Handbook of Deep-Sea Hydro Thermal Vent Fauna

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


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


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


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


6

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.“


7

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.


Remarks


8

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.


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]


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


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).

17


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).

Bibliography

In addition to scientific reports, there have been many ex-cellent reviews of the biology of vent and other chemosynthet-ic communities that provide entrée into the primary literatureand deliver a historical perspective on the exploration of hy-drothermal-vent biota. A selection of these reviews is listedhere.

CHILDRESS J.J. & C.R. FISHER (1992): The biology of hydrothermal vent ani-mals: Physiology, biochemistry, and autotrophic symbioses. —Oceanogr. Mar. Biol. 30: 337-441.

DESBRUYÈRES D. & M. SEGONZAC (1997): Handbook of Deep-sea Hydrother-mal Vent Fauna. — Ifremer éd., Plouzané, France: 1-279.

GEBRUK A.V. (Ed.; 2002): Biology of Hydrothermal Systems. — Moscow,KMK Press: 1-543.

GEBRUK A.V., GALKIN S.V., VERESHCHAKA A.L., MOSKALEV L.I. & A.J. SOUTHWARD

(1997): Ecology and biogeography of the hydrothermal vent faunaof the Mid-Atlantic Ridge. — Adv. Mar. Biol. 32: 93-144.

GRASSLE J.F. (1985): Hydrothermal vent animals: Distribution and biology.— Science 229: 713-717.

JOLLIVET D. (1996): Specific and genetic diversity at deep-sea hydrother-mal vents: An overview. — Biodiv. Conserv. 5: 1619-1653.

JUNIPER S.K. & V. TUNNICLIFFE (1997): Crustal accretion and the hot ecosys-tem. — Philos. Trans. R. Soc. Lond. A 355: 450-474.

TUNNICLIFFE V. (1991): The biology of hydrothermal vents: ecology andevolution. — Oceanogr. Mar. Biol. 29: 319-407.

TUNNICLIFFE V. (1992): Hydrothermal vent communities of the deep sea. —Amer. Sci. 80: 336-349.

TUNNICLIFFE V., FOWLER C.M.R. & A.G. MCARTHUR (1996): Plate tectonic his-

tory and hot vent biogeography. — In: MACLEOD C.J., TYLER P.A. &

C.L. WALKER (Eds.): Tectonic, Magmatic, Hydrothermal and Biological

Segmentation of Mid-Ocean Ridges. J. Geol. Soc. Lond. Spec. Publ.

118: 225-238.

TUNNICLIFFE V., MACARTHUR A.G. & D. MCHUGH (1998): A biogeographicalperspective of the deep-sea hydrothermal vent fauna. — Adv. Mar.Biol. 34: 353-442.

18


TUNNICLIFFE V., JUNIPER S.K. & M. SIBUET (2003): Reducing environments of

the deep-sea floor. — In: TYLER P.A. (Ed.): Ecosystems of the Deep

Oceans. Ecosystems of the World 28, Elsevier, Amsterdam: 81-110.

VAN DOVER C.L. (1995): Ecology of Mid-Atlantic Ridge hydrothermal

vents. — J. Geol. Soc. Lond. Spec. Publ. 87: 257-294.

VAN DOVER C.L. (2000): The Ecology of Deep-Sea Hydrothermal Vents. —

Princeton University Press, Princeton, NJ: 1-424.

ReferencesAnonymous (1989): Hydrothermal vent communities found in lake. —

Mar. Pollut. Bull. 20: 540-541.

ARP A.J. & J.J. CHILDRESS (1983): Sulfide binding by the blood of the hy-

drothermal vent tube worm Riftia pachyptila. — Science 219: 295-

297.

ARP A.J., CHILDRESS J.J. & C.R. FISHER (1984): Metabolic and blood gas trans-

port characteristics of the hydrothermal vent bivalve Calyptogena

magnifica. — Physiol. Zool. 57: 648-662.

AUZENDE J.-M., HASHIMOTO J., FIALA-MEDIONI A. & S. OHTA (1997): Étude

géologique et biologique in situ de deux zones hydrothermales du

bassin de Manus (Papouasie Nouvelle-Guinée). — C. R. Acad. Sci.

(Ser. 2a) 325: 585-591.

BALLARD R.D. (1977): Notes on a major oceanographic find. — Oceanus

20: 35-44.

BARBIERI R., ORI G.G. & CAVALAZZI B. (2004): A Silurian cold-seep ecosystem

from the Middle Atlas, Morocco. — Palaios 19: 527-542.

BARRIGA F.J.A.S., FOUQUET Y., ALMEIDA A., BISCOITO M.J., CHARLOU J.L., COSTA

R.L.P., DIAS A., MARQUES A.M.S.F., MIRANDA J.M.A., OLU K., PORTEIRO F. &

M.G.P.S. QUEIROZ (1998): Discovery of the Saldanha Hydrothermal

Field on the FAMOUS segment of the MAR (36°30’N). — Eos, Trans.

Amer. Geophys. Union 79: F67.

BELTENEV V.E., NESCHERETOV A.V., IVANOV V.N., V SHILOV.V., ROZHDESTVENSKAYA

I.I., SHAGIN A.A., STEPANOVA T.V., ANDREEVA I.A., SEMENOV Y.P., SERGEEV

M.B., CHERKASHEV G.A., BATUEV B.N., SAMOVAROV M.L., KROTOV A.G. &

V.F. MARKOV (2004): New hydrothermal ore field in the axial zone of

the Mid-Atlantic Ridge. — Dokl. Akad. Nauk 397: 78-82.

BLACK M.B., LUTZ R.A. & R.C. VRIJENHOEK (1994): Gene flow among vesti-

mentiferan tube worm (Riftia pachyptila) populations from hy-

drothermal vents in the eastern Pacific. — Mar. Biol. 120: 33-39.

BLAKE J.A. (1985): Polychaeta from the vicinity of deep-sea geothermal

vents in the Eastern Pacific. I. Euphrosinidae, Phylodocidae, Hesion-

idae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae and Mal-

danidae. — Proc. Biol. Soc. Wash. 6: 67-102.

BOGDANOV Y.A., SAGALEVITCH A.M., CHERNYAEV E.C., ASHADZE A.M., GURVITCH

E.G., LUKASHIN V.N., IVANOV G.V. & V.N. PERESYPKIN (1995): Hydrother-

mal field at 14°45’N on the Mid-Atlantic Ridge. — Dokl. Akad. Nauk

343: 353-357.

BOGDANOV Y.A., BORTNIKOV N.S., VIKENTEV I.V., GURVITCH E.G. & A.M. SAGALE-

VITCH (1997): A new type of modern mineral forming system: Black

smokers of the hydrothermal filed at 14°45’N on the Mid-Atlantic

Ridge. — Geol. Ore Deposits 39: 68-90.

BOTH R., CROOK K., TAYLOR B., BROGAN S., CHAPPELL B.W., FRANKEL E., LIU L.,

SINTON J. & D. TIFFIN (1986): Hydrothermal chimneys and associated

fauna in the Manus back-arc basin, Papua New Guinea. — Eos,

Trans. Amer. Geophys. Union 67: 489-490.

CAMPBELL K.A. (in press): Hydrocarbon seep and hydrothermal vent

palaeoenvironments and palaeontology: Past developments and

future research directions. — Palaeogeogr. Palaeoclimatol.

Palaeoecol.

CAMPBELL K.A., FARMER J.D. & D. DES MARAIS (2002): Ancient hydrocarbonseeps from the Mesozoic convergent margin of California: Carbon-ate geochemistry, fluids, and palaeoenvironments. — Geofluids 2:63-94.

CASM (Canadian American Seamount Expedition; 1985): Hydrothermalvents on an axis seamount of the Juan de Fuca Ridge. — Nature313: 212-214.

CAVANAUGH C.M., GARDINER S.L., JONES M.L., JANNASCH H.W. & J.B. WATERBURY

(1981): Prokaryotic cells in the hydrothermal vent tube worm Riftiapachyptila: Possible chemoautotrophic symbionts. — Science 213:340-342.

COPLEY J., TYLER P.A., MURTON B.J. & C.L. VAN DOVER (1997): Spatial and in-terannual variation in the faunal distribution at Broken Spur ventfield (29°N, Mid-Atlantic Ridge). — Mar. Biol. 129: 723-733.

COPLEY J., TYLER P.A., VAN DOVER C.L., SCHULZ A., SULANOWSKA M., SINGH S. &P. DICKSON (1999): Effects of drilling on the TAG hydrothermal ventcommunity, 26°N Mid-Atlantic Ridge. — P.S.Z.N. Mar. Ecol. 20: 291-306.

CORLISS J.B., BAROSS J.A. & S.E. HOFFMAN (1980): An hypothesis concerningthe relationship between submarine hot springs and the origin oflife on Earth. — Eos, Trans. Amer. Geophys. Union 61: 996.

CORLISS J.B., DYMOND J., GORDON L.I., EDMOND J.M., VON HERZEN R.P., BALLARD

R.D., GREEN K., WILLIAMS D., BAINBRIDGE A., CRANE K. & T.H. VAN ANDEL

(1979): Submarine thermal springs on the Galapagos Rift. — Science203: 1073-1083.

CRANE K., HECKER B. & V. GOLUBEV (1991): Heat flow and hydrothermalvents in Lake Baikal, USSR. — Eos, Trans. Amer. Geophys. Union 72:585, 588-589.

DANDO P.R., STUEBEN D. & S.P. VARNAVAS (1999): Hydrothermalism in theMediterranean Sea. — Progr. Oceanogr. 44: 333-367.

DESBRUYÈRES D. (1995): Temporal variations of deep-sea hydrothermalcommunities at 13°N/EPR. — InterRidge News 4: 6-10.

DESBRUYÈRES D. & L. LAUBIER (1980): Alvinella pompejana gen. sp. nov.,Ampharetidae aberrant des sources hydrothermales de la ride East-Pacifique. — Oceanologica Acta 3: 267-274.

DESBRUYÈRES D., CRASSOUS P., GRASSLE J., KHRIPOUNOFF A., REYSS D., RIO M. &M. VAN PRAET (1982): Données écologiques sur un nouveau site d’hy-drothermalisme actif de la ride du Pacifique oriental. — C. R. Acad.Sci. 295: 489-494.

DESBRUYÈRES D., HASHIMOTO J. & M.-C. FABRI (in press): Composition andbiogeography of hydrothermal vent communities in Western Pacif-ic back arc basins, in Back-Arc Spreading Systems: Geological, Bio-logical, Chemical and Physical Interactions. — Geophys. Monogr.

DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA & the Scientific Parties of Bi-olau and Starmer Cruises (1994): Deep-sea hydrothermal communi-ties in Southwestern Pacific back-arc basins (the North Fiji and LauBasins): Composition, microdistribution and food web. — Mar. Ge-ol. 116: 227-242.

DESBRUYÈRES D., ALMEIDA A., BISCOITO M., COMTET T., KHRIPOUNOFF A., LE BRIS

N., SARRADIN P.M. & M. SEGONZAC (2000): A review of the distributionof hydrothermal vent communities along the northern Mid-AtlanticRidge: dispersal vs. environmental controls. — Hydrobiologia 440:201-216.

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): Variations in deep-sea hy-drothermal vent communities on the Mid-Atlantic Ridge near theAzores plateau. — Deep-Sea Res. I 48: 1325-1346

EDMONDS H.N., MICHAEL P.J., BAKER E.T., CONNELLY D.P., SNOW J.E., LANGMUIR

C.H., DICK H.J.B., MUEHE R., GERMAN C.R. & D.W. GRAHAM (2003): Dis-covery of abundant hydrothermal venting on the ultraslow-spread-ing Gakkel ridge in the Arctic Ocean. — Nature 421: 252-256.

19


EINSELE G., GIESKES J.M., CURRAY J., MOORE D.M., E AGUAYO., AUBRY M.-P.,

FORNARI D., GUERRERO J., KASTNER M., KEITS K., LYLE M., MATOBA Y., MOLI-

NA-CRUZ A., NIEMITZ J., RUEDA J., SAUNDERS A., SCHRADER H., SIMONEIT B. &

V. VACQUIER (1980): Intrusion of basaltic sills into highly porous sedi-

ments, and resulting hydrothermal activity. — Nature 283: 441-445.

EMBLEY R.W., LUPTON J.E., MASSOTH G., URABE T., TUNNICLIFFE V., BUTTERFIELD

D.A., SHIBATA T., OKANO O., KINOSHITA M. & K. FUJIOKA (1998): Geolog-

ic, chemical, and biological evidence for Recent volcanism at 17.5°S;

East Pacific Rise. — Earth Planetary Sci. Lett. 163: 131-147.

EMBLEY R.W., BAKER E.T., CHADWICK W.W. Jr., LUPTON J.E., RESING K.A., MAS-

SOTH G.J. & K. NAKAMURA (2004): Explorations of Mariana Arc volca-

noes reveal new hydrothermal systems — Eos, Trans. Amer. Geo-

phys. Union 85: 37.

ENRIGHT J.T., NEWMAN W.A., HESSLER R.R. & J.A. MCGOWAN (1981): Deep-

ocean hydrothermal vent communities. — Nature 289: 219-221.

FAUGERES J.C., DESBRUYÈRES D., GONTHIER E., GRIBOULARD R., POUTIERS J., DE

RESSEGUIER A. & G. VERNETTE (1987): Sedimentary and biological wit-

ness of the present tectonic activity of the Barbados accretion

wedge. — C. R. Acad. Sci. (Ser. 2) 305: 115-119.

FELBECK H. (1981): Chemoautotrophic potential of the hydrothermal vent

tube worm Riftia pachyptila JONES (Vestimentifera). — Science 213:

336-338.

FIALA-MÉDIONI A. (1984): Mise en évidence par microscopie électronique

à transmission de l’abondance de bactéries symbiotiques dans la

branchie de mollusques bivalves de sources hydrothermales pro-

fondes. — C. R. Acad. Sci. 298: 487-492.

FOUQUET Y., ONDREAS H., CHARLOU J.-L., DONVAL J.P., RADFORD-KNOERY J., COS-

TA I., LOURENCO N. & M.K. TIVEY (1995): Atlantic lava lakes and hot

vents. — Nature 377: 201.

FOUQUET Y., CHARLOU J.L., ONDREAS H., RADFORD-KNOERY H., DONVAL J., DOU-

VILLE J.P., APPRIOU E., CAMBON R., PELL P., LANDURE H.J.Y., NORMAND A.,

PONSEVERA E., GERMAN C.R., PARSON L. M., BARRIGA F.J.A.S., COSTA I.M.A.,

RELVAS J.M.R.S. & A. RIBEIRO (1997): Discovery and first submersible in-

vestigations on the Rainbow hydrothermal field on the MAR

(36°14’N). — Eos, Trans. Amer. Geophys. Union 78: 832.

FOUQUET Y., CAMBON P., CHARLOU J.L., BARRIGA F.J.A.S., ONDREAS H., BAYON G.,

HENRY K., MURPHY P., ROUXEL O., POROSHINA I., DONVAL J.P. & J. ETOUBLEAU

(in press). Geodiversity of hydrothermal processes along the Mid-

Atlantic Ridge – Ultramafic-hosted mineralization: a new type of

oceanic Cu-Zn-Co-Au VMS deposit. — Economic Geol.

FRICKE H., GIERE O., STETTER K., ALFREDSSON G.A., KRISTJANSSON J.K., STOFFERS P.

& J. SVAVARSSON (1989): Hydrothermal vent communities at the shal-

low subpolar Mid-Atlantic Ridge. — Mar. Biol. 102: 425-429.

FUSTEC A., DESBRUYÈRES D. & S.K. JUNIPER (1987): Deep-sea hydrothermal

vent communities at 13°N on the East Pacific Rise: Microdistribution

and temporal variations. — Biol. Oceanogr. 4: 121-164.

GAL’CHENKO V.F., PIMENOV N.V., LEIN A.Y., GALKIN S.V., MOSKALEV L.I. & M.V.

IVANOV (1989): Autotrophic CO2-assimilation in tissues of shrimp

Rimicaris exoculata from a hydrothermal area on the Mid-Atlantic

Ridge. — Dokl. Akad. Nauk 308: 1478-1481.

GAL’CHENKO V.F., PIMENOV N.V., LEIN A.Y., GALKIN S.V., MILLER Y.M. & M.V.

IVANOV (1992): Mixotrophic type of feeding of Olgaconcha tufari

BECK (Gastropoda: Prosobranchia) from the active hydrothermal

field of the Manus Basin (Bismarck Sea). — Dokl. Akad. Nauk 323:

776-781.

GEBRUK A.V., PIMENOV N.V. & A.S. SAVVICHEV (1992): Bacterial “farm“ on the

mouthparts of shrimp. — Priroda 6: 37-39.

GEBRUK A.V., KUZNETSOV A.P., NAMSARAEV B.B. & Y.M. MILLER (1993): Role of

bacterial component in feeding of deep-water benthic fauna in

Froliha Bay, Lake Baikal. — Izvestija Acad. Nauk Ser. Biol. 6: 903-908.

GEBRUK A.V., CHEVALDONNÉ P., SHANK T., LUTZ R.A. & R.C. VRIJENHOEK (2000):

Deep-sea hydrothermal vent communities of the Logatchev area

(14°45’N, Mid-Atlantic Ridge): Diverse biotopes and high biomass.

— J. Mar. Biol. Ass. U.K. 80: 383-394.

GEISTDOERFER P., AUZENDE J.-M., BATIZA R., BIDEAU D., CORMIER M.-H., FOUQUET

Y., LAGABRIELLE Y., SINTON J. & P. SPADEA (1995): Hydrothermalisme et

communautés animales associés sur la dorsale du Pacifique oriental

entre 17°S and 19°S (Campagne NAUDUR, Décembre 1993). — C. R.

Acad. Sci. 320: 47-54.

GOFFREDI S.K., CHILDRESS J.J., DESAULNIERS N.T. & F.J. LALLIER (1997): Sulfide ac-

quisition by the vent worm Riftia pachyptila appears to be via up-

take of HS^, rather than H2S. — J. Exp. Biol. 200: 2609-2616.

GOFFREDI S.K., WARÉN A., ORPHAN V.J., VAN DOVER C.L. & R.C. VRIJENHOEK

(2004): Novel forms of structural integration between microbes. —

Appl. Environm. Microbiol. 70: 3082-3090.

GOVENAR B., FREEMAN M., BERGQUIST D.C., JOHNSON G.A. & C.R. FISHER (2004):

Composition of a one-year-old Riftia pachyptila community follow-

ing a clearance experiment: Insight to succession patterns at deep-

sea hydrothermal vents. — Biol. Bull. 207: 177-182.

GRASSLE J.F., BERG C.J., CHILDRESS J.J., GRASSLE J.P., HESSLER R.R., JANNASCH H.,

KARL D.M., LUTZ R.A., MICKEL T.J., RHOADS D.C., SANDERS H.L., SMITH K.L.,

SOMERO G.N., TURNER R.D., TUTTLE J.H., WALSH P.J. & A.J. WILLIAMS

(1979): Galapagos ’79: Initial findings of a deep-sea biological

quest. — Oceanus 22: 2-10.

GUINOT D. & L.A. HURTADO (2003): Two new species of hydrothermal vent

crabs of the genus Bythograea from the southern East Pacific Rise

and from the Galapagos Rift (Crustacea, Decapoda, Brachyura,

Bythograeidae). — C. R. Acad. Sci. 326: 423-439.

HASHIMOTO J., JOLLIVET D. & the KAIYO 88 Shipboard Party (1989): The hy-

drothermal vent communities in the North Fiji Basin: Results of

Japan-France cooperative research on board KAIYO 88. — La Mer

27: 62-71.

HASHIMOTO J., OHTA S., TANAKA T., HOTTA H., MATSUZAWA S. & H. SAKAI (1989):

Deep-sea communities dominated by the giant clam, Calyptogena

soyoae, along the slope foot of Hatsushima Island, Sagami Bay, cen-

tral Japan. — Palaeogeogr. Palaeoclimatol. Palaeoecol. 71: 179-192.

HASHIMOTO J., OHTA S., FIALA-MEDIONI 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., S TSUCHIDA., ISHIBASHI J., HENRY 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): Hydrothermal vent communities in the Manus Basin, Papua

New Guinea: Results of the BIOACCESS cruise ’96 and ’98. — Inter-

Ridge News 8: 12-18.

HASHIMOTO J., OHTA S., GAMO T., CHIBA H., YAMAGUCHI T., TSUCHIDA S., OKU-

DAIRA T., WATABE H., YAMANAKA T. & M. KITAZAWA (2001): First hy-

drothermal vent communities from the Indian Ocean discovered. —

Zool. Sci. 18: 717-721.

HAYMON R.M. & R.A. KOSKI (1985): Evidence of an ancient hydrothermal

vent community: Fossil worm tubes in Cretaceous sulfide deposit of

the Samail Ophiolite, Oman. — Proc. Biol. Soc. Wash. 6: 57-65.

HAYMON R.M., KOSKI R.A. & C. SINCLAIR (1984): Fossils of hydrothermal vent

worms from Cretaceous sulfide ores of the Samail Ophiolite, Oman.

— Science 223: 1407-1409.

HAYMON R.M., FORNARI D.J., VON DAMM K.L., LILLEY M.D., PERFIT M.R., ED-

MOND J.M, SHANKS W.C. III, LUTZ R.A., GREBMEIER J.M., CARBOTTE S.,

WRIGHT C.D., MCLAUGHLIN E., SMITH M., BEEDLE N. & E. OLSON (1993):

Volcanic eruption of the mid-ocean ridge along the East Pacific Rise

crest at 9°45-52’N: Direct submersible observations of seafloor phe-

nomena associated with an eruption event in April 1991. — Earth

Planetary Sci. Lett. 119: 85-101.

20


HECKER B. (1985): Fauna from a cold sulfur-seep in the Gulf of Mexico:Comparison with hydrothermal vent communities and evolutionaryimplications. — Proc. Biol. Soc. Wash. 6: 465-473.

HERZIG P., HANNINGTON M., MCINNES B., STOFFERS P., VILLINGER H.W., SEIFERT R.,BINNS R. & T. LIEBE (1994): Submarine volcanism and hydrothermalventing studied in Papua, New Guinea. — Eos, Trans. Amer. Geo-phys. Union 75: 513-, 515-516.

HESSLER R.R. & P.F. LONSDALE (1991): Biogeography of Mariana Trough hy-drothermal vent communities. — Deep-Sea Res. A 38: 185-199.

HESSLER R.R. & W.M. SMITHEY (1983): The distribution and communitystructure of megafauna at the Galapagos Rift hydrothermal vents.— In: RONA P.A., BOSTRÖM K., LAUBIER L. & K.L. Jr. SMITH (Eds.): Hy-drothermal Processes at Seafloor Spreading Centers, Plenum Press,NY: 735-770.

HESSLER R.R., SMITHEY W.M. Jr. & C.H. KELLER (1985): Spatial and temporalvariation of giant clams, tube worms and mussels at deep-sea hy-drothermal vents. — Bull. Biol. Soc. Wash. 6: 411-428.

HESSLER R.R., LONSDALE P. & J. HAWKINS (1988): Patterns on the ocean floor.— New Scientist 1605: 47-51.

HURTADO L.A., LUTZ R.A. & R.C. VRIJENHOEK (2004): Distinct patterns of ge-

netic differentiation among annelids of eastern Pacific hydrother-

mal vents. — Mol. Ecol. 13: 2603-15

JENKINS C.D., WARD M., TURNIPSEED M., OSTERBERG J. & C.L. VAN DOVER (2002):

The digestive system of the hydrothermal vent polychaete Gala-

pagomystides aristata (Phyllodocidae): Evidence for hematophagy?

— Invertebr. Biol. 121: 243-254.

JONES, M.L. (1981): Riftia pachyptila JONES: Observations on the vesti-

mentiferan worm from the Galapagos Rift. — Science 213: 333-336.

JONES M.L. & C.F. BRIGHT (1985): Appendix 1. Bibliography of hydrother-

mal vents and related areas. Their biotas, ecological parameters

and ancillary data. — Proc. Biol. Soc. Wash. 6: 495-538.

JUNIPER S.K., TUNNICLIFFE V. & D. DESBRUYÈRES (1990): Regional-scale features

of Northeast Pacific, East Pacific Rise, and Gulf of Aden vent com-

munities. — In: MCMURRAY G.R. (Ed.): Gorda Ridge: A Seafloor

Spreading Center in the United States Exclusive Economic Zone.

Springer, NY: 265-278.

JUNIPER S.K., TUNNICLIFFE V. & E.C. SOUTHWARD (1992): Hydrothermal vents in

turbidite sediments on a Northeast Pacific spreading centre: Or-

ganisms and substratum at an ocean drilling site. — Canad.J. Zool.

70: 1792-1809.

JUNIPER S.K., BIRD D.F., SUMMIT M., PONG VONG M. & E.T. BAKER (1998): Bac-

terial and viral abundances in hydrothermal event plumes over

northern Gorda Ridge. — Deep-Sea Res. II 45: 2739-2749.

KARL D.M., BRITTAIN A.M. & B.D. TILBROOK (1989): Hydrothermal and mi-

crobial processes at Loihi Seamount, a mid-plate hot-spot volcano.

— Deep-Sea Res. A 36: 1655-1673.

KELLEY D.S., KARSON J.A., BLACKMAN D.K., FRUEH-GREEN G.L., BUTTERFIELD D.A.,

LILLEY M.D., OLSON E.J., SCHRENK M.O., ROE K.K., LEBON G.T. & P. RIV-

IZZIGNO (2001): An off-axis hydrothermal vent field near the Mid-At-

lantic Ridge at 30°N. — Nature 412: 145-149.

KLEINSCHMIDT M. & R. TSCHAUDER (1985): Shallow-water hydrothermal vent

systems off the Palos Verdes Peninsula, Los Angeles County, Califor-

nia. — Proc. Biol. Soc. Wash. 6: 485-488.

KOJIMA S. (2002): Deep-sea chemoautosynthesis-based communities in

the Northwestern Pacific. — J. Oceanogr. 58: 343-363.

KOJIMA S., SEGAWA R., FUJIWARA Y., HASHIMOTO J. & S. OHTA (2000): Genetic

differentiation of populations of a hydrothermal vent-endemic gas-

tropod, Ifremeria nautilei, between the North Fiji Basin and the

Manus Basin revealed by nucleotide sequences of mitochondrial

DNA. — Zool. Sci. 17: 1167-1174.

KOJIMA S., SEGAWA R., FUJIWARA Y., FUJIKURA K., OHTA S. & J. HASHIMOTO

(2001): Phylogeny of hydrothermal-vent-endemic gastropods

Alviniconcha spp. from the western Pacific revealed by mitochon-

drial DNA sequences. — Biol. Bull. Mar. Biol.Lab. Woods Hole 200:

298-304.

KOJIMA S., FUJIKURA K., OKUTANI T. & J. HASHIMOTO (2003): Phylogenetic re-

lationship of Alviniconcha gastropods from the Indian Ocean to

those from the Pacific Ocean (Mollusca: Provannidae) revealed by

nucleotide sequences of Mitochondrial DNA. — Venus 63: 65-68.

KUZNETSOV A.P., MASLENNIKOV V.V. & V.V. ZAIKOV (1993): Hydrothermal fau-

na of the Silurian paleoocean in South Ural. — Izvestiya Akademii

Nauk, Ser. Biol. 4: 525-534.

LANGMUIR C., HUMPHRIS S., FORNARI D., VAN DOVER C., VON DAMM K., TIVEY

M.K., COLODNER D., CHARLOU J.-L., DESONIE D., WILSON C., FOUQUET Y.,

KLINKHAMMER G. & H. BOUGAULT (1997): Hydrothermal vents near a

mantle hot spot: The Lucky Strike vent field at 37°N on the Mid-At-

lantic Ridge. — Earth Planetary Sci. Lett. 148: 69-91.

LAUBIER L. & D. DESBRUYÈRES (1985): Oases at the bottom of the ocean. —

Endeavour 9: 67-76.

LAUBIER L., OHTA S. & M. SIBUET (1986): Decouverte de communautes ani-

mals profondes durant la campagne franco-japonaise KAIKO de

plongees dans les fosses de subduction autour du Japon. — C. R.

Acad. Sci. (Ser. 3) 303: 25-29.

LE PENNEC M. & A. HILY (1984): Anatomie, structure et ultra structure de

la branchie d’un Mytilidae des sites hydrothermaux du Pacifique

Oriental. — Oceanol. Acta 7: 517-523.

LE BRIS N., GOVENAR B., LE GALL C. & C.R. FISHER (2006): Variability of physi-

co-chemical conditions in 9°50’N EPR diffuse flow vent habitats. —

Mar. Chem. 98: 167-182.

LEVESQUE C., JUNIPER S.K. & J. MARCUS (2003): Food resource partitioning

and competition among alvinellid polychaetes of Juan de Fuca

Ridge hydrothermal vents. — Mar. Ecol. Progr. Ser. 246: 173-182.

LITTLE C.T.S. & R.C. VRIJENHOEK (2003): Are hydrothermal-vent animals liv-

ing fossils? — Trends Ecol. Evol. 18: 582-588.

LITTLE C.T.S., HERRINGTON R.J., MASLENNIKOV V.V., MORRIS N.J. & V.V. ZAYKOV

(1997): Silurian hydrothermal-vent community from the southern

Urals, Russia. — Nature 385: 146-148.

LISTER C.R.B. (1972): On the thermal balance of a mid-ocean ridge. —

Geophys. J. Roy. Astronom. Soc. 26: 515-535.

LONSDALE P. (1977): Clustering of suspension-feeding macrobenthos near

abyssal hydrothermal vents at oceanic spreading centers. — Deep-

Sea Res. 24: 857-863.

LUTHER G.W. III, ROZAN T.F., TAILLEFERT M., NUZZIO D.B., DI MEO C., SHANK T.M.,

LUTZ R.A. & S.C. CARY (2001): Chemical speciation drives hydrother-

mal vent ecology. — Nature 410: 813-816.

LUTZ R.A., SHANK T.M., FORNARI D.J., HAYMON R.M., LILLEY M.D., VON DAMM

K.L. & D. DESBRUYÈRES (1994): Rapid growth at deep-sea vents. — Na-

ture 371: 663-664.

MARSH A.G., MULLINEAUX L.S., YOUNG C.M. & D.T. MANAHAN (2001): Larval

dispersal potential of the tubeworm Riftia pachyptila at deep-sea

hydrothermal vents. — Nature 411: 77-80

MEVEL 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): La

ride du Snake Pit (dorsale médio-Atlantique, 23°22’N); résultats

préliminaires de la campagne Hydrosnake. — C. R. Acad. Sci. (Ser. 2)

308: 545-552.

MICHELI F., PETERSON C.H., JOHNSON G.A., MULLINEAUX L.S., MILLS S.W., SANCHO

G., FISHER C.R. & H.S. LENIHAN (2002): Predation structures communi-

ties at deep-sea hydrothermal vents. — Ecol. Monogr. 72: 365-382.

21


MULLINEAUX L.S., FISHER C.R., PETERSON C.H. & S.W. SCHAEFFER (2000): Tube-worm succession at hydrothermal vents: use of biogenic cues to re-duce habitat selection error? — Oecologia 123: 275-284.

MULLINEAUX L.S., PETERSON C.H., MICHELI F. & S.W. MILLS (2003): Successionalmechanism varies along a gradient in hydrothermal fluid flux atdeep-sea vents. — Ecol. Monogr. 73: 523-542.

MURTON B.J., KLINKHAMMER G., BECKER K., BRIAIS A., EDGE D., HAYWARD N., MIL-LARD N., MITCHELL I., ROUSE I., RUDNICKI M., SAYANAGI K., SLOAN H. & L.PARSON (1994): Direct evidence for the distribution and occurrenceof hydrothermal activity between 27°N-30°N on the Mid-AtlanticRidge. — Earth Planetary Sci. Lett. 125: 119-128.

NAAR D.F., HEKINIAN R., SEGONZAC M., FRANCHETEAU J., & the Pito Dive Team(2004): Vigorous venting and biology at Pito Seamount, Easter Mi-croplate. — In: GERMAN C.R., LIN J., L.M. PARSON (Eds.): Mid-OceanRidges: Hydrothermal Interactions Between the Lithosphere andOceans. Amer. Geophys. Union, Washington, D.C.: 305-318.

NELSON D.C., HAYMON R.M., LILLEY M. & R. LUTZ (1991): Rapid growth of un-usual hydrothermal bacteria observed at new vents during the AD-VENTURE dive program on the EPR crest at 9°45’-52’N. — Eos, Trans.Amer. Geophys. Union 72: 481.

ODP Leg 106 Scientific Party (1986): Drilling the Snake Pit hydrothermalsulfide deposit on the Mid-Atlantic Ridge, 23°22’N. — Geology 14:3739-3762.

OHTA S. & L. LAUBIER (1987): Deep biological communities in the subduc-tion zone of Japan observed by bottom photographs taken duringNautile dives in the KAIKO project. — Earth Planetary Sci. Lett. 83:329-342.

OKUTANI T. & K. EGAWA (1985): The first underwater observation on livinghabitat and thanatocoenoses of Calyptogena soyoae in bathyaldepth of Sagami Bay. — Venus 49: 83-91.

OKUTANI T., HASHIMOTO J. & T. SASAKI (2004): New gastropod taxa from hy-drothermal vent (Kairei Field) in the central Indian Ocean. — Venus63: 1-12.

OUDIN E. & G. CONSTANTINOU (1984): Black smoker chimney fragments inCyprus sulphide deposits. — Nature 308: 349-353.

PAULL C.K., HECKER B., COMMEAU R., FREEMAN-LYNDE R.P., NEUMANN C., CORSO

W.P., GOLUBIC S., HOOK J.E., SIKES E. & J. CURRAY (1984): Biological com-munities at the Florida Escarpment resemble hydrothermal venttaxa. — Science 226: 965-967.

POWELL M.A. & G.N. SOMERO (1983): Blood components prevent sulfidepoisoning of respiration of the hydrothermal vent tube worm Rif-tia pachyptila. — Science 219: 297-299.

POWELL M.A. & G.N. SOMERO (1986): Adaptations to sulfide by hydrother-mal vent animals: Sites and mechanisms of detoxification and me-tabolism. — Biol. Bull. Mar. Biol.Lab. Woods Hole 171: 274-290.

PRADILLON F., SHILLITO B., YOUNG C.M. & F. GAILL (2001): Developmental ar-rest in vent worm embryos. — Nature 413: 698-699.

Rise Project Group (1980): East Pacific Rise: Hot springs and geophysicalexperiments. — Science 207: 1421-1433.

RONA P.A., KLINKHAMMER G., NELSON T.A., TREFRY J.H. & H. ELDERFIELD (1986):Black smokers, massive sulfides and vent biota at the Mid-AtlanticRidge. — Nature 321: 33-37.

SAGALEVITCH A.M., TOROKHOV P.V., MATVEENKOV V.V., GALKIN S.V. & L.I.MOSKALEV (1992): Hydrothermal manifestations of the submarine Pi-ip’s Volcano. — Izvestiya RAN, Ser. Geol. 9: 104-114.

SARRAZIN J. & S.K. JUNIPER (1999): Biological characteristics of a hydrother-mal edifice mosaic community. — Mar. Ecol. Progr. Ser. 185: 1-19.

SARRAZIN J., ROBIGOU V., JUNIPER S.K. & J.R. DELANEY (1997): Biological andgeological dynamics over four years on a high-temperature sulfidestructure at the Juan de Fuca Ridge hydrothermal observatory. —Mar. Ecol. Progr. Ser. 153: 5–24.

SEGONZAC M., DE SAINT LAURENT M. & B. CASANOVA (1993): L’énigme du com-

portement trophique des crevettes Alvinocarididae des sites hy-

drothermaux de la dorsale médio-atlantique. — Cah. Biol. Mar. 34:

535-571.

SELIVERSTOV N.I., AVDEIKO G.P., IVANENKO A.N., SHKIRA V.A. & S.A. HUBUNAIA

(1986): New submarine volcano in the western part of the Aleutian

Island Arc. — Volcanol. Seismol. 4: 3-16.

SHANK T.M., FORNARI D.J., VON DAMM K.L., LILLEY M.D., HAYMON R.M. & R.A.

LUTZ (1998): Temporal and spatial patterns of biological community

development at nascent deep-sea hydrothermal vents (9°50’N, East

Pacific Rise). — Deep-Sea Res. II 45: 465-515.

SHILLITO B., JOLLIVET D., SARRADIN P.-M., RODIER P., LALLIER F., DESBRUYÈRES D. &

F. GAILL (2001): Temperature resistance of Hesiolyra bergi, a poly-

chaetous annelid living on deep-sea vent smoker walls. — Mar. Ecol.

Progr. Ser. 216: 141-149.

SIBUET M. & K. OLU (1998): Biogeography, biodiversity and fluid depend-

ence of deep-sea cold-seep communities at active and passive mar-

gins. — Deep-Sea Res. II 45: 517-567.

SIRENKO B.I., PETRYASHOV V.V., RACHOR E. & K. HINZ (1995): Bottom bio-

cenoses of the Laptev Sea and adjacent areas. — Ber. Polar-

forsch./Rep. Polar Res. 176: 211-221.

SMITH C.R. & A.R. BACO (2003): Ecology of whale falls at the deep-sea

floor. — Oceanogr. Mar. Biol. 41: 311–354.

SMITH C.R., KUKERT H., WHEATCROFT R.A., JUMARS P.A. & J.W. DEMING (1989):

Vent fauna on whale remains. — Nature 323: 251-253.

STEIN J.L., CARY S.C., HESSLER R.R., OHTA S., VETTER R.D., CHILDRESS J.J. & H. FEL-

BECK (1988): Chemoautotrophic symbiosis in a hydrothermal vent

gastropod. — Biol. Bull. Mar. Biol. Lab. Woods Hole 174: 373-378.

STEIN C.A., STEIN S. & A.M. PELAYO (1995): Heat flow and hydrothermal cir-

culation. — In: HUMPHRIS S.E., ZIERENBERG R.A., MULLINEAUX L.S. & R.E.

THOMSON (Eds.): Seafloor Hydrothermal Systems: Physical, Chemical,

and Geochemical Interactions. Geophys. Monogr. 91, Amer. Geo-

phys. Union, Washington, DC: 425-445.

SUESS E., CARSON B., RITGER S.D., MOORE J.C., JONES M.L., KULM L.D. & G.R.

COCHRANE (1985): Biological communities at vent sites along the sub-

duction zone off Oregon. — Proc. Biol. Soc. Wash. 6: 475-484.

TALWANI M., WINDISCH C.C. & M.L. LANGSETH (1971): Reykjanes ridge crest:

A detailed geophysical study. — J. Geophys. Res. 76: 473-517.

TANAKA T., HOTA H., SAKAI H., ISHIBASHI J., OOMORI T., IZAWA E. & N. ODA

(1990): Occurrence and distribution of the hydrothermal deposits in

the Izena Hole, central Okinawa Trough. — JAMSTEC Deep Sea Res.

6: 11-26.

TARASOV V.G. (in press): Effects of shallow-water hydrothermal venting

on biological communities of coastal marine ecosystems of the

western Pacific. — Adv. Mar. Biol.

TARASOV V.G., PROPP M.V., PROPP L.N., ZHIRMUNSKY A.V., NAMSARAEV B.B., GOR-

LENKO V.M. & D.A. STARYNIN (1990): Shallow-water gasohydrothermal

vents of Ushishir Volcano and the ecosystem of Kraternaya Bight

(the Kurile Islands). — P.S.Z.N. Mar. Ecol. 11: 1-23.

TIERCELIN J.-J., PFLUMIO C., CASTREC M., BOULEGUE J., GENTE P., ROLET J., COUSSE-

MENT C., STETTER K.O., HUBER R., BUKU S. & W. MIFUNDI (1993): Hy-

drothermal vents in Lake Tanganyika, East African Rift system. —

Geology 21: 499-502.

TSURAMI M. & V. TUNNICLIFFE (2001): Characteristics of a hydrothermal vent

assemblage on a volcanically active segment of Juan de Fuca Ridge,

northeast Pacific. — Canad. J. Fisheries Aquat. Sci. 58: 530-542.

TUNNICLIFFE V. (1988): Biogeography and evolution of hydrothermal-vent

fauna in the eastern Pacific Ocean. — Proc. Roy. Soc. Lond. B 233:

347-366.

22


TUNNICLIFFE V., JUNIPER S.K. & M.E. DE BURGH (1985): The hydrothermal vent

community on Axial Seamount, Juan de Fuca Ridge. — Proc. Biol.

Soc. Wash. 6: 453-464.

TUNNICLIFFE V., JUNIPER S.K. & M. SIBUET (2003a): Reducing environments of

the deep-sea floor. — In: P. TYLER (Ed.): Ecosystems of the Deep

Oceans. Elsevier, Amsterdam: 81-110.

TUNNICLIFFE V., M BOTROS., DE BURUGH M.E., DONET A., JONSON H.P., JUNIPER S.K.

& R.E. MCDUFF (1986): Hydrothermal vents of Explorer Ridge, north-

east Pacific. — Deep-Sea Res. A 33: 401-412.

TUNNICLIFFE V., EMBLEY R.W., HOLDEN J.F., BUTTERFIELD D.A., MASSOTH G.J. &

S.K. JUNIPER (1997): Biological colonization of new hydrothermal

vents following an eruption on Juan de Fuca Ridge. — Deep-Sea

Res. I 44: 1627-1644.

TUNNICLIFFE V., BAROSS J.A., GEBRUK A.V., GIERE O., HOLLAND M.E., KOSCHINSKY

A., REYSENBACH A.-L. & T.M. SHANK (2003b): Group Report: What are

the interactions between biotic processes at vents and physical,

chemical and geological conditions? — In: HALBACK P.E., TUNNICLIFFE V.

& J.R. HEIN (Eds.): Energy and Mass Transfer in Marine Hydrothermal

Systems. Dahlem Press, Berlin: 251-270.

TURNIPSEED M., JENKINS C.D. & C.L. VAN DOVER (2004): Community structure

in Florida Escarpment seep and Snake Pit (Mid-Atlantic Ridge) vent

mussel beds. — Mar. Biol. 145: 121-132.

TYLER P.A., GERMAN C.R., RAMIREZ-LLODRA E. & C.L. VAN DOVER (2003): Un-

derstanding the biogeography of chemosynthetic ecosystems. —

Oceanologica Acta 25: 227-241.

VAN DOVER C.L. (2003): Variation in community structure within hy-

drothermal vent mussel beds of the East Pacific Rise. — Mar. Ecol.

Progr. Ser. 253: 55–66.

VAN DOVER C.L., GRASSLE J.F. & M. BOUDRIAS (1990): Hydrothermal vent fau-

na of Escanaba Trough (Gorda Ridge). — In: MCMURRAY G.R. (Ed.):

Gorda Ridge: A Seafloor Spreading Center in the United States Ex-

clusive Economic Zone. Springer-Verlag, NY: 253-264.

VAN DOVER C.L., FRY B., GRASSLE J.F., HUMPHRIS S. & P.A. RONA (1988): Feed-

ing biology of the shrimp Rimicaris exoculata at hydrothermal vents

on the Mid-Atlantic Ridge. — Mar. Biol. 98: 209-216.

VAN DOVER C.L., SZUTS E.Z., CHAMBERLAIN S.C. & J.R. CANN (1989): A novel eye

in ‘eyeless’ shrimp from hydrothermal vents of the Mid-Atlantic

Ridge. — Nature 337: 458-460.

VAN DOVER C.L., DESBRUYÈRES D., SEGONZAC M., COMTET T., SALDANHA L., FIALA-

MEDIONI A. & C. LANGMUIR (1996): Biology of the Lucky Strike hy-

drothermal field. — Deep-Sea Res. I 43: 1509-1529.

VAN DOVER C.L., HUMPHRIS S.E., FORNARI D., CAVANAUGH C.M., COLLIER R., GOF-

FREDI S.K., HASHIMOTO J., LILLEY M.D., REYSENBACH A.L., T SHANK.M., VON

DAMM K.L., BANTA A., GALLANT R.M. & R.C. VRIJENHOEK (2001): Bio-

geography and ecological setting of Indian Ocean hydrothermal

vents. — Science 294: 818-823.

VAN DOVER C.L., GERMAN C.R., SPEER K.G., PARSON L.M. & R.C. VRIJENHOEK

(2002): Evolution and biogeography of deep-sea vent and seep in-

vertebrates. — Science 295: 1253-1257.

VINE F.J. & D.H. MATTHEWS (1963): Magnetic anomalies over oceanic

ridges. — Nature 199: 947-949.

VINOGRADOVA N. (1979): The geographical distribution of the abyssal and

hadal (ultra-abyssal) fauna in relation to vertical zonation of the

ocean. — Sarsia 64: 41-50.

VRIJENHOEK R.C. (1997): Gene flow and genetic diversity in naturally frag-

mented metapopulations of deep-sea hydrothermal vent animals.

— J. Heredity 88: 285-293.

WEISS R.F., LONSDALE P.F., LUPTON J.E., BAINBRIDGE A.E. & H. CRAIG (1977): Hy-drothermal plumes in the Galapagos Rift. — Nature 267: 600-603.

WARÉN A. & P. BOUCHET (2001): Gastropoda and Monoplacophora fromhydrothermal vents and seeps; new taxa and records. — The Veliger44: 116-231.

WARÉN A., BENGTSON S., GOFFREDI S.K. & C.L. VAN DOVER (2003): A hot-ventgastropod with iron-sulphide dermal sclerites. — Science 302: 1007.

WHITE S.N., CHAVE A.D., REYNOLDS G.T. & C.L. VAN DOVER (2002): Ambientlight emission from hydrothermal vents on the Mid-Atlantic Ridge.— Geophys. Res. Lett. 29: 10.1029/2002GL014977.

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.

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24

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


25

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


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


28

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.


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.


30

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.


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.


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].


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.


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.


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


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


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.


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


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:



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).



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:



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).



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.


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:




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.


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.



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).


J. VACELET & M. SEGONZAC Denisia 18 (2006): 42–43


Reference:


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


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.


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.


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.


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:



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).



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.


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.


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


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


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


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.


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


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.


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.


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:



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


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.



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.


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


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.


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


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.


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).



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).


Reference:




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.


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).



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.


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


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.


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.



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.


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).



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.


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


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


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


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.


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.


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.


Reference:



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.



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.


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.



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.


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


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).


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.


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.



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.


Chondrophellia cf. coronata (VERRILL, 1883)

Reference:



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.


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.


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.


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


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.


References:

SCHELTEMA A.H. (2000) Argonauta XIV(2): 15-25.


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).



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).


L. SALVINI-PLAWEN Denisia 18 (2006): 78–79


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.

79


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


Leptochiton tenuidontus SAITO & OKUTANI, 1990

Reference:

SAITO H. & T. OKUTANI (1990) Venus 49(3): 165-179.


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.


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.


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.


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]


1: N. gordensis, exterior, interior and lateral views; by R. von Cosel & A. Le Goff.


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).


Paralepetopsis ferrugivora WARÉNWW & BOUCHET, 2001 “rust-eating limpets”


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].


1: Exterior, interior and lateral views; by R. von Cosel, A. Le Goff, P. Briand & A. Warén.



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].


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.


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].


1 top:G. emarginatus;middle: G. patuluswith incrustations;bottom: G. spiralis.All with exterior,interior and lateralview; by R. von Cosel& A. Le Goff.


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


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


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.


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


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


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.

89


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.

90


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”


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.


Species DistributionP.PP marianae MCLEAN, 1989 Mariana Back-Arc BasinP. midatlantica MCLEAN, 1992 Mid-Atlantic Ridge: 38-15°N



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.

92


Pyropelta MCLEAN & HASZPRUNAR, 1987


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].


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


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.


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.


Fucaria WARÉNWW & BOUCHET, 1993

1: View ofdifferentsides ofseveralspecimens;by R. vonCosel.


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].



Leptogyra inflata WARÉNWW & BOUCHET, 1993


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).



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].


1 top: P. thorvaldssoni, apertural,abapertural and lateral view; by R. von Cosel & A. Le Goff;

right: P. valvatoides; by P. Briand.


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


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.


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].


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


Temnocinclis euripes MCLEAN, 1989 “slit-limpet”


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].


1: Different views; by R. von Cosel & P.PP Briand.

2: Specimen, dorsal view; by R. von Cosel & P. Briand.



Temnozaga parilis MCLEAN, 1989


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.


1: Habitus; by A. Warén & P.PP Briand.



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].


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.


Helicrenion reticulatum WARÉNWW & BOUCHET, 1993


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:



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


Vetulonia phalcata WARÉNWW & BOUCHET, 1993


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:



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


Cyathermia naticoides WARÉNWW & BOUCHET, 1989

1: Apertural, abapertural and lateral views; by R. von Cosel, A. Le Goff & P. Briand.


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.


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].



Lacunoides WARÉNWW & BOUCHET, 1989


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.


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).


Species DistributionL. exquisitus Galapagos Spreading Center

L. vitreus Juan de Fuca Ridge: Axial Seamount – Ashes vent field


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].


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.


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


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.


Neomphalus fretterae MCLEAN, 1981


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].


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.



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].


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.


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.


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].


1 top left, right and bottom: Dorsal, ventral and apertural view of P.PP planispira; by R. von Cosel & A. Le Goff.


Species DistributionP. depressa WARÉN & BOUCHET, 1993 Lau Back-Arc Basin: Hine HinaP.PP planispira WARÉN & BOUCHET, 1989 East Pacific Rise: 21°N-17°S


Symmetromphalus MCLEAN, 1990

References:

BECK L. (1992) Ann. Nat.hist. Mus. Wien B 93: 243-257.MCLEAN J. (1990) Nautilus 104: 77-86.


1: S. regularis; by R. von Cosel & P.PP Briand.



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


Ctenopelta porifera WARÉNWW & BOUCHET, 1993


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.


References:

WARÉNW A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [33-35].WARÉNW A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [170].


1: Several specimens; by R. von Cosel.

Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae


Depressigyra globulus WARÉNWW & BOUCHET, 1989


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.


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].


1: Several specimens; by R. von Cosel © MNHN.



Echinopelta fistulosa MCLEAN, 1989 “tapersnout limpet”


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].


1: Two specimens in various views; by R. von Cosel.TT



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].


1: All specimens H. hirta; D. Brabant © MNHN and P. Briand © Ifremer.


Species DistributionH. hirta MCLEAN, 1989 East Pacific Rise: 13-21°NH. tufari BECK, 2002 East Pacific Rise: 21°S


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.


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.


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


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:




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.


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


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.



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.

117


Peltospira MCLEAN, 1989 “tapersnout limpets”


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].


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.


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


Rhynchopelta concentrica MCLEAN, 1989 “tapersnout limpet”


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.


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].


1 top: Exterior, lateral and interior view; by R. von Cosel & A. Le Goff; bottom: Exterior, lateral and interi-or view; by P. Briand.




“scaly foot gastropod”


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.


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.



Olgasolaris BECK, 1992


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.


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


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:


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.



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].

123


Eosipho THIELE, 1929


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


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.

125


Speculator cariosus WARÉNWW & BOUCHET, 2001


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.


1: Apertural view; after WARÉNWW &BOUCHET (2001).

2: Enlargement of aperture; after WARÉNWW & BOUCHET (2001).

Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Cerithiopsidae


Laeviphitus desbruyeresi WARÉNWW & BOUCHET, 2001


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].


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


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.


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].


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.


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


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.




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.

131


Provanna DALL, 1918

1 top left: P. laevis; top right: P. variabilis; bottom left: P. ios; bottom right: P. buccinoides; by P. Briand.


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.



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


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


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


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.


Mollusca, Gastropoda, Caenogastropoda, Vitrinellidae


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:




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].


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.


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].


1: Two specimens: left dorsal view, right ventral view; by A. Warén.

Mollusca, Gastropoda, Heterobranchia, Xylodisculidae


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


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.


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.


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.


1: Bathymodiolus azoricus from Rainbow, Mid-Atlantic Ridge, Atos cruise © Ifremer.


Calyptogena edisonensis OKUTANI, KOJIMA & KIM, 2003

Reference:

OKUTANI T.K., KOJIMA S. & D. KIM (2004) Venus 63: 29-32.


1: Several specimens; by R. von Cosel.

Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae


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.


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.


1: Habitus; by R. von Cosel.



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).


Calyptogena gigas DALL, 1896


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].


2: In situ; by courtesy of K. Juniper.




Calyptogena magnifica BOSS & TURNER, 1980


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.


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:


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.


1: Habitus of a specimen; by R. von Cosel.



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).


Calyptogena okutanii KOJIMA & OHTA, 1997


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].





Calyptogena solidissima OKUTANI, HASHIMOTO & FUJIKURA, 1992


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.



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


Bathymodiolus aduloides HASHIMOTO & OKUTANI, 1994


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


Bathymodiolus azoricus COSEL & COMTET, 1998


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.


R. VON COSEL Denisia 18 (2006): 151–152


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


Bathymodiolus brevior COSEL, MÉTIVIER & HASHIMOTO, 1994

Reference:

COSEL R. VON, METIVIER B. & J. HASHIMOTO (1994) Veliger 37(4): 374-392 [375-380].


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.



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.



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.



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.


Reference:

COSEL R. VON, METIVIER B. & J. HASHIMOTO (1994) Veliger 37(4): 374-392 [380-386].



Bathymodiolus japonicus HASHIMOTO & OKUTANI, 1994


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.


2: In situ view of a mussel bed © JAMSTEC.


1: Paratype MNHN Paris, exterior of both valves, interiorof right valve; by R. von Cosel & P. Lozouet © MNHN.


Bathymodiolus marisindicus HASHIMOTO, 2001


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.


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.



Bathymodiolus platifrons HASHIMOTO & OKUTANI, 1994


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.


2: In situ view of a mussel bed © JAMSTEC.


1: Paratype MNHN Paris, exterior and interior of right valve; by R. von Cosel & P. Lozouet © MNHN.


Bathymodiolus puteoserpentis COSEL, MÉTIVIER & HASHIMOTO, 1994


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.


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.



Bathymodiolus septemdierum HASHIMOTO & OKUTANI, 1994


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.


2: In situ © JAMSTEC.


1: Paratype MNHN, interior and exterior of right valve;by R. von Cosel & P. Lozouet © MNHN.


Bathymodiolus thermophilus KENK & WILSON, 1985

1: In situ views; cruise Hope © Ifremer; top right, by P. Briand © Ifremer.


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).


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


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.


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


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.


Gigantidas horikoshii HASHIMOTO & YAMANE, 2005


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.


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


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.


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).


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.


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


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


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


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


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.



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



Nematoda

1: Thalassomonhystera sp. from East Pacific Rise: 9°N, Tica; by M. Bright.

173


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.


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:




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.


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.


Reference:



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.


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.


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.


Reference:




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.


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.


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.


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.


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.


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.


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.



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.



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


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.


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.



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.


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.



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.


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.


References:

BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.TUNNICLIFFE V. (1988) Proc. R. Soc. Lond. B 233: 347-366.



Ophryotrocha platykephale BLAKE, 1985

2: Prostomium, dorso-lateral view (SEM) © Ifremer.


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


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


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.


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.


Reference:

BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.

J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 192


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.


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


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.



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.


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


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.

197


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


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.


1A: Anterior end, dorsal view; B: Right parapodium from middle segment, anterior view; by courtesy of J. Blake.



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.


F. PLEIJEL Denisia 18 (2006): 200–201


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).

201


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.


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


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.

203


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.


1A: Anterior end, dorsal view with proboscis everted; B: Middle parapodium, posterior view; by courtesy of J. Blake.



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.


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.


1A: Anterior end, dorsal view, proboscis everted; B: Right parapodium from chaetiger 6, anterior view; by courtesy of J. Blake.



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


Mytilidiphila enseiensis MIURA & HASHIMOTO, 1993



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.


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.



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).




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.


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.



Shinkai longipedata MIURA & OHTA, 1991


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.


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.


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.



Shinkai semilonga MIURA & HASHIMOTO, 1996


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).


T. MIURA Denisia 18 (2006): 210–211


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.

211


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.


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


Reference:


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


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:



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.


Eulalia papillosa (BLAKE, 1985)

Reference:



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.


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.


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.



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.


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.


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.


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.



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.


2: In vivo, ventral view of a male specimen;by P. Briand © Ifremer.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 219–220


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.

220


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.




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


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:



1: Dorsal view; by P. Briand © Ifremer. 2: Ventral view of the posterior segments, female;by P. Briand © Ifremer.



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


Branchinotogluma segonzaci (MIURA & DESBRUYÈRES, 1995)


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.



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


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.


Reference:

MIURA T. & D. DESBRUYÈRES (1995) Proc. Biol. Soc. Wash. 108: 583-595.


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.



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.


1: In situ; by courtesy of S.K. Juniper.



Reference:


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


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:



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.



Branchipolynoe pettibonae MIURA & HASHIMOTO, 1991


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.


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.



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.


2: Tip of upper neurochaetae(SEM) © Ifremer.

1: Proboscis, frontal view (SEM) © Ifremer.



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


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.


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.



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.


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.



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.


Reference:

DESBRUYÈRES D. & S. HOURDEZ (2000) Cah. Biol. Mar. 41: 47-54.


6: Distal part of a neuropodial chaeta(SEM) © Ifremer.


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.


3: Specimen on a smoker wall; East Pacific Rise: 13°N, 2630 m. Cruise Phare Ifremer.

2: Prostomium, dorsal view Ifremer.




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.


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.



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.




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


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.


References:

PETTIBONE M.H. (1984) Proc. Biol. Soc. Wash. 97: 849-863.VAN DOVER C.L. (2002) Mar. Ecol. Progr. Ser. 230: 137-158.


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.


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.



2: Habitus, in vivo Ifremer.

5: Prostomium, dorsal view (SEM) Ifremer.

3: Proboscis, frontal view (SEM) Ifremer.

4: Elytron, posterior border(SEM) Ifremer



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.


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


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.


1: Habitus Ifremer.

2: Posterior part Ifremer.

3: Parapodia, dorsal view Ifremer.




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


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.


Reference:

MIURA T. (1994) Proc. Biol. Soc. Wash. 107: 532-543.


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).


2: Anterior end, dorsal viewelytra removed; from MIURA (1994).


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.


2: Proboscis, frontal view (preserved specimen) © Ifremer.



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


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.


T. MIURA & D. DESBRUYÈRES Denisia 18 (2006): 249–250


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.


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


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.


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.


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.


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


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


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


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.


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.


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.



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.


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.


2: Branchial regionin lateral view; byE. Southward.


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.


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.


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.


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.


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.



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.


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.


Ridgeia piscesae JONES, 1985

1: Population in situ (Endeavour segment); by courtesy of S.K. Juniper.



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.



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.




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


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.


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.


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.


Tevnia jerichonana JONES, 1985 ”Jericho worm“


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.


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.



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.


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


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:



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


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:



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.



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).


Size: Small, probably no more than 4 mm long, 0.6 mm wide,for 25 segments.


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:




Orbiniella hobsonae BLAKE & HILBIG, 1990

Size: Small, threadlike, up to 4.5 mm long, 0.2 mm wide, for 25chaetigers.


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.


1: Anterior end, dorsolateral view (SEM) © Ifremer. 2: Barbed acicular spines (SEM) © Ifremer.


References:

BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.TUNNICLIFFE V. (1988) Proc. R. Soc. Lond. B 233: 347-366.



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).


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.



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.



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.



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).


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.


Reference:

MACIOLEK N.J. (1981) Proc. Biol. Soc. Wash. 94: 826-837.



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).


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.



Lindaspio southwardorum BLAKE & MACIOLEK, 1992


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


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


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.


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.


J.A. BLAKE Denisia 18 (2006): 278–279


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:


279


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.


1: Habitus (preserved specimen) Ifremer. Box: Distalpart of notopodial hooded hook (SEM) Ifremer.

2: Anterior part (SEM) Ifremer.



Xandaros acanthodes MACIOLEK, 1981

Reference:



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).


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.



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.


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.


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.


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.



4: Specimen in vivo; by Dugornay © Ifremer.

284


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.


Reference:

DESBRUYÈRES D. & L. LAUBIER (1991) Ophelia Suppl. 5: 31-45.


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.


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.


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.



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.


2: Male buccal apparatus (SEM) Ifremer.

3: Cross cut of a tube wall, show-ing epibiotic bacteria on the innerpart (SEM) Ifremer.


1: Habitus;by V. Martin.

Box: Anterior part,dorsal view (SEM)

Ifremer.


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.


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.


1: Drawing of P. p. pandorae;by V. Martin Ifremer.


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.


1: Habitus; by V. Martin © Ifremer.


2: Anterior portion (SEM); by V. Martin © Ifremer.


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


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.


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.


1: Habitus; by V. Martin;box preserved specimen© Ifremer.

Reference:

DESBRUYÈRES D. & L. LAUBIER (1993) Proc. Biol. Soc. Wash. 106: 225-236.



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.


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.



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.


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.


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.


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.


3: Population in situ (Endeavour segment); by courtesy of S.K. Juniper.

1: Anterior part (preserved specimen) Ifremer.


2: Population in situ (Endeavour seg-ment); by courtesy of S.K. Juniper.


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.


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



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.


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



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


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


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.


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


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.


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).



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.


Reference:

CHILD C.A. & M. SEGONZAC (1996) Proc. Biol. Soc. Wash. 109(4): 664-676.


1: Holotype female. A: Trunk, dorsal view; B: Trunk, lateral view; C: Palp; D: Oviger; E: Third leg; from CHILD & SEGONZAC (1989).



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.


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].


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.


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.


1: Male. A: Trunk, dorsal view; B: Third leg, with enlargement of cement gland; C: Palp; D: Oviger; from GORDON (1944).


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.


Sericosura venticola CHILD, 1987

Reference:

CHILD C.A. (1987) Proc. Biol. Soc. Wash. 100(4): 896-899.


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).


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.


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.


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.


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.



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.



2: Adult maleendopod of secondantenna; by Kornicker.

1: Adult male carapace from right side; by Kornicker.

3: Adult male sixth limb; by Kornicker.


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.


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.


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.


Polycopetta pax KORNICKER & HARRISON-NELSON, 2005


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.


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).


Prionotoleberis styx KORNICKER, 1991

Reference:

KORNICKER L.S. (1991) Smithson. Contr. Zool. 516: 1–46.


2: Sixth limb; by Kornicker.

4: Mandible; by Kornicker.

Arthropoda, Crustacea, Ostracoda, Myodocopa, Myodocopida, Cylindroleberidae

1: Complete carapace showing central adductormuscles; by Kornicker.


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.


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.


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.


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.


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.


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


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


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


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


Uptionyx verenae CONROY-DALTON & HUYS, 2000

Reference:

CONROY-DALTON S. & R. HUYS (2000) Cah. Biol. Mar. 41(4): 343-397.


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.


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.



Jamstecia terazakii LEE & HUYS, 2000


1: Female habitus, lateral;from LEE & HUYS (2000).


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.


References:

LEE W. & R. HUYS (2000) Zool. J. Linn. Soc. 129: 1-71.LEE W. & R. HUYS (2001) Zool. J. Linn. Soc. 131: 249.



Scabrantenna yooi LEE & HUYS, 2000


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.


Reference:

LEE W. & R. HUYS (2000) Zool. J. Linn. Soc. 129: 1-71.



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.


Reference:

HUMES A.G. (1989) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 11, section A (4): 829-849.


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).


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.


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).


Hyphalion captans HUMES, 1987


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.


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.


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).


Barathricola rimensis HUMES, 1999


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.


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).


Heptnerina confusa IVANENKO & DEFAYE, 2004


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.


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).


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


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


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.



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).


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.


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.


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.



Chasmatopontius thescalus HUMES, 1990


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.



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).


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.



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.


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.


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.



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.


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.




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).


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).



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.



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.



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).


Rhogobius HUMES, 1987


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.


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).


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.



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).


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).


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.



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.


Stygiopontius HUMES, 1987


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).


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


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


Ecbathyrion prolixicauda HUMES, 1987


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.


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).


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.



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.


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


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).


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.


Arthropoda, Crustacea, Copepoda, Cyclopoida, Lubbockiidae

1: Male; A: Habitus, dorsal; B: Rostrum, lateral; C: Antenna; D: Mandible; E: Maxilla; F: Maxilliped; from HUMES (1987).


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.


1: Male; A: Habitus, dorsal; B: Oral cone, lateral;C: Antenna; from HUMES (1988).


Stenhelia gundulae WILLEN, 2003


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.


Arthropoda, Crustacea, Copepoda, Harpacticoida, Miraciidae

1A: Female habitus, dorsal; B: Maxilliped; C: Leg 5; D: Male habitus, dorsal; E: Leg 5; from WILLEN (2003).


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.


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).


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).


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).


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.



Grievella shanki FERRARI & MARKHASEVA, 2000

Reference:

FERRARI F.D. & E.L. MARKHASEVA (2000) Proc. Biol. Soc. Wash. 113 (4): 1079-1088.


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).


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.


Isaacsicalanus paucisetus FLEMINGER, 1983


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).


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.


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).


Smacigastes micheli IVANENKO & DEFAYE, 2004

Reference:

IVANENKO V.N. & D. DEFAYE (2004) Cah. Biol. Mar. 45: 255-268.


Arthropoda, Crustacea, Copepoda, Harpacticoida, Tegastidae

1A: Female habitus, lateral; B: Male habitus, lateral; from IVANENKO & DEFAYE (2004).


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).



Cholidya polypi FARRAN, 1914


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.


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).


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).


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.



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.


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


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


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.


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


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.



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.


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.



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).



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.


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.


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.



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.


2: Group of 16 paratypes; from BUCKERIDGE (2000).


1: Habitus; scale bar 1 cm© T. Yamaguchi.


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.


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).



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.


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.



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.


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


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.


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


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.


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


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


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.


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.


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.


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.


Reference:



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


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).


Reference:



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


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.


Reference:



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


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.


Reference:

LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.C.R CUNHA (in press) Zootaxa.


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


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


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


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.


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


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.


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


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.


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].


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.


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.


Reference:

BELLAN-SANTINI D. & M. THURSTON (1996) J. Nat.Hist. 30: 685-702.


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


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.


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


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.


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.


Reference:

MARTIN J.W. & G. PETTIT (1998) Bull. Mar. Sci. 63: 189-198.


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


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.


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).


Luckia striki BELLAN-SANTINI & THURSTON, 1996


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.


Reference:



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


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.


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.


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.


Arthropoda, Crustacea, Amphipoda, Gammaridea, Ischyroceridae

1: Female 3.4 mm, with both antennules and antennae lost; after SHAW (1989).


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.


Cyclocaris tahitensis STEBBING, 1888


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].



Euonyx mytilus BARNARD & INGRAM, 1990

Reference:

BARNARD J.L. & C. INGRAM (1990) Smithson. Contr. Zool. 499: 1-80.


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).


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.


Hirondellea glutonis BARNARD & INGRAM, 1990

Reference:



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).



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.


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.


1A: Head; B: Pleon; C: Gnathopod 1; D: Gnathopod 2; E: Pereopod 7; F: Uropod 3; G: Telson; from BARNARD & INGRAM (1990).



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.


1A: Head; B: Pleon; C: Gnathopod 1; D: Gnathopod 2; E: Pereopod 7; F: Uropod 3; G: Telson; from BARNARD & INGRAM (1990).



Tectovalopsis diabolus BARNARD & INGRAM, 1990

Reference:



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).



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.


Tectovalopsis wegeneri BARNARD & INGRAM, 1990



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.



Transtectonia torrentis BARNARD & INGRAM, 1990

Reference:



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).



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.


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.


Reference:

MARTIN J., FRANCE S.C. & C.L. VAN DOVER (1993) Pacific Can. J. Zool. 71: 1724-1732.


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


Pardalisca endeavouri SHAW, 1989

Reference:

SHAW P. (1989) Can. J. Zool. 67(8): 1882-1890.


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.


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.


1: Male 3.4 mm; after SHAW (1989).

Arthropoda, Crustacea, Amphipoda, Gammaridea, Sebidae


Steleuthera ecoprophycea BELLAN-SANTINI & THURSTON, 1996


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:



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


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].


Arthropoda, Crustacea, Amphipoda, Gammaridea, Stenothoidae

1: Habitus; from BARNARD (1966).


Stenothoe menezgweni BELLAN-SANTINI, 2005


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.


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).



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.


1A: Habitus; B: Gnathopod 1; C: Gnathopod 2; D: Uropod 1; E: Uropod 2; F: Uropod 3; G: Telson; from BELLAN-SANTINI (2005).



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].


1A: Habitus; B: Gnathopod 1; C: Uropod 3; D: Telson; from BARNARD & INGRAM (1990).

Arthropoda, Crustacea, Amphipoda, Lysianassoidea, Uristidae


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).


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


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


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


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.


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:


T. KOMAI & M. SEGONZAC Denisia 18 (2006): 414

1: Carapace and cephalic appendages, lateral.

2: Third to sixth abdominal somites, lateral.


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.


Alvinocaris longirostris KIKUCHI & OHTA, 1995

Size: Largest female 16.9 mm. Maximal total length ca. 85 mm.


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.


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.



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.


1: Body, lateral; enlargement x5.

3: Telson anduropods, dorsal;

enlargement x10.1-3: from WILLIAMS

& CHACE (1982).


2: Anterior part of carapaceand cephalic appendages; en-largement x10.


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.


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.


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


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).


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.



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.



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).


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.


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.



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.


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).


1: Body, lateral; scale bar 10 mm.


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.


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.



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.


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.



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).


2: Front, dorsal view.

3: Telson and right uropod,dorsal view (setae omitted).


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.


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).


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.


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.


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.


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.



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).


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.


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.



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.


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.



Shinkaicaris leurokolos (KIKUCHI & HASHIMOTO, 2000)

Size: An ovigerous female carapace length 12.5 mm.


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:



3: Telson and uropods,dorsal view. 1-3: Scale bar 1.0 mm;from KIKUCHI &HASHIMOTO (2000).


1: Holotype female in lateral view.

2: Anterior region,dorsal view.


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


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


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.


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


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


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


Paralomis verrilli (BENEDICT, 1895)


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.


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


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.


Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

1: Holotype male, carapacelength 7.2 mm; dorsal view;scale bar 1 mm; from BABA & TÜRKAY (1992).


Munidopsis acutispina BENEDICT, 1902

4: Specimen from Lost City; cruise Exomar; by P. Briand © Ifremer.


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.


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).


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).


Munidopsis alvisca WILLIAMS, 1988

2: Carapace, lateral view; scale bar 5 mm; by A.B. Williams.



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.


1: Carapace and abdomen, dorsalview; holotype carapace length23.7 mm (including rostrum);scale bar 5 mm; by A.B. Williams.


Munidopsis diomedeae (FAXON, 1893)


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.


1: Male, carapace length ca. 30 mm(with rostrum), habitus, dorsal view;from FAXON (1895).


Munidopsis exuta MACPHERSON & SEGONZAC, 2005

3: Specimen from Snake Pit; cruise Hydrosnake; by P. Briand© Ifremer.


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.


1: Carapace, dorsal view; holotype, carapace length34.7 mm; scale bar 5 mm; from MACPHERSON & SEGON-ZAC (2005).


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).


Munidopsis lauensis BABA & DE SAINT LAURENT, 1992

3: Munidopsis sp. (probably M. lauensis); cruise TUIM 07; by courtesy of C.R. Fisher.


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.


1: Carapace and abdomen, dorsal view; holo-type male, carapace length 10.8 mm; scalebar 1 mm; by K. Baba & M. de Saint Laurent.


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.


Munidopsis lentigo WILLIAMS & VAN DOVER, 1983

2: Carapace, lateral view; femaleparatype, carapace length 11.9 mm;from WILLIAMS & VAN DOVER (1983).



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.



Munidopsis marianica WILLIAMS & BABA, 1989

2: Rostrum and anterior part ofcarapace, dorsal view.


References:

BABA K. (2005) Galathea Report 20: 1-317.WILLIAMS A.B. & K. BABA (1989) Fish. Bull. 87: 899-910.


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.


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.


Munidopsis sonne BABA, 1995

3: Telson.


1: Carapace and abdomen, dorsal view.


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.


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.



Munidopsis starmer BABA & DE SAINT LAURENT, 1992

3: Telson; scale bar 1 mm.


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.


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.


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.


Munidopsis WHITEAVES, 1874


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.


1: Carapace, dorsal view; scale bar 5 mm;from BABA (2005).

2: Right, carapace, lateral view; bar 5 mm; from BABA (2005).



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.


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.



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).


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


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


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.



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.



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.



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).



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.



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


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


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.


1: Habitus; by H. Encarnação © MMF, 2000.

Chordata, Vertebrata, Chondrichthyes, Chimaeriformes, Chimaeridae


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.


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.


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


Ilyophis saldanhai KARMOVSKAYA & PARIN, 1999

R. CAUSSE, M. BISCOITO & P. BRIAND Denisia 18 (2006): 494-495

1: Holotype; from KARMOVSKAYA & PARRIN (1999).


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.


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


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.



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).


Reference:

SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.


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


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.



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.


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.


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.


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


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.


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.


1: Holotype, 422 mm standard length; by P. Drukker-Brammall, 1988.

Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae


Pachycara rimae ANDERSON, 1989


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.


1: Holotype, 403 mm standard length; © K. Klitz, 1989.



Pachycara saldanhai BISCOITO & ALMEIDA, 2004


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.


1: Habitus; by H. Encarmaçao © MMF, 2002.

2: At Rainbow vent field; cruiseMarvel © Ifremer.

3: At Rainbow vent field; cruise Marvel © Ifremer.



Pachycara thermophilum GEISTDOERFER, 1994


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.



Pyrolycus manusanus MACHIDA & HASHIMOTO, 2002


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.


1: Holotype; from MACHIDA & HASHIMOTO (2002).

2: At Manus Back-Arc Basin; cruise Bioaccess © JAMSTEC. 3: At Manus Back-Arc Basin; cruise Bioaccess © JAMSTEC.



Thermarces cerberus ROSENBLATT & COHEN, 1986


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


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


Careproctus hyaleius GEISTDOERFER, 1994


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


Trachyscorpia cristulata echinata (KOEHLER, 1896) “spiny scorpionfish”


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.



1: Menez Gwen; cruise Saldanha © Ifremer & FCT.

Chordata, Vertebrata, Osteichthyes, Scorpaeniformes, Sebastidae

2: Habitus; from HUREAU & LITVIENKO (1984).


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.



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


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


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


Fig. 1: Atlantic-EastPacific regions.

Fig. 2: Atlantic-EastPacific sites.

516


517

Fig. 3: Indian-West Pacificregions.

Fig. 4: Indian-West Pacific sites.


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


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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

,Hyd

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

,Hyd

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


Phyl

um

,C

lass

,O

rder

,Fa

mil

ySp

ecie

sD

istr

ibu

tio

nPa

ge

Mol

lusc

a,So

leno

gast

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


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


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


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


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


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

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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


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

orer

;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


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


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


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


Phyl

um

,C

lass

,O

rder

,Fa

mil

ySp

ecie

sD

istr

ibu

tio

nPa

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

paul

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

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464

530


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

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gonz

acia

mes

atla

ntic

a(W

ILLI

AM

S,19

88)

MA

R:LO

,LS,

MG

,RB,

SP,T

AG

473

Art

hrop

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Crus

tace

a,D

ecap

oda,

Varu

nida

eX

enog

raps

uste

stud

inat

usN

G,H

UA

NG

&H

O,2

000

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naw

aA

:NET

474

Echi

node

rmat

a,Cr

inoi

dea,

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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

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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

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SEN

&K

ORE

N,1

883

MA

R:LS

;non

-ven

tlo

cati

ons

479

Echi

node

rmat

a,H

olot

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idea

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rido

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eCh

irid

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hydr

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rmic

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OV

&G

EBRU

K,2

000

EPR:

17°S

-21°

S;M

anus

,NFi

ji48

0Ec

hino

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ata,

Oph

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idea

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iuri

da,O

phia

ctid

aeO

phia

ctis

tyle

riST

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R&

SEG

ON

ZAC,

2005

MA

R:M

G,3

8°N

482

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node

rmat

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phiu

roid

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rida

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iuri

dae

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TYLE

Ret

al.,

1995

MA

R;no

n-ve

ntlo

cati

ons

483

Echi

node

rmat

a,O

phiu

roid

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rida

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iuri

dae

Spin

ophi

ura

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veti

STÖ

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&SE

GO

NZA

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R:13

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18°S

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SEG

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ZAC,

2006

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13°N

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ORE

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GH,1

985

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alap

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487

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data

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ndri

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yes,

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aeri

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affi

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(CA

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O,1

868)

MA

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RB;n

on-v

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loca

tion

s49

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mae

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ydro

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ARD

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MA

NN,1

990

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R:LS

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tlo

cati

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492

Chor

data

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eich

thye

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min

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gosa

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ate

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ount

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493

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data

,Ost

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s,A

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bran

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ophi

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haiK

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Chor

data

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mob

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ifor

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drop

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ESQ

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1810

MA

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497

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data

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7M

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MG

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om,L

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;non

-ven

tlo

cati

ons

499

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data

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mes

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hiti

dae

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mic

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1990

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ER&

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inpr

ess

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17°S

501

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data

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s,Pe

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es,Z

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idae

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502

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oarc

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989

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apag

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aniB

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data

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erm

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GEI

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idae

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506

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mar

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es,L

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idae

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proc

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hyal

eius

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ER,1

994

EPR:

13°N

-9°N

509

Chor

data

,Ost

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s,Sc

orpa

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tida

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achy

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pia

cris

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taec

hina

ta(K

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LER,

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)M

AR:

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;non

-ven

tlo

cati

ons

510

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data

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ryph

aeno

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atus

HEC

TOR,

1875

EPR:

9°N

;Gal

apag

os;M

AR:

LS,R

B,SP

;51

1no

nve

ntlo

cati

ons

531


532

Cnidaria, Anthozoa


533

Mollusca, Gastropoda


534

Mollusca, Bivalvia


535

Mollusca, Cephalopoda


536

Annelida, Polychaeta


537

Annelida, Polychaeta


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.


539

Arthropoda, Copepoda

Arthropoda, Ostracoda


540

Arthropoda, Decapoda, Brachyura


541

Vertebrata, Osteichthyes


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


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


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


Documents

Handbook of Deep-Sea Hydro Thermal Vent Fauna