine eel with small broad-based pec-toral fins, their tips not sharply pointed. Dorsal-fin origin for-ward, about one head length behind a vertical through the gillslits. Fleshy, papillose snout slightly overhangs tip of lower jaw.Gill slits separate, small (about equal to diameter of eye), cres-centic, oriented nearly vertically on the ventrolateral surface.Vertebrae 127-134.
Biology: Benthopelagic; carnivorous, feeding on shrimps andpossibly other invertebrates. Oviparous. At Vailulu’u, living increvices and holes in a low-temperature (10°C) diffuse vent
field with substantial coverage of microbial mat. Not found innon-venting habitats at the same depth nearby. Swims shortdistances into the water column, probably to feed.
Distribution: General: Widespread at tropical latitudes. West-ern Atlantic Ocean, from off the Carolinas to the Caribbean.Southwestern Indian Ocean, Canal of Mozambique. PacificOcean, Hawaii. At vents, this species has been reported onlyfrom warm-water vents at the peak of a volcanic cone, Nafan-ua, at Vailulu’u Seamount, Samoan Archipelago. Depth range:260-775 m.
Reference:
ROBINS C.H. & C.R. ROBINS (1989) in: BÖHLKE E. (Ed.) Fishes of the Western North Atlantic. Part 9(1): 207-253.
C. YOUNG & M. BISCOITO Denisia 18 (2006): 493
1: Habitus; from ROBIN & ROBIN (1989).
2: In situ view at Vailulu’u Seamount; by C.Young © OIMB.
Chordata, Vertebrata, Osteichthyes, Anguilliformes, Synaphobranchidae
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Ilyophis saldanhai KARMOVSKAYA & PARIN, 1999
R. CAUSSE, M. BISCOITO & P. BRIAND Denisia 18 (2006): 494-495
1: Holotype; from KARMOVSKAYA & PARRIN (1999).
Chordata, Vertebrata, Osteichthyes, Anguilliformes, Synaphobranchidae
Size: Total length 410 mm (Atlantic specimens), and 335 mm(Pacific specimen).
Color: Uniformly brownish.
Morphology: Body elongate, compressed and naked. Origin ofdorsal fin above the 11th pore of the lateral line and behindend of pectoral fin, origin of anal fin after the 32nd pore of thelateral line, both fused with caudal fin (fig. 1). Head well dif-ferentiated from the rest of the body, its profile slightly convexat level of posterior margin of orbit. Head 3.3 times in pre-anallength. Snout length moderate (2.65 times in head). Fleshy tipof snout with two pairs of well-developed plicae (fig. 2). Twosharply pronounced medial folds descending from tip of snout;at its lower edge, then bending horizontally to the right and leftof median line of snout. Two other shorter lyre-shaped folds sit-uated laterally to two median folds. Supra-orbital canal withthree pores, postorbital canal with two pores, infra-orbital canalwith five pores, preoperculo-mandibular canal with eight pores,six before the commisure, the seventh at the level of the com-misure and the eigth on the preopercle. Mouth opening
straight, its rictus situated behind a vertical through posteriormargin of orbit. Mouth cavity covered with longitudinal foldsbearing numerous papillae that differ in size and shape. Teethin jaws conical, pointed, slightly curved, bent inside, and close-ly set in rows forming a band on each jaw. Teeth increasing insize in inner rows and medially on each jaw. Teeth on palatelarger. Teeth on vomer forming two irregular longitudinal rows(Fig. 2). Eyes small, round, covered with a thin membrane,their diameter 6.7 times in head and 2.5 times in snout. Ante-rior nostrils tubular, directed forward, bearing small flaps.Openings of posterior nostrils simple and round, their indentedmargins slightly turned out. Gill slits semicircular, situated hor-izontally at lower part of head before bases of pectoral fins(CAUSSE et al. 2005; KARMOVSKAYA & PARIN 1999).
Biology: This species is benthopelagic, associated with ventcommunities. Carnivorous, feeding on hydrothermal vent in-vertebrates (small shrimps and polychaetes).
Distribution: East Pacific Rise: 21°S, hydrothermal vent siteGromit, and MAR: Broken Spur vent field.
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References:
CAUSSE R., BISCOITO M. & P. BRIAND (2006) Cybium 29(4): 413-416.KARMOVSKAYA E.S. & PARIN N.V. (1999) J. Ichthyol. 39(5): 353-362.NIELSEN J. & D.C. COHEN (2005) Cybium 29(4): 395-398.PARIN N.V. (1995) Vopr. Ihtiol. 35(5): 698-701.
2: In vivo, after collection from East Pacific Rise: 21°S, Gromit site; cruise Biospeedo, by Briand © Ifremer.
3: In situ, cruise Biospeedo © Ifremer.
495
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Thermobiotes mytilogeiton GEISTDOERFER, 1991
Size: Up to 247 mm.
Color: In formalin pale brownish, beige in vivo with abdomendarker.
Morphology: Eel-shaped, no scales, no pectoral fins, lateral linepresent. Dorsal fin origin behind level of anus. Teeth on bothjaws, small and needle-like. Vertebrae 132.
Biology: Dwelling among mussels.
Distribution: Lau Back-Arc Basin: Valu Fa Rise.
References:
GEISTDOERFER P. (1991) C. R. Acad. Sci. Paris, Sér. III 312: 91-97.MEUNIER F.J. & P. GEISTDOERFER (1991) Cybium 15: 83-87.
P. BRIAND Denisia 18 (2006): 496
1: Total view; by P. Briand © Ifremer. 2: Anterior part in lateral view; by P. Briand © Ifremer.
4: In situ view of a specimen on a mussel bed of Bathymodio-lus brevior, at Lau Basin; cruise Biolau © Ifremer.
3: Anterior end in ventral view; by P. Briand © Ifremer.
Chordata, Vertebrata, Osteichthyes, Anguilliformes, Synaphobranchidae
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Gaidropsarus RAFINESQUE, 1810
Size: To about 360 mm standard length.
Color: Generally reddish pink with a distinct white mark oncheeks.
Morphology: Elongate gadoid fish with two dorsal fins, the firstwith a moderately elongate first finray, followed by fine shortrays. A single anal fin. Second dorsal and anal fins long andmoderately high. Three barbels, one on chin and one on eachanterior nostril. Pre-anal length longer than post-anal (exclud-ing the caudal fin). Snout longer than twice the eye diameter.Forty seven vertebrae (16+31).
Remark: A specimen was collected with the “Nautile” sub-mersible during Diva 2 cruise and its description is in press.Other specimens caught in the Bay of Biscay and the RockallTrough, currently under study, may eventually prove to belongto the same yet undescribed species.
Biology: Very few data. Benthic. The specimens observed atLucky Strike were usually inside crevices, on the mussel beds,well inside the active hydrothermal vent field. Carnivorous,feeding on hydrothermal vent crustaceans (alvinocarididshrimps).
Distribution: Mid-Atlantic Ridge: Lucky Strike.
Reference:
SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 497
1: By Luiz Saldanha © LMG, 1986.2: At Lucky Strike; cruise Seahma © FCT & Ifremer.
3: At Lucky Strike; cruise Diva © FCT & Ifremer. 4: Anterior part in dorsal view; by P. Briand © Ifremer.
Chordata, Vertebrata, Osteichthyes, Gadiformes, Lotidae
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Lepidion schmidti SVETOVIDOV, 1936
References:
COHEN D.M. (1984) in WHITEHEAD P.J.P., BAUCHOT M.L., HUREAU J.-C., NIELSEN J. & E. TORTONESE (Eds.) Fishes of the North-eastern Atlantic and the Mediter-ranean 2: 713-723.
SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 498
3: At Lucky Strike vent field; cruise Marvel © Ifremer.
1: At Rainbow vent field; cruise Atos © Ifremer.
2: At Lucky Strike vent field; cruise Victor Première © Ifremer.
Chordata, Vertebrata, Osteichthyes, Gadiformes, Moridae
Size: Up to 1230 mm standard length.
Morphology: Head conical and robust. Posterior nostril aheadof erye. Chin barbel present, as long as snout. Orbit 3.5-5.8 inhead length. Second ray of first dorsal fin very elongate. Later-al line well marked until near the origin of caudal peduncle.
Remark: One specimen caught with the “Nautile“ at Rainbowand others at Lucky Strike with bottom long lines.
Biology: Probably benthopelagic on continental slope. AtRainbow, Lucky Strike and Menez Gwen, specimens werefound lying on the bottom, amongst rocks or in crevices, in theinner periphery of the active fields (over brown sulphide de-posits).
Distribution: General Eastern North Atlantic: in the Bay ofBiscay and West of Ireland; Pacific Ocean: Sagami Bay, Japan;Mid-Atlantic Ridge: Menez Gwen. Depth range: 900-2300 m.
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Cataetyx laticeps KOEFOED, 1927
Size: To about 650 mm standard length.
Color: Greyish brown.
Morphology: Body elongate. Head dorsoventrally flattened,with a strong opercular spine. Dorsal and anal fins confluentwith caudal. Dorsal finrays 91-107; anal finrays 74-87; pectoralfinrays 22-29; pelvic finrays 1.
Remarks: Specimens were collected with the submersible atLucky Strike. A second species of this genus was recorded fromRainbow (a single specimen caught in a fish trap) and so farcannot be assigned to any of the previously known species ofCataetyx.
Biology: Benthic species. At Lucky Strike, specimens occurringoften in pairs on sulphide deposits amongst shells of dead mus-sels, on the inner border of the hydrothermal field. Carnivo-rous. At Lucky Strike, feeding on hydrothermal vent crus-taceans (crab Segonzacia mesatlantica, alvinocaridid shrimps).Viviparous.
Distribution: Generally known from a few localities in theNortheast Atlantic and western Mediterranean. Found also inthe Azores and along West Africa to the Cape of Good Hope.Possibly also in the Gulf of Mexico. Mid-Atlantic Ridge:Menez Gwen, Lucky Strike, Mount Saldanha, Menez Hom, andRainbow. Depth range: 900-2830 m.
References:
NIELSEN J.G. (1984) in WHITEHEAD P.J.P., BAUCHOT M.L., HUREAU J.-C., NIELSEN J. & E. TORTONESE (Eds.) Fishes of the North-eastern Atlantic and the Mediter-ranean 3: 1153-1157.
SALDANHA L. (1994) Cybium 18(4): 460-462.SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 499
1: At Lucky Strike, Tower Eiffel vent field; cruise Exomar ©Ifremer.
Chordata, Vertebrata, Osteichthyes, Ophidiiformes, Bythitidae
4: At Lucky Strike, Tower Eiffel vent field; cruise Exomar © Ifremer.
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Thermichthys hollisi (COHEN, ROSENBLATT & MOSER 1990)
Size: 218-304 mm standard length.
Morphology: A bythitid genus with joined vertical fins, shortbody and blunt snout. Body with imbricate scales and headnaked. Mouth terminal, upper jaw ending well behind eye. Eyeshorter than snout. Opercular spine strong. Palatine with teeth.Developed rakers on anterior arch 3. Rays in dorsal fin 122-124, anal fin 88, causal fin 12, pectoral fin 36-37 and ventral fin1. Vertebrae 76, precaudal 20-21.
Biology: One specimen was caught associated with hydrother-mal vents. A specimen of T. hollisi was seen eating a zoarcid fish
(Thermarces-like) with half of its length, during cruiseBiospeedo. In the video images, only the posterior half of theprey-fish has been swallowed and the anterior part is slowlymoving.
Distribution: Known from two specimens from the GalapagosSpreading Center and Southern East Pacific Rise: 17°S, siteHobbs.
References:
COHEN D.M., ROSENBLATT R.H. & H.G. MOSER (1990) Deep-Sea Res. A 37: 267-283.NIELSEN J.G. & D.M. COHEN (2005) Cybium 29(4): 395-398.
J.G. NIELSEN & D.M. COHEN Denisia 18 (2006): 500
1 top: Specimen taken in vivo, onboard, dorsal view; bottom: dorsolateral view; East Pacific Rise: 17°S, site Hobbs, cruise Biospeedo © Ifremer.
Chordata, Vertebrata, Osteichthyes, Ophidiiformes, Bythitidae
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Ventichthys biospeedoi NIELSEN, MOLLER & SEGONZAC, in press
Reference:
NIELSEN J.G. & P.R. MOLLER & M. SEGONZAC (in press) Zootaxa.
J.G. NIELSEN, MOLLER P.R. & M. SEGONZAC Denisia 18 (2006): 501
1: Couple of specimens caught at East Pacific Rise: 17°S, site Oasis; cruise Biospeedo, by P. Briand © Ifremer.
2: In situ view of several speci-mens on shimmering water, onmussel bed Bathymodiolusthermophilus, with stalkedbarnacle (Neolepas n. sp.) andsea anemaone (Chondrophelliacf. coronata); East Pacific Rise:17°S, site Oasis; cruiseBiospeedo © Ifremer.
Chordata, Vertebrata, Osteichthyes, Ophidiiformes, Ophidiidae
Size: Maximum standard length 282 mm.
Morphology: Body robust, very small, overlapping scales onhead and body, thick skin, and four indistinct lateral lines; dor-sal fin origin above tip of pectorals, basis of pelvic fins belowhind margin of opercle; head broad with blunt snout; strong op-ercular spine covered by thick skin; upper jaw ends just behindeye; teeth granular, one median basibranchial tooth patch; an-terior gill arch with 10-11 long rakers; number of rays in dorsalfin 80-89, caudal fin 8, anal fin 64-72, pelvic fin 2, pectoral fin24-25; number of vertebrae 16-17+36.
Biology: Species abundant, living in shimmering vent fluidswith temperatures between 2 and 7°C, among hydrothermalcommunity: mytilid and clam bivalves, stalked barnacles.Necrophagous.
Distribution: East Pacific Rise: 17°S, site Oasis, but probablythe same species occurs on northern and southern sites.
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Pachycara gymninium ANDERSON & PEDEN, 1988
Size: Up to about 420 mm standard length.
Color: In life dark brown, head and pectoral fin darker, almostblack. Margins of vertical fins and peritoneum black.
Morphology: Body short, deep, broader in cross section, whencompared with its congeners. Head large, ovoid. Pelvic finspresent (4.8-11.3% of head length). Mediolateral branch of lat-eral line originating in pectoral axil just posterior to verticalthrough pectoral base. Scales absent on nape, or, if present, notextending anterior to line connecting anterodorsal edges of gillslits. Vertebrae 102-109. Anal fin origin associated with verte-brae 27-31.
Biology: Benthic over brown and green mud bottoms. Carniv-orous, eating amphipods, isopods and polychaetes. Oviparous, agravid female caught in February.
Distribution: General: Northeast Pacific Ocean, off the QueenCharlotte Islands, British Columbia, south of Guadalupe Is-land, Mexico and Gulf of California. Juan de Fuca Ridge: Cryp-to Vent Field, Axial Seamount; Endeavour Segment; Ham-mond’s Hell vent. Depth range: 1575-3219 m.
References:
ANDERSON M.E. & A.E. PEDEN (1988) Proc. Calif. Acad. Sci. 46(3): 83-94.TUNNICLIFFE V., MCARTHUR A.G. & D. MCHUGH (1998) Adv. Mar. Biol. 34: 353-451.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 502
1: Holotype, 422 mm standard length; by P. Drukker-Brammall, 1988.
Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae
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Pachycara rimae ANDERSON, 1989
Size: Up to 403 mm standard length.
Color: Uniformly light brown, eye and abdomen bluish.
Morphology: Head deep, rounded, somewhat shorter than sim-ilarly sized congeners. Pelvic fins nublike, of two soft rays. Ver-tebrae 93. Dorsal-fin rays 86, anal-fin rays 70. Lateral line ofmediolateral branch only. Dorsal fin origin associated with ver-tebra 8. Pseudobranchs absent.
Remark: Data available are not sufficient to determinewhether this species is endemic to hydrothermal vent environ-ments or not.
Distribution: Galapagos Spreading Center. Known only fromthe holotype.
References:
ANDERSON M.E. (1989) Proc. Calif. Acad. Sci. 46(10): 221-242.COHEN D.M. & R.L. HAEDRICH (1983) Deep-Sea Res. 30(4A): 371-379.COHEN D.M., ROSENBLATT R.H. & R.L. HAEDRICH (1985) Biol. Soc. Wash. Bull. 6: 229-230.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 503
1: Holotype, 403 mm standard length; © K. Klitz, 1989.
Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae
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Pachycara saldanhai BISCOITO & ALMEIDA, 2004
Size: Up to 256 mm total length.
Color: In life, light brownish grey, head, dorsal and anal finsdarker. When preserved light brown, body with conspicuouswhitish scale pockets.
Morphology: Body elongate and compressed, scaled. Mouthsubterminal, no crests on chin. Dorsal fin origin over pectoralfins. Pelvic fins present. Lateral line with two branches. Dorsalfin rays 108-115, anal fin rays 90-95, vertebrae 117-123.
Biology: Benthic, not very numerous over sulphide depositsand among mussels inside the active field. Food: hydrothermalvent crustaceans.
Distribution: Up to present restricted to Mid-Atlantic Ridge:Rainbow.
Reference:
BISCOITO M. & A.J. ALMEIDA (2004) Copeia 3: 562-568.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 504
1: Habitus; by H. Encarmaçao © MMF, 2002.
2: At Rainbow vent field; cruiseMarvel © Ifremer.
3: At Rainbow vent field; cruise Marvel © Ifremer.
Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae
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Pachycara thermophilum GEISTDOERFER, 1994
Size: Up to 388 mm total length.
Color: Light brown with pinkish hues. Border of fins darker.
Morphology: Eelpout shaped. Pelvic fin rays 2. Mediolaterallateral line only, originating just behind posterior-most postor-bital pore. Scales absent on nape. Dorsal fin rays 105-107. Analfin rays 86-89. Total vertebrae 113-114. Dorsal fin origin asso-ciated with vertebrae 7-8.
Biology: Benthic, in areas with active smokers, swimming insea water 5-20°C, away from rocks and along smokers nearcrowds of shrimps Rimicaris exoculata and Chorocaris chacei.Carnivorous, feeding on hydrothermal vent shrimps.
Distribution: Mid-Atlantic Ridge: Snake Pit and TAG. Depthrange: 3500-3700 m.
References:
ANDERSON M.E. & H. BLUHM (1996) Trans. R. Soc. Afr. 51: 219-227.GEISTDOERFER P. (1994) Cybium 18(2): 109-115.PARIN N.V. (1995) J. Ichthyol. 35(9): 328-332.
P. GEISTDOERFER Denisia 18 (2006): 505
2: In situ at Snake Pit © Ifremer.
1: Habitus; by E. Heemstra © JLB Smith Inst. Ichth., 1996.
3: Habitus; by P. Briand © Ifremer.
Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae
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Pyrolycus manusanus MACHIDA & HASHIMOTO, 2002
Size: Up to 170 mm total length.
Color: Light brown or beige.
Morphology: Eeelpout shaped. Suborbital bones 6, canal with6-7 pores; flesh gelatinous; dermal papillae absent from head;gill slit large, extending ventrally beyond pectoral fin base;pelvic fins present, each with 2-3 rays; scales, lateral line andpseudobranch absent; pyloric caeca present; oral valve weak;interorbital pore absent; postorbital pores 3; occipital pores 1;palatopterygoid series weak; vomerine and palatine teeth pres-ent; pectoral fin rays 16-17; caudal fin rays 8-10; vertebrae 22-23+56-59=78-81.
Biology: Seen in areas close to active smokers and/or in andaround diffuse vent fluids. Carnivorous, alvinocaridid shrimpswere found in stomach contents of some specimens.
Distribution: Manus Back-Arc Basin: Pacmanus and Desmossites.
References:
HASHIMOTO J., OHTA S., FIALA-MÉDIONI A., AUZENDE J.-M., KOJIMA S., SEGONZAC M., FUJIWARA Y., HUNT J.-C., GENA K., MIURA T., KIKUCHI T., YAMAGUCHI T., TODA T.,CHIBA H., TSUCHIDA S., ISHIBASHI J., HENRI K., ZBINDEN M., PRUSKI A., INOUE A., KOBAYASHI H., BIRRIEN J.-L., NAKA J., YAMANAKA T., LAPORTE C., NISHIMURA K.,YEATS C., MALAGUN S., KIA P., OYAIZU M. & T. KATAYAMA (1999) InterRidge News 8: 12-18.
MACHIDA Y. & J. HASHIMOTO (2002) Ichthyol. Res. 49: 1-6.
J. HASHIMOTO Denisia 18 (2006): 506
1: Holotype; from MACHIDA & HASHIMOTO (2002).
2: At Manus Back-Arc Basin; cruise Bioaccess © JAMSTEC. 3: At Manus Back-Arc Basin; cruise Bioaccess © JAMSTEC.
Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Thermarces cerberus ROSENBLATT & COHEN, 1986
Size: Up to 370 mm total length.
Color: Whitish or pinkish.
Morphology: Eelpout shaped, head and body laterally com-pressed, naked, covered with mucous, without scales. Lips dis-tinct, thick and fleshy, continuous and smooth; oral valves ob-solete. Head pores large and conspicuous. Occipital pores ab-sent. Teeth in both jaws stout, conical and pointed. No lateralline. Pelvic fins absent. Pectoral fins small, rounded, with rayscovered by thick skin. Vertebrae: 93-97.
Remarks: 1. Another close species, T. andersoni ROSENBLATT &COHEN, 1986, was described from Galapagos Rift, but its syn-onymy with T. cerberus is debated. 2. A new species of Ther-
marces was found at East Pacific Rise: 9°N, showing strong mor-phological and color differences. It is being described elsewhere(M. Biscoito & M. Segonzac, unpublished data).
Biology: Observed in areas of active smokers, associated withRiftia pachyptila or on smoker walls. Very often stays in diffuseventing areas, amongst rubble. Carnivorous, feeding on smallinvertebrates, mainly amphipods and limpets, but also poly-chaetes. Occasionally they have been seen biting the gills of R.pachyptila.
Distribution: Galapagos Spreading Center; East Pacific Rise:9°N, 13°N and 21°N; probably also at South East Pacific ventsites.
M. BISCOITO & M. SEGONZAC Denisia 18 (2006): 507-508
1: Fresh specimen collected at East Pacific Rise: 13°N; by P. Briand © Ifremer.
Chordata, Osteichthyes, Perciformes, Zoarcidae
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
2: In situ specimens from East Pacific Rise: 13°N, among vestimentiferan (Riftia pachyptila) and mussel bed of Bathymodiolusthermophilus; cruise Phare © Ifremer.
References:
GEISTDOERFER P. (1986) Bull. Mus. Natl. Hist. Nat. Paris, Sér. 4A 8: 969-980.GEISTDOERFER P. (1996) Oceanol. Acta 19(5): 539-548.ROSENBLATT R.H. & D.M. COHEN (1986) Trans. San Diego Soc. Nat. Hist. 21: 71-79.
508
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Careproctus hyaleius GEISTDOERFER, 1994
Size: Up to 112 mm standard length.
Color: Whitish to pinkish; translucent.
Morphology: Small-sized fish, tadpole-shaped, with a roundand globular head accounting for 22% of total length.Branchial aperture small (32% of head length) and located inthe upper part of the body. Teeth simple, hooked and all simi-lar. Body flaccid, covered by thick layer of mucous. Bare andfragile skin. Pelvic fins modified into a sucking disc.
Biology: Frequently observed at the boundary of active ventfields and occasionally among Riftia or within diffuse ventingareas.
Distribution: East Pacific Rise: 9° N to 13° N.
References:
BISCOITO M., SEGONZAC M., ALMEIDA A.J., GEISTDOERFER P., TURNSIPSEED M. & C. VAN DOVER (2002) Cah. Biol. Mar. 43: 359-362.GEISTDOERFER P. (1994) Cybium 18(3): 325-333.
P. BRIAND Denisia 18 (2006): 509
1: Dorsal view; by P. Briand © Ifremer.
2: Ventral view; by P. Briand © Ifremer. 3: View in situ of a specimen (righ), with a zoarcid Thermarcescerberus, tubeworms Riftia pachyptila and serpulids Lami-natubus alvini; East Pacific Rise: 13°N. Cruise Hot 96 © Ifremer.
Chordata, Vertebrata, Osteichthyes, Scorpaeniformes, Liparidae
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Trachyscorpia cristulata echinata (KOEHLER, 1896) “spiny scorpionfish”
Size: Up to 500 mm standard length.
Color: Reddish, dorsal fin with bluish pigment.
Morphology: Head large. Orbit large, much wider than snoutlength. Well developed spines on head. Dorsal fin ray with 12spines and 8-9 rays. Pectoral fin with characteristic shape(longest rays near upper part of fin) and with 20-23 rays.
Biology: Benthic. At Menez Gwen on pillow lavas, off the ac-tive sites. Carnivorous. One anguilliform fish found inside thestomach of a specimen collected at Menez Gwen. Oviparous.
Distribution: General: Eastern Atlantic, from Ireland south-ward to Mauritania. Mid-Atlantic Ridge: Menez Gwen.
References:
HUREAU J.-C. & N.I. LITVINENKO, (1984) in WHITEHEAD P.J.P., BAUCHOT M.-L., HUREAU J.-C., NIELSEN J. & E. TORTONESE (Eds.) Fishes of the North-eastern Atlanticand the Mediterranean 3: 1211-1229.
SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 510
1: Menez Gwen; cruise Saldanha © Ifremer & FCT.
Chordata, Vertebrata, Osteichthyes, Scorpaeniformes, Sebastidae
2: Habitus; from HUREAU & LITVIENKO (1984).
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Coryphaenoides armatus (HECTOR, 1875) “armed grenadier”
Size: To at least 800 mm total length.
Color: Generally brownish to reddish brown, fin membranesbrownish, mouth and gill cavity blackish.
Morphology: Ventral parts of head mostly naked, includingsnout, most ventral surfaces of suborbital space, ventral preop-ercular margin and anterior part of mandible. Premaxillaryteeth in one or two rows, one row on mandible. Inner gill rak-ers on first arch 11 to 14. First dorsal fin with two spines and 8-10 rays, pectoral fins rays I+17-21. Pelvic fins 10. Anus close toanal fin origin. No light organ.
Biology: Benthopelagic. Carnivorous, feeding on benthic ani-mals (amphipods, isopods, cumaceans), also pelagic animals(mysids, euphausids and other crustaceans, echinoderms,cephalopods and fishes). Two specimens were collected withbottom long lines at Lucky Strike and one at Snake Pit, the lat-ter had Rimicaris exoculata in its stomach. Oviparous.
Distribution: General: Worldwide. Marginal to the SouthernOcean. Mid-Atlantic Ridge: Lucky Strike, Snake Pit, Rainbow;Galapagos Spreading Center; East Pacific Rise: 9°N. Depthrange: 282-4700 m.
1: Habitus; from GÜNTHER (1887).
2: In situ from Mid-Atlantic Ridge;cruise Atos © Ifremer.
Chordata, Vertebrata, Osteichthyes, Macrouridae
References:
COHEN D.M. (1990) in QUÉRO J.-C., HUREAU J.-C., KARRER C., POST A. & L. SALDANHA (Eds.) Check-list of the Fishes of the Eastern Tropical Atlantic 2: 541-563.
COHEN D.M., T. INADA, T. IWAMOTO & N. SCIALABBA (1990) FAO Species Catalogue. FAO Fish. Synop. 125(10): 1-442.GEISTDOERFER P. (1988) Oceanol. Acta, 8 n° sp.: 125-130.GEISTDOERFER P. (1991) C. R. Acad. Sci. Paris, Sér. III 312: 91-97.GÜNTHER A. (1987) Challenger Reports, Zool. 22: 1-268.MARQUES A & A.J. ALMEIDA (2000) InterRidge News 9(2): 16-17.MERRETT N.R. & N.B. MARSHALL (1981) Progr. Oceanogr. 9: 185-244.
M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 511
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Major known deep-sea hydrothermal vent fieldsOcean Ridge / BAB Field Lat. Long. Depth (m) SitesAtlantic Jan-Mayen Ridge Kolbeinsey 67°05’N 18°43’W 90
Mid-Atlantic Ridge Menez Gwen 37°51’N 31°31’W 840-865Lucky Strike 37°17’N 32°16’W 1620-1730 Eiffel Tower, Statue of Liberty,
Sintra, Bairro-Alto, Isabel, Petitchimiste, Fantôme, Montségur,Hélène, Pico, Nuno, Elisabeth
Mount Saldanha 36°33’N 33°28’W 2300Rainbow 36°13’N 33°54’W 2260-2350Lost City 30°07’N 42°07’W 750-900Broken Spur 29°10’N 43°10’W 3050-3875 Saracen’s Head, Wasp’s nest,
SpireTAG 26°08’N 44°49’W 3635-3670 Mir zone, Kremlin Snake Pit 23°23’N 44°58’W 3480 Moose, Beehive, The NailLogatchev 14°45’N 44°58’W 2930-3020 Irina Mound, Main Mound,
Anna-Louise Ashadze-2 12°59’N 44°54’W 3200-3300Ashadze-1 12°58’N 44°52’W 4100-4200Ascension 4°48’S 12°22’W 3100
Mediterranean Gulf of Naples Capo Misseno 40°50’N 14°05’E coastalTyrrhenian Sea Capo Palinuro 40°01’N 15°16’E 9Aegian Sea Milos 36°40’N 24°23’E 0-115
Indian Central Indian Ridge Edmond Vent Field 23°52’S 69°35’E 3290-3320Kairei Vent Field 25°19’S 70°02’E 2415-2460
Gulf of Aden 11°57’N 43°40’E 1400-1600North East Pacific Southern Explorer Magic Mountain 49°46’N 130°15’W 1797 Zooarium, Dead Chimney,
Ridge Zoo Chimney Field Recordbreaker, ZeusMagic Mountain 49°46’N 130°15’W 1778 Stump, Tubeworm, Beercan, Merlin Mound Limpet, AnhydriteMagic Mountain 49°46’N 130°16’W 1778 Einstein, Eastern Island, Mystic Mound Ridge TopMagic Mountain 49°46’N 130°15’W 1785 Anhydrite 1-4, Magestic, Magestic Chimney Obelisk, Digit Field
Juan de Fuca Ridge Middle Valley 48°27’N 128°42’W 2400Endeavour Segment 48°N 129°04’W 2200Sasquatch FieldEndeavour Segment 47°58’N 129°04’N 2200 GrendlSalty Dawg FieldEndeavour Segment 47°57’N 129°06’W 2200 S & M, Peanut, Puffer, HulkMain EndeavourEndeavour Segment 47°57’N 129°07’W 2200-2220 Godzilla
High Rise FieldEndeavour segment 47°55’N 129°08’W 2200 Faulty Towers ComplexMothra Vent FieldCoAxial Segment 46°19’N 129°42’W 2291 Source site, Floc site , Flow site,
Mongo vent, Beard vent, Churchvent
Axial Volcano 45°59’N 130°02’W 1580 Shepherd ventCASM Vent FieldAxial Volcano 45°55’N 129°59’W 1530-1544 Magnesia, Oxide, Blue nose, Easy, Lava Flow 98 45°57’N Milky, Old worms, Minisnow, nSRZ Roof, The Pit, Old flow, Nascent,
Cloud, Snail, Snow, Circ, Castle,Village, Pillar, Joystick, Iron City,Bag City, Crevice, Coquille ventfield, Dying
D. DESBRUYÈRES Denisia 18 (2006) 513–517
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Ocean Ridge / BAB Field Lat. Long. Depth (m) SitesAxial Volcano 45°56’N 130°00W 1547 White, Virgins Daughter, Virgin, Ashes Vent Field Marshmallow, Mushroom, Gol-
lum, Inferno, Daves, Hairdo,Medusa, Ropos, Phoenix, Styx,Crack, Steve Mound
Cleft Segment 44°39’N 130°15’W 2270 Fountain(Southern Juan de Fuca)
Gorda Ridge Sea Cliff Vent Field, 42°15’N 126°42’W 2700Northern Gorda RidgeEscanaba Trough 41°00’N 127°29’W 3250
East Pacific Southern California Palos Verdes 33°42’N 118°19’W 1-10White Point
Gulf of California Guaymas Basin 27°00’N 111°24’W 2000East Pacific Rise 21°N 20°50’N 109°06’W 2615 Clam Acres (Northern) 13°N 12°48’N 103°56’W 2635 Pogosud, Pogonord, Totem,
12°50’N 103°57’W Genesis, Elsa9°N (Venture Hydro- 9°48’N 104°17’W 2500 Biovent, M-vent, Riftia Field, thermal Field) 9°51’N Mussel Bed, East Wall, Tube-
worm Pilar, Tevnia Hole, Brasou-cade, Tica
Galapagos Ridge Galapagos Spreading 00°47’N 86°07’W 2450-2490 Rose Garden, Mussel bed, Small Center 00°49’N 86°13’W Fry, Garden of Eden
East Pacific Rise 7°S 7°21’S 107°47’W 2735-2752 White Christmas, Sarah’spring, (Southern) 7°25’S Petite cheminée, Last hope
14°S 13°58’S 112°28’W 2623-2635 Pagoda, Smokin’Shank,13°59’S Alvinella chimney, Lucky Eric
17°24’S 17°24’S 113°12’W 2575-2590 Stanley, Robbie Roost, S-vent, 17°25’S Nadir, Oasis, Rehu Marka
17°34’S 17°34’S 113°14’W 2590-2600 Calvin, Hobbs, Suzie, Miss 17°35’S Wormwood
18°S 18°24’S 113°24’W 2636-2680 Fromveur, Cathedral, Sojourn,18°36’S Animal Farm
21°S 21°25’S 114°17’W 2800-2850 Les trois diffuseurs, Bordreaux, 21°34’S Grande cheminée, Krasnov, Gro-
mit, Brandon, Preston.23°S 23°32’S 115°34’W 2598 Needles
Pacific-Antarctic 31°S 31°09’S 111°55’W 2330 Fred’s Fortress, Nolan’s NookRidge
Saguaro Field 31°51’S 112°02’W 2235Foundation 37°35’S 110°35’W 2220German Flats - 38°S 37°47’S 110°55’W 2216 Sebastian’s Steamer, Pâle Etoile,
Annie’s Anthill, Serpulid CityWest Pacific Bering Sea Piyp Submarine 55°41’N 167°27’E 352-450
VolcanoKuril Islands Kraternaya Bight 47°31’N 152°49’E 49Japan Kagoshima Bay 31°39’N 130°48’E 82-110Izu Ogasawara Arc Myojin Knoll 32°07’N 139°51’E 1300-1400(Bonin)
Sumisu Caldera 31°28’N 140°04’E 670-690Suiyo Seamount 28°34’N 140°39’E 1367Kaikata Seamount 26°42’N 141°04’E 460-910Nikko Seamount 23°06’N 142°40’E 430-600
Okinawa Trough Minami-Ensei Knoll 28°24’N 127°38’E 640-720127°39’E
North Iheya Knoll 27°47’N 126°54’E 990-1070Iheya Ridge 27°33’N 126°59’E 1400-1430Izena Cauldron 27°16’N 127°04’E 1310-1580
127°05’EHatoma Knoll 28°24’N 123°50’E 1470-1520
Okinawa Arc Northeastern Taiwan 24°50’N 121°60’E 20-180 Tashi fishing groundsMariana Mariana Trough 18°12’N 144°42’E 3595Back-Arc Basin
Anemone Heaven 18°02’N 144°45’E 3676
514
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
515
Ocean Ridge / BAB Field Lat. Long. Depth (m) SitesSnails Pit
Alice Spring Field 13°23’N 143°55’E 1450 Forecast vent fieldMariana Volcanic Arc Daikoku Seamount 21°19’N 144°12’E 378 Fish Spa, AA, Bluff, Bottomless
pit, Bubbles, Cactus flower, Cra-ter tuff luck, Fish crack
Kagusa 2 Seamount 21°37’N 143°38’E 387-393 Barnacle Boulders, Mat Ridge,Crabe slope, Cracked vent, FlatBottom, Hairy mat, Hairy rock,Pinnacle, SW eels, Whale rock,Yellow overlord
North West Eifuku 21°29’N 144°02’E 1573-1604 More Mussels, Top Towers, Seamount Champagne, Near Fouling, Fou-
ling Heights, Sulphur spicules,Bacterial balls, Cliff house, Floc,Ice cream, Ice Fall, Ice sheet,Mussel mound, Walter grenade
East Diamante 15°56’N 145°40’E 179-457 Barnacle Beach, Diamante Fo-Seamount rest, Central Cone (Boulder
Vent), Central Cone (Aquarium),Fe-Mn Crust, Intense Diffuse,Floc Storm, Basket case, Five to-wers, Limpets, Snail flange, Softy
North West Rota 14°36’N 144°46’E 516-599 Brimstone Pit, Cnidaria area, Seamount Dark sands, Eastern Fault, Fault
Shrimp, Flank vent, Gastros,High Flow, Iceberg, Loose sands,Pit Edge, Scarp top, Shimmeringsands, Shimmering vent, Shrim-p’s peak, Snowcone, Sulfursand mats, White mat, Whitewall, Yellow granules, Yellowtop
Tabar-Feni Volcanic Edison Seamount 03°01’S 152°03’E 1450Fore-ArcManus Back-Arc Vienna Woods 03°09’S 150°16’E 2500Basin
Desmos Cauldron 03°41’S 151°52’E 1860-2000 Onsen, Genge-baPacmanus Complex 03°43’S 151°40’E 1700 Field D, Mont Blanc, Field F,
Field E, Black smoker, ChimneyForest, Field G
Intra-plate Loihi Seamount 18°57’N 155°16’W 969seamounts
Vailulu’u Seamount 14°13’S 169°04’W 600-1000North Fiji White Lady 16°59’S 173°55’E 2000 LHOS, White Lady, Back-Arc Basin Kaiyo Chimney
Mussel Valley 18°49’S 173°29’E 2700Lau Back-Arc Basin Kilo Moana 20°03’S 176°08’W 2618
Tow Cam 20°06’S 176°34’W 2703ABE 20°46’S 176°11’W 2148Tu’i Malila 21°59’S 176°34’W 1887Vaï Lili 22°13’S 176°37’W 1764-1707Hine Hina 22°32’S 176°43’W 1832-1887
Kermadec-Arc Macauley Caldera 30°12’S 178°27’E 200-504Giggenbach 30°02’S 178°43’E 144-175SeamountBrothers Seamount 34°52’S 179°04’E 1197Rumble III Seamount 35°44’S 178°28’E 200Rumble V Seamount 36°08’S 178.11’E 367-755
Bay of Plenty Calypso Vents 37°39S 177°07’E 159-200
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Fig. 1: Atlantic-EastPacific regions.
Fig. 2: Atlantic-EastPacific sites.
516
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
517
Fig. 3: Indian-West Pacificregions.
Fig. 4: Indian-West Pacific sites.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Abbreviation Ridge, Back-Arc Basin Abbreviation FieldAzores Azores Plateau
CIR Central Indian Ridge EVF Edmond Vent Field
KVF Kairi Vent Field
EPR East Pacific Rise 21°N-23°S
Explorer Southern Explorer Ridge MM Magic Mountain
Galapagos Galapagos Ridge Galapagos Spreading Center
Guaymas Gulf of California Guaymas Basin
Gorda Gorda Ridge
Izu Izu Ogasawara Arc KS Kaikata Seamount
MK Myojin Knoll
NS Nikko Seamount
SS Suijo Seamount
JFR Juan de Fuca Ridge AV-AVF Axial Volcano, Ashes Vent Field
AV-CASM Axial Volcano, CASM Vent Field
CAS CoAxial Segment
ES Endeavour Segment
MV Middle Valley
S-JFR Southern Juan de Fuca Ridge (Cleft Segment)
Kagoshima Japan KB Kagoshima Bay
Kermadec Kermadec Arc BS Brothers Seamount
RIIIS Rumble III Seamount
RVS Rumble V Seamount
Lau Lau Back-Arc Basin VF Valu Fa Ridge
HH Hine Hina
MAR Mid-Atlantic Ridge A1 Ashadze-1
BS Broken Spur
LC Lost City
LO Logatschev
LS Lucky Strike
MG Menez Gwen
MS Mount Saldanha
RB Rainbow
SP Snake Pit
TAG TAG
Mariana Mariana Back-Arc-Basin
Manus Manus Back-Arc Basin
NFiji North Fiji Back-Arc Basin WL White Lady
MV Mussel Valley
Okinawa A Okinawa Arc NET North Eastern Taiwan
Okinawa T Okinawa Trough HK Hatoma Knoll
IR Iheya Ridge
MEK Minami-Ensei Knoll
PAR Pacific-Antarctic Ridge 31°S-38°S
Tabar-Feni Tabar-Feni Volcanic Fore-Arc ES Edison Seamount
Vailulu Intra-plate Seamounts VS Vailulu’u Seamount
Abbreviations, ridges, back-arc basins and vent fields
M. BRIGHT Denisia 18 (2006): 518
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
519
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on-v
ent
loca
tion
s41
Pori
fera
,Dem
ospo
ngia
e,Po
ecilo
scle
rida
,Cla
dorh
izid
aeCh
ondr
ocla
dia
lam
padi
glob
usV
ACE
LET,
inpr
ess
EPR:
17°S
42
Pori
fera
,Dem
ospo
ngia
e,Po
ecilo
scle
rida
,Cla
dorh
izid
aeCl
ador
hiza
abys
sico
laG
.O.S
ARS
,187
2M
AR:
LOno
n-ve
ntlo
cati
ons
44
Pori
fera
,Dem
ospo
ngia
e,Po
ecilo
scle
rida
,Cla
dorh
izid
aeCl
ador
hiza
sego
nzac
iVA
CELE
T,in
pres
sEP
R:17
°S45
Pori
fera
,Dem
ospo
ngia
e,Po
ecilo
scle
rida
,Gui
tarr
idae
Euch
elip
lum
apr
isti
naTO
PSEN
T,19
09M
AR:
LS,R
B46
Pori
fera
,Hex
acti
nella
,Lys
saci
nosi
da,C
aulo
phac
idae
Caul
opha
cus
cyan
aeBO
URY
-ESN
AU
LT&
DE
VO
S,19
88EP
R:13
°N47
Cnid
aria
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rozo
a,A
ntho
athe
cata
,Can
dela
brid
aeCa
ndel
abru
mph
rygi
umFA
BRIC
IUS,
1780
MA
R:LS
,RB;
non-
vent
loca
tion
s49
Cnid
aria
,Hyd
rozo
a,A
ntho
athe
cata
,Can
dela
brid
aeCa
ndel
abru
mse
rpen
tari
iSEG
ON
ZAC
&V
ERV
OO
RT,1
995
MA
R:SP
50
Cnid
aria
,Hyd
rozo
a,A
ntho
athe
cata
,Eud
endr
iidae
Eude
ndri
umpl
anum
BON
NEV
IE,1
898
MA
R:LO
,LS;
non-
vent
loca
tion
s51
Cnid
aria
,Hyd
rozo
a,A
ntho
athe
cata
,Tub
ular
iidae
Ecto
pleu
rala
rynx
ELLI
S&
SOLA
ND
ER,1
786
MA
R:M
G;n
on-v
ent
loca
tion
s52
Cnid
aria
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rozo
a,Le
ptot
heca
ta,H
alop
teri
dida
ePo
lypl
umar
iafl
abel
lata
G.O
.SA
RS,1
874
MA
R:M
G;n
on-v
ent
loca
tion
s53
Cnid
aria
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rozo
a,Le
ptot
heca
ta,H
ebel
lidae
Hal
isip
honi
aar
ctic
aK
RAM
P,19
32M
AR:
LO54
Cnid
aria
,Hyd
rozo
a,Le
ptot
heca
ta,L
afoe
idae
Gra
mm
aria
abie
tina
SARS
M.,
1850
MA
R:M
G;n
on-v
ent
loca
tion
s55
Cnid
aria
,Hyd
rozo
a,Le
ptot
heca
ta,L
afoe
idae
Lafo
eadu
mos
aFL
EMIN
G,1
820
MA
R:RB
;non
-ven
tlo
cati
ons
56
Cnid
aria
,Hyd
rozo
a,Le
ptot
heca
ta,L
afoe
idae
Zygo
phyl
axec
hina
taCA
LDER
&V
ERV
OO
RT,1
998
MA
R:M
G57
Cnid
aria
,Hyd
rozo
a,Le
ptot
heca
ta,L
afoe
idae
Zygo
phyl
axle
loup
iRA
MIL
&V
ERV
OO
RT,1
992
MA
R:RB
;non
-ven
tlo
cati
ons
58
Cnid
aria
,Hyd
rozo
a,Le
ptot
heca
ta,S
ertu
lari
idae
Hyd
rallm
ania
falc
ata
LIN
NA
EUS,
1758
MA
R:LS
;non
-ven
tlo
cati
ons
59
Cnid
aria
,Hyd
rozo
a,Le
ptot
heca
ta,S
ertu
lari
idae
Sert
ular
ella
tene
llaA
LDER
,185
6M
AR:
LS;n
on-v
ent
loca
tion
s60
Cnid
aria
,Hyd
rozo
a,Le
ptot
heca
ta,S
ertu
lari
idae
Sym
plec
tosc
yphu
sba
thya
lisV
ERV
OO
RT,1
972
MA
R:LS
;non
-ven
tlo
cati
ons
61
Cnid
aria
,Hyd
rozo
a,Le
ptot
heca
ta,T
iara
nnid
aeSt
egol
aria
geni
cula
taA
LLM
AN,1
888
MA
R:LS
,RB;
non-
vent
loca
tion
s62
Cnid
aria
,Hyd
rozo
a,Si
phon
opho
ra,R
hoda
liida
eTh
erm
opal
iata
raxa
caPU
GH,1
983
EPR,
Gal
apag
os63
Cnid
aria
,Scy
phoz
oa,S
taur
omed
usae
,Ele
uthr
ocar
pida
eLu
cern
aria
jane
tae
COLL
INS
&D
ALY
,200
5EP
R:21
°N-7
°S64
Cnid
aria
,Ant
hozo
a,A
ctin
iari
a,A
ctin
osto
lidae
Act
inos
tola
VER
RILL
,188
3EP
R:13
°N66
Cnid
aria
,Ant
hozo
a,A
ctin
iari
a,A
ctin
osto
lidae
Cyan
anth
eahy
drot
herm
ala
DO
UM
ENC
&V
AN
PRA
ET,1
986
EPR:
13°N
67
Cnid
aria
,Ant
hozo
a,A
ctin
iari
a,A
ctin
osto
lidae
Mar
acti
sri
mic
ariv
ora
FAU
TIN
&BA
RBER
,199
9M
AR:
A1,
SP,T
AG
68
Cnid
aria
,Ant
hozo
a,A
ctin
iari
a,A
ctin
osto
lidae
Mar
iana
ctis
byth
ios
FAU
TIN
&H
ESSL
ER,1
989
Mar
iana
69
Cnid
aria
,Ant
hozo
a,A
ctin
iari
a,A
ctin
osto
lidae
Pacm
anac
tis
hash
imot
oiLÓ
PEZ-
GO
NZÁ
LES,
ROD
RIG
UEZ
&SE
GO
NZA
C,20
05M
anus
70
Cnid
aria
,Ant
hozo
a,A
ctin
iari
a,A
ctin
osto
lidae
Para
ntho
sact
isde
nhar
togi
LÓPE
Z-G
ON
ZÁLE
S,RO
DRI
GU
EZ&
SEG
ON
ZAC,
2003
Gua
ymas
71
Cnid
aria
,Ant
hozo
a,A
ctin
iari
a,Bo
loce
roid
idae
Bolo
cero
ides
daph
neae
DA
LY,i
npr
ess
EPR:
9°N
-23°
S;PA
R:32
°S72
Cnid
aria
,Ant
hozo
a,A
ctin
iari
a,H
orm
athi
idae
Chon
drop
helli
acf
.cor
onat
a(V
ERRI
L,18
83)
EPR:
13°N
,7°S
-23°
S;PA
R:32
°S73
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Phyl
um
,C
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mil
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Mol
lusc
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leno
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res,
Cavi
belo
nia,
Sim
roth
ielli
dae
Hel
icor
adom
enia
acre
dem
aSC
HEL
TEM
A,2
000
EPR:
21°N
;17°
S;G
alap
agos
76
Mol
lusc
a,So
leno
gast
res,
Cavi
belo
nia,
Sim
roth
ielli
dae
Hel
icor
adom
enia
bisq
uam
aSC
HEL
TEM
A,2
000
EPR:
21°N
77
Mol
lusc
a,So
leno
gast
res,
Cavi
belo
nia,
Sim
roth
ielli
dae
Hel
icor
adom
enia
juan
iSCH
ELTE
MA
&K
UZI
RIA
N,1
991
Expo
rer,
Gor
da,J
FR:E
S78
Mol
lusc
a,Po
lypl
acop
hora
,Neo
lori
cata
,Isc
hnoc
hito
nida
eTh
erm
ochi
ton
undo
cost
atus
SAIT
O&
OK
UTA
NI,
1990
Oki
naw
aT:
IR80
Mol
lusc
a,Po
lypl
acop
hora
,Neo
lori
cata
,Lep
toch
iton
idae
Lept
ochi
ton
tenu
idon
tus
SAIT
O&
OK
UTA
NI,
1990
Oki
naw
aT:
IR81
Mol
lusc
a,G
astr
opod
a,Pa
telli
gast
ropo
da,N
eole
peto
psid
aeEu
lepe
dops
isvi
trea
MCL
EAN,1
990
EPR:
21°N
-17°
S;G
alap
agos
83
Mol
lusc
a,G
astr
opod
a,Pa
telli
gast
ropo
da,N
eole
peto
psid
aeN
eole
peto
psis
dens
ata
MCL
EAN,1
990
EPR:
13°N
;Gal
apag
os84
Mol
lusc
a,G
astr
opod
a,Pa
telli
gast
ropo
da,N
eole
peto
psid
aeN
eole
peto
psis
gord
ensi
sM
CLEA
N,1
990
EPR:
20°N
,5°S
;Gor
da,
84
Mol
lusc
a,G
astr
opod
a,Pa
telli
gast
ropo
da,N
eole
peto
psid
aeN
eole
peto
psis
occu
lta
MCL
EAN,1
990
EPR:
21°N
84
Mol
lusc
a,G
astr
opod
a,Pa
telli
gast
ropo
da,N
eole
peto
psid
aeN
eole
peto
psis
verr
uca
MCL
EAN,1
990
EPR:
21°N
84
Mol
lusc
a,G
astr
opod
a,Pa
telli
gast
ropo
da,N
eole
peto
psid
aePa
rale
peto
psis
ferr
ugiv
ora
WA
RÉN
&BO
UCH
ET,2
001
MA
R:LS
85
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeCl
ypeo
sect
uscu
rvus
MCL
EAN,1
989
Expl
orer
,JFR
86
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeCl
ypeo
sect
usde
lect
usM
CLEA
N,1
989
EPR:
21°N
,13°
N;G
alap
agos
86
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeG
orgo
lept
isem
argi
natu
sM
CLEA
N,1
988
EPR:
21°N
-9°N
87
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeG
orgo
lept
ispa
tulu
sM
CLEA
N,1
988
EPR:
13°N
;Gal
apag
os87
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeG
orgo
lept
issp
iral
isM
CLEA
N,1
988
EPR:
13°N
87
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
sat
lant
icus
WA
RÉN
&BO
UCH
ET,2
001
MA
R:38
°N-2
3°N
88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
sco
rrug
atus
MCL
EAN,1
993
JFR
88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
scr
ista
tus
MCL
EAN,1
988
EPR:
21°N
,13°
N;G
alap
agos
88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
sel
evat
usM
CLEA
N,1
988
EPR:
21°N
-17°
S;G
alap
agos
;Lau
;Mar
iana
;NFi
ji88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
sfu
cens
isM
CLEA
N,1
988
JFR
88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
sga
lrif
tens
isM
CLEA
N,1
988
EPR:
9°N
;Gal
apag
os88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
sgu
aym
asen
sis
MCL
EAN,1
988
Gua
ymas
88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
sja
poni
cus
OK
UTA
NIFU
JIK
URA
&SA
SAK
I,19
93O
kina
wa
T88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
snu
xO
KU
TAN
IFU
JIK
URA
&SA
SAK
I,19
93O
kina
wa
T88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
sov
alis
MCL
EAN,1
988
EPR:
21°N
-17°
S;G
alap
agos
88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
spu
stul
osus
MCL
EAN,1
988
EPR:
21°N
-17°
S;G
alap
agos
88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
ssc
hrol
liBE
CK,1
993
Man
us88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aeLe
peto
drilu
ste
vnia
nus
MCL
EAN,1
991
EPR:
11°N
88
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aePs
eudo
rim
ula
mar
iana
eM
CLEA
N,1
989
Mar
iana
91
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Lep
etod
rilid
aePs
eudo
rim
ula
mid
atla
ntic
aM
CLEA
N,1
992
MA
R:37
°N-1
4°N
91
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Pyr
opel
tida
ePy
rope
lta
cory
mba
MCL
EAN
&H
ASZ
PRU
NA
R,19
87G
uaym
as93
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Pyr
opel
tida
ePy
rope
lta
mus
aica
MCL
EAN
&H
ASZ
PRU
NA
R,19
87JF
R93
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Pyr
opel
tida
ePy
rope
lta
yam
ato
SASA
KIO
KU
TAN
I&
FUJI
KU
RA,2
003
Izu
93
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Pyr
opel
tida
ePy
rope
lta
bohl
eiBE
CK,1
996
Taba
r-Fe
ni:E
S93
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Ske
neid
aeBr
ucei
ella
glob
ulus
WA
RÉN
&BO
UCH
ET,1
993
Lau,
NFi
ji94
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Ske
neid
aeFu
cari
am
ysta
xW
ARÉ
N&
BOU
CHET
,200
1Ta
bar-
Feni
:ES
95
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Ske
neid
aeFu
cari
ast
riat
aW
ARÉ
N&
BOU
CHET
,199
3JF
R95
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Ske
neid
aeLe
ptog
yra
infl
ata
WA
RÉN
&BO
UCH
ET,1
993
Lau
96
520
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Ske
neid
aePr
otol
ira
thor
vald
sson
iWA
RÉN,1
996
MA
R:SP
;non
-ven
tlo
cati
ons
97
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Ske
neid
aePr
otol
ira
valv
atoi
des
WA
RÉN
&BO
UCH
ET,1
993
MA
R:LS
,MG
97
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Sut
ilizo
nida
eSu
tiliz
ona
pter
odon
WA
RÉN
&BO
UCH
ET,2
001
MA
R:SP
98
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Sut
ilizo
nida
eSu
tiliz
ona
thec
aM
CLEA
N,1
989
EPR:
13°N
98
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Sut
ilizo
nida
eSu
tiliz
ona
tunn
iclif
fae
WA
RÉN
&BO
UCH
ET,2
001
JFR:
ES98
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Sut
ilizo
nida
eTe
mno
cinc
liseu
ripe
sM
CLEA
N,1
989
JFR
99
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Sut
ilizo
nida
eTe
mno
zaga
pari
lisM
CLEA
N,1
989
EPR:
21°N
100
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Tro
chid
aeBa
thym
arga
rite
ssy
mpl
ecto
rW
ARÉ
N&
BOU
CHET
,198
9EP
R:13
°N,2
1°S
101
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Tro
chid
aeH
elic
reni
onre
ticu
latu
mW
ARÉ
N&
BOU
CHET
,199
3La
u:H
H10
2
Mol
lusc
a,G
astr
opod
a,Ve
tiga
stro
poda
,Tro
chid
aeVe
tulo
nia
phal
cata
WA
RÉN
&BO
UCH
ET,1
993
NFi
ji10
3
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeCy
athe
rmia
nati
coid
esW
ARÉ
N&
BOU
CHET
,198
9EP
R:21
°N-9
°N10
4
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeLa
cuno
ides
exqu
isit
usW
ARÉ
N&
BOU
CHET
,198
9G
alap
agos
105
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeLa
cuno
ides
vitr
eus
WA
RÉN
&BO
UCH
ET,2
001
JFR:
AV-
AV
F10
5
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeM
elan
odry
mia
aura
ntia
caH
ICK
MA
N,1
984
EPR:
13°N
,7°S
-23°
S;PA
R:32
°S10
6
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeM
elan
odry
mia
brig
htae
WA
RÉN
&BO
UCH
ET,1
993
JFR:
ES10
6
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeM
elan
odry
mia
gale
rona
eW
ARÉ
N&
BOU
CHET
,199
3EP
R:13
°N10
6
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeN
eom
phal
usfr
ette
rae
MCL
EAN,1
981
EPR:
21°N
-9°N
;Gal
apag
os10
7
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aePa
chyd
erm
iala
evis
WA
RÉN
&BO
UCH
ET,1
989
EPR:
21°N
-17°
S10
8
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aePa
chyd
erm
iasc
ulpt
aW
ARÉ
N&
BOU
CHET
,199
3La
u,N
Fiji
108
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aePl
anor
bide
llade
pres
saW
ARÉ
N&
BOU
CHET
,199
3La
u:H
H10
9
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aePl
anor
bide
llapl
anis
pira
WA
RÉN
&BO
UCH
ET,1
989
EPR:
21°N
-17°
S10
9
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeSy
mm
etro
mph
alus
hage
niBE
CK,1
992
Man
us11
0
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Neo
mph
alid
aeSy
mm
etro
mph
alus
regu
lari
sM
CLEA
N,1
990
Mar
iana
110
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eCt
enop
elta
proi
fera
WA
RÉN
&BO
UCH
ET,1
993
EPR:
13°N
111
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eD
epre
ssig
yra
glob
ulus
WA
RÉN
&BO
UCH
ET,1
989
JFR
112
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eEc
hino
pelt
afi
stul
osa
MCL
EAN,1
989
EPR:
13°N
,21°
S11
3
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eH
irto
pelt
ahi
rta
MCL
EAN,1
989
EPR:
13°N
-21°
S11
4
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eH
irto
pelt
atu
fari
BECK
,200
2EP
R:21
°S11
4
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eLi
rape
xco
stel
lata
WA
RÉN
&BO
UCH
ET,2
001
MA
R:LS
115
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eLi
rape
xgr
anul
aris
WA
RÉN
&BO
UCH
ET,1
989
EPR:
21°N
-9°N
115
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eLi
rape
xhu
mat
aW
ARÉ
N&
BOU
CHET
,200
1EP
R:21
°N11
5
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eN
odop
elta
hem
inod
aM
CLEA
N,1
989
EPR:
21°N
,13°
N11
6
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eN
odop
elta
rign
eae
WA
RÉN
&BO
UCH
ET,2
001
EPR:
13°N
116
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eN
odop
elta
subn
oda
MCL
EAN,1
989
EPR:
13°N
116
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
ePe
ltos
pira
delic
ata
MCL
EAN,1
989
EPR:
13°N
-9°N
118
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
ePe
ltos
pira
lam
ellif
era
WA
RÉN
&BO
UCH
ET,1
989
EPR:
13°N
118
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
ePe
ltos
pira
oper
cula
taM
CLEA
N,1
989
EPR:
21°N
-9°N
,17°
S11
8
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
ePe
ltos
pira
smar
agdi
naW
ARÉ
N&
BOU
CHET
,200
1M
AR:
38°N
-15°
N11
8
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
eRh
ynch
opel
taco
ncen
tric
aM
CLEA
N,1
989
EPR:
21°N
-17°
S11
9
Mol
lusc
a,G
astr
opod
a,N
eom
phal
ina,
Pelt
ospi
rida
e„s
caly
foot
gast
ropo
d“CI
R12
0
521
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Phyl
um
,C
lass
,O
rder
,Fa
mil
ySp
ecie
sD
istr
ibu
tio
nPa
ge
Mol
lusc
a,G
astr
opod
a,N
erit
imor
pha,
Phen
acol
epad
idae
Olg
asol
aris
tollm
anni
BECK
,199
2M
anus
121
Mol
lusc
a,G
astr
opod
a,N
erit
imor
pha,
Phen
acol
epad
idae
Shin
kaile
pas
bria
ndiW
ARÉ
N&
BOU
CHET
,200
1M
AR:
LO,L
S,M
G12
2
Mol
lusc
a,G
astr
opod
a,N
erit
imor
pha,
Phen
acol
epad
idae
Shin
kaile
pas
kaik
aten
sis
OK
UTA
NISA
ITO
&H
ASH
IMO
TO,1
989
Izu:
KS
122
Mol
lusc
a,G
astr
opod
a,N
erit
imor
pha,
Phen
acol
epad
idae
Shin
kaile
pas
moj
inen
sis
SASA
KIO
KU
TAN
I&
FUJI
KU
RA,2
003
Izu
122
Mol
lusc
a,G
astr
opod
a,N
erit
imor
pha,
Phen
acol
epad
idae
Shin
kaile
pas
tufa
riBE
CK,1
992
NFi
ji12
2
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Bu
ccin
idae
Eosi
pho
auze
ndei
WA
RÉN
&BO
UCH
ET,2
001
EPR:
17°S
124
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Bu
ccin
idae
Eosi
pho
desb
ruye
resi
nipp
onen
sis
OK
UTA
NI&
OH
TA,1
993
Oki
naw
aT
124
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Bu
ccin
idae
Eosi
pho
desb
ruye
resi
NIP
PON
ENSI
SO
KU
TAN
I&
FUJI
WA
RA,2
000
Lau,
Mar
iana
,NFi
ji12
4
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Ce
rith
iops
idae
Spec
ulat
orca
rios
usW
ARÉ
N&
BOU
CHET
,200
1Ex
pore
r:M
M12
6
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,El
achi
sini
dae
Laev
iphi
tus
desb
ruye
resi
WA
RÉN
&BO
UCH
ET,2
001
MA
R:M
G-R
B12
7
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
eA
lvin
ocon
cha
hess
leri
OK
UTA
NI&
OH
TA,1
988
CIR:
KV
F;La
u;M
aria
na;N
Fiji
128
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
eD
esbr
uyer
esia
canc
ella
taW
ARÉ
N&
BOU
CHET
,199
3La
u,N
Fiji
129
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
eD
esbr
uyer
esia
mar
iana
ensi
sO
KU
TAN
I,19
90M
anus
129
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
eD
esbr
uyer
esia
mar
isin
dica
OK
UTA
NIH
ASH
IMO
TO&
SASA
KI,
2004
CIR
129
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
eD
esbr
uyer
esia
mel
anio
ides
WA
RÉN
&BO
UCH
ET,1
993
NFi
ji12
9
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
eD
esbr
uyer
esia
spin
osa
WA
RÉN
&BO
UCH
ET,1
993
Lau,
Man
us12
9
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
eIf
rem
eria
naut
ileiB
OU
CHET
&W
ARÉ
N,1
991
Lau,
Man
us,N
Fiji
130
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
ePr
ovan
nabu
ccin
oide
sW
ARÉ
N&
BOU
CHET
,199
3La
u,N
Fiji
132
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
ePr
ovan
nagl
abra
OK
UTA
NITS
UCH
IDA
&FU
JIK
URA
,199
2O
kina
wa
T;no
n-ve
ntlo
cati
ons
132
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
ePr
ovan
naio
sW
ARÉ
N&
BOU
CHET
,198
6EP
R:21
°N-1
7°N
,13°
N;G
alap
agos
132
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
ePr
ovan
nala
evis
WA
RÉN
&PO
ND
E,19
91G
uaym
as;J
FR;n
on-v
ent
loca
tion
s13
2
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
ePr
ovan
nam
uric
ata
WA
RÉN
&BO
UCH
ET,1
986
EPR:
21°N
;Gal
apag
os;L
au;N
Fiji
132
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
ePr
ovan
nana
ssar
iaef
orm
isO
KU
TAN
I,19
90M
aria
na,M
anus
132
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
ePr
ovan
nase
gonz
aciW
ARÉ
N&
PON
DER
,199
1CI
R13
2
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Pr
ovan
nida
ePr
ovan
nava
riab
ilis
WA
RÉN
&BO
UCH
ET,1
986
JFR;
non-
vent
loca
tion
s13
2
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,Ri
ssoi
dae
Alv
ania
sten
olop
haBO
UCH
ET&
WA
RN,1
993
MA
R:LS
,MG
133
Mol
lusc
a,G
astr
opod
a,Ca
enog
astr
opod
a,V
itri
nelli
dae
Neu
sas
mar
shal
liSY
KES
,192
5M
AR:
MG
134
Mol
lusc
a,G
astr
opod
a,H
eter
obra
nchi
a,H
yalo
gyri
nida
eH
yalo
gyri
nagl
obul
aris
WA
RÉN
&BO
UCH
ET,2
001
JFR:
ES13
5
Mol
lusc
a,G
astr
opod
a,H
eter
obra
nchi
a,H
yalo
gyri
nida
eH
yalo
gyri
nagr
assl
eiW
ARÉ
N&
BOU
CHET
,199
3G
uaym
as13
5
Mol
lusc
a,G
astr
opod
a,H
eter
obra
nchi
a,O
rbit
este
llida
eLu
rifa
xvi
treu
sW
ARÉ
N&
BOU
CHET
,200
1M
AR:
MG
,LS
136
Mol
lusc
a,G
astr
opod
a,H
eter
obra
nchi
a,X
ylod
iscu
lidae
Xyl
odis
cula
anal
oga
WA
RÉN
&BO
UCH
ET,2
001
MA
R:LS
,MG
137
Mol
lusc
a,G
astr
opod
a,H
eter
obra
nchi
a,X
ylod
iscu
lidae
Xyl
odis
cula
maj
orW
ARÉ
N&
BOU
CHET
,200
1N
Fiji
137
Mol
lusc
a,G
astr
opod
a,Pr
osob
ranc
hia,
Turr
idae
Phym
orhy
nchu
sca
rina
tus
WA
RÉN
&BO
UCH
ET,2
001
MA
R:23
°N-1
5°N
138
Mol
lusc
a,G
astr
opod
a,Pr
osob
ranc
hia,
Turr
idae
Phym
orhy
nchu
shy
fifl
uxiB
ECK,1
996
NFi
ji,no
n-ve
ntlo
cati
ons
138
Mol
lusc
a,G
astr
opod
a,Pr
osob
ranc
hia,
Turr
idae
Phym
orhy
nchu
sm
ajor
WA
RÉN
&BO
UCH
ET,2
001
EPR:
13°N
-9°N
138
Mol
lusc
a,G
astr
opod
a,Pr
osob
ranc
hia,
Turr
idae
Phym
orhy
nchu
sm
oska
levi
SYSO
EV&
KA
NTO
R,19
95M
AR:
26°N
-23°
N13
8
Mol
lusc
a,G
astr
opod
a,Pr
osob
ranc
hia,
Turr
idae
Phym
orhy
nchu
sov
atus
WA
RÉN
&BO
UCH
ET,2
001
MA
R:37
°N-1
5°N
138
Mol
lusc
a,G
astr
opod
a,Pr
osob
ranc
hia,
Turr
idae
Phym
orhy
nchu
sst
arm
eriO
KU
TAN
I&
OH
TA,1
993
Man
us,N
Fiji
138
Mol
lusc
a,G
astr
opod
a,Pr
osob
ranc
hia,
Turr
idae
Phym
orhy
nchu
sw
aren
iSY
SOEV
&K
AN
TOR,
1995
Taba
r-Fe
ni:E
S13
8
522
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Mol
lusc
a,G
astr
opod
a,N
udib
ranc
hia,
Den
dron
otid
aeD
endr
onot
usco
mte
tiV
ALD
ES&
BOU
CHET
,199
8M
AR:
LS14
0
Mol
lusc
a,Bi
valv
ia,H
eter
odon
ta,V
esic
omyi
dae
Caly
ptog
ena
edis
onen
sis
OK
UTA
NI,
KO
JIM
A&
KIM
,200
3Ta
bar-
Feni
:ES
142
Mol
lusc
a,Bi
valv
ia,H
eter
odon
ta,V
esic
omyi
dae
Caly
ptog
ena
exte
nta
KRY
LOVA
&M
OSK
ALE
V,1
996
Gor
da;n
on-v
ent
loca
tion
s14
3
Mol
lusc
a,Bi
valv
ia,H
eter
odon
ta,V
esic
omyi
dae
Caly
ptog
ena
giga
sD
ALL
,189
6G
uaym
as,J
FR;n
on-v
ent
loca
tion
s14
4
Mol
lusc
a,Bi
valv
ia,H
eter
odon
ta,V
esic
omyi
dae
Caly
ptog
ena
mag
nifi
caBO
SS&
TURN
ER,1
980
EPR:
21°N
-23°
S;G
alap
agos
145
Mol
lusc
a,Bi
valv
ia,H
eter
odon
ta,V
esic
omyi
dae
Caly
ptog
ena
nank
aien
sis
OK
UTA
NI,
KO
JIM
A&
ASH
I,19
96O
kina
wa
T:IR
;non
-ven
tlo
cati
ons
146
Mol
lusc
a,Bi
valv
ia,H
eter
odon
ta,V
esic
omyi
dae
Caly
ptog
ena
okut
anii
KO
JIM
A&
OH
TA,1
997
Oki
naw
aT:
IR;n
on-v
ent
loca
tion
s14
7
Mol
lusc
a,Bi
valv
ia,H
eter
odon
ta,V
esic
omyi
dae
Caly
ptog
ena
solid
issi
ma
OK
UTA
NI,
HA
SHIM
OTO
&FU
JIK
URA
,199
2O
kina
wa
T:M
EK14
8
Mol
lusc
a,Bi
valv
ia,P
roto
bran
chia
,Sol
emyi
dae
Ach
arx
alin
aeM
ÉTIV
IER
&CO
SEL,
1993
Lau:
HH
149
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
adul
oide
sH
ASH
IMO
TO&
OK
TUA
NI,
1994
Oki
naw
aT:
MEK
,IR;
non-
vent
loca
tion
s15
0
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
azor
icus
COSE
L&
COM
TET,
1998
MA
R:LS
,MG
,RB
151
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
brev
ior
COSE
L,M
ÉTIV
IER
&H
ASH
IMO
TO,1
994
Lau,
NFi
ji15
3
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
elon
gatu
sCO
SEL,
MÉT
IVIE
R&
HA
SHIM
OTO
,199
4N
Fiji
154
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
japo
nicu
sH
ASH
IMO
TO&
OK
TUA
NI,
1994
Oki
naw
aT:
MEK
,IR;
non-
vent
loca
tion
s15
5
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
mar
isin
dicu
sH
ASH
IMO
TO,2
001
CIR:
KV
F15
6
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
plat
ifro
nsH
ASH
IMO
TO&
OK
TUA
NI,
1994
Oki
naw
aT:
IR,I
zena
Cald
ron;
157
non-
vent
loca
tion
s
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
pute
oser
pent
isCO
SEL,
MÉT
IVIE
R&
HA
SHIM
OTO
,199
4M
AR:
LO,S
P15
8
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
sept
emdi
erum
HA
SHIM
OTO
&O
KTU
AN
I,19
94Iz
u:M
K,S
S15
9
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eBa
thym
odio
lus
ther
mop
hilu
sK
ENK
&W
ILSO
N,1
985
EPR:
13°N
-21°
S16
0
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eG
igan
tida
sgl
adiu
sCO
SEL
&M
ARS
HA
LL,2
003
Ker
mad
ec:R
IIIS,
RVS
162
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Myt
ilida
eG
igan
tida
sho
riko
shii
HA
SHIM
OTO
&YA
MA
NE,
2005
Izu:
KS
163
Mol
lusc
a,Bi
valv
ia,P
teri
omor
phia
,Pec
tini
dae
Bath
ypec
ten
vulc
aniS
CHEI
N-F
ATT
ON,1
985
EPR:
13°N
-9°N
;PA
R:32
°S,3
8°S;
Gal
apag
os16
4
Mol
lusc
a,Bi
valv
ia,P
tero
mor
phia
,Pec
tini
dae
Sine
pect
ense
gonz
aciS
CHEI
N,i
npr
ess
Man
us16
5
Mol
lusc
a,Ce
phal
opod
a,O
ctop
oda,
Cirr
oteu
thid
aeCi
rrot
haum
am
urra
yiCH
UN,1
911
EPR,
non-
vent
loca
tion
s16
7
Mol
lusc
a,Ce
phal
opod
a,O
ctop
oda,
Cirr
oteu
thid
aeCi
rrot
euth
ism
agna
HO
YLE
,188
5M
AR:
LO;E
PR:1
7°S;
non-
vent
loca
tion
s16
8
Mol
lusc
a,Ce
phal
opod
a,O
ctop
oda,
Gri
mpo
teut
hida
eG
rim
pote
uthi
sRO
BSO
N,1
932
EPR,
MA
R,no
n-ve
ntlo
cati
ons
169
Mol
lusc
a,Ce
phal
opod
a,O
ctop
oda,
Oct
opod
idae
Vul
cano
ctop
ushy
drot
herm
alis
GO
NZÁ
LES,
GU
ERRA
,RO
CHA
&BR
IAN
D,2
002
EPR:
13°N
,23°
S17
0
Mol
lusc
a,Ce
phal
opod
a,O
ctop
oda,
Oct
opod
idae
Bent
hoct
opus
GRI
MPE
,192
1G
orda
,JFR
,non
-ven
tlo
cati
ons
171
Mol
lusc
a,Ce
phal
opod
a,O
ctop
oda,
Oct
opod
idae
Gra
nele
done
JOU
BIN,1
918
Expl
orer
,Gal
apag
os,J
FR,K
erm
adec
,MA
R17
2
Nem
atod
a,A
deno
phor
ea,D
esm
odor
ida,
Des
mod
orid
aeD
esm
odor
aal
bert
iVER
SCH
ELD
E,G
OU
RBA
ULT
&V
INCX
,199
8G
uaym
as17
4
Nem
atod
a,A
deno
phor
ea,D
esm
odor
ida,
Des
mod
orid
aeD
esm
odor
am
arci
VER
SCH
ELD
E,G
OU
RBA
ULT
&V
INCX
,199
8La
u:H
H17
5
Nem
atod
a,A
deno
phor
ea,D
esm
odor
ida,
Des
mod
orid
aeD
esm
odor
ella
balt
eata
DEC
RAEM
ER&
GO
URB
AU
LT,1
998
Gua
ymas
176
Nem
atod
a,A
deno
phor
ea,D
esm
odor
ida,
Des
mod
orid
aeD
esm
odor
ella
spin
eaca
udat
aD
ECRA
EMER
&G
OU
RBA
ULT
,199
8G
uaym
as17
7
Nem
atod
a,A
deno
phor
ea,C
hrom
ador
ida,
Dra
cone
mat
idae
Ceph
aloc
haet
osom
apa
cifi
cum
noti
umD
ECRA
EMER
&G
OU
RBA
ULT
,199
7La
u17
8
Nem
atod
a,A
deno
phor
ea,C
hrom
ador
ida,
Dra
cone
mat
idae
Din
etia
nyct
erob
iaD
ECRA
EMER
&G
OU
RBA
ULT
,199
7EP
R:21
°N17
9
Nem
atod
a,Se
cern
ente
a,Sp
irur
ida,
Cyst
idic
olid
aeM
orav
ecne
ma
sego
nzac
iJU
STIN
E,CA
SSO
NE
&PE
TTER
,200
2M
AR:
LO,S
P18
0
Aca
ntho
ceph
ala,
Pala
eaca
ntho
ceph
ala,
Echi
norh
ynch
ida,
Hyp
oech
inor
hync
hus
ther
mac
eriB
URO
N,1
988
EPR:
13°N
181
Hyp
oech
inor
hync
hida
e
Nem
erti
ni,H
oplo
nem
erti
ni,M
onos
tilif
era,
fam
ilyun
know
nTh
erm
onem
erte
sva
lens
ROG
ERS,
GIB
SON
&TU
NN
ICLI
FFE,
1996
JFR
183
523
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Phyl
um
,C
lass
,O
rder
,Fa
mil
ySp
ecie
sD
istr
ibu
tio
nPa
ge
Ann
elid
a,Po
lych
aeta
,Eun
icid
a,A
mph
inom
idae
Arc
hino
me
rosa
cea
BLA
KE,
1985
Gal
apag
os18
6A
nnel
ida,
Poly
chae
ta,E
unic
ida,
Dor
ville
idae
Exal
lopu
sju
mar
siBL
AK
E,19
85G
uaym
as18
7A
nnel
ida,
Poly
chae
ta,E
unic
ida,
Dor
ville
idae
Oph
ryot
roch
aak
esso
niBL
AK
E,19
85EP
R,G
alap
agos
,Gua
ymas
188
Ann
elid
a,Po
lych
aeta
,Eun
icid
a,D
orvi
lleid
aeO
phry
otro
cha
glob
opal
pata
BLA
KE
&H
ILBI
G,1
990
JFR
189
Ann
elid
a,Po
lych
aeta
,Eun
icid
a,D
orvi
lleid
aeO
phry
otro
cha
plat
ykep
hale
BLA
KE,
1985
Gua
ymas
190
Ann
elid
a,Po
lych
aeta
,Eun
icid
a,D
orvi
lleid
aePa
roug
iaw
olfi
BLA
KE
&H
ILBI
G,1
990
JFR
192
Ann
elid
a,Po
lych
aeta
,Eun
icid
a,Eu
nici
dae
Euni
cepu
lvin
opal
pata
FAU
CHA
LD,1
982
EPR:
21°N
,13°
N19
3A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,C
ryso
peta
lidae
Thra
umas
tos
diet
eriW
ATS
ON,2
001
Lau,
Nfi
ji;no
n-ve
ntlo
cati
ons
194
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Gly
ceri
dae
Gly
cera
bran
chio
poda
MO
ORE
,191
1G
uaym
as;n
on-v
ent
loca
tion
s19
5A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,G
lyce
rida
eG
lyce
rate
ssel
ata
GRU
BE,1
863
MA
R:LS
;non
-ven
tlo
cati
ons
196
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Hes
ioni
dae
Am
phid
urop
sis
axia
lens
is(B
LAK
E&
HIL
BIG,1
990)
EPR:
9°N
;JFR
:AV-
AV
F,M
V19
8A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,H
esio
nida
eH
esio
deir
iagl
abra
BLA
KE
&H
ILBI
G,1
990
JFR:
AV-
AV
F,A
V-CA
SM19
9A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,H
esio
nida
eH
esio
lyra
berg
iBLA
KE,
1985
EPR:
21°N
,9°N
200
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Hes
ioni
dae
Hes
iosp
ina
vest
imen
tife
raBL
AK
E,19
85EP
R;Ex
plor
er;J
FR;G
alap
agos
202
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Hes
ioni
dae
Ner
eim
yra
alvi
nae
BLA
KE,
1985
Gal
apag
os;G
uaym
as20
4A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,H
esio
nida
eSi
rsoe
gras
slei
(BLA
KE,
1985
)G
uaym
as20
5A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,N
auti
linie
llida
eIh
eyom
ytili
dico
latr
iden
tatu
sM
IURA
&H
ASH
IMO
TO,1
996
Oki
naw
aT
206
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Nau
tilin
ielli
dae
Myt
ilidi
phila
ense
iens
isM
IURA
&H
ASH
IMO
TO,1
993
Oki
naw
aT
207
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Nau
tilin
ielli
dae
Myt
ilidi
phila
okin
awae
nsis
MIU
RA&
HA
SHIM
OTO
,199
3O
kina
wa
T20
8A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,N
auti
linie
llida
eSh
inka
ilon
gipe
data
MIU
RA&
OH
TA,1
991
Oki
naw
aT
209
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Nau
tilin
ielli
dae
Shin
kais
emilo
nga
MIU
RA&
HA
SHIM
OTO
,199
6O
kina
wa
T:M
EK21
0A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,N
erei
dae
Ner
eis
pisc
esae
BLA
KE
&H
ILBI
G,1
990
JFR
212
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Ner
eida
eN
erei
ssa
nder
siBL
AK
E,19
85EP
R;G
alap
agos
;Gua
ymas
214
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Phy
llodo
cida
eEu
lalia
papi
llosa
(BLA
KE,
1985
)EP
R:21
°N21
5A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
hyllo
doci
dae
Gal
apag
omys
tide
sar
ista
taBL
AK
E,19
85EP
R:13
°N,9
°N;G
alap
agos
216
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Phy
llodo
cida
ePr
otom
ysti
des
vere
nae
BLA
KE
&H
ILBI
G,1
990
Expl
orer
,JFR
217
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Bath
ykur
ilagu
aym
asen
sis
PETT
IBO
NE,
1898
Gua
ymas
218
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Bran
chin
otog
lum
ahe
ssle
riPE
TTIB
ON
E,19
85EP
R:9°
N;G
alap
agos
;Gua
ymas
219
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Bran
chin
otog
lum
am
aria
nus
(PET
TIBO
NE,
1989
)M
anus
221
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Bran
chin
otog
lum
asa
nder
siPE
TTIB
ON
E,19
85EP
R:21
°N-1
7°S;
Gal
apag
os;G
uaym
as22
3A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eBr
anch
inot
oglu
ma
sego
nzac
i(M
IURA
&D
ESBR
UY
ÈRES
,199
5)La
u,M
anus
,NFi
ji22
5A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eBr
anch
inot
oglu
ma
trif
urcu
s(M
IURA
&D
ESBR
UY
ÈRES
,199
5)La
u,N
Fiji
227
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Bran
chin
otog
lum
atu
nnic
liffa
ePE
TTIB
ON
E,19
88Ex
plor
er,J
FR:A
V,ES
,S-J
FR22
8A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eBr
anch
iplic
atus
cupr
eus
PETT
IBO
NE,
1985
EPR:
21°N
-9°N
;Gua
ymas
230
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Bran
chip
olyn
oepe
ttib
onae
MIU
RA&
HA
SHIM
OTO
,199
1Iz
u,O
kina
wa
T;N
Fiji,
Lau
231
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Bran
chip
olyn
oese
epen
sis
PETT
IBO
NE,
1986
MA
R:LS
;non
-ven
tlo
cati
ons
232
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Bran
chip
olyn
oesy
mm
ytili
daPE
TTIB
ON
E,19
84EP
R;G
alap
agos
234
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Iphi
onel
lari
sens
isPE
TTIB
ON
E,19
86EP
R:21
°N,9
°N23
5A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eLe
pido
noto
podi
umat
alan
tae
DES
BRU
YÈR
ES&
HO
URD
EZ,2
000
EPR:
13°N
-17°
S23
6
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Lepi
dono
topo
dium
fim
bria
tum
PETT
IBO
NE,
1983
EPR:
21°N
-9°N
237
524
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Lepi
dono
topo
dium
joui
nae
DES
BRU
YÈR
ES&
HO
URD
EZ,2
000
MA
R:LS
,MG
238
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Lepi
dono
topo
dium
pisc
esae
PETT
IBO
NE,
1986
Expl
orer
,Gor
da,J
FR:A
V,ES
,S-J
FR23
9A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eLe
pido
noto
podi
umri
ften
sePE
TTIB
ON
E,19
84EP
R:21
°N-9
°N;G
alap
agos
241
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Lepi
dono
topo
dium
will
iam
sae
PETT
IBO
NE,
1984
EPR:
21°N
-9°N
;Gal
apag
os24
2A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eLe
vens
tein
iella
iris
HO
URD
EZ&
DES
BRU
YÈR
ES,2
003
MA
R:LS
,RB
243
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Pol
ynoi
dae
Leve
nste
inie
llapl
icat
aH
OU
RDEZ
&D
ESBR
UY
ÈRES
,200
0EP
R:9°
N24
4A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eTh
erm
iphi
one
fijie
nsis
MIU
RA,1
994
Lau,
NFi
ji24
6A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eTh
erm
iphi
one
tufa
riH
ART
MA
NN-S
CHRÖ
DER
,199
2EP
R:7°
S-21
°S24
7A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eTh
erm
opol
ynoe
bran
chia
taM
IURA
,199
4La
u,M
anus
,NFi
ji24
9A
nnel
ida,
Poly
chae
ta,P
hyllo
doci
da,P
olyn
oida
eVa
mpi
ropo
lyno
eem
bley
iMA
RCU
S&
HO
URD
EZ,2
002
JFR:
AV
251
Ann
elid
a,Po
lych
aeta
,Phy
llodo
cida
,Syl
lidae
Spha
eros
yllis
ridg
iens
isBL
AK
E&
HIL
BIG,1
990
Expl
orer
,JFR
252
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Se
rpul
idae
Lam
inat
ubus
alvi
niTE
NH
OV
E&
ZIBR
OW
IUS,
1986
EPR:
21°N
-23°
S;G
alap
agos
253
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Se
rpul
idae
Prot
ishy
drot
herm
ica
TEN
HO
VE
&ZI
BRO
WIU
S,19
86EP
R:21
°N,1
3°N
,17°
S;G
alap
ago;
PAR:
38°S
254
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Si
bogl
inid
aeA
lays
iasp
iral
isSO
UTH
WA
RD,1
991
Lau:
VF
255
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Si
bogl
inid
aeA
rcov
esti
aiv
anov
iSO
UTH
WA
RD&
GA
LGIN
,199
7La
u,M
anus
256
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Si
bogl
inid
aeLa
mel
libra
chia
barh
amiW
EBB,
1969
JFR:
MV;
non-
vent
loca
tion
s25
7A
nnel
ida,
Poly
chae
ta,S
abel
lida,
Sibo
glin
idae
Lam
ellib
rach
iaco
lum
naSO
UTH
WA
RD,1
991
Lau:
VF
258
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Si
bogl
inid
aeLa
mel
libra
chia
sats
uma
MIU
RA,T
SUK
AH
ARA
&H
ASH
IMO
TO,1
997
Japa
n:K
agos
him
a;N
anka
iTro
ugh
259
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Si
bogl
inid
aeO
asis
iaal
vina
eJO
NES
,198
5EP
R:21
°N-9
°N26
0A
nnel
ida,
Poly
chae
ta,S
abel
lida,
Sibo
glin
idae
Ridg
eia
pisc
esae
JON
ES,1
985
Expl
orer
;JFR
;Gua
ymas
261
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Si
bogl
inid
aeRi
ftia
pach
ypti
laJO
NES
,198
1EP
R,G
alap
agos
,Gua
ymas
262
Ann
elid
a,Po
lych
aeta
,Sab
ellid
a,Si
bogl
inid
aeSi
phon
obra
chia
laue
nsis
SOU
THW
ARD
,199
1La
u:V
F26
4A
nnel
ida,
Poly
chae
ta,S
abel
lida,
Sibo
glin
idae
Tevn
iaje
rich
onan
aJO
NES
,198
5EP
R:13
°N-2
1°S
265
Ann
elid
a,Po
lych
aeta
,Sco
leci
da,M
alda
nida
eN
icom
ache
arw
idss
oniB
LAK
E,19
85EP
R:21
°N,1
3°N
;Gal
apag
os;M
aria
na?
266
Ann
elid
a,Po
lych
aeta
,Sco
leci
da,M
alda
nida
eN
icom
ache
vent
icol
aBL
AK
E&
HIL
BIG,1
990
Gua
ymas
;Exp
lore
r;JF
R:A
V-A
VF,
S-JF
R26
7A
nnel
ida,
Poly
chae
ta,S
cole
cida
,Orb
iniid
aeLe
itos
colo
plos
pach
ybra
nchi
atus
BLA
KE
&H
ILBI
G,1
990
JFR
268
Ann
elid
a,Po
lych
aeta
,Sco
leci
da,O
rbin
iidae
Orb
inie
llaac
icul
ata
BLA
KE,
1985
Gal
apag
os26
9A
nnel
ida,
Poly
chae
ta,S
cole
cida
,Orb
iniid
aeO
rbin
iella
hobs
onae
BLA
KE
&H
ILBI
G,1
990
MA
R:RB
,TA
G27
0A
nnel
ida,
Poly
chae
ta,S
cole
cida
,Orb
iniid
aeSc
olop
los
ehle
rsiB
LAK
E,19
85G
alap
agos
271
Ann
elid
a,Po
lych
aeta
,Spi
onid
a,Ch
aeto
pter
idae
Spio
chae
topt
erus
SARS
,185
3M
AR:
RB,T
AG
272
Ann
elid
a,Po
lych
aeta
,Spi
onid
a,Sp
ioni
dae
Laon
ice
athe
cata
SIG
VALD
AD
ÓTT
IR&
DES
BRU
YÈR
ES,2
003
MA
R:LS
,LO
273
Ann
elid
a,Po
lych
aeta
,Spi
onid
a,Sp
ioni
dae
Laub
ieri
ellu
sgr
assl
eiM
ACI
OLE
K,1
981
Gal
apag
os27
4A
nnel
ida,
Poly
chae
ta,S
pion
ida,
Spio
nida
eLi
ndas
pio
dibr
anch
iata
BLA
KE
&M
ACI
OLE
K,1
992
Gua
ymas
275
Ann
elid
a,Po
lych
aeta
,Spi
onid
a,Sp
ioni
dae
Lind
aspi
oso
uthw
ardo
rum
BLA
KE
&M
ACI
OLE
K,1
992
JFR:
MV
276
Ann
elid
a,Po
lych
aeta
,Spi
onid
a,Sp
ioni
dae
Prio
nosp
iosa
nder
siM
ACI
OLE
K,1
981
Gal
apag
os27
8A
nnel
ida,
Poly
chae
ta,S
pion
ida,
Spio
nida
ePr
iono
spio
unila
mel
lata
SIG
VALD
AD
OTT
IR&
DES
BRU
YÈR
ES,2
003
MA
R:LS
,RB,
SP28
0A
nnel
ida,
Poly
chae
ta,S
pion
ida,
Spio
nida
eX
anda
ros
acan
thod
esM
ACI
OLE
K,1
981
Gal
apag
os28
1A
nnel
ida,
Poly
chae
ta,T
ereb
ellid
a,A
lvin
ellid
aeA
lvin
ella
caud
ata
DES
BRU
YÈR
ES&
LAU
BIER
,198
6EP
R:21
°N-1
7°S
282
Ann
elid
a,Po
lych
aeta
,Ter
ebel
lida,
Alv
inel
lidae
Alv
inel
lapo
mpe
jana
DES
BRU
YÈR
ES&
LAU
BIER
,198
0EP
R:21
°N-2
3°S
283
Ann
elid
a,Po
lych
aeta
,Ter
ebel
lida,
Alv
inel
lidae
Para
lvin
ella
(Mir
alvi
nella
)ba
cter
ioco
laD
ESBR
UY
ÈRES
&LA
UBI
ER,1
991
Gua
ymas
285
Ann
elid
a,Po
lych
aeta
,Ter
ebel
lida,
Alv
inel
lidae
Para
lvin
ella
(Mir
alvi
nella
)de
laD
ETIN
OVA
,198
8JF
R:A
V,ES
286
525
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Phyl
um
,C
lass
,O
rder
,Fa
mil
ySp
ecie
sD
istr
ibu
tio
nPa
ge
Ann
elid
a,Po
lych
aeta
,Ter
ebel
lida,
Alv
inel
lidae
Para
lvin
ella
(Mir
alvi
nella
)he
ssle
riD
ESBR
UY
ÈRES
&LA
UBI
ER,1
989
Mar
iana
,Man
us28
7A
nnel
ida,
Poly
chae
ta,T
ereb
ellid
a,A
lvin
ellid
aePa
ralv
inel
la(N
auta
lvin
ella
)pa
ndor
aeD
ESBR
UY
ÈRES
&LA
UBI
ER,1
986
JFR
288
Ann
elid
a,Po
lych
aeta
,Ter
ebel
lida,
Alv
inel
lidae
Para
lvin
ella
(Nau
talv
inel
la)
unid
enta
taD
ESBR
UY
ÈRES
&LA
UBI
ER,1
993
Lau,
NFi
ji28
9A
nnel
ida,
Poly
chae
ta,T
ereb
ellid
a,A
lvin
ellid
aePa
ralv
inel
la(P
aral
vine
lla)
fijie
nsis
DES
BRU
YÈR
ES&
LAU
BIER
,199
3La
u,N
Fiji
291
Ann
elid
a,Po
lych
aeta
,Ter
ebel
lida,
Alv
inel
lidae
Para
lvin
ella
(Par
alvi
nella
)gr
assl
eiD
ESBR
UY
ÈRES
&LA
UBI
ER,1
982
EPR;
Gal
apag
os;G
uaym
as29
2A
nnel
ida,
Poly
chae
ta,T
ereb
ellid
a,A
lvin
ellid
aePa
ralv
inel
la(P
aral
vine
lla)
palm
ifor
mis
DES
BRU
YÈR
ES&
LAU
BIER
,198
6Ex
pore
r,G
orda
,JFR
293
Ann
elid
a,Po
lych
aeta
,Ter
ebel
lida,
Alv
inel
lidae
Para
lvin
ella
(Par
alvi
nella
)su
lfin
cola
DES
BRU
YÈR
ES&
LAU
BIER
,199
3Ex
plor
er;J
FR;G
uaym
as29
4A
nnel
ida,
Poly
chae
ta,T
ereb
ellid
a,A
mph
aret
idae
Am
athy
slu
tziD
ESBR
UY
ÈRES
&LA
UBI
ER,1
996
MA
R:BS
,LS,
SP29
5A
nnel
ida,
Poly
chae
ta,T
ereb
ellid
a,A
mph
aret
idae
Am
phis
amyt
haga
lapa
gens
isZO
TTO
LI,1
983
Man
us,M
aria
na,N
Fiji,
Oki
naw
aT
296
Ann
elid
a,O
ligoc
haet
a,H
irud
inea
,Pis
cico
lidae
Bath
ybde
llasa
wye
riBU
RRES
ON,1
981
EPR:
14°S
,17°
S;G
alap
agos
297
Art
hrop
oda,
Ara
chni
da,A
cari
form
es,H
alac
arid
aeCo
pido
gnat
hus
papi
llatu
sK
RAN
TZ,1
982
Lau,
NFi
ji29
9A
rthr
opod
a,A
rach
nida
,Aca
rifo
rmes
,Hal
acar
idae
Hal
acar
ellu
sau
zend
ei(B
ART
SCH,1
990)
MA
R:BS
,LS
300
Art
hrop
oda,
Pycn
ogon
ida,
,Am
mot
heid
aeA
mm
othe
ave
rena
eCH
ILD,1
987
Expl
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;JFR
:AV,
ES30
1A
rthr
opod
a,Py
cnog
onid
a,,A
mm
othe
idae
Seri
cosu
raco
chle
ifov
eaCH
ILD,1
989
Mar
iana
302
Art
hrop
oda,
Pycn
ogon
ida,
,Am
mot
heid
aeSe
rico
sura
cyrt
oma
CHIL
D&
SEG
ON
ZAC,
1996
EPR:
13°N
303
Art
hrop
oda,
Pycn
ogon
ida,
,Am
mot
heid
aeSe
rico
sura
hete
rosc
ela
CHIL
D&
SEG
ON
ZAC,
1996
MA
R:LS
,LO
;MG
,RB,
SP30
4A
rthr
opod
a,Py
cnog
onid
a,,A
mm
othe
idae
Seri
cosu
ram
itra
taG
ORD
ON,1
944
MA
R:SP
;non
-ven
tlo
cati
ons
305
Art
hrop
oda,
Pycn
ogon
ida,
,Am
mot
heid
aeSe
rico
sura
vent
icol
aCH
ILD,1
987
JFR:
ES30
6A
rthr
opod
a,Cr
usta
cea,
Ost
raco
da,H
aloc
ypri
dae
Arc
hico
ncho
ecia
(Arc
hico
ncho
ecia
)ch
avtu
riK
ORN
ICK
ER&
HA
RRIS
ON-N
ELSO
N,2
005
EPR:
9°N
307
Art
hrop
oda,
Crus
tace
a,O
stra
coda
,Hal
ocyp
rida
eBa
thyc
onch
oeci
ade
evey
aeK
ORN
ICK
ER,1
969
Gua
ymas
,non
-ven
tlo
cati
ons
308
Art
hrop
oda,
Crus
tace
a,O
stra
coda
,Hal
ocyp
rida
eBa
thyc
onch
oeci
apa
ulul
aD
EEV
EY,1
968
Gua
ymas
,non
-ven
tlo
cati
ons
309
Art
hrop
oda,
Crus
tace
a,O
stra
coda
,Phi
lom
edid
aeEu
philo
med
escl
imax
KO
RNIC
KER
,199
1Ex
plor
er;J
FR31
0A
rthr
opod
a,Cr
usta
cea,
Ost
raco
da,P
olyc
opid
aePo
lyco
pett
apa
xK
ORN
ICK
ER&
HA
RRIS
ON-N
ELSO
N,2
005
EPR:
9°N
311
Art
hrop
oda,
Crus
tace
a,O
stra
coda
,Cyl
indr
oleb
erid
aePr
iono
tole
beri
sst
yxK
ORN
ICK
ER,1
991
EPR:
21°N
312
Art
hrop
oda,
Crus
tace
a,O
stra
coda
,Pon
tocy
prid
idae
Thom
onto
cypr
isbr
ight
aeM
AD
DO
CKS,
2006
EPR:
9°N
313
Art
hrop
oda,
Crus
tace
a,O
stra
coda
,Pon
tocy
prid
idae
Thom
onto
cypr
isgo
llner
aeM
AD
DO
CKS,
2006
EPR:
9°N
314
Art
hrop
oda,
Crus
tace
a,O
stra
coda
,Cyt
heru
rida
eX
yloc
ythe
reva
nhar
teni
MA
DD
OCK
S,20
06EP
R:13
°N,9
°N31
5A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Aeg
isti
dae
And
rom
asta
xm
uric
atus
CON
ROY-
DA
LTO
N&
HU
YS,
1999
Gal
apag
os31
8A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Aeg
isti
dae
And
rom
asta
xce
phal
ocer
atus
LEE
&H
UY
S,20
00O
kina
wa
T31
8A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Anc
orab
olid
aeU
ptio
nyx
vere
nae
CON
ROY-
DA
LTO
N&
HU
YS,
2000
JFR:
MV
319
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,A
rges
tida
eA
rges
toid
espr
ehen
silis
HU
YS
&CO
NRO
Y-D
ALT
ON,1
997
Gal
apag
os32
0A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Arg
esti
dae
Jam
stec
iate
raza
kiiL
EE&
HU
YS,
2000
Oki
naw
aT
321
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,A
rges
tida
eSc
abra
nten
nayo
oiLE
E&
HU
YS,
2000
Oki
naw
aT
322
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,A
ster
oche
rida
eCh
eram
omyz
onab
yssa
leH
UM
ES,1
989
EPR:
13°N
323
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,A
ster
oche
rida
eCo
lloch
erid
esbr
ychi
usH
UM
ES,1
999
JFR:
S-JF
R32
4A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Cla
usid
iidae
Hyp
halio
nca
ptan
sH
UM
ES,1
987
Gua
ymas
325
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,C
yclo
pini
dae
Bara
thri
cola
rim
ensi
sH
UM
ES,1
999
JFR:
CAS
326
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,C
yclo
pini
dae
Hep
tner
ina
conf
usa
IVA
NEN
KO
&D
EFA
Y,2
004
MA
R:LS
327
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Aph
otop
onti
usac
anth
inus
HU
MES
&LU
TZ,1
994
Gal
apag
os32
8A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eA
phot
opon
tius
arcu
atus
HU
MES
,198
7EP
R:9°
N;G
alap
agos
328
526
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Aph
otop
onti
usat
lant
eus
HU
MES
,199
6M
AR:
LS,M
G32
8A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eA
phot
opon
tius
bacu
liger
usH
UM
ES,1
987
EPR:
21°N
;Gal
apag
os32
8A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eA
phot
opon
tius
flex
ispi
naH
UM
ES,1
987
EPR:
13°N
328
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Aph
otop
onti
usfo
rcip
atus
HU
MES
,198
7G
orda
;JFR
;Gua
ymas
328
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Aph
otop
onti
ushy
dron
auti
cus
HU
MES
,198
9EP
R:13
°N32
8A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eA
phot
opon
tius
limat
ulus
HU
MES
,198
7EP
R:21
°N,9
°N;G
alap
agos
328
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Aph
otop
onti
usm
amm
illat
usH
UM
ES,1
987
EPR:
13°N
,21°
N,1
3°N
;Gal
apag
os;G
uaym
as32
8A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eA
phot
opon
tius
prob
olus
HU
MES
,199
0G
alap
agos
328
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Bent
hoxy
nus
spic
ulif
erH
UM
ES,1
984
Expo
rer,
Gor
da,J
FR:A
V33
0A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eBe
ntho
xynu
stu
mid
iset
aH
UM
ES,1
989
EPR:
13°N
330
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Ceut
hoec
etes
acan
thot
hrix
HU
MES
,198
7EP
R;G
alap
agos
331
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Ceut
hoec
etes
alig
erH
UM
ES&
DO
JIRI
,198
0EP
R;G
alap
agos
331
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Ceut
hoec
etes
cris
tatu
sH
UM
ES,1
987
EPR;
Gal
apag
os33
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eCe
utho
ecet
esin
trov
ersu
sH
UM
ES,1
987
EPR;
Gal
apag
os33
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eCh
asm
atop
onti
usth
esca
lus
HU
MES
,199
0La
u,M
aria
na33
2A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eD
iriv
ultu
ssp
inig
ulat
usH
UM
ES,1
999
Taba
r-Fe
ni:E
S33
3A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eEx
rim
ado
licho
pus
HU
MES
,198
7EP
R:13
°N33
4A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eEx
rim
asi
ngul
arH
UM
ES,1
987
EPR:
21°N
334
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Fiss
uric
ola
cari
tus
HU
MES
,198
7EP
R:21
°N33
5A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eH
umes
ipon
tius
arth
uriI
VAN
ENK
O&
FERR
ARI
,200
3JF
R33
6A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eN
ilva
tori
fera
HU
MES
,198
7EP
R:21
°N,1
3°N
;Gal
apag
os33
7A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eRh
ogob
ius
cont
ract
usH
UM
ES,1
987
EPR:
21°N
,13°
N;G
alap
agos
338
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Rhog
obiu
spr
essu
lus
HU
MES
,198
9G
alap
agos
338
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Rhog
obiu
sra
punc
ulus
(HU
MES
&SE
GO
NZA
C,19
98)
EPR:
9°N
338
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Rim
ipon
tius
med
iosp
inif
erH
UM
ES,1
996
MA
R:BS
,LO
,SP
339
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Scot
oece
tes
intr
orsu
sH
UM
ES,1
987
EPR:
13°N
,9°N
340
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sap
posi
tus
HU
MES
,198
9EP
R:21
°N,1
3°N
341
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sbr
evis
pina
HU
MES
,199
1La
u34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
cinc
tige
rH
UM
ES,1
987
EPR:
21°N
,13°
N,9
°N34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
clad
arus
HU
MES
,199
6M
AR:
BS,T
AG
341
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sfl
exus
HU
MES
,198
7EP
R:9°
N;G
uaym
as34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
hisp
idul
usH
UM
ES,1
987
EPR:
21°N
,13°
N,9
°N34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
latu
lus
HU
MES
,199
6M
AR:
SP34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
laue
nsis
HU
MES
,199
1La
u34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
mir
usH
UM
ES,1
996
EPR:
9°N
;MA
R:SP
341
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sm
ucro
nife
rus
HU
MES
,198
7EP
R:9°
N;G
uaym
as34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
paxi
llife
rH
UM
ES,1
989
EPR:
21°N
,9°N
341
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
spe
ctin
atus
HU
MES
,198
7M
aria
na,M
AR:
BS,S
P,TA
G34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
quad
risp
inos
usH
UM
ES,1
987
Gor
da,E
xplo
rer,
JFR
341
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sre
gius
HU
MES
,199
6M
AR:
SP34
1A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Dir
ivul
tida
eSt
ygio
pont
ius
rim
ivag
usH
UM
ES,1
997
MA
R:LS
341
527
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Phyl
um
,C
lass
,O
rder
,Fa
mil
ySp
ecie
sD
istr
ibu
tio
nPa
ge
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sse
ntif
erH
UM
ES,1
987
EPR:
21°N
,13°
N,9
°N34
1
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sse
rrat
usH
UM
ES,1
996
MA
R:SP
341
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sst
abili
tus
HU
MES
,199
0EP
R:13
°N,M
aria
na34
1
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
ste
res
HU
MES
,199
6M
AR:
SP34
1
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,D
iriv
ulti
dae
Styg
iopo
ntiu
sve
rruc
ulat
usH
UM
ES,1
987
EPR:
21°N
,9°N
341
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,E
cbat
hyri
onti
dae
Ecba
thyr
ion
prol
ixic
auda
HU
MES
,198
7EP
R:21
°N,1
3°N
,9°N
;Gal
apag
os34
3A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Ere
bona
ster
idae
Am
bilim
bus
alta
lis(H
UM
ES&
HU
YS,
1992
)JF
R34
4A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Ere
bona
ster
idae
Am
bilim
bus
arcu
scel
esti
sIV
AN
ENK
OD
EFA
YE
&H
UY
S,20
05M
AR:
RB34
4A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Ere
bona
ster
idae
Am
bilim
bus
tuer
kayi
(MA
RTIN
EZ-A
RBIZ
U,1
999)
NFi
ji34
4A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Lao
phon
tida
eBa
thyl
aoph
onte
azor
ica
LEE
&H
UY
S,19
99M
AR:
LS,M
G34
5A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Lao
phon
tida
eBa
thyl
aoph
onte
paci
fica
LEE
&H
UY
S,19
99EP
R:17
°S34
5A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Lub
bock
iidae
Lait
mat
obiu
scr
init
usH
UM
ES,1
987
Gua
ymas
346
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,M
egap
onti
idae
Hya
lopo
ntiu
sbo
xsha
lliH
UM
ES,1
988
Gal
apag
os34
7A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Mir
aciid
aeSt
enhe
liagu
ndul
aeW
ILLE
N,2
003
Taba
r-Fe
ni:E
S34
8A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Mis
ophr
iidae
Mis
ophr
iops
islo
ngic
auda
taBO
XSH
ALL
,198
3JF
R:CA
S34
9A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Onc
aeid
aeO
ncae
apr
aecl
ara
HU
MES
,198
8EP
R:21
°N,1
3°N
,9°N
;Gal
apag
os;G
uaym
as35
0A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Sco
leci
tric
hida
eG
riev
ella
shan
kiFE
RRA
RI&
MA
RKH
ASE
VA,2
000
EPR:
21°S
351
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,S
pino
cala
nida
eIs
aacs
ical
anus
pauc
iset
usFL
EMIN
GER
,198
3EP
R:21
°N35
2A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Teg
asti
dae
Smac
igas
tes
mic
heli
IVA
NEN
KO
&D
EFA
YE,
2004
MA
R:LS
353
Art
hrop
oda,
Crus
tace
a,Co
pepo
da,T
isbi
dae
Chol
idya
poly
piFA
RRA
N,1
914
JFR:
MV;
non-
vent
loca
tion
s35
4A
rthr
opod
a,Cr
usta
cea,
Cope
poda
,Tis
bida
eG
enes
isvu
lcan
octo
pusi
LÓPE
Z-G
ON
ZALE
Z,BR
ESCI
AN
I&
HU
YS,
2000
EPR:
13°N
355
Art
hrop
oda,
Crus
tace
a,Ci
rrip
edia
,Chi
onel
asm
atoi
dae
Eoch
ione
lasm
usoh
taiY
AM
AG
UCH
I,19
90N
Fiji,
Lau,
Man
us35
8A
rthr
opod
a,Cr
usta
cea,
Cirr
iped
ia,C
hion
elas
mat
oida
eEo
chio
nela
smus
paqu
ensi
sYA
MA
GU
CHI&
NEW
MA
N,1
997
EPR:
17°S
359
Art
hrop
oda,
Crus
tace
a,Ci
rrip
edia
,Eol
epad
idae
Ash
inka
ilepa
sse
epio
phili
aYA
MA
GU
CHI,
NEW
MA
N&
HA
SHIM
OTO
,200
4Sa
gam
iBay
(ven
tpa
rt)
360
Art
hrop
oda,
Crus
tace
a,Ci
rrip
edia
,Eol
epad
idae
Leuc
olep
aslo
nga
SOU
THW
ARD
&JO
NES
,200
3Ta
bar-
Feni
:ES
361
Art
hrop
oda,
Crus
tace
a,Ci
rrip
edia
,Eol
epad
idae
Neo
lepa
sra
panu
iiJO
NES
,199
3EP
R:23
°S36
2A
rthr
opod
a,Cr
usta
cea,
Cirr
iped
ia,E
olep
adid
aeN
eole
pas
zevi
nae
NEW
MA
N,1
979
EPR:
21°N
-9°N
363
Art
hrop
oda,
Crus
tace
a,Ci
rrip
edia
,Eol
epad
idae
Vul
cano
lepa
sos
heai
(BU
CKER
IDG
E,20
00)
Ker
mad
ec:B
S36
4A
rthr
opod
a,Cr
usta
cea,
Cirr
iped
ia,E
olep
adid
aeV
ulca
nole
pas
pare
nsis
SOU
THW
ARD
,200
5PA
R:37
°S-3
8°S
365
Art
hrop
oda,
Crus
tace
a,Ci
rrip
edia
,Neo
brac
hyle
padi
dae
Neo
brac
hyle
pas
relic
aN
EWM
AN
&YA
MA
GU
CHI,
1995
Lau
366
Art
hrop
oda,
Crus
tace
a,Ci
rrip
edia
,Neo
verr
ucid
aeIm
bric
aver
ruca
yam
aguc
hiiN
EWM
AN,2
000
Lau:
HH
367
Art
hrop
oda,
Crus
tace
a,Ci
rrip
edia
,Neo
verr
ucid
aeN
eove
rruc
abr
achy
lepa
dofo
rmis
NEW
MA
N,1
989
Mar
iana
368
Art
hrop
oda,
Crus
tace
a,Le
ptos
trac
a,N
ebal
iidae
Dah
lella
cald
arie
nsis
HES
SLER
,198
4EP
R:21
°N-1
8°S;
Gal
apag
os36
9A
rthr
opod
a,Cr
usta
cea,
Cum
acea
,Bod
otri
idae
Atl
anto
cum
abi
dent
atum
LED
OY
ER,1
988
EPR:
13°N
;non
-ven
tlo
cati
ons
370
Art
hrop
oda,
Crus
tace
a,Cu
mac
ea,B
odot
riid
aeBa
thyc
uma
brev
iost
re(N
ORM
AN,1
879)
MA
R:LS
371
Art
hrop
oda,
Crus
tace
a,Ta
naid
acea
,Fam
.ind
et.
Arm
atur
atan
ais
atla
ntic
usLA
RSEN
,BLA
ZEW
ICZ-
PASZ
KO
WY
CZ&
CUH
NA,i
npr
ess
MA
R:LS
373
Art
hrop
oda,
Crus
tace
a,Ta
naid
acea
,Col
lete
idae
Lept
ogna
thie
llafr
agili
sLA
RSEN
,BLA
ZEW
ICZ-
PASZ
KO
WY
CZ&
CUH
NA,i
npr
ess
MA
R:LS
374
Art
hrop
oda,
Crus
tace
a,Ta
naid
acea
,Lep
toch
eliid
aeM
esot
anai
sst
yxis
LARS
EN,B
LAZE
WIC
Z-PA
SZK
OW
YCZ
&CU
HN
A,i
npr
ess
MA
R:LS
375
Art
hrop
oda,
Crus
tace
a,Ta
naid
acea
,Not
otan
aida
eO
besu
tana
issi
grid
iLA
RSEN
,BLA
ZEW
ICZ-
PASZ
KO
WY
CZ&
CUH
NA,i
npr
ess
MA
R:LS
376
Art
hrop
oda,
Crus
tace
a,Ta
naid
acea
,Not
otan
aida
eTy
phlo
tana
isin
cogn
itus
LARS
EN,B
LAZE
WIC
Z-PA
SZK
OW
YCZ
&CU
HN
A,i
npr
ess
MA
R:LS
377
528
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Art
hrop
oda,
Crus
tace
a,Ta
naid
acea
,Pse
udot
anai
dae
Pseu
dota
nais
vuls
ella
BIRD
&H
OLD
ICH,1
989
MA
R:LS
;non
-ven
tlo
cati
ons
378
Art
hrop
oda,
Crus
tace
a,Is
opod
a,Is
chno
mes
idae
Het
erom
esus
calc
arCU
NH
A&
WIL
SON,i
npr
ess
MA
R:LS
381
Art
hrop
oda,
Crus
tace
a,Is
opod
a,Is
chno
mes
idae
Het
erom
esus
cten
obas
ius
CUN
HA
&W
ILSO
N,i
npr
ess
MA
R:LS
380
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,A
mpe
lisci
dae
Am
pelis
caro
mig
iBA
RNA
RD,1
954
EPR,
Gua
ymas
,non
-ven
tlo
cati
ons
383
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,A
mph
iloch
idae
Git
anop
sis
alvi
naBE
LLA
N-S
AN
TIN
I&
THU
RSTO
N,1
996
MA
R:LS
384
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,A
orid
aeA
uton
oelo
ngic
orni
sCH
EVRE
UX,1
909
MA
R:LS
385
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Ca
lliop
iidae
Ora
dare
alo
ngim
ana
(BO
ECK,1
871)
JFR;
non-
vent
loca
tion
s38
6A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Capr
ellid
aeCa
prel
laba
thyt
atos
MA
RTIN
&PE
TTIT
,199
8JF
R38
7A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Eusi
rida
eBo
uvie
rella
curt
iram
aBE
LLA
N-S
AN
TIN
I&
THU
RSTO
N,1
996
MA
R:LS
388
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Eu
siri
dae
Luck
iast
riki
BELL
AN-S
AN
TIN
I&
THU
RSTO
N,1
996
MA
R:LS
389
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Is
chyr
ocer
idae
Bonn
iere
llaco
mpa
rM
YER
S&
CUN
HA,2
004
MA
R:LS
390
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Is
chyr
ocer
idae
Bonn
iere
llalin
eari
sBA
RNA
RD,1
964
JFR,
Expl
orer
,non
-ven
tlo
cati
ons
391
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Ly
sian
assi
dae
Apo
tect
onia
hete
rost
egos
BARN
ARD
&IN
GRA
M,1
990
Gal
apag
os39
2A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Lysi
anas
sida
eCy
cloc
aris
tahi
tens
isST
EBBI
NG,1
888
Gua
ymas
,non
-ven
tlo
cati
ons
393
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Ly
sian
assi
dae
Euon
yxm
ytilu
sBA
RNA
RD&
ING
RAM
,199
0EP
R:13
°N,G
alap
agos
394
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Ly
sian
assi
dae
Hir
onde
llea
glut
onis
BARN
ARD
&IN
GRA
M,1
990
Gal
apag
os,E
PR39
5A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Lysi
anas
sida
eO
rcho
men
e(A
byss
orch
omen
e)ab
ysso
rum
STEB
BIN
G,1
888
Gal
apag
os,n
on-v
ent
loca
tion
s39
6A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Lysi
anas
sida
eO
rcho
men
e(A
byss
orch
omen
e)di
stin
ctus
BIRS
TEIN
&V
INO
GRA
DO
V,1
960
EPR:
13°N
,non
-ven
tlo
cati
ons
397
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Ly
sian
assi
dae
Tect
oval
opsi
sdi
abol
usBA
RNA
RD&
ING
RAM
,199
0EP
R:13
°N39
8A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Lysi
anas
sida
eTe
ctov
alop
sis
weg
ener
iBA
RNA
RD&
ING
RAM
,199
0EP
R:13
°N39
9A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Lysi
anas
sida
eTr
anst
ecto
nia
torr
enti
sBA
RNA
RD&
ING
RAM
,199
0EP
R:13
°N40
0A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Oed
icer
otid
aeBa
thym
edon
curt
ipal
pus
VIN
OG
RAD
OV,1
993
EPR
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,Pa
rdal
isci
dae
Hal
ice
hesm
onec
tes
MA
RTIN
,FRA
NCE
&V
AN
DO
VER
,199
3EP
R:9°
N40
1A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Pard
alis
cida
ePa
rdal
isca
ende
avou
riSH
AW
,198
9Ex
plor
er40
2A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Phox
ocep
halid
aeH
arpi
niop
sis
fulg
ens
BARN
ARD
,196
0EP
R,G
uaym
asA
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Sebi
dae
Seba
prof
undu
sSH
AW
,198
9Ex
plor
er40
3A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Steg
ocep
halid
aeSt
eleu
ther
aec
opro
phyc
eaBE
LLA
N-S
AN
TIN
I&
THU
RSTO
N,1
996
MA
R:LS
404
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,St
enot
hoid
aeM
etop
a(P
rom
etop
a)sa
msi
luna
BARN
ARD
,196
6G
uaym
as,n
on-v
ent
loca
tion
s40
5A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Sten
otho
idae
Sten
otho
em
enez
gwen
iBEL
LAN-S
AN
TIN
I,20
05M
AR:
MG
406
Art
hrop
oda,
Crus
tace
a,A
mph
ipod
a,St
enot
hoid
aeTo
rom
etop
asa
ldan
hae
BELL
AN-S
AN
TIN
I,20
05M
AR:
MG
;LS;
RB40
7A
rthr
opod
a,Cr
usta
cea,
Am
phip
oda,
Uri
stid
aeVe
ntie
llasu
lfur
isBA
RNA
RD&
ING
RAM
,199
0G
alap
agos
;EPR
408
Art
hrop
oda,
Crus
tace
a,Eu
phau
siac
ea,E
upha
usiid
aeTh
ysan
oess
apa
rva
HA
NSE
N,1
905
MA
R:RB
;non
-ven
tlo
cati
ons
409
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Nem
atoc
arci
nida
eN
emat
ocar
cinu
sbu
ruko
vsky
iKO
MA
I&
SEG
ON
ZAC,
2005
EPR:
17°S
,23°
S;PA
R:31
°S41
1A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,A
lvin
ocar
idid
aeA
lvin
ocar
isbr
evit
elso
nis
KIK
UCH
I&
HA
SHIM
OTO
,200
0O
kina
wa
T:M
EK41
3A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,A
lvin
ocar
idid
aeA
lvin
ocar
isdi
ssim
ilis
KO
MA
I&
SEG
ON
ZAC,
2005
Oki
naw
aT:
MEK
414
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Alv
inoc
aris
long
iros
tris
KIK
UCH
I&
OH
TA,1
995
Ker
mad
ec:B
S;O
kina
wa
T:IR
,HK
;41
5no
n-ve
ntlo
cati
ons
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Alv
inoc
aris
lusc
aW
ILLI
AM
S&
CHA
CE,1
982
EPR:
9°N
;Gal
apag
os41
6A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,A
lvin
ocar
idid
aeA
lvin
ocar
ism
arke
nsis
WIL
LIA
MS,
1988
MA
R:BS
,LO
,LS,
RB,S
P,TA
G41
7A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,A
lvin
ocar
idid
aeA
lvin
ocar
isni
wa
WEB
BER,
2004
Ker
mad
ec:B
S,RV
S41
9A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,A
lvin
ocar
idid
aeA
lvin
ocar
isw
illia
msi
SHA
NK
&M
ART
IN,2
003
MA
R:M
G42
0
529
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Phyl
um
,C
lass
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rder
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mil
ySp
ecie
sD
istr
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ge
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Chor
ocar
isch
acei
(WIL
LIA
MS
&RO
NA,1
986)
MA
R:LO
,LS,
RB,S
P,TA
G42
1A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,A
lvin
ocar
idid
aeCh
oroc
aris
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exa
MA
RTIN
&SH
AN
K,2
005
EPR:
17°S
422
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Chor
ocar
isva
ndov
erae
(MA
RTIN
&H
ESSL
ER,1
990)
Lau,
Mar
iana
,Man
us,N
Fiji
423
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Mir
ocar
isfo
rtun
ata
(MA
RTIN
&CH
RIST
IAN
SEN,1
995)
MA
R:BS
,LO
,LS,
MG
,RB,
SP,T
AG
424
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Mir
ocar
isin
dica
KO
MA
I,M
ART
IN,Z
ALA
,TSU
CHID
A&
HA
SHIM
OTO
,in
pres
sCI
R:K
VF,
EVF
425
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Nau
tilo
cari
ssa
intl
aure
ntae
KO
MA
I&
SEG
ON
ZAC,
2005
Lau,
NFi
ji42
6A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,A
lvin
ocar
idid
aeO
paep
ele
loih
iWIL
LIA
MS
&D
OBB
S,19
95H
awai
i:Lo
ihiS
eam
ount
427
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Rim
icar
isex
ocul
ata
WIL
LIA
MS
&RO
NA,1
986
MA
R:LS
,RB,
SP,T
AG
428
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Alv
inoc
arid
idae
Rim
icar
iska
irei
WA
TABE
&H
ASH
IMO
TO,2
002
CIR:
KV
F,EV
F42
9A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,A
lvin
ocar
idid
aeSh
inka
icar
isle
urok
olos
(KIK
UCH
I&
HA
SHIM
OTO
,200
0)O
kina
wa
T:M
EK43
0A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,H
ippo
lyti
dae
Lebb
eus
cari
natu
sD
ESA
INT
LAU
REN
T,19
84EP
R:13
°N43
1A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,N
ephr
opid
aeTh
ymop
ides
laur
enta
eSE
GO
NZA
C&
MA
CPH
ERSO
N,2
004
MA
R:SP
433
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Para
pagu
rida
ePa
ragi
opag
urus
vent
ilatu
sLE
MA
ITRE
,200
4O
kina
wa
A:N
ET43
5A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,Li
thod
idae
Para
lom
ishi
rtel
laD
ESA
INLA
URE
NT
&M
ACP
HER
SON,1
997
Lau:
HH
;NFi
ji:W
L43
6A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,Li
thod
idae
Para
lom
isve
rrill
i(BE
NED
ICT,
1895
)JF
R:A
V;no
n-ve
ntlo
cati
ons
437
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
ida
mag
nian
tenn
ulat
aBA
BA&
TÜRK
AY,1
992
Lau:
HH
438
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
idop
sis
acut
ispi
naBE
NED
ICT,
1902
MA
R:LC
;non
-ven
tlo
cati
ons
439
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
idop
sis
alvi
sca
WIL
LIA
MS,
1988
Expl
orer
:MM
;Gua
ymas
;JFR
440
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
idop
sis
diom
edea
e(F
AX
ON,1
893)
EPR
441
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
idop
sis
exut
aM
ACP
HER
SON
&SE
GO
NZA
C,20
05M
AR:
SP,T
AG
442
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
idop
sis
laue
nsis
BABA
&D
ESA
INT
LAU
REN
T,19
92La
u:V
F,H
H;N
Fiji:
WL
443
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
idop
sis
lent
igo
WIL
LIA
MS
&VA
ND
OV
ER,1
983
EPR:
21°N
444
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
idop
sis
mar
iani
caW
ILLI
AM
S&
BABA
,198
9M
anus
445
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Gal
athe
idae
Mun
idop
sis
sonn
eBA
BA,1
995
NFi
ji44
6A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,G
alat
heid
aeM
unid
opsi
sst
arm
erBA
BA&
DE
SAIN
TLA
URE
NT,
1992
NFi
ji:W
L44
7A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,G
alat
heid
aeM
unid
opsi
sW
HIT
EAV
ES,1
874
Gal
apag
os;E
PR44
8A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,G
alat
heid
aeSh
inka
iacr
osni
eriB
ABA
&W
ILLI
AM
S,19
98O
kina
wa
A:N
ET;O
kina
wa
T:IR
,NIK
;Tab
ar-F
eni:
ES;
449
Ryuk
uIs
land
sA
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,K
iwai
dae
Kiw
ahi
rsut
aM
ACP
HER
SON,J
ON
ES&
SEG
ON
ZAC,
2005
PAR:
38°S
450
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Chir
osty
lidae
Uro
ptyc
hus
bica
vus
BABA
&D
ESA
INT
LAU
REN
T,19
92N
Fiji:
MV
452
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Chir
osty
lidae
Uro
ptyc
hus
edis
onic
usBA
BA&
WIL
LIA
MS,
1998
Taba
r-Fe
ni:E
S45
3A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,Ch
iros
tylid
aeU
ropt
ychu
sth
erm
alis
BABA
&D
ESA
INT
LAU
REN
T,19
92N
Fiji:
WL;
non-
vent
loca
tion
s45
4A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,M
ajid
aeM
acro
rego
nia
mac
roch
ira
SAK
AI,
1978
Expl
orer
,JFR
,non
-ven
tlo
cati
ons
456
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Port
unid
aeBa
thyn
ecte
sm
arav
igna
(PRE
STA
ND
REA,1
839)
MA
R:LS
,MG
;non
-ven
tlo
cati
ons
457
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Ger
yoni
dae
Chac
eon
affi
nis
MIL
NE-
EDW
ARD
S&
BOU
VIE
R,18
94M
AR:
MG
,LS;
non-
vent
loca
tion
s45
8A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,By
thog
raei
dae
Allo
grae
ato
men
tosa
GU
INO
T,H
URT
AD
O&
VRI
JEN
HO
EK,2
002
PAR:
31°S
459
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Byth
ogra
eida
eA
usti
nogr
aea
alay
seae
GU
INO
T,19
90La
u:H
H,V
L;M
anus
;NFi
ji:W
L46
0A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,By
thog
raei
dae
Aus
tino
grae
aro
drig
ueze
nsis
TSU
CHID
A&
HA
SHIM
OTO
,200
2CI
R:K
VF,
EVF
462
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Byth
ogra
eida
eA
usti
nogr
aea
will
iam
siH
ESSL
ER&
MA
RTIN
,198
9M
aria
na46
3A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,By
thog
raei
dae
Aus
tino
grae
ayu
noha
naTA
KED
A,H
ASH
IMO
TO&
OH
TA,2
000
Izu
464
530
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Byth
ogra
eida
eBy
thog
raea
gala
page
nsis
GU
INO
T&
HU
RTA
DO
,200
3G
alap
agos
465
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Byth
ogra
eida
eBy
thog
raea
inte
rmed
iaD
ESA
INT
LAU
REN
T,19
88G
alap
agos
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Byth
ogra
eida
eBy
thog
raea
laub
ieri
GU
INO
T&
SEG
ON
ZAC,
1997
EPR:
11°S
-23°
S;PA
R:31
°S46
6A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,By
thog
raei
dae
Byth
ogra
eam
icro
psD
ESA
INT
LAU
REN
T,19
89EP
R:21
°N-2
3°S;
PAR:
31°S
-38°
S;G
alap
agos
468
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Byth
ogra
eida
eBy
thog
raea
ther
myd
ron
WIL
LIA
MS,
1980
EPR:
21°N
-18°
S;G
alap
agos
469
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Byth
ogra
eida
eBy
thog
raea
vrije
nhoe
kiG
UIN
OT
&H
URT
AD
O,2
003
PAR:
31°S
-38°
S47
0A
rthr
opod
a,Cr
usta
cea,
Dec
apod
a,By
thog
raei
dae
Cyan
agra
eapr
eada
tor
DE
SAIN
TLA
URE
NT,
1984
EPR:
21°N
,13°
N,1
4°S,
18°S
472
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Byth
ogra
eida
eSe
gonz
acia
mes
atla
ntic
a(W
ILLI
AM
S,19
88)
MA
R:LO
,LS,
MG
,RB,
SP,T
AG
473
Art
hrop
oda,
Crus
tace
a,D
ecap
oda,
Varu
nida
eX
enog
raps
uste
stud
inat
usN
G,H
UA
NG
&H
O,2
000
Oki
naw
aA
:NET
474
Echi
node
rmat
a,Cr
inoi
dea,
Cyrt
ocri
nida
,Hyo
crin
idae
Hyo
crin
usbi
scoi
toiR
OU
X,2
004
EPR:
13°N
;non
-ven
tlo
cati
ons
475
Echi
node
rmat
a,Cr
inoi
dea,
Cyrt
ocri
nida
,Hyo
crin
idae
Laub
ieri
crin
uspe
ntag
onal
isRO
UX,2
004
NFi
ji;no
n-ve
ntlo
cati
ons
476
Echi
node
rmat
a,A
ster
oide
a,Br
isin
gida
,Bri
sing
idae
Bris
inga
ende
cacn
emos
ASB
JØRN
SEN,1
856
MA
R:LO
;non
-ven
tlo
cati
ons
477
Echi
node
rmat
a,A
ster
oide
a,Br
isin
gida
,Fre
yelli
dae
Frey
ella
PERR
IER,
1885
EPR:
13°N
;PA
R;La
u,N
Fiji
478
Echi
node
rmat
a,Ec
hino
idea
,Ech
inid
aeEc
hinu
sal
exan
driD
AN
IELS
SEN
&K
ORE
N,1
883
MA
R:LS
;non
-ven
tlo
cati
ons
479
Echi
node
rmat
a,H
olot
huro
idea
,Chi
rido
tida
eCh
irid
ota
hydr
othe
rmic
aSM
IRN
OV
&G
EBRU
K,2
000
EPR:
17°S
-21°
S;M
anus
,NFi
ji48
0Ec
hino
derm
ata,
Oph
iuro
idea
,Oph
iuri
da,O
phia
ctid
aeO
phia
ctis
tyle
riST
ÖH
R&
SEG
ON
ZAC,
2005
MA
R:M
G,3
8°N
482
Echi
node
rmat
a,O
phiu
roid
ea,O
phiu
rida
,Oph
iuri
dae
Oph
ioct
enel
laac
ies
TYLE
Ret
al.,
1995
MA
R;no
n-ve
ntlo
cati
ons
483
Echi
node
rmat
a,O
phiu
roid
ea,O
phiu
rida
,Oph
iuri
dae
Spin
ophi
ura
jolli
veti
STÖ
HR
&SE
GO
NZA
C,20
06EP
R:13
°N-9
°N,1
7°S-
18°S
;PA
R:38
°S48
4Ec
hino
derm
ata,
Oph
iuro
idea
,Oph
iuri
da,O
phia
cant
hida
eO
phio
lam
ina
epra
eST
ÖH
R&
SEG
ON
ZAC,
2006
EPR:
13°N
,9°N
,17°
S-18
°S48
5Ch
aeto
gnat
ha,S
pade
llida
eCa
lispa
della
alat
aCS
AN
OVA
&M
ORE
AU,2
005
MA
R:LS
486
Hem
icho
rdat
a,En
tero
pneu
sta,
,Sax
ipen
diid
aeSa
xipe
ndiu
mco
rona
tum
WO
OD
WIC
K&
SEN
SEN
BAU
GH,1
985
EPR:
21°N
-17°
S;PA
R:38
°S;G
alap
agos
487
Chor
data
,Cho
ndri
chth
yes,
Chim
aeri
form
es,C
him
aeri
dae
Hyd
rola
gus
affi
nis
(CA
PELL
O,1
868)
MA
R:LS
,MS,
RB;n
on-v
ent
loca
tion
s49
1Ch
orda
ta,C
hond
rich
thye
s,Ch
imae
rifo
rmes
,Chi
mae
rida
eH
ydro
lagu
spa
llidu
sH
ARD
Y&
STEH
MA
NN,1
990
MA
R:LS
,Men
ezH
om,M
S;RB
;non
-ven
tlo
cati
ons
492
Chor
data
,Ost
eich
thye
s,A
ngui
llifo
rmes
,Syn
apho
bran
chid
aeD
ysom
min
aru
gosa
GIN
SBU
RG,1
951
Intr
a-pl
ate
Seam
ount
:Vai
lulu
493
Chor
data
,Ost
eich
thye
s,A
ngui
llifo
rmes
,Syn
apho
bran
chid
aeIly
ophi
ssa
ldan
haiK
ARM
OV
SKA
YA&
PARI
N,1
999
EPR:
21°S
;MA
R:BS
494
Chor
data
,Ost
eich
thye
s,G
adif
orm
es,S
ynap
hobr
anch
idae
Ther
mob
iote
sm
ytilo
geit
onG
EIST
DO
RFER
,199
1La
u:V
V49
6Ch
orda
ta,O
stei
chth
yes,
Gad
ifor
mes
,Lot
idae
Gai
drop
saru
sRA
FIN
ESQ
UE,
1810
MA
R:LS
497
Chor
data
,Ost
eich
thye
s,G
adif
orm
es,M
orid
aeLe
pidi
onsc
hmid
tiSV
ETO
VID
OV,1
936
MA
R:M
G;n
on-v
ent
loca
tion
s49
8Ch
orda
ta,O
stei
chth
yes,
Oph
idiif
orm
es,B
ythi
tida
eCa
taet
yxla
tice
psK
OEF
OED
,192
7M
AR:
MG
,MS,
Men
ezH
om,L
S,RB
;non
-ven
tlo
cati
ons
499
Chor
data
,Ost
eich
thye
s,O
phid
iifor
mes
,Byt
hiti
dae
Ther
mic
hthy
sho
llisi
(CO
HEN
,RO
SEN
BLA
TT&
MO
SER,
1990
)EP
R:17
°S,2
1°S;
Gal
apag
os50
0Ch
orda
ta,O
stei
chth
yes,
Oph
idiif
orm
es,O
phid
iidae
Vent
icht
hys
bios
peed
oiN
IELS
EN,M
OLL
ER&
SEG
ON
ZAC,
inpr
ess
EPR:
17°S
501
Chor
data
,Ost
eich
thye
s,Pe
rcif
orm
es,Z
oarc
idae
Pach
ycar
agy
mni
nium
AN
DER
SON
&PE
DEN
,198
8JF
R:A
V,ES
;non
-ven
tlo
cati
ons
502
Chor
data
,Ost
eich
thye
s,Pe
rcif
orm
es,Z
oarc
idae
Pach
ycar
ari
mae
AN
DER
SON,1
989
Gal
apag
os50
3Ch
orda
ta,O
stei
chth
yes,
Perc
ifor
mes
,Zoa
rcid
aePa
chyc
ara
sald
aniB
ISCO
ITO
&A
LMEI
DA,2
004
MA
R:RB
504
Chor
data
,Ost
eich
thye
s,Pe
rcif
orm
es,Z
oarc
idae
Pach
ycar
ath
erm
ophi
lum
GEI
STD
ORF
ER,1
994
MA
R:SP
,TA
G50
5
Chor
data
,Ost
eich
thye
s,Pe
rcif
orm
es,Z
oarc
idae
Pyro
lycu
sm
anus
anus
MA
CHID
A&
HA
SHIM
OTO
T,20
02M
anus
506
Chor
data
,Ost
eich
thye
s,Pe
rcif
orm
es,Z
oarc
idae
Ther
mar
ces
cerb
erus
ROSE
NBL
ATT
&CO
HEN
,198
6EP
R:21
°N,1
3°N
,9°N
;Gal
apag
os50
7
Chor
data
,Ost
eich
thye
s,Sc
orpa
enif
orm
es,L
ipar
idae
Care
proc
tus
hyal
eius
GEI
STD
ORF
ER,1
994
EPR:
13°N
-9°N
509
Chor
data
,Ost
eich
thye
s,Sc
orpa
enif
orm
es,S
ebas
tida
eTr
achy
scor
pia
cris
tula
taec
hina
ta(K
OEH
LER,
1896
)M
AR:
MG
;non
-ven
tlo
cati
ons
510
Chor
data
,Ost
eich
thye
s,G
adif
orm
es,M
acro
urid
aeCo
ryph
aeno
ides
arm
atus
HEC
TOR,
1875
EPR:
9°N
;Gal
apag
os;M
AR:
LS,R
B,SP
;51
1no
nve
ntlo
cati
ons
531
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
532
Cnidaria, Anthozoa
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
533
Mollusca, Gastropoda
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
534
Mollusca, Bivalvia
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
535
Mollusca, Cephalopoda
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
536
Annelida, Polychaeta
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
537
Annelida, Polychaeta
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
538
Arthropoda, Decapoda, Carididae
Arthropoda, Amphipoda
Total length (TL): from the tip of the rostrum to the tip of the telson. Cephalothorax length (CL): from the level of the posteriormargin of the orbit to the midpoint of the posterodorsal margin of the carapace.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
539
Arthropoda, Copepoda
Arthropoda, Ostracoda
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
540
Arthropoda, Decapoda, Brachyura
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
541
Vertebrata, Osteichthyes
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Genus index
Abyssocladia, 35, 37f., 519Abyssorchomene, 396f.Abyssotherma, 31, 33, 519Acharax, 149, 523Achelia, 305Actinostola, 66, 71, 519Adula, 207Agathotanais, 372Alaysia, 255, 525Allograea, 455, 459, 470, 530Alvania, 133, 522Alvinella, 14, 27, 104, 106, 108f., 116, 118,
182, 200, 282ff., 341, 399, 525Alvinocaris, 360, 410, 413-421, 529Alvinoconcha, 15, 82, 128, 130, 422, 425,
480, 522Amathys, 295, 526Ambilimbus, 344, 528Ammothea, 301, 306, 526Ampelisca, 383, 529Amphicrossus, 344Amphiduropsis, 198, 524Amphisamytha, 130, 183, 190, 296, 526Andromastax, 318, 526Anisopes, 304Aphotopontius, 316, 328f., 338, 526f.Apotectonia, 392, 529Arborammina, 32Archiconchoecia, 307, 526Archinome, 183, 185, 524Arcovestia, 18, 70, 256, 525Argestoides, 320, 526Armaturatanais, 372f., 528Asbestopluma, 39ff., 519Ashinkailepas, 356, 360, 362f., 528Atlantocuma, 370, 528Austinograea, 35, 425, 455, 460-466, 530Autonoe, 385, 529Barathricola, 326, 526Bathybdella, 29, 297, 526Bathyconchoecia, 308f., 526Bathycuma, 371, 528Bathykurila, 218, 524Bathylaophonte, 345, 528Bathymargarites, 101, 521Bathymedon, 529Bathymodiolus, 13, 15, 29, 68ff., 80f., 83, 85,
91, 97, 115, 125, 137, 141, 150-161,194, 206, 208, 231, 243f., 297, 327,353, 359, 379, 385, 411, 421, 424f.,429, 430, 450, 458, 461, 469, 473,483ff., 491, 496, 501, 508, 523
Bathynectes, 457, 530Bathypecten, 141, 164, 523Beggiatoa, 188, 205Benthoctopus, 166, 171, 523Benthomisophria, 349Benthoxynus, 330, 340, 527
Boloceroides, 65, 72, 519Bonnierella, 390f., 529Bouillonia, 48Bouvierella, 382, 388, 529Brachylepas, 366Branchinotogluma, 183, 219-229, 301, 524Branchiplicatus, 230, 524Branchipolynoe, 14, 160, 162, 231-234, 524Brisinga, 477, 531Bruceiella, 94, 520Bythograea, 411, 455, 463, 465-472, 530f.Calispadella, 486, 531Callinectes, 297Calocarcinus, 455Calyptogena, 13, 77, 80f., 101, 106, 141-148,
209f., 215, 219, 230, 297, 334, 337,360, 480, 484, 523
Candelabrum, 46, 49f., 140, 424, 519Caprella, 387, 529Carcinonemertes, 29Careproctus, 509, 531Cataetyx, 499, 531Caulophacus, 47, 519Cephalochaetosoma, 178, 523Cerianthus, 72Ceuthoecetes, 29, 331, 527Chaceon, 455, 458, 530Chasmatopontius, 332, 527Cheramomyzon, 323, 526Chionelasmus, 358Chionoecetes, 297Chiridota, 480, 531Cholidya, 29, 354, 528Chondrocladia, 36, 42f., 519Chondrophellia, 73, 164, 501, 519Chorocaris, 15, 341, 404, 419-423, 466, 505,
530Cirroteuthis, 166, 168, 523Cirrothauma, 167, 523Cladorhiza, 44f., 519Clypeosectus, 86, 520Collocherides, 324, 526Copidognathus, 299, 526Coryphaenoides, 442, 511, 531Ctenopelta, 111, 521Cyanagraea, 14, 297, 455, 472, 531Cyananthea, 65, 67, 519Cyathermia, 104, 521Cyclocaris, 393, 529Cyclopina, 316Dahlella, 369, 528Dendronotus, 140, 523Depressigyra, 82, 112, 521Desbruyeresia, 129, 522Desmodora, 174f., 523Desmodorella, 176f., 523Dinetia, 179, 523Dirivultus, 316, 333, 527
Dysommina, 493, 531Ecbathyrion, 343, 528Echinopelta, 113, 521Echinus, 479, 531Ectopleura, 48, 52, 519Eochionelasmus, 356-359, 362f., 366, 480,
528Eosipho, 124f., 459, 522Erebonaster, 344Euchelipluma, 46, 519Eudendrium, 51, 519Eulalia, 215, 524Eulepedopsis, 83, 520Eunice, 193, 524Euonyx, 394, 529Euphilomedes, 310, 526Exallopus, 187, 524Exrima, 334, 527Fissuricola, 335, 527Freyella, 478, 531Fucaria, 95, 520Gaidropsarus, 497, 531Galapagomystides, 18, 216, 524Genesis, 29, 170, 355, 528Gigantidas, 162f., 419, 523Gitanopsis, 384, 529Glycera, 195ff., 524Gordotanais, 372Gorgoleptis, 87, 520Grammaria, 55, 519Graneledone, 166, 172, 523Grievella, 351, 528Grimpoteuthis, 166, 169, 523Halacarellus, 300, 526Halecium, 48Halice, 170, 382, 401, 529Halisiphonia, 54, 519Harmothoe, 183Harpiniopsis, 529Helicoradomenia, 75-79, 520Helicrenion, 102, 521Heptnerina, 316, 327, 526Hesiodeiria, 199, 524Hesiolyra, 183, 200f., 524Hesiospina, 183, 202f., 524Heteromesus, 379ff., 529Hirondellea, 395, 529Hirtopelta, 114, 521Humesipontius, 336, 527Hyalogyrina, 135f., 522Hyalopontius, 347, 528Hydractinia, 48Hydrallmania, 59, 519Hydrolagus, 489, 491f., 531Hyocrinus, 475, 531Hyphalion, 325, 526Hypoechinorhynchus, 29, 181, 523Hysterothylacium, 29
542
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Ifremeria, 15, 82, 110, 121, 130f., 227, 249,436, 460, 480, 522
Iheyomytilidicola, 206, 524Ilyophis, 494f., 531Imbricaverruca, 357, 362, 367, 528Iphionella, 235, 524Isaacsicalanus, 352, 528Jamstecia, 321, 526Johanssonia, 297Kiwa, 450f., 530Lacunoides, 105, 521Laeviphitus, 127, 522Lafoea, 56, 519Laitmatobius, 346, 528Lamellibrachia, 257ff., 333, 410, 525Laminatubus, 125, 164, 183, 253, 509, 525Laonice, 273, 525Laubiericrinus, 476, 531Laubieriellus, 274, 525Lebbeus, 360, 431f., 461, 530Leitoscoloplos, 268, 525Lepetodrilus, 15, 67, 82, 88ff., 152, 520Lepidion, 498, 531Lepidonotopodium, 183, 236-242, 524f.Leptochiton, 80f., 520Leptognathia, 372Leptognathiella, 374, 528Leptogyra, 96, 520Leucolepas, 357, 361f., 365, 528Levensteiniella, 183, 243ff., 525Lindaspio, 275ff., 525Lirapex, 115, 521Lucernaria, 64, 519Luckia, 382, 389, 529Luffammina, 32, 519Lurifax, 136, 522Macroregonia, 386f. 456, 530Maractis, 68, 462, 519Marianactis, 69, 156, 158, 426, 429, 519Melanodrymia, 106, 521Mesotanais, 375, 528Methanotrix, 32Metopa, 405, 529Miralvinella, 285ff.Mirocaris, 424f., 530Misophriopsis, 349, 528Moravecnema, 29, 180, 523Munida, 438, 530Munidopsis, 70, 411, 434, 439-448, 530Mytilidiphila, 207f., 524Myzobdella, 297Nautalvinella, 288ff.Nautilocaris, 426, 460, 530Nautilus, 166Nematocarcinus, 411f., 466, 529Neobrachylepas, 356f., 362, 366, 528Neolepas, 73, 357, 361ff., 528Neolepetopsis, 84f., 520Neomphalus, 107, 521Neoverruca, 356ff., 362, 366, 368, 528Nereimyra, 204, 524Nereis, 212ff., 524Neusas, 134, 522Nicomache, 266f., 525Nilva, 337, 527
Nodopelta, 116f., 521Nuculana, 344Oasisia, 260, 288, 525Obesutanais, 376, 528Olgaconcha, 15, 130Olgasolaris, 121, 131, 522Oncaea, 316, 350, 528Opaepele, 419, 427, 530Ophiactis, 482, 531Ophioctenella, 483, 531Ophiolamina, 481, 484f., 531Ophryotrocha, 188-191, 524Opisthotrochopodus, 219, 221, 228Oradarea, 386, 529Orbiniella, 269f., 525Orchomene, 396f., 529Pachycara, 68, 158, 180, 502-505, 531Pachydermia, 108, 521Pacmanactis, 70, 519Paragiopagurus, 435, 530Paralepetopsis, 84f., 520Paralomis, 360, 436f., 530Paralvinella, 15, 183, 285-294, 332, 525f.Paranthosactis, 71, 519Parasicyonis, 68Pardalisca, 402, 529Parougia, 183, 192, 524Periclimenes, 410Peltospira, 118, 521Phymorhynchus, 68, 138f., 156, 361, 522Planorbidella, 109, 521Polycirrus, 183Polycopetta, 311, 526Polyplumaria, 53, 519Prionospio, 278ff., 525Prionotoleberis, 312, 526Prometopa, 405Protis, 254, 525Protolira, 97, 152, 521Protomystides, 217, 524Provanna, 93, 132, 522Pseudorimula, 91f., 520Pseudotanais, 372, 378, 529Pyrolycus, 506, 531Pyropelta, 93, 520Rhogobius, 338, 527Rhynchopelta, 119, 521Rickettsia, 29Ridgeia, 89, 106, 126, 172, 261, 293, 455,
525Riftia, 14, 29, 33, 67, 71, 88, 90, 101, 104,
106, 111, 119, 170, 182f., 202, 215,219, 230, 262f., 278, 297, 307, 311,313ff., 403, 411, 416, 424, 431f., 464,468f., 484f., 508f., 525
Rimicaris, 17, 68, 156, 339, 341, 404, 417,419, 428f., 462, 505, 511, 530
Rimipontius, 339, 527Rimitantalus, 29Saxipendium, 14, 487, 531Scabrantenna, 322, 526Scoloplos, 271, 525Scotoecetes, 340, 527Seba, 403, 529Segonzacia, 455, 463, 473, 499, 531
Sericosura, 302-306, 526Sertularella, 60, 519Shinkai, 209ff., 524Shinkaia, 434, 449, 530Shinkaicaris, 413f., 419, 430, 530Shinkailepas, 122f., 522Sinepecten, 141, 165, 523Siphonalia, 435Siphonobrachia, 264, 525Sirsoe, 205, 524Smacigastes, 353, 528Speculator, 126, 522Speleophriopsis, 349Sphaerosyllis, 252, 525Spinophiura, 484f., 531Spiochaetopterus, 272, 525Stegolaria, 62, 519Steleuthera, 404, 529Stenhelia, 348, 528Stenothoe, 406, 529Stygiopontius, 341f., 527f.Sutilizona, 98, 521Symmetromphalus, 110, 521Symplectoscyphus, 61, 519Tectovalopsis, 398f., 529Temnocinclis, 99, 521Temnozaga, 100, 521Tevnia, 65, 88, 265, 288, 525Thalassomonohystera, 173Thermaloniscus, 379Thermarces, 181, 500, 507ff., 531Thermichthys, 500, 531Thermiphione, 246ff., 525Thermobiotes, 496, 531Thermochiton, 80f., 520Thermonemertes, 183, 523Thermopalia, 14, 63, 519Thermopolynoe, 249f., 525Thomontocypris, 313f., 526Thraumastos, 194, 524Thymopides, 433, 442, 530Thysanoessa, 409, 529Tisbe, 354Torometopa, 407, 529Trachyscorpia, 510, 531Transtectonia, 400, 529Tubularia, 48, 53Typhlotanais, 372, 377, 528Uptionyx, 319, 526Uroptychus, 452ff., 530Vampiropolynoe, 251, 525Ventichthys, 501, 531Ventiella, 382, 408, 529Vesicomya, 141Vetulonia, 103, 521Vulcanoctopus, 166, 170, 355, 523Vulcanolepas, 145, 355f., 361, 364f., 528Xandaros, 281, 525Xenograpsus, 455, 474, 531Xylocythere, 315, 526Xylodiscula, 137, 522Zooarium, 458Zygophylax, 48, 57f., 519
543
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544
Author index
Almeida, 182, 491f., 497ff., 502ff., 510f.
Baba, 434f., 438, 440, 444ff., 448f., 452ff.
Bamber, 301-306
Barthélémy, 409
Bartsch, 299f.
Bellan-Santini, 382-408
Biscoito, 13-25, 457f., 489-495, 497ff., 502ff., 507f., 510f.
Blake, 187-193, 212f., 215ff., 252, 266-271, 274-279, 281
Böggemann, 195
Bouchet, 83f., 94f., 99-104, 107, 111ff., 119f., 124f., 130, 133ff.,137, 140
Boury-Esnault, 47
Briand, 457, 494, 496, 509
Bright, 6-9, 27f., 174-179, 518
Buron de, 29f., 181
Burreson, 297
Casanova B., 409
Casanova J.-P., 486
Causse, 494f.
Cohen, 500
Corbera, 370f.
Cosel, 80f., 83, 85, 94, 96, 99-104, 107, 111ff., 119f., 126f., 130,134f., 137, 141-148, 151-154, 156, 158, 162
Cunha, 372-381, 385, 390
Daly, 64, 72
Defaye, 319-327, 332, 335ff., 339f., 343, 346-355
Desbruyeres, 6-9, 13-25, 149, 184f., 190-194, 196f., 212ff., 218-224, 227-230, 232-245, 247-250, 252f., 256, 260, 262f., 266,268, 272f., 280, 282-296, 487, 513-517
Gebruck, 13-25, 480
Geistdoerfer, 484, 505
Gonzalez, 170
Guerra, 166, 168, 170
Guinot, 455, 459ff., 465-473
Haney, 369
Harrison-Nelson, 307-312
Hashimoto, 13-25, 148, 150, 155, 157, 159, 163, 429, 464, 506
Holzmann, 31ff.
Hourdez, 218f., 223, 227, 235ff., 242ff., 247, 251
Hurtado, 459, 465
Ivanenko, 316-355
Jaume, 386, 391-395, 398ff., 402-405
Jones, 362
Justine, 180
Komai, 410-428, 430ff.
Kornicker, 307-312
Kudenov, 186
Larsen, 373-377
Lemaitre, 435
Lopez-Gonzalez, 65-73
Lutz, 145, 160f.
Macpherson, 433-449, 452ff., 456
Maddocks, 313ff.
Mah, 477f.
Miura, 206-211, 225f., 231, 246, 249f., 259
Möller, 501
Moreau, 486
Myers, 385, 390
Newman, 356-368
Ng, 474
Nielsen, 500f.
Okutani, 138f.
Pleijel, 198-205
Pugh, 63
Ramirez-Llodra, 479
Rodríguez, 70f.
Roux, 475f.
Salvini-Plawen, 75-79
Schein, 164f.
Segonzac, 6-9, 29f., 37-46, 49-69, 73, 140, 167-170, 253f., 297,356ff., 378, 411-414, 416-424, 426, 428, 430, 433, 436f., 439,441ff., 447, 450f., 455, 460f., 463, 466-472, 474, 477, 478, 482,485, 501, 507
Senz, 183
Southward A.J., 356-368
Southward E.C., 255, 257f., 260f., 264f.
Stöhr, 481ff., 485
Tsuchida, 462
Tunnicliffe, 13-25
Tyler, 13-25, 479, 483
Vacelet, 35-46
Van Dover, 13-25,
Verschelde, 174-177
Vervoort, 48-62
Villanueva, 166f., 169
Voight, 166, 171f., 339
Vrijenhoek, 145, 160, 459
Warén, 82-123, 126-137
Wilson, 380f.
Yamaguchi, 356-368
Young, 493
Zekely, 174-179
Zibrovius, 254
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