Sie sind auf Seite 1von 503

© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.

at

Handbook
of Deep-Sea
Hydrothermal
Vent Fauna
Second completely revised edition
Editors:
Daniel Desbruyères, Michel Segonzac & Monika Bright
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Impressum

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: bio-linz@landesmuseum.at
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 bio.buch@landesmuseum.at
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)
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Contents

Abstract, Résumé, Zusammenfassung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5


Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Photo credits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Contributors and adresses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Milestones in the discovery of hydrothermal-vent faunas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Hydrothermal vent meiofauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Hydrothermal vent parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

Species descriptions

Granuloreticulosa, Foraminifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Porifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35
Demospongiae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Hexactinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

Cnidaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .48
Hydrozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Scyphozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Anthozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

Mollusca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Solenogastres . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Polyplacophora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Gastropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Patelligastropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Vetigastropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Neomphalina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Neritimorphoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
Caenogastropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Heterobranchia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Prosobranchia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Nudibranchia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Bivalvia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Heterodonta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Protobranchia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Pteriomorphia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Cephalopoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166

Nematoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173

Acanthocephala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

Nemertini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Polychaeta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Eunicida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .186
Phyllodocida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
Sabellida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
Scolecida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266
Spionida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272
Terebellida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Oligochaeta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297

Arthropoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .298
Arachnida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298
Pycnogonida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .301
Crustacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
Ostracoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
Copepoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316
Cirripedia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 356
Leptostracta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .369
Cumacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .370
Tanaidacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .372
Isopoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Amphipoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
Ephausidacae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409
Decapoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 410
Caridea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .410
Astacidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .433
Anomura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
Brachyura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455

Echinodermata
Crinoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475
Asteroidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
Echinoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 479
Holothuroidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480
Ophiuroidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481

Chaetognatha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 486
Hemichordata, Enteropneusta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487
Chordata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 489
Chondrichthyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 489
Osteichthyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 493

Major known deep-sea hydrothermal vent fields . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 513


Abbreviations, ridges, back-arc basins and vent fields . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 518
Species list . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 519
Schematic drawings of major taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 532
Genus index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 542
Author index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 544
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Abstract
The second extensively expanded edition of the “Hand- schematic drawings, which aim to help non-specialists to iden-
book of Deep-Sea Hydrothermal Vent Fauna“ gives on over- tify the animals. 86 authors contributed with their expertise to
view of our current knowledge on the animals living at hydro- create a comprehensive database on animals living at hydro-
thermal vents. The discovery of hydrothermal vents and pro- thermal vents, which contains information on the morphology,
gresses made during almost 30 years are outlined. A brief intro- biology, and geographic distribution of more than 500 current-
duction is given on hydrothermal vent meiofauna and parasites. ly described species belonging to one protist and 12 animal
Geographic maps and a table of mid-ocean ridges and back-arc phyla. It comprises also the largest collection of more than
basins with the major known hydrothermal vent fields, their lo- 1000 pictures of hydrothermal vent animals taken in situ with
cation and depth range and the most prominent vent sites are submersibles, in vivo after collection, and with various dissec-
provided. Higher taxa are presented individually with informa- tion, light, and scanning electron microscopes after fixation
tion on the current taxonomic and biogeographic status, the and preparations.
number of species described, recommendations for fixation, and

Résumé
La seconde édition, considérablement étendue et améliorée champs hydrothermaux le long des rides océaniques et dans les
du « Handbook of Deep-Sea Hydrothermal Vent Fauna » don- bassins arrière-arc sont proposées assorties d’un tableau qui
ne un panorama de notre connaissance sur les animaux qui vi- donne leur position géographique, leur profondeur et les sur-
vent autour des sources hydrothermales océaniques. Les avan- noms des principaux sites. 86 auteurs ont contribué à la créa-
cées faites depuis leurs découvertes il y a trente ans environ, tion d’une base de données sur les animaux présents dans cet
donnent lieu à une mise en perspective rédigée par des acteurs écosystème surprenant, en donnant des informations sur la
des campagnes océanographiques. Une brève introduction est morphologie, la biologie et la distribution géographique de plus
proposée sur la méiofaune et les parasites, puis chaque taxon su- de 500 espèces appartenant à 12 phylums animaux. Cette base
périeur est présenté avec des rappels sur la taxinomie, des in- de données, présentée sous forme de fiches individuelles, re-
formations sur sa distribution géographique, des représentations groupe une série unique de plus de 1000 illustrations compor-
schématiques de la morphologie permettant à des non-spécia- tant dessins au trait, photographies des organismes dans leur
listes de comprendre la nomenclature, des informations sur le milieu, vues d’animaux isolés après récoltes et micro-photogra-
nombre d’espèces présentes, des conseils pour la fixation et la phies prises au microscope électronique à balayage.
conservation des échantillons. Des cartes de distribution des

Zusammenfassung
Die zweite, wesentlich erweiterte Auflage des „Handbook die Anzahl der beschriebenen Arten, Methoden zur Fixierung
of Deep-Sea Hydrothermal Vent Fauna“ gibt einen Überblick und schematischen Zeichnungen, die es dem Nicht-Spezialis-
über den momentanen Stand der Forschung. Die Entdeckung ten die Identifizierung erleichtern sollen. Über 80 Autoren ga-
der Hydrothermalquellen, Fortschritte der letzten beinahe 30 ben ihren Beitrag zur Erstellung einer umfassenden Datenbank
Jahre, sowie Meiofauna und Parasiten der Hydrothermalquellen mit mehr als 500 Tieren aus einem Protisten-Stamm und 12
werden kurz umrissen. Alle ozeanischer Rücken und Back-Arc Tierstämmen und Informationen zu deren Morphologie, Biolo-
Basins mit ihren bekannten Hydrothermalquellfeldern, ihre ge- gie und Verbreitung. Darüber hinaus werden die Tiere der
naue Lage und Tiefe, sowie ihre benannten Fundstellen werden Hydrothermalquellen in mehr als 1000 Bildern, Unterwasser-
in geographischen Karten und einer Tabelle gezeigt. Höhere aufnahmen von bemannten und unbemannten U-Booten, in
Taxa werden individuell dargestellt mit Information über den vivo Aufnahmen und diversen licht- und elektronenmikrosko-
momentanen taxonomischen und biogeographischen Stand, pischen Aufnahmen, dargestellt.

Key words: Hydrothermal vents, deep sea, mid-ocean ridge, back-arc basin, hydrothermal vent fauna.

5
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

« Toute partie dans la chaise qui ne se voyait pas était tout


aussi 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 view
are as perfectly crafted as the visible ones. This ist the very
principle of the cathedrals.“

Foreword
Deep-sea hydrothermal vents and their associated fauna work has been conducted for three decades producing more
were discovered in 1977. The presence of these rich oases of li- than 500 fully identified species of which descriptions or
fe in the food-limited environment of basaltic Mid-Oceanic records are dispersed throughout the scientific literature. Nev-
Ridges fostered the interest of the marine biologists to explore ertheless the list of animal species living at vents is still evolv-
the deep sea using submersibles with a focus on fragmented ha- ing every week; new species are described from additional sam-
bitats. During the intervening three decades, a large number of ples in known locations and from new locations, descriptions
cruises using deep submergence vehicles (HOV or ROV) were are revisited, sibling species are detected based on molecular
devoted to the biological study of the hydrothermal communi- taxonomy, and new combinations are proposed based on stud-
ties on the Pacific, Atlantic, and Indian Ridges as well as in ies of large series and molecular information. The technology of
Back-Arc Basins and Volcanic Arcs of the Pacific Rim of Fire. imaging in situ greatly improved in the last few years giving
These series of cruises were mainly devoted to the understan- considerable information on the shape, colors, behavior, and
ding (1) the distribution, the phylogeny, and the dispersal of micro-distribution of living animals. The dialogue between
this specialized fauna which is restricted to this fragmented “ex- classical and molecular taxonomy brought new insights and
treme” habitat along the world’s oceanic ridges and (2) to the clarifies the biogeographic patterns of vent communities as well
niche characterization and the study of the physiological adap- as their relationships with other deep-sea reduced habitats
tations of the organisms (or symbiotic associations) living in (cold seeps, food falls, and minimum oxygen zones).
the mixing zone between the superheated fluid and the cold
In the foreword of the first edition of the “Handbook of
deep-sea water. This endeavour remains unique in its extent
Deep-Sea Hydrothermal Vent Fauna” (Ifremer Editions, 1997),
and intensity in the history of biological exploration of the
we proposed to publish periodic addenda to the first edition
world’s oceans since the early studies of the deep-sea by the pio-
based on new published observations and we invited contribu-
neering expeditions of the nineteenth century. However, be-
tors to help us by sending information and illustration. The re-
cause of logistic constrains, this accomplishment is still far from
sults far exceeded our hopes and we realized soon that a new is-
providing an exhaustive overview of the composition and dis-
sue of the handbook would be fully justified. At the same time,
tribution of hydrothermal vent fauna; some Ridges (South At-
the Biologiezentrum der Oberösterreichischen Landesmuseen
lantic and the three Indian Ridges) still deserve obviously mo-
(Biology Center of the Upper Austrian Museums) was prepar-
re attention and the study of remote targets as high latitude rid-
ing an exhibition on deep-sea hydrothermal vent life planned
ges (Arctic and Antarctic Seas) or isolated basins (South Sand-
for April 2006 and was looking for authors willing to publish a
wich and Andaman Sea) would bring valuable information on
review on vent biology. It was the perfect opportunity for us to
the biogeography of the vent fauna. Multiyear surveys on the
restart the collection of contributions from the worldwide net-
East Pacific Rise and the Juan de Fuca Ridge have also demon-
work of taxonomists involved in the study of hydrothermal
strated that the hydrothermal environment is submitted to
vent samples. We want to thank here all the contributors who
temporal instability due to volcanic and tectonic events; this
participated in the wording of this new issue.
instability greatly affects the patterns of the ecosystem deve-
lopment and produces a variation in specific composition of This book, published under the auspices of InterRidge and
communities only detected by temporal series of sampling. Census of Marine Life/ChEss, is an overview of the present tax-
onomic knowledge of the deep-sea hydrothermal vent fauna,
Reliable morphological identification of collected speci-
intended for scientists and submersible pilots working at sea,
mens presently remains the inescapable base of most of the bi-
but also for students and the general public curious about the
ological work conducted at vents on animals, including the mo-
life in the ocean.
lecular approach. Because of the taxonomic novelty of deep-sea
hydrothermal vent communities, an impressive descriptive D. Desbruyères, M. Segonzac & M. Bright

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© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Remarks
Collection of animals as well as in situ photographs and vi- We stress our belief that our successes in understanding the
deos of animals from a deep-sea environment are mainly achie- fauna living at hydrothermal vents will be highly influenced by
ved through the use of manned and unmanned submersibles. the way we approach solving these issues. So we recommend for
Valuable sampled animals are often a by-product of large bulk taxonomy, ecology and outreach purposes, that researchers ta-
samples, fixation and storage sometimes are less than optimal ke the time on the bottom to video and photograph, that scien-
for all taxa, and the time for taking pictures while on board is tists onboard research vessels take high quality photographs of
often scarce. Also, it is often due to pure luck that in situ animals before preservation as well as follow instructions for
photographs and videos are taken at the decision of chief scien- proper fixation and storage, and that they get into contact with
tists, pilots, and researchers on board as they must interrupt taxonomists for identification and appropriate procedures for
their ongoing tasks and use valuable bottom time to document voucher specimens. Moreover, we hope that more taxonomists
animals in situ. Good quality pictures taken on the bottom pro- will be invited to research cruises to better support our goal of
vide a unique set of information on the micro-distribution, be- learning more about hydrothermal vents and their inhabitants.
havior, and biological interactions. Even those photographs, Recent experiences demonstrated that the presence of taxono-
which are in low resolution or out of focus, are highly valuable mists on cruises improved by far our knowledge of the faunal
because they give us an impression of where these animals oc- composition of vent communities and fostered interactions
cur and how they behave naturally. with other disciplines.
D. Desbruyères, M. Segonzac & M. Bright

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© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Acknowledgements
We would like to thank all the contributors whose know- We express our heartful thanks to the chief scientists Fern-
ledge of the fauna and taxonomic expertise allowed us to revi- ando Barriga, Nadine Le Bris, Charles R. Fisher, Francoise
se and supplement the second edition of this book. We are gra- Gaill, Anne Godfroy, Jun Hashimoto, S. Kim Juniper, François
teful to Gerhard Aubrecht, the head/director of the Biologie- Lallier, Marvin D. Lilley, Pierre-Marie Sarradin, Tim Shank,
zentrum der Oberösterreichischen Landesmuseen (Biology Verena Tunnicliffe, Karen Von Damm, Cindy L. Van Dover
Center of the Upper Austrian Museums), for making this pu- and Robert C. Vriejenhoek, who allowed us to have access to
blication possible and for his continuous support throughout samples, data, and in situ pictures. We are also grateful to the
the entire process. Special thanks to Erna Aescht (Biology many contributors who shared their pictures with us.
Center) for the editing, Eva Rührnößl (Linz) for the page-ma-
This book was supported by the Federal Government of
king, Heidemarie Grillitsch (University Vienna) for the sche-
Upper Austria, the Biology Center of the Upper Austrian Mu-
matic drawings, Ingrid Kolar (University Vienna) for checking
seums, Ifremer, ChEss (especially the coordinators of the ChEss
the numerous references, and Patrick Briand (Ifremer) for his
project Eva Ramirez-Llodra and Maria Baker), InterRidge, the
help with the pictures, Violaine Martin (Ifremer) for many dra-
Austrian Science Foundation, and the Faculty of Life Sciences,
wings, and Philippe Crassous (Ifremer) for his technical exper-
University of Vienna.
tise with the SEM preparations.
D. Desbruyères, M. Segonzac & M. Bright

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Photo credits
Auger, V. – Canadian Scientific Submersible Facility, Canada Lozouet, P. – National Muséum d’Histoire Naturelle, Paris,
Biscoito, M. – Museu Municipal do Funchal, Portugal France
Blake, J. – ENSR Marine & Coastal Center, USA Lutz, R. A. – Rutgers University, USA
Blaston, P. – University of Otago, New Zealand Maddocks, R. M. – University of Houston, USA
Briand, P. – Ifremer, Brest, France Maestrati, P. – Muséum National d’Histoire Naturelle, Paris,
Bright, M. – University of Vienna, Austria France
Burreson, E. – College of William and Mary, Williamsburg, Marcus, J. – University of Victoria, Canada
USA Martin, V. – Ifremer, Brest, France
Chan, T. Y. – National Taïwan Ocean University, Taïwan McLean, J. – Museum of Natural History, Los Angeles, USA
Cosel, R. von – Muséum National d’Histoire Naturelle, Paris, Mills, S.W. – Woods Hole Oceanographic Institution, USA
France Miura, T. – JAMSTEC, Nagasaki, Japan
Cunha, M.R. – Centro das Zonas Costeiras e do Mar, Universi- Mullineaux, L. – Woods Hole Oceanographic Institution, USA
dad, Portugal Ng, N. K. – National University of Singapore, Republic of Sin-
Fautin, D. – Natural History Museum and Biodiversity Rese- gapore
arch Center, Kansas, USA Okata, Y. – JAMSTEC, Nagasaki, Japan
Dejouannet, J.-F. – IRD (Institut Recherche Développement) Pleijel, F. – University of Göteborg, Sweden
Paris, France Rouse, G. – University of Adelaide, Australia
Drukker-Brammall, P. – Smith Institute of Ichthyology, Gra- Roux, M. – Université de Reims, France
hamstown, South Africa Sagalevitch, A.M. – Shirshov Institute of Oceanology, Mos-
Dugornay O. – Ifremer, Brest, France cow, Russia
Encarnação, H. – Museu Municipal do Funchal, Portugal Saldanha, L. † – LMG, Lisbonne, Portugal
Fifis, A. – Ifremer, Brest, France Shagin, A. – PMGE (Mining Intitute, St Petersburg), Russia
Fisher, C. R. – The Pennsylvania State University, USA Smith Jr, K. – Scripps Institution of Oceanography, USA
Glover, A. – The Pennsylvania State University, USA Southward, E.C. – Plymouth Marine Laboratory, United King-
Guinot, D. – Muséum National d’Histoire Naturelle, Paris, dom
France Stöhr, S. – Swedish Museum of Natural History, Sweden
Harrison-Nelson, E. – National Museum of Natural History, Takeda, M. – JAMSTEC, Japan
Smithsonian Institution, USA Tivey, M. – Woods Hole Oceanographic Institution, USA
Hashimoto, J. – JAMSTEC, Nagasaki, Japan Tsuchida, S. – JAMSTEC, Japan
Heemstra, E. – Smith Institute of Ichthyology, Grahamstown, Tunnicliffe, V. – University of Victoria, Canada
South Africa
Van Dover, C.L. – College of William and Mary, Williamsburg,
Hessler, R.R. – Scripps Oceanographic Institution, USA USA
Hourdez, S. – Station Biologique Roscoff, France Verschelde, D. – Universiteit Gent, Belgium
Ivanenko, V. N. – Moscow State University, Russia Vrijenhoek, R. – Monterey Bay Aquarium Research Institute,
Juniper, K. S. – Université du Québec, Montréal, Canada USA
Klitz, K. – National Museum of Natural History, Smithsonian Warén, A. – Swedish Museum of Natural History, Sweden
Institution, USA Wilson, G.D.F. – Australian Museum, Syndney, Austrialia
Kornicker, L.S. – National Museum of Natural History, Smith- Young, C.M. – Origon Institute of Marine Biology, USA
sonian Institution, USA
Zierenberg, R. – University of California, Davis, USA
Kudenov, J.D. – University Alaska, Anchorage, USA
Le Goff, A. – Muséum National d’Histoire Naturelle, Paris,
France
Lindsay, S. C – Australian Museum, Syndney, Austrialia
Lopez-Gonzalez, P. – Universidad de Sevilla, Spain

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© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Contributors and addresses


Almeida, Armando J. – Laboratório Marítimo da Guia, IMAR, Burreson, Eugene M. – Department of Environmental and
Faculdade de Ciências da Universidade de Lisboa, Estrada Aquatic Animal Health, Virginia Institute of Marine Scien-
do Guincho, 2750-642 Cascais, Portugal aalmeida@fc.ul.pt ce, College of William and Mary, PO. Box 1346, Gloucester
Baba, Keiji – Kumamoto University, Faculty of Education, 2-40- Point, VA 23062 USA gene@vims.edu
1 Kurokami, Kumamoto 860-8555, Japan Casanova, Bernadette – Université de Provence, Biologie Ani-
keiji5962@yahoo.co.jp male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé-
Bamber, Roger – Environment: Coastal & Marine, The Natural dex 3, France bioplank@newsup.univ-mrs.fr
History Museum, Cromwell Road, London SW7 5BD, Uni- Casanova, Jean-Paul – Université de Provence, Biologie Ani-
ted Kingdom r.bamber@nhm.ac.uk male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé-
Barthélémy, Roxane – Université de Provence, Biologie Ani- dex 3, France bioplank@newsup.univ-mrs.fr
male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé- Causse, Romain – Muséum National d’Histoire Naturelle, La-
dex 3, France bioplank@newsup.univ-mrs.fr boratoire d’Ichthyologie, 43, rue Cuvier, 75231 Paris cedex
Bartsch, Ilse – Biologische Anstalt Helgoland, Zentrale Ham- 05, France causse@mnhn.fr
burg, Notkestrae 31, D-22607 Hamburg, Germany Cohen, Daniel M. – California Academy of Sciences, Golden
bartsch@meeresforschung.de Gate Park, San Francisco, CA 95521, USA dmco3@com-
Bellan-Santini, Denise – Université d’Aix-Marseille 2, Station cast.net
Marine d’Endoume-Luminy, Rue de la Batterie des Lions, Corbera, Jordi – Carrer Gran, 90, 08310 Argentona, Spain cor-
13007 Marseille, France bellan@com.univ-mrs.fr bera@sct.ictnet.es
Biscoito, Manuel – Museu Municipal do Funchal, Rua da Mou- Cosel, von Rudo – Département Systématique et Évolution,
raria, 31, 9000 Funchal, Madeira, Portugal manuel.biscoi- Muséum National d’Histoire Naturelle, Taxonomie-Collec-
to@mail.cm-funchal.pt tion Mollusques, 55 rue Buffon, 75231 Paris, France co-
Blake, James A. – ENSR Marine & Coastal Center, 89 Water sel@cimrs1.mnhn.fr
Street, Woods Hole, MA 02543, USA jablake@ix.net- Cunha, Marina R. – Centro das Zonas Costeiras e do Mar, Uni-
com.com versidad, Departamento de Biologia, 3810-193 Aveiro, Por-
Böggemann, Markus – Habsburgerallee 37, 60385 Frankfurt am tugal mcunha@bio.ua.pt
Main, Germany mboeggem@aol.com Daly, Marymegan – Dept. Evolution, Ecology and Organismal
Bouchet, Philippe – Département Systématique et Évolution, Biology, Ohio State University, 315 Kinnear RD, Columbus
Muséum National d’Histoire Naturelle, Taxonomie-Collec- OH 43212, USA daly.66@osu.edu
tion Mollusques, 55 rue Buffon, 75231 Paris, France pbou- Defaye, Danielle – Département Milieux et Peuplements Aqua-
chet@mnhn.fr tiques, Muséum National d’Histoire Naturelle, 61, rue de
Boury-Esnault, Nicole – Université d’Aix-Marseille 2, Centre Buffon, 75005 Paris, France ddefaye@cimrs1.mnhn.fr
d’Océanologie de Marseille, Station Marine d’Endoume - Desbruyères, Daniel – Département «Etude des écosystèmes pro-
Luminy, Rue de la Batterie des Lions, 13007 Marseille, Fran- fonds», Ifremer, Centre de Brest, BP 70, 29280 Plouzané,
ce esnault@com.univ-mrs.fr France Daniel.Desbruyeres@ifremer.fr
Briand, Patrick – Département «Etude des écosystèmes pro- Gebruk, Andrey V. – Laboratory of Ocean Benthic Fauna, P.P.
fonds», Ifremer, Centre de Brest, BP 70, 29280 Plouzané, Shirshov Institute of Oceanology, Nakhimovsky Pr., 36,
France patrick.briand@ifremer.fr Moscow 117997, Russia agebruk@sio.rssi.ru
Bright, Monika – Department of Marine Biology, University of Geistdoerfer Patrick – Laboratoire d’Ichthyologie, 43, rue Cu-
Vienna, Althanstr. 14, A-1090 Vienna, Austria vier, 75231 Paris, France Patrick.Geistdoerfer@
monika.bright@univie.ac.at cimrs1.mnhn.fr
Buron, de Isaure – Division of Natural Sciences and Enginee- Gonzalez, Angel – Ecología y Biodiversidad Marina (ECOBIO-
ring, University of South Carolina Spartanburg, Spartan- MAR), Instituto de Investigaciones Marinas (CSIC), C/
burg, SC 29303, USA deburoni@cofc.edu Eduardo Cabello 6, 36208 Vigo, Spain afg@iim.csic.es

10
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Guerra, Angel – Ecología y Biodiversidad Marina (ECOBIO- Larsen, Kim – Department of Invertebrates, Zoological Mu-
MAR), Instituto de Investigaciones Marinas (CSIC), C/ seum, Universitetsparken 15, 2100 KBH, Denmark ta-
Eduardo Cabello 6, 36208 Vigo, Spain brc1@iim.csic.es naids@hotmail.com
Guinot, Danièle – Département Milieux et Peuplements Aqua- Lemaitre, Rafael – Smithsonian Institution, Department of Sys-
tiques, Muséum National d’Histoire Naturelle, 61, rue de tematic Biology, NMHN, MRC 163, PO Box 37012, Was-
Buffon, 75005 Paris, France guinot@mnhn.fr hington, DC 20013-7012, USA lemaitre.rafael@
Haney, Todd – Department of Organismic Biology, Ecology, and nmnh.si.edu
Evolution, University of California Los Angeles, 621 Char-
Lopez-Gonzalez, Pablo – Departamento de Fisiologia y Biologia
les E. Young Drive South, Box 1606, Los Angeles, Califor-
Animal, Facultad de Biologia, Universidad de Sevilla, Rei-
nia 90095-1606, USA haney@ucla.edu
na Mercedes, 6, 41012 Sevilla , Spain pjlopez@us.es
Harrison-Nelson, Elisabeth – Department of Invertebrate Zoo-
logy, National Museum of Natural History, Smithsonian In- Lutz, Richard A. – Center for Deep-Sea Biology & Biotechno-
stitution, MRC 163, Washington, D. C. 20013-7012, USA logy, Rutgers University, Dudley Road, New Brunswick, NJ
nelsone@si.edu 08903-0231, USA rlutz@imcs.rutgers.edu
Hashimoto, Jun – Japan Marine Science and Technology Cen- Macpherson, Enrique – CSIC/CEAB, Camí Santa Bàrbara, s/n,
ter (JAMSTEC) 2-15, Natsushima-Cho, Yokosuka, 237, Ja- 17300 Blanes, Girona, Spain macpherson@ceab.csic.es
pan junh@net.nagasaki-u.ac.jp
Maddocks, Rosalie F. – Department of Geosciences, Room 312
Holzmann, Maria – Department of Paleontology, University of Science & Research Bldg. 1, University of Houston, Hous-
Vienna, Althanstr. 14, A-1090 Vienna, Austria maria.holz- ton, TX 77204-5007, USA RMaddocks@uh.edu
mann@univie.ac.at
Mah, Christopher – Department. of Invertebrate Zoology, Na-
Hourdez, Stephan – Station Biologique, 29680 Roscoff, France tional Museum of Natural History, MRC-163, PO Box
shourdez@sj2.seanet.in
37012, Smithsonian Institution, Washington DC 20013,
Hurtado, Luis – University of Arizona, 310 Biosciences West, USA brisinga@gmail.com
P.O. Box 210088, Tucson, AZ, 85721-0088, USA lhurta-
Miura, Tomoyuki – Faculty of Fisheries, Kagoshima University,
do@u.arizona.edu
4-50-20, Shimoarata, Kagoshima, 890 Japan
Ivanenko, Viatcheslav N. – Department of Invertebrate Zoolo- miura@ugs.agri.kagoshima-u.ac.jp
gy, Biological Faculty, Moscow State University, Moscow
119899, Russia ivanenko.slava@mail.ru Möller, Peter R. – Zoological Museum, University of Copenha-
gen Universitetsparken 15, 2100 KBH, Denmark PDRMol-
Jaume, Damian – IMEDEA (CSIC-UIB), Instituto Mediterra-
ler@zmuc.ku.dk
neo de Estudios Avanzados, C/ Miquel Marques 21, 07190-
Esporles (Mallorca, Illes Balears), Spain d.jaume@uib.es Moreau, Xavier – Université de Provence, Biologie Animale
Jones, Diana S. – Western Australian Museum, Locked Bag 49. (Plancton), 3, place Victor Hugo, 13331 Marseille, Cédex 3,
WELSHPOOL DC, Western Australia 6986, Australia dia- France bioplank@newsup.univ-mrs.fr
na.jones@museum.wa.gov.au Myers, Alan A. – Department of Zoology, Ecology and Plant
Justine, Jean-Lou – Institut Développement Recherche (IRD), Science, National University of Ireland, Cork, Republic of
Nouméa, Nouvelle-Calédonie, France jean- Ireland alanmyers@crustacea.net
lou.justine@noumea.ird.nc Newman, William A. – Scripps Institution of Oceanography, La
Komai, Tomoyuki – Natural History Museum and Institute Chi- Jolla, CA 92093-0202, USA wnewman@ucsd.edu
ba, 955-2, Aoba-cho, Chuo-ku, Chiba 260-8682, Japan ko-
Ng, Ngan Kee – Department of Biological Sciences, National
mai@chiba-muse.or.jp
University of Singapore, 14, Science Drive 4, Singapore
Kornicker, Louis S. – Department of Invertebrate Zoology, Na- 117543, Republic of Singapore ngankee@nus.edu.sg
tional Museum of Natural History, Smithsonian Institution,
MRC 163, Washington, D. C. 20013-7012, USA kor- Nielsen, Jorgen G. – Department of Invertebrates, Zoological
nickl@si.edu Museum, Universitetsparken 15, 2100 KBH, Denmark
JGNielsen@zmuc.ku.dk
Kudenov, Jerry D. – Department Biological Sciences, Universi-
ty Alaska Anchorage, 3211 Providence Drive, Anchorage, Okutani, Takashi – Tokyo University of Fisheries, Konan 4-5-7,
Alaska 99508, USA afjdk@uaa.alaska.edu Minato-ku, Tokyo 108, Japan okutani@jamstec.go.jp

11
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Pleijel, Fredrik – Tjärnö Marine Biological Laboratory, Depart- Van Dover, Cindy Lee – Biology Department, The College of
ment of Marine Ecology, Göteborg University, SE-452 96 William & Mary Williamsburg, VA 23187, USA
Strömstad, Sweden fredrik.pleijel@tmbl.gu.se clvand@wm.edu
Pugh, Phillip R. – School of Ocean and Earth Science, National Verschelde, Dominick – Universiteit Gent, Museum voor Dier-
Oceanography Centre, European Way, Southampton, SO14 kunde, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium do-
3ZH, United Kingdom prp@noc.soton.as.uk minick.verschelde@ugent.be

Ramirez-Llodra, Eva – Institut de Ciències del Mar, CMIMA- Vervoort, Willem – National Museum of Natural History, P.O.
CSIC, Passeig Marítim de la Barceloneta 37-49, E-08003 Box 9517, 230 RA Leiden, The Netherlands vervoort@na-
Barcelona, Spain ezr@cucafera.cmima.csic.es turalis.nnm.nl

Rodríguez, Estefanía – Departamento de Fisiologia y Biologia Villanueva, Roger – Instituto de Ciencias del Mar (CSIC)Insti-
Animal, Facultad de Biologia, Universidad de Sevilla Reina tuto de Ciencias del Mar (CSIC), Paseo Juan de Borbon s/n
Mercedes, 6, 41012 Sevilla, Spain fani@us.es E-08039 Barcelona, Spain roger@icm.csic.es
Voight, Janet R. – Department of Zoology, The Field Museum of
Roux, Michel – Laboratoire des Sciences de la Terre, Centre de
Natural History, Roosevelt Road at Lake Shore Drive, Chi-
Recherches Agronomiques, 2 esplanade Roland Garros,
cago, IL 60605, USA jvoight@fieldmuseum.org
51100 Reims, France michel.roux@univ-reims.fr
Vrijenhoek, Robert C. – Monterey Bay Aquarium Research In-
Salvini-Plawen, von Luitfried – Department of Evolutionary
stitute, 7700 Sandholdt Road, Moss Landing, CA 95039-
Biology, University of Vienna, Althanstr. 14, A-1090 Vien-
9644, USA vrijen@ahab.rutgers.edu
na, Austria luitfried.salvini-plawen@univie.ac.at
Warén, Anders – Swedish Museum of Natural History, Depart-
Schein, Elisabeth – Laboratoire des Sciences de la Terre, Centre
ment of Invertebrate Zoology, Box 50 007, Stockholm, S-
de Recherches Agronomiques, 2 esplanade Roland Garros,
10405 Sweden anders.waren@nrm.se
51100 Reims, France elisabeth.schein@univ-reims.fr
Wilson, George D. F. – Australian Museum, 6 College Street,
Segonzac, Michel – Département «Etude des écosystèmes pro- Sydney NSW 2010, Australia buzw@austmus.gov.au
fonds», Ifremer, Centre de Brest, BP 70, 29280 Plouzané,
Yamaguchi, Toshiyuki – Chiba University, Marine Biosystems
France segonzac@ifremer.fr
Research Center 1-33, Yayoi-cho, Inage, Chiba 263-8522,
Senz, Wolfgang – Department of Evolutionary Biology, Univer- Japan tyamaguc@earth.s.chiba-u.ac.jp
sity of Vienna, Althanstr. 14, A-1090 Vienna, Austria
Young, Craig – Oregon Institute of Marine Biology, University
Southward, Alan J. – Plymouth Marine Laboratory, Citadel of Oregon, Charleston, OR 97420, USA cmyoung@uore-
Hill, Plymouth, PL1 2PB, United Kingdom gon.edu
100721.3720@compuserve.com
Zekely, Julia – Department of Marine Biology, University of
Southward, Eve C. – Plymouth Marine Laboratory, Citadel Hill, Vienna, Althanstr. 14, A-1090 Vienna, Austria
Plymouth, PL1 2PB, United Kingdom 100721.3720@com- j.zekely@gmx.at
puserve.com Zibrovius, Helmut – Université d’Aix-Marseille 2, Station Ma-
Stöhr, Sabine – Swedish Museum of Natural History, Depart- rine d’Endoume, Rue de la Batterie des Lions, 13007 Mars-
ment of Invertebrate Zoology, Box 50 007, Stockholm SE, eille, France zibrowius@com.univ-mrs.fr
10405 Sweden sabine.stohr@nrm.se
Tsuchida, Shinji – Marine Ecosystems Research Department, Ja-
pan Marine Science and Technology Center, 2-15 Natsushi-
ma-cho, Yokosuka, Kanagawa, 237-0061, Japan tsuchi-
das@jamstec.go.jp
Tunnicliffe, Verena – School of Earth and Ocean Sciences, and
Department of Biology, University of Victoria, Victoria
V8W 2Y2, Canada verenat@uvic.ca
Tyler, Paul A. – Department of Oceanography, Southampton
Oceanography Centre, Southampton, SO14 3ZH, United
Kingdom P.A.Tyler@soton.ac.uk
Vacelet, Jean – Station Marine d’Endoume, Rue de la Batterie
des lions, 13007 Marseille, France jvacelet@com.univ-
mrs.fr

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Milestones in the discovery of hydrothermal-vent faunas

Seafloor Spreading and Hot Springs During Alvin dives at the Galapagos Spreading Center in
1977, geologist Jack Corliss first described vent mussels (al-
Any account of the discovery of hydrothermal-vent faunas
though he erred in his initial identification). His difficulty in
must begin with the geology of seafloor spreading centers. The
finding the words to describe what was before him, a vista no
symmetry of magnetic anomalies on either side of the mid-ocean
one had ever seen before, is evident:
ridges that girdle the globe and the correspondence of the anom-
aly patterns with the pattern of magnetic reversals on Earth con- “They are abalone shells. They are shells. They are big
firmed the process of seafloor spreading and led to general ac- shells. They are living. … attached shellfish….”
ceptance of plate tectonic theory in the early 1960s (VINE & The wonder in his voice is captured on the audio record of
MATTHEWS 1963). The bathymetric relief of mid-ocean ridges the dive.
was understood to be a consequence of the thermal buoyancy of
hot rock in volcanic systems. Conductive heat loss was expect- Further into the dive series, geochemist John Edmond first
ed to be greatest at the axis of these linear volcanoes and to di- sounds incredulous as he provided the first description of giant
minish along transects away from the ridge crest, but heat-flow worms, and then frustrated as his observations were limited by
measurements collected by placing a vertical array of thermistors the green light of the thallium iodide bulb on Alvin:
into seafloor sediments consistently documented a heat-deficit “There are big ones [worms] out there. Looks like an Indian
near the ridge axis. This heat deficit suggested that conductive [Native American] headdress. There are four of them in a row,
heat loss was not the only operative mode of cooling (STEIN et right outside my view port. I wonder what color they are. You
al. 1995). Geologists hypothesized that the convective heat can’t tell. They may be red.”
driven by cells of seawater percolating into the crust and subse-
quent heating and buoyancy-driven flux of fluids out of the crust Vent animals were useful as flux indicators of hydrothermal
would account for the missing heat. Thus the presence of hot activity for geologists prospecting for vents, but geologists also
springs on the seafloor with temperatures as great as 300°C was appreciated the need for an explanation of the tremendous bio-
anticipated (TALWANI et al. 1971; LISTER 1972; WOLERY & SLEEP mass of animals and of the means by which species could be
1976), and the first unequivocal evidence of warm-water, buoy- maintained at vents in the face of inevitable local extinctions.
ant plumes was collected by May 1976 using the Scripps Institu- Following the first Alvin dive series to Galapagos vents, micro-
tion of Oceanography Deep-Tow vehicle (WEISS et al. 1977). biologist John Baross postulated that the millimolar concentra-
While systematic geophysical studies predicted the existence of tions of hydrogen sulfide in vent fluids were a source of reduced
hydrothermal vents on the seafloor, the discovery of their atten- sulfur for free-living chemolithoautotrophic, sulfur-oxidizing
dant chemosynthetic ecosystems was unimagined. bacteria (cited in CORLISS et al. 1979). Chemolithoautotrophs
are primary producers that use the energy from the oxidation of
reduced compounds like hydrogen sulfide to yield ATP. That
Strange Animals at Hydrothermal Vents on ATP is then used to fix inorganic carbon (CO2) into organic
the Galapagos Spreading Center carbon. This contrasts with photosynthetic processes, where it
The study of chemosynthetic ecosystems in the deep sea is light energy that is harvested to produce ATP. Subsequent
dates back to a Black and white photograph of large white clam biochemical steps in the production of organic carbon can be
shells lying within cracks in a pavement of Black basalt that was identical in chemoautotrophs and photoautotrophs. Discovery
captured by the Deep-Tow camera system at 13:20:39 (GMT) on of vents thus led us to understand for the first time that com-
29 May 1976 at 0°47‘84‘‘N, 86°09‘18‘‘W (Galapagos Spreading plex food webs could be dependent on microbial chemosyn-
Center, eastern Pacific Ocean; LONSDALE 1977). Given that bi- thetic primary production. The initial dives to deep-sea vents
valves are generally suspension feeders, the diet of the clams was also sparked the hypothesis that life on Earth may have origi-
inferred to have been organic particulates concentrated by bot- nated at submarine hot springs (CORLISS et al. 1980).
tom-water currents (convection cells) induced by hydrothermal The triptych of charismatic vent organisms – clams (Calyp-
activity (LONSDALE 1977; ENRIGHT et al. 1981). togena magnifica), mussels (Bathymodiolus thermophilus), and gi-

C.L. VAN DOVER, M. BISCOITO, A. GEBRUK, J. HASHIMOTO, V. TUNNICLIFFE,


P. TYLER & D. DESBRUYÈRES Denisia 18 (2006) 13–25
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

ant tubeworms (Riftia pachyptila) – has sustained scientific inter- pompejana) and commensal polynoid polychaetes (Branchipolynoe
est since their discovery (BALLARD 1977) and the first field in- symmytilida) is unimpeded across the microplates (HURTADO et
vestigations by biologists (GRASSLE et al. 1979). These organ- al. 2004).
isms, together with “dandelions” (siphonophores: Thermopalia
Samples from American and French cruises to Galapagos,
taraxaca), “spaghetti worms” (enteropneusts: Saxipendium coro-
21°N, and 13°N vent sites between 1979 and 1985 became the
natum), limpets, crabs, amphipods, polychaetes, and other ani-
foundation for a noteworthy series of studies, in which the re-
mals collected from Galapagos vents in the earliest collections
markable anatomy, physiology, and trophic ecology of many
(HESSLER & SMITHEY 1983), established the existence of a fauna
representative species of vent organisms was established (see
endemic to and specialized for life in chemosynthetic ecosys-
JONES & BRIGHT 1985 for an exhaustive bibliography for this
tems. They provided a baseline against which all other vent fau-
period). The important role of endosymbiotic, sulfur-oxidizing
nas would be compared. Our understanding of species zonation,
bacteria in the nutrition of tubeworms (Riftia pachyptila), for ex-
of food webs, and even of succession of megafaunal species at
ample, was established in 1981 through morphological (JONES
vents in the eastern Pacific dates back to reports from these first
1981), ultra-structural (CAVANAUGH et al. 1981), and biochem-
geological and biological expeditions to the Galapagos Spread-
ical studies (FELBECK et al. 1981). The paradox of large animals
ing Center.
living in waters with sulfide concentrations normally toxic to
metazoans was also largely resolved by 1985, with descriptions
Early Investigations on the East Pacific Rise of detoxification mechanisms, including symbiont consumption
of sulfide, sulfide-binding proteins in tubeworms (ARP & CHIL-
Early opportunities for ecological comparisons came with dis-
DRESS 1983; POWELL & SOMERO 1983) and clams (ARP et al.
covery and description of vent fields at 21°N [RISE Program in
1984), and high activities of sulfide-oxidizing enzymes in super-
1979 (Rise Project Group 1980); Oasis Expedition (HESSLER et
ficial cell layers (POWELL & SOMERO 1986).
al. 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- Northeast Pacific Vents
ities between the invertebrate faunas at Galapagos and NEPR
The 1983 exploration of hydrothermal vents at Axial
vent fields indicated that, despite the restricted, insular nature of
Seamount on the Juan de Fuca Ridge (CASM 1985; TUNNI-
the benthic vent communities, exchange between populations
CLIFFE et al. 1985) and subsequent studies of vent faunas on the
on the two ridge axes was sufficient to sustain relatively large
Explorer (TUNNICLIFFE et al. 1986) and Gorda Ridges (VAN
species ranges (Rise Project Group 1980). The NEPR vent fields
DOVER et al. 1990) in the northeast Pacific provided the first
included a habitat – the warm-to-hot (20 to >110 °C), leaky sur-
evidence that the hydrothermal-vent fauna was not globally
faces of Black smoker chimneys – not present in Galapagos vent
cosmopolitan at the species level (CASM 1985). Faunal al-
fields. At least some of the difference between Galapagos and
liances between the EPR and the NE Pacific vent systems are
NEPR species lists derives from the addition of species adapted to
recognized at the level of genus and higher (TUNNICLIFFE 1988),
the warmer waters of the chimney habitat (DESBRUYÈRES &
although several major taxonomic groups commonly found at
LAUBIER 1980, DESBRUYÈRES et al. 1982; FUSTEC et al. 1987), in-
hydrothermal vents on the EPR are so far conspicuously absent
cluding the Pompeii worms (Alvinella pompejana and A. caudata)
(e.g., alvinocarid shrimp, lysianassid amphipods) at NE Pacific
and a brachyuran crab (Cyanagraea praedator). Faunal similarities
vents. Some alliances between the East Pacific Rise and NE Pa-
along the East Pacific Rise are now known to extend from the
cific vent faunas may reflect the paleotectonic history of the
northern limit of the ridge system (Guaymas Basin) to 19°S (JU-
East Pacific Rise and NE Pacific ridge system, which were once
NIPER et al. 1990; BLACK et al. 1994; GEISTDOERFER et al. 1995).
part of a single continuous ridge system before the override of
Vents in Guaymas Basin are exceptional in that they are one of
the North American Plate (TUNNICLIFFE 1988).
the few localities along the mid-ocean ridge system where hy-
drothermal activity and volcanic eruptions take place in associa- Subsequent explorations along the Juan de Fuca Ridge re-
tion with thick layers of pelagic and terrigenous sediment rather vealed vent assemblages in a wide variety of venting conditions:
than bare basalt (EINSELE et al. 1980). The nominal transition of extensive Black smoker fields at Endeavour (e.g., SARRAZIN et
the southern East Pacific Rise to the Pacific Antarctic Ridge oc- al. 1997), sedimented sulfide mounds at Middle Valley (e.g., JU-
curs south of the Easter and Juan Fernandez microplates, i.e., at NIPER et al. 1992), and recent eruptive lavas of the Cleft Seg-
~37°S. This region lies at the boundary between the Indo-Pacif- ment (e.g., TSURAMI & TUNNICLIFFE 2001). From Explorer
ic and Antarctic marine biogeographic provinces (VINOGRADO- Ridge to Gorda Ridge, eight major vent fields provide biologists
VA 1979); the region also appears to be a boundary region for with settings that vary in depth, age, substratum and the rela-
some vent taxa (HURTADO et al. 2004). Bathymodiolid mussels tive importance of volcanism versus tectonism. In recent years,
and bythograeid crabs from 32°S, for example, are sister species studies of community dynamics on sulfide edifices of Juan de
to their northern counterparts (GUINOT & HURTADO 2003; WON Fuca vents have given us a view of temporally and spatially
et al. 2003b), but gene flow in alvinellid polychaetes (Alvinella shifting species populations in response to physical and chemi-

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cal changes in the environment and to biological interactions 1991). Back-arc spreading centers are zones of extension of
(e.g., SARRAZIN & JUNIPER 1999). The proximity of NE Pacific oceanic crust between active and remnant volcanic arcs associ-
vent sites to coastal ports has fostered studies that examine tem- ated with subduction zones. Because back-arc systems tend to
poral change and community comparisons. be short, relatively young, and isolated (i.e., separated by large
distances from the nearly continuous mid-ocean ridge systems),
fauna distinct from that of EPR vents, but evolved to occupy
Discoveries in the Atlantic
similar niches was postulated (HESSLER & LONSDALE 1988).
Discovery of shrimp- (Rimicaris exoculata) dominated vent Dense clusters of large, “hairy gastropods” (Alviniconcha hessleri)
fields at TAG (36°N; RONA et al. 1986) and Snake Pit in 1985 with chemoautotrophic endosymbionts in their gills (STEIN et
(23°N; ODP Leg 106 Scientific Party 1986; MEVEL et al. 1989) al. 1988) proved to be one of the most distinctive species, be-
on the Mid-Atlantic Ridge (MAR) highlighted the fact that longing to a new genus and having trophic attributes not previ-
similar types of organisms might be found at hydrothermal vents ously known within the Gastropoda.
throughout the world’s oceans (e.g., alvinocarid shrimp, bathy-
Although Mariana vent organisms belonged to undescribed
modioliform mussels, siphonostome copepods), but that the
species, more than half of the genera found there were already
species, and even many genera, are different from one ocean
known from vent fields in the eastern Pacific. A mussel in the
basin to another. Major taxonomic groups familiar from EPR
genus Bathymodiolus dominated the biomass; other familiar an-
vents (e.g., vestimentiferan tubeworms, alvinellid polychaetes,
imal types included polychaete worms in the genus Paralvinella
stalked barnacles) are so far unknown at the MAR vents. In
and limpets in the genus Lepetodrilus. Affinities of some Mari-
contrast to the dynamic succession of macrofaunal species ob-
ana taxa were shared with species known from Atlantic hy-
served in some NE Pacific vent habitats (e.g., SARRAZIN & JU-
drothermal vents – for example, shrimp in the genus Chorocaris.
NIPER 1999) and of megafaunal invertebrates at EPR vents (e.g.,
There were no surprises in terms of fidelity to a particular niche
SHANK et al. 1998), communities at MAR vent fields are re-
or microhabitat in genera shared between Mariana and other
markably stable on decadal time scales (COPLEY et al. 1997,
mid-ocean ridge vents; the implication is that speciation has
1999).
largely been passive within these genera, a consequence of iso-
The visual impact of vent megafauna is different between lation by distance and barriers to dispersal, rather than of adap-
MAR and eastern Pacific vents: motile shrimp at MAR vents tive radiations into new niches. The Mariana studies provided
were described by John Edmond as looking “like maggots the first incontestable evidence for faunal interchange and for
swarming on a hunk of rotten meat”; sessile worms and bivalves barriers or filters to dispersal of species between Pacific back-arc
at EPR and NE Pacific vents lend the sites a more garden-like basins and the mid-ocean ridge system (HESSLER & LONSDALE
serenity, as evoked by names like Rose Garden and Garden of 1988). They also provided compelling evidence for the poten-
Eden. There are differences as well in the details of the bacter- tial for discovery of unanticipated taxa (in this case, the hairy
ial-invertebrate symbioses implicated in the nourishment of the gastropods) as new geographic regions are explored.
dominant megafauna between the two ocean basins. Episym-
Subsequent explorations of other southwestern Pacific
bionts likely contribute to the nutrition of rimicarid shrimp on
back-arc systems [Manus (AUZENDE et al. 1997; HASHIMOTO et
the MAR (VAN DOVER et al. 1988; GAL’CHENKO 1989; GEBRUK
al. 1999); North Fiji and Lau Basins (HASHIMOTO et al. 1989;
et al. 1992; SEGONZAC et al. 1993), whereas tubeworms, clams,
DESBRUYÈRES et al. 1994); New Ireland Basin (HERZIG et al.
and mussels that dominate EPR vents rely on endosymbionts. 1994)] demonstrated that some species have restricted distribu-
MAR mussels have dual endosymbionts (methanotrophs and tions while other species, such as provannid gastropods, are
thiotrophs), while EPR mussels have only thiotrophic en- shared among basins. Molecular techniques have now distin-
dosymbionts in their gills (FIALA-MEDIONI 1984; LE PENNEC & guished four species of alviniconchid snails: Alviniconcha hessleri
HILY 1984). Discovery of shrimp swarms at Mid-Atlantic Ridge at Mariana vents, two Alviniconcha species in the Manus and
vents ultimately led to the description of modified eyes in Rim- North Fiji Basins, and a fourth Alviniconcha species at Indian
icaris exoculata and other alvinocarid shrimp, eyes that are Ocean vents (KOJIMA et al. 2003; OKUTANI et al. 2004). In con-
adapted for detecting dim sources of light (VAN DOVER et al. trast to the basin-scale differentiation of alviniconchid gas-
1989; WHITE et al. 2002). tropods in the southwestern Pacific, Ifremeria nautilei (synony-
mous with Olgaconcha tufari) so far is only known to inhabit
Western Pacific Explorations vent sites in the Manus, North Fiji, and Lau Basin region (DES-
BRUYÈRES et al. 1994; HASHIMOTO et al. 1999). The relatively
Hydrothermal vents in Manus Basin were discovered in narrow distributional range of Ifremeria nautilei might be attrib-
1985 (BOTH et al. 1986), the same year that vents on the Mid- uted its lower dispersal ability compared to that of Alviniconcha
Atlantic Ridge were reported (RONA et al. 1986). Further at- species (KOJIMA et al. 2000, 2001). Ifremeria nautilei is also the
tention was brought to the southwestern Pacific in 1987 with only known gastropod that supports dual symbioses, with
the first description of vent faunas from the Mariana back-arc methane- and sulfur-oxidizing bacteria in gill bacteriocytes
spreading center (HESSLER et al. 1988, HESSLER & LONSDALE (GAL’CHENKO et al. 1992).

15
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Japanese biological explorations of deep-sea hydrothermal Hydrothermal vents also occur in fresh-water lakes. Fresh-
systems in the northwestern Pacific began in 1988 with the dis- water hydrothermal systems, similar to those in the ocean, are
covery of active hydrothermal vents at Izena Hole (Okinawa related either to rift zones, like lakes Baikal (CRANE et al. 1991)
Trough; TANAKA et al. 1990) by scientists using Shinkai 2000. and Tanganyika (TIERCELIN et al. 1993), or volcanic activity in
Subsequent work discovered venting in yet another setting on subduction zones, e.g. Crater Lake in Oregon (Anonymous
the arc volcanos of the Izu-Bonin seamount chain south of 1989). No examples of fresh-water, vent-specific metazoa are
Japan (KOJIMA 2002). Recent exploration of the Mariana Arc known. Bacterial mats are common at all known fresh-water
reveals variability in venting conditions and associated com- vents. Mineral structures analogous to Black smoker chimneys,
munities (EMBLEY et al. 2004). Together, the southwestern and reaching up to 10 m in height, can also develop (e.g., in Crater
northwestern Pacific hydrothermal fields represent a fertile re- Lake). The Russian-operated Pisces submersible studies in Lake
gion for the study of population genetics and the biogeography Baikal documented aggregations of non-vent fauna in associa-
and evolution of vent taxa (DESBRUYÈRES et al. 2006). tion with bacterial mats; carbon derived from bacterial
methane oxidation is incorporated into local food web of the
benthic fauna (GEBRUK et al. 1993).
Discovery of Seeps and Other
Chemosynthetic Settings
Eruptions and the Hydrothermal Cycle
The 1980s included other milestones: Seep communities
discovered first in 1984 at the base of the Florida Escarpment in 1991 was the year of the eruption at 9°50’N on the East Pa-
the Gulf of Mexico (PAULL et al. 1984; HECKER 1985) and in cific Rise (HAYMON et al. 1993) and the beginning of a long-
Sagami Bay (OKUTANI & EGAWA 1985; HASHIMOTO et al. term study of hydrothermal cycles in this region (SHANK et al.
1989), and then in association with subduction zone settings off 1998). The bloom of bacteria that marked the commencement
Oregon (SUESS et al. 1985), in the northwestern Pacific of the cycle (NELSON et al. 1991; HAYMON et al. 1993) and oth-
(LAUBIER et al. 1986; OHTA & LAUBIER 1987), and off Barbados er biological indications of recent volcanic activity were subse-
(FAUGERES et al. 1987), taught us that vent-like taxa and troph- quently observed at other locales, including the NE Pacific
ically complex, chemosynthetically based ecosystems are not Ridges (e.g., TUNNICLIFFE et al. 1997; JUNIPER et al. 1998) and at
restricted to hydrothermal settings. Discovery of chemosyn- 17°S on the East Pacific Rise (EMBLEY et al. 1998). Repeated vis-
thetic ecosystems and of vent- and seep-like taxa associated its to developing vent communities documented the rapidity
with whale skeletons on the seafloor (SMITH et al. 1989; re- with which colonization of new sites of venting takes place (e.g.,
viewed in SMITH & BACO 2003) underscored the need to adopt well-established colonies of tubeworms within one year; TUNNI-
a broad view of chemosynthetic faunas. Vent taxa were also dis- CLIFFE et al. 1997; SHANK et al. 1998) and confirmed earlier re-
covered in diverse volcanic and hydrothermal settings, includ- ports of rapid growth rates in vent species (LUTZ et al. 1994).
ing alvinocarid shrimp at a mid-plate, hot-spot volcano (Loihi Careful re-sampling of evolving vents allows insights into troph-
seamount; KARL et al. 1989; WILLIAMS & DOBBS 1995), alvinel- ic strategies and the role of biological interactions such as com-
lids, mussels, crabs, and alvinocarid shrimp at Pito Seamount petition (LEVESQUE et al. 2003).
(NAAR et al. 2004), and relatively shallow (489 m) populations
The 9°50N vent field became a favored field site for eco-
of vesicomyid clams at Piips volcano (SELIVERSTOV et al. 1986;
logical studies for several reasons: (1) the eruption took place
SAGALEVITCH et al. 1992). Thus we learn that vent and vent-
there, (2) its relatively simple topography has been thoroughly
like taxa and trophically complex, chemosynthetically based
mapped, (3) its vents support a variety of foundation species
ecosystems are not restricted to mid-ocean ridge hydrothermal
with different habitat preferences (e.g., tubeworm clumps, mus-
settings, and that an understanding of the evolution and bio-
sel beds, alvinellid galleries, serpulid and anemone fields), (4)
geography of vent taxa can only be understood in the broader
there is a long record of contemporaneous and complementary
context of the evolution and biogeography of allied taxa at oth-
geological and geochemical studies, and (5) the transit time
er reducing environments (SIBUET & OLU 1998; VAN DOVER et
from major ports in Mexico is relatively short. The 15+ years of
al. 2002; TUNNICLIFFE et al. 2003a).
research at 9°50‘N since the eruption have resulted in impor-
Other shallow hydrothermal settings were found to lack en- tant new insights into successional sequences (SHANK et al.
demic invertebrate species and do not support invertebrates 1998), physiological adaptations to the vent environment (e.g.,
with endosymbiotic chemoautotrophic bacteria [e.g., subtidal GOFFREDI et al. 1997; SHILLITO et al. 2001), reproductive at-
vents off the coast of Southern California (KLEINSCHMIDT & tributes of vent invertebrates (e.g., MARSH et al. 2000; PRADIL-
TSCHAUDER 1985); vents of the volcanic arc in the Mediter- LON et al. 2001), biological interactions among species (e.g.,
ranean (reviewed in DANDO et al. 1999); vents on the Kolbein- MULLINEAUX et al. 2000; MICHELI et al. 2002; MULLINEAUX et
sy (Olafsson et al. 1989) and Jan Mayen ridges (FRICKE et al. al. 2003), animal-chemical relationships (e.g., LUTHER et al.
1989) north of Iceland; vents in the crater of Ushishir volcano 2001; LE BRIS et al. 2006), population genetics (reviewed in
of the Kamchatka region (TARASOV et al. 1990, TARASOV VRIJENHOEK 1997), and community structure (e.g., VAN DOVER
2006)]. 2003; GOVENAR et al. 2004).

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One of the longest records of time-series studies at vents is meager [except for dense shrimp Rimicaris exoculata populations
that for 13°N on the East Pacific Rise, where French and Amer- at Rainbow], possibly due to the distinctive fluid chemistries as-
ican scientists have made intermittent observations since 1982 sociated with these sites (KELLEY et al. 2001).
(reviewed by DESBRUYÈRES 1995). At this site, a previously
“dead” vent area with relic populations resumed activity in
The Indian Ocean
1987, and followed the successional pattern described for 9°N by
SHANK et al. (1998). The presumption is that there was a sub- By 2000, a basic understanding of vent faunas of the north-
surface dyking event (i.e., an upward migration of molten rock ern Mid-Atlantic Ridge, the East Pacific Rise, the Northeast
through a fissure, but without overflow onto the seabed) in 1987 Pacific, and back-arc basins in the western Pacific was estab-
that delivered heat to the system and restored the hydrothermal lished. One of the biggest missing pieces of the biogeographic
circulation at this site. jigsaw puzzle was the ridge system in the Indian Ocean. Cyana
dives in the Gulf of Aden in 1984 (1400-1600 m) had docu-
mented low-temperature flow dominated by shrimp, anemones,
More Atlantic Discoveries
and galatheid squat lobsters, but specimens were not collected
Attention returned to the Mid-Atlantic Ridge in 1992 with (JUNIPER et al. 1990). Japanese scientists brought attention to
the chance collection of vent animals in a rock-dredge haul, and the Indian Ocean in 2000, using the ROV Kaiko to explore the
in 1993 during dives at the site that came to be known as Lucky Kairei hydrothermal field on the Central Indian Ridge, just
Strike at 37°N (VAN DOVER et al. 1996; LANGMUIR et al. 1997). north of the Rodriguez Triple Junction (HASHIMOTO et al.
Although Lucky Strike shares a few species with the TAG and 2001). Shrimp closely related to Rimicaris exoculata of Atlantic
Snake Pit vent sites, most species at Lucky Strike were new to vents dominate the biomass of the Kairei field so that the site
science. This site and the nearby Menez Gwen vents explored in resembles TAG and Snake Pit, but other taxa, including hairy
1993 (FOUQUET et al. 1995; DESBRUYÈRES et al. 2001) appear to gastropods and mussels, are more closely allied with Pacific vent
belong to a biogeographic province different from that of the faunas. These observations supported the hypothesis that the
more southerly TAG and Snake Pit faunas. Variables associated Indian Ocean ridges serve as a link between Atlantic and Pa-
with the depth differential between the two provinces (<1700 m cific vent faunas (HASHIMOTO et al. 2001). Further explorations
at Lucky Strike and Menez Gwen vs. >3000 m at TAG, Snake of the Kairei Field (VAN DOVER et al. 2001) led to the discov-
Pit) have been implicated in the biogeographic differentiation ery of a remarkable scaly-footed gastropod with no close al-
of the two vent faunas (VAN DOVER et al. 1996; DESBRUYÈRES et liance to any known Pacific or Atlantic vent genera (Waren et
al. 2000, 2001). al. 2003). Unlike other mollusks at vents that house autotroph-
Continuing geological and geochemical exploration for hy- ic endosymbionts in epithelial bacteriocytes of their gills, the
drothermal activity on the Mid-Atlantic Ridge between the Ve- scaly-footed gastropod hosts its symbionts in a hypertrophied
ma transform fault at 11°N and the Azores hotspot (38°N) re- esophageal gland (GOFFREDI et al. 2004).
sulted in the first submersible dives to hydrothermal sites at Lo-
gatchev in 1993 (BOGDANOV et al. 1995; GEBRUK et al. 2000), Fossil Records
Broken Spur in 1993 (MURTON et al. 1995), and Rainbow in
1997 (FOUQUET et al. 1997). The Logatchev site at 14°45’N was The first accounts of fossil vent invertebrates comes from
the first vent discovery where the host rock was ultramafic (i.e., studies of Cretaceous Oman and Cyprus ophiolites that yielded
of mantle rather than crustal origin; BOGDANOV et al. 1997; well-preserved tubes of worms, including what might be a ves-
FOUQUET et al. 2006). Logatchev represents the current south- timentiferan (HAYMON et al. 1984; OUDIN & CONSTANTINOU
ern limit of published information on vent faunas along the 1984; HAYMON & KOSKI 1985). Invertebrates have occupied
Mid-Atlantic Ridge, although a new active vent field known as vents (KUZNETSOV 1993; LITTLE et al. 1997) and seeps (BARBI-
Ashadze at 12°38’N has been reported (BELTENEV et al. 2004), as ERI et al. 2004) at least since the Silurian, but the ancient (old-
have vent fields south of the equator (C. German, pers. comm). er than the early Cretaceous) assemblages described so far were
Logatchev is notable as the only known site on the Mid-At- dominated by now-extinct families of brachiopods, monopla-
lantic Ridge that supports large populations of vesicomyid clams; cophorans, bivalves, and gastropods (reviewed in CAMPBELL
in other aspects, it has a species list similar to that found at 2006). There is little evidence to support the view that vents
Snake Pit (e.g., TURNIPSEED et al. 2004). Broken Spur (29°10’N) (or seeps) have served as a refuge during global extinction
is a hybrid zone for mussels, where two species of mussels (Ba- events for most taxa (reviewed in WARÉN & BOUCHET 2001;
thymodiolus puteoserpentis, dominant at southern locales, and B. LITTLE & VRIJENHOEK 2003), although there may be exceptions
azoricus, known from the northern vent sites) co-occur and in- (YAMAGUCHI et al. 2004). As in extant chemosynthetic com-
ter-breed (WON et al. 2003a). Rainbow (FOUQUET et al. 1997), munities, fossil records include indications of successional re-
Saldanha (BARRIGA et al. 1998), and Lost City (KELLEY et al. sponses to changing conditions at a given site (CAMPBELL et al.
2001) vent fields, like Logatchev, are hosted by ultramafic rocks. 2002), and paleobiogeographic patterns that are likely to reflect
Their megafaunal and macrofaunal communities are generally past plate tectonic configurations (CAMPBELL 2006).

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© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Prospects for Exploration and Discovery Life) was conceived to promote international efforts to improve
our understanding of the identity and biogeography of the fau-
There remain a number of oceanic regions where we can
na of these reducing environments (TYLER et al. 2003). ChEss
anticipate discovery of entire communities of new species at hy-
also promotes tests of hypotheses regarding potential barriers
drothermal vents (VAN DOVER et al. 2002; TYLER et al. 2003).
and filters to dispersal of species whose adults are restricted to
Of these, the Arctic Ocean has long been of great interest: its
these environments. Through ChEss and other ocean explo-
relative youth and degree of bathymetric isolation from the rest ration activities, we look forward to the next decade of discov-
of the deep waters of the world’s oceans suggest that the vent ery that will undoubtedly add many new species, genera, and
fauna of the region might provide insight into the invasibility families to the compendium of taxa so carefully compiled here.
of vent environments by metazoans. Hydrothermal vents on the
Gakkel Ridge in the Arctic Ocean have been localized (ED-
MONDS et al. 2003), but their faunas remain to be photo-docu- Acknowledgements
mented and sampled. Shells of vesicomyid clams (Archivesica We thank our many shipboard colleagues – Captains, pilots,
sp.), indicative of reducing environments, occur in dredge sam- ships’ crews, scientists, technicians, and students – who have
ples from depths of ~2000 m near the Gakkel Ridge and seem contributed to the discovery of deep-sea hot springs and of the
likely to have been associated with a hydrothermal site when strange animals that live at them. There are too many individ-
the animals were alive (SIRENKO et al. 1995). uals to cite by name; we honor their efforts with images of some
Other sites of considerable biogeographic and taxonomic of the many vehicles that have given us such remarkable access
interest are isolated ridge systems, such as the Scotia Ridge in to the seafloor (Fig. 1, 2).
the southwest Atlantic, the Cayman Rise in the Caribbean, and
the Andaman back-arc ridge in the northeast Indian Ocean. Bibliography
Even on the mid-ocean ridge system, there remain large stretch-
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segments is the Mid-Atlantic Ridge north of the Azores – here cellent reviews of the biology of vent and other chemosynthet-
the ridge axis dives from shallowest depths back down to more ic communities that provide entrée into the primary literature
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VAN DOVER C.L. (2003): Variation in community structure within hy-


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VAN DOVER C.L., GRASSLE J.F. & M. BOUDRIAS (1990): Hydrothermal vent fau-
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Gorda Ridge: A Seafloor Spreading Center in the United States Ex-
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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
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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
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VAN DOVER C.L., DESBRUYÈRES D., SEGONZAC M., COMTET T., SALDANHA L., FIALA-
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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.

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drothermal plumes in the Galapagos Rift. — Nature 267: 600-603.

23
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

A B

C D

E F
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 Hawaii
Undersea Research Laboratory, U.S.A.; F: Shinkai 6500 (6500 m), operated by JAMSTEC, Japan.

24
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

A B

C D
Fig. 2: A selection of remotely-operated vehicles (and
maximum depth rating) used in deep-ocean exploration.
A: ROPOS (5000 m), operated by the Canadian Scientific
Submersible 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 the
Woods Hole Oceanographic Institution, U.S.A.; E: Kaiko
(11000 m), operated by JAMSTEC, Japan.

E F

25
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Hydrothermal vent meiofauna


Meiofauna, small animals and protists, which pass through known. The few ostracod species described exclusively come
a net with 1 mm mesh size and are retained on a net with 63 from the eastern Pacific (DEEVEY 1969; KORNICKER 1969, 1991,
µm (or smaller) mesh size (GIERE 1993), is part of the hy- KORNICKER & HARRISON-NELSON 2005; MADDOCKS 2006). The
drothermal vent community. In general, very little is known few tanaidaceans and the two isopod species were collected at
about the diversity of vent meiofauna. So far, only a few phyla the site Lucky Strike at the Mid-Atlantic Ridge (CUNHA &
such as the Arthropoda (Acariformes, Copepoda, Cumacea, WILSON, in press; LARSEN et al., in press); tanaidaceans were al-
Isopoda, Ostracoda, Tanaidacea), the foraminiferan Granu- so observed at the site Rainbow (M. Segonzac, pers. comm.)
loreticulosa, and the Nematoda have been described from Only one mite is described from Lau and North Fiji Back-Arc
vents. In addition, Plathelminthes and three protist phyla in- Basins (KRANTZ 1982) and the second species comes from the
cluding amoebae, colonial chrysophytes and ciliates are known Mid-Atlantic Ridge (BARTSCH 1990). Similarily, one cumacean
but have yet to be described (SMALL & GROSS 1985; M. B., and one foraminiferan species each are decribed from the East
pers. obs.). In contrast, other marine habitats, such as sedi- Pacific Rise (BRÖNNIMANN et al. 1998; CORBERA this volume),
ments from shallow waters to the deep sea harbor far more di- the second species each come from the Mid-Atlantic Ridge
verse meiofauna. For example gnathostomulids, gastrotriches, (KAMENSKAYA et al. 2002; CORBERA this volume). The few des-
loriciferans, tardigrades, and small-sized representatives of hy- cribed free-living nematode species exclusively were collected at
droids, bryozoans, nemertines, rotifers, or gastropods have not sedimented vent sites from the East Pacific Rise, the Guaymas
yet been discovered (HIGGINS & THIEL 1988; GIERE 1993). Basin, and the Lau and the North Fiji Back-Arc Basins (DE-
CRAEMER & GOURBAULT 1998; VERSCHELDE et al. 1998).
All together, two foraminiferan, seven nematode, nine ost-
racod, two acari, two cumacean, six tanaidacean, two isopod, To date there is not a single site for which the entire meio-
and 78 copepod species are included in this edition. Only the fauna community structure is known. The picture emerging
Copepoda are known to a greater extent (IVANENKO & DEFAYE, from the few meiofauna studies points to a vent meiofauna,
this volume). Currently, meiofauna species contribute to about which is low in abundance and diversity (DINET et al. 1988;
20% of the total diversity at vents. However, at large, the diver- SHIRAYAMA 1992; VANREUSEL et al. 1997; TSURUMI et al. 2003).
sity of meiofauna is unknown. There are entire mid-ocean rid- Comprehensive studies are urgently needed not only to under-
ges or back-arc basins known for a relatively long time, in which stand the community structure of meiofauna but also to provi-
not a single species of ostracod, tanaidacean, mite, or nematode de insight in the biodiversity of the entire vent community and
is described. Most known species are reported from a single site. their trophic interactions.
The biogeographic range of meiofauna species is virtually un-

1, 2: Folliculinid ciliates from the East Pacific Rise: 9°N; by M. Bright.

M. BRIGHT Denisia 18 (2006): 27–28


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To our knowledge, vent meiofauna can be found in virtual- As there is no single technique available for extracting and
ly all vent habitats, from inactive bare basalt or sediments to fixing the entire meiofauna community quantitatively, the usu-
diffuse flow areas up to Alvinella pompejana DESBRUYÈRES & ally applied compromise is either to sort through the entire
LAUBIER, 1980 aggregations located on black smokers. Many sample including the sediment or to use various centrifugation
vent meiofauna animals such as copepods, nematodes, and ost- techniques in order to separate organic from inorganic materi-
racods survive the transit from the vent environment at depths al. Fixation then is done by using 4% buffered formalin. Howe-
of more than 2500 m to the water surface and can be maintai- ver, it has to be kept in mind that mostly the more robust so-
ned without pressure at 4°C for at least a few days (M. B., pers. called “hard” meiofauna such as copepods and nematodes is
obs.). extracted and fixed and “soft” meiofauna such as plathelmin-
thes or gastrotrichs, which is often not separated from the sedi-
Quantitative sampling of meiofauna is usually carried out in
ment grains and/or is more delicate and sometimes requires ot-
soft sediments with various types of corers. Hard substrate sam-
her fixation media, is lost. For storage, 70-80% ethanol is re-
pling and as well as sampling of megafauna aggregations such as
commended (see HIGGINS & THIEL 1988).
tubeworms clumps or mussel beds requires some sort of special
designed devices such as the “mussel pot” (VAN DOVER 2002)
or the “Bushmaster Jr” (GOVENAR et al. 2005). Abundance and
biomass are standardized to 10 cm-2 surface area.

References:
BARTSCH I. (1990) Bull. Mus. Natl Hist. Nat., Paris, 4è sér., A. 12: 69-73.
BRÖNNIMANN P., VAN DOVER C. & J.E. WHITTAKER (1989) Micropaleontology 35: 142-149.
CUNHA M.R. & G.D.F. WILSON (in press) Zootaxa.
DECRAEMER W. & N. GOURBAULT (1997) Zool. Scr. 26: 1-12.
DEEVEY G.B. (1968) Proc. Biol. Soc. Wash. 81: 539-570.
DINET A., GRASSLE F. & V. TUNNICLIFFE (1988) Oceanol. Acta 85: 7-14.
GIERE O. (1993) Meiobenthology, the Microscopic Fauna in Aquatic Sediments. Springer Verlag: Berlin: 1-328.
GOVENAR B., LE BRIS N., GOLLNER S., GLANVILLE J., APERGHIS A.B., HOURDEZ S. & C.R. FISHER (2005) Mar. Ecol. Prog. Ser. 305: 67-77.
HIGGINS R.P. & H. THIEL (1988) Introduction to the Study of Meiofauna. Smithsonian Institution Press. Washington D.C., London: 1-488.
KAMENSKAYA O.E., BAGIROV N.E. & T.G. SIMDIANOV (2002) Adaptation Aspects of Evolution of Marine Fauna. Collected Proc. Moscow VNIRO Publ. House:
144-152 [in Russian].
KORNICKER L.S. & E. HARRISON-NELSON (2005) Zootaxa 1071: 19-38.
KRANTZ G.W. (1982) Can. J. Zool. 6: 1728-1731.
LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.
MADDOCKS R.F. (2006) Micropaleontology 51: 345-372.
SHIRAYAMA Y. (1992) Proc. JAMSTEC Symp. Deep Sea Res.: 287-290.
SMALL E.B. & M.E. GROSS (1985) Bull. Biol. Soc. Wash. 6: 401-410.
TSURUMI M., DE GRAAF R. C. & V.TUNNICLIFFE (2003) J. Mar. Biol. Ass. U. K. 83: 469-477.
VAN DOVER C.L. (2002) Mar. Ecol. Prog. Ser. 230: 137-158.
VANREUSEL A., VAN DE BORSCHES I. & F. THIERMANN (1997) Mar. Ecol. Prog. Ser. 157: 207-219.
VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.

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Hydrothermal vent parasites


Currently, very few macroparasites are known from hydro- In brief, most often parasites must carefully looked for in or-
thermal deep-sea vents. Only four copepods, Genesis vulcanoc- der to be noticed. Hosts are various and diverse and range from
topusi (East Pacific Rise, LOPEZ-GONZALEZ et al. 2000), Cholidya the smallest of copepods to the largest of fish.
polypi (Endeavour Segment, HUMES & VOIGHT 1997), Ceuthoe-
Since numerous parasites have a complex life cycle invol-
cetes aliger and Rimitantalus hirsutus (Galapagos Spreading Cen-
ving more than one host, a better knowledge of the parasite
ter, HUMES & DOJIRI 1980), one leech Bathybdella sawyeri (East vent fauna would help researchers to understand the trophic
Pacific Rise, BURRESON 1981; BURRESON & SEGONZAC, in structure present at each vent. Because trophically transmitted
press), one acanthocephalan Hypoechinorhynchus thermaceri parasites are in many cases known to alter intermediate host
(East Pacific Rise, BURON 1988), and one nematode, Moravec- behavior to increase capture by definitive hosts (LAFFERTY &
nema segonzaci (Mid-Atlantic Ridge, JUSTINE et al. 2002) have MORRIS 1996) these organisms may play significant roles in in-
been reported from these sites. Significantly, each of these spe- creasing or directing the flow of energy into higher trophic le-
cies was new to science. At least one monogenean and two di- vels at the vents. For example, acanthocephalans at the East
geneans are currently being described by various specialists Pacific Rise use fish as definitive hosts and appear to be trans-
from material collected on crabs and fishes collected from sou- mitted via amphipod ingestion (I. B., pers. obs.): evidence from
thern East Pacific Rise sites and each of these also appear to be other systems indicates that such infected amphipods exhibit
previously unreported species. altered behavior (BAUER et al. 2005). Significantly, at the At-
Because the concentration of potential hosts in a vent eco- lantic sites these crustaceans were found to contain nematode
system could possibly increase parasite transmission and diversi- larvae of the ascarid genus Hysterothylacium although the adults
ty, BURON & MORAND (2004) hypothesized that the low diversi- were not found (A. Petter, unpublished data).
ty of macroparasites presently reported from these sites is likely a Furthermore, non-trophically transmitted parasites present
reflection of these organisms having been overlooked by vent at vents may also pose interesting problems for researchers. For
biologists due to their inconspicuous nature. In support of this instance, the leech Batybdella sawyeri, living among Riftia pachyp-
idea, the use of molecular tools has shown the presence of a high tila and mussel Bathymodiolus thermophilus at the Galapagos Spre-
diversity of parasitic protists genetically linked to apicomplexans, ading Center, raises an interesting specificity problem: although
perkinsozoans, syndiniales, and kinetoplastids (MOREIRA & LO- a parasite of fish, it has never been found on such hosts but was
PEZ-GARCIA 2003). Additionally, Rickettsia-like inclusions have observed on vent crabs and bivalves from the southern East Pa-
also been found in numerous limpets from various vents (TERLIZ- cific Rise (BURRESON & SEGONZAC, in press). Because of the well
ZI et al. 2004), and in mussels from Snake Pit and Logatchev si- known specificity of many parasite-host relationships the recog-
tes (WARD et al. 2004) are further indications that parasite di- nition and identification of vent parasites may play an important
versity is vastly underreported from these important sites. role in deciphering vent colonization. In this regard it is impor-

1: Hirudinea Bathybdella sawyeri from East Pacific Rise: 17°S; cruise 2: Nemertea Carcinonemertes sp., semi-parasite on
Biospeedo (Ifremer, CNRS). bythograeid crabs from Lau Basin; cuise TUIMO6MV
(MBARI); by G. Rouse.

I. DE BURON & M. SEGONZAC Denisia 18 (2006): 29–30


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

tant to note the presence of carcinonemertids (nemertean In summary, the study of parasitism in isolated vent com-
worms) at vents. These worms are known to feed on the eggs of munities should be emphasized not just because it would furt-
shallow water decapods and were discovered on several vent crab her increase our understanding of the unique parasite fauna
species (SHIELDS 2001). Given the depth (1800-2400 m) and iso- present at these sites, but also because it would allow us to ad-
lated nature of vent communities, the origin of vent colonization dress and answer important questions regarding the coloniza-
by carcinonemertids has been raised and it has been speculated tion of vents and vent community structure as well as allow us
that infections were acquired from a more shallow-water host to identify and recognize new strategies for parasite survival.
(see SHIELDS 2001 for review). Such egg predators were recently Parasites may be located almost anywhere on or in a host.
collected on several bythograeid crab species from western back- Ideally, parasites should be isolated from their host while still
arc basins, and northern and southern East Pacific Rise (J. fresh and then prepared in a specific manner dependent on the
Shields & M. Segonzac, unpublished data). Moreover, because of taxon collected (see, for example, PRITCHARD & KRUZE 1982
the isolated nature of the vent ecosystems these sites will likely for macroparasite fixation techniques). Importantly, parasites
provide excellent sources of material for the study of the proces- collected from formaldehyde fixed hosts, such as museum spe-
ses of co-evolution in a relatively simple trophic system. cimens, are most often of no use to taxonomic study.

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.

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Granuloreticulosa, Foraminifera

Members of the order Foraminifera are widely distributed in ments. Some foraminiferal species that have been collected in
marine settings and can be found from polar regions to tropical vent regions also belong to planktonic species whose empty
latitudes and from shallow water environments to the abyssal tests are accumulating on the sea floor. So far, only two end-
plains of the deep sea. More than 300 benthic foraminiferal ta- emic foraminiferal species have been described from vent re-
xa have been reported from deep-sea hydrothermal vent re- gions, which are listed in the present work.
gions, but most of them are not restricted to these environ-

1: Abyssotherma pacifica from East Pacific Rise: 9°N, Riftia Field; by M. Bright.

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


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Granuloreticulosa, Foraminifera, Allogromida, Arboramminidae

Luffammina atlantica KAMENSKAYA, BAGIROV & SIMDIANOV, 2002

close to the genus Arborammina but differs from it by weak


Size: Up to 2 cm in length.
branching of the upper test, absence of bead-like structures on
Morphology: Agglutinated foraminifer with soft cylindrical the test and absence of globogerins inside the test.
tests are attached to the substrate by extended basal part. Up-
Biology: Found on the surface of relict hydrothermal chimneys.
per extended part of the test may be dichotomously divided.
Test wall consists of fine detritus and agglutinated coccolithes. Distribution: Mid-Atlantic Ridge, site Rainbow.
Surface of the test is covered by a thick layer of Methanotrix-like
bacteria. Inner volume of the test with fine agglutinated parti-
cles, diffuse cytoplasm and cavities. The genus Luffammina is

1: Test; scale bar 1 mm; 2: Test surface covered with Methanotrix-like bacteria; scale
by O. Kamenskaya. bar 100 µm; by O. Kamenskaya.

3: Specimens (arrows) on hydrothermal chimneys; 4: Specimens (arrows) on hydrothermal chimneys;


by O. Kamenskaya. by O. Kamenskaya.

Reference:
KAMENSKAYA O.E., BAGIROV N.E. & T.G. SIMDIANOV (2002) in Adaptation Aspects of Evolution of Marine Fauna. Coll. Proc., Moscow VNIRO Publ. House:
144-152 [in Russian].

M. HOLZMANN Denisia 18 (2006): 32

32
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Granuloreticulosa, Foraminifera, Textulariida, Remaneicidae

Abyssotherma pacifica BRÖNNIMANN, VAN DOVER & WHITTAKER, 1989

Size: Maximum and minimum test diameter, 940 and 830 µm, Biology: According to BRÖNNIMANN et al. (1989), A. pacifica
respectively (holotype). occurs in artificial recruitment arrays placed in the vicinity of
deep-sea (2600 m) hydrothermal springs in the East Pacific.
Morphology: Test free, a low watchglass-shaped trochospire.
Water temperature immediately surrounding the arrays was
Adult chambers spirally elongate, umbilically asymmetric and
near ambient. Abyssotherma pacifica occurs also associated with
mushroom-shaped; interior subdivided by secondary septa
Riftia pachyptila and on bare basalt, suggesting a wide range of
formed by infolding of inner organic sheet. Aperture double:
temperature and sulfide tolerance (pers. comm. Bright).
primary opening interiomarginal, in strongly incurved anterior
flank of septum, anteriorly directed; secondary opening in axi- Distribution: East Pacific Rise: 21°N (BRÖNNIMANN et al.
al-sutural position at tip of posterior flank of septum, posterior- 1989) and 9°N (pers. comm. Bright).
ly directed, also interiomarginal. Wall imperforate, consisting
of agglutinated layer between outer and inner organic sheets.

1: Umbilical view x 80 2: Spiral view x 80 3: Close-up of agglutinated test, spi-


© Micropaleontology. © Micropaleontology. ral view x 125 © Micropaleontology.

References:
BRÖNNIMANN P., VAN DOVER C. & J.E. WHITTAKER (1989) Micropaleontol. 35: 142-149.
LEE J.J., ANDERSON O.R., KARIM B. & J. BERI (1991) Micropaleontol. 37: 303-312.
JONASSON K.E. & C.J. SCHRÖDER-ADAMS (1996) J. Foraminiferal Res. 26: 137-149.
KAMINSKI M.E. (2000) Proc. 5th Intern. Workshop on agglutinated Foraminifera. Grzybowski Foundation Special Publication 7: 185-219.
MICHELI F., PETTERSON C.H., MULLINEAUX L.S., FISHER C.R., MILLS S.W., SANCHO G., JOHNSON G.A. & H.S. LENIHAN (2002) Ecol. Monogr. 72(3): 365-382.
MULLINEAUX L.S., PETTERSON C.H., MICHELI F.& S.W. MILLS (2003) Ecol. Monogr. 73(4): 523-542.

M. HOLZMANN Denisia 18 (2006): 33


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Porifera

Deep-sea representatives of the Porifera (sponges) belong Both hexactinellids and cladorhizids are not members of
mainly to the classes Hexactinellida and to a special family of the true vent communities. Up to now, they have never been
Demospongiae, class Poecilosclerida, the Cladorhizidae, al- observed in the immediate environment of active smokers.
though many other Demospongiae are able to live in bathyal However, a few hexactinellids and numerous cladorhizids occur
environments. Hexactinellida and Cladorhizidae display two at few distance from the active vents in high diversity, and at a
very different life strategies allowing them to survive in deep- high proportion of undescribed species in the case of
sea conditions where life is difficult for sponges, which are fil- cladorhizids (VACELET, in press). These carnivorous sponges
ter-feeders living on tiny particles. The hexactinellids have a benefit, at least in part, from a general distant enrichment
highly developed filter-feeding system with large cavities lined around the vents At the Lau sites, cladorhizid specimens caught
by extremely thin living tissue to maximize the volume of fil- tens of larval zoe of crab Austinograea alaysae (M. Segonzac,
tered water and the ability for retention of particles. Converse- pers. obs.). It cannot be excluded, however, that they belong to
ly, the Cladorhizidae have lost their filter-feeding mode and re-
a “non-vent” fauna taking advantage only of the presence of
ly on carnivory of relatively large prey, mostly crustaceans, a
hard substrata offered by the lava issued at rapidly spreading ar-
very unusual and unexpected mode of feeding in sponges. This
eas and that their high diversity is due to a higher sampling ef-
strategy allows cladorhizids to reach the hadal zone with a
fort in these areas.
depth record of 8840 m, whereas the hexactinellids and “nor-
mal” demosponges do not exceed 7000 m.

1: Abyssocladia sp. from Kilo Moana, Lau Back-Arc Basin, TUIM 05 cruise © M. Tivey.

J. VACELET Denisia 18 (2006): 35


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The cladorhizids are generally small sponges with a mor- These deep-sea sponges are highly fragile. The hexactinel-
phology rather unusual in sponges. They are erect, with a pin- lids, although supported by a highly developed siliceous skele-
nate or symmetrical shape, and their aquiferous system, long ton, have very thin, partly syncytial living tissue which is very
considered as a diagnostic characteristic of the phylum Porifera, difficult to preserve properly. They must be preserved in good
has been discarded in most genera. A special mention, howev- histological or cytological fixatives as soon as possible after col-
er, should be made for the genus Chondrocladia, in which a lection. The carnivorous sponges have very fragile appendages
canal system and choanocyte chambers are maintained, but that are generally only partially preserved during collection.
serve to inflate large spheres on which the prey is captured. The Observation of their morphology from manned submersibles or
prey, mostly small crustaceans, are passively captured by means ROVs is important. For instance, a long time was needed to re-
of hook-like cheloid microscleres lining the surface of the alize that a mysterious organism with pedunculate, translucent
sponge and catching the setae or appendages of various inver- spheres actually corresponds to the genus Chondrocladia as it
tebrates, acting as a Velcro cover. In the absence of a digestive was known from preserved specimens, in which the spheres
cavity, cells act individually to digest the prey. were contracted. Their histology and reproduction is poorly
known and could be studied only from specimens preserved in
The classification of the carnivorous sponges is presently
formalin or fixatives for electron microscopy. The present un-
rather problematic (HAJDU & VACELET 2002). They all belong
certainties in the distinction of evolutionary lines in carnivo-
to the order Poecilosclerida, and most are classified in the fam-
rous sponges require more information on their molecular char-
ily Cladorhizidae, with several genera dinstinguished mainly by
acters, and the preservation of fragments in alcohol is highly
the microsclere spicules. These microscleres, however, belong
desirable.
to different evolutionary lines of the Poecilosclerida. Further-
more, it appears that a carnivorous feeding habit also exists in
some other poecilosclerids displaying a similar morphology, but
classified in different families due to their different cheloid mi-
croscleres. Molecular characters have not yet been used for the
distinction of the evolutionary lines of these sponges.

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.
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Porifera, Demospongiae, Poecilosclerida, Cladorhizidae

Abyssocladia dominalba VACELET, in press

Size: 30 mm high, with a body 10 mm in diameter. Biology: On a dead smoker near an active site, water tempera-
ture 2.6°C at the site of collection. A carnivorous feeding habit
Color: White in alcohol.
has been demonstrated in another species of the genus.
Morphology: A thin peduncle bearing an ovoid or subspherical
Distribution: North-Fiji Back Arc Basin: site White Lady.
body made of radiating fascicles. No aquiferous system. Skele-
ton: Peduncle made of longitudinally arranged long fusiform
styles; radiating fascicles made of fusiform styles and smaller
styles 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 end
tridentate, the other end with fused alae, 9.5-11 µm; sigman-
cistra in two classes, 30-40 µm and 9.5-12.5 µm.

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).

Reference:
VACELET J. (in press) Zool. J. Linn. Soc.

J. VACELET & M. SEGONZAC Denisia 18 (2006): 37


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Porifera, Demospongiae, Poecilosclerida, Cladorhizidae

Abyssocladia naudur VACELET, in press

substrongyles or strongyles of the basal coating, fusiform, bent


Size: Up to 40 mm high, with thin lateral filaments 6 mm long.
or slightly flexuous, 30-825 x 8-30 µm; abyssochelae with
Color: White in situ, yellowish gray to clear brown in alcohol. frontal alae roughly parallelepipedal, nearly in contact with the
opposite frontal ala, 48-72 µm; sigmancistra in two classes, 15-
Morphology: Small erect sponge, forming a flattened axis with
19 µm and 5-8 µm.
numerous lateral filaments, frequently with a bud-like branch-
ing process. Filaments regularly arranged in two lateral rows, al- Biology: Several specimens collected from a dead smoker, a few
ternating on each side. No visible aquiferous system. Skeleton: meters from active black smokers. Presumably with a carnivo-
main axis of fusiform styles longitudinally arranged, lined by rous feeding habit.
substrongyles at the base; axis of the processes conical at the
Distribution: East Pacific Rise: 17°S.
base, with the styles anchored by their head entirely crossing
the 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;

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).

Reference:
VACELET J. (in press) Zool. J. Linn. Soc.

J. VACELET & M. SEGONZAC Denisia 18 (2006): 38


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Porifera, Demospongiae, Poecilosclerida, Cladorhizidae

Asbestopluma agglutinans VACELET, in press

is, 1550-2100 x 30-35 µm; mycalostyles of the filaments, 370-


Size: Axis 4 cm high and 0.8-1 mm in diameter, lateral fila-
780 x 8.5-17 µm; styles or strongyles of the base, 220-535 x 15-
ments up to 5-6 mm long.
42 µm; acanthotylostrongyles, 65-165 x 0.8-2.3 µm; aniso-
Color: Brown in alcohol. chelae, 32-36 µm and 9.8-10.5 µm; sigmancistras, 23-28 µm.
Morphology: Small erect sponge, consisting of a flattened axis Biology: Collected on a dead smoker and a basalt fragment.
smooth on the base, then with biserially arranged filaments
Distribution: East Pacific Rise: collected at 18°S and 14°S.
arising perpendicularly to the axis in two opposite series. Fila-
ments thick, cylindrical at the basis, then abruptly reduced to a
thin spicular axis. No visible aperture. Skeleton: axis of large
fusiform styles longitudinally arranged, surrounded by a felt-
work of acanthotylostrongyles including numerous skeletons of
radiolarians and foraminiferans; base with a cover of tangential
flexuous styles or strongyles. Spicules: fusiform styles of the ax-

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, scale
bar 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: Anisochelae
2, scale bar 2 µm; J: Anisochela 2, back view, scale bar 2 µm; K: Sigmancistra, scale bar 2.9 µm; from VACELET (in press).

Reference:
VACELET J. (in press) Zool. J. Linn. Soc.

J. VACELET & M. SEGONZAC Denisia 18 (2006): 39


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Porifera, Demospongiae, Poecilosclerida, Cladorhizidae

Asbestopluma formosa VACELET, in press

dense outer cover of microstrongyles, filaments with an axis of


Size: Approximately 8 cm high and 16 cm wide, with stem and
smaller styles; base of the stem with a cover of special microty-
branches up to 1 mm in diameter.
lostyles and short fusiform substrongyles. Spicules: Styles, 200-
Color: White in situ and after preservation. 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 x
Morphology: Erect, fan shaped, consisting of a short stem from
5-7 µm; microtylostyles, minutely spinose, 25-45 x 4-7 µm;
which arise branches which divide dichotomously three or four
anisochelae sometimes in rosettes, 72-90 µm and 10-15 µm.
time in a single plane, the last branches being long and paral-
lel. Terminal branches flattened, bearing on both sides numer- Biology: On a fossil chimney ca. 50-100 m distant from an ac-
ous, regularly spaced thin filaments. Flattened enlargements tive site White Lady. Presumably with a carnivorous feeding
present at each dichotomy containing numerous reproduction habit.
bodies. No visible apertures or aquiferous system. Skeleton:
Distribution: North-Fiji Back-Arc Basin.
Stem and branches with an axis of large fusiform styles with a

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: Microstrongyle
and detail of the head, scale bars 6.2 µm and 1.5 µm;
2: Collection of the holotype by the arm of “Nau- E: Two anisochelae 1 and an immature one, scale bar
tile“ submersible, 1997 m; from VACELET (in press) 10 µm; F: Two microtylostyles, scale bar 4.3 µm; G:
© Ifremer/Starmer 1. Anisochelae 2, scale bar 2 µm; from VACELET (in press).

Reference:
VACELET J. (in press) Zool. J. Linn. Soc.

J. VACELET & M. SEGONZAC Denisia 18 (2006): 40


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Porifera, Demospongiae, Poecilosclerida, Cladorhizidae

Asbestopluma pennatula (SCHMIDT, 1875)

slightly fusiform with an acerate end, straight, fusiform, 380-


Size: 100 mm high. Lateral filaments up to 2.5 mm x 0.1-0.2
520 µm x 7-10 µm. Microtylostrongyles coating the surface of
mm, possibly longer but easily broken. Specimens up to 180
the axis near its basis, slightly flexuous, with a short spination,
mm with filaments 7 mm long have been recorded near Ice-
40-130 x 0.5-1 µm. Palmate anisochelae I, moderately abun-
land.
dant, longitudinally arranged with the teeth upwards on the fil-
Color: Cream in situ and in alcohol. aments, with well marked lateral teeth in the small end, 33-40
µm. Palmate anisochelae II, very abundant, perpendicular to
Morphology: Erect axis with distinct groups of lateral filaments
the surface of the filaments with the large tooth upwards, 9-10
arising perpendicularly, regularly spaced along the stalk. No
µm. Sigmancistras very abundant, 17-25 µm.
aquiferous system. Skeleton: spicular axis of styles in the stalk
and in lateral filament, covered at the basis of the sponge by a Biology: Attached to solid substrata in the bathyal zone.
coating of spinose microtylostrongyles. Spicules: Styles of the
Distribution: North Sea, Arctic, Bay of Biscay. Collected at
axis of the stalk, fusiform, 700-920 x 17-22 µm, shorter and
Mid-Atlantic Ridge: Lucky Strike and Menez Gwen.
thicker at the basis of the axis where they gradually become
substrongyles 200-540 x 18-30 µm. Styles of the filaments,

1A: Specimen from Lucky Strike, scale bar 5 mm; B: Style of the filament, scale bar 50 µm; C: Style of the filament and
head 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.

References:
LUNDBECK W. (1905) The Danish Ingolf-Expedition 6(2): 1-219.

J. VACELET & M. SEGONZAC Denisia 18 (2006): 41


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Porifera, Demospongiae, Poecilosclerida, Cladorhizidae

Chondrocladia lampadiglobus VACELET, in press

140 µm, alae 25 µm long, and 20-32 µm, alae 10-11 µm long;
Size: Up to approximately 50 cm high, with inflated spheres 3-
sigmas 45-120 x 2-3 µm.
5 cm in diameter.
Biology: The collected specimen was rooted in sediment be-
Color: White in life, yellowish white to clear brown in alcohol.
tween pillow lava, near active hydrothermal sites, but in an
Morphology: Large stalked sponge, composed of a rhizoid fixa- area still with low density of animal life. Sponges of similar
tion system, a cylindrical stalk ending in an enlarged, ovoid morphology have been often observed on various sites of the
body from which radiate in all directions secondary branches, East Pacific Rise, either rooted in sediment or attached to pil-
each ending in translucent sphere in the living animal, in an ir- low lava, always at some distance from the rich animal com-
regular swelling including crustacean debris on the preserved munities of the active hydrothermal sites. Their identification
specimen. Aquiferous system present, with large choanocyte to C. lampadiglobus cannot be ascertained from external mor-
chambers and canals ending in the inflatable spheres. Skeleton: phology alone, and several species may be present in this large
stalk and branches made of large fusiform styles longitudinally geographic zone. Carnivorous mode of feeding.
arranged, covered with a feltwork of rugose tylostyles; terminal
Distribution: East Pacific Rise: collected at 17°S. Sponges of
swellings made of smaller styles with an outer cover of numer-
similar morphology observed in various sites extending from
ous microscleres. Spicules: fusiform styles, 700-4750 x 15-75 µm
23°S to 13°N, 2600-3000 m deep.
and 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-

1: ROV Tiburon/2003, dive 556, 20°47.03’N, 109°08.98’W, 2555 m; by R. 2A: Holotype; scale bar 17 mm; B: Tylostyle of the
Vrijenhoek © MBARI. cover of the stalk; scale bars: 42 µm and 4.2 µm; C:
Style of the stalk; scale bar 30 µm; D: Style of the
body surface; scale bar 63 µm; E: Anchorate isochelae
1; scale bar 14 µm; F: Anchorate isochelae 1; scale bar
9.6 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


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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).

Reference:
VACELET J. (in press) Zool. J. Linn. Soc.

43
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Porifera, Demospongiae, Poecilosclerida, Cladorhizidae

Cladorhiza abyssicola G.O. SARS, 1872

unguiferate anisochelae with five teeth at each end, 21-25 µm;


Size: Up to 75 mm high.
sigmas, 78-100 µm; sigmancistras, 40-42 µm.
Color: Cream white in alcohol.
Biology: A common species in the bathyal and abyssal zones,
Morphology: Erect, with numerous lateral branches arising at usually anchored in mud by the roots, with a carnivorous mode
nearly right angle from a central axis and bearing short second- of feeding. A near relative, C. methanophila VACELET & BOURY-
ary processes, generally anchored in the sediment by richly ESNAULT, 2002, is both carnivorous and symbiotic with
branched roots. No aquiferous system. Main skeleton: polyspic- methanophilous bacteria.
ular fibres of styles in the axis of stem, branches, lateral process-
Distribution: Atlantic N.E., Arctic, Mediterranean, Mid-At-
es and roots. Spicules: Styles, fusiform, 390-730 µm x 14-22 µm;
lantic Ridge.

1A: Type specimen from Lofoten, 550 m, approximately 60 mm high, from SARS (1872); B: Specimen from MAR, Logatchev
vent site, DiversExpedition, dive 3668, 3012 m, with oocytes and embryo; scale bar 3 mm; C: Style and anisochelae; scale
bar 30 µm; D: Anisochela; scale bar 5 µm; E: Sigmas; scale bar 25 µm; F: Sigmancistra; scale bar 8 µm.

References:
LUNDBECK W. (1905) The Danish Ingolf-Expedition 6(2): 1-219.
SARS G.O. (1872) in Kongelige Norske Universitet (Ed.) Spongiae. Volume I. Brøgger & Christie, Christiania, Norway: 1-82.

J. VACELET & M. SEGONZAC Denisia 18 (2006): 44


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Porifera, Demospongiae, Poecilosclerida, Cladorhizidae

Cladorhiza segonzaci VACELET, in press

processes, slightly fusiform, 380-990 x 14-23 µm; anchorate/un-


Size: Axis up to 32 mm high and 0.4-0.5 mm in diameter, with
guiferate anisochelae numerous, with five lanceolate alae and
lateral processes 4-9 mm long.
well developed fimbriae at the large end, and three fang-like
Color: In situ white; cream to clear brown in alcohol, with the alae at the small end, 15-17.5 µm; sigmas 50-80 x 1-1.5 µm; sig-
basal portion of the stem darker. mancistras without notch, 20-25 x 2.0 µm.
Morphology: Small erect sponge, forming an unbranched axis Biology: 11 specimens collected from a dead smoker, a few me-
with numerous lateral processes arranged in opposed series at ters from active black smokers. Presumably carnivorous mode of
right angle all around the axis. No aquiferous system. Skeleton: feeding.
main axis of fusiform styles longitudinally arranged; axis of the
Distribution: East Pacific Rise: 17°S.
processes conical at the base, with the styles anchored by their
head reaching the centre of the stem and the point outwardly
directed. Spicules: Styles of the axis of the stem and lateral

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).

Reference:
VACELET J. (in press) Zool. J. Linn. Soc.

J. VACELET & M. SEGONZAC Denisia 18 (2006): 45


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Porifera, Demospongiae, Poecilosclerida, Guitarridae

Euchelipluma pristina TOPSENT, 1909

700 x 6-14 µm; palmate isochelae, 75-130 µm, in two size cate-
Size: 14-120 mm high, shaft 3-4 mm wide, lateral filaments up
gories in some specimens; placochelae I, 60-90 µm; placochelae
to at least 8 mm long and 70-100 µm in diameter.
II, 100-130 µm, absent in the type specimen; sigmancistra I, 11-
Color: Clear brown in alcohol. 14 µm; sigmancistra II, 22-30 µm, sometimes absent.
Morphology: Erect, pennaceous, composed of a short peduncle Biology: Type specimens collected from 91 m. Fixed on solid
and a flattened shaft bearing symmetrically paired lateral fila- substrata up to 4960 m deep. The presence of lateral filaments
ments, broken and reduced to their basis in most specimens. No covered by microscleres regularly arranged with the teeth up-
aquiferous system. Skeleton a spicular axis of styles, condensed wards, and the absence of canal system and apertures in the col-
at the basis and divided in parallel fibres upward; lateral fila- lected specimens suggest a carnivorous mode of feeding, similar
ments with a twisted axis of subtylostyles. Microscleres most of- to that developed in Cladorhizidae. This is supported by the
ten regularly arranged at the surface of the shaft and of the fil- presence of debris of small crustaceans in the best preserved
aments, with the teeth outwardly directed. Spicules: Styles of specimens from Barbados (cruise Manon © Ifremer).
the axis, fusiform with an obtuse point, 900-1550 x 15-40 µm;
Distribution: Cape Verde Islands (91 m); Barbados (4960 m);
strongyles in the basis of the peduncle, slightly curved or flexu-
Mid-Atlantic Ridge: Lucky Strike; Rainbow.
ous, 210-490 x 14-22 µm; subtylostyles in the filaments, 330-

1A: Two specimens


from Barbados; scale
bar 4 mm;
B: Head of a subty-
lostyle, Barbados;
scale bar 10 µm;
C: Subtylostyles,
placochelae I and II,
isochela, sigmancistras
I and II, Barbados;
scale bar 50 µm;
D: Immature
placochela, 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.

References:
TOPSENT E. (1909) Bull. Inst. Océanogr. Monaco 151: 1-23.
TOPSENT E. (1928) Résult. Camp. Sci. Prince Albert Ier de Monaco 74: 1-376.

J. VACELET & M. SEGONZAC Denisia 18 (2006): 46


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Porifera, Hexactinella, Lyssacinosida, Caulophacidae

Caulophacus cyanae BOURY-ESNAULT & DE VOS, 1988

Size: Up to 50 cm Distribution: East Pacific Rise: 13°N. On vertical walls, inac-


tive sulphide edifices and basaltic pillars in the graben.
Morphology: Mushroom-like body, white with a solid stalk and
Caulophacus-like Porifera occur along the northern and south-
a convex discoid upper part; ectosomal skeleton of small pen-
ern East Pacific Rise and Pacific-Antarctic Ridge: 38°S, but in
tactine, pinnular dermalia and of strong pentactine hypoder-
fewer abundance.
malia. Choanosomal megascleres of hexactines, pentactines
and rhabdodiactines. Microscleres are discohexasters of three
sizes. Large choanocyte chambers composed of anucleate
choanocytes.

1: Some specimens in situ; cruise Hope @ Ifremer.

2: Spicules (SEM); after BOURY-ESNAULT & DE VOOS (1988). 3: A preserved specimen; by P. Briand @ Ifremer.

Reference:
BOURY-ESNAULT N. & L. DE VOS (1988) Oceanol. Acta 8: 51-60.

N. BOURY-ESNAULT Denisia 18 (2006): 47


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Cnidaria, Hydrozoa, Leptolida (= Hydroida)

The leptolid fauna of hydrothermal vents of oceanic ridge Collecting activities at or near Mid-Atlantic vents areas
sites is still poorly investigated and understood. Fragmentary have currently provided material for proper identification of 3
colonies of leptolids from oceanic ridges in the Pacific have so anthoathecates and 23 leptothecates; 4 more leptothecates
far made it possible to identify Zygophylax cervicornis (NUTTING, could only be identified to the genus (CALDER & VERVOORT
1905), Zygophylax sp. and Halecium tenellum HINCKS, 1861. 1998). One more species (Hydrallmania falcata) was dead when
While from vents on the Mid-Atlantic Ridge only a single an- collected and accidentally introduced; all remaining leptothe-
thoathecate leptolid, Candelabrum serpentarii SEGONZAC & cates belong to the deep-water fauna and are only party re-
VERVOORT, 1995 can be considered to be a true companion of stricted to the Atlantic.
the vent community. A second species, Candelabum phrygium
Because of the technique used to collect animals at ridge
(FABRICIUS, 1780) is a North Atlantic deep-sea inhabitant oc-
sites, only rarely do complete animals or colonies become avail-
casionally found associated with the hydroythermal vent fauna.
able for scientific study. Anthoathecates in particular are hard
Another anthoathecate, Bouillonia sp. was found on the to collect and they must be narcotized prior preservation. So
Mid-Atlantic Ridge in the Gulf of Guinea area; it probably rep- far, only a single complete animal of Candelabrum serpentarii has
resents the deep-water leptolid community without direct been obtained, in addition to many incomplete individuals,
affinities to hydrothermal vents (SVOBODA, STEPANJANTS & which have been studied. Anthoathecates are best preserved in
LJUBENKOV, in press). Photographs taken during surveys of Mid- 6% formalin (after prior relaxation by adding MgSO4 solution
Atlantic vent areas show a third species of anthoathecate, oc- to the seawater), but Leptothecates get very brittle in formalin
casionally in great numbers; it may represent a species of Tubu- and are best preserved in 70% ethanol. These preservation
laria or Ectopleura. There is presently no material available to techniques only apply to material for routine taxonomic inves-
substantiate that guess. A probably undescribed species of Hy- tigations; more sophisticated methods are necessary for modern
dractinia has also been observed. DNA techniques.

References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 1-65.
SEGONZAC M. & W. VERVOORT (1995) Bull. Mus. Natl. Hist. Nat., Paris (4)17(1-2): 31-64.
SVOBODA A., STEPANJANTS S. & J. LJUBENKOV (in press) Zool. Meded., Leiden.

W. VERVOORT Denisia 18 (2006): 48


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Cnidaria, Hydrozoa, Leptolida, Anthoathecata, Candelabridae

Candelabrum phrygium FABRICIUS, 1780

20.5 x 8.2-9.9 µm; small desmonemes 8.2-9.0 x 6.4 x 6.6 µm;


Size: 7-9 cm.
large desmonemes 12.5-13.0 x 9.0-9.8 µm; stenoteles 10.6-11.5
Color: White to brownish. x 8.2-9.8 µm.
Morphology: Body in extended condition slender, composed of Biology: Attached to rocks or other solid objects. Lucky Strike
foot, blastostyle bearing region and trunk. Foot lobed and flat- specimens were attached to flange formations and occasionally
tened part of body, attaching animal to rocks or bivalve mol- to organic support (shells of living bivalves). Carnivorous ani-
luscs. Blastostyle bearing region occupying one third to one mal, feeding on shrimps and other small crustaceans. Dioecious,
quarter part of body, with 10-15 large, tubular blastostyles, bear- development of larva unknown, but probably young polyp de-
ing male or female gonophores in various stages of development velops inside female gonophore.
and dispersed capitate tentacles; apex of each blastostyle with
Distribution: Circumarctic, occurring both in Atlantic and Pa-
circle of 4-5 such tentacles. Remainder of body forming elon-
cific Oceans. Mid-Atlantic Ridge: Lucky Strike and Rainbow.
gated, tubular trunk, completely covered by capitate tentacles.
Mouth at distal end of trunk. Nematocysts: Haplonemes 19.7-

1: Several animals in situ


on substrate; cruise Diva 2
© Ifremer.

2: Preserved specimen on small rock in Lucky Strike area, Mid-Atlantic Ridge;


by P. Briand © Ifremer.

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


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Cnidaria, Hydrozoa, Hydroida, Anthomedusae, Candelabridae

Candelabrum serpentarii SEGONZAC & VERVOORT, 1995

Size: 10 cm. Biology: Attached to rocks (pillow lava or sulfide rocks) in


vicinity of active vents. Probably exclusively carnivorous, feed-
Color: white, whitish or pale orange.
ing on pericarideans and shrimps. Reproduction: Monoecious
Morphology: Robust, gelatinous body composed of foot, blas- with hermaphroditic gonophores; development of larva un-
tostyle bearing region and trunk, attached to pillow lava or sul- known, but probably young polyp (actinula) developing in fe-
fide rocks by means of (flattened) foot covered by thick, dark male gonophore.
brown perisarc. Blastostyles on proximal part of body large,
Distribution: Mid-Atlantic Ridge: Snake Pit.
bearing a number of male and female gonophores in various
stages of development; elongated trunk completely covered
with capitate tentacles, in extended condition several mm
long. Mouth at distal end of trunk. Nematocysts: Haplonemes
16.5-18 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.

2: In situ, Snake Pit area, Mid-Atlantic Ridge;


cruise Hydrosnake © Ifremer.

1: In situ, Snake Pit area, Mid-Atlantic Ridge; cruise Hydrosnake


© Ifremer.

Reference:
SEGONZAC M. & W. VERVOORT (1995) Bull. Mus. Natl Hist. Nat., Paris, 4e sér. 17(1-2): 31-64.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 50


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Cnidaria, Hydrozoa, Leptolida, Anthoathecata, Eudendriidae

Eudendrium planum BONNEVIE, 1898

one or two chambers, attached to non-reduced hydranths. Fe-


Size: Colonies up to 90 mm high, branching reduced.
male gonophores undescribed. Nematocysts: macrobasic eury-
Morphology: Colonies slender, stiff, sparsely branched, hy- teles 12.9-14.1 x 5.5-6.4 µm; heterotrichous microbasic euryte-
drorhiza creeping, growth monopodial with terminal hydranth. les 7.5-8.4 x 3.9-4.6 µm.
Stem stiff, upright, basally polysiphonic but largely monosi-
Biology: Attached to fixed object like stones or rocks. At
phonic; branching irregular, more or less in one plane, pedicels
Lucky Strike site, attached to flanges where mussels are fixed.
alternate, long. Perisarc thick, yellowish, thinning out distally,
Probably a dioecious species feeding carnivorously.
terminating at bases of hydranths. Annulations present at base
of pedicels and branches, occasionally elsewhere but perisarc Distribution: Deep water of the north-eastern Atlantic; Mid-
mostly smooth. Hydranths about 500 µm long, urn-shaped, Atlantic Ridge: Logatchev, Lucky Strike, Rainbow.
with large, flared hypostome. Tentacles filiform, numbering
about 20. Gonophores fixed sporosacs. Male gonophores with

1 right side: Part of hydrocaulus with


pedicel and (damaged) hydranth; scale
bar 0.5 mm; left side: Male gonophore
attached to base of hydranth; scale bar
0.25 mm; from CALDER & VERVOORT (1998).

Reference:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 51


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Cnidaria, Hydrozoa, Leptolida, Anthoathecata, Tubulariidae

Ectopleura larynx ELLIS & SOLANDER, 1786

small stenoteles 6.7-7.3 x 5.4-5.7 mm; large stenoteles 10.6-


Size: Height of individual polyps up to 50 mm.
21.1 x 9.5-10.1 µm; mastigophores 8.4-9.6 x 3.5-4.0 µm;
Morphology: Colonies formed by branched hydrocauli or by O-isorhiras 9.2-9.4 x 8.4-8.7 µm.
development of actinulae on individual hydrocauli arising from
Biology: Attached to fixed objects but frequently developing
creeping, twisted stolon with wrinkled, straw-coloured perisarc
into a free-living, detached colony as the result of fragmenta-
attached to fixed objects. Diameter of hydrocaulus about 400
tion of the original colony. Monoecious or dioecious colonies.
Ìm, perisarc firm basally, gradually thinning about distally and
The eggs develop inside the female gonophore and become free
terminating just below bulbous base of terminal, vasiform hy-
in the actinula stage. As the possibilities for dispersion of the
dranth. Height of hydranth about 1.7 mm, width 1.2 m; one
actinulae are limited many settle on or in the vicinity of the
whorl of short oral tentacles, up to 750 µm long and one whorl
elder.
of much longer aboral tentacles, up to 2.800 µm long.
Gonophores fixed sporosacs, developing just above insertion of Distribution: Mid-Atlantic Ridge: Menez Gwen. Sub-arctic,
aboral tentacles on short, slender dichotomously branches blas- temperate and subtropical waters of the northern Atlantic,
tostyles. Both male and female gonophores are eumedusoid from the littoral zone down to considerable depth in the ba-
without remnants of radial canals but with three rudimentary thyal zone.
tentacles. Nematocysts: Desmonemes 3.8-4.6 x 3.0 x 3.6 µm;

1: Hydranth with female gonophores, seen from above 2: Tubularia or Ectopleura sp. in situ, from Menez Gwen vent
(oral view); from CALDER & VERVOORT (1998). field at the boundary of active area; cruise Ataos © Ifremer.

Reference:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 52


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Halopterididae

Polyplumaria flabellata G.O. SARS, 1874

axis of hydrotheca at 45∞ to length axis of internode; hy-


Size: Height of colony up to 350 mm, spread almost equal.
drothecal rim circular and smooth, perpendicular to hydrothe-
Morphology: Stolonal mass of fine tubules supporting a rigid, cal length axis; lateral (flanking) nematothecae on short
pinnately branched colony of very regular structure. Stem poly- apophyses near hydrothecal axil. Hydrocladia occasionally
siphonic and repeatedly branched, branches in opposite pairs; forked, secondary branch springing from base of first hydrothe-
branches originating from secondary tubes; primary axis visible ca. Depth of hydrotheca 140-190 µm, diameter at rim 135-160
in polysiphonic parts in front of bundle of secondary tubules, µm. Female and male gonothecae of same shape, curved ovoid,
divided by oblique nodes into internodes of varied length, each attached to apophyses by means of short pedicel; basal part of
internode with one to five apophyses, alternately directed gonothecae with 6-9 nematothecae. Aperture circular, laterally
obliquely upwards and to left and right. Primary and secondary disposed, turned towards hydrocladium, closed by circular lid.
tubules with nematothecae, each apophysis with a ‘mamelon’ Length of gonothecae 610-750 µm; diameter 340-400 µm. Ne-
and three nematothecae; in addition two to four nematothecae matocysts: Not studied in detail.
between two successive apophyses. Hydrocladia about 3 mm
Biology: Usually on soft bottoms (off vent areas). Serves as host
long, supported by apophyses, slightly curved, first internode
for many epizoic leptolid species.
athecate, with one nematothecae; all following internodes the-
cate, with one proximal, a pair of flanking, one distal and sev- Distribution: Mid-Atlantic Ridge: near Menez Gwen, 997-696
eral additional and variously distributed nematothecae on dis- m. Moderately deep to deep waters of tropical, temperate, and
tal part of internode. All nematothecae bithalamic with deeply northern parts of the eastern Atlantic.
scooped adcauline wall. Hydrotheca cylindrical with almost
parallel walls, adcauline wall adnate for half its length, length

1 bottom: Colony; from COR-


NELIUS (1995); right: Central axis
of branch with apophysis and
base of hydrocladium; scale
bar 0.25 mm; left: Hydrocladial
internode with hydrotheca and
nematothecae; scale bar 0.25
mm; top: Gonothecae; scale
bar 0.10 mm; from CALDER &
VERVOORT (1998).

References:
ANSÍN AGÍS J., RAMIL F. & W. VERVOORT (2001) Zool. Verh., Leiden 333: 3-268.
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
CORNELIUS P.F.S. (1995) North-West European Thecate Hydroids and their Medusae, 2. Synopses of the British Fauna (New Series) 50: vii, 1-386.
RAMIL F. & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 53


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Hebellidae

Halisiphonia arctica KRAMP, 1932

short, ringed pedicel, distally open, truncated, rim notched.


Size: Length of hydrotheca about 1 mm; length of pedicel up to
Gonophores developing free medusae that are so far unde-
3 mm.
scribed. Nematocysts: Unknown.
Morphology: Stolonal colony with hydrothecae and gonothe-
Biology: Attached to firm substrate. Although it seems clear
cae developing from branching and anastomosing stolons. Hy-
that the gonophore develops free medusae these are unde-
drothecae pedicellate, pedicel up to 3 mm long, smooth, with
scribed so far. The presense of free medusae in Halisiphonia
some proximal annulations, merging into almost tubular to
brings it close to such species of Hebella that also have a free
slightly conical hydrotheca, separated from pedicel by quite
medusa in their life-cycle.
thin diaphragm. Hydrothecal rim slightly everted, frequently
renovated. Hydranth unknown. Gonothecae big, flattened, Distribution: Mid-Atlantic Ridge: Logatchev.
about 2.5 mm long, spade-shaped, proximally narrowing into a

1: Colony composed of hydrothecae of varied lengths and a gonotheca on stem of 2: Gonotheca with developing medusa;
hydroid. Length of hydrotheca, including pedicel, 0.8- 5.0 mm; from KRAMP (1932). scale bar 0.5 mm; from SCHUCHERT (2001).

References:
KRAMP P.L. (1932) Medd. om Grønland 79: 1-86.
SCHUCHERT P. (2001) Medd. Grønland Biosci. 53: 1-184.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 54


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Lafoeidae

Grammaria abietina M. SARS, 1850

rim 270-306 µm. Gonothecae aggregated into ovoid coppiniae


Size: Colonies from the Mid-Atlantic Ridge are a few cm high;
with many strongly curved tubules projecting from between
in boreal and subarctic waters colonies may reach a size of 100
gonothecae. Coppiniae hermaphroditic, female gonothecae
mm.
amphora-shaped, narrowing into short neck distally; male
Morphology: Irregularly branched, erect colonies with polysi- gonothecae globular and pedicellate. Eggs developing in an
phonic stems and branches, only distal parts of branches mono- acrocyst projecting from female gonothecae. Nematocysts: Not
siphonic. In these parts hydrothecae placed in four, six or described in detail, “large and small nematocysts“ (BROCH
(rarely) eight longitudinal series with hydrothecae of adjacent 1918) being present.
rows alternate; in quadriseriate colonies hydrothecae in oppo-
Biology: Colonies developing on solid substrata. Eggs brooded
site pairs and decussate. Polysiphony caused by copious devel-
in an acrycyst.
opment of secondary tubules obscuring structure of polysiphon-
ic stems and branches. Hydrothecae tubular, half of adcauline Distribution: Mid-Atlantic Ridge: Menez Gwen. Generally
wall adnate to stem or branch, curved outwards, rim circular, recognized as a circumpolar species, extending its distribution
not flared, frequently renovated. No nematophores or nema- southwards in the deep water of the northern Atlantic.
tothecae. Total length of hydrotheca 1.1-1.3 mm, diameter at

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).

References:
BROCH H. (1918) Danish Ingolf-Expedition 5(7): 1-205.
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
CORNELIUS P.F.S. (1995) North-West European Thecate Hydroids and their Medusae, 1. Synopses of the British Fauna (New Series) 50: vii, 1-347.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 55


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Lafoeidae

Lafoea dumosa FLEMING, 1820

lowish. Gonothecae aggregated into ovoid, several mm long


Size: Erect colony up to 100 mm high; length of hydrotheca
coppinia on stem or branches, hermaphroditic, with many
varied between 150 and 800 µm, diameter at rim 100-200 µm;
strongly curved accessory tubes protruding from between
greatest dimensions in specimens from cold water.
gonothecae. Gonothecae amphora-shaped, strongly adnate.
Morphology: Stolonal or erect colonies, often a combination of Presence of acrocyst unknown. Coppiniae rare, reproduction
both; erect colonies very irregularly branched, often with anas- apparently frequently vegetative. Nematocysts: Isorhiza’s of two
tomoses, mono- or polysiphonic, the latter being composed of size classes (about 21 and 16 µm length, not in same colony!)
several adnate parallel tubes all bearing hydrothecae on all and a smaller unidentified capsule, 6 µm long (SCHUCHERT
sides, occasionally in more or less opposite and decussate 2001).
arrangement. Hydrotheca tubular to elongated conical, fre-
Biology: On fixed objects, also epizoic. Although a well-dis-
quently asymmetrical, rim circular, usually renovated. Hy-
tributed species few details of its biology are available.
drotheca narrowing basally into frequently almost impercepti-
ble pedicel; other colonies may have a well developed, twisted Distribution: Mid-Atlantic Ridge: Rainbow. Shallow to deep
hydrothecal pedicel. Hydranth extensile, with about 20 amph- waters of all oceans [concept of species as defined by CORNELIUS
icoronate tentacles around conical proboscis, attached deep in- (1975)].
side hydrothecae by means of ring of desmocytes; coenosarc yel-

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).

References:
CORNELIUS P.F.S. (1975) Bull. Br. Mus. Nat. Hist., Zool. 28: 373-426.
CORNELIUS P.F.S. (1995) North-West European Thecate Hydroids and their Medusae, 1. Synopses of the British Fauna (New Series) 50: vii, 1-347.
RAMIL F. & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.
SCHUCHERT P. (2001) Medd om Grønland. Bioscience 53: 1-184.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 56


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Lafoeidae

Zygophylax echinata CALDER & VERVOORT, 1998

ly asymmetrical by bulging adcauline wall. Proximal portion of


Size: Height of erect colony 25 mm.
hydrotheca narrowing into short, plump pedicel; diaphragm
Morphology: Colony erect, stem strongly polysiphonic, mono- distinct, oblique. Length of hydrotheca (without renovations)
siphonic only in upper parts and there slightly geniculate. Hy- 250-400 µm, rim moderately everted, circular, frequently reno-
drocladia short, alternately arranged on either side of axis, bis- vated, diameter 94-117 µm. Gonothecae aggregated into char-
eriate, with axillar hydrotheca bearing two opposite nema- acteristic, oval coppinia, 3.5 mm long, 2 mm high, surrounding
tothecae at base of pedicel. Hydrocladia may become polysi- axis basally. Gonothecae flask-shaped, strongly adnate, tapering
phonic by apposition of secondary tubes and presence of nema- distally into a sharply pointed structure with a lateral aperture.
tothecae in axillar hydrothecae may become obscured. Hy- Nematophorous ramules protrude from between gonothecae,
drothecae on axis and hydrocladia alternately arranged, placed curving over coppinia, bearing both nematothecae and some
on distinct, swollen apophysis with a pair of opposite nema- hydrothecae. Nematocysts: Undescribed.
tothecae. Besides axillar hydrotheca there are two hydrothecae
Biology: Attached to solid substrate like rocks or stones.
between two successive hydrocladia. Nematothecae small,
elongated tumbler-shaped, narrowing proximally into short Distribution: Mid-Atlantic Ridge: near Menez Gwen.
pedicel, length 42-68 µm, diameter at rim 24-31 µm, renova-
tions frequent. Hydrothecae small, slender vase-shaped, slight-

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 of
nematophorous ramule with hydrotheca and two nematothecae, scale bar 0.1 mm; bottom row middle: Three gonothecae and
nematophorous ramule from coppinia, scale bar 0.25 mm; right: Apophysis of hydrocladium with pedicel of hydrotheca and
two nematothecae, scale bar 0.1 mm; after CALDER & VERVOORT (1998).

Reference:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 57


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Lafoeidae

Zygophylax leloupi RAMIL & VERVOORT, 1992

lodged, on apophyses of hydrothecae, cylindrical, proximally


Size: Height of colony up to 140 mm.
rounded and shortly pedicellate. Gonothecae aggregated into
Morphology: Structure of colony in principal as in Z. echinata loose coppinia without nematophorous ramules, ovoid, distally
CALDER & VERVOORT, 1998, but colony generally bigger, with with two or three short tubular processes each with terminal
stronger, repeatedly bifurcated stem, with longer hydrocladia aperture. Coppinia surrounding proximal part of stem. Nema-
and distinctly pedicellate, big hydrothecae that are distinctly tocysts: Not described.
frontally directed. Length of hydrotheca (from diaphragm on-
Biology: Attached to solid substrata, among mussel bed.
wards, without renovations) 500-650 µm, diameter at rim 280-
340 µm; renovations frequent. Pedicel long, wrinkled or with Distribution: Mid-Atlantic Ridge: Rainbow; seamount at Seg-
septa, 270-250 µm; diaphragm distinct but thin, occasionally ment 38°N, 919 m.
duplicated. Aperture of hydrotheca slightly tilted, rim everted,
diameter 210-325 µm. Nematothecae scarce, frequently dis-

1 left: Monosiphonic part of stem


with three hydro-thecae; middle:
Hydrotheca and nematotheca; right:
Gonotheca isolated from coppinia.
Length of hydrotheca about 0.8
mm, length of gonotheca 1.0-1.4
mm; from RAMIL & VERVOORT (1992).

References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
RAMIL F. & VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 58


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Sertulariidae

Hydrallmania falcata LINNAEUS, 1758

aperture on short distal tube, rather wide. Length of gonothe-


Size: Full grown colonies may reach a length of 650 mm; usual-
cae 1.2-1.7 mm; greatest diameter 550-700 µm. Nematocysts:
ly much smaller.
Not described in detail.
Morphology: Colony lax, with characteristic, spirally twisted,
Biology: Frequent on soft bottoms in shallow to moderately
monosiphonic stem, supporting branches of 20-40 mm length
deep waters, particularly in temperate regions. One of the few
with pinnately arranged hydrocladia. Each hydrocladium divid-
species of Leptolida was (and probably still is) commercially ex-
ed into internodes, supporting closely packed groups of 3-10 hy-
ploited. Gonothecae produced in early spring. Colonies serve
drothecae, alternately inclined left or right to a moderate de-
the attachment for mussel spat.
gree. Hydrotheca more or less tubular, largely free, slightly nar-
rowing distally, aperture circular, operculum two-flapped. Total Distribution: Mid-Atlantic Ridge: Lucky Strike. Shallow to
length of hydrotheca 300-400 µm; greatest diameter 130-190 moderately deep waters of the temperate and northern At-
µm; diameter at rim 80-140 µm. Male and female gonothecae lantic; not a deep water species.
similar, elongated pear-shaped, narrowing proximally into short
pedicel attaching gonothecae to internode of hydrocladium;

1: Colony, height about 9 cm; 2 from left to right: Normally developed hydrocladium; hydrocladium
from CORNELIUS (1995). from young colony (similar arrangement of hydrothecae occasionally
also in older colonies); hydrotheca; gonothecae; scale bars 0.5 mm;
from CORNELIUS (1995).

References:
CORNELIUS P.F.S. (1995) North-West European Thecate Hydroids and their Medusae, 2. Synopses of the British Fauna (New Series) 50: VII, 1-386.
RAMIL F & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 59


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Sertulariidae

Sertularella tenella ALDER, 1856

cae undistinguishable, ovoid, 900-2.000 µm long, greatest di-


Size: Erect colonies about 20 mm high or smaller.
ameter 550-1.000 µm, with 8-10 transverse annulations; apex
Morphology: Monosiphonic, occasionally branched stem aris- with 3-4 low cusps surrounding a small, circular aperture. Ne-
ing from firm, tortuous stolon attached to solid substrate or matocysts: Not studies in detail.
host; stem geniculate, composed of slender internodes with a
Biology: Frequently on foliate Bryozoa and epizoic on other hy-
few basal rings; nodes indistinct. Hydrotheca with almost fee
droids.
abcauline wall, almost cylindrical to slightly swollen in proxi-
mal half, with six annulations, best visible on abcauline side; Distribution: Mid-Atlantic Ridge: Lucky Strike, Sintra. Cir-
depth 450-840 µm, greatest diameter 220-370 µm. Rim with cumglobal in deeper parts of the littoral zone to greater depths
four low cusps; operculum four-flapped, flaps attached in em- (1000-2000 m).
bayments between marginal cusps. Female and male gonothe-

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.

References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
RAMIL F. & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 60


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Sertulariidae

Symplectoscyphus bathyalis VERVOORT, 1972

lum observed. Gonotheca pyriform, 1.7 mm high; greatest di-


Size: Height of colony up to 80 mm.
ameter at two-thirds of distance from top, with about seven
Morphology: Erect colony, rising from creeping stolon. Proxi- transverse ribs, petering our proximally. Orifice at end of a short
mal parts of stem and branches polysiphonic; branching irregu- tube in middle of apical ‘field’ formed by distalmost rib. Pedicel
lar, basal part of stem ahydrothecate. Monosiphonic parts of quite short, attaching gonothecae to internode at hydrothecal
colony composed of long, slender, geniculate internodes sepa- base. Nematocysts: Not studies in detail.
rated by nodes sloping in alternate directions. Hydrothecae
Biology: On solid substrates in deep ocean water.
950-1.200 µm deep, greatest diameter about 500 µm, placed at
end of internode, one-fifth of abcauline wall adnate, curved Distribution: Mid-Atlantic Ridge: Lucky Strike, Sintra site.
outwards, slender, abcauline wall convex, abcauline wall Deep water of Atlantic (Bay of Biscay) and Pacific (off Chile
bulging in proximal third. Hydrothecal rim not everted, occa- and SE of New Caledonia).
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-

1 from left to right: Monosiphonic part of stem with three hydrothecae; part of monosiphonic
stem with hydrotheca and gonotheca (drawn from slide; gonotheca slightly compressed); scale
bars 0.5 mm; from CALDER & VERVOORT (1998).

References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
VERVOORT W. (1972) Zool. Verh., Leiden 120: 3-247.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 61


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Cnidaria, Hydrozoa, Leptolida, Leptothecata, Tiarannidae

Stegolaria geniculata ALLMAN, 1888

widest in mid-region, attached to axial tube amongst hydrothe-


Size: Height of colonies up to 10 cm.
cae. Distal part narrowing fairly sharply, curving outwards as a
Morphology: Ramified, irregularly shaped, erect colonies with short tube with circular aperture. Condition of gonophore un-
thick, geniculate, polysiphonic stems, resulting from geniculate known; development of a free medusa phase is not unlikely. Ne-
axial tube with hydrothecae becoming covered by many acces- matocysts: Not described.
sory tubules; ultimate ramifications monosiphonic. Stem basal-
Biology: Attached to solid objects like stones or rocks. Re-
ly with stolonal tubes attaching colony to firm substrate. Hy-
silient colonies frequently without living tissue in deep water
drothecae alternately arranged, in same plane as hydrocaulus
hauls; abraded specimens difficult to recognize. Though this is
and branches, adnate to axial tube and immersed to varying de-
by no means a rare species in deep oceanic waters the structure
gree, curving outwards, gradually widening from base onwards;
of polyp and gonophore still remain unknown because of the
walls smooth, abcauline wall concave, free adcauline wall con-
scarcity of well preserved material. Reproduction and mode of
vex to nearly straight. Operculum a roof-shaped enclosure with
feeding unknown.
two longitudinally pleated valves, suspended in U-shaped em-
bayments of hydrothecal margin. Perisarc firm on stem and Distribution: Mid-Atlantic Ridge: Lucky Strike and Rainbow.
branches, yellowish-brown, thinning out considerably along Deep water of the Atlantic Ocean; probably also in deep water
walls of hydrothecae and becoming almost hyaline distally; dis- of Indian and Pacific Oceans or replaced by closely allied
tal part of many hydrothecae collapsed. No diaphragm; hy- species [Stegolaria operculata (NUTTING, 1905)].
dranths attached to internal wall of hydrotheca by means of cir-
cular collar of tissue. Gonothecae elongated sack-shaped,

2 from left to right: Renovated hydrotheca on


1 from left to right: Monosiphonic top part of colony; polysiphonic part of colony; scale bar 0.25 mm;
terminal hydrotheca; hydrotheca on monosiphonic part branch with empty gonotheca and two damaged
of stem; length of hydrotheca 1.3-1.6 mm; from RAMIL & hydrothecae; scale bar 0.5 mm;
VERVOORT (1992). from CALDER & VERVOORT (1998).

References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 3-65.
RAMIL F. & W. VERVOORT (1992) Zool. Verh., Leiden 277: 3-262.

W. VERVOORT & M. SEGONZAC Denisia 18 (2006): 62


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Cnidaria, Hydrozoa, Siphonophora, Rhodaliidae

Thermopalia taraxaca PUGH, 1983 “Galapagos dandelion“

Size: Height of main body up to 40 mm. Biology: The animals are very obvious structures as they float a
few centimetres above the substrate, and are attached, like
Morphology: Ball-like structure, with an apical, smooth-walled
tethered hot-air balloons, by means of numerous lines (tenta-
float (pneumatophore), pinkish-orange in colour. Immediately
cles).
below this is a corona of flimsy, swimming bells; but these are
only apparent if the animal is actively swimming. The main Distribution: Galapagos Spreading Center and East Pacific
body consists of a mass of orange-coloured feeding polyps (gas- Rise. T. taraxaca or relatives have been observed on the Juan de
trozooids), together with other structures. The specific charac- Fuca Ridge but never collected.
teristics are rather esoteric, and include a smooth-walled au-
rophore; cormidia attached directly to the main body of the
corm; and the absence of gonopalpons.

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; by 4: In situ from Galapagos Spreading Center; by
courtesy of R.R. Hessler. courtesy of R.R. Hessler.

Reference:
PUGH P.R. (1983) Philos. Trans. R. Soc. Lond. 301(1105): 165-300.

P.R. PUGH Denisia 18 (2006): 63


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Cnidaria, Scyphozoa, Stauromedusae, Eleuthrocarpidae

Lucernaria janetae COLLINS & DALY, 2005 “stalked jellyfish”

Size: Calyx diameter to 100 mm, height to 60 mm; peduncle Biology: On basalt at vent periphery or areas of diffuse flow.
height to 60 mm. Many individuals of varying sizes co-occur; may dominate local
fauna. Co-occurs with tubeworm Tevnia, sea anemones
Color: Creamy greenish-white to orangish-white.
(Cyananthea sp.), and vent-associated decapods (LUTZ et al.
Morphology: Extremely large stauromedusan with goblet- 1998; HALANYCH et al. 1999). Aggregations comprised of spec-
shaped calyx and eight equa-distant clusters of capitate second- imens of varying sizes and age stages. Smallest juveniles may be
ary tentacles. Adults have no perradial anchors (primary tenta- attached to basal end of adults.
cles); juveniles may have small, ovoid perradial anchors.
Distribution: East Pacific Rise: 7°S to 21°N. A similar form
was observed at Pacific-Antarctic Rise, 38°S, 2330 m (cruise
PAR5, April 2005).

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 Pacific
1: Aggregation in situ (East Pacific Rise: 9°N, 2540 m); by R. Zierenberg. Rise: 9°N, 2540 m; FMNH 10327).

References:
COLLINS A.G. & M. DALY (2005) Biol. Bull. 208: 221-330.
HALANYCH K.M., TIEGER M., O’MULLAN G.D., LUTZ R. & R.C. VRIJENHOEK (1999) InterRidge News 8: 23-27.
LUTZ R.A., DESBRUYÈRES D., SHANK T.M. & R.C. VRIJENHOEK (1998) Deep-Sea Res. II 45: 329-334.

M. DALY Denisia 18 (2006): 64


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Cnidaria, Anthozoa, Actiniaria

According to the current knowledge, the sea anemones, the study of the internal anatomy and cnidocysts is essential for
with eight presently known species, constitute about 1.6% of a correct identification, sea anemones should be carefully treat-
the total number of identified vent species. Several other mor- ed. If specimens are obtained in good condition infrequent at
photypes have been photographed in situ or are currently under best they can be relaxed adding menthol’s crystals on the sur-
study. Thus the number of sea anemone species will probably face of the water. Fixation is best in a big volume of 10%
increase slowly but continually when taxonomic studies con- buffered formalin for at least a week, then they can be trans-
tinue and new vent field are discovered. ferred to 70% ethanol or maintained in 10% buffered formalin
10%. It is highly recommended to inject a solution of concen-
Although the total number of sea anemone species report-
trated 20-30% formalin into the gastrovascular cavity. This will
ed for these environments is apparently not very significant –
facilitate proper fixation of the internal structures used for his-
compared with other groups like gastropods or polychaetes, for
tological studies.
example – the number of species living in a specific vent field
is not higher that two or three these few species are sometimes For molecular studies, either parts of animals or if dissection
so abundant that they form characteristic fields completely of some tentacles is possible in large animals, fixation in 100%
covered by individuals, sometimes equidistantly separated. alcohol and several changes of the fixation medium or freezing
are recommended. It is very important that the rest of the spec-
Most of the species reported belong to the family Actinos-
imen from which the tentacles or fragments have been removed
tolidae (six species), while only one species each of Hormathi-
is fixed in formalin as described above. Thus, molecular as well
idae and Boloceroididae are known to date. One of the acti-
as traditional taxonomic studies can be carried out using the
nostolid species (Actinostola sp.) is only known at the generic
same voucher specimen. Material entirely frozen or fixed in
level at vents. However, this genus is well known from non-
ethanol is seldom useful for the current taxonomic work.
vent environments. The other five actinostolid species were
originally described from hydrothermal vents and were also
used as types of their monotypic genera. The specific status of
the single hormathiid species is currently under revision.
The term sea anemones it is widely used to refer to soft
hexacorallians of the orders Actiniaria (including Ptycodac-
tiaria), Zoanthidea and Corallimorpharia. Members of the first
two orders have been collected from these chemoautotrophic
environments although, for the moment, only identifications of
actiniarian specimens are available.
The current knowledge of sea anemone species associated
with vents is far from completed, most of the characters used to
identify the different genera being anatomic (histology) or mi-
croscopic (types and sizes of cnidocysts). In addition, several
species 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 video
records a difficult task. At present, molecular studies on vent
sea anemone lack. Such studies as well as studies on reproduc-
tion, feeding, and other biological aspects should be carried out
with the appropriate number of specimens per species in the
next decades.
Sea anemones are usually delicate animals. Also those with 1: Boloceroides daphnae from East Pacific Rise: 17°S,
thick body walls and a cuticle are internally high delicate. As Biospeedo cruise © Ifremer.

P. LÓPEZ-GONZÁLEZ Denisia 18 (2006): 65


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Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae

Actinostola VERRILL, 1883

definitive conclusion. Although in the figure 1 of that paper,


Size: Diameter of the pedal disc, 90 mm; mouth, 70 mm; col-
the name A. callosa is included, it was probably a mistake dur-
umn height, 60 mm; length of contracted tentacles, 11 mm.
ing the editorial process of the manuscript. DOUMENC & VAN
Color: Column, tentacles, pharynx, and mesenteries yellowish PRAËT (1986: 63) also reported the presence of four cycles of
white in situ. tentacles, this number of cycles corresponding to 48 tentacles
(6+6+12+24). However, the number of tentacles seen in the in
Morphology: Column cylindrical, with conspicuous mesogloeal
situ photographs suggest five cycles. Additional material is
tubercles arranged regularly in longitudinal rows; tentacles
needed to study the specific pertinence and other anatomical
arranged in four or five cycles, with an apical orifice, aboral side
details.
of tentacles often thickened at the base; sphincter weak,
mesogloeal, thus tentacles can not be completely retracted; Biology: As other sea anemones, probably feeds on caught prey,
longitudinal musculature of tentacles mesogloeal; mesenteries with a relative wide prey size-range. All other biological aspects
arranged according to Actinostola-rule; oldest cycles of mesen- are unknown. Often observed within the gravel, attached at
teries sterile. the top of lava pillars.
Remarks: DOUMENC & VAN PRAËT (1986) discussed the possi- Distribution: East Pacific Rise: 13°N.
ble specific identity of the single specimen examined without a

2: In situ specimen; cruise


Hot 96 © Ifremer.

1: In situ specimen;
cruise Hope 99 © Ifremer.

Reference:
DOUMENC D. & M. VAN PRAËT (1986) Oceanol. Acta 8: 61-68.

P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 66


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Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae

Cyananthea hydrothermala DOUMENC & VAN PRAËT, 1986

Size: Diameter of pedal disc, 10 mm, height, 35 mm; tentacles, Remarks: This species was insufficiently described to be cor-
ca. 25 mm. rectly diagnosed or compared with other actinostolid genera
and species. The familial pertinence was tentative after the sup-
Color: Whitish in situ, column and tentacles dark brownish,
posed absence of acontia in the single fragment available to
mesenteries pale beige.
study by DOUMENC & VAN PRAËT (1986). Material collected in
Morphology: Column smooth, with distal margin distinctly the type locality should be re-described in order to precise the
marked; tentacles occupying most oral disc, reaching near to taxonomic status of this genus and species.
the mouth; more mesenteries distally than proximally; sphinc-
Biology: As other sea anemones, probably feeds on caught prey,
ter mesogloeal, strong, on the endodermic face of the
with a relative wide prey size-range. All other biological aspects
mesogloeal layer; retractor musculature diffuse and weak; longi-
are unknown. The association with bacterial symbionts has not
tudinal musculature of tentacles ectodermic; radial musculature
been demonstrated. Lives on the walls of the black smokers at
of the oral disc ecto-mesogleal.
temperatures of 12-20°C and on sulphide rocks.
Distribution: East Pacific Rise: 13°N.

1: Two specimens taken onboard from the East Pacific Rise: 13°N, cruise Phare 2: In situ specimen taken from the East Pacific
© Ifremer. Rise: 13°N, among siboglinids Riftia pachyptila
and limpets Lepetodrilus elevatus, cruise Phare
© Ifremer.

Reference:
DOUMENC D. & M. VAN PRAËT (1986) Oceanol. Acta 8: 61-68.

P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 67


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Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae

Maractis rimicarivora FAUTIN & BARBER, 1999

spite of that report, there is no formal morphological descrip-


Size: Diameter of the pedal disc, up to 55 mm; mouth, up to 30
tion of this poorly known species from Mid-Atlantic Ridge ma-
mm; column height, up to 20 mm; length of contracted tenta-
terial. At present, all other material subsequently studied from
cles, up to 25 mm.
the Mid-Atlantic Ridge corresponds with the description of
Color: Column, tentacles, pharynx and mesenteries yellowish Maractis rimicarivora. Although Parasicyonis ingolfi was included
white in situ. in the first edition of the Handbook of deep-sea hydrothermal
vent fauna (DESBRUYÈRES & SEGONZAC 1997), at present, we
Morphology: Column conical in contraction, with transversal
decided not to include this species until new material will be
furrows due to contraction; sphincter mesogloeal, strong; tenta-
studied.
cles about 100 in number, closely placed at the rim; longitudi-
nal musculature of tentacles ectodermal; no mastigophore ne- Biology: As other sea anemones, probably feeds on caught prey,
matocysts in tentacles; mesenteries hexamerously arranged, but with a relative wide prey size-range. This species was reported
not according Actinostola-rule; equal number of mesenteries preying on shrimps Rimicaris exoculata. All other biological as-
distally and proximally, 4 cycles, only the first cycle perfect, all pects are unknown. Observed in peripheral regions of active
stronger mesenteries fertile; parietobasilar musculature only dif- black smokers, attached to the crumbly substratum of oxidized
ferentiate in the stronger cycles. sulfide.
Remarks: SEGONZAC (1992) reported Parasicyonis ingolfi CAR- Distribution: Mid-Atlantic Ridge: TAG, Snake Pit-Elan, and
GREN, 1942 from Snake Pit, Mid-Atlantic Ridge. This species Ashadze 1.
was described before the discovery of hydrothermal vents. De-

1: Specimens taken in situ by the


Russian Polar Marine Expedition at
the site Ashadze, among
chaetopterid 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), mytilid
bivalves (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: 14-16
FAUTIN D.G. & B.R. BARBER (1999) Proc. Biol. Soc. Wash. 112(3): 624-631.
LÓPEZ-GONZÁLEZ P.J., RODRÍGUEZ E., GILI J.-M. & M. SEGONZAC (2003) Zool. Verh. Leiden 345: 215-243.
SEGONZAC M. (1992) C. R. Acad. Sci. Paris, Sér. III 314: 593-600.
SEGONZAC M. (1997) in DESBRUYÈRES D. & M. SEGONZAC (Eds.) Handbook of Deep-sea Hydrothermal Vent Fauna, Ifremer: 31.

P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 68


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Cnidaria, Anthozoa, Actiniaria, Actinostolidae

Marianactis bythios FAUTIN & HESSLER, 1989

Size: In life, typical column length and oral disc diameter about Remark: Another sea-anemone, morphologically similar to M.
125 mm in animal with 50 mm pedal disc diameter; length of bythios and attributed to M. aff. bythios, dominates the periph-
inner tentacles about equal to oral disc diameter. Dimensions of ery of black smokers at the Kairei vent field (Central Indian
contracted specimens 25-30% of expanded ones. Ocean).
Color: White in situ. Biology: High abundance in the immediate vicinity of low tem-
perature vents. No morphological adaptations to this habitat
Morphology: Adherent pedal disc; smooth column; tentacles
detected. Probably carnivorous, feeding on vent shrimps.
about 60 in number tapering, pointed, the outer ones of which
are much shorter than the inner. Mesenteries not arranged ac- Distribution: Mariana Back-Arc Basin.
cording to Actinostola-rule; first cycle perfect; stronger ones ga-
metogenic. Tentacles without basal thickenings, no microbasic
b-mastigophores in tentacles but with microbasic amastigo-
phores; fewer than mesenteries. Marginal sphincter mesogloeal;
longitudinal musculature of tentacles ectodermal; parietobasi-
lar musculature distinct.

1: Specimens taken in situ in the Mariana Back-Arc Basin; 2: Marianactis cf. bythios taken in situ at Central Indian
by courtesy of Daphne Fautin. Ocean, Kairei vent field; in the center, mytilid bivalves
(Bathymodiolus marisindicus); by courtesy of J. Hashimoto
© JAMSTEC.

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.

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Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae

Pacmanactis hashimotoi LÓPEZ-GONZÁLEZ, RODRÍGUEZ & SEGONZAC, 2005

rule, first and second cycles of mesenteries perfect, all stronger


Size: Diameter of the pedal disc, 15 mm; column distally, 30
ones fertile; same number of mesenteries distally and proximal-
mm; column height, 20 mm; length of contracted tentacles, 10
ly; parietobasilar musculature not differentiated.
mm. All measurements from preserved and contracted speci-
mens. Biology: As other sea anemones, probably feeds on caught prey,
with a relative wide prey size-range. All other biological aspects
Color: Preserved material of a whitish color. Living animals are
are unknown. This species has been observed on living mussels
whitish to pinkish, with oral disc light brown.
or rocks, within 10 cm of a crack emitting a transparent fluid at
Morphology: Column smooth, distal part including oral disc ca 40°C. Some anemones have been observed outside the area
wider than mid-column and pedal disc, not divisible into sca- of thermal influence, where isolated mytilid bivalves (Bathy-
pus and scapulus; sphincter distinctly marked distally as a modiolus sp.) and siboglinid tubeworms (Arcovestia ivanovi
prominent circumferential marginal ridge, mesogloeal, relative- SOUTHWARD & GALKIN, 1997) also occurred and where the
ly weak; tentacles about 100 in number, of uniform thickness seawater temperature was 2.8°C.
along entire length, with microbasic b-mastigophores; longitu-
Distribution: Manus Back-Arc Basin: Fields D and E; probably
dinal tentacle and oral disc circular musculature ectodermal,
at Lau Back-Arc Basin.
that of tentacles equally well developed on all sides; mesenter-
ies hexamerously arranged, but not according to Actinostola-

1: Preserved specimen collected from Manus Basin


(cruise BIOACCESS’98), specimen showing partially
protruded 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 cut
longitudinally, showing wide oral disc and mar- 3: Probably the same species on basalt covered by zoantharian, with
ginal ridge formed by the sphincter; galatheid crabs Munidopsis spp.; in situ view from Lau Back-Arc Basin;
from LÓPEZ-GONZÁLEZ et al. (2005). Cruise TUIM7; by courtesy of C.R. Fisher.

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


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Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae

Paranthosactis denhartogi LÓPEZ-GONZÁLEZ, RODRÍGUEZ & SEGONZAC, 2003

tentacles with mastigophore nematocysts; mesenteries hexam-


Size: Diameter of the pedal disc, 35 mm; mouth, 16 mm; col-
erously arranged, but not according Actinostola-rule; same num-
umn height, 15 mm; tentacles, 8 mm length. All measurements
ber of mesenteries distally and proximally, four cycles, only the
from preserved and contracted specimens.
fist cycle perfect, all stronger mesenteries fertile, parietobasilar
Color: Preserved material of a brownish color, the tentacles musculature not differentiate as a separate lamella.
darker. Living animals show a brown crown of tentacles, while
Biology: As other sea anemones, probably feeds on caught prey,
the column is dirty white.
with a relative wide prey size-range. All other biological aspects
Morphology: Column smooth, dome-shaped to cylindrical in are unknown. This species has been observed living among Rif-
preserved state; tentacles about 70 in number, of uniform thick- tia, on sediment where 10 cm depth a temperature of 98°C was
ness along entire length; sphincter distinctly marked as a recorded.
prominent marginal ring, mesogloeal; longitudinal tentacle
Distribution: East Pacific Guaymas Basin.
musculature ectodermal, equally well developed on all sides;

2: In situ view of specimens among


tubeworms Riftia pachyptila; cruise
Guaynaut © Ifremer.

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.

Reference:
LÓPEZ-GONZÁLEZ P.J., RODRÍGUEZ E., GILI J.-M. & M. SEGONZAC (2003) Zool. Verh. Leiden 345: 215-243.

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Cnidaria, Anthozoa, Actiniaria, Boloceroididae

Boloceroides daphneae DALY, in press “giant sea anemone”

Size: Diameter of column 0.1-0.5 m, diameter of tentacle crown Biology: Solitary specimens attached to boulders and cliff faces;
0.5-2.0 m. long, tapering tentacles stream in current or contact sediment,
shed when animal is disturbed. Prey unknown but cnidae ex-
Morphology: Very large, pale pink to deep purple sea anemone
tremely large.
with crown of extremely long, strongly tapering tentacles. Col-
umn with ectodermal longitudinal muscles and numerous large Distribution: East Pacific Rise: 9°N to 17°S; the specimens ob-
(~ 100 µm) spirocysts. Not more tentacles than mesenteries. served in southern areas belong probably to the same species.
Preserved specimens may lack tentacles, having only circular
openings on oral disc where tentacles had been attached.
Remark: Possibly misidentified as a species of Cerianthus in
DESBRUYÈRES & SEGONZAC (1997): accompanying photograph
resembles this species but description does not.

3: Spirocyst from the column ectoderm; scale bar 40 µm.

1: Habitus, in situ, from East Pacific Rise: 17°S; cruise Biospeedo 4: Cross section through the column wall, showing
© Ifremer. longitudinal 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 extruded
through the mouth and most of
the tentacles missing; by M. Daly.

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


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Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Hormathiidae

Chondrophellia cf. coronata (VERRILL, 1883)

Size: Diameter of the pedal disc, 27 mm; column distally, 30 Remarks: Despite of the report of this species at Pacific hy-
mm; column height, 30 mm; length of contracted tentacles, 4 drothermal vent areas, the correspondence of Atlantic and Pa-
mm. All measurements from preserved and contracted speci- cific material attributed to this species is unclear. Chondrophel-
mens. lia coronata is known from the North Atlantic. Pacific material
assigned to this species shows similar cnidae biometry, but is
Color: Column brownish due to the presence of cuticle, tenta-
different in external morphology. This species was also report-
cles orange in living specimens, to pale rose in preserved state.
ed – with doubts – from the coast of Chile. Specimens from hy-
Morphology: Pedal disc sharp, covered by cuticle; column di- drothermal vents agree well with the drawings illustrated by
visible into scapus and scapulus, the former with a cuticle, with DOUMENC & VAN PRAËT (1988: figure 2). A revision of the
12 rows of three high pointed tubercules distally, tubercles not species described in this genus by López-González & Segonzac
clearly isolated from each other, with smaller tubercles between is in progress, and will give more information on the specific
the principal rows; cuticle thick, especially on the tubercles; pertinence of the specimens collected at hydrothermal vents.
sphincter mesogloeal, alveolar; tentacles up to 96 in number,
Biology: As other sea anemones, probably feeds on caught prey,
the last circle can be incomplete; about similar number of
with a relative wide prey size-range. All other biological aspects
mesenteries distally and proximally, 6 pairs of perfect but ster-
are unknown. Very abundant mainly around the Southern East
ile mesenteries; acontia present on oldest mesenteries; third
Pacific Rise sites, settled on sulphide deposits or basalts.
and following cycles of mesenteries without filaments and
acontia; without cinclidies. Distribution: East Pacific Rise: 13°N, 7°S to 32°S.

1: In situ view showing numerous specimens among stalked barnacles (Neolepas n. sp.), ophidiid fish
and galatheid crabs; cruise Biospeedo © Ifremer.

Reference:
DOUMENC D. & M. VAN PRAËT (1986) Oceanol. Acta 8: 61-68.

P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 73


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Mollusca, Solenogastres

The aplacophoran molluscs are represented in the fauna as- Recorded hot vent Solenogastres range between 1.5-6 mm
sociated with the hydrothermal vent biotope by the Solenogas- length and are – characteristically for all aplacophoran mol-
tres (neomeniomorphs); members of the second class Caudo- luscs – covered by small aragonitic sclerites. Because these scle-
foveata (chaetodermomorphs) have not yet been evidenced. rites are important specific as well as supraspecific characters,
Solenogastres show a narrowed body, glide upon their restrict- the collected animals should be preserved in 70% ethanol (or
ed foot (pedal groove), and are carnivorous. They include in formalin and soon transferred to ethanol) to save the scle-
about 245 known species. However, neither these nor the rites for determination. Furthermore, specimens should be pre-
presently three described species from the hot vent areas docu- served as soon as possible after sampling, because internal or-
mented here reflect the true biodiversity. Several new species gans (as important as the sclerites) rapidly undergo histolysis
under description and additional representatives come from the (as was the case in quite a number of specimens from the above
East Pacific Rise 13°N (Genesis, Parigo, Elsa), 9°N (Tica), records).
18°S, and 21°S; and the Mid-Atlantic Ridge, Rainbow.

1: Helicoradomenia sp. 2 from southern East Pacific Rise: 21°S; cruise Biospeedo © Ifremer.

L. SALVINI-PLAWEN Denisia 18 (2006): 75


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Mollusca, Solenogastres, Cavibelonia, Simrothiellidae

Helicoradomenia acredema SCHELTEMA, 2000

Size: Up to 3 mm. Remarks: There is no information concerning the internal soft


organs. Due to the similarities or even possible identities of
Morphology: Solenogastres of cylindric body somewhat taper-
hard parts in different species (two new species are under de-
ing at both ends, posterior end bluntly rounded and somewhat
scription), the conspecifity of specimens far off the type locali-
flattened dorsoventrally; appearance similar to H. juani, but
ty (see distribution) needs to be confirmed according to inter-
shorter and fuzzier. Dominant sclerites slender and up to 190
nal organisation. See also H. juani.
μm long with distal end rounded, in part with distal swelling;
dorsofrontal sensory pit often obvious as transverse slit. Radula Distribution: East Pacific Rise: 21°N (type locality), 17°24’S;
in sheath distinctly bipartite, paired ventral radula sack at end Galapagos Spreading Center.
curved; radula plates with 5-7 denticles, the lateral-most dis-
tinctly longer and close to next one. Paired copulatory stylet
apparatus with two elements each.

1A, B: Holotype in lateral (A) and ventral (B) views (anterior end at left); arrow points to pedal pit; C: Mantle sclerites; main
type elongate and often with distal swelling; D: Two radula plates in dorso-frontal view, above from a right row, below from a
left row; after SCHELTEMA (2000).

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

L. SALVINI-PLAWEN Denisia 18 (2006): 76


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Mollusca, Solenogastres, Cavibelonia, Simrothiellidae

Helicoradomenia bisquama SCHELTEMA, 2000

Size: Up to 3.5 mm. Remarks: Though there is poor information concerning the in-
ternal soft organs, the present species is within Helicoradomenia
Morphology: Solenogastres of cylindric body slightly tapering
(see H. juani) well-defined by the characters of the hard parts.
at anterior end, posterior end truncate, appearance rough and
bumpy. Sclerites up to 135 μm long, most of which broad and Biology: From vent clam (Calyptogena) or vestimentiferan (Rif-
thickest at margins as if formed of two joined spicules, often tia) beds.
with bifurcate distal end. Dorsofrontal sensory pit not obvious.
Distribution: East Pacific Rise: 21°N.
Radula biserial with paired ventral sack, at end spirally en-
rolled; radula plates with nine denticles with increasing length
towards lateral. Paired copulatory stylet apparatus with seven
(3+3+1) elements each.

1A, B: Holotype in lateral (A) and ventral view (B, anterior end at left) with part of respiratory papillae protruding from mantle
cavity; 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 SCHELTEMAL (2000).

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

L. SALVINI-PLAWEN Denisia 18 (2006): 77


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Mollusca, Solenogastres, Cavibelonia, Simrothiellidae

Helicoradomenia juani SCHELTEMA & KUZIRIAN, 1991

Size: Up to 5 mm. Remarks: Helicoradomenia is particularly defined by a biserial


radula with paired ventral sack, by solid, elongate sclerites in
Morphology: Solenogastres of cylindric body somewhat enlarg-
one layer and by a dorsofrontal sensory pit. Specific characters
ing posteriorly, anterior end rounded, posterior end somewhat
refer to detailed shape of hard parts (sclerites, radula plates,
flattened ventroterminally; appearance fuzzy due to one layer of
copulatory stylets) as well as to soft internal organs (configura-
solid, elongate, slightly curved sclerites, distally with blunt
tion of foregut, of accessory genital organs and of pallial cavi-
point, and measuring posteriorly up to 200 μm in length. Dor-
ty). There are other hydrothermal vent representatives under
sofrontal sensory pit obvious as transverse slit. Radula biserial
description belonging to Helicoradomenia and to different gen-
with paired ventral sack, at end spirally enrolled; radula plates
era (see Fig. 2-7).
with 5-6 denticles, the lateral-most distinctly longer and close
to next one. Mouth opening within posterior atrium/vestibu- Biology: Carnivorous, but probably not on Cnidaria.
lum; with multicellular dorsal foregut gland and with short
Distribution: Juan de Fuca Ridge: Endeavour Segment; Explor-
esophagus; one pair of shortly stalked seminal receptacle.
er Ridge; Gorda Ridge.
Paired copulatory stylet apparatus with two elements each.

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 row
in 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


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2: H. cf. acredema, in vivo specimen from East Pacific Rise: 3: Helicoradomenia sp. 1, in vivo specimen from East Pacific Rise:
9°N, site Tica; by M. Bright. 9°N, site Tica; by M. Bright.

4: Helicoradomenia sp. 2, body length 2 mm, in ventral 5: Simrothiellidae gen. et sp. 1, body length 3 mm, lateral view,
view,
w anterior end at left showing slit-like dorsofrontal sen- anterior end at left; mantle in part with dorsal incrustation of or-
sory pit and atrio-buccal area, pedal groove (foot) wide; ange iron oxide; from East Pacific Rise: 13°N, site PP Hot 3 (Elsa);
from East Pacific Rise: 18°S; cruise Biospeedo © Ifremer. cruise Phare © Ifremer.

6: Helicoradomenia sp. 3, body length 2.3 mm, lateral view 7: Two specimens of Simrothiellidae sp. in vivo from North Fiji
(anterior at left), mantle with incrustation by yellow-orange Back-Arc Basin, site White Lady; cruise TUIM06MV (June 2005,
deposits; from East Pacific Rise: 21°S; cruise Biospeedo MBARI) © G. Rouse.
© Ifremer.

Reference:
SCHELTEMA
L A.H. & A.M. KUZIRIAN (1991) Veliger 34: 195-203.

79
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Mollusca, Polyplacophora, Neoloricata, Ischnochitonidae

Thermochiton undocostatus SAITO & OKUTANI, 1990

zoid in the tail valves. Valves and girdle white, exposed surface
Size: Length up to 13 mm.
with rusty brown deposits. Radula ca. 4 mm long in the 13 mm
Morphology: Animal elliptical, little elevated, with a sub-cari- specimen, with 170 densely packed teeth rows.
nated back. Girdle rather broad, with imbricating obliquely set,
Biology: Found together with the other species Lepidochiton
convex, finely ribbed scales, which possess a granular dorsal
tenuidontus among bivalves Bathymodiolus and Calyptogena, si-
edge and with slender and smooth, marginal spiculae. Tegmen-
boglinid tubeworms and shrimps. It is not yet known whether
tum (dorsally visible part of the valves) sculptured with some-
the two species are endemic to vents or they also occur else-
what irregular, concentrically arranged undulatin costae, strong
where.
towards the outer margins and weaker and more irregular near
the apices. Articulamentum (layer beneath tegmentum) at the Distribution: Okinawa Trough: Iheya Ridge.
middle of the intermediate valves with a transverse callus, su-
tural laminae roughly triangular in intermediate valves, trape-

1 left: Dorsal view; top right: Head valve, exterior and interior; bottom: Valve 4, anterior view; after SAITO & OKUTANI (1990).

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

R. VON COSEL Denisia 18 (2006): 80


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Mollusca, Polyplacophora, Neoloricata, Leptochitonidae

Leptochiton tenuidontus SAITO & OKUTANI, 1990

portion of each valve, sutural laminae roughly sub-triangular


Size: Length up to 16 mm.
and widely separated. Radula long, 7.3 mm in the 16 mm spec-
Color: Valves pale buff with some blackish deposits, girdle buff imen, with 193 densely packed teeth rows in the holotype.
in preserved specimens.
Biology: Found together with the other species Thermochiton
Morphology: Animal oblong, moderately elevated, with an undocostatus among bivalves Bathymodiolus and Calyptogena, si-
evenly rounded back. Girdle narrow, with elongate calcareous boglinid tubeworms and shrimps. It is not yet known whether
scales, each ornamented with 7-8 longitudinal ribs, and with the two species are endemic to vents or they also occur else-
long, smooth, needle-like marginal spiculae. Tegmentum (dor- where.
sally visible part of the valves) sculptured with elongated,
Distribution: Okinawa Trough: Iheya Ridge.
close-set granules, arranged in longitudinal series. Articula-
mentum (layer beneath tegmentum) with callus at the middle

1 left: Dorsal view; top right, head valve, exterior and interior; bottom: Valve
V IV, anterior and dorsal
view; after SAITO & OKUTANI
T (1990).

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

R. VON COSEL Denisia 18 (2006): 81


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Mollusca, Gastropoda

Currently about 60 genera and more than 100 species of gas- it is recommended that voucher specimens are deposited in a
tropods have been recognized in hydrothermal communities in recognized scientific museum collection. Many natural history
the Indo-Pacific, Indian and Atlantic Oceans. That makes the museums readily accept such deposits. For morphological work
gastropods the most species-rich group of vent animals thanks to formalin-fixed specimens are preferred, although also ethanol-
all researchers who have sent specimens to taxonomists for iden- preserved specimens can be used. Frozen specimens are usually
tification. The gastropods are probably also the group where too macerated to show details in tentacles and epithelia. As al-
knowledge on distribution and zoogeography is the best. ways when calcareous shells are involved, do not forget to
buffer the formalin, for example with a tea spoon of borax per
Although the present handbook aims to present a fairly
litre 4% formalin.
complete overview with diagnostic information and illustra-
tion, it is important to stress several points: (4) Beginning genetic work has revealed existence of isolat-
ed genotypes or populations within some of the previously rec-
(1) The identification of several genera of small gastropods
ognized species. In some species this is accompanied by morpho-
is difficult
f and is a matter of specialist work.
logical differences and/or geographic separation (e.g. Lepetodrilus
(2) The limpets can be assigned to genus based on shell and fucensis JOHNSON et al., in press). In others (e.g. Alviniconcha hess-
external morphology only, but many can be confusingly similar leri) there are no noticeable differences in morphology and their
on superficial examination by an untrained eye. It is sometimes geographic distribution is not easily explicable. It should be re-
even easier to identify limpets to species than to genus. membered that distribution, dispersal and speciation of deep-sea
(3) For any work deemed to have results of more than tem- animals still is “terra incognita“ and should be explored.
porary importance and where species identification is involved, (5) This manual may give the impression that the fauna is ad-
equately known. However, many vent inhabitants are still known
from a single or very few specimens, and almost every cruise to
the site at 13°N on the East Pacific Rise (probably the best
known gastropod fauna) has resulted in one or a couple of unde-
scribed species; new species are also likely to be discovered by new
expeditions at other “well-known“ and “well-collected“ sites.
Many gastropods and other hard-bodied animals from vent
sites and other reducing environments may have thick crusts of
deposited materials covering the surface. These may be black or
reddish 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 growth
in protected cracks and crevices, whereas exposed areas seem to
be grazed with only a short stub of bacterial filaments in border
zones. Exposed areas have smooth and hard surface, but the
process behind this transition seems unknown. The description
of colours of the gastropod shells and periostracum is based on
shells 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-
1: Ifremeria nautilei from Kilo Moana, Lau Back-Arc Basin, TU- tent be host specific since their appearance differs
f between host
IM 07 cruise © C.R. Fisher. species. They have only been observed in vent environments.

References:
JOHNSON P., VRIJENHOEK R.C. & A. WARÉN (in press) Bio. Bull.
SASAKI T., OKUTANI T. & K. FUJIKURA (2005) Venus 64: 87-133.

A. WARÉN Denisia 18 (2006): 82


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Mollusca, Gastropoda, Patelligastropoda, Lepetopsoidea, Neolepetopsidae

Eulepetopsis vitrea MCLEAN, 1990 “translucent limpets”

Size: Shell length up to 17 mm. Biology: Specimens have been collected on the basalt substra-
tum and on the mussel Bathymodiolus. Genus endemic to vents.
Morphology: Shell oblong, very flat; apex one-quarter shell
Larval development lecithotrophic with planktonic dispersal
length from anterior end; shell smooth on superficial examina-
stage.
tion, with fine radial striae on closer inspection. Shells im-
mersed in water or ethanol nearly transparent, dried shells Distribution: East Pacific Rise: 21°N to 17°S; Galapagos
white; interior with metallic sheen when viewed at angle. No Spreading Center.
operculum. No appendage along foot margin
Remarks: An unnamed species known from the Kairei Vent
Field (Indian Ocean).

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

References:
FRETTER V. (1990) J. Zool. Lond. 222: 529-555.
MCLEAN J. (1990) J. Zool. Lond. 222: 485-528 [503].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [123].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 83


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Mollusca, Gastropoda, Patelligastropoda, Lepetopsoidea, Neolepetopsidae

Neolepetopsis MCLEAN, 1990 “symmetrical limpets”

Species Distribution
N. densata MCLEAN, 1990 East Pacific Rise: 12-13°N, Galapagos Spreading Center
N. gordensis MCLEAN, 1990 Gorda Ridge: 41°N, S Gulf of California: 20°N; possibly also off Peru: 5°S
N. occulta MCLEAN, 1990 East Pacific Rise: 21°N
N. verruca MCLEAN, 1990 East Pacific Rise: 21°N

Size: Shell length up to 4-7.5 mm (varies with species). Biology: Genus endemic to vents and seeps; some species have
been collected on inactive chimneys devoid of other megafau-
Morphology: Shells oblong, depressed; apex 1/3 to 2/5 shell
nal species. Larval development lecitotrophic with planktonic
length from anterior end; sculpture of strong beads produced at
dispersal stage. As in Paralepetopsis, species demarcation based
intersections of radial and concentric ribs. Species differ in de-
on morphology is close to impossible since the species are fea-
tails of sculpture. No operculum. No appendage on sides of foot.
ture-less, variable and often badly corroded (except N. densa-
Remarks: Demarcation of genus against Paralepetopsis uncer- ta).
tain.

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

References:
FRETTER V. (1990) J. Zool Lond. 222: 529-555.
MCLEAN J. (1990) J. Zool. Lond. 222: 485-528 [492, 496].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [122-123]

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 84


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Mollusca, Gastropoda, Patelligastropoda, Lepetopsoidea, Neolepetopsidae

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

ronments in the Atlantic and eastern Pacific and on whale


Size: Shell length up to 17 mm.
skeletons off California. They are difficult to identify due to
Color: Shell semi-transparent, white or slightly brown; interior their feature-less external morphology and often corroded
whitish. shells. Several species adjust the shell to the substrate with a
slender outline and concave shell base, when living on tubes.
Morphology: Shell oblong, sturdy, depressed, without trace of
coiling; apex one-third shell length from anterior end, always Biology: Specimens have been collected on and among Bathy-
worn; shell with rough surface, numerous radial ribs. No oper- modiolus. Intestine filled with orange-brown matter of granular
culum. No remarkable appendage or extension on head or foot structure with nematode and ciliate fragments. Larval develop-
margin. Eyes absent. ment lecithotrophic with planktonic dispersal stage
Remarks: Demarcation of genus against Neolepetopsis uncer- Distribution: Mid-Atlantic Ridge: Lucky Strike.
tain. Other species of Paralepetopsis are common in seep envi-

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

Reference:
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [123-125].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 85


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Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Lepetodrilidae

Clypeosectus MCLEAN, 1989 “slit-limpets”

Species Distribution
C curvus MCLEAN, 1989
C. Explorer and Juan de Fuca Ridges
C. delectus MCLEAN, 1989 East Pacific Rise: 13°N and 21°N, Galapagos Spreading Center

Biology: Found with siboglinid tubeworms, details of habitat


Size: Shell length up to 5-8 mm (varies with species).
not known. Genus endemic to vents.
Morphology: Shells of limpet form with oblique, elongate slit;
sculptured by fine radiating ribs. Slit open at margin. No oper-
culum. Eyes absent.

1: C. delectus; top: Exteri-


or and interior view; bot-
tom left, same specimen,
lateral view; by A. Le
Goff © MNHN; bottom
right: Another, inclined
view; P. Briand © Ifremer.

2: Larvae; by courtesy
of L. Mullineaux.

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].
WARÉN A. & P. BOUCHET (2001) Veliger 44 (2): 116-231 [155].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 86


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Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Lepetodrilidae

Gorgoleptis MCLEAN, 1988 “dimorphic limpets”

Species Distribution
G. emarginatus MCLEAN, 1988 East Pacific Rise: 9-21°N
G. patulus MCLEAN, 1988 East Pacific Rise: 13°N, Galapagos Spreading Center
G. spiralis MCLEAN, 1988 East Pacific Rise: 13°N

Size: Shell length up to 3-9 mm (varies with species). Biology: Details of habitat unknown. Genus endemic to vents.
Larval development lecithotrophic with planktonic dispersal
Morphology: Shells depressed, ear-shaped, with a distinctly
stage
coiled initial whorl; sculpture of beaded or imbricate radial ribs.
Operculum small, not closing the shell. Male with penis formed
by expansion of snout on left side. Five pairs of long epipodial
tentacles.

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

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].
WARÉN A. & P. BOUCHET Veliger 44(2): 116-231 [154-155].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 87


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Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Lepetodrilidae

Lepetodrilus MCLEAN, 1988 “dimorphic limpets”

Species Distribution
L. atlanticus WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: 38-23°N
L. corrugatus MCLEAN, 1993 Juan de Fuca Ridge
L. cristatus MCLEAN, 1988 East Pacific Rise: 13°, 21°N, Galapagos Spreading Center
L. elevatus MCLEAN, 1988 East Pacific Rise: 21°N-17°S, Galapagos Spreading Center, North Fiji Basin,
Lau Basin, Mariana Basin
L. fucensis MCLEAN, 1988 Juan de Fuca Ridge
L. galriftensis MCLEAN, 1988 East Pacific Rise: 9°N, Galapagos Spreading Center
L. guaymasensis MCLEAN, 1988 Guaymas Basin
L. japonicus OKUTANI, FUJIKURA & SASAKI, 1993 Okinawa Basin
L. nux OKUTANI, FUJIKURA & SASAKI, 1993 Okinawa Basin
L. ovalis MCLEAN, 1988 East Pacific Rise: 21°N-17°S, Galapagos Spreading Center
L. pustulosus MCLEAN, 1988 East Pacific Rise: 21°N-17°S, Galapagos Spreading Center
L. schrolli BECK, 1993 Manus Basin
L. tevnianus MCLEAN, 1991 East Pacific Rise: 11°N
Further species Indian Ocean hydrothermal vents

boglinid tubes that there is no doubt that a primary habitat of


Size: Shell length up to 6-20 mm (varies with species).
these limpets is directly on Riftia and Tevnia
T . Association of
Morphology: Shells of limpet form; coiling distinct to indis- chemoautotrophic bacteria with the gill confirmed for Lepeto-
tinct. Apex posterior, lower than highest shell elevation, slight- drilus fucensis, which has been found in densities of 400000 m-2.
ly to strongly projecting. Species differ by shell proportions and The genus occurs in vents and seeps. Development with free-
sculpture. No operculum. Male usually with penis near base of swimming lecithotrophic larvae with planktonic dispersal
right cephalic tentacle. Three pairs of short epipodial tentacles. stage.
Biology: At East Pacific Rise sites, specimens of L. elevatus Remark: Differ
f ent genetic types, probably species, occur, espe-
have been collected in such abundance from washings of si- cially of the forms similar to L. elevatus and L. schrolli.

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.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 88–90


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2: L. cristatus, on an active chimney, near an alvinellid colony; East Pacific Rise: 13°N; cruise Phare. P. Briand © Ifremer.

3: L. fucensis, among tubeworms Ridgeia picesae, Juan de Fuca Ridge; by courtesy of K. Juniper.

89
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4 top: L. elevatus; middle: L. ovalis; bottom: L. pustulosus. All with exterior, interior and lateral view; by R. von Cosel & A. Le
Goff; in situ views, limpets on tubes of Riftia pachyptila © Ifremer/Phare.

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.
WARÉN A. & P.P BOUCHET (2001) Veliger 44(2): 116-231 [143-154].

90
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Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Lepetodrilidae

Pseudorimula MCLEAN, 1989 “slit-limpets”

Species Distribution
P marianae MCLEAN, 1989
P. Mariana Back-Arc Basin
P. midatlantica MCLEAN, 1992 Mid-Atlantic Ridge: 38-15°N

Size: Shell length up to 5-8 mm (varies with species). Biology: On rocks. P. midatlantica also occurs on Bathymodiolus
mussels. Genus endemic to vents. Development without plank-
Morphology: Shells of limpet form with oblique, elongate slit;
totrophic larvae.
sculptured by fine radiating ribs. Slit closed at margin. No op-
erculum. Eyes absent.

1: P. midatlantica; top: Dorsal, lateral and ventral view, no precision of origin; by R. von Cosel & A. Le Goff; middle: Specimen
from Logatchev (Mid-Atlantic Ridge), dorsal and ventral view; bottom: specimen from Snake Pit; by A. Warén.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 91–92


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2: P. midatlantica; by P. Briand.

References:
MCLEAN J. (1989) Nat. Hist. Mus. Los Angeles Cty., Contrib. Sci. 407: 1-29 [22].
MCLEAN J. (1992) Nautilus 106: 115-118 [116].
WARÉN A. & P.
P BOUCHET (2001) Veliger 44 (2): 116-231 [155-157].

92
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Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Pyropeltidae

Pyropelta MCLEAN & HASZPRUNAR, 1987

Species Distribution
P bohlei BECK, 1996
P. Lihir Volcano, Edison Seamount, West Pacific
P. corymba MCLEAN & HASZPRUNAR, 1987 Oregon Margin: 45°N, Guaymas Basin
P. musaica MCLEAN & HASZPRUNAR, 1987 Juan de Fuca Ridge: 45°56’N, off California: 36-33°N, Jalisco Block: 20°N
P yamato SASAKI, OKUTANI & FUJIKURA, 2003
P. Izu-Ogasawara Arc

washings of vestimentiferan tubes. Pyropelta corymba lives on


Size: Shell length up to 5 mm.
shells of Provanna spp., grazing bacteria. The genus occurs on
Morphology: The family is defined by anatomical characters vents, seeps and also on whale bones. Larval development
and the shells are rather featureless, cap-shaped, with central lecithotrophic with planktonic dispersal stage. The featureless
apex; sculpture unknown, exterior surface deteriorated. No op- shell, always badly corroded, makes identification very uncer-
erculum. Foot with a pair of posterior epipodial tentacles. tain.
Biology: Specimens have been collected on sulphide crust in
the vents and on surrounding rocks, and they are absent in

1: P.
P corymba
r ; top left: Inclined and dorsal view of a specimen; right: The same specimen; SEM by A. Warén; bottom left: Lateral
view of another specimen, from Guaymas; by P. Briand © Ifremer; bottom right: Another specimen; by J. McLean.

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].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [125-129].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 93


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Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Skeneidae

Bruceiella globulus WARÉN & BOUCHET, 1993

Size: Shell diameter up to 2.3 mm. Remark: Another species of this genus, B. athlia WARÉN &
BOUCHET, 2001, was described from cold seeps of the Aleutian
Morphology: Shells globular, almost as high as broad, with low
Trench, Shumagin Site, 54°18’N, 157°12’W, 4808 m.
spire, rather robust, surface almost smooth. Umbilicus deep and
wide. Whorls rounded, with a deep suture, aperture circular, al- Biology: At hydrothermal vents.
most or completely detached from the following whorl. Proto-
Distribution: North Fiji and Lau Back-Arc Basins.
conch with spiral sculpture, about 0.65 whorls, with a diameter
of about 340 μm. Operculum round, multispiral and thin.

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, specimen
4, apical view,
w 1.9 mm; both cruise Starmer II North Fiji Back-Arc Basin; after WARÉN & BOUCHET (1993).

References:
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231.

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Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Skeneidae

Fucaria WARÉN & BOUCHET, 1993

Species Distribution
F mystax WARÉN & BOUCHET, 2001 S of Lihir, Edison Seamount, West Pacific
F. striata WARÉN & BOUCHET, 1993 Juan de Fuca Ridge

erculum. No eyes. Right neck-lobe fused with eye-lobe and


Size: Diameter up to 5.8 mm (F. mystax) and 10.6 mm (F. striata).
equipped with marginal tentacles.
Color: White, greenish or brownish.
Biology: Species of Fucaria occur only in seeps and vents. Food
Morphology: Shells turbinate, about as high as broad, rather consists of the detritus layer on the bottom (stomach contents
sturdy (only known with top of the shell eroded), almost closed consist of mineral particles, sponge spicules and radiolarian
umbilicus; operculum with central nucleus. Surface smooth (F. fragments in a matrix of mucus and detritus). Larval develop-
mystax) or sculptured by spiral lirae and groves (F. striata). ment lecithotrophic with planktonic dispersal stage.
Aperture rounded, completely closed by horny, multispiral op-

1: View of
different
sides of
several
specimens;
by R. von
Cosel.

References:
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [15-17].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [136-137].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 95


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Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Skeneidae

Leptogyra inflata WARÉN & BOUCHET, 1993

2.25 whorls, with about nine low and indistinct spiral ribs and
Size: Shell diameter up to 1.32 mm.
axial growth lines. Periostracum thin and transparent. Opercu-
Morphology: Shell very small, skeneiform, broader than high, lum thin, multispiral, brownish.
with evenly rounded whorls. Umbilicus deep and broad.
Biology: Only known from hydrothermal vent sites.
Whorls with a deep suture. Protoconch with 0.6 whorls, regu-
larly coiled, the initial part with an irregular net sculpture, the Distribution: Lau Back-Arc Basin.
remaining half smooth; diameter 200 μm. Teleoconch with

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 WARÉN & BOUCHET (1993).

3: Umbilicus; after WARÉN & BOUCHET (1993). 2: Protoconch; after WARÉN & BOUCHET (1993).

Reference:
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.

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Mollusca, Gastropoda, Vetigastropoda, Cocculiniformia, Skeneidae

Protolira WARÉN & BOUCHET, 1993

Species Distribution
P thorvaldssoni WARÉN, 1996
P. Mid-Atlantic Ridge: Snake Pit, to Iceland
P. valvatoides WARÉN & BOUCHET, 1993 Mid-Atlantic Ridge: Menez Gwen and Lucky Strike

Size: Shell diameter up to 4.2 mm. Biology: Found among Bathymodiolus, on sediment and on
rocks, Protolira thorvaldssoni also on whale bones, from which it
Morphology: Shells small, fragile, turbinate, about as high as
was originally described. Food consists of the detrital surface
broad or broader than high, with spirally arranged micro-tuber-
layer on the bottom (stomach contents consist of a mixture of
cles on the protoconch (spire almost always corroded) and an
organic material and mineral particles, scattered sponge
almost smooth, globular shell with open umbilicus and a deep
spicules, polychaete bristles, diatoms and crustacean frag-
suture. Aperture circular, completely closed by horny opercu-
ments). Larval development lecithotrophic with planktonic
lum. Shell whitish, greenish, brownish; surface covered by
dispersal stage.
thick periostracum and mineral deposits. Right neck-lobe con-
tinuous with eye-lobe. Right anterior corner of propodium
drawn out into a spirally coiled tentacle.

1 top: P. thorvaldssoni, apertural,


abapertural and lateral view;
by R. von Cosel & A. Le Goff;
right: P. valvatoides; by P. Briand.

References:
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [22-26].
WARÉN A. & P. BOUCHET (2001) Veliger 44 (2): 116-231 [138-139].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 97


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Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Sutilizonidae

Sutilizona MCLEAN, 1989 “slit-limpet”

Species Distribution
S. pterodon WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: Snake Pit,
S. theca MCLEAN, 1989 East Pacific Rise: 12-13°N
S. tunnicliffae WARÉN & BOUCHET, 2001 Juan de Fuca Ridge: Endeavour Segment

Size: Shell length up to 2.4 mm. Biology: Details of habitat not known. Genus endemic to
vents. Larval development lecithotrophic with planktonic dis-
Morphology: Shells fragile, limpet-like with long, moderately
persal stage.
oblique 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. Operculum
small and thin, vestigial, multispiral with central nucleus.

1: S. pterodon; two specimens, dorsal views and apertural view (SEM); by A. Warén.

References:
MCLEAN J. (1989) Nat. Hist. Mus. Los Angeles Cty., Contrib. Sci. 407: 1-29 [15].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [141-143].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 98


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Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Sutilizonidae

Temnocinclis euripes MCLEAN, 1989 “slit-limpet”

Size: Shell length up to 4 mm. Biology: Details of habitat not known. Larval development
lecithotrophic with planktonic dispersal stage.
Morphology: Coiled limpet with long, moderately oblique slit;
reticulate sculpture formed by intersection of fine radiating ribs Distribution: Juan de Fuca Ridge.
and concentric ridges. Operculum small, vestigial.

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


P. Briand.

2: Specimen, dorsal view;


by R. von Cosel & P. Briand.

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].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [143].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 99


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Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Sutilizonidae

Temnozaga parilis MCLEAN, 1989

Size: Shell length up to 4 mm. Biology: Details of habitat not known. Monotypic genus en-
demic in vents. Larval development lecithotrophic with plank-
Morphology: Symmetrical limpet with median, oblique, elon-
tonic dispersal stage.
gate slit; sculptured by strong radiating ribs bearing raised
scales. Operculum small, vestigial. Distribution: East Pacific Rise: 21-13°N.

1: Habitus; by A. Warén & P


P. Briand.

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.

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Mollusca, Gastropoda, Vetigastropoda, Trochoidea, Chilodontidae

Bathymargarites symplector WARÉN & BOUCHET, 1989

Size: Shell up to 11 mm. Biology: Specimens have been obtained from rubble samples
and washings of Riftia and Calyptogena. Monotypic genus en-
Color: Off-white.
demic to vents. Food consists of the detrital surface layer on the
Morphology: Shell turbinate, about as high as broad, almost bottom. Stomach content consists of mineral particles, sponge
smooth, without umbilicus. Umbilical area and columella cov- spicules, polychaete bristles, crustacean and diatom fragments,
ered by a solid callus in adult specimens. Interior with a well de- radiolarian tests in a matrix of undefined organic matter. Larval
veloped nacreous layer. Eyes at base of cephalic tentacles, but development lecithotrophic with planktonic dispersal stage.
pigment spot of variable shape, sometimes double or absent.
Distribution: East Pacific Rise: 13°N and 21°N.
Male with left neck lobe modified to function as a penis.

1 top: Specimen 1, apertural, abapertural and lateral view by A. Le Goff © MNHN;


bottom: Specimen 2, inclined dorsal view,
w apertural view,w inclined basal view;
by P. Briand © Ifremer.

2: Larva; by courtesy
of L. Mullineaux.

References:
WARÉN A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [87-94].
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [11-13].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 101


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Mollusca, Gastropoda, Vetigastropoda, Trochoidea, Trochidae

Helicrenion reticulatum WARÉN & BOUCHET, 1993

set granular incremental axial lines. Animal not known, radula


Size: Shell diameter up to 1.56 mm.
with unusual few marginal teeth for a neomphalid radula type.
Morphology: Shell very small, skeneiform, like a vitrinellid, de-
Biology: Known only from hydrothermal vent sites.
pressed, fragile, with an almost smooth surface. Umbilicus
widely open, without any basal area or spiral sculpture. Whorls Distribution: Only known from Lau Back-Arc Basin: Hine Hi-
with a deep suture. Protoconch with 0.6 whorls, the initial part na vent field.
with sharp spiral lines and irregularly spaced axial lines, form-
ing a large-meshed net sculpture, later half smooth. Teleoconch
with about 1.5 rapidly increasing whorls, with very fine, close-

1 top: holotype apertural view,


w 1.56 mm; bottom: paratype 1.10 mm, apical and basal view;
Lau Basin; from WARÉN & BOUCHET (1993).

2: Protoconch showing reticulated sculpture; Lau 3: Protoconch showing umbilicus; Lau Basin;
Basin; from WARÉN & BOUCHET (1993). from WARÉN & BOUCHET (1993).

Reference:
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 102


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Mollusca, Gastropoda, Vetigastropoda, Trochoidea, Trochidae

Vetulonia phalcata WARÉN & BOUCHET, 1993

Size: Shell diameter up to 1.24 mm. Biology: At vent sites; it is likely that the species feeds on su-
perficial detritus and bacterial film on the bottom, like many
Morphology: Shell trochiform, thin and fragile, with regular
other deep water archaegastropods.
and sharp axial ribs parallel to the outer lip, ribs basally distin-
cly flexuous. Deep, narrow umbilicus which has an indistinct, Distribution: North Fiji Back-Arc Basin.
steeply ascending spiral rib. Whorls slightly depressed, with a
deep suture. Surface between the axial ribs almost smooth. Pro-
toconch with 0.65 whorls, strongly corroded, with a diameter of
at least 200 μm; teleoconch with 2.45 whorls; operculum
round, multispiral, thin and transparent. Animal not known.

1: Holotype, 1.06 mm; left apical view; right apertural view: collected during the cruise Starmer II; after WARÉN & BOUCHET (1993).

Reference:
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.

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Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Cyathermia naticoides WARÉN & BOUCHET, 1989

Size: Shell diameter up to 7 mm. Biology: Specimens have been found in abundance in washings
of Riftia tubes, more rarely with Alvinella tubes. Monotypic
Color: White.
genus endemic to vents. Probably a filter-feeder, possibly in
Morphology: Shell globular, regularly coiled, without sculpture. combination with cleaning the worms’ tubes of bacterial
Adults with deep notch in the lower part of the outer lip. Very growth. Larval development lecithotrophic with planktonic
large bipectinate gill. Left cephalic tentacle modified to a penis. dispersal stage.
Distribution: East Pacific Rise: 9-21°N.

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

2: Larva; by courtesy
of L. Mullineaux.

References:
WARÉN A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [69-70].
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22 (1): 1-90 [33].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [158].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 104


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Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Lacunoides WARÉN & BOUCHET, 1989

Species Distribution bilicus not present or indistinct. Whorls round with a deep su-
L. exquisitus Galapagos Spreading Center ture. Protoconch with diameter of 160 μm (L. exquisitus ) or
L. vitreus Juan de Fuca Ridge: 180 μm (L. vitreus), about 0.5 whorls, the initial part with ir-
Axial Seamount – Ashes vent field regular net sculpture, later half smooth. Teleoconch with 2.25
(L. exquisitus ) or about 2 (L. vitreus) round whorls. Operculum
Size: Shell diameter up to 2.5 mm. multispiral, thin and without color, slightly larger than the
Morphology: Shells small, thin and fragile, colourless and aperture.
transparent, globular, with low, depressed spire, large aperture Biology: Near hydrothermal vents on mussel beds or on other
and rapidly increasing diameter of the whorls. Surface with hard substrate.
fine, dense, sometimes slightly irregularly spaced, sharp axial
lines or low lamellae and still finer, dense spiral striations. Um-

1: L. exquisitus, top specimen 1, apertural 3: L. vitreus, top holotype, apertural view,


view,
w 3.4 mm; bottom specimen 2, apical 2.4 mm; bottom paratype, apical view, w 2.2
view, 2.3 mm; from WARÉN & BOUCHET (1989). mm; from WARÉN & BOUCHET (1989).

2: L. exquisitus, close-up of protoconch of 4: L. vitreus: close-up of protoconch of


the same specimen; from WARÉN & BOUCHET the same paratype; from WARÉN &
(1989). BOUCHET (1989).

References:
WARÉN A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102.
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 105


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Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Melanodrymia HICKMAN, 1984

Species Distribution
M. aurantiaca HICKMAN, 1984 East Pacific Rise: 13°N, 21°N-17°S
M. brightae WARÉN & BOUCHET, 1993 Juan de Fuca Ridge: Endeavour segment
M. galeronae WARÉN & BOUCHET, 2001 East Pacific Rise: 13°N
Melanodrymia sp. “rust covered“ East Pacific Rise: 13°N

Size: Shell diameter up to 3.5 mm. Biology: Genus with three species endemic in vents. The
species are locally common in washings of Riftia, Ridgeia, Ca-
Color: Rusty orange (M. aurantiaca) or whitish (M. brightae).
lyptogena and Alvinella. Genus endemic to vents. Food consists
Morphology: Shells depressed (M. aurantiaca, M. brightae) or of the detrital surface layer of the bottom. Larval development
higher than broad (M. galeronae) with one or two strong pe- lecithotrophic with planktonic dispersal stage.
ripheral keels. Umbilicus open. Surface of shell above and be-
low keel covered by raised riblets. Most specimens have a thick
mineral crust.

1: Top:
T M. aurantiaca; by R. von Cosel; middle: M. aurantiaca; 2: M. aurantiaca;
by P. Briand © Ifremer; bottom: M. galeronae; by A. Warén. by P. Briand © Ifremer.

References:
HASZPRUNAR G. (1989) Acta Zool. 70: 175-186.
HICKMAN C.S. (1984) Zool. Scr. 13: 19-25 [19-20].
WARÉN A. & P. BOUCHET 1989) Zool. Scr. 18: 67-102 [75].
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22(1): 1-90 [41-44].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [158-161].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 106


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Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Neomphalus fretterae MCLEAN, 1981

Size: Shell diameter up to 30 mm. Biology: Occurs locally in dense aggregations on the walls of
the vents. The Neomphalidae are endemic to vents. Probably
Morphology: Shell cap-shaped with coiled apical whorl, apex
filter feeder. Larval development lecithotrophic with plankton-
subcentral, sculptured with fine radiating ribs. No trace of op-
ic dispersal stage.
erculum. Head with long neck, no snout, left cephalic tentacle
enlarged. Distribution: Common at Galapagos Spreading Center, rare at
the East Pacific Rise: 9-21°N.

2: Habitus; by R. von Cosel & P. Briand.

3: In situ specimens, from East Pacific


1: Habitus; by R. von Cosel & P. Briand. Rise: 13°N © Ifremer.

References:
FRETTER V., GRAHAM A. & J. MCLEAN (1981) Malacologia 21: 337-361.
MCLEAN J. (1981) Malacologia 21: 291-336 [294].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [162].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 107


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Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Pachydermia WARÉN & BOUCHET, 1989

Species Distribution
P laevis WARÉN & BOUCHET, 1989
P. East Pacific Rise: 21°N -17°S
P. sculpta WARÉN & BOUCHET, 1993 North Fiji and Lau Back-Arc Basins

Size: Shell height up to 4.6 mm. Biology: Many specimens found in washings of tubes of Alvinel-
la and siboglinids but both species seems to occur also on other
Morphology: Shell small and rather fragile, regularly coiled, up
substrates. Genus endemic to vents. Stomach contents indicate
to 3.5 whorls, with part of the body whorl disjunct and with cir-
detritus feeding. Larval development lecithotrophic with
cular aperture. Surface of protoconch net-sculptured, teleo-
planktonic dispersal stage.
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 mineral
deposits. Operculum multispiral, closing the aperture com-
pletely.

1 top left to right: Apertural, abapertural and lateral view of P. laevis; by A. Le Goff © MNHN; bottom: another
specimen; by P P. Briand © Ifremer.

References:
WARÉN A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [75-80].
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [40-41].
WARÉN A. & P. BOUCHET (2001) Veliger 44 (2): 116-231 [161-162].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 108


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Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Planorbidella WARÉN & BOUCHET, 1989

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

erculum multispiral, completely closing the aperture. Speci-


Size: Shell diameter up to 5.1 mm (P. planispira) and 1.56 mm
mens may be partly encrusted with mineral deposits.
(P. depressa).
Biology: Found in washings of tubes of Alvinella and siboglin-
Morphology: Shells flat-spired, regularly coiled, aperture al-
ids. Genus endemic to vents. Stomach contained only unde-
most circular and strongly prosocline, umbilicus very wide and
fined organic material. Larval development lecithotrophic with
deep. Surface smooth (P. planispira) or with strong, oblique ax-
planktonic dispersal stage.
ial ribs and fine spiral cords (P. depressa), protoconch with fine-
mesh net sculpture. Last part of the body whorl detached. Op-

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

References:
WARÉN A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [81-82].
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [35-39].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [162].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 109


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Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Symmetromphalus MCLEAN, 1990

Species Distribution male enormously distended, bearing a deep sperm groove con-
S. hageni BECK, 1992 Manus Back-Arc Basin nected to neck groove. Gill large, overlying head.
S. regularis MCLEAN, 1990 Mariana Back-Arc Basin Biology: Symmetrophalus regularis occurs in dense aggregations
Symmetromphalus sp. North Fiji and Lau Back-Arc Basins on the walls of vents; the species at Lau and Fiji lives on mus-
sels and on the subsutural ramp of Ifremeria nautilei. Endemic to
Size: Shell length up to 14-21 mm (varies with species). vents. Sexually dimorphic, with males much smaller. Feeding
Morphology: Shells of limpet form with coiled apical whorl, biology not known. Pallial furrow and especially epipodial ten-
sculptured by finely beaded radial ribs. Operculum small, ves- tacles with growth of large filiform bacteria. Larval develop-
tigial. Head with long neck, no snout, left cephalic tentacle of ment lecithotrophic with planktonic dispersal stage.

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


P. Briand.

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

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 110


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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Ctenopelta porifera WARÉN & BOUCHET, 1993

Size: Shell length up to 10 mm. Biology: Genus with a single species endemic to vents. Speci-
mens have been rarely collected in washings of tubes of Tevnia
Morphology: Shell depressed, ear-shaped, sculptured with a
and Riftia or sulphide crusts. The setae of the foot and the
dozen spirally arranged rows of hollow conical tubercules con-
strange tubular processes on the back of the shell may be in-
nected via pore to the interior of the shell. Surrounding the tu-
volved in some kind of symbiosis with chemosynthetic organ-
bercules are soft, tubular hollow appendages of organic materi-
isms (?). Sexes separate, females larger than males. Larval de-
al. No eyes. Sides of foot and epipodium finely setose; posterior
velopment lecithotrophic with planktonic dispersal stage.
part of visceral mass carrying warts corresponding to the pores
in the shell. Distribution: East Pacific Rise: 13°N.

1: Several specimens; by R. von Cosel.

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

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 111


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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Depressigyra globulus WARÉN & BOUCHET, 1989

Size: Shell diameter up to 5.4 mm. Biology: On vestimentiferan tubes, extremely common. Occa-
sionally, near acidic outflows, the calcareous layer may be dis-
Color: Shell greenish to brownish, interior not nacreous. Pe-
solved and the living animal is surrounded only by the strong
riostracum thick, brownish green.
periostracum. Monotypic genus endemic to vents. Stomach
Morphology: Shell broader than high with rather large body contains amorphous organic matter. Larval development
whorl, about three whorls; aperture subcircular with an indis- lecithotrophic with planktonic dispersal stage.
tinct shallow basal notch. Umbilicus reduced to a small chink,
Distribution: Juan de Fuca Ridge.
suture deep. Protoconch strongly ridged, teleoconch with irreg-
ular incremental lines, otherwise smooth. Operculum multispi-
ral, densely coiled, with central nucleus. Animal with tentacles
of even size in both sexes and a snout of approximately even
width.

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

References:
BATES
A B. & V.
V TUNNICLIFFE (2006) Mar. Ecol. Prog. Ser. 305: 1-15.
WARÉN A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [80-81].
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [35].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [173].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 112


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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Echinopelta fistulosa MCLEAN, 1989 “tapersnout limpet”

Size: Shell length up to 9 mm. Biology: Common on outer face of active black smokers.
Monotypic genus endemic to vents. Larval development
Morphology: Shell of limpet form, apex close to posterior mar-
lecithotrophic with planktonic dispersal stage.
gin but left of center. Sculpture with widely spaced tubular
spines. Periostracum thick. Large specimens coated with rust Distribution: East Pacific Rise: 21°N, 13°N.
coloured iron deposits. No operculum. Mantle edge bearing nu-
merous, crowded and elongate tentacles.

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

References:
FRETTER V. (1989) J. Zool. Lond. 218: 123-169.
MCLEAN J. (1989) Zool. Scr. 18: 49-66 [58-60].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [170].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 113


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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Hirtopelta MCLEAN, 1989

Species Distribution
H. hirta MCLEAN, 1989 East Pacific Rise: 13-21°N
H. tufari BECK, 2002 East Pacific Rise: 21°S

Size: Shell length up to 13 mm. Biology: Genus endemic to vents. Gill and intestine morphol-
ogy indicate that Hirtopelta uses the gill for chemosynthetic
Color: Olive-brown.
purposes. Larval development lecithotrophic with planktonic
Morphology: Shells more or less depressed, ear-shaped, sculp- dispersal stage.
tured by raised, scale-like projections arranged along growth
lines. 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.

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

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].
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [35].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [169].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 114


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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Lirapex WARÉN & BOUCHET, 1989

Species Distribution
L. costellata WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: Lucky Strike, Snake Pit
L. granularis WARÉN & BOUCHET, 1989 East Pacific Rise: 9-21°N
L. humata WARÉN & BOUCHET, 1989 East Pacific Rise: 21°N

posits. Operculum multispiral with central nucleus, completely


Size: Shell height up to 3.4 mm.
closing the aperture. Head with simple tentacles and a snout of
Morphology: Shells very small to small, valvatoid, regularly even width.
coiled, about three whorls, rather sturdy, aperture almost circu-
Biology: On soft bottom and among Bathymodiolus. Genus en-
lar, umbilicus narrow to wide, suture deep, last part of body
demic to vents. Stomach contains amorphous organic material
whorl often detached. Protoconch with sculpture of strong spi-
but rarely sponge spicules, crustacean remains. Larval develop-
ral ridges, teleoconch with axial ribs, variable from species to
ment lecithotrophic with planktonic dispersal stage.
species. Shell whitish to brownish with more or less thick pe-
riostracum; specimens are often encrusted with mineral de-

1 top: Apertural view of two specimens of L. costatus; by A. Warén; bottom: Same species; 2: Larvae; by courtesy
by P.
P Briand. of L. Mullineaux.

References:
WARÉN A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [84-86].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [170-171].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 115


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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Nodopelta MCLEAN, 1989 “tapersnout limpets“

Species Distribution
N. heminoda MCLEAN, 1989 East Pacific Rise: 13°N, 21°N
N. rigneae WARÉN & BOUCHET, 2001 East Pacific Rise: 13°N
N. subnoda MCLEAN, 1989 East Pacific Rise: 13°N

Size: Shell length up to 20 mm. Biology: Closely associated with black smokers, recovered from
aggregations of Alvinella. Genus endemic to East Pacific Rise
Color: Beige to olive-brown.
vents. Larval development lecithotrophic with planktonic dis-
Morphology: Shells limpet-shaped, depressed, apex close to persal stage.
posterior margin but not overhanging it. Sculpture finely
clathrate with scattered imbricate nodes. Periostracum thick.
No eyes, no operculum. Mantle margin with transverse ridges
aligned perpendicular to mantle edge.

1 top: N. heminoda; middle: N. subnoda; bottom: N. rigneae. All with exterior,


r interior and lateral view. T
Top and
middle row by R. von Cosel & A. Le Goff; bottom row by A. Warén.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 116–117


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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; East
Pacific Rise: 13°N, Phare cruise © Ifremer.

References:
FRETTER V. (1989) J. Zool. Lond. 218: 123-169.
MCLEAN J. (1989) Zool. Scr. 18: 49-66 [53-56].
WARÉN A. & P.P BOUCHET (2001) Veliger 44(2): 116-231 [168-169].

117
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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Peltospira MCLEAN, 1989 “tapersnout limpets”

Species Distribution
P delicata MCLEAN, 1989
P. East Pacific Rise: 9-13°N
P. lamellifera WARÉN & BOUCHET, 1989 East Pacific Rise: 13°N
P. operculata MCLEAN, 1989 East Pacific Rise: 9-21°N, 17°S
P smaragdina WARÉN & BOUCHET, 2001
P. Mid Atlantic Ridge: 15-38°N

(P. delicata). Eyes absent. Epipodium bearing club-shaped


Size: Shell length up to 12 mm.
processes of irregular size, along opercular lobe.
Color: Olive-brown.
Biology: Specimens have been collected from washings of
Morphology: Shells depressed, ear-shaped, with a distinctly Alvinella and appear to be closely associated with active smok-
coiled initial whorl; sculptured by numerous concentric lamel- ers. Genus endemic to vents. Larval development lecithotroph-
lae (species differ
f in sculpture). Strong periostracum. Opercu- ic with planktonic dispersal stage.
lum small, not closing the shell (P. operculata) or even absent

1 top: Four specimens of P.


smaragdina; by A. Le Goff
© MNHN; middle: Five specimens
of P. operculata; from these the
upper three specimens by A. Le
Goff © MNHN and the lower two
specimens with lateral and exteri-
or view; by P. Briand © Ifremer;
bottom: Three specimens of P.
delicata, interior,
r lateral and
exterior; by A. Le Goff © MNHN.

References:
FRETTER V. (1989) J. Zool. Lond. 218: 123-169.
MCLEAN J. (1989) Zool. Scr. 18: 49-66 [51-53].
WARÉN A. & P. BOUCHET (1989) Zool. Scr. 18: 67-102 [84-85].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [165-168].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 118


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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Rhynchopelta concentrica MCLEAN, 1989 “tapersnout limpet”

Size: Shell length up to 13 mm. Biology: Associated with the tubes of Riftia. Genus endemic to
vents. Larval development lecithotrophic with planktonic dis-
Color: Yellowish-brownish.
persal stage.
Morphology: Shell of limpet form, apex projecting close to the
Distribution: East Pacific Rise: 21-17°N.
posterior margin. Sculpture of fine concentric ridges and radial
striae. Periostracum thin. No operculum. Epipodium and man-
tle edge simple, without modification.

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

2: Larva; by courtesy
of L. Mullineaux.

References:
FRETTER V. (1989) J. Zool. Lond. 218: 123-169.
MCLEAN J. (1989) Zool. Scr. 18: 49-66 [57-58].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [170].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 119


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Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

“scaly foot gastropod”

Size: Shell length up to 50 mm. Remark: This genus and species is still not formally named, but
its conspicuous morphology and interesting symbiosis makes it
Morphology: Shell globular, ear-shaped, distinctly coiled;
desirable to include this novelty.
sculptured by numerous spiral ribs and growth lines. Perios-
tracum thick, dark olive brown. Operculum absent in adult. Biology: The scaly-foot gastropod harbours thiotrophic γ-pro-
γ
Cephalic tentacles thick, conical without eyes. Epipodium teobacteria in an enormously enlarged oesophageal gland. It is
strongly reduced, consisting of a series of inconspicuous tuber- a sedentary organism firmly attached to rocks at the base of
cles on the side of the foot above the scales. Operculum modi- black smoker chimneys. Genus endemic to vents. Sexes sepa-
fied into several hundred horizontally aligned scales, arranged rate, sperm transfer by spermatophores. Development
in a roof tile fashion along the sides of the foot. Scales covered lecithotrophic, presumably with a planktonic dispersal stage.
by thick layers of quite pure pyrite and greigite (iron sulphides)
Distribution: Indian Ocean: Rodriguez Triple Junction.
deposited in a very uniform way, indicating active participation
by the snail in the process.

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.

References:
WARÉN A., BENGTSSON S., GOFFREDI S.K. & C. VAN DOVER (2003) Science 302: 1007.
GOFFREDI S.K., WARÉN A., ORPHAN V.J., VAN DOVER C.L. & R.C. VRIJENHOEK (2004) Appl. Environ. Microbiol. 70(5): 3082-3090.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 120


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Mollusca, Gastropoda, Neritimorpha, Neritoidea, Phenacolepadidae

Olgasolaris BECK, 1992

Species Distribution shell. Small, vestigial operculum. Animal with large oral lobe,
O. tollmanni BECK, 1992 Manus Back-Arc Basin penis near right cephalic tentacle. Eyes rudimentary; similar
Olgasolaris sp. North Fiji and Lau Back-Arc Basins size in veliger larva and adult.
Biology: Genus endemic to vents. Feeds by grazing bacterial
Size: Shell diameter up to 13 mm. mats from surfaces of sulphide chimneys and of shells of bi-
Color: Animal pinkish-reddish when alive. valves and gastropods. The reddish colour of live animals is
probably caused by hemoglobin. Larval development with
Morphology: Shells almost circular, of limpet shape, depressed, planktotrophic larvae. Egg capsules, 1 mm diameter, deposited
with subcentral apex; sculptured by very fine, radiating, beaded on shells of other molluscs, often in large numbers on Ifremeria.
ribs; narrow shelf-like septum on the posterior inner side of

1: O. tollmanni; top left, bottom right, bottom left: exterior view, top right: Interior view; middle lateral view; by R. von Cosel.

Reference:
BECK L. (1992) Ann. Nat.hist. Mus. Wien B 93: 259-275.

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Mollusca, Gastropoda, Neritimorpha, Neritoidea, Phenacolepadidae

Shinkailepas OKUTANI, SAITO & HASHIMOTO, 1989

Species Distribution
S. briandi WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: Lucky Strike, Menez Gwen, Logatchev
S. kaikatensis OKUTANI, SAITO & HASHIMOTO, 1989 Kaikata Seamount
S. mojinensis SASAKI, OKUTANI & FUJIKURA, 2003 Ogasawara Ridge
S. tufari BECK, 1992 Manus Back-Arc Basin
Shinkailepas sp. North Fiji Back-Arc Basin
Further undescribed species Mariana Back-Arc Basin, Indian Ocean, East Pacific Rise:

right cephalic tentacle; very small eyes present in larvae and at


Size: Shell length up to 11 mm.
least some species.
Color: Animal bright red infreshly dead and living specimens.
Biology: Genus endemic to vents. The reddish colour of fresh
Morphology: Shells symmetrical, of limpet shape, depressed, animals is probably caused by hemoglobin. Development with
with posteriorly inclined apex; sculptured by radiating ribs; planktotrophic larvae. Egg capsules common on shells and oth-
shelf-like septum on the posterior inner side of shell. Small, er hard surfaces.
vestigial operculum. Animal with large oral lobe, penis near

1: S. briandi: exterior,
r interior and lateral views; by R. von Cosel & A. Le Goff.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 122–123


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2: S. briandi: exterior, interior view;


by P. Briand.

3: S. briandi: eggs; by P. Briand. 4: S. briandi: in situ; 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].
WARÉN A. & P.P BOUCHET (2001) Veliger 44(2): 116-231 [174-177].

123
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Mollusca, Gastropoda, Caenogastropoda, Muricoidea, Buccinidae

Eosipho THIELE, 1929

Species Distribution
E. auzendei WARÉN & BOUCHET, 2001 East Pacific Rise: 17-23°S; Pacific-Antarctic Ridge: 31-38°S
E. desbruyeresi OKUTANI & OHTA, 1993 Mariana, North Fiji and Lau Basins
E. desbruyeresi nipponensis OKUTANI & FUJIWARA, 2000 Okinawa Trough, Ogasawara

Size: Shell length up to 70 mm. Biology: A small radiation of buccinids living at vents; provi-
sionally placed in the genus Eosipho, a genus known from
Color: Chestnut brown to black.
sunken drift wood and normal bathyal environments. A relat-
Morphology: Large whelks with solid, smooth shell. Spire al- ed species lives at Caribbean seeps. Buccinidae are carnivorous
ways truncated (early whorls dissolved) in subadults and adults. or scavengers and E. desbruyeresi has been collected in quanti-
Thick corneous operculum smaller than aperture. 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.

P. BOUCHET Denisia 18 (2006): 124–125


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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 worm 4: E. desbruyeresi in situ, among mytilid Bathymodiolus bre-
Laminatubus alvini from southern East Pacific vior from Lau Back-Arc Basin, cruise TUIM07; by courtesy of
Rise, cruise Biospeedo © Ifremer. C.R. Fisher.

References:
OKUTANI T. & S. OHTA (1993) Venus 52: 217-221.
OKUTANI T. & Y. FUJIWARA (2000) Venus 59: 123-128 [125-126].
WARÉN A. & P.
P BOUCHET (2001) Veliger 44(2): 116-231 [190-191].

125
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Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Cerithiopsidae

Speculator cariosus WARÉN & BOUCHET, 2001

Size: Shell height up to 8.3 mm. Biology: The only known specimen was collected together with
tubeworms Ridgeia piscesae, but nothing is known about its diet.
Color: Brownish yellow.
Distribution: Northern Pacific, known only from Explorer
Morphology: Shell tall and very slender, rather fragile, with
Ridge: Magic Mountain (Steve 4 vent).
high spire and small, rounded aperture with obliquely drawn-
out siphonal canal. Whorls distinctly convex. Surface with four
strong spiral cords and two more on the body whorl and with
somewhat 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 strongly
excentric nucleus.

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

1: Apertural view; after WARÉN &


BOUCHET (2001).

Reference:
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 126


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Mollusca, Gastropoda, Caenogastropoda, Elachisinidae

Laeviphitus desbruyeresi WARÉN & BOUCHET, 2001

Size: Shell height up to 1.8 mm. Biology: Among Bathymodiolus and in sediment. Genus known
from vents and driftwood (unpubl.). Development with plank-
Morphology: Shell very small, not especially fragile, about four
totrophic larvae.
whorls, with shallow suture, narrow umbilical crevice and an-
teriorly bluntly rounded and posteriorly slightly pointed aper- Distribution: Mid-Atlantic Ridge: Menez Gwen to Rainbow.
ture, smooth teleoconch and distinctly demarcated and cancel- Genus also known from Japan (Laeviphitus japonicus OKUTANI,
late protoconch. Shell surface usually with strong ferrugineous FUJIKURA & SASAKI, 1993) and the Marianas (unpubl.). Larvae
deposits. Operculum thin, yellowish brownish, paucispiral. common at the East Pacific Rise: 13°N, but no juvenile or adult
specimens have been found.

1: The same specimen with and without ferrugineous layer; scale bar 0.5
mm; by A. Warén.

2: Different views of larvae; by L. Mullineaux.

References:
OKUTANI T. FUJIKURA K. & T. SASAKI (1993) Bull. Natn. Sci. Mus., Tokyo A 19(4): 123-143.
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [182].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 127


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Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae

Alviniconcha hessleri OKUTANI & OHTA, 1988

pertrophied and specialized gill cells contain endosymbiotic


Size: Shell height up to 85 mm.
chemoautotrophic bacteria of the sulphur cycle. Reduced di-
Color: Yellowish. gestive tract as compared to related species of the same family.
Morphology: Shell globose, rather elastic, spectacularly orna- Distribution: A single morphospecies in the Western Pacific:
mented with regularly spirally arranged periostracal hairs. Mariana, North-Fiji and Lau Back-Arc Basins; Indian Ocean:
Aperture with a shallow sinus in front. Operculum horny and Rodriguez Triple Junction, Kairei hydrothermal field. Never-
ovo-quadrate. theless, A. hessleri from the type location is genetically different
from the three Alviniconcha populations collected at other
Biology: Genus endemic in vents. Usually in stacks around
places.
vent openings where they are exposed to warm (up to 13°C),
sulphide-rich (up to 750 μM) water. Scattered specimens live
on side of chimneys and attain a very large size. The gill is hy-

2: Dorsal and ventral view; by P


P. Briand © Ifremer.
1: Apertural view; by P.
P Briand © Ifremer.

3: In situ specimen in the Lau Back-Arc


Basin; Biolau cruise © Ifremer. 4: In situ population at Lau Back-Arc Basin; Biolau cruise © Ifremer.

References:

DENIS F., JOLLIVET D. & D. MORAGA (1993) Biochem. Syst. Ecol. 21: 431-440. OKUTANI T., HASHIMOTO J. & T. SASAKI (2004) Venus 63: 1-11.
ENDOW K. & S. OHTA (1989) Bull. Jap. Soc. Microb. Ecol. 3: 73-82. OKUTANI T. & S. OHTA (1988) Venus 47: 1-9.
HEALY J. (1992) Bull. Mus. Natl. Hist. Nat. 14: 273-291. STEIN J.L., CARY S.C.,HESSLER R.R., OHTA S., VETTER R.D., CHILDRESS J.J. & H. FEL-
KOJIMA S., FUJIKURA K., OKUTANI
T T. & J. HASHIMOTO (2003) Venus 63: 65-68. BECK (1988) Biol. Bull. 174: 373-378.
KOJIMA S., OHTA S., FUJIWARA Y. & J. HASHIMOTO (1999) JAMSTEC J. Deep-Sea WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90.
Res. 14: 501-506.

A. WARÉN Denisia 18 (2006): 128


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Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae

Desbruyeresia WARÉN & BOUCHET, 1993

Species Distribution
D. cancellata WARÉN & BOUCHET, 1993 North Fiji and Lau Back-Arc Basins
D. marianaensis (OKUTANI & FUJIKURA, 1990) Mariana Back-Arc Basin
D. marisindica OKUTANI, HASHIMOTO & SASAKI, 2004 Central Indian Ridge
D. melanioides WARÉN & BOUCHET, 1993 Manus and Lau Back-Arc Basins
D. spinosa WARÉN & BOUCHET, 1993 North Fiji Back-Arc Basin
D. sp. aff. spinosa Mariana Back-Arc Basin

ters of the sculpture. Eyes reduced or absent. Right pallial ten-


Size: Shell height up to 10-12 mm.
tacle absent.
Morphology: Shells regularly coiled, distinctly slender, with
Biology: Genus endemic in vents; detritus feeders; develop-
high spire and small, rounded aperture. Sculpture consisting of
ment unknown.
axial ribs, spiral cords, knobs and occasionally short spines. Tip
often corroded, shell often encrusted. Species differ in charac-
1 top row left: D. spinosa;
middle: D. melanoides; right:
D. cancellata, holotype coated for
SEM; by R. von Cosel & A. Le Goff;
bottom row left: D. spinosa;
middle and right: D. provanna;
by P
P. Briand.

References:
OKUTANI T. & K. FUJIKURA (1990) Venus 49: 83-91.
OKUTANI T., HASHIMOTO J. & T. SASAKI (2004) Venus 63: 1-11.
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [71-73].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 129


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Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae

Ifremeria nautilei BOUCHET & WARÉN, 1991

Alviniconcha, the gill and circulatory system are hypertrophied


Synonym: Olgaconcha tufari BECK, 1991.
and the alimentary system is unexpectedly small. Two types of
Size: Up to 95 mm. bacterial symbionts are present in the gills, one dominant sul-
phide-oxidizing bacteria and one in lower abundance, likely
Color: Brown (juvenile) to black (adult).
methane-oxidizing. Different commensal polychaetes were
Morphology: Shell with about two whorls, umbilicus, nearly found inside the pallial cavity (scale worms) and the umbilicus
oval aperture, and a conspicuous subsutural ramp; periostracum (Amphisamytha cf. galapagensis). The female broods the larvae
thickened, faintly glossy, dissolved at the apex. While the shell in a brood chamber in the foot (A. Warén, unpublished). De-
is growing at the aperture, the apex dissolves. Front and sides of velopment probably lecithotrophic.
foot conspicuously light blue in living specimens.
Distribution: North Fiji, Lau, Manus Back-Arc Basins.
Biology: Genus with a single species endemic in vents, usually
in massive heaps at the edge of Alviniconcha stacks. As in

1: Adult specimens, apertural and apical view; juvenile specimens, apertural view; by P. Maestrati.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 130–131


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2 top: Living specimens in situ, inhabited by limpets Olgasolaris sp.; bottom: Left by P. Briand © Ifremer; right by
Biolau cruise © Ifremer.

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. WARÉN (1991) C. R. Acad. Sci. Paris, Sér. III 312: 495-501.
DESBRUYÈRES D., ALAYSE
A -DANET A.-M. & S. OHTA
T (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.
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [64-71].
WINDOFFER R. & O. GIERE (1997) Biol. Bull. 193: 381-392.

131
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Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae

Provanna DALL, 1918

Species Distribution
P. buccinoides WARÉN & BOUCHETT, 1993 North Fiji and Lau Back-Arc Basins
P glabra OKUTANI, TSUCHIDA & FUJIKURA, 1992
P. Sagami Bay, Okinawa Basin
P. ios WARÉN & BOUCHET, 1986 Galapagos, East Pacific Rise: 13-21°N, 17°S
P. laevis WARÉN & PONDER, 1991 Guaymas Basin
P. muricata WARÉN & BOUCHET, 1986 North Fiji and Lau Back-Arc Basin, Galapagos Sprea-
ding Center, East Pacific Rise: 21°N
P. nassariaeformis OKUTANI, 1990 Mariana and Manus Back-Arc Basins
P. segonzaci WARÉN & PONDER, 1991 Lau Back-Arc Basin
P. variabilis WARÉN & BOUCHET, 1986 Juan de Fuca Ridge, Oregon Margin

in characters of the sculpture. Tip often corroded, shells often


Size: Shell height up to 10-12 mm.
encrusted. Eyes reduced or absent. Right pallial tentacle present.
Color: Olive-brown to dark brown or greenish, often covered
Biology: Genus known from vents, seeps and sunken drift wood
by thick mineral deposits.
(only the vent species are listed herein). Detritus feeders; devel-
Morphology: Shells regularly coiled, moderately slender, with opment without planktotrophic larvae in species with known
rather high spire and rounded aperture. Surface smooth or with protoconchs. One species from seeps has adelphophagy and
sculpture consisting of axial ribs and spiral cords. Species differ
f hatches in the crawling stage (A. Warén, unpublished).

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

References:
OKUTANI
T T. & K. FUJIKURA (1990) Venus 49: 83-91.
OKUTANI T., TSUCHIDA S. & K. FUJUKURA (1992) Venus 51: 137-148.
WARÉN A. & P. Bouchet (1986) Zool. Scr. 15: 157-164.
WARÉN A. & P.P BOUCHET (1989) Zool. Scr. 18: 67-102 [94-95].
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [74-76].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [178-179].
WARÉN A. & W.F. PONDER (1991) Zool. Scr. 20: 27-56.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 132


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Mollusca, Gastropoda, Caenogastropoda, Rissoidae

Alvania cf. stenolopha BOUCHET & WARÉN, 1993

Size: Shell height up to 2.6 mm. Remark: It is not sure whether or not this species is a member
of the vent fauna; the species was described from a locality
Morphology: Shell small, conical, thin and fragile, with blunt
about 150 km northeast of Menez Gwen.
spire and large aperture, outer lip not thickened. Whorls dis-
tinctly and evenly convex, suture well-marked. Surface with Biology: Development lecithotrophic; each egg capsule con-
very fine but distinct spiral lines, 4-8 stronger spiral cords at tains a single juvenile. At vent sites encountered on sulphide
and below the periphery and distinct, evenly spaced sharp and rocks or at the base of black smokers, partly with Hydrozoa.
narrow axial ribs, 20 on the last whorl, which end abruptly at
Distribution: Mid-Atlantic Ridge: Lucky Strike, Menez Gwen
the first to third spiral cord. Protoconch dome-shaped, with 1.4
and surroundings.
whorls, diameter about 460 μm, teleoconch with 2.4 whorls.

1 left: Specimen 1, 2.6 mm, apertural view; right: Specimen 2, 2.3 mm; Menez Gwen;
after WARÉN & BOUCHET (2001).

References:
BOUCHET P. & A. WARÉN (1993) Boll. Malacol. Suppl. 3: 579-840.
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231.

A. WARÉN & P. BOUCHET Denisia 18 (2006): 133


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Mollusca, Gastropoda, Caenogastropoda, Vitrinellidae

Neusas marshalli SYKES, 1925

Size: Shell diameter up to 2.06 mm. Remarks: The species was also collected from a non-hy-
drothermal locality off Portugal, 39°42’N, 09°43’W, 1092-1993
Morphology: Shell planispiral, resembling a planorbid, with
m (type locality). Two other species of the genus are known;
rounded and almost smooth whorls with a deep suture. Proto-
they are confined to non-vent localities off New Caledonia and
conch tall-spired and obliquely inserted, with slightly more
off New Zealand.
than two whorls, smooth. Teleoconch with about three slightly
irregularly coiled whorls. Operculum corneous, multispiral, Biology: No data. Apparently the species is not obligatorily
round with central nucleus. confined to vents.
Distribution: Mid-Atlantic Ridge: Menez Gwen.

1 top: Apertural view; bottom left: Basal view; bottom right: Apical view (note the inclined protoconch); from Menez Gwen;
after WARÉN & BOUCHET (2001).

Reference:
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 134


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Mollusca, Gastropoda, Heterobranchia, Hyalogyrinidae

Hyalogyrina MARSHALL, 1988

Species Distribution
H. globularis WARÉN & BOUCHET, 2001 Juan de Fuca Ridge: Endeavour Segment
H. grasslei WARÉN & BOUCHET, 1993 Guaymas Basin

cylindrical snout. Additional tentacles behind the normal


Size: Maximum height ca 3.4 mm.
cephalic ones of variable development, possibly changing with
Morphology: Shell small, rather fragile, depressed, globular or age. Foot large, broad and flat, rounded or truncated posterior-
more tall-spired, 2.5-4 whorls on the teleoconch, vitreous and ly and shallowly bilobed anteriorly, no propodium distinguish-
fragile, with deep suture, deep umbilicus, aperture rounded. Ini- able.
tial part of protoconch sculptured by small, crowded pits, later
Biology: Epifaunal grazers. Genus known from vents, seeps,
smooth, teleoconch smooth. Operculum round, transparent
whalebone and driftwood. Development unknown.
and colorless to brown, multispiral with distinct growth lines
and central nucleus. Animal with thick tentacles and almost

1: H. globularis,
apertural and dor-
sal view of a speci-
men; Juan de Fuca
Ridge, Endeavour
Segment, Clam
Bed; by A. Warén.

2: H. globularis, 3: H. umbellifera,
critical point early protoconch;
dried specimen; A. Warén.
front view of
head-foot, shell
and pallial skirt
removed; Juan
de Fuca Ridge,
Endeavour Seg-
ment, Clam
Bed; by A.
Warén.

References:
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [49-52].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [200-207].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 135


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Mollusca, Gastropoda, Heterobranchia, Orbitestellidae

Lurifax vitreus WARÉN & BOUCHET, 2001

cylindrical tentacles. Foot with well-demarcated propodium, no


Size: Diameter up to 2.8 mm.
appendages except metapodial lobes.
Morphology: Shell small, rather fragile, depressed conical, five
Biology: Epifaunal. Genus known from vents and seeps. Devel-
whorls, vitreous, with shallow suture, broad and deep umbili-
opment unknown.
cus. Protoconch smooth, teleoconch with spiral striae and
stronger ribs and radiating flexuous incremental lines. Surface Distribution: Mid-Atlantic Ridge: Menez Gwen and Lucky
often covered by thick crusts of rust. Operculum stiff, almost Strike. Genus with three species endemic to vents and seeps.
transparent, multispiral with six whorls and central nucleus. One at Lau and Fiji Back-Arc Basins. Lurifax japonicus from
Animal with well developed eyes at the dorsal base of simple Sumisu Caldera, Southern Japan.

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


P. Briand @ Ifremer.

References:
SASAKI T. & T. OKUTANI (2005) Venus 63: 121-124.
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [207-208].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 136


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Mollusca, Gastropoda, Heterobranchia, Xylodisculidae

Xylodiscula analoga WARÉN & BOUCHET, 2001

Size: Diameter up to 3 mm. Biology: Found among Bathymodiolus and on sediment. Genus
known from vents, seeps and biogenic substrates. Development
Morphology: Shell small, fragile, subplanispiral, 2.5 whorls
unknown.
(teleconch), with rather deep suture, very wide and deep um-
bilicus and subradial and slightly prosocline aperture. Proto- Distribution: Mid-Atlantic Ridge: Menez Gwen and Lucky
conch always corroded, teleoconch with incremental lines, Strike. Another species, Xylodiscula major WARÉN & BOUCHET,
otherwise smooth. Surface with a thick, yellowish-brownish pe- 1993 was collected from North Fiji Back-Arc Basin.
riostracum. Operculum thin and transparent, round, smooth,
multispiral with central nucleus.

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

References:
WARÉN A. & P. BOUCHET (1993) Zool. Scr. 22: 1-90 [35].
WARÉN A. & P. BOUCHET (2001) Veliger 44(2): 116-231 [208, 210].

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 137


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Mollusca, Gastropoda, Prosobranchia, Conoidea, Turridae

Phymorhynchus DALL, 1908

Species Distribution
P carinatus WARÉN & BOUCHET, 2001
P. Mid-Atlantic Ridge: 23-15°N
P. hyfifluxi BECK, 1996 North Fiji Basin
P. major WARÉN & BOUCHET, 2001 East Pacific Rise: 13-9°N
P moskalevi SYSOEV & KANTOR, 1995
P. Mid-Atlantic Ridge: 26-23°N
P. ovatus WARÉN & BOUCHET, 2001 Mid-Atlantic Ridge: 37-15°N
P. 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 wareni SYSOEV & KANTOR, 1995
P. Edison Seamount, Lihir Is, West Pacific

chodaeum. Toxoglossate radula teeth hollow, needle-like in


Size: Shell height to 72 mm.
shape with basal expansion. Distal tip sharp, monocuspidate
Morphology: Shells short, obese fusiform or bucciniform. Shell with a slit.
surface white with strong spiral ribs (about 8-10 on penultimate
Biology: Often observed at the periphery of dying vents, prob-
whorl and some 25-30 on body whorl including base) which
ably predator on molluscs and scavenger. Genus known from
overlie weak growth lines. Spire roundly conical, but body
seeps and vents as well as free-living. Larval development
whorl well inflated, occupying about 50% of shell length. Aper-
planktotrophic.
ture wide, lunate, with crenulated outer lip corresponding to
spiral ribs. Columellar lip almost straight. Siphonal canal open,
not twisted. No operculum present. Head with big rhyn-

1 top row, left to right: Two


specimens of P. carinatus; one
specimen of P. moskalevi; by A.
Warén (SNHM) & P. Briand
(Ifremer); bottom row: Three
specimens of P.
P ovatus, from
Mid-Atlantic Ridge; by R. von
Cosel & A. Le Goff (MNHN).

T. OKUTANI Denisia 18 (2006): 138–139


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2 left to right: One specimen of P. major from East Pacific Rise 13°N, and two specimens of P.
starmeri from North Fiji Back-Arc Basin; by P. Briand (Ifremer).
3: Phymorhynchus sp. from East Pacific
Rise: 18°S, cruise Biospeedo © Ifremer;
middle: Phymorhynchus sp. from Juan de
Fuca Ridge; by K. Juniper; bottom: Larva
from Juan de Fuca Ridge; by K. Juniper &
L. Mullineaux.

References:
BECK L. (1996) Arch. Moll. 126(1/2): 109-115.
BOUCHET P. & A. WARÉN (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.
WARÉN A. & P.
P BOUCHET (2001) Veliger 44(2): 116-231 [192-199].
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Mollusca, Gastropoda, Nudibranchia, Dendronotidae

Dendronotus comteti VALDÈS & BOUCHET, 1998

lamellae. The margin of the rhinophoral sheath has four un-


Size: Up to 5 mm.
branched tentacular papillae of even size.
Color: Unknown in life. Preserved specimens: uniformly pale
Biology: First species of nudibranch recorded with certainty
cream background color, without traces of colored spots or
from a vent site. Living among mussel bed of Bathymodiolus
lines.
azoricus, but probably not restricted to this environment. Its oc-
Morphology: The frontal velum has four short papillae, two on currence at vents can be explained on the one hand by the
each side of the body midline. Medial pair longer and always presence of prey such as hydrozoa (Candelabrum phrygium), and
branched, but outer two only branched in the larger specimens. on the other hand by the absence of predators.
2-4 pairs of cerata on each side of the dorsum. When branched,
Distribution: Mid-Atlantic Ridge: Lucky Strike, Eiffel Tower.
the cerata form three conical processes. Rhiniphores have 7-8

1A: Dorsal view of the holotype (SEM); scale bar 1 mm; B: Details of the anterior region of
the 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 VALDÈS & BOUCHET (1998).

Reference:
VALDÈS A. & P. BOUCHET (1998) Deep-Sea Res. II 45: 319-327.

P. BOUCHET & M. SEGONZAC Denisia 18 (2006): 140


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Mollusca, Bivalvia

The Solemyidae are represented at hydrothermal vent or carbon seeps or hydrothermal vents. In the past, species of Vesi-
cold seep biotopes by the genus Acharax. Eight named species comyidae have been assigned to different genera or subgenera
as well as several still unnamed species are known, but only a and many of the large species are now commonly placed in the
single species has been collected at hydrothermal vents (Lau genus Calyptogena (sensu lato). Recently however, some au-
Back-Arc Basin). Representatives of this genus can grow to rel- thors have revised tentatively all Vesicomyidae in the sole
atively large sizes between 10 and 22 cm. They live deeply genus Vesicomya (in a broad sense) pending future supra-specif-
buried in soft sediment. At least some species are characterized ic revisions based mainly on molecular research. Anyway, shell
by the absence of a digestive tract, and their nourishment relies and soft part morphology remain important and key characters
exclusively on their chemosynthetic symbiotic bacteria. are among others hinge dentition, shell size and shape and pres-
Of the large mussels living at hydrothermal vents or cold ence or absence of a well-developed pallial sinus. Herein, the
seeps, 20 species have been currently described, 18 of them in genus Calyptogena is maintained. The assignment of a vesi-
the genus Bathymodiolus. Of these, 10 are known from hy- comyid species to Calyptogena differs with authors, and the
drothermal vents and are covered herein. Several other large supra-specific systematics of the Vesicomyidae is still far from
mussels are still awaiting description. All known Bathymodiolus settled. Most of the larger species have been described from
have a larval shell of about 0.5 mm or smaller with set-off pro- cold seep biotopes, but of the 28 named species currently treat-
toconch 1; in general, the protoconch has a distinct rose color. ed as Calyptogena, seven were found inhabiting also (or exclu-
sively) hydrothermal vents. Not very much is known of the lar-
The Vesicomyidae are a rather diverse family in size, shape val development of vesicomyid clams, but many are known to
and species number: at the moment 84 recently named species have lecithotrophic development.
are known, but there are still numerous undescribed species
currently under study by several authors. The family comprises The Pectinidae are represented by two species, Bathypecten
on one hand small species (genus Vesicomya sensu stricto), vulcani, present at the Galapagos Spreading Center, the north-
which occur in the deep sea but are not necessarily associated ern East Pacific Rise 13°N and the southern East Pacific Rise
with cold seep and/or hot vent habitats and on the other hand, 38°S and Sinepecten segonzaci, from the Manus Back-arc Basin.
most of the numerous medium-sized or large to very large More sampling need of these small bivalves, difficult to see in
species are confined to reducing sediments, cold seeps, hydro- situ, to understand the processes of their distribution.

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

R. VON COSEL Denisia 18 (2006): 141


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Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae

Calyptogena edisonensis OKUTANI, KOJIMA & KIM, 2003

Size: Shell length up to 100 mm. Biology: At hydrothermal vents.


Morphology: Shell rather large, moderately thick, elongate- Distribution: Western Pacific: Edison Seamount.
oval, dorsal and ventral margin almost parallel in their middle
part. Umbones not prominent, prosogyrous, situated anteriorly
at 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 plate
strong but not very broad, typical for Calyptogena, with three
cardinals in each valve.

1: Several specimens; by R. von Cosel.

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

R. VON COSEL Denisia 18 (2006): 142


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Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae

Calyptogena extenta KRYLOVA & MOSKALEV, 1996

Size: Shell length up to 246 mm. Biology: At cold seeps and hot vents. The specimens live
buried in the sediment but with the posterior part (2/3 of shell
Morphology: Shell very large and very elongate, thick, slightly
length) free, they are inclined at about 50° to the sediment sur-
inequivalve, somewhat irregular, bean-shaped, curved with
face and are capable of moving around by means of their large
markedly concave ventral margin and somewhat convex dorsal
muscular foot, leaving short tracks in the sediment.
margin, gaping anteriorly and posteriorly. Umbones not promi-
nent, prosogyrous, situated anteriorly at 1/6 of shell length. Distribution: Gorda Ridge (hydrothermal vents); Monterey
Surface with low growth ridges, periostracum brownish-olive, Canyon: 36°35’N, 122°30’5’’W, 3041 m (cold seeps).
persistent only posteriorly and near margins. Hinge plate rather
narrow but strong, with three cardinals in the left valve and
two cardinals in the right valve.

1: Habitus;
by R. von Cosel.

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.

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Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae

Calyptogena gigas DALL, 1896

Size: Shell length up to 125 mm. Biology: At hot vents and cold seeps, collected between 550
and 2610 m.
Morphology: Shell large, regularly oval-oblong, rather thin-
shelled, very inflated, with broadly rounded anterior and poste- Distribution: Gulf of California, 1567 m (type locality); Guay-
rior margin. Ventral margin almost straight, dorsal margin mas Basin; Juan de Fuca Ridge.
slightly convex. Umbones not prominent, prosogyrous. Surface
with fine irregular growth lines, sometimes more or less eroded
in the umbonal region. Periostracum pale olive to dark brown,
often eroded on the earlier parts of the valves. Inside with very
short and broad pallial sinus. Hinge plate short and rather
strong, with three cardinals in the left valve and three cardinals
in the right valve.

1: Habitus; by R. von Cosel.

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

References:
COAN E.V., SCOTT P.V. & F.R. BERNARD (2000) in COAN E.V., SCOTT P.V. & F.R. BERNARD (Eds.) Bivalve Seashells of Western North America: 336-343.
DALL W.H. (1896) Proc. U.S. Natl. Mus. 18(1034): 7-20 [18-19].
KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406 [400].

R. VON COSEL Denisia 18 (2006): 144


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Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae

Calyptogena magnifica BOSS & TURNER, 1980

often encountered living within basaltic rubble associated with


Size: Shell length up to 263 mm.
such warm cracks and crevices. The presence of this species is
Morphology: Shell very large, variable, oval to elongate-oval in correlated with elevated levels of hydrogen sulphide. The soft
outline, inequilateral and equivalve, usually without gape; tissue is coloured dark red when retrieved living due to the
valves white, entirely aragonitic; periostracum present, but fre- presence of intracellular hemoglobin. The thick and large gills
quently eroded away on older portions of shell; umbos prosogy- contain sulfur oxidizing chemoautotrophic symbionts. The
rous, sometimes partially enrolled; lunule and escutcheon vari- large rugose foot is often seen protruding when the clams are
able, present or absent; ligament external, opisthodetic, and viewed in life position.
parivincular; foot strong and rugose, with byssal gland.
Distribution: Entire Northern East Pacific Rise and Southern
Biology: At hot vents only. The life habit of the adult is semi- East Pacific Rise: 21°N to 22°S; Galapagos Spreading Center.
epibiotic on bare basalt, living in cracks and crevices from
which hydrothermal fluids emanate; early life history stages are

2: In situ; by courtesy of K. Smith Jr.

1: In situ, among barnacle bed of


Vulcanolepas n. sp. and chiridotid
holothurian; southern East Pacific Rise:
17°S, Biospeedo cruise © Ifremer.
3: Exterior, interior and dorsal view of left
valve, specimen from 21°N;
by R. von Cosel & P. Lozouet.

References:
ARP A. J., CHILDRESS J.J. & C.J. FISHER (1984) Physiol. l Zool. 57: 648-662. HURTADO L.A., MATEOS M., LUTZ R.A. & R.C. VRIJENHOEK (2003) Appl. Envi-
BOSS K. & R. TURNER (1980) Malacologia 20: 161-194. ronm. Microbiol. 69(4): 2058-2064.
CHILDRESS J.J., FISHER C.R., FAVUZZI J.A. & N.K. SANDERS (1991) Physiol. Zool. KENNISH M. & R. LUTZ (1992) Rev. Aquat. Sci. 6: 29-66.
64: 1444-1470. KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406
FATTON E. & M. ROUX (1981) C. R. Acad. Sc. Paris, Série III 293: 63-68. [400].
FIALA-MÉDIONI A. & C. METIVIER (1986) Mar. Biol. 90: 215-222. LUTZ R.A., FRITZ L.W. & R.M. CERRATO (1988) Deep-Sea Res. I 35: 1793-1810.
FISHER C. et al. (1988) Deep-Sea Res. 35: 1811-1831. 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.

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Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae

Calyptogena nankaiensis OKUTANI, KOJIMA & ASHI, 1996

Size: Shell length up to 191 mm. Biology: At cold seeps and hot vents, on seep sites sometimes
co-occurring with C. soyoae OKUTANI, 1957.
Morphology: Shell very large, solid, elongate-oval, with slight-
ly concave middle part of ventral margin and with posterior Distribution: Nankai Trough (cold seeps); Okinawa Trough,
part slightly higher than anterior part. Anterior margin nar- North Iheya Knoll (hot vents).
rowly rounded, posterior margin broadly rounded. Umbones
prosogyrous, almost subterminal, situated anteriorly at about
1/7 of shell length. Surface with fine, irregular, close-set growth
lines. Periostracum thick, straw-coloured, sometimes more or
less eroded. Hinge plate strong, with three cardinals in right
and left valve.

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

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.

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Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae

Calyptogena okutanii KOJIMA & OHTA, 1997

Size: Shell length up to 120 mm. Remarks: Calyptogena okutanii and C. soyoae are sibling species
with slight but constant morphological and molecular differ-
Morphology: Shell large, thick and solid, elongate-oval in out-
ences
line, with slightly concave middle part of ventral margin. Um-
bones low, prosogyrous. Surface with irregular, close-set com- Biology: At cold seeps and hot vents, at some seep sites co-oc-
marginal growth ridges and lines. Periostracum thin and brown- curring with C. soyoae OKUTANI, 1957.
ish, but often more or less erodedon the earlier parts of the
Distribution: Sagami Bay and Nankai Trough (cold seeps); Ok-
valves. Hinge plate strong and rather broad with two strong
inawa Trough, Iheya Ridge (hydrothermal vents).
cardinals in both valves.

1: Habitus; by R. von Cosel.

References:
KOJIMA S. & S. OHTA (1997) Venus 56: 189-195.
KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406 [400].

R. VON COSEL Denisia 18 (2006): 147


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Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae

Calyptogena solidissima OKUTANI, HASHIMOTO & FUJIKURA, 1992

Size: Shell length up to 128.5 mm. Biology: At hot vents and cold seeps. Living clams bury them-
selves about one half to two-thirds of the shell length into the
Morphology: Shell large, thick and robust, oblong-oval in out-
sediment. A temperature anomaly of 0.3°C was recorded 30 cm
line; with ventral margin slightly concave in its middle part.
below the white-stained bottom surface. The thick and large
Umbones prosogyrous, not prominent. Surface with irregular
gills contain sulphur oxidizing chemoautotrophic endosym-
growth lines and very fine and densely spaced radial threads,
bionts.
visible under a lens only. Periostracum extremely thin and dull
yellowish, but generally eroded and persistent only near the Distribution: Mid-Okinawa Trough: Minami-Ensei Knoll.
margins. Hinge plate strong, with three cardinals in each valve,
sub-umbonal pit present.

1: Habitus of two specimens; by R. von Cosel.

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


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Mollusca, Bivalvia, Protobranchia, Solemyoida, Solemyidae

Acharax alinae MÉTIVIER & COSEL, 1993

Size: Shell length up to 135 mm (including periostracum). Biology: In reducing sediments; found buried 20-30 cm deep in
pale-coloured coarse sediment at the base of an isolated large si-
Morphology: Shell elongate-oval. Dorsal margin straight. An-
boglinid tube in a seep area at the edge of a hydrothermal vent.
terior and posterior margin rounded. Ventral margin nearly
Known from vents only. Gills very likely harbouring chemosyn-
straight or somewhat concave. Umbos at posterior third, broad
thetic symbiosis. Lecitrotrophic development, most probably
and flattened, beaks eroded. Periostracum very strong, dark
no, or extremely short, free swimming larval phase. Larval shell
brown, extending far beyond the calcified part of the valve.
(no separate protoconch I & II) 1.35 mm long and 0.68 mm
Animal with very large gills and a voluminous, roughly cylin-
high.
drical foot which distally ends in a pedal disk. Digestive tract
absent. Distribution: Lau Basin: Valu Fa Ridge, Hine Hina.

1: Holotype, exterior and interior of both valves; by P. Maestrati © MNHN; bottom: Another specimen with soft parts;
by P. Briand © Ifremer.

Reference:
MÉTIVIER B. & R. VON COSEL (1993) C. R. Acad. Sci. Paris, Sér. III 316: 229-237.

D. DESBRUYÈRES Denisia 18 (2006): 149


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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus aduloides HASHIMOTO & OKUTANI, 1994

Size: Shell length up to 96 mm. Biology: This species is found at both hydrothermal vent and
cold seeps. The specimens are found with their anterior end
Morphology: Shell large, rather thick and solid. Umbones sub-
thrust into diffused vents and attached with byssus to outcrops
terminal, at about one-tenth of the shell length. Ligament
close to the vent openings. Periostracum colour is variable ac-
strong. Extremely thick and large ctenidia with long demi-
cording to habitat. Development with planktotrophic larvae.
branchs. No extreme mantle fusion, valvular siphonal mem-
brane short. Anterior adductor muscle scar located in front of Distribution: Central Japan: Sagami Bay; Okinawa Trough:
the umbo. Posterior adductor muscle scar rounded trapezoid. Minami-Ensei Knoll and Iheya Ridge, 710 m to 1389 m.
Gut with a single clockwise loop.

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

2: In situ © JAMSTEC.

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


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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus azoricus COSEL & COMTET, 1998

Size: Shell length up to 121 mm. Biology: In dense clusters byssally attached to hard substrate
around the hydrothermal vent, mostly on the walls and flanges
Morphology: Shell variable, more or less elongate-modio-
of active edifices, at temperatures from 6° to up to 30°C. En-
liform, beaks subterminal but close to the anterior margin. Ven-
demic to vents. Development with long planktonic larval
tral margin straight to more or less concave. Postero-dorsal
phase; protoconch is 0.5 mm long.
margin slightly to markedly convex, occasionally straight. Lig-
ament plate slightly arched. Exterior with dense irregular Distribution: Mid-Atlantic Ridge: Menez Gwen, Lucky Strike,
growth lines and growth waves. Periostracum dull, warm chest- Rainbow (hybrids between B. azoricus and B. puteoserpentis
nut brown, in umbonal region and often also postero-dorsally were observed at Broken Spur vent field).
lighter brown. Anterior byssus retractor scar in the umbonal
cavity, 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, narrow
and rather strong.

1: Holotype MNHN, exterior and interior of both valves; by R. von Cosel & P. Lozouet © MNHN.

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


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2: Post-larvae fixed on gastropod probably Protolyra thorvaldssoni (det. A. Warén); by


P. Briand © Ifremer.

3: In situ at Menez-Gwen vent field, with 4: In situ at Rainbow vent field with strong
limpets Lepetodrilus atlanticus on the sulphide deposit; Atos cruise © Ifremer.
shells; Atos cruise © Ifremer.

References:
COMTET T., LE PENNEC M. & D. DESBRUYÈRES (1999) J. Mar. Biol. Ass. U.K. 79: 1149-1150.
COMTET T., JOLLIVET D., KHRIPOUNOFF A., SEGONZAC M. & D.R. DIXON (2000) Limnol. Oceanogr. 45: 1655-1661.
COSEL R. VON, MÉTIVIER B. & J. HASHIMOTO (1994) Veliger 42(3): 218-248 [220-231].
FIALA-MÉDIONI A., MCKINESS Z., DANDO P., BOULEGUE J., MARIOTTI A., ALAYSE-DANET J., ROBINSON J. & C. CAVANAUGH (2002) Mar. Biol. 141: 1035-1043.
O’MULLAN G.D., MAAS P.A.Y., LUTZ R.A. & R.C. VRIJENHOEK (2001) Mol. Ecol. 10: 2819-2831.

152
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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus brevior COSEL, MÉTIVIER & HASHIMOTO, 1994

Size: Shell length up to 143 mm. Biology: In dense clusters byssally attached to hard bottom
around the hydrothermal vents at temperatures up to 18°C. En-
Morphology: Shell oval-wedge shaped, stout, beaks subtermi-
demic to vents. Development with long planktonic larval
nal. Ventral margin straight to more or less concave. Postero-
phase; protoconch is 0.4 mm long.
dorsal margin slightly convex to almost straight. Ligament plate
slightly arched. Exterior smooth, with irregular growth lines. Distribution: North Fiji and Lau Back-Arc Basins.
Periostracum dull, dark brown, in umbonal region lighter
brown. Anterior retractor scar at the anterior part of the um-
bonal cavity. Anterior part of posterior byssus retractor muscle
scar at 2/3 of the ligament. Animal with large gills. Mantle
lobes on anterior half of ventral side separate. Valvular siphonal
membrane short, narrow and rather strong.

1: Holotype MNHN, exterior of both valves, interior


of 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; Biolau
cruise © Ifremer.

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

R. VON COSEL Denisia 18 (2006): 153


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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus elongatus COSEL, MÉTIVIER & HASHIMOTO, 1994

Size: Shell length up to 155 mm. Biology: Byssally attached to lava around diffuse vents, with
absence of massive hydrothermal deposits and vent fluid tem-
Morphology: Shell elongate-wedge shaped, slender, beaks well
perature not exceeding 8.5°C. Endemic to vents. Development
subterminal. Ventral margin straight or slightly convex. Liga-
not known but most probably with planktonic larval phase.
ment plate somewhat arched to straight. Exterior with irregular
growth lines and narrow to broad irregular, concentric grooves, Distribution: North Fiji Back-Arc Basin.
mostly on the ventral part. Periostracum glossy, light chestnut
brown, in umbonal region lighter brown. Anterior retractor
scar at the anterior part of the umbonal cavity. Anterior part of
posterior 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.

1: Several specimens, exterior, interior, dorsal view; bottom: Juveniles, exterior and interior of right valve; from
North Fiji Back-Arc Basin, Mussel Valley site; cruise Starmer 2; by P. Maestrati © MNHN & P. Briand © Ifremer.

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

R. VON COSEL Denisia 18 (2006): 154


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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus japonicus HASHIMOTO & OKUTANI, 1994

Size: Shell length up to 107 mm. Biology: This species is found at both hydrothermal vents and
cold seeps. It is the most abundant and conspicuous organism of
Morphology: Shell rather thick, stout, moderately tumid. Um-
the hydrothermal vent community on the Minami-Ensei Knoll.
bones subterminal, at about 5% anterior of the shell length.
Development not known but most probably with planktonic
Posterior end of ligament abrupt. Extremely thick and large
larval phase.
ctenidia with long demibranchs. No extreme mantle fusion,
valvular siphonal membrane short. Anterior byssus retractor Distribution: Off Hatsushima site and Sagami Bay, central
muscle scar located in the anterior part of the umbonal cavity. Honshu, 1170 m. Mid-Okinawa Trough: Minami-Ensei Knoll
Pallial line smooth, concave in adult, but slightly concave in and Iheya Ridge.
juvenile and sub-adult.

1: Paratype MNHN Paris, exterior of both valves, interior


of right valve; by R. von Cosel & P. Lozouet © MNHN. 2: In situ view of a mussel bed © JAMSTEC.

References:
HASHIMOTO J. & T. OKUTANI (1994) Venus 53(2): 61-83.
HASHIMOTO J. HASHIMOTO J., OHTA S., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. 42(4): 577-598.

J. HASHIMOTO Denisia 18 (2006): 155


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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus marisindicus HASHIMOTO, 2001

Size: Shell length up to 86 mm. Biology: In dense beds byssally attached to hard bottom along
shimmering crevices near black smokers. Endemic to vents. De-
Morphology: Shell oval-modioliform, stout, beaks subterminal.
velopment with long planktonic larval phase.
Ventral margin almost straight to slightly concave. Postero-dor-
sal margin somewhat convex. Ligament plate slightly arched. Distribution: Indian Ocean: Rodriguez Triple Junction, Kairei
Exterior with dense fine commarginal lines. Periostracum hydrothermal field.
smooth and glossy, strong, dark brown to blackish brown, in
young 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 the
whole ventral margin. Valvular siphonal membrane short, nar-
row and rather strong.

2: In situ view of a small


mussel bed, among numerous
sea anemones Mariactis cf.
bythios, alvinocaridid shrimps
Rimicaris kairei and gastro-
pod Phymorhynchus sp.
© JAMSTEC.

1: Some specimens, exterior and interior of valves; by R. von Cosel & P. Lozouet © MNHN.

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.

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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus platifrons HASHIMOTO & OKUTANI, 1994

Size: Shell length up to 115.6 mm. Biology: This species is found at both hydrothermal vents and
cold seeps. In Sagami Bay, B. japonicus is occasionally sympatric
Morphology: Shell rather thick and solid. Umbones often near-
in the same habitat. Periostracum colour variable according to
ly subterminal, but mostly on the same plane with anterior end
habitat. Development not known but most probably with
of the shell. No extreme mantle fusion, valvular siphonal mem-
planktonic larval phase.
brane short. Extremely thick and large ctenidia with long demi-
branchs. Distribution: Okinawa Bank and off Hatsushima site in Saga-
mi Bay, central Honshu, 1180 m; Mid-Okinawa Trough: Iheya
Ridge and Izena Caldron.

1: Paratype MNHN Paris, exterior


and interior of right valve;
by R. von Cosel & P. Lozouet © MNHN.

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

Reference:
HASHIMOTO J. & T. OKUTANI (1994) Venus 53(2): 61-83.

J. HASHIMOTO Denisia 18 (2006): 157


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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus puteoserpentis COSEL, MÉTIVIER & HASHIMOTO, 1994

posterior third of the ligament, near the end. Animal with large
Size: Shell length up to 141 mm.
gills. Mantle lobes on anterior half of ventral side separate.
Morphology: Shell oval-wedge shaped, rather stout, beaks sub- Valvular siphonal membrane short.
terminal. Ventral margin straight to very weakly convex. Pos-
Biology: Bysally attached to sulfur blocks immediately around
tero-dorsal margin markedly convex. Ligament plate slightly
diffuse venting of water. Endemic to vents. Development un-
arched in anterior part, straighter in posterior part. Exterior
known, most probably with long planktonic larval phase.
smooth, with pronounced irregular growth lines. Periostracum
dark brown and rather glossy. Anterior byssus retractor scar on Distribution: Mid-Atlantic Ridge: TAG, Snake Pit and Lo-
anterior part of the umbonal cavity, in front of the beaks. An- gatchev.
terior part of posterior byssus retractor muscle scar under the

1 top: Holotype MNHN, exterior of both valves; mid- 2: In situ view of a mussel bed, with sea
dle: Other specimen, interior and exterior of left valve; anemones Maractis cf. rimicarivora and
bottom: Interior of left valve of holotype; by R. von zoarcid fish Pachycara cf. thermophilum;
Cosel & P. Lozouet © MNHN. cruise Microsmoke © Ifremer.

References:
CAVANAUGH C. M., WIRSEN C.O. & H.W. JANNASCH (1992) Appl. Environ. Microbiol. 58: 3799-3803.
COSEL R. VON, METIVIER B. & J. HASHIMOTO (1994) Veliger 37(4): 374-392 [387-389].
O’MULLAN G.D., MAAS P.A.Y., LUTZ R.A. & R.C. VRIJENHOEK (2001) Mol. Ecol. 10: 2819-2831.

R. VON COSEL Denisia 18 (2006): 158


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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus septemdierum HASHIMOTO & OKUTANI, 1994

Size: Shell length up to 124 mm. Biology: At hydrothermal vents emitting fluids over 310°C
around the living beds. Endemic to vents. The shell surfaces are
Morphology: Shell large, rather thin but solid, inflated, ellipti-
covered with numerous filamentous bacteria. Periostracum col-
cal, more or less compressed. Umbones subterminal, less than
or changes according to growth stages. Development not
one-eighth anterior of the shell length. Ligament weak. Ex-
known but most probably with planktonic larval phase.
tremely thick and large ctenidia with long demibranchs. No ex-
treme mantle fusion, valvular siphonal membrane short. Ante- Distribution: Izu Ogasawara Arc: Suiyo Seamount and Mokuyo
rior adductor muscle scar located below the umbo. Anterior Seamount.
part of the posterior byssus retractor muscle scar under posteri-
or end of ligament, at about four-fifths of ligament length.

1: Paratype MNHN, interior and exterior of right valve;


by R. von Cosel & P. Lozouet © MNHN.

2: In situ © JAMSTEC.

Reference:
HASHIMOTO J. & T. OKUTANI (1994) Venus 53(2): 61-83.

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Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus thermophilus KENK & WILSON, 1985

Size: Shell length up to 18.4 cm. Biology: The life habit of the adult is epibiotic on bare basalt
and other hard substrates (e.g. tubes of vestimentiferans) asso-
Morphology: Shell smooth, modioliform, with subterminal um-
ciated with deep-sea hydrothermal vents. The presence of this
bones; periostracum present and straw-yellow to brown in
species is correlated with elevated levels of hydrogen sulfide.
colour; external surface lacks sculpture and is dull white be-
Endemic to vents. Protoconch 0.4 mm in length, indicative of
neath periostracum; hinge edentulous; ligament opisthodetic,
a planktotrophic larval stage with a high dispersal capability.
paricincular, strong, extending most of the length of the dorsal
Paired ctenidia consist of inner and outer demibranchs, each
margin; well developed byssus. Valvular siphonal membrane
with descending and ascending lamellae. The gills contain sul-
long and stretching towards ventrally; on anterior part fusion of
phur-oxidizing chemoautotrophic symbionts. A commensal
inner mantle fold reaching ventrally to 1/3 shell length, to-
polychaete (Branchipolynoe symmytilida) is frequently found
gether with siphonal membrane leaving rather short byssus
within the mantle cavity of the mussel.
opening in the middle.
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. von
Cosel & R. Vrijenhoek, unpublished data).

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

R. LUTZ & R. VRIJENHOEK Denisia 18 (2006): 160–161


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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.

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.

161
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Mollusca, Bivalvia, Pteriomorphia, Mytiloida, Mytilidae

Gigantidas gladius COSEL & MARSHALL, 2003

Size: Shell length up to 31.6 cm. Biology: Protoconch II about 0.4 mm in length, which indi-
cates a long planktonic larval stage with a high dispersal capa-
Morphology: Shells large to very large, aduliform, extremely
bility. The extremely enlarged gills contain sulphide oxidizing
long, elongate and sword-shaped, rather thin for the size but
chemoautotrophic symbiotic bacteria. A commensal poly-
solid, with flattened and very broad umbones situated well sub-
chaete of the genus Branchipolynoe was encountered within the
terminally in the anterior half of the valves. Ventral margin
mantle cavity of most specimens. Mussels of the genus Giganti-
well concave. Periostracum strong and dark brown; valves dull
das are found on warm seeps near active submarine volcanoes
white beneath the periostracum. Inner mantle folds separate
and therefore are included herein. The species lives in dense
along whole length of the ventral margin their edges frilled
populations at sulphur-rich hydrothermal seepings, they are
along the posterior fifth of the shell length or less. Valvular
partly buried in sediment.
siphonal membrane almost absent, but a deep cleft between in-
ner mantle fold of right and left valve. Attachment point of the Distribution: Southwestern Pacific: Rumble III and Rumble V
anterior foot-byssus retractor muscle directly above the anteri- submarine volcanoes, S-Kermadec Ridge, New Zealand; West-
or adductor scar and united with it; posterior byssus retractor ern Pacific: Kaikata Seamount, SW of Ogasawara (Bonin) Is-
complex multibundle with two principle diverging muscle bun- land.
dles and two additional thin bundles.

1: Two paratypes (MNHN), exterior and interior of right valve, exterior of left valve, dorsal view; by D. Brabant © MNHN.

References:
COSEL R. VON & B.R. MARSHALL (2003) Nautilus 117: 31-46.
HASHIMOTO J. & M. HORIKOSHI (1989) Deep-Sea Newsletter 15: 32-34.
HASHIMOTO J. & T. YAMANE (2005) Venus 64: 1-10.
JONES W.J., WON Y.J., MAAS P.A.Y., SMITH P.J., LUTZ R.A. & R.C. VRIJENHOEK (2006) Mar. Biol. 148: 841-851.

R. VON COSEL Denisia 18 (2006): 162


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Mollusca, Bivalvia, Pteriomorphia, Mytiloida, Mytilidae

Gigantidas horikoshii HASHIMOTO & YAMANE, 2005

Anterior retractor scar situated in front of umbonal cavity; pos-


Size: Shell length up to 19.5 cm.
terior byssus retractor complex with two principle diverging
Morphology: Shells large, aduliform, long and slender, rather muscle bundles.
thick, solid. Ventral margin markedly concave in adults, giving
Biology: The mussels were observed on sandy bottom with
the shell a curved appearance. Umbonal cavity large and
warm water seepage (18°C). The species live in dense popula-
swollen. Periostracum strong chocolate or dark brown, um-
tions, partly buried in the sediment.
bonal region light brown; valves dull white beneath the perios-
tracum. Inner mantle folds entirely separate, terminating ante- Distribution: Kaikata Seamount, SW of Ogasawara.
riorly on anterior adductor. No valvular siphonal membrane.

1: Muscular system; after HASHIMOTO & YAMANE (2005).

2 top: Dorsal view; bottom: Lateral view; 3: Specimens in situ, from Kaikata Seamount;
after HASHIMOTO & YAMANE (2005). by J. Hashimoto © JAMSTEC.

References:
HASHIMOTO J. & M. HORIKOSHI (1989) Deep-Sea Newsletter 15: 32-34.
HASHIMOTO J. & T. YAMANE (2005) Venus 64: 1-10.

J. HASHIMOTO Denisia 18 (2006): 163


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Mollusca, Bivalvia, Pteriomorphia, Eupteriomorphia, Pectinidae

Bathypecten vulcani SCHEIN-FATTON, 1985

Size: Up to 17 mm. Biology: Can be abundant at the periphery of vents (30-50 in-
dividuals m-2) with mussels and gastropods. Byssally attached in
Morphology: Small, thin, flattened, left (upper) valve more
diffuse venting areas with a low temperature anomaly. Usual bi-
convex than right (lower); smooth, but early stage with con-
valve filter-feeding status. Non planktotrophic larval develop-
centric undulations, more developed and regular on upper
ment.
valve; hinge line straight, posterior auricle not delimited, ante-
rior auricle above a distinct byssal notch in lower valve. Distribution: Galapagos Spreading Center; East Pacific Rise:
9°N and 13°N; Pacific-Antarctic Ridge: 32°S, observed at 38°S.

1-4: Specimens from East Pacific Rise: 13°N, cruise HOT 96.

1: Specimen viewed from the right (lower) side; 2: Same specimen viewed from the left (upper) side;
by P. Briand. by P. Briand.

4: In situ view of a population of B. vulcani on basalt,


among polychaete serpulid tubeworms (Laminatubus
3: Lower (right) side of the antero-dorsal region showing alvini), and small sea anemones, probably Chon-
the anterior auricle above the byssal notch; by P. Briand. drophellia cf. coronata; by courtesy of R. Vrijenhoek.

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


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Mollusca, Bivalvia, Pteriomorphia, Eupteriomorphia, Pectinidae

Sinepecten segonzaci SCHEIN, in press

Size: Up to 28 mm. Biology: Byssally attached on glassy basalt near vents where the
temperature ranged between 2.8°C and about 40°C.
Morphology: Shell thin, lower (right) valve flat; upper (left)
valve convex. Commarginal ridges or lamellae covering the Distribution: Manus Back-Arc Basin. A juvenile specimen of
whole of the upper valve, and the lower valve only after the ju- the same genus was collected at North-Fiji Back-Arc Basin,
venile stage. Posterior auricle poorly delimited. Anterior auri- close to the vent field White Lady in June 2005 (cruise TU-
cle of the lower valve above a byssal notch which is widely IM06MV, R. Vrijenhoek © MBARI).
open in the juvenile, but gradually closing and finally over-
lapped by its lower edge in the adult.

1-3: Specimens collected by J. Hashimoto, cruise Bioaccess 98; from SCHEIN (in press).

1: Outer view of a juvenile left valve (SEM). 2: Lower side of a juvenile specimen.

3: Upper side (left) and lower side (right) of large adult holotype.

Reference:
SCHEIN E. (in press) Zootaxa.

E. SCHEIN Denisia 18 (2006): 165


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Mollusca, Cephalopoda

The living cephalopods, commonly the finned and finless their maintenance under suitable aquaria conditions can an-
octopuses, squid and cuttlefishes with the vampire squid and swer this and other intriguing questions on the behaviour of
Nautilus, comprise a well-defined class of Mollusca. Although these deep-sea cephalopods.
these marine predators have undergone an extensive evolu-
After specimen collection, a tissue sample from arm muscu-
tionary radiation, only benthic incirrate octopuses of Granele-
lature should be taken by slitting the skin on the dorsal side of
done and Benthoctopus and benthopelagic cirrate octopus
the arm to expose the muscle. Opening the skin avoids per-
species of Cirroteuthis and Grimpoteuthis have been reported
turbing the ventral suckers and minimizes contamination from
near vents, and only one octopod genus is recognized as en-
the skin. The tissue sample should be frozen or preserved in
demic to hydrothermal vents, Vulcanoctopus on the East Pacif-
95% ethanol for subsequent analysis. The specimen should be
ic Rise. Octopods may form an ecologically important compo-
then placed in a 6% solution of buffered formalin in seawater,
nent of the vent ecosystem as opportunistic rather than en-
if possible with the arms extended. The specimen can be kept
demic predators, but they remain under-collected in the vent
in the formalin for several days before being shifted to ethanol.
habitat. The largest single cause is the difficulty in capturing
The specimens should be examined for any parasites and their
these animals. Manipulators can grab large individuals of some
locations should be noted. The presence of adaptations to hy-
species, especially cirrate and incirrate octopuses of Granele-
drothermal vent habitat, such as high concentrations of heavy
done, and suction samplers and traps effectively collect Vul-
metals in the tissues or presence of amoebocytes clots in the ve-
canoctopus specimens, but individuals of Benthoctopus tend to be
nous system and in the renal sacs, should be carefully exam-
much more wary. Even when secured, specimens can be severe-
ined.
ly damaged.
A second problem limiting our knowledge of these animals
is that, even with the specimen in the hand, species-level dif-
ferences in these little-known groups are often subtle at best
and rely heavily on internal characters of sexually mature
males. As gravid females of some species are suggested to con-
1, 2: Two in situ views of
gregate in rocky areas, such as mid-ocean ridges, collections
an unknown cephalopod
tend to be female-biased, although in Vulcanoctopus hydrother- taken at the Mid-Atlantic
malis almost all known specimens are male. Based on isolated Ridge, north of Rainbow;
deep-sea specimens, current species boundaries may be found to cruise Flores © Ifremer.
be artificial, but only after careful reassessment of type speci-
mens and newly collected specimens.
Videotape from research submarines and remotely operated
vehicles have documented a wide repertoire of cirrate octopus
behaviours 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 basic
as whether the animals can change colour, a notable character
of shallow-water octopods. Bioluminescence has been observed
in some cirrate species and may be a more widespread phenom-
enon than suspected. Careful collection of live specimens and

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


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Mollusca, Cephalopoda, Octopoda, Cirroteuthidae

Cirrothauma murrayi CHUN, 1911

or two eggs are stored in the distal oviduct, nearly to be re-


Size: Total length up to 940 mm; mantle length 220 mm, from
leased, measuring to 14 x 8.9 mm. This indicates that once sex-
sampled specimens; may be longer from observations at the East
ual maturity it is attained, eggs are probably released one or two
Pacific Rise: 13°N.
at a time, following a continuous spawning strategy of repro-
Color: In situ photographs show cirroteuthids with purple, red duction. No data exist about the feeding habits of this species
and/or brown color on both oral and dorsal surfaces. and no predators are reported. Despite their small size, the eyes
of C. murrayi are probably sufficient to detect the biolumines-
Morphology: External features: body relatively elongate (i.e.,
cence produced by other animals and the wide aperture of the
not compressed) and gelatinous. Eyes degenerate, cup-like,
cornea allows for detection of flashes over wide angles and for
without lenses or iris, embedded within the jelly of the skin,
greater sensitivity. Locomotion it is basically by the fins and the
look like small black balls. Fins large, wide, longer that head
animal becomes streamlined during fin swimming.
width. Mantle aperture closed around a long, slender funnel.
Each arm bears a single longitudinal row of suckers alternating Distribution: Specimens of this species have been collected in
with paired, very long cirri. Suckers strongly modified in barrel- Atlantic, Pacific and Arctic Oceans. The animals were seen at
like form. Intermediate web present, linking arms to primary great depths (1500-5000 m), often near the bottom, but some-
web. Internal features: butterfly-shaped shell. times 300-450 m above it. In the NE Atlantic COLLINS et al.
(2001) collected 27 specimens, most of them below 3000 m
Biology: The spermatophores of the males are small, rounded,
depth. Always one or two specimens observed at the East Pa-
simple in structure, and stored in the oviducal gland of the fe-
cific Rise: 13°N-17°S, around the vents. Probably the same
male, indicating an internal fertilization. In gravid females, the
species observed near other vents along the East Pacific Rise.
ovarian oocytes reach 200 in number and are present in differ-
ent sizes and stages of development, from 0.4-9 mm long. One

1: Specimen in situ, East Pacific Rise: 13°N; cruise Biocyarise 2: Specimen in situ, East Pacific Rise: 17°S; cruise Biospeedo
© Ifremer. © Ifremer.

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


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Mollusca, Cephalopoda, Octopoda, Cirroteuthidae

Cirroteuthis magna HOYLE, 1885

They can be considered as abyssopelagic animals, although they


Size: Up to 1300 mm total length. The largest known cirrate
can also rest on the bottom, where they probably feed. Its vo-
octopod.
luminous and operative eyes indicate that these animals are
Morphology: Deep-sea animals of very great size (up to 1300 able to detect light produced by themselves, other animals of
mm TL). Butterfly-like shell. Shell Width Index: 26-31. Very the same species, or by potential predators and prey. This
voluminous eyes (Eye-ball Diameter Index: 39-43), with large species, and other related ones, have been observed to swim at
lenses (Lens Diameter Index: 12-18). Arm length up to 940 2.2 km/h and 0.46 km/h. It occurs near hydrothermal vents, but
mm. Primary web inserting at different levels on the oral and was rarely observed at the Mid-Atlantic Ridge.
aboral ends of the dorso-lateral and ventro-lateral arms on both
Distribution: South Indian (2557 m) and Pacific (1500 m) to
sides, and at the same levels on both ends of the dorsal and ven-
subtropical North Atlantic (1350 m). One specimen (1300 mm
tral arms. Very long non-retractile cirri (Cirrus Length Index:
length) was caught at the Mid-Atlantic Ridge, near Logatchev,
96-71); the first cirri commences between the fourth and fifth
3351 m. Another specimen was captured near a vent site in the
suckers. Three types of suckers on all arms: cylindro-conical
southern East Pacific Rise: 17°S, 2574 m (M. Lilley & K. Van
form and those with the acetabulum highly deformable on the
Damm, chief Scientists; J. Voight, det.).
first 2/3 and barrel-shaped on the rest of the arm; no enlarged
suckers in male or female.
Biology: C. magna is, as other cirrate octopods, a cephalopod,
typically adapted to the deep-sea environment in the abyssal
ecosystem. These gelatinous animals are neutrally buoyant.

1: Ventral view of mature male


(220 mm). Reconstruction
based on the specimen 2, 3: In situ views of the specimen caught north of Logatchev; cruise Faranaut © Ifremer. The
captured and video images; specimen swans by moving its fins a few meters of the bottom, a locomotion mode named
from GUERRA et al. (1998). fin-swimming; from VILLANUEVA et al. (1997).

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


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Mollusca, Cephalopoda, Octopoda, Grimpoteuthidae

Grimpoteuthis ROBSON, 1932

Size: Up to 115 mm mantle length. Biology: Stomach contents in G. wuelkeri and G. boylei showed
polychaetes, copepods, amphipods and isopods. Mature females
Color: Skin of red brown, orange or purple color in fresh spec-
have few and large eggs in oviducts, measuring from 10-18 mm
imens. Usually oral side more pigmented that dorsal side.
in length, suggesting a continuous spawning mode as observed
Morphology: External features: body gelatinous and bell- in other cirrate octopods. Individuals of Grimpoteuthis sp. have
shaped. Fins medium to large, with distinct lobe near the ante- been observed resting and crawling on the sea bottom, swim-
rior fin insertion. Thick primary web. Intermediate web absent. ming basically by the use of fins and also by arm-web contrac-
Each arm bears a single longitudinal row of suckers alternating tions. As a response to disturbance, individuals have been ob-
with paired, medium-sized cirri. Internal features: optic lobe served in web inversion, with arms and web upturned, oral sur-
spherical, optic nerves pass though white body as a single boun- face facing outward, completely covering mantle, head and fins.
dle of fibres. Shell U-shaped with other edges of lateral walls
Distribution: Specimens of this genus has been collected in the
parallel, not tapered to single fine points. The genus comprises
Atlantic and Pacific Oceans. In the NE Atlantic, abundances
14 species.
of Grimpoteuthis sp. ranged from 1.3-25 individuals km-2 at
depths from 1500-4850 m (COLLINS et al. 2001).

1: In situ specimen taken at East Pacific Rise: 13°N, 2: In situ specimen taken at Mid-Atlantic Ridge,
site Elsa; cruise HOPE 99 © Ifremer. north of Rainbow; cruise MARVEL © Ifremer.

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


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Mollusca, Cephalopoda, Octopoda, Octopodidae

Vulcanoctopus hydrothermalis GONZÁLEZ, GUERRA, PASCUAL & BRIAND, 1998

Size: Up to 56 mm mantle length, 235 mm total length and 45 Biology: This benthic species has characters that represent ei-
g body weight. ther adaptations to the deep-sea (absence of ink sac, loss of the
anal flaps, eye without iris and optic chiasma) or to a hy-
Color: In situ, white, eyes black.
drothermal vent habitat (eyes are covered by a thin semi-
Morphology: Body semi-translucent with a muscular consisten- translucent skin, high concentrations of metals and presence of
cy; mantle pear-shaped and posteriorly mitre-like; presence of a amoebocytes clots in the venous system and in the renal sacs).
large white body (which covers the eye, the optic nerves and It inhabits an isolated extreme environment among aggrega-
the optic lobe), an inflated black bulb (dark swelling), in prox- tions of tubeworms Riftia pachyptila, Alvinellidae polychaetes or
imal third of the intestine, a crop, and a multilobulate digestive mussels, very close to the chimneys (2-10°C); also observed on
gland; absence of an ink sac. Arms 1.5-4.3 times mantle length. the pillow lava at several meters from the active areas. No pre-
Two rows of suckers on each arm. Arm formula typically 1.2.4.3 dation over these species was observed. Octopuses forage on ba-
or 2.1.4.3. No enlarged suckers. Maximum depth of the largest thypelagic amphipods, apparently targeting their attacks based
sector of the web about 22% of the longest arm. Gills with 7-8 on contact with the swarming amphipods (e.g. Halice hes-
lamellae per demibranch. Right arm III hectocotylized (HA monectes). Some male specimens were parasitised by Genesis
1.5-2.1 times mantle length) in males. Ligula short (8-10% of vulcanoctopusi, a species of cholidynid harpacticoid (copepod).
HA), lance-shaped and without transverse ridges. Calamus rep-
Distribution: East Pacific Rise: collected at 13°N (Fig. 2) and
resents 30-50% of the ligula length in fully mature specimens.
21°S (Fig. 1); observed at 23°S (Fig. 3).
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.

2: Specimen observed
on tubes of Riftia at
East Pacific Rise:
13°N; cruise HOPE’99
© Ifremer.

3: Specimen observed
at East Pacific Rise:
23°S in April 2005;
cruise PAR 5
© MBARI.

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.

References:
GONZÁLEZ A.F., GUERRA A., PASCUAL S. & P. BRIAND (1998) Cah. Biol. Mar. 39: 169-184.
GONZÁLEZ A.F., GUERRA A., ROCHA F. & P. BRIAND (2002) Bull. Mar. Sci. 71(1): 289-298.
LÓPEZ-GONZÁLEZ P.J., BRESCIANI J., HUYS R., GONZÁLEZ A.F., GUERRA A. & S. PASCUAL (2000) Cah. Biol. Mar. 41: 241-253.
NIXON M. & J.Z. YOUNG (2003) The Brain and Lives of Cephalopods. Oxford Univ. Press: 1-368.
NORMAN M.D & F.G. HOCHBERG (2005) Phuket marine biol. Center Res. Bull. 66: 127-154.
ROCHA F., GONZÁLEZ A.F., SEGONZAC M. & A. GUERRA (2002) Cah. Biol.Mar. 43: 299-302.
VOIGHT J.R. (2005) J. Mar. Biol. Ass. U.K. 85: 985-988.

A. GONZÁLEZ, A. GUERRA & M. SEGONZAC Denisia 18 (2006): 170


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Mollusca, Cephalopoda, Octopoda, Octopodidae

Benthoctopus GRIMPE, 1921

Size: Total lengths can approach one meter, but typically 50 cm Biology: Members of Benthoctopus are typically more active and
or less. more apt to jet away from submersibles and ROVs than are
other deep-sea octopus, making submersible-collected speci-
Color: Variable, from light to red to violet or deep purple. Re-
mens rare. These octopuses sometimes extend their dorsal arms
verse counter-shading (the dorsal surface lighter than ventral
vertically into the water column, perhaps to enhance chemore-
surface) is pronounced in some species, absent in others.
ception. Egg-brooding females may aggregate on hard substrate
Morphology: The genus is poorly delineated and is likely not where they often sit with the suckered surfaces of their arms
monophyletic; it includes octopuses without an ink sac with facing away from the rock.
two rows of arm suckers. Field identification of members of this
Distribution: Octopuses of the genus Benthoctopus occur world-
genus relies on their smooth skin, double rows of suckers
wide, typically between about 400 m rarely to as deep as 2000
(which are distinct from zig-zag sucker rows of octopus of
m. Octopuses of Benthoctopus have been seen near vents at Juan
Graneledone), the comparatively narrow heads and mantles
de Fuca Ridge including Endeavour segment, and Gorda Ridge
and, in some species, the mantle being lighter in dorsally than
(where they brood eggs), and cold seeps off South America.
ventrally. 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 enlarged
suckers contribute to species identification.

2: Individual on the bottom.

1: Brooding females; with numerous ophiuroids, probably Ophiuridae © MBARI.

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.

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Mollusca, Cephalopoda, Octopoda, Octopodidae

Graneledone JOUBIN, 1918

Size: Adults from shallow in the depth range of the genus (800- Biology: Hydrothermal vent polychaetes and gastropods were
1000 m) can exceed 1 m in length; those from in deepest part found in the gut of an octopus from Axial Volcano. Egg-brood-
of the range (2800 m) are only about half as long. ing females sit with their ventral (suckered) surface toward the
hard substrate to which their eggs are attached; groups of egg-
Morphology: Large octopuses with a very broad head and man-
brooding females can form locally dense aggregations. On open,
tle, suckers arranged in a single or at times zig-zag row on each
sedimented seafloor the octopuses may probe the sediment with
arm; skin texture warty (warts more prominent in specimens
their arms to locate infaunal prey.
from the greater depths), with conspicuous papillae over the
eyes which can make the eyes appear larger. Individuals tend to Distribution: Specimens of the genus Graneledone have been
be uniformly colored, but individuals vary in color from violet collected from Juan de Fuca and Explorer Ridges. Photos docu-
to orange to blue. These octopuses tend to move with their ment an individual of the genus near clams at the Galapagos
arms curled dorsally in what appears to be a semi-protective po- Spreading Center, near cold seeps off Oregon and California
sition. and on the Kermadec Arc at Monowai Caldera (http://ocean-
explorer.noaa.gov/explorations/05fire/logs/april12/april12.html).
Remark: Identifying species of Graneledone is difficult, but eggs
Members of the genus, also known from near Antarctica and
collected in the Northeast Pacific at near 2660 m depth were
South Africa, may associate with chemosynthetic areas.
shorter than those from near 1500 m depth indicating two dis-
tinct species are present.

1: Graneledone sp. with tubeworms Ridgeia; by courtesy of V. Tunnicliffe, HiRise Expedition.

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


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Nematoda

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

173
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Nematoda, Adenophorea, Desmodorida, Desmodoridae, Desmodorinae

Desmodora alberti VERSCHELDE, GOURBAULT & VINCX, 1998

Size: 750-1150 µm. Biology: Living in heterogeneous sediment covered with bacte-
ria. Specific structure of buccal cavity points to epistrate feeder.
Morphology: Desmodorid with rounded head capsule and long,
slender, conical tail. Fine somatic setae; absence of subcephalic Distribution: Guaymas Basin.
setae. Body cuticle annulated. Amphid cryptospiral. Buccal
cavity with large dorsal tooth, ventral teeth not observed.
Cylindrical pharynx with muscular endbulb, bipartite cuticular
valves in terminal bulb of pharynx. Large globular sperm cells
and short bent spicule in males. Females with vulva at approx-
imately 56% of body length.

3: Holotype male
posterior end;
scale bar 50 µm;
by D. Verschelde.

2: Holotype male anteri-


or end; scale bar 50 µm;
1: Holotype male total; 4: Allotype female total; scale bar by D. Verschelde.
scale bar 50 µm; 50 µm; by D. Verschelde.
by D. Verschelde.

5: Head capsule (SEM); scale bar 1 µm;


by D. Verschelde.

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


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Nematoda, Adenophorea, Desmodorida, Desmodoridae, Desmodorinae

Desmodora marci VERSCHELDE, GOURBAULT & VINCX, 1998

Size: 2110-2760 µm. Biology: Colleted with mussels and Solenidae. Maximal tem-
perature below 20°C. Specific structure of buccal cavity points
Morphology: Desmodorid with long slender body. Head cap-
to epistrate feeder.
sule. Slender conico-cylindrical tail. Fine somatic setae. Body
cuticle annulated. Amphid cryptospiral. Buccal cavity with Distribution: Lau Back-arc Basin, Hine Hina site.
large dorsal tooth and one or two ventral teeth hard to distin-
guish. Cylindrical pharynx with muscular endbulb. Spicules of
males with complex capitulum. Vulva of females located at
50-55% of body length.

2: Paratype male
anterior end;
scale bar 50 µm;
by D. Verschelde.

3: Holotype male
posterior end;
scale bar 50 µm;
1: Holotype male
by D. Verschelde.
total;
scale bar 50 µm;
by D. Verschelde.

4: Allotype female total; scale bar 50 µm;


by D. Verschelde.

Reference:
VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.

D. VERSCHELDE, J. ZEKELY & M. BRIGHT Denisia 18 (2006): 175


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Nematoda, Adenophorea, Desmodorida, Desmodoridae, Desmodorinae

Desmodorella balteata VERSCHELDE, GOURBAULT & VINCX, 1998

Size: 870-1080 µm. Biology: Living in heterogeneous sediment covered with bacte-
ria. Specific structure of buccal cavity points to epistrate feed-
Morphology: Desmodorid with long head capsule and short,
er.
conical tail approximately 10% of total body length. Slender
body annuli. Two pairs of lateral rows of distinct spines among Distribution: Guaymas Basin.
other longitudinal rows of spines. Absence of subcephalic setae.
Amphid multispiral, buccal cavity with cheilorhabdia in
cheilostome, 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 body
length.

1: Holotype male to-


tal, note epizoic suc-
torians attached to
body cuticle;
scale bar 50 µm;
by D. Verschelde.

5: Total view female (SEM); scale


3: Holotype male bar 100 µm; by D. Verschelde.
posterior end; scale bar
50 µm; by D. Verschelde.

2: Holotype male
anterior end;
scale bar 50 µm; 4: Allotype female total; 6: Head capsule male (SEM);
by D. Verschelde. scale bar 50 µm; by D. Verschelde. scale bar 1 µm; by D. Verschelde.

Reference:
VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.

D. VERSCHELDE, J. ZEKELY & M. BRIGHT Denisia 18 (2006): 176


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Nematoda, Adenophorea, Desmodorida, Desmodoridae, Desmodorinae

Desmodorella spineacaudata VERSCHELDE, GOURBAULT & VINCX, 1998

Size: 745-956 µm. Biology: Living in heterogeneous sediment covered with bacte-
ria. Specific structure of buccal cavity points to epistrate feed-
Morphology: Desmodorid with smooth three-part rounded
er.
head capsule and short, conical tail. Slender body annuli. Two
pairs of lateral rows of distinct spines among other longitudinal Distribution: Guaymas Basin.
rows 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 filiform
spicules and postcloacal thorns on the tail. Vulva in females at
approximately 62% of body length.

1: Holotype male
total;
scale bar 50 µm; 3: Holotype male
by D. Verschelde. posterior end;
scale bar 50 µm;
by D. Verschelde.

4: Allotype
female total;
scale bar 50 µm;
by D. Verschelde.

2: Holotype male anterior end; scale bar 50 µm; 5: SEM micrograph head capsule male;
by D. Verschelde. scale bar 1 µm; by D. Verschelde.

Reference:
VERSCHELDE D., GOURBAULT N. & M. VINCX (1998) J. Mar. Biol. Ass. U.K. 78: 75-112.

D. VERSCHELDE, J. ZEKELY & M. BRIGHT Denisia 18 (2006): 177


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Nematoda, Adenophorea, Chromadorida, Draconematidae, Prochaetosomatinae

Cephalochaetosoma pacificum notium DECRAEMER & GOURBAULT, 1997

Size: 575-850 µm. Biology: Hard substrate with oxide deposits and bacterial mats.
Temperature 5-17°C. Specific structure of buccal cavity points
Morphology: Draconematid with coarsely striated cuticle ex-
to epistrate feeder.
cept in head region. Helmet present. Amphid spiral. Buccal
cavity well developed, with a dorsal tooth and two small sub- Distribution: Lau Back-Arc Basin.
ventral teeth. Pharynx with terminal bulb. Numerous cephalic
adhesion tubes with non-swollen base and open tip located pos-
terior head region and extending to about two head diamters
along the cervical region; posterior adhesion tubes with bell-
shaped end, all located anterior to cloacal opening/anus. Tail
cylindro-conoid with numerous annules. Four pairs of cloacal
setae. Females with vulva at about 48% of body length. Males
with spicule length 40-50 µm.

1: Holotype male total; scale bar 20 µm;


by W. Decraemer.

2: Paratype female total;


scale bar 20 µm; by W. Decraemer.

Reference:
DECRAEMER W. & N. GOURBAULT (1997) Zool. Scr. 26: 1-12.

J. ZEKELY & M. BRIGHT Denisia 18 (2006): 178


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Nematoda, Adenophorea, Chromadorida, Draconematidae, Prochaetosomatinae

Dinetia nycterobia DECRAEMER & GOURBAULT, 1997

Size: 320-490 µm. Biology: Sedimented vent site. Specific structure of buccal cav-
ity points to deposit feeder.
Morphology: Draconematid with short, stout and sigmoid to
epsilonematoid habitus. Body annulation reaching the lip re- Distribution: East Pacific Rise: 21°N.
gion and surrounding the small, spiral amphid. Cuticle not
thickend in head region. Buccal cavity narrow and unarmed.
Pharynx with terminal bulb with thickend cuticle. Cephalic
adhesion tubes without expanded base located in cervical re-
gion; posterior tubes with bell-shaped end. In females vulva at
45% of body length.

3: Paratype 4th
stage juvenile
female;
scale bar 20 µm:
by W. Decraemer.

1: Holotype male
total;
scale bar 20 µm;
by W. Decraemer.

2: Paratype female
total; scale bar 20 µm;
by W. Decraemer.

Reference:
DECRAEMER W. & N. GOURBAULT (1997) Zool. Scr. 26: 1-12.

J. ZEKELY & M. BRIGHT Denisia 18 (2006): 179


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Nematoda, Secernentea, Spirurida, Habronematoidea, Cystidicolidae

Moravecnema segonzaci JUSTINE, CASSONE & PETTER, 2002

360 µm long, distal extremity pointed with a cuticular spur at


Size: Male, 5 mm long, female, 4-10 mm long.
23 µm from end. Gubernaculum absent. Four pairs of precloa-
Morphology: Body filiform. Oral opening dorsoventrally elon- cal pedunculate papillae arranged in two groups, Six pairs of
gated, hexagonal in shape. Pseudolabia rudimentary. Four sub- postcloacal pedunculate papillae. Female: amphidelphic. Vulva
median buccal processes on margin of oral opening. Four sub- at about two thirds from cephalic apex. Eggs larvated in vagina,
median papillae and two lateral amphids. Buccal cavity long, 37-42 x 25 µm with 1-5 thin filaments, 40-140 µm long, aris-
dilated dorsoventrally at anterior extremity. Oesophagus divid- ing from a small plug at each pole. Tail 65-95 µm long.
ed into short anterior muscular and long posterior glandular
Biology: Parasitic in the intestine of the zoarcid fish Pachycara
parts. Deirids tiny, spine-like, located just anterior to posterior
thermophilum GEISTDOERFER, 1994.
end of buccal cavity. Nerve ring and excretory pore located at
anterior and posterior thirds, respectively, of muscular oesoph- Distribution: Mid-Atlantic Ridge: Logatchev; Snake Pit-
agus. Tail rounded in both sexes. Male: Caudal alae present. Moose.
Area rugosa absent. Spicules unequal: short spicule arcuate,
rounded at distal extremity, 80-100 µm long; long spicule 260-

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).

Reference:
JUSTINE J.-L., CASSONE J. & A. PETTER (2002) Folia Parasitol. 49: 299-303.

J.-L. JUSTINE Denisia 18 (2006): 180


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Acanthocephala, Palaeacanthocephala, Echinorhynchida, Hypoechinorhynchidae

Hypoechinorhynchus thermaceri BURON, 1988

Size: 3.5 to 5 mm long by 0.4 to 0.7 mm wide. Biology: Intestinal parasite of zoarcid fish Thermarces cerberus.
No intermediate host was identified but other species from the
Color: White, or sometimes yellowish.
same family have amphipods for intermediate hosts. Gono-
Morphology: Globular proboscis armed with ~2 rows of 2-3 choric with a slight sexual dimorphism. Attaches to intestinal
hooks each. Trunk unarmed. Testis arranged in tandem. Six ce- mucosa using the proboscis.
ment glands. Female genital pore subterminal.
Distribution: East Pacific Rise: 13°N.
Remarks: Needs to be carefully isolated from the mucosa by
dissecting the intestine around the proboscis. For morphologi-
cal studies, worms should be placed in distilled water for about
one hour in order to ensure optimal evagination of the pro-
boscis and fixed in AFA. For molecular studies, fix the worms
directly in 70% alcohol.

2: Proboscis, face view (SEM); by I. de Buron. 3: Proboscis, lateral view (SEM); by I. de Buron.

1: Female (left) and male (right); 4: In situ specimens in the gut of a zoarcid 5: In situ specimen of a digenean in the in-
by I. de Buron. fish Thermarces cerberus; by I. de Buron. testine of a zoarcid fish Thermarces cerberus,
among specimens of Hypoechinorhynchus
thermaceri; by I. de Buron.

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


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Nemertini, Hoplonemertini, Monostilifera

Thermonemertes valens ROGERS, GIBSON & TUNNICLIFFE, 1996

Size: 5 cm. Remarks: Another yet undescribed species of Thermonemertes


was found at East Pacific Rise: 9°N from inactive sulfide chim-
Color: Bright red; after fixation uniformly flesh colored.
neys and at 13°N from Riftia pachyptila aggregations.
Morphology: The head is spatuloate and dorsoventrally flat-
Biology: Lives on rocks and bacterial mats; not associated with
tened. A single cephalic furrow on the ventral surface towards
vestimentiferan tubeworms. The diet remains unknown. Might
the rear of the head. Mouth and rhynchodeum open into a sin-
be able to swim. It has been suggested that this species is an ear-
gle opening behind the tip of the head. The body is slender and
ly colonizer of vent communities, but is displaced in aging com-
dorsoventrally compressed. In the posterior half the ventral sur-
munities.
face appears distinctly concave.
Distribution: Juan de Fuca Ridge.

2: Thermonemertes sp.; live specimen from East Pacific Rise:


9°N; by M. Bright.

1: T. valens (preserved specimen); left anterior end, right total; 3: Thermonemertes sp.; live specimen anterior end with par-
from ROGERS et al. (1996). tially extruded proboscis; from East Pacific Rise: 13°N; by M.
Bright

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


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Annelida, Polychaeta1

According to different countings, the polychaete worms, with lows recognizing three different new species of Archinome and one
111 presently described species, constitute 18-20% of the total new genus of Amphinomidae among specimens previously identi-
number of species fully identified from vent samples. 30% of these fied as A. rosacea (J. Kudenov, pers. comm.) Conversely, new
bristle worms are belonging to scale worms (Polynoidae), 10% to records of previously described species from new locations (e.g. Hes-
alvinellids, 9% to siboglinids, 6% to spionids, 5% to hesionids and iolyra bergi on Mid-Atlantic Ridge vents or Hesiospina vestimentifera
dorvilleids. Other families account for less than 5% of described from Lau Basin) question the dispersal and allopatric speciation in
annelid species. The present recorded species are largely (94%) response to major vicariance events. Because of the influence of
considered as ‘endemic’ of the deep-sea hydrothermal vent envi- joint occurrences in biogeographic analyses, the solution of these
ronment (or according to Andrey Gebruk, vent ‘obligate’), less taxonomic riddle must be seriously considered and we urge scien-
than 4% are shared with other deep-sea reduced environments and tists working at sea to focus on these widespread morphotypes and
ca. 2% are ‘regular’ deep-sea species. However this high ratio of en- to build up parallel samples for classical and molecular taxonomy.
demism will probably be revised downward when the sampling of Polychaetes are delicate and fragile animals, and special care
the peripheral areas will make progress accounting for opportunis- should be taken when handling them. To be described, the speci-
tic groups as dorvilleids or spionids which could be thriving in ad- mens must be unfragmented and as intact as possible and damaged
jacent disturbed habitats. specimens may be misidentified. Thus a gentle sorting is desirable as
Because of their odd biology and ecology, two species of an- soon as possible after recovery of the samples, using soft pliers for
nelids Riftia pachyptila (‘the giant tube worm’) and Alvinella pompe- macroscopic individuals and gentle sieving for others (avoid stack
jana (‘the Pompeii worm’) became emblematic of the vent research of sieves and split the samples before sieving). Specimens may be re-
and are still considered as ‘biological models’. Numerous research laxed prior to preservation (7.5% of magnesium chloride) and tubes
projects dealing with the functioning of endosymbiosis in Riftia of tubicolous annelids must be opened; fixation is best in buffered
pachyptila and the physiological adaptation of Alvinella pompejana formalin 5-10%. After a suitable time of fixation, depending of the
to its extreme habitat are presently underway. size (in general < 24 hours), the worms may be transferred to 80%
Nevertheless, the basic taxonomy of vent annelids is far to be ethyl or isopropyl alcohol (formaldehyde even buffered, is a very
completed. Recent cruises which occurred in the Western Pacific poor preservative).
(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 from
Lau Basin are being described belonging to Levensteiniella,
Branchinotogluma, Lepidonotopodium and Harmothoe as well as one
species 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, several
species of scale worms, one species of alvinellid and one surprising
new genus of spionid are currently under description. New families
for this environment were found and the description of new species
of flabelligerid and sphaerodorid are under wording. Former collec-
tions of worms from Mid-Atlantic Ridge and East Pacific Rise are
still not exhaustively studied and several families (e.g. dorvilleid,
spionid, capitellid, cirratulid) need further taxonomic work. 1: Laminatubus alvini; by courtesy of R. A. Lutz.
The molecular identification of sibling species among wide- 1
Even if the question is still strongly debated and the molecular information is
spread morphotypes (e.g. Amphisamytha galapagensis or Archinome not consistent, we chose herein, following ROUSE & FAUCHALD (1997) to in-
clude the Pogonophora (Perviata and Vestimentifera) within the family Si-
rosacea) lead ‘classical’ taxonomists to dig back throughout former
boglinidae (Polychaeta: Sabellida). Contrarily SOUTHWARD et al. (2005) chose
collections looking for new diagnostic characters. For example, a a more conservative standpoint and retain the class ‘Pogonophora’ within An-
joint work between morphologists and molecular taxonomists al- nelida, waiting for more conclusive information (HALANYCH 2005).

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.

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Annelida, Polychaeta, Eunicida, Amphinomidae

Archinome rosacea (BLAKE, 1985)

conspicuous denticles confined to tips. Largest notochaetae


Size: Up to 13 mm long, 5.8 mm wide.
slightly less robust than largest neurochaetae. Ratio of long to
Morphology: Body short, fusiform, roughly trapezoidal in cross- short prongs of bifurcate notochaetae ranging from 5.3-10.7: 1;
section, with 18 segments. Prostomium with paired dorsal and neurochaetae from 2.1-4.7: 1; notoaciculae from 20.5-22: 5: 1;
inconspicuous median antenna. Caruncle fused through and neuroaciculae around 36: 1. Branchiae present from
chaetiger 2, cantilevered to posterior margin of chaetiger 3 and chaetiger 3, palmate. Pygidium opening on dorsum of last two
anterior margin of chaetiger 4 (sometimes to chaetiger 5). chaetigers; cirrus digitiform, recurved anterodorsally.
Paired palps similar in shape, just slightly longer than paired
Biology: Ubiquitous; associated with tube worms and in mussel
prostomial antennae. Peristomium glandular, ciliated; chordate
beds. Carnivore and scavenger with dietary preferences for
shape divided into two lobes by a long, deep midventral groove,
polychaetes, crustaceans; also some molluscs.
extending through anterior lip of mouth into pharynx. Mouth
ventral, opening at junction of chaetigers 1-2. Chaetae simple, Distribution: Galapagos Spreading Center, Rose Garden,
calcareous and bifurcate. Long prongs of bifurcate chaetae near- Northern East Pacific Rise.
ly straight, smooth to distally serrated with around three in-

2: Branchiae; by Kudenov. 5: Bifurcate chaetae with


short prongs; by Kudenov.

1: Specimens from East Pacific Rise: 13°N;


by P. Briand © Ifremer. 3: Prostomium and caruncle, 4: Branchiae (SEM) © Ifremer.
dorsal view; by Kudenov.

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


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Annelida, Polychaeta, Eunicida, Dorvilleidae

Exallopus jumarsi BLAKE, 1985

Size: Small, about 3 mm long, 0.6 mm wide for 26 segments. Biology: From fine sediments among hydrothermal mounds.
Densities of 2667 individuals m-2 were recorded from mats of
Morphology: Prostomium wider than long, rounded along an-
Beggiatoa sp.
terior margin; with two annulated dorsal antennae and two
short, smooth lateral palps. Chaetiger 1 modified, without dor- Distribution: Guaymas Basin.
sal and ventral cirri, but bearing two types of heavy modified
simple 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, no
ventral 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.

1A: Anterior end, dorsal view; B: Same, ventral view; C: Heavy 2: Maxillary apparatus, showing maxillary carriers and indi-
modified chaetae from chaetiger 1; E: Group of composite fal- vidual plates. Labels on maxillary apparatus indicate maxil-
cigers from anterior chaetiger; F: Normal simple chaetae from lary carriers (MC) and maxillae (1-7); from BLAKE (1985).
anterior chaetigers; from 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


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Annelida, Polychaeta, Eunicida, Dorvilleidae

Ophryotrocha akessoni BLAKE, 1985

Size: Up to 8.5 mm long, 1.1 mm wide, for 55 segments. Biology: An abundant, opportunistic epifaunal species occur-
ring with siboglinids, mussels, and clams. In the Guaymas
Morphology: A small, compact species. Color in alcohol light
Basin, O. akessoni dominated in sediments saturated with pe-
tan. Prostomium broadly rounded anteriorly with two smooth
troleum hydrocarbons. Densities of 3,222 individuals per m2
antennae and two similar palps. Segments 1-2 achaetous, fol-
were recorded from mats of Beggiatoa sp. in the Guaymas Basin.
lowed by similar appearing chaetigers. Each segment with sin-
gle ciliary band encircling body. Pygidium with two anal cirri. Distribution: Widespread at the East Pacific Rise: 21°N, 13°N,
Each chaetiger with long dorsal cirrus and short, stubby ventral 9°N, Galapagos Spreading Center and Guaymas Basin.
cirrus. 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.

1: Specimen in vivo, apical 2: Maxillary apparatus: left juvenile, right


view  Ifremer adult; by courtesy of J. Blake.

3: Anterior part, lateral view (SEM)  Ifremer 4: Composite falciger


(SEM)  Ifremer

References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 8: 103-116.
GRASSLE J.F. (1988) Adv. Mar. Biol. 23: 301-366.
PETRECCA R.F & J.F. GRASSLE. (1989) Proc. Gorda Ridge Symp.: 279-283.

J.A. BLAKE Denisia 18 (2006): 188


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Annelida, Polychaeta, Eunicida, Dorvilleidae

Ophryotrocha globopalpata BLAKE & HILBIG, 1990

dorsolateral anal cirri. Mandibles elongate, rodlike, with short,


Size: Up to 7 mm long, 1.0 mm wide, with 40+ segments.
rounded, serrated cutting edge and large, unsclerotized lateral
Color: In alcohol opaque white. wings giving mandible triangular outline. Maxillae with P-type
forceps and eight pairs of free denticles with numerous small
Morphology: Body wide, dorsoventrally compressed, blunt an-
teeth along cutting edge.
teriorly, tapering towards pygidium; with long, prominent para-
podia. Prostomium rounded, twice as wide as long; with two Biology: An epifaunal species associated with smokers and ac-
long, antennae; with two biarticulate palps bearing wide, glob- tive vents. Mature females with about 6-8 eggs in each para-
ular palpophores and slender palpostyles; eyes absent. Peris- podium and sometimes 3-4 additional ones in large ventral
tomium with two achaetous rings about as long as following pouches formed from ventral body wall; egg diameters from 64-
chaetigers. Parapodia uniramous, with triangular acicular lobe, 120 µm, those in pouches larger (mean = 102 µm) than those
low prechaetal lobe and retractable ventral chaetal lobe; dorsal in the parapodium (mean = 81 µm). Mature males with sperm
and ventral cirri absent. Chaetae of three kinds: (1) supraacic- from chaetiger 3 or 4 to end of body, visible in parapodia and
ular fascicle of long, flat capillaries with fine subdistal serration; under dorsal body wall.
(2) subacicular fascicle of heterogomph falcigers with distally
Distribution: Juan de Fuca Ridge.
serrated shaft and smooth, narrow, distally blunt blades; (3) in-
feriormost smooth capillary. Pygidium rounded with two long

1A: Anterior end dorsal view; B: Posterior end, dorsal view; C: Middle seg- 1: Dorsolateral view of the anterior part (SEM)
ments of ovigerous female with ventral pouches, ventral view; D: Parapodium © Ifremer.
anterior view; E: Supraacicular chaetae; F: Subacicular falcigers; G: Ventralmost
capillary; from BLAKE & HILBIG (1990).

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

J.A. BLAKE Denisia 18 (2006): 189


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Annelida, Polychaeta, Eunicida, Dorvilleidae

Ophryotrocha platykephale BLAKE, 1985

Size: Up to 40 mm long, 2.5 mm wide, with more than 150 seg- Remarks: (1) The illustration of the parapodium in the origi-
ments. nal description of O. platykephale (BLAKE 1985: Fig. 14C) was
shown upside down and misinterpreted. This error was later
Morphology: Body dorsoventrally compressed throughout,
noted by BLAKE & HILBIG (1990) and corrected by SOLÍS WEISS
thickest anteriorly; posterior segments appearing ragged from
& HILBIG (1992). The original figure has been corrected here.
projecting parapodia. Color in life and alcohol opaque white.
(2) A closely related morphotype of O. platykephale is present
Prostomium flattened, wider than long, with cirriform, distally
in the same samples from the Guaymas basin hydrothermal
tapering antennae and palps, both of similar length; eyes ab-
mounds. It differs externally from O. platykephale having a peri-
sent. Peristomium with single achaetous ring. Uniramous para-
stomium with two achaetous rings and a globose dorsal cirrus
podia projecting far away from body, with pre- and postchaetal
(see Fig. 2-6). Its mandibles differ also from O. platykephale be-
lobes short and rounded anteriorly, becoming elongate by about
ing more spindly and curved. Its specific status remains to be
chaetiger 15. Dorsal cirri from chaetiger 17-19, becoming long,
discussed (likely morphological variants or different stages of
digitiform, bifid in posterior chaetigers. Ventral cirri short, re-
O. platykephale).
tractile from chaetiger 3 to end of body. Dorsal branchiae from
chaetiger 14; ventral branchiae from chaetiger 2; ventral Biology: Living specimens of O. platykephale were observed
branchiae longer than dorsal ones. Large parapodial glands swimming among a colony of the ampharetid Amphisamytha
present between dorsal cirrus and dorsal parapodial base. galapagensis. The general habitat was of a seep site warmed by
Chaetae include dorsal fascicle of simple spines and ventral fas- percolating hot water, but away from chimneys.
cicle of compound spinigers. Maxillae with large P-type forceps
Distribution: Guaymas Basin hydrothermal mounds.
and seven pairs of free denticles.

1A: Anterior end, dorsal


view; B: Same, ventral view;
C: Left middle parapodium;
D-G: chaetae; adapted from
BLAKE (1985).

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

J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 190–191


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3: Medial parapodia, dorso-lateral view showing branchia and 5: Anterior parapodia, dorso-lateral view showing the two
globular dorsal cirri (SEM) © Ifremer. chaetal fascicles (SEM) © Ifremer.

4: Medial parapodia, dorso-lateral view showing the two 6: Composite blunt-tipped chaetae (SEM) © Ifremer.
chaetal fascicles (SEM) © Ifremer

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.

191
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Annelida, Polychaeta, Eunicida, Dorvilleidae

Parougia wolfi BLAKE & HILBIG, 1990

supraacicular fascicle simple, with subdistal serrations and fur-


Size: Up to 11.5 mm long, 1.0 mm wide, with about 55 seg-
cate tip (seldom observed by D. DESBRUYERE); capillaries ab-
ments.
sent. Subacicular fascicle with 10-15 compound falcigers with
Color: In alcohol opaque yellowish-white. subdistally serrated shaft and bifid, distally hooded blade. Py-
gidium small, with terminal anus surrounded by heavily ciliat-
Morphology: Body long, cylindrical, with projecting parapodia,
ed, rounded papillae. Mandibles triangular, with heavily sclero-
tapering within last 5-10 segments towards pygidium. Segmen-
tized handle and transparent lateral wings; cutting edge with
tal ciliation absent (present on specimens observed by DD).
outer hook-shaped tooth and about five large, rounded and five
Prostomium rounded anteriorly, triangular to pear-shaped, with
smaller, irregular teeth. Maxillae with superior basal plates and
large, flattened palps, longer than greatest prostomial width;
30-35 superior and inferior free denticles; maxillary carriers and
antennae unknown. Peristomium with two rings, as long as fol-
inferior basal plates absent.
lowing chaetigers. Parapodia in chaetiger 1 uniramous, follow-
ing chaetigers subbiramous; neuropodium with low, square Biology: Collected from active vents.
prechaetal lobe and longer, conical postchaetal lobe; notopodi-
Distribution: Juan de Fuca Ridge.
um with conspicuous acicula in long basal part and short, con-
ical distal article; ventral cirri short, cirriform. Chaetae of

1: Prostomium, dorso-lateral view (SEM)


© Ifremer.

2: Anterior part, ventral view (SEM) 4: Middle parapodia, dorsal view (SEM)
© Ifremer. © Ifremer.

3: Three compound falcigers of the sub- 5: Simple chaetae of 6: Simple chaetae of


acicular fascicle with subdistally serrated the supraacicular fas- the supraacicular fasci-
shaft and bifid, distally hooked blade cicle with a furcate cle with subdistal serra-
(SEM) © Ifremer. tip (SEM) © Ifremer. tions (SEM) © Ifremer.

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

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


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Annelida, Polychaeta, Eunicida, Eunicidae

Eunice pulvinopalpata FAUCHALD, 1982

three protruding dark brown or black aciculae; with long dorsal


Size: A large species, 450 mm long, 12 mm wide, for more than
cirrus and short, thick ventral cirrus. Chaetae include bidentate
350 chaetigers.
compound hooded hooks, limbate capillaries, pectinate
Morphology: A large, robust eunicid. Prostomium short, trans- chaetae, bidentate subacicular hooks, and acicula.
verse, with pair of large palps along anterior edge; with five
Biology: Found on smoker walls in rusted areas; probably a car-
weakly articulated antennae. Peristomium with two achaetous
nivore.
rings, with separation between the two indistinct; a pair of peri-
stomial cirri on second ring. Branchiae from chaetiger 3-4 to Distribution: East Pacific Rise: 21°N and 13°N.
end of body; branchiae mostly pectinate, with multiple fila-
ments. Parapodia all similar, with rounded acicular lobe bearing

5: Anterior part of the proboscis (SEM)


© Ifremer.

1: Anterior part, dorsal view; 3: Anterior part, ventral view;


by P. Briand © Ifremer. by P. Briand © Ifremer.

6: Bidentate compound hooded hooks


(SEM) © Ifremer.

4: Parapodium, SEM view


2: Midpart, dorsal view; showing one of the dark
by P. Briand © Ifremer. brown acicula © Ifremer. 7: Pectinate chaeta (SEM) © Ifremer.

References:
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 8: 103-116.
FAUCHALD K. (1982) Proc. Biol. Soc. Wash. 95: 871-877.

J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 193


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Annelida, Polychaeta, Phyllodocida, Chrysopetalidae

Thraumastos dieteri WATSON, 2001

the ventral edge of the prostomium. Mid-body notopodia with


Size: > 10 mm maximum size (type truncated).
lateral and main paleal fascicles. Neuropodia with subacicular
Morphology: Anterior mid-body rectangular, long tapered an- fascicle of heterogomph falcigerous neurochaetae. Glandular
teriorly, numerous segments. Pale gold coloured palae fans pads present.
(brown areas present). Prostomium small quadrate compressed
Biology: Common in washings of Bathymodiolus brevior.
between anterior segments. Subulate median antennae, with
short ceratophore, inserted medially on anterior edge of the Distribution: North Fiji and Lau Basins and cold seep south of
prostomium, dorsally to two similar sized lateral antennae. Lihir Papua New Guinea.
Prostomial cirri with filiform tips. Two ovoid palps inserting on

1: Habitus in vivo  Ifremer 4: Prostomium, ventral view (SEM)  Ifremer.

5: Neuropodial spinigerous falciger (SEM)


 Ifremer.
2: Prostomium, dorsal view (SEM)  Ifremer.

6: Detail of two median paleae (SEM)


3: Notopodial fan, middle body (SEM)  Ifremer.  Ifremer.

References:
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.
WATSON C. (2001) The Beagle, Records of the Museums and Art Galleries of the Northern Territory 17: 57-66.

D. DESBRUYÈRES Denisia 18 (2006): 194


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Annelida, Polychaeta, Phyllodocida, Glyceridae

Glycera branchiopoda MOORE, 1911

Synonym: Glycera profundi CHAMBERLIN, 1919. Biology: Burrows in soft sediments; not endemic. Glycerids,
sometimes called bloodworms, are long-bodied active burrow-
Size: Body up to 90 mm with up to 137 segments.
ing predatory worms, which lives in soft sediments.
Morphology: Conical prostomium long with 8-9 rings. Pro-
Distribution: Pacific coast of North America, Central Ameri-
boscis with two types of papillae: (1) numerous tall thin, with
can Trench; off central Oregon and Gulf of California deep
straight, median longitudinal ridge and very small papillae sur-
basins including Guaymas Basin hydrothermal mounds. Depth
rounding core; (2) isolated shorter broader oval to globular
range 10-2562 m.
papillae without ridge. Aileron with pointed triangular base.
First two parapodia uniramous; following parapodia biramous.
Two slender triangular to digitiform praechaetal lobe. One
shorter, rounded, sometimes blunt triangular postchaetal lobe.
Dorsal cirri distinctly visible from 3rd parapodium, oval to glob-
ular. Posterior parapodial lobes prolonged. Branchiae absent.

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.

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


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Annelida, Polychaeta, Phyllodocida, Glyceridae

Glycera tesselata GRUBE, 1863

to digitiform prechaetal lobes; neuropodial lobe usually slightly


Size: Up to 76 mm long with 181 segments.
longer and wider than notopodial lobe. Dorsal cirrus from third
Morphology: Proboscidial papillae mainly digitiform with parapodium, conical to oval; inserted on body wall far above
straight, median, longitudinal ridge; aileron with deeply incised parapodial base. Branchiae absent.
bases; parapodia of mid-body with two short rounded
Biology: Glycerids are considered to be carnivorous worms.
postchaetal lobes; branchiae absent. Mid-body segments bian-
nulate. Conical prostomium consisting of about 8-9 rings. Pro- Distribution: Northwestern and northeastern Atlantic, Gulf
boscis with two types of papillae: (1) numerous digitiform papil- of Mexico, Caribbean Sea, Mediterranean Sea, Red Sea,
lae with straight, median, longitudinal ridge; (2) isolated, south coast of Africa, Indo-Pacific, northwestern Pacific;
shorter and broader conical papillae with more or less distinct- 2000-4066 m; Mid-Atlantic Ridge: Lucky Strike.
ly straight, median longitudinal ridges. First two parapodia uni-
ramous; following parapodia biramous. Two slender triangular

1: Top left, proboscis and anterior 2: Same part, lateral view (SEM) © 3: Proboscidial papillae (SEM)
part in a living specimen; Ifremer. © Ifremer.
by P. Briand © Ifremer.

D. DESBRUYÈRES Denisia 18 (2006): 196–197


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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) 8: Notochaetae, median parapodia (SEM)
© Ifremer. © Ifremer.

Reference:
BÖGGEMANN M. (2002) Abhandl. Senkenberg. Naturforsch. Ges. Frankfurt 555: 1-249.

197
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Annelida, Polychaeta, Phyllodocida, Hesionidae

Amphiduropsis axialensis (BLAKE & HILBIG, 1990)

rophores. Neuropodial lobes and neurochaetae present from


Size: All known specimens incomplete, anterior end fragments
segment 5, notopodial lobes and notochaetae from segment 6.
up to 8 mm long for 21 segments.
Notopodial lobes conical, pointed. Notochaetae include many
Color: Preserved animals yellow-white, pigmentation absent or fine capillaries with two rows of small teeth, three to four ven-
with dark brown-black pigmentation on lips, laterally on pros- trally situated curved chaetae with distal serration on ventral
tomium, palpophores and nuchal organs, distally on proboscis, side, and two (rarely one) medially situated stout emerging aci-
and on venter. cular chaetae. Neuropodial lobes conical, much larger than no-
topodial lobes. Neurochaetae all compound. Ventral cirri dis-
Morphology: Prostomium rounded rectangular, wider than
tally inserted on neuropodium.
long, posteriorly with small median incision. Palpophores very
short, palpostyles of equal width or with slightly enlarged Remarks: The new record from East Pacific Rise: 9°N is based
rounded tips. Paired antennae as long as palps, tapered. Both on a single specimen from the expedition Hope’99 (Nautile
palpostyles and paired antennae very long and slender. Median dive 1372, 2500 m, 09°46’17’’N, 104°21’70’’W); although not
antenna much shorter than paired antennae, with tapering tip, in good condition it agrees well with A. axialensis.
inserted medio-dorsally on prostomium. Eyes absent. Proboscis
Distribution: Juan de Fuca Ridge: Axial Seamount and Middle
smooth and unarmed, ending anteriorly in ring of cilia but
Valley; East Pacific Rise: 9°N (new record; see Remarks).
without papillae. Dorsal cirri and cirrophores on segment 1–5
larger than on following segments, ventral cirri on segment 1–4
larger and longer than on following segments, with distinct cir-

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

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


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Annelida, Polychaeta, Phyllodocida, Hesionidae

Hesiodeira glabra BLAKE & HILBIG, 1990

segment 1–3 larger than on following segments. Dorsal cirri


Size: No complete specimens known. Anterior end of 21 seg-
connected by several muscle bands. Ventral cirri on segment 4
ments 5.6 mm long.
very small. Neuropodial lobes and neurochaetae from segment
Morphology: Body shape cylindrical with flattened venter. 5, notopodial lobes and notochaetae from segment 6. No-
Prostomium rounded rectangular, wider than long, with anteri- tochaetae include capillaries with two rows of teeth and a few
or median incision (Fig. 1A). Palpophores prominent, annulat- furcate chaetae. Neurochaetae all compunds, with indentate
ed, palpostyles tapered with pointed tips. Paired antennae tips. Ventral cirri inserted distally on neuropodium.
cylindrical with short pointed tips, median antenna small,
Distribution: Juan de Fuca Ridge: Axial Seamount.
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

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

References:
BLAKE J.A. & B. HILBIG (1990) Pac. Sci. 44: 219-253.
PLEIJEL F. (1998) Zool. Scr. 27: 89–163.

F. PLEIJEL Denisia 18 (2006): 199


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Annelida, Polychaeta, Phyllodocida, Hesionidae

Hesiolyra bergi BLAKE, 1985 “caterpillar worm“

large number of lyriform, tapered spinulose, and subdistally en-


Size: Up to 80 mm for 100 segments.
larged (and often twisted) spinulose. Neurochaetae all com-
Morphology: Body anteriorly truncate and posteriorly tapered, pound, dark to black in color (possibly due to some inclu-
dorso-ventrally flattened. Prostomium rounded rectangular, sions?), with unidentate tips. Ventral cirri inserted medially on
slightly wider than long. Palpophores short, stout; palpostyles neuropodium, with distinct cirrophores.
much longer and thinner than palpophores, cylindrical with
Biology: On smoker walls, often visiting tubes of Alvinella pom-
rounded tips. Paired antenna similar to palpostyles; median an-
pejana. Can occur in large numbers. Epibiotic filamentous bac-
tenna absent. Eyes absent. Outside of everted proboscis with
teria on parapodia and chaetae. Egg diameter about 125 µm.
pointed papillae, increasing in size distally. Terminal ring of
proboscis with large number of pointed papillae. Inside of pro- Distribution: East Pacific Rise: 9°N and 21°N, and a closely re-
boscis with large number of denticles in band on ventral side; lated species on Mid-Atlantic Ridge: at the Azores Triple
dorsal and ventral teeth present, dorsal much larger. Dorsal cir- Junction (expeditions Diva 1 and 2, Marvel) and 22°N
ri and cirrophores on segment 1–5 and ventral cirri and cir- (Microsmoke). Newly collected specimens from Mid-Atlantic
rophores on segment 1–4 larger and longer than on following Ridge agree well with H. bergi. Whether they should be referred
segments. Neuropodial lobes and neurochaetae from segment 5, to a new species or not cannot be stated at present; they are
notopodial lobes and notochaetae from segment 6. No- currently best referred to as “Hesiolyra sp.”.
tochaetae surrounded by large pre- and postchaetal lamellae,
smaller lamellae also surrounding neurochaetae. Notochaetae
of four kinds: one-two serrated flattened (difficult to detect),

2: Population on a smoker wall, visit-


ing Alvinella’s tubes (East Pacific Rise
13°N, 2630 m). Phare cruise  Ifremer.

1: Habitus in vivo; by P. Briand 3: Proboscis, frontal view (SEM)


 Ifremer.  Ifremer.

F. PLEIJEL Denisia 18 (2006): 200–201


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4A: Habitus; by V. Martin © Ifremer; B-D: Denticles of the proboscis; E: Left parapodium of
segment 13; by courtesy of J. Blake; F: Lyriform notochaeta; G: Tapered spinulose notochaeta;
H: Compound neurochaeta (B-D & F-H © F. Pleijel).

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.

201
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Annelida, Polychaeta, Phyllodocida, Hesionidae

Hesiospina vestimentifera BLAKE, 1985

rophores on segment 1–5 larger than on following segments,


Size: Up to 36 mm long for 45 segments.
ventral cirri on segment 1–4 larger and longer than on follow-
Color: Live specimens yellowish transparent, with yellow-or- ing segments, with distinct cirrophores. Neuropodial lobes and
ange gut, dorsal and parapodial blood vessels red. Preserved neuropodia from segment 5, notopodial lobes and notochaetae
specimens yellowish. (except hooks; see below) absent from all segments. Notopodia
from segment 8–9 to last segments with single stout, straight to
Morphology: Body anteriorly truncate and posteriorly tapered.
slightly curved protruding hook, emerging on frontal side of cir-
Prostomium rounded trapezoid, anteriorly wider (Fig. 5A).
rophore. Neuropodial lobes elongated triangular. Neurochaetae
Palpophores long; palpostyles much shorter than palpophores,
of two kinds: one to two stout, bent acicular chaetae, usually
varying in shape from ovoid to elongated, tips rounded. Paired
shorter than shafts of compounds, and c. 50 compounds with
antenna thin, evenly tapering to rounded tips, slightly shorter
unidentate tips. Ventral cirri inserted subdistally on neuropodi-
or of same length as palps. Median antenna absent. Eyes absent.
um, without distinct cirrophores.
Everted proboscis basally with papillae appearing in c. 10–15
poorly defined rows, with three to five conical, pointed papillae Biology: Commonly associated with the siboglinid Riftia. Egg
in each row. Papillae increasing in size distally. Anterior end of size 100–120 µm in diameter.
proboscis with 10 conical papillae. Dorsal and ventral pair of
Distribution: Explorer Ridge, Juan de Fuca Ridge, Galapagos
jaw plates inside terminal ring of papillae, both pairs with two
Spreading Center, and East Pacific Rise.
basally jointed lamellae, dorsal pair anteriorly rounded, ventral
pair pointed (PLEIJEL, 2004, fig. 10A, B). Dorsal cirri and cir-

1A: Anterior end, dorsal view with proboscis everted; B: Middle left parapodium, ante-
rior view; C: Notopodium, showing aciculum and protruding acicular spine; by courtesy
of J. Blake.

F. PLEIJEL Denisia 18 (2006): 202–203


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2: Habitus, dorsal view; by P. Briand  Ifremer. 3: Anterior part in vivo; by P. Briand  Ifremer.

4: Parapodia, dorso-lateral view  Ifremer. 3: Prostomium, frontal view  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
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Annelida, Polychaeta, Phyllodocida, Hesionidae

Nereimyra alvinae BLAKE, 1985

Size: Length of largest observed anterior fragment 3 mm for 20 Biology: Associated with mussels at the Galapagos Rift and
segments. sorted from sediment in box cores from the Guaymas Basin.
Morphology: Prostomium wider than long. Palpophores cylin- Remarks: Poorly known and requires redescription (the type
drical, palpostyles thinner, basally inflated with tapering tips. material is in bad condition and the above description is based
Paired antennae thinner and longer than palpostyles, median on the original one); the affinities with other members of
antenna absent. Eyes absent. Everted proboscis smooth, anteri- Nereimyra is uncertain.
or end with ring of papillae, jaws absent. Dorsal cirri and cir-
Distribution: Galapagos Spreading Center, Guaymas Basin.
rophores on segment 1–3 (possible 1–4) longer than on follow-
ing segments, ventral cirri (and cirrophores?) on segment 1–3
larger than on following segments. Noto- and neuropodial lobes
and noto- and neurochaetae from segment 4. Notochaetae all
capillaries, neurochaetae unidentate capillaries and com-
pounds. Ventral cirri inserted subdistally on neuropodium.

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

Reference:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.

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Annelida, Polychaeta, Phyllodocida, Hesionidae

Sirsoe grasslei (BLAKE, 1985)

segments and with developed cirrophores (also on segment 3).


Synonym: Orseis grasslei BLAKE, 1985.
Notopodial lobes and notochaetae absent from all segments.
Size: Length 8–9 mm for 32 segments (holotype). Neuropodial lobes and neurochaetae present from segment 2.
Neurochaetae all unidentate compounds. Ventral cirri subdis-
Morphology: Body shape anteriorly truncate and posteriorly ta-
tally inserted on neuropodium.
pered. Prostomium rounded rectangluar, slightly wider than
long. Palpophores short and large, palpostyles much longer, Remarks: Poorly known, requires redescription.
evenly tapered and pointed. Paired antennae similar to pal-
Biology: Densities of 2,844 individuals per m2 have been
postyles but slightly smaller, median antenna similar to paired
recorded from mats of Beggiatoa (PETRECCA & GRASSLE 1989).
ones but much smaller, inserted on dorsal, posterior part of
Eggs large, about 250 µm in diameter.
prostomium. Eyes absent. Everted proboscis smooth, unarmed,
anterior end with ring of 10 small pointed papillae. Dorsal cir- Distribution: Guaymas Basin.
ri of segment 1–2 or 1–3 (uncertain) larger than on following
segments, ventral cirri on segment 1–2 larger than on following

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

References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.
PETRECCA R.F & J.F. GRASSLE (1989) Proc. Gorda Ridge Symp.: 279-283.
PLEIJEL F. (1998) Zool. Scr. 27: 89-163.

F. PLEIJEL Denisia 18 (2006): 205


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Annelida, Polychaeta, Phyllodocida, Nautiliniellidae

Iheyomytilidicola tridentatus MIURA & HASHIMOTO, 1996

edge of the main fang; occurring up to five on each parapodi-


Size: Up to 20 mm in length. Up to 2.3 mm in width.
um. Additional neurochaetae two kinds; tridentate chaetae on
Color: Preserved specimens pale. outer base of chaetal lobes, simple slender than neuropodial
hooks, shorter than other chaetae, with blunt main tooth on
Morphology: Body flattened ventrally and arched dorsally, with
one side of distal tip and two shorter blunt teeth on opposite
106 segments. Prostomium short with a pair of short cirriform
side, occurring 10-20 per parapodium; minute slender chaetae
antennae, without eyes. Second or tentacular segment com-
on inner side of chaetal lobe, enlarged subdistally, with single
pletely fused to prostomium, lacking dorsal cirri and neu-
distal mucronate spin, longer and more numerous than triden-
rochaetae, defined by ventral cirri. Parapodia subbiramous
tate chaetae. Pygidium rounded, without anal cirri.
throughout the body, with very short dorsal and ventral cirri.
Notopodia slightly inflated, supported by single stout acicula. Biology: Living in the mantle cavity of the deep-sea mussel Ba-
Neuropodia supported by a single stout neuroacicula, project- thymodiolus aduloides. Foregut with well muscularized part; sto-
ing from the parapodial base, slightly elongated on middle and modeum may be eversible. The species is thought to be para-
posterior segments. Ventral cirri on ventro-posterior side of sitic.
neuropodia, decreasing in size on middle and posterior parapo-
Distribution: Okinawa Trough.
dia. Neuropodial hooks on dorsal side of chaetal lobes, stout,
slightly inflated subdistally, with minute projection on cutting

1A: Anterior end, dorsal view; B: Posterior end, dorsal view; C: 2A: Neuropodial hooks; B: Tridentate chaetae; C: Minute mu-
Anterior end, ventral view; D: Same enlarged; E: Parapodium cronate chaetae; D: Anterior end of a juvenile; from MIURA &
of the 12th segment (or parapodium 11); F: Parapodium 40, HASHIMOTO (1996).
anterior view; from MIURA & HASHIMOTO (1996).

Reference:
MIURA T. & J. HASHIMOTO (1996) Publ. Seto Mar. Biol. Lab. 37(3/6): 257-274.

T. MIURAS Denisia 18 (2006): 206


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Annelida, Polychaeta, Phyllodocida, Nautiliniellidae

Mytilidiphila enseiensis MIURA & HASHIMOTO, 1993

short ventral cirri and stout acicula; ventral hooks simple, thin,
Size: Up to 28 mm in length. Up to 1.5 mm in width.
occurring more than 35 as maximal number of posterior para-
Color: Preserved specimens pale. podia. Pygidium simple, without anal cirri.
Morphology: Body long, vermiform, flattened ventrally and Biology: Living in the mantle cavity of a burrowing deep-sea
arched dorsally, with up to 164 segments; first segment tentac- mussel (Adula?). Foregut with well muscularized part; sto-
ular, chaetigerous. Prostomium short with a pair of short cirri- modeum may be eversible. The species is thought to be para-
form antennae, without eyes. Second segment well fused with sitic.
prostomium, defined by embedded acicula and ventral cirri as
Distribution: Okinawa Trough.
well as projected simple hooks. Notopodia very short, with dor-
sal cirri and stout acicula. Neuropodia cylindrical, with very

2: Anterior part in dorsal view;


by T. Miura.

1A: Anterior end, dorsal view; B: Posterior end, dorsal view; C: Anterior
end enlarged, dorsal view; D: Same, ventral view; E: Parapodium of the 11th
segment, anterior view; F: Parapodium of the 27th segment, anterior view; 3: Details of the prostomium;
G: Hooks; from MIURA & HASHIMOTO (1993). by T. Miura.

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


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Annelida, Polychaeta, Phyllodocida, Nautiliniellidae

Mytilidiphila okinawaensis MIURA & HASHIMOTO, 1993

short ventral cirri and stout acicula; ventral hooks simple, thin,
Size: Up to 15 mm in length. Up to 1.4 mm in width.
ocuring more than 30 as maximal number on posterior parapo-
Color: Preserved specimens pale. dia. Pygidium rounded, without anal cirri.
Morphology: Body long, vermiform, flattened ventrally and Biology: Living in the mantle cavity of a deep-sea mussel Ba-
arched dorsally, with about 100 segments; first segment tentac- thymodiolus japonicus. Foregut with well muscularized part; sto-
ular, achaetous. Prostomium short with ventrally a pair of cirri- modeum may be eversible. The species is thought to be para-
form antennae, without eyes. Second or tentacular segment sitic.
well fused with prostomium, achaetous, defined by embedded
Distribution: Okinawa Trough.
acicula and ventral cirri. Notopodia very short, with dorsal cir-
ri and stout acicula. Neuropodia short cylindrical, with very

1A: anterior end, dorsal view; B: Same enlarged, dorsal view; C: Same, ventral view; D: Posterior end, dorsal
view; 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).

References:
HASHIMOTO J., OHTA S., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. 42: 577-598.
MIURA T. & J. HASHIMOTO (1993) Zool. Soc. 10: 169-174.

T. MIURA Denisia 18 (2006): 208


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Annelida, Polychaeta, Phyllodocida, Nautiliniellidae

Shinkai longipedata MIURA & OHTA, 1991

simple, stout, 2-10 as maximal number on anterior parapodia or


Size: Up to 200 mm in length. Up to 6.0 mm in width.
occuring singly on successing parapodia.
Color: In alcohol light greenish brown; parapodia darker than
Biology: Living in the mantle of a giant clam species of the
trunk (small individuals pale or colorless).
genus Calyptogena. Foregut with well muscularized part; sto-
Morphology: Body long, vermiform, flattened ventrally and modaeum may be eversible. The species is thought to be para-
arched dorsally, with about 242 segments (excluding preanal sitic.
achaetous segments); first segment tentacular, chaetigerous.
Distribution: Okinawa Trough.
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

1: Ventral view of the anterior part. 3: Female as found in the clams’ mantle cavity; by courtesy
of T. Miura.

4: Male as found in the clams’ mantle cavity; by courtesy of


2: Variations in parapodia and chaetae. T. Miura.

References:
MIURA T. & J. HASHIMOTO (1996) Publ. Seto Mar. Biol. Lab. 37(3/6): 257-274.
MIURA T. & S. OHTA (1991) Zool. Soc. 8: 383-387.

T. MIURA Denisia 18 (2006): 209


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Annelida, Polychaeta, Phyllodocida, Nautiliniellidae

Shinkai semilonga MIURA & HASHIMOTO, 1996

Neuropodial hooks simple, stout, slightly curved on distal end;


Size: Up to 100 mm in length. Up to 3.9 mm in width.
number of hooks per parapodium, about 15 on parapodia 1-3,
Color: The female is deep purplish brown on parapodia and the more than 25 on parapodia 4-6 as maximum, then decreasing
male whitish. to reach about ten on parapodium 10, 5-8 on parapodia 50-200.
Morphology: Body long, vermiform, flattened ventrally and Remark: A closely related species has been sampled at Manus
arched dorsally, with about 267 segments (excluding preanal Back-Arc Basin: Desmos Cauldron (Shinkai 2K # 916). It dif-
achaetous segments). Prostomium short, with a pair of short an- fers from S. semilonga by the shape of the neuropodial hooks,
tennae, without eyes. Tentacular segment well fused with pros- which have a tridentate apical end.
tomium, with dorsal cirri and ventral cirri, neuroacicula, and
Biology: Living in the valve cavity of a giant clam Calyptogena
numerous neuropodial hooks. Foregut with strongly muscular-
solidissima. The species is thought to be parasitic.
ized part. Pygidium rounded, without anal cirri.
Parapodia subbiramous, with dorsal and cirri. Notopodia slight- Distribution: Okinawa Trough: Minami-Ensei Knoll.
ly elongated, supported by single, very thin notoacicula; dorsal
cirri short, on distal end of notopodia. Neuropodia well devel-
oped, supported by single stout neuroacicula; ventral cirri on
ventro-posterior side of neuropodia, shorter than dorsal cirri.

1A: Anterior end, dorsal view; B: Same, ventral view; C: Same, lateral view; D: Posterior end, dorsal view; E: Neuropodial hooks
from parapodia 51 (right) and 200 (left); F: Parapodium; from MIURA & HASHIMOTO (1996).

T. MIURA Denisia 18 (2006): 210–211


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2: Shinkaia sp., middle segments dorsal view (SEM); Manus 3: Shinkaia sp., distal view of posterior hooks (SEM); Manus
Basin © Ifremer. Basin © Ifremer.

4: Shinkaia sp., tridentate hooks from anterior parapodia 5: Shinkaia sp., tridentate hooks from anterior parapodia
(SEM); Manus Basin © Ifremer. (SEM); Manus Basin © Ifremer.

Reference:
MIURA T. & J. HASHIMOTO (1996) Publ. Seto Mar. Biol. Lab. 37(3/6): 257-274.

211
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Annelida, Polychaeta, Phyllodocida, Nereididae

Nereis piscesae BLAKE & HILBIG, 1990

basally, tapering distally; ventral cirrus narrow, cirriform. No-


Size: A large, robust species, some fragments up to 5 cm long;
topodial acicular lobes small in both anterior and posterior seg-
largest complete specimens 75 mm long, 7 mm wide, for 100
ments. Neuropodial acicular lobes conical throughout. Four
segments.
types of chaetae present. Anterior notochaetae all homogomph
Morphology: Prostomium triangular, narrowing anteriorly to spinigers replaced from segments 30-35 by 1-2 heavy ho-
rounded tip bearing two short antennae; eyes absent, but two mogomph falcigers with elongated blades bearing 6-8 blunt
pairs of sunken depressions representing positions of missing denticles [not seen in specimens from type locality studied by
eyes present; palps thickened, fleshy, each bearing two finger- D. Desbruyères, probably worn falcigers]; upper neurochaetae
like palpostyles. Peristomium enlarged, inflated, surrounding including heterogomph spinigers and heterogomph falcigers;
prostomial structures; four pairs of tentacular cirri. Jaws golden, lower neurosetae include homogomph spinigers and hetero-
pointed, each with 78 teeth. Paragnaths all conical, arranged as gomph falcigers; blades of all spinigers with thin denticles on
follows: I – two large cones in vertical row; II – 12 large cones cutting edge; falcigers with thin, pointed denticles; shafts of all
in nearly circular arc; III – 21-22 small and large cones in clus- chaetae smooth; internally ornamented with dark brown to
ter; IV – 19-20 small and large in three distinct clusters black camerations.
arranged in three transverse rows; V – 0; VI – 46 large pointed
Biology: An epifaunal species associated with siboglinids. The
cones; VII – two small cones; VIII – two groups of two small
species is probably an omnivore, feeding on filamentous bacte-
cones. Parapodia of middle and posterior segments with upper
ria.
notopodial ligule becoming enlarged and flattened, with dis-
tinct dorsal curvature and tapering triangular tip, with thick- Distribution: Juan de Fuca Ridge.
ened glandular border, bearing thin dorsal cirrus; lower notopo-
dial ligule tapering, not enlarged; neuropodial ligule thickened

1: Anterior end, ventral view (SEM) © Ifremer. 2: Anterior end, lateral view (SEM) © Ifremer.

J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 212–213


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3: Enlarged notopodial ligule of middle segments (SEM) 4: Anterior notochaetae homogomph spinigers (SEM)
© Ifremer. © Ifremer.

5: Heterogomph falcigers of 6: Heavy homogomph falciger of poste- 7: Lower neurochaetae of posterior segments in-
upper neurochaetae (SEM) rior notopods (SEM) cluding heterogomph falcigers and homogomph
© Ifremer. © Ifremer. spinigers (SEM) © Ifremer.

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

213
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Annelida, Polychaeta, Phyllodocida, Nereididae

Nereis sandersi BLAKE, 1985

ent: anterior chaetae homogomph spinigers replaced by 4-6


Size: Largest specimen 95 mm.
heavily homogomph falcigers. Lower neurochaetae include ho-
Morphology: Parapodia elongated in posterior segments, pro- mogomph spinigers and heterogomph falcigers; blades of all
viding ragged appearance to worm. Body terminated in two spinigers with thin denticles on cutting edge.
long anal cirri, each with basal swelling. Jaws black to golden,
Biology: On smoker’s walls, dwelling inside active/inactive de-
with 10-12 blunt teeth. Paragnaths all conical, arranged as fol-
posits. Some nereids are carnivorous, others are grazers of bac-
lows: I – 4-6 in vertical cluster; II – about 30 small cones in an
terial mats.
arc; III, IV – 85-100 small cones in dense field; V – 0; VI 4-5
dark large, curved cones in cluster; VII – two irregular rows of Distribution: Galapagos Spreading Center, Northern East Pa-
about 10 cones each; VIII – 1-2 large colored cones and 3-4 cific Rise, Guaymas Basin.
cones. Parapodia of segments 1-2 uniramous, following parapo-
dia biramous with two black acicula. Four types of chaetae pres-

1: Anterior part; by 2: Anterior part in 3: Prostomium, frontal view (SEM)  Ifremer.


V. Martin  Ifremer. vivo  Ifremer.

6: Posterior parapodia elon-


gated in vivo  Ifremer.

4: Specimen on a smo- 5: Everted proboscis showing jaws 7: Homogomph falcigers


ker wall (EPR 13°N); and paragnaths (SEM)  Ifremer. (SEM)  Ifremer.
cruise Phare  Ifremer.

Reference:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 6: 67-101.

D. DESBRUYÈRES Denisia 18 (2006): 214


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Annelida, Polychaeta, Phyllodocida, Phyllodocidae

Eulalia papillosa (BLAKE, 1985)

each with shaft bearing single medial tooth and fine lateral
Size: Up 13 mm long, 0.8 mm wide, for 75 segments.
denticles, blade with finely serrated cutting edge. Pygidium
Color: In alcohol opaque white with brown pigment granules with two long broad flattened cirri.
on antennae, cirri, and body surface.
Remarks: E. papillosa was originally described as a species of
Morphology: Prostomium pear-shaped, slightly wider than long Protomystides. However, the median antenna was misinterpret-
with four frontal antennae; with short, clavate median antenna ed as a papilla and when correctly interpreted, the species
on posterior one-fourth. With four pairs of tentacular cirri on agrees with the definition of Eulalia.
three segments with following formula: 1 +S1/1 + S1/n; ventral
Biology: An epifaunal species associated with Riftia and Calyp-
tentacular cirrus of segment 2 short, broad, pointed; other three
togena. The species probably feeds on small epifaunal inverte-
tentacular cirri long, tapering, cirriform. Dorsal cirri from seg-
brates.
ment 4 longer than wide, oval-shaped; ventral cirri asymmetri-
cal, not pointed, shorter than podial lobe. Parapodia uniramous Distribution: East Pacific Rise at 21°N.
with single internal aciculum and 10-12 compound spinigers,

1A: Anterior end, dorsal view with pharynx everted; B: Left parapodium from posterior region, anterior view; C-D: Composite
chaetae; from BLAKE (1985).

Reference:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.

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Annelida, Polychaeta, Phyllodocida, Phyllodocidae

Galapagomystides aristata BLAKE, 1985

bearing 4-8 heavy compound chaetae with heavy shaft and


Size: Up to 20 mm long, 1.2 mm wide, with about 90 segments.
thin, aristate-like narrow blade. Pygidium with pair of long,
Morphology: Color dark brown to light tan, with dark brown thick cirri.
spots on prostomium, antennae, tentacular cirri, and dorsal
Biology: An epifaunal species associated with siboglinid tubes
ventral cirri. Prostomium wider than long bearing four long,
and mussels. The species is probably a scavenger, or a possible
cirriform frontal antennae shifted ventrolaterally. Proboscis
predator on small invertebrates. This worm was proposed to be
narrow, smooth. Segment 1 dorsally fused to prostomium, free
an hematophagous.
ventrally; segments 2-3 distinct dorsally and ventrally. Three
pairs of long, cirriform tentacular cirri with following formula: Distribution: Galapagos Spreading Center; East Pacific Rise:
1 + S1/1 +S0/n. Segment 3 without dorsal cirrus. Dorsal cirri from 13°N, 9°N; common species.
segment 4 long, thickened, fleshy, tapering distally. Ventral cir-
ri small, oval subequal to podial lobe. Parapodia uniramous

1: Habitus in vivo (dorsal); 2: Ventral view (SEM) 3: Middle parapodia, right side
by P. Briand  Ifremer.  Ifremer. (SEM)  Ifremer.

4: Shaft of a composite chaetae (SEM) 5: Aristate-like composite chaetae (SEM)


 Ifremer.  Ifremer.

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.

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Annelida, Polychaeta, Phyllodocida, Phyllodocidae

Protomystides verenae BLAKE & HILBIG, 1990

both dorsal and ventral cirri with dense concentrations of


Size: Up to 27 mm long, 2.1 mm wide, for 75 segments.
brown pigment spots. Parapodia uniramous bearing single acic-
Color: Body dark brown to light tan with dark spots on pros- ula and fascicle of 8-12 compound spinigers of two types: (1)
tomium, all cirri, and body segments. large, heavy, brown in color, (2) small, thin, non-colored; both
with shaft bearing large, pointed terminal tooth and subapical
Morphology: Prostomium trapezoidal, broadly rounded anteri-
notch from which thin blade emerges; tip of shaft with fine
orly, as wide as long, V-shaped posteriorly; with four long, cirri-
teeth along one edge. Pygidium bearing two long, cirriform anal
form antennae, ventral pair shifted ventrolaterally. Anterior
cirri.
segments free from one another and prostomium. With three
pairs of long, tapering, cirriform tentacular cirri with following Biology: The species is likely an epifaunal associate of vesti-
formula: 1 + S1/1 + S1/v. Dorsal cirri from segment 4, each short, mentiferans and other larger invertebrates found at vent sites.
oval, thickened, about 2-3 times as long as wide. Dorsal cirri of The species is probably predatory on small invertebrates.
anterior segments long, extending well beyond podial lobe;
Distribution: Explorer Ridge and Juan de Fuca Ridge.
those of middle segments smaller, shifted to higher location on
body wall; ventral cirri shorter than ventral cirri throughout;

1A: Anterior end in dorsal view; B: Right middle parapodium; C-F: Composite chaetae; 2: Habitus; scale bar 1 mm;
from BLAKE & HILBIG (1988). by courtesy of V. Tunnicliffe.

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

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Annelida, Polychaeta, Phyllodocida, Polynoidae

Bathykurila guaymasensis PETTIBONE, 1989

projecting acicular lobes on lower side. Neuropodia with coni-


Size: Up to 8 mm in length.
cal projecting prechaetal acicular lobe and shorter, rounded
Color: Pinkish red. postchaetal lobe. Notochaetae numerous, short to longer form-
ing radiating bundles stouter than neurochaetae, with row of
Morphology: Body fusiform, flattened, with 15 segments (first
widely spaced teeth along one side and blunt tips. Neu-
achaetous). Elytra seven pairs on segments 2, 4, 5, 7, 9, 11, and
rochaetae very numerous forming fan-shaped bundles; neu-
13, large, oval, covering dorsum, delicate, with larger conical to
rochaetae with two rows of long spines. Pair of long ventral
rounded tubercles variable in size on posterior and lateral bor-
papillae present in males on segment 11, sometimes extending
ders with smaller ones scattered on surface. Prostomium oval,
far beyong posterior end. Posterior two segments smaller, en-
deeply bilobed with triangular anterior continuations bearing
closed in elytragerous segment 13. Pygidium long with pair of
small frontal filaments. First or tentacular fused to prostomium,
slender short anal cirri.
not visible dorsally; tentaculophores lateral to prostomium,
each with prominent acicular lobe on inner side, without Biology: Found also on whale carcasses.
chaetae, and pair of subulate tentacular cirri, shorter than
Distribution: Guaymas Basin.
palps; upper lip large bilobed. Notopodia rounded basally with

3: Prostomium, dorsal view (SEM);


by courtesy of A. Glover.

1: Dorsal view in vivo; 2: Ventral view in vivo;


by courtesy of A. Glover. by courtesy of A. Glover.

4: Elytra (SEM); 5: Distal part of sexual male ventral 6: Anterior end, ventral view (SEM);
by courtesy of A. Glover. papillae; (SEM); by courtesy of A. by courtesy of A. Glover.
Glover.

Reference:
PETTIBONE M. (1989) Proc. Biol. Soc. Wash. 102(1): 154-168.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 218


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchinotogluma hessleri PETTIBONE, 1985

posterior two segments, with pair of anal cirri. Six pairs of small
Synonym: Opisthotrochopodus alvinus PETTIBONE, 1985.
rounded segmental lamellae near ventral bases of neuropodia of
Size: Up to 16 mm. segments 11-16.
In males (described as Opisthotrochopodus alvinus), segment 12
Morphology: Segments 21. Body flattened, tapering slightly
with a greatly extended papilla, parapodia of posterior four seg-
anteriorly and posteriorly, with parapodia longer than body
ments modified and directed posteriorly differing from one an-
width. Dorsum with transverse ciliated bands. Elytra 10 pairs.
other. Parapodia of chaetigerous segment 18 smaller than pre-
Elytra covering dorsum, round to oval with branched “veins“.
ceding and following parapodia. Notopodium represented as a
Dorsal cirri with short cylindrical cirrophores and long tapered
small acicular lobe with a bundle of short smooth acicular
smooth styles extending beyong tips of chaetae. Groups of ar-
chaetae and delicate rounded flattened lamella. Neuropodium
borescent branchiae attached on the lateral sides of elytro-
without neurochaetae. Modified parapodia of segment 19 with-
phores and dorsal tubercles. Branchiae begining on segment 3
out chaetae with small elytrophore and elongate-oval elytron.
as two small groups and becoming larger posteriorly. Prostomi-
Parapodium 20 greatly modified. Notopodium represented by
um bilobed, oval anterior lobes triangular with delicate frontal
achaetous elongated cylindrical acicular lobe with distal style of
filaments. Median antenna with bulbous ceratophore. Palp
dorsal cirrus. Neuropodium forming large cyclindrical acicular
stout, tapering, smooth. First segment with two tentacular cir-
lobe with projecting conical acicular process and enclosing cir-
ri. Notopodium with hood or bract encircling small bundle of
cle of chaetae of several types. Segment 21 with right and left
chaetae. Everted pharynx with three dorsal and two ventral
parapodia closely approximated medially and directed posteri-
papillae, two pairs of prominent jaws with minute denticles on
orly. Notopodium consisting of expanded thin lamella and
the inner border. Notopodia with short conical acicular lobes
thickened acicular lobes without notochaetae. Neuropodium
and on elytrigerous segments, with prominent bracts enclosing
also with expanded thin lamella and thickened acicular lobes
notochaetae dorsally and posteriorly. Notochaetae acicular
with a small bundle of needle-like neurochaetae. Pygidium con-
much stouter than neurochaetae, smooth or with 1-2 rows of
sisting of small ovale lobe without anal cirri.
short spines. Neurochaetae slender, very numerous, forming
fan-shaped bundles. Few upper neurochaetae with more promi- Biology: Found in Riftia and Calyptogena washes.
nent spines, with tip flattened and finely spinous. Tips slightly
Distribution: East Pacific: from Guaymas Basin to East Pacific
hooked, with finely spinous hood.
Rise 9°N, Galapagos Spreading Center.
In females pygidium consisting of small squarish, to rounded
lobes wedged between bases of posteriorly-directed parapodia of

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


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

3: Prostomium, apical view (SEM) 4: Jaws and proboscis papillae, apical view
© Ifremer. (SEM) © Ifremer.

5: Notochaetae (SEM) © Ifremer.

6: Ventral view of the posterior part of a 7: Modified segments 18-21, ventral view 8: Upper neurochaetae (SEM)
male specimen showing the elongated papil- (SEM) © Ifremer. © Ifremer.
la of the segment 12 (SEM) © Ifremer.

9: Parapodium (SEM) 10: Hooked neurochaetae from chaetiger 20 11: Bract second segment (SEM)
© Ifremer. (SEM) © Ifremer. © Ifremer.

Reference:
PETTIBONE M.H. (1985) Proc. Biol. Soc. Wash. 98: 447-469.

220
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchinotogluma marianus (PETTIBONE, 1989)

segment 15 more or less modified. Parapodia 16 and following


Synonym: Opisthotrochopodus marianus PETTIBONE, 1989.
parapodia without branchiae. Parapodium 17 with a neu-
Size: Up to 6 mm in length. Up to 3 mm in width. ropodium shorter than ventral cirrus. Parapodium 18 with
small notopodial acicular lobe fused to cirrophore, conical neu-
Morphology: Body flattened, with 21 segments (first
ropodium shorter than ventral cirrus. Parapodium 19 with a
achaetous). Dorsum with transverse ciliated bands. Elytra 10
small elytron, inflated oval notopodium; neuropodium with
pairs on segments 2, 4, 5, 7, 9, 11, 13, 15, 17 and 19. Elytra cov-
small bundle of long capillary neurochaetae. Parapodium 20 di-
ering the dorsum, except for posterior modified segments (ely-
rected posteriorly, with small notopodial acicular lobe fused to
tra very small on segments 17 and 19). Branchiae delicate, ar-
cirrophore of dorsal cirrus. Notochaetae long, wider subdistally
borescent, on lateral bases of elytrophores and dorsal tubercles
spinuous, tapering to capillary tips. Parapodium 21 directed
and dorsal bases of notopodia, beginning on segment 3 as two
posteriorly, with notopodial acicular lobe fused to cirrophore of
small groups, becoming larger and continuing to segment 15 as
the dorsal cirrus, without notochaetae; neuropodial conical aci-
a small group. Muscular pharynx encircled distally by four pairs
cular lobe with slender capillary neurochaetae and a large oval
of papillae, dorsal with two medial ones larger and lateral ones
lateral lamella.
smaller, ventral with four small papillae; two pairs of curved
reddish jaws occupying most of opening, without denticles on Remark: A closely related species was sampled in North Fiji
inner border. Parapodia 2-16 biramous with notopodia shorter Back-Arc Basin. It differs from PETTIBONE’s (1989) original de-
than neuropodia. Notopodia long, conical with notochaetae scription by the shape of the muscular pharynx which is encir-
short smooth, or longer with faint spinuous rows. Larger neu- cled by four double large papillae instead of four pairs of papil-
ropodia with prechaetal lobe long, subconical, with projecting lae, with two dorsomedial ones larger.
acicular process, postchaetal lobe shorter rounded. Neu-
Biology: Female unknown.
rochaetae with slightly hooked tips. Elongated ventral papillae
on segment 12 extending posteriorly to segment 15. six pairs of Distribution: Mariana Back-Arc Basin.
short, flat ventral lamellae on segments 13-18. Parapodia from

D. DESBRUYÈRES Denisia 18 (2006): 221–222


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

1: Muscular pharynx in distal view (SEM) 2: Posterior chaetigers (ventral view) showing the transformed lateral
© Ifremer. lamella of the 21st segment (SEM) © Ifremer.

3: Ventral papillae of the 12th segment (SEM) © Ifre- 4: Long notochaetae of the 20th segment, wider subdistally spinuous and
mer. tapering to capillary tips © Ifremer.

1-4: Branchinotogluma aff. marianus from North Fiji Back-Arc Basin; cruise Starmer.

Reference:
PETTIBONE M.H. (1989) Proc. Biol. Soc. Wash. 102(1): 137-153.

222
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchinotogluma sandersi PETTIBONE, 1985

short and acicular, superior neurochaetae with two rows of


Synonym: B. grasslei PETTIBONE, 1985.
prominent spines and a slightly hooked tip, lower neurochaetae
Size: Up to 60 mm in length. with flattened spinous hooked hood. Parapodia of the posterior
three segments modified directed posteriorly and enclosed in
Color: Pink to light red in living specimens. Golden-colored
parapodia of segment 18, all with ventral cirri but branchiae
chaetae.
lacking. Parapodia of elytrigerous segment 19 biramous with ra-
Morphology: Body rectangular flattened, tapering slightly an- mi similar in length; notopodia with thickened dorsal bract.
teriorly and posteriorly. Dorsum with transverse ciliated bands. Parapodia of segment 20 much smaller and enclosed in parapo-
Elytra 10 pairs, large covering dorsum, round to oval, stiff, dia of segment 19. Notopodial acicular lobe fused to cirrophore
opaque. Each segment from 4rd chaetiger with two groups of del- of dorsal cirrus. Notochaetae few, short, stout, curved with two
icate arborescent compact branchiae, with terminal filaments. rows of distal spines. Upper neurochaetae with scattered long
A single group of branchiae on 3rd segment. Prostomium spines and rounded tips. Segment 21 similar to 20th but lacking
bilobed, anterior lobes cylindrical with a terminal filament. notochaetae. Four pairs of segmental ventral papillae on seg-
Median antenna with a short style. Palps stout tapered smooth. ment 12-15. Three pairs of rounded ventral lamellae on seg-
Tentaculophores with two pairs of tentacular cirri, dorsal ones ments 16-18.
as long as palps. Muscular pharynx showing five papillae around In females (described as B. grasslei) segments 19-21 not trans-
opening (3 dorsal and two ventral). Jaws denticled in their in- formed. With or without five pairs of small squarish segmental
ner border. ventral papillae near ventral bases of neuropodia 11-15.
In males, parapodia of segments 2-18 biramous. Notopodia con-
Distribution: Galapagos Spreading Center, East Pacific Rise:
ical with projecting acicular processes, without bracts except on
21°N to 17°S, Guaymas Basin.
segment 2. Neuropodia with long conical prechaetal acicular
lobes, postchaetal lobes rounded and shorter. Notochaetae

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

Reference:
PETTIBONE M.H. (1985) Proc. Biol. Soc. Wash. 98: 447-469.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 223–224


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

6: Upper neurochaetae of the 20th segment,


3: Frontal view of the prostomial part (SEM) © Ifremer. male (SEM) © Ifremer.

7: Notopodial bract on segment 2 (SEM)


4: Ventral view of the posterior segments, male (SEM) © Ifremer. © Ifremer.

5: Ventral view of the segmental papillae, male (SEM) © Ifremer. 8: Subacicular neurochaeta (SEM) © Ifremer.

224
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchinotogluma segonzaci (MIURA & DESBRUYÈRES, 1995)

squarish ventral lamellae on following segments 13-17; lacking


Size: Up to 49 mm in length. Up to 19 mm in width.
both papillae on female specimens. On males, parapodium of
Morphology: Body flattened, with 21 segments (first segment 20 modified with reduced notopodium and elongated
achaetous). Elytra 10 pairs on segments 2, 4, 5, 7, 9, 11, 13, 15, neuropodium, lacking notochaetae. Notopodial acicular lobe
17 and 19. Branchiae arborescent, separated into two groups on fused to cirrophore with ventral lamellar expansion and em-
segments 3-19, or on segments 3-21 on half specimens. Muscu- bedded notoacicula. Neuropodium with lamellar expansion
lar pharynx encircled distally by four pairs of small papillae, folding small bundle of chaetae. Parapodium of segment 21
subequal in size, bearing lateral lamellar expansions with four strongly modified on males, lacking chaetae. Notopodial acicu-
distal papillae, and numerous small papillae forming proximal lar lobe fused to cirrophore of the short conical dorsal cirrus,
band. Two pairs of jaws without denticulation on the inner bor- ventral lamellar expansion enlarged.
der. Dorsal cirri on non-elytrigerous segments with short cylin-
Biology: Free living polynoids, found on whitish parts of active
drical cirrophores, and short styles with rounded tips, extending
chimney walls. Active predators of shrimps and bacterial mat
to tips of neurochaetae. Notopodia subconical, with projecting
grazers.
acicular lobe. Neuropodia subconical, deeply notched on upper
part. On males, pair of segmental papillae at the most extend- Distribution: North Fiji, Lau and Manus Back-Arc Basins.
ing to the base of ventral cirrus on segment 12; five pairs of

1: Specimen on active smoker walls; cruise TUIM07, Lau 2: Prostomium in dorsal view (SEM). Notice the proximal band of
Basin; by courtesy of C.R. Fisher. papillae on the pharynx © Ifremer.

T. MIURA Denisia 18 (2006): 225–226


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

3: Second left parapodia in dorsal view, with the typical dor- 4: Ventral view of the left side of the 12th and 13th segments
sal cirrus (SEM) © Ifremer. in male showing the ventral papilla and the first flat semioval
ventral lamella (SEM) © Ifremer.

5: Upper part of the pharynx with two dorsal rounded papil- 6: Lower part of the pharynx showing two small papillae
lae (SEM) © Ifremer. (SEM) © Ifremer.

7: Ventral view of the posterior end in male (SEM) © Ifremer. 8: Ventral view of the posterior end in female (SEM) © Ifremer.

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

226
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchinotogluma trifurcus (MIURA & DESBRUYÈRES, 1995)

present on segments 13-17. Segments 18-21 modified with re-


Size: Small species up to 15 mm in length and 7 mm in width.
duced parapodia. Segment 18 with very long dorsal cirri. Seg-
Color: Bright red in vivo. Preserved specimens pale. ment 19 with achaetous notopodia bearing broad dorsal lamel-
lar expansion. Segment 20 modified strongly, with long
Morphology: Body short with 21 segments, including the ten-
achaetous notopodia bearing bifurcate distal end, dorsal digiti-
tacular segment. Elytra 10 pairs, large imbricated, oval smooth.
form expansion and ventral fringed lamella, club-shaped neu-
Dorsal cirri with very long cirriform styles and tapered tips.
ropodia fringed by trifurcate neurochaetae. Segment 21 re-
Branchiae arborescent in two groups, present on segments 3-21
duced in size achaetous, with uniramous parapodia, notopodia
(females) or 3-20 (males). Prostomium bilobed. Anterior lobes
digitiform with very short cirri and basal lamellar expansion.
prominent, without frontal filaments. Tentacular segment
Anal cirri short.
achaetous not distinct dorsally. Muscular pharynx encircled by
Females lack posterior modified parapodia and ventral lamel-
four large papillae, subequal in size; two pairs of jaws with den-
lae, but bear ventral papillae on segment 11.
ticulations on inner border. Notopodia long, conical, with sev-
eral notochaetae; neuropodia long, conical, deeply notched on Biology: Very often found in washings of Ifremeria nautilei; no
upper part. evidence of co mmensalisms.
In males, ventral segmental papillae on segment 12, as long as
Distribution: North Fiji and Lau Back-Arc Basins.
segment; five pairs of flat semioval ventral segmental lamellae

3: Trifurcate neurochaetae
1: Living specimens dorsal and ventral view; by courtesy of F. 2: Ventral papillae on segment from segment 20, male (SEM)
Pleijel. 11, female (SEM) © Ifremer. © Ifremer.

4: Anterior part in dorso-anterior view 5: Ventral view of the posterior part, fe- 6: Dorsal view of the posterior part, male
(SEM) © Ifremer. male (SEM) © Ifremer. (SEM) showing the transformed segments
© Ifremer.

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

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 227


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchinotogluma tunnicliffae (PETTIBONE, 1988)

on anterodorsal side of a large neuropodium. Pygidium consist-


Synonym: Opisthotrochopodus tunnicliffae PETTIBONE, 1988,
ing of small squarish lobe with pair of anal cirri.
Branchinotogluma grasslei sensu PETTIBONE, 1988 (specimens
In males (described as Opisthotrochopodus): Biramous parapodia
from Northeast Pacific).
on segments 3-17 similar with smaller notopodium on an-
Size: Up to 31 mm. terodorsal side of a large neuropodium. Notochaetae stouter
than neurochaetae acicular with two rows of spines. Neu-
Morphology: Segment 21, elytra 10 pairs. Body flattened ven-
rochaetae long, slender with slightly hooked tips. Cirrigerous
trally, arched dorsally, with parapodia longer than body width.
parapodium of segment 18 much smaller, with long dorsal cir-
Elytra large, overlapping covering dorsum. Elytra smooth, stiff,
rus projecting posteriorly and single arborescent branchiae.
somewhat vaulted around place of attachment to elytrophores.
Segment 19 with smaller elytrophores approaching medially,
Elytrophores large, projecting posteriorly, with arborescent
with small parapodia hidden from view dorsally, notopodia with
branchiae attached on lateral sides. Dorsal tubercules elongate,
delicate lamella on dorsal side, notochaetae stout acicular with
projecting posteriorly with arborescent branchiae on distal
two rows of spines. Segment 20 with parapodia greatly modi-
part. Prostomium bilobed, anterior lobes subtriangular, with
fied. Neuropodia enlarged, inflated with projecting conical aci-
long frontal filaments; median antenna with bulbous cer-
cular lobe with neurochaetae projecting from wide opening and
atophore in anterior notch, subulate style with long slender tip,
ventral cirrus attached near the base. Neurochaetae consisting
shorter than tentacular cirri; palp stout, tapered, smooth. Ten-
of four stout acicular harpoon chaetae and long slender
taculophores lateral to prostomium, without chaetae, each with
chaetae. Neuropodia inflated and extended according to the
pair of tentacular cirri, dorsal tentacular cirrus longer than palp,
specimens (even according to the side of the same specimen).
ventral one shorter. Second segment forming posterior lip of
Segment 21 with elongated cirrophores fused medially and to
the mouth, bearing biramous parapodia and first pair of
the notopodial lobe. Ventral segmental papillae long tapering
elytrophores. Notopodia with hood or bract on anterodorsal
on segment 12, shorter and thicker with slender distal part on
side encircling small bundle of notochaetae and conical acicu-
segments 13-15 reduced to a rounded lamellae on 16 and 17.
lar lobe; ventral buccal cirri similar to tentacular cirri longer
than following ventral cirri. Muscular pharynk showing five Distribution: Explorer Ridge; Juan de Fuca Ridge: Endeavour
papillae around opening (3 dorsal and 2 ventral); two paires of segment; Axial Seamount and Southern Juan de Fuca Ridge.
jaws minutely denticulated.
In females (described as Branchinotogluma grasslei): Biramous
parapodia on segments 3-21 similar with smaller notopodium

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

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 228–229


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

2: Proboscis, left anterior view showing the jaws 3: Posterior part, male (SEM)
(SEM) © Ifremer. © Ifremer.

4: Parapodium (SEM) © Ifremer. 5: Neurochaetae 6: Stout acicular harpoon chaetae of


(SEM) © Ifremer. the 20th segment (SEM) © Ifremer.

Reference:
PETTIBONE M.H. (1988) Proc. Biol. Soc. Wash. 101: 192-208.

229
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchiplicatus cupreus PETTIBONE, 1985

to dorsal sides of notopodia and dorsal cirrophores. Parapodia


Size: 50 mm in length, 15 mm in width.
biramous. Notopodia shorter than neuropodia. Notochaetae
Morphology: Segments 31-35. Prostomium truncate. No eyes, numerous forming radiating bundles short to long. Notochaetae
no lateral antenna. Elytra 12 pairs on segments 2, 4, 5, 7, 9, 11, much stouter than neurochaetae, acicular, smooth with double
13, 15, 17, 19, 21 and 23. Up to 12 posterior segments without rows of spines near distal bare tips. Neurochaetae very numer-
elytra. Elytra large covering dorsum excepted for posterior seg- ous, slender, forming fan-shaped bundles. Subacicular neu-
ments. Elytra stiff, smooth, opaque, with « veins » Posterior rochaetae more slender, with shorter spines and slightly hooked
border of elytra with variable number of small rounded tuber- bare tips.
cules. Elytra surface with minute sensory papillae. Branchiae
Biology: From Riftia and Calyptogena washings.
begin on segment 3 of unique type, formed of flattened elon-
gated sacs, deeply folded and convoluted attached by broad Distribution: East Pacific Rise: 9°N to 21°N; Guaymas Basin.
bases to lateral sides of elytrophores and dorsal tubercules and

1: Dorsal view; by P. Briand  Ifremer. 2: Prostomium (SEM)  Ifremer.

3: Elytron (SEM) 4: Close up of the surface of 5: Folded branchiae (SEM)


 Ifremer. the elytron (SEM)  Ifremer.  Ifremer.

Reference:
PETTIBONE M.H. (1985) Proc. Biol. Soc. Wash. 98: 150-157.

D. DESBRUYÈRES Denisia 18 (2006): 230


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Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchipolynoe pettibonae MIURA & HASHIMOTO, 1991

Size: Up to 43 mm in length. Up to 19 mm in width. Biology: Living in the mantle cavity of the deep-sea vent mus-
sels (Bathymodiolus platyfrons, B. japonicus, B. brevior, B. elonga-
Morphology: Body flattened, with 21 segments (first
tus).
achaetous). Elytra 10 pairs on segments 2, 4, 5, 7, 9, 11, 13, 15,
17 and 19. Branchiae well developed, arborescent on segment Distribution: Izu Ogasawara Arc, Okinawa Trough, North Fiji
3-21; separated into upper larger and lower smaller groups. No- and Lau Back Arc Basins.
topodia very small, digitiform, with a few notochaetae. Neu-
ropodia large, long, with slightly bilobed acicular lobe and
rounded postchaetal lobe. Ventral segmental papillae long; two
pairs on segments 11 and 12.

1: Habitus © Greg Rouse (TUIM06).

2: Anterior part, ventral view from 3: Parapodium, 11th segment, 4: 11th and 12th segments,
a specimen of Hine Hina, Lau right side, anterior view ventral view 
Bassin  Briand/Ifremer.  Briand/Ifremer. Briand/Ifremer.

References:
COSEL R. & B. MÉTIVIER (1994) Veliger 37: 374-392.
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.
HASHIMOTO J., OHTA S., FUJIKURA K. & T. MIURA (1995) Deep-Sea Res. I 42: 577-598.
MIURA T. & J. HASHIMOTO (1991) Proc. Biol. Soc. Wash. 104: 166-174.

T. MIURA Denisia 18 (2006): 231


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Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchipolynoe seepensis PETTIBONE, 1986

Size: Up to 31 mm in length and 13 mm in width. Remark: According to CHEVALDONNÉ et al. (1998) B. seepensis
from the Gulf of Mexico and B. seepensis from Mid-Atlantic
Morphology: Body short, spindle-shaped, tapered anteriorly
Ridge appear to have recently diverged and must be considered
and posteriorly, flattened ventrally and arched dorsally. 21 seg-
as two isolated phylogenetic species, although no morphologi-
ments with 10 pairs of elytra attached on segments 2, 4, 5, 7, 9,
cal differences have been found on large series of specimens.
11, 13, 15, 17 and 19. Elytra moderatly large, oval, covering lat-
eral thirds of body on larger specimens and nearly covering Biology: Commensal within the mantle cavity of mussels. Two
body of juveniles. Branchiae well developped, arborescent. individuals (male and female) are frequently observed within a
Prostomium oval, bilobed, with rounded anterior lobes lacking single mussel. Sexual dimorphism with females larger than
frontal filaments. Median antenna lacking distinct cer- males; females having two pairs of ventral papillae on segments
atophore. Palp rather short, thick, smooth and tapered. Thick 11 and 12. Sex ratio 0.5-0.7: 1. Females contain mature sperm
muscular pharynx, five pairs of small delicate oval papillae and there is evidence of internal fertilization. Inferred mode of
around opening and two pairs of small jaws lacking denticulate development lecitotrophic or direct. Asynchronous gametoge-
base. Lower neurochaetae slender and having hooked tips with nesis, rapid oogenesis.
some longer lateral denticles.
Distribution: Hypersaline seeps at the base of Florida Escarp-
ment, 26°02’24’’N and 84°54’48’’W; Mid-Atlantic Ridge:
Lucky Strike.

1: Proboscis, frontal view (SEM) © Ifremer. 2: Tip of upper neurochaetae


(SEM) © Ifremer.

D. DESBRUYÈRES Denisia 18 (2006): 232–233


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

3: Habitus, dorsal view; by A. Fifis © Ifremer.

4: Habitus, ventral view; by A. Fifis © Ifremer.

References:
CHEVALDONNÉ P., JOLLIVET D., FELDMAN R. A., DESBRUYÈRES, d. & LUTZ R. A. & R. C. VRIEJENHOEK (1998) Cah. Biol. Mar. 39: 347-350.
DESBRUYÈRES D., ALAYSE A.M., ANTOINE E., BARBIER G., BARRIGA F., BISCOITO M., BRIAND P., BRULPORT J.P., COMTET T., CORNEC L., CRASSOUS P., DANDO P., FABRI M.C.,
FELBECK H., LALLIER F., FIALA-MÈDIONI A., CONÇALVES J., MÈNARD F., KERDONCUFF J., PATCHING J., SALDANHA L. & P.M. SARRADIN (1994) InterRidge News 3(2):
18-19.
HOURDEZ S. & C. JOUIN-TOULMOND (1998) Zoomorphology 118: 225-233.
HOURDEZ S., LALLIER F.H., GREEN B.N. & A. TOULMOND (1999) Proteins 34: 427-434.
JOLLIVET D., COMTET T., CHEVALDONNÉ P., HOURDEZ S. DESBRUYÈRES D. & D. DIXON (1998) Cah. Biol. Mar. 39: 359-362.
PETTIBONE M.H. (1986) Proc. Biol. Soc. Wash. 99(3): 444-451.
VAN DOVER C.L., TRASK J., GROSS J. & A. KNOWLTON (1999) Mar. Ecol. Progr. Ser. 181: 201-214.

233
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Branchipolynoe symmytilida PETTIBONE, 1984

Size: Up to 50 mm. Biology: All worms were collected from mussels and were
found occupying the anterior and ventral parts of the mantle
Morphology: Scale worm dwelling commensal of Bathymodiolus
cavity. A study of the digestive contents led to hypothesize a
thermophilus; body flattened, segments 21 first achaetous. Elytra
nutrition based on mussel gills and pseudofaeces.
and elytrophores 10 paires on segments 2, 4, 5, 7, 9, 11, 13, 15,
17 and 19. Branchiae well developped, arborescent on all para- Distribution: Galapagos Spreading Center, East Pacific Rise
podia from segment 2. Notopodia small, with few short acicu- (not collected at 21°N and Guaymas Basin).
lar notochaetae. Neuropodia short truncate, without projecting
acicular lobes; with very numerous neurochaetae including up-
per stout, slightly hooked ones and very slender ones. Adults
with very small elytra. 0/2 pairs of ventral papillae on segments
11 and 12.

2: Gill in vivo; by P. Briand  Ifremer.

1: Habitus; by V. Martin  Ifremer.

3: Ventral view showing the two pairs of


papillae; by P. Briand  Ifremer.

References:
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.
HOURDEZ S., LALLIER F., MARTIN-JEZEQUEL V. WEBER R.E. & A. TOULMOND (1999) Proteins 34: 427-434.
PETTIBONE M.H. (1984) Proc. Biol. Soc. Wash. 97: 226-239.

D. DESBRUYÈRES Denisia 18 (2006): 234


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Annelida, Polychaeta, Phyllodocida, Polynoidae

Iphionella risensis PETTIBONE, 1986

are straw-colored and bipinnate feathered, with a slender axis


Size: Up to 11 mm in length, width 6 mm including chaetae.
and lateral spines. The upper group of neurochaetae are similar
Color: Reddish. to notochaetae. The rest of neurochaetae are stouter, with
close-set spinous rows on the basal enlarged part.
Morphology: Body ovate greatly flattened. Thirteen pairs of
elytra, 28 segments. The elytra are reniform, their surface cov- Biology: Uncommon, in mussels and clams washings. More
ered by polygonal areas enclosing small areolae. Prostomium common among stalked cirripeds.
bilobed forming to separate rounded lobes, with anterolateral
Distribution: East Pacific Rise: 21°N, 9°N. A posteriorly in-
bulbous extensions fused to the facial tubercle. Antennae and
complete, unique specimen was sampled at Mid-Atlantic
eyes lacking. Long cylindrical tentaculophores emerge lateral
Ridge: Lucky Strike and could correspond to I. risensis or a
to the prostomium and palps each with a stout aciculum. Short
closely related species.
papillate dorsal and ventral tentacular cirri. Small oval medial
nodules are found on segments 4 and 5, variable in number and
position among studied specimens. Very numerous notochaetae

2: Surface of elytron in vivo  Ifremer. 3: Prostomium, dorsal view (SEM)  Ifremer.

1: Habitus in vivo  Ifremer. 4: Surface of elytron (SEM)  Ifremer. 5: Notochaetae (SEM)  Ifremer.

References:
PETTIBONE M.H. (1986) Smithson. Contr. Zool. 428: 1-43.
VAN DOVER C.L. (2002) Mar. Ecol. Progr. Ser. 230: 137-158.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 235


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Annelida, Polychaeta, Phyllodocida, Polynoidae

Lepidonotopodium atalantae DESBRUYÈRES & HOURDEZ, 2000

pairs of unequal bulbous papillae encircle the opening of the


Size: Up to 9 mm in length for 23 segments.
extended pharynx. The three dorsal papillae are pear-shaped,
Color: Red to pinkish, notopodial chaetae are straw colored, longer than the other ones. The two pairs of hooked jaws are
neuropodial chaetae are light brown. minutely serrated with numerous teeth. The notopodium is
subconical with acicular lobes and is enclosed anterodorsally by
Morphology: The body is short, suboval in outline, flattened
flaring bracts. The neuropodium is deeply cleft in the upper and
dorsoventrally, slightly tapered and rounded anteriorly and pos-
lower part. The notochaetae are numerous forming thick radi-
teriorly. The 11 pairs of elytra are located on segments 2, 4, 5,
ating bundles. The notochaetae have two rows of spines and
7, 9, 11, 13, 15, 17, 19 and 21. They cover the dorsum. The ely-
bare tips. The neurochaetae are numerous; they have two rows
tra are subreniform, overlapping with 6-12 macrotubercules
of numerous spines and bare tips. On one third of the speci-
raised on the posterior border. The elytral surface appears
mens (males), there are four ventral pairs of elongated papillae
smooth but is covered with numerous globular or clavate mi-
on segments 11-14, extending to the tip of neurochaetae.
cropapillae. The surface of both elytrophores and dorsal tuber-
There is a pair of very long ventral anal cirri.
cles have bands or tufts of cilia. The prostomium is bilobed, the
anterior lobes subtriangular with a frontal filament, lateral an- Distribution: East Pacific Rise: 13°N to 17°S.
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

5: Elytron from the middle part of the


body (SEM) © Ifremer.
3: Anterior part in dorsal view (SEM) © Ifremer.

1: Habitus in vivo © Ifremer.

2: Ventral view in vivo of the


segments 11-14 showing the
sexual elongated papillae 4: Frontal view of the distal part of the proboscis 6: Distal part of a neuropodial chaeta
© Ifremer. (SEM) © Ifremer. (SEM) © Ifremer.

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

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 236


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta, Phyllodocida, Polynoidae

Lepidonotopodium fimbriatum PETTIBONE, 1983

microtubercules. The prostomium is bilobed, the anterior lobes


Size: Maximum observed 35 mm in length, 18 mm in width.
subtriangular, each with a small frontal filament. The biramous
Color: Living specimens pinkish or rustish to whitish in color, parapodia have a shorter notopodia located on the anterodor-
neurochaetae brown to black. sal sides of the longer neuropodia. The distal margins of the no-
topodial acicular lobes, notopodial bracts, and neuropodial
Morphology: Body stout, short, rectangular in outline, flat-
lobes are fimbriated with slender papillae.
tened ventrally, strongly arched dorsally, and slightly tapered
and rounded anteriorly and posteriorly. The elytra are thick; Biology: Mainly observed abundant on smoker walls. Likely a
leathery, imbricated, covering the dorsum. There are 11 paires carnivorous species (amphipods observed crushed by the jaws).
of elytra attached on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19
Distribution: East Pacific Rise: 21°N to 9°N.
and 21. Except for the first and the last pairs, each elytron has
two raised smooth macrotubercules on the posterior one-third.
The elytra appears smooth but is covered with numerous round

2: Prostomium, dorsal view  Ifremer.

1: Habitus; by V. Martin  Ifremer.

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


Cruise Phare  Ifremer.

Reference:
PETTIBONE M.H. (1983) Proc. Biol. Soc. Wash. 96(3): 392-399.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 237


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Annelida, Polychaeta, Phyllodocida, Polynoidae

Lepidonotopodium jouinae DESBRUYÈRES & HOURDEZ, 2000

numerous teeth. The biramous parapodia have a shorter no-


Size: Maximum observed 14 mm in length and width 7 mm.
topodia located on the anterodorsal sides of the longer neu-
Color: Light brown after preservation. ropodia. The notopodium is enclosed by a flaring bract. The
neuropodium is deeply cleft on the upper and lower parts. The
Morphology: Body short, suboval in outline, flattened
notochaetae are stouter than neurochaetae. They have two
dorsoventrally, slightly tapered and rounded anteriorly and pos-
rows of alternating teeth. The dorsal neurochaetae have one
teriorly. The elytra are opaque, oval to subreniform, imbricate,
row of spines and straight tips. The ventral ones have two rows
covering the dorsum. There are 11 pairs of elytra attached on
of numerous spines. 0/5 pairs of elongated papillae on segments
segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19 and 21. The elytra ap-
11-15.
pears smooth to the naked eye. Under SEM, the elytral surface
in the non-overlapping region appears covered with numerous Biology: Mainly observed in washings of mussels. Likely a car-
globular or clavate micropapillae. The prostomium is bilobed, nivorous/omnivorous species.
the anterior lobes subtriangular, each with a small frontal fila-
Distribution: Mid-Atlantic Ridge: Azores Triple Junction,
ment. Six pairs of unequal pear-shaped papillae and two medi-
Menez Gwen, and Lucky Strike.
an papillae encircle the opening of the extended pharynx. The
two pairs of dorsal and ventral jaws are minutely serrated with

3: Prostomium, fronto-dorsal view


(SEM)  Ifremer.

6: Anterior elytron (SEM)  Ifremer.

4: Proboscis everted showing four


1: Habitus (preserved specimen) pairs of jaws with numerous teeth 7: Clavate micropapilla at the surface of
 Ifremer. (SEM) © Ifremer. elytron (SEM)  Ifremer.

2: Segments 11-15, ventral view


© Ifremer. 5: Upper neurochaetae (SEM)  Ifremer. 8: Notochaetae (SEM)  Ifremer.

Reference:
DESBRUYÈRES D. & S. HOURDEZ (2000) Cah. Biol. Mar. 41: 399-405.

D. DESBRUYÈRES Denisia 18 (2006): 238


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Annelida, Polychaeta, Phyllodocida, Polynoidae

Lepidonotopodium piscesae PETTIBONE, 1988

on posterior lower side and deep notch on posterior upper side.


Size: Maximum observed 29 mm in length, 13 mm in width.
Notochaetae with two rows of spines and blunt tapered and
Color: Living specimens brownish to tan in color, elytra cov- “hairy“ tips. Very numerous neurochaetae with slightly hooked
ered with bacterial filaments giving them a white color. tips and fine spiny rows on cutting edge. On some specimens,
neuropodia of segment 13 differing by the presence of only 1-2
Morphology: Body sturdy, elongated oval, rounded anteriorly
stout, reddish sabre like chaetae. 0/5 elongate papillae on seg-
and posteriorly, flattened ventrally, arched dorsally. The elytra
ments 11-15.
are large, overlapping, covering the dorsum, thick, stiff, opaque.
There are 11 pairs of elytra attached on segments 2, 4, 5, 7, 9, Biology: Widely dispersed species, but low in abundance. Ob-
11, 13, 15, 17, 19 and 21. Elytral surface thickly covered with served at the periphery of vents, crawling on the rocks, active
opaque rounded to conical microtubercles, especially thick on sulphide and vestimentiferan tubes, grazing on the bacterial
somewhat transverse areas in the middle of elytra and along mats and protozoans. Trophic generalist or/and carnivorous.
borders with scattered globular and filiform micropapillae. The
Distribution: Explorer Ridge; Juan de Fuca Ridge: Endeavour
prostomium is bilobed, the anterior lobes prominent, each with
segment, Axial Seamount and Southern Juan de Fuca Ridge;
a long frontal filament. The biramous parapodia have a shorter
Gorda Ridge.
notopodia located on the anterodorsal sides of the longer neu-
ropodia. Neuropodium diagonally truncate, with shallow notch

2: Prostomium, dorsal (SEM)  Ifremer.

1: Habitus in vivo  Ifremer. 3: Elytron (SEM)  Ifremer.

D. DESBRUYÈRES Denisia 18 (2006): 239–240


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

5: Elongated ventral papillae on segments 11-15 (SEM)


 Ifremer.

4: Proboscis and jaws (SEM)  Ifremer.

6: Sabre-like supra acicular neurochaetae on segment 13 7: Elytron surface (SEM)  Ifremer.


(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.

240
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Lepidonotopodium riftense PETTIBONE, 1984

have a variable number of oval projections near the posterior


Size: Maximum observed 13 mm in length and 7 mm in width.
border. The prostomium is bilobed, the anterior lobes subtrian-
Color: No color in preservative. gular, each with a small frontal filament. The biramous parapo-
dia have a shorter notopodia located on the anterodorsal sides
Morphology: Body short, suboval in outline, flattened
of the longer neuropodia. The neuropodium is diagonally trun-
dorsoventrally, slightly tapered and rounded anteriorly and pos-
cated and deeply cleft on the upper part. 0/2 pairs of elongated
teriorly. The elytra are oval to subreniform, imbricate, covering
papillae on segments 11-12.
the dorsum. There are 11 pairs of elytra attached on segments
2, 4, 5, 7, 9, 11, 13, 15, 17, 19 and 21. The elytra appear smooth Biology: Mainly observed in washings of mussels, clams and
with branching “veins“ emanating from the place of attach- tube worms. Likely a carnivorous/omnivorous species.
ment to the elytrophores. Near the posterior and lateral borders
Distribution: East Pacific Rise: 21°N to 9°N and Galapagos
of the elytra, scattered micropapillae with short tapered or
Spreading Center.
cylindrical bases. On some specimens from the EPR, the elytra

3: Opening of the pharynx


encircled by nine pairs of
bulbous papillae © Ifremer.
6: Prostomium, dorsal view (SEM) © Ifremer.

1: Habitus (preserved specimen)


© Ifremer.

4: Oval projections of the


posterior border of one elytron
(SEM) © Ifremer.

2: Ventral view (preserved speci-


men) showing the two pairs of
elongated papillae on segments
11-12 attached basally on the 5: Globular micropapillae
ventroposterior sides of the at the surface of elytron (SEM) 7: Two pairs of dorsal and ventral jaws minutely
neuropodia. © Briand/Ifremer. © Ifremer. serrated 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.

D. DESBRUYÈRES Denisia 18 (2006): 241


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta, Phyllodocida, Polynoidae

Lepidonotopodium williamsae PETTIBONE, 1984

topodia located on the anterodorsal sides of longer neuropodia.


Size: 26 mm in length, 19 mm in width. Largest up to 36 mm
The notopodia are subconical with projecting acicular lobes
in length.
hidden by the numerous notochaetae and enclosed dorsally by
Color: Brownish to tan. well developed large flaring bracts. The distal margins of the
notopodial acicular lobes, notopodial bracts and neuropodial
Morphology: Body short, stout, rectangular in outline, 24-26
lobes are fimbriate with slender papillae. 0/2 elongated papillae
segments. 11 pairs of elytra attached eccentrically. The elytra
ventral on segments 11-12.
are large, imbricate, covering the dorsum. They are thick, stiff
opaque; their dorsal surface is nearly covered with rounded to Biology: Ubiquitous at vents.
conical microtubercules (inner anterior part) and rounded ele-
Distribution: Galapagos Spreading Center; East Pacific Rise:
vations at the distal posterior part. The prostomium is bilobed,
21°N to 9°N.
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-

2: Habitus, in vivo  Ifremer. 3: Proboscis, frontal view (SEM)  Ifremer.

4: Elytron, posterior border


1: Habitus; by V. Martin  Ifremer. (SEM)  Ifremer 5: Prostomium, dorsal view (SEM)  Ifremer.

References:
DESBRUYÈRES D. & S. HOURDEZ (2000) Cah. Biol. Mar. 41: 47-54.
PETTIBONE M.H. (1984) Proc. Biol. Soc. Wash. 97: 849-863.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 242


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Annelida, Polychaeta, Phyllodocidae, Polynoidae

Levensteiniella iris HOURDEZ & DESBRUYÈRES, 2003

curved and appears as a parrot beak. The parapodia are bira-


Size: Maximum observed 21 mm in length for 25 segments.
mous. Notopodia are shorter in length to the neuropodia and
Color: Living specimens transluscent or greenish with gold col- located on their anterior-dorsal side. Notopodia are cylindrical
ored chaetae. Elytra and chaetae often coated with mineral de- in shape, with the bundle of notochaetae radiating on the dor-
posits. sal side, close to the tip. The acicular lobe forms a pointed,
scoop-like ligule located underneath the notochaetae. The
Morphology: Body short, tapering anteriorly and posteriorly.
neuropodia are not cleft in the upper and lower part, but the
There are 11 pairs of elytra (they are deciduous). Elytra overlap
acicular lobe forms a pointed, scoop-like ligule on the dorsal-
anteriorly and posteriorly, and cover the mid-dorsal line. Short
anterior side. Notochaetae are stouter than the neurochaetae.
dorsal cirri on the non-elytra bearing segments. Elytra are
Notochaetae are short, curved on one side, with rows of teeth
translucent, smooth, oval, and bear posterior, raised, irregular-
appearing as scales on the curved side. They taper to blunt tips.
ly-spaced macrotubercles. The elytral surface appears smooth or
The neurochaetae are numerous, forming a fan-shaped bundle
slightly covered of small scattered bumps. The cirri are as long
and bear two lateral rows of spines on one side. They are
as the parapodia, tapering to tips. The prostomium is bilobed,
unidentate and and slightly hooked. Ventral cirri are short and
with cephalic peaks and frontal filaments as long as 1/3 of the
tapered, located at the base of the neuropodia. The pygidium is
prostomium length. Lateral antennae are absent and the medi-
small, rounded, bearing a pair of anal cirri. All the ventral
an antenna is approximately as long as the prostomium, tapers
papillae are inconspicuous, except of segment 11 where they
close the tip and bears a terminal filament. The median anten-
are very developed in some of the specimens.
na is inserted in the anterior notch of the prostomium and has
a short, spherical ceratophore. Palps are short and thick, ap- Biology: Among Bathymodiolus azoricus beds.
proximately 1.5 times the length of the prostomium, and bear
Distribution: Mid-Atlantic Ridge: Rainbow, Lucky Strike (un-
terminal filaments. When dissected, the pharynx opening re-
common species).
vealed seven pairs of papillae, all similar in size. There are two
pairs of jaws which do not bear any teeth but the tip is slightly

1: Habitus, dorsal view  Ifremer. 2: Prostomium, dorsal view (SEM)  Ifremer.

3: Posterior part of elytron 4: Acicular lobe of notochaetae 5: Ventral pair papillae


(SEM)  Ifremer. (SEM)  Ifremer. 11th segment  Ifremer.

Reference:
HOURDEZ S. & D. DESBRUYÈRES (2003) Cah. Biol. Mar. 44: 13-21.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 243


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Annelida, Polychaeta, Phyllodocida, Polynoidae

Levensteiniella plicata HOURDEZ & DESBRUYÈRES, 2000

has two pairs of straight jaws with up to 20 teeth. Both rami of


Size: Maximum size observed 57 mm in length, width 21 mm
the parapodia well developed; the notopodia shorter than the
including chaetae.
neuropodia are rounded and the acicula project from acicular
Color: Pinkish in vivo, light brown when preserved. Chaetae lobes. The notochaetae are numerous, stouter than neu-
are straw colored. rochaetae. They are straight, flattened, tapering to blunt tips
and with scales on one side. The neurochaetae are slender, nu-
Morphology: Body long suboval, flattened dorso-ventrally. 11
merous, forming a fan-shaped bundle. They have two longitu-
pairs of elytra. The elytra are smooth, thick and with a promi-
dinal rows of spines on one side and their tip is bare and
nent longitudinal fold. They leave uncovered the middle part
straight. 0/2 elongated papillae on 11 and 12.
of the body and the posterior end. The tapering tip of the dor-
sal cirrus does not extend beyong the neurochaetae. The pros- Biology: Living among mussel beds of Bathymodiolus ther-
tomium is trapezoid, deeply bilobed with frontal terminal fila- mophilus.
ments on the cephalic peaks. The ceratophore of the median
Distribution: East Pacific Rise: 9°N.
antennae is short, cylindrical, and inserted in the anterior
notch. The style is shorter than the palps, which are stout,
shorts and with a terminal articulated filament. The proboscis

2: Posterior part  Ifremer.

1: Habitus  Ifremer. 3: Parapodia, dorsal view  Ifremer.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 244–245


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

4: Right parapodium, anterodorsal view (SEM)  Ifremer.

6: Elongated ventral
5: Prostomium, frontal view; median antenna papillae 11 and 12
removed (SEM)  Ifremer. segments  Ifremer.

7: Notochaetae (SEM)  Ifremer. 8: Neurochaetae (SEM)  Ifremer.

References:
HOURDEZ S. & D. DESBRUYÈRES (2000) Cah. Biol. Mar. 41: 97-102.
VAN DOVER C.L. (2002) Mar. Ecol. Progr. Ser. 230: 137-158.

245
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Thermiphione fijiensis MIURA, 1994

14, thereafter in single row, absent posterior to segment 20.


Size: Up to 20 mm in length and 10 mm in width.
Elytrophores bulbous, transversaly elongated; places of attach-
Morphology: Body flattened, with 30 or 31 segments. Elytra 14 ment with latero-posterior extensions. Dorsal tubercles on cir-
pairs on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26 and rigerous segments bulbous, prominent, transversally elongated,
27; elytra on segment 2 oval, with fringes of short papillae cov- continuous with enlarged cirrophores of dorsal cirri. Dorsal and
ered with filamentous bacteria; other elytra elongated subreni- ventral cirri with short clavate papillae. Branchiae absent. No-
form, lateral borders with fringes of short papillae; elytral sur- topodia subconical, with projecting acicular lobes hidden by
face covered with hexagonal or polygonal areas with secondary numerous notochaetae. Neuropodia large truncate. Hooked
areolae. Prostomium partially fused to tentacular segment, and neurochaetae occurring on segment 3.
withdrawn in anterior segments. Prostomium bilobed forming
Biology: Free living scale-worm. Epibiotic filamentous bacteria
separate rounded lobes with anterolateral extensions. Tentacu-
on parapodia.
lar segment with long cylindrical tentaculophores lateral to
prostomium; each with a single aciculum. Nodular papillae aris- Distribution: North Fiji and Lau Back-Arc Basins.
ing from dorsum of anterior segments; more than 20 papillae on
anterior segments; arranged in two or more rows before segment

2: Anterior end, dorsal view


elytra removed;
from MIURA (1994).

3: Anterior
(segment 2) and
median (segment
5) elytra with de-
tails of the hexag-
onal areas and
marginal papillae;
from MIURA (1994).

1: Dorsal view; by P. Briand © Ifremer.

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

T. MIURA Denisia 18 (2006): 246


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Annelida, Polychaeta, Phyllodocida, Polynoidae

Thermiphione tufari HARTMANN-SCHRÖDER, 1992

found on segments 4 (1), 5 (2) and two following segments (2)


Size: Up to 30 mm in length, width 7.5 mm including chaetae.
but variable in number according to specimens. Segment 3 not
Color: Reddish. visible dorsally. Elytrophores bulbous, transversally elongated.
Dorsal tubercles on cirrigerous segments transversally elongat-
Morphology: Body ovate greatly flattened. Fourteen pairs of
ed and striated. Dorsal and ventral cirri with short clavate
elytra (…23, 26, 27), 29-32 segments. The elytra are reniform,
papillae. Notochaetae forming radiating bundles of dense tufts,
their surface are covered by polygonal areas with secondary are-
shorter than neurochaetae. Hooked neurochaetae first present
olae; elytrae ciliated at the outer and posterior borders. Pros-
from chaetiger 4.
tomium bilobed forming two separate rounded lobes, with an-
terolateral bulbous extensions fused to the facial tubercle. An- Biology: Rare, found at the periphery of active vents, with
tennae and eyes lacking. Long cylindrical tentaculophores stalked barnacles.
emerge lateral to the prostomium and palps each with an aci-
Distribution: East Pacific Rise: 7°S to 21°S.
culum as well as one or two chaetae. Short papillae on dorsal
and ventral tentacular cirri. Small oval medial nodules are

1: Anterior part, dorsal view © Ifremer. 2: Proboscis, frontal view (preserved specimen) © Ifremer.

D. DESBRUYÈRES & S. HOURDEZ Denisia 18 (2006): 247–248


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

3: Median elytron © Ifremer. 4: Elytra withciliated border (SEM) © Ifremer.

5: Polygonal area with a central areola (SEM) © Ifremer. 6: Medial parapodia, dorsal view (SEM) © Ifremer.

7: Notochaetae (SEM) © Ifremer. 8: Ventral cirrus (SEM) © Ifremer.

Reference:
HARTMANN-SCHRÖDER G. (1992) Helgol. Meeresunters. 46: 389-403.

248
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Thermopolynoe branchiata MIURA, 1994

seven pairs of bulbous papillae. Dorsal and ventral pairs of jaws


Size: Up to 53 mm in length. Up to 20 mm in width.
fused medially each with up to 15 teeth on basal sides. No-
Morphology: Body flattened, with 27 segments (first topodia subconical with projecting acicular lobes and large flar-
achaetous). Elytra 11 pairs on segments 2, 4, 5, 7, 9, 11, 13, 15, ing bracts. Neuropodia diagonally truncate, deeply notched on
17, 19 and 21, large, covering dorsum, stiff, rough with numer- upper part; distal margins fimbriated with slender papillae cov-
ous brownish papillae. Branchiae well developed, arborescent ered with numerous filamentous bacteria. Ventral segmental
on segment 3-26; separated into two groups on anterior and papillae long; three pairs present on segments 12, 13 and 14 and
posterior sides of notopodia on segment 3 and on some less de- lacking in half of the specimens.
veloped posterior segments; forming single large branchial areas
Biology: Free living. Found in washings of Ifremeria and mus-
encircling central parts of notopodia on other fully developed
sels. Epibiotic bacteria on parapodia.
parapodia. Prostomium bilobed. Anterior lobes prominent,
cylindrical, with small frontal filaments. Mouth opening situat- Distribution: North Fiji, Lau Basin, and Manus Back-Arc
ed between segments 1 and 2. Muscular pharynx encircled by Basins.

4: Prostomium and buccal region, frontal view (SEM)


1: Habitus in vivo; by courtesy of C. Rouse (cruise TUIM06). © 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


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

7: Middle notochaetae (SEM) © Ifremer.

10: Jaw (dissection) (SEM) © Ifremer.

8: Flaring bracts first chaetigers, frontal view (SEM) © Ifremer.

11: Median elytrophorous chaetiger (SEM) © Ifremer.

9: Arborescent branchiae forming large branchial areas (SEM)


© Ifremer. 12: Median cirriphorous chaetiger (SEM) © Ifremer.

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

250
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Annelida, Polychaeta, Phyllodocida, Polynoidae

Vampiropolynoe embleyi MARCUS & HOURDEZ, 2002

Size: Up to 52 mm. Biology: All worms were collected from bare basalt on vent pe-
ripheries, after the 1998 eruption of Axial Volcano. The species
Morphology: Scale worm found on bacterial mats; body flat-
is thought to feed on bacterial mats and may be a pioneer
tened, segments 43-45 first achaetous. Elytra (smooth) and
species.
elytrophores 10 paires on segments 2, 4, 5, 7, 9, 11, 13, 15, 17
and 19. Dorsal tubercles developed as gills on cirrigerous seg- Distribution: Juan de Fuca Ridge: Axial Volcano.
ments starting on segment 6. Prostomium bilobed, strong sharp
acicular lobes on segment 1. Notochaetae stouter than neu-
rochaetae. No jaws, keratinized teeth inside the pharynx.
Mouth opening with papillae. No elongated ventral papillae.

1: Paratype, dorsal
view of the whole
specimen;
from MARCUS &
HOURDEZ (2002).

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.

2: Keratinized teeth at the junction between the mouth opening


and the pharynx (SEM); by courtesy of J. Marcus and S. Hourdez.

5: Dorsale view of the anterior part of the worm, showing


3: Dorsal view of the segments 11-14 (SEM); by courtesy of J. Mar- the strong aciculae protrudring from the first segment
cus and S. Hourdez. (SEM); by courtesy of J. Marcus & S. Hourdez.

Reference:
MARCUS J. & S. HOURDEZ (2002) Proc. Biol. Soc. Wash. 111(2): 341-349.

S. HOURDEZ Denisia 18 (2006): 251


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Annelida, Polychaeta, Phyllodocida, Syllidae

Sphaerosyllis ridgiensis BLAKE & HILBIG, 1990

chaetiger 2; ventral cirri long, cirriform. Chaetae including sin-


Size: Small species, holotype 3.1 mm long for 23 chaetigerous
gle, long dorsalmost simple chaetae with bidentate tips and 8-
segments, 1 mm wide.
10 compound falcigers with bidentate tips and conspicuous ser-
Morphology: Prostomium weakly notched on anterior margin, rations on blade. Pharynx red in color, occupying tentacular
about as long as wide, clearly fused with tentacular segment; segment, and bearing single middorsal tooth sometimes emerg-
medial antenna inserted at point of folded merger of tentacular ing from oral opening; proventriculus occupying chaetigers 1-4,
segment with prostomium: lateral antennae inserted nearly in tan colored, with about 20 rows of muscle cells.
line with medial antenna; antennae glandular, with bulbous
Biology: Unknown. The syllids are known to be suctorial-feed-
bases and narrow tapering tips. Eyes lacking. Tentacular cirri
ers.
present laterally on tentacular segment similar to antennae;
parapodia similar throughout body, conical with single imbed- Distribution: Explorer Ridge, Juan de Fuca Ridge.
ded aciculum lacking papillae; dorsal cirri glandular, with bul-
bous bases and narrow tapering tips; without dorsal cirri on

2: Anterior part, ventral view 3: Median parapodium, dorsal view


(SEM) © Ifremer. (SEM) © Ifremer.

4: Proventricle (SEM) 5: Compound falcigers with bidentate tips (SEM)


1: Habitus, dorsal view; by courtesy of J. Blake. © Ifremer. © Ifremer.

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

J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 252


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Annelida, Polychaeta, Sabellida, Serpulidae

Laminatubus alvini TEN HOVE & ZIBROWIUS, 1986

Branchial crown about 1/2 of the animal. Branchial radioles in


Size: Tube, until 100 mm long, 5.3 mm wide.
a short spirale. Collar with capilaries and special chaetae, five
Morphology: The tube is thick-walled, smooth, subtriangular pairs of subsequent bundles. Abdomen with up to 109 seg-
in cross-section. It has a wide flattened area of attachment; the ments.
lateral parts are solids, not with the alveolar structure as in. A
Biology: At the border of active vent fields. Filter feeding or-
crest-like, sometimes undulating medial keel is the only orna-
ganisms (mixotrophic?).
mentation present. The wall of the tube comprises two distinct
layers. The operculum is radially or slightly bilaterally symmet- Distribution: Galapagos Spreading Center; East Pacific Rise:
rical. It consist of a bulbous proximal part, which may be some- 21°N to 23°S.
what conical, and a rounded to inverted saucer-like distal cap.

3: Tubes of specimens from East Pacific Rise


2: Habitus in vivo; 13°N  Ifremer.
by P. Briand  Ifremer.

1: Habitus opercu-
lum missing;
by V. Martin 4: Geniculated chaeta 5: Thoracic uncini 6: Population in situ from East Pacific Rise
 Ifremer. from anterior abdomen (SEM)  Ifremer. 13°N; cruise Phare  Ifremer.
(SEM)  Ifremer.
7: Two special collar
chaetae and one
capillary chaeta below;
from TENHOVE & ZIBROVIUS
(1986).

References:
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Biol. Soc. Wash. Bull. 6: 103-116.
TEN HOVE H.A. & H. ZIBROWIUS (1986) Zool. Scripta 15(1): 21-31.

D. DESBRUYÈRES & M. SEGONZAC Denisia 18 (2006): 253


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Annelida, Polychaeta, Sabellida, Serpulidae

Protis hydrothermica TEN HOVE & ZIBROWIUS, 1986

not been observed. The wall of the tube is homogenous, and


Size: Tube, up to 70 mm long, 3.8 mm wide, the thickness of
does not consist of two distinct layers. The surface is lusterless.
the wall is ca 0.2 mm; length of the body 47 mm, width of the
Older tube parts are frequently encrusted by yellowish-brown
thorax up to 2 mm.
deposits. No operculum.
Morphology: The tube is semicircular in cross-section, adher-
Distribution: Galapagos Spreading Center; East Pacific Rise:
ing relatively flatly to the substrate. If a flattened area of at-
21°N, 13°N, 17°S; Pacific-Antarctic Ridge: 38°S.
tachment is present, it is very small. The tube is rather indis-
tinctive, somewhat rugged. Very careful illumination reveals
that, in addition to the faint medial ridge, there may be a pair
of shallow grooves or ridges laterally, making the cross-section
of the tube slightly trapezoidal. Free anterior tube parts have

2: Anterior part of tube; from TEN


HOVE & ZIBROWIUS (1986).

1 left to right: Special


collar chaeta, capillary 4: Specimen removed from tube; East Pacific Rise: 13°N;
collar chaeta and capil- by D. Desbruyères © Ifremer.
lary chaeta; from TEN
HOVE & ZIBROWIUS (1986).

3: Tube of specimen on
the capitulum of
stalked banacle Vul-
canolepas sp., from East
Pacific Rise: 17°S; cruise 5: Colony of tubes on an artificial substrate left several
Biospeedo; by P. Briand months on the vent site Parigo, East Pacific Rise: 13°N; cruise
© Ifremer. Hydronaut; by P. Briand © Ifremer.

Reference:
TEN HOVE H.A. & H. ZIBROWIUS (1986) Zool. Scr. 15(1): 21-3.

H. ZIBROWIUS & M. SEGONZAC Denisia 18 (2006): 254


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Annelida, Polychaeta, Sabellida, Siboglinidae

Alaysia spiralis SOUTHWARD, 1991 “small spiral tube-worm“

Size: Tube length max. 300 mm; diameter 0.7-1.0 mm. Biology: Tubes are fixed to rocks close to mussel beds in areas
of diffuse venting; temperature about 7°C. Endosymbiotic bac-
Color: White/grey, semi-transparent.
teria present, probably sulphur-oxidisers.
Morphology: Anterior part of the tube coiled like a corkscrew,
Distribution: Valu Fa Ridge in the Lau Back-Arc Basin.
rising from a sinuous basal region. Surface smooth, with narrow
flanges 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, forming
lamellae, composed of two types: one pair of sheath lamellae,
composed of filaments without pinnules, surrounds the inner
lamellae made of pinnulate filaments.

1 left: Anterior region of larger tube; right: Outer lamellae, inner filaments and obturaculum, dorsal view; from 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.
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


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Annelida, Polychaeta, Sabellida, Siboglinidae

Arcovestia ivanovi SOUTHWARD & GALKIN, 1997

rates prominent lateral ridges. The branchial filaments are


Size: The sinuous tubes are up to 200 mm long, the anterior
grouped in numerous paired lamellae, parallel to the obturacu-
part is up to 3 mm in diameter.
lum. The collar at the anterior end of the vestimental region is
Color: The semi transparent tube wall is light-tan to olive- long, covering the base of the branchial lamellae. The lateral
green (grey in preserved specimens). vestimental folds meet at the dorsal mid line, enclosing the dor-
sal vestimental space, and join posteriorly to form an entire
Morphology: The tube has an anterior funnel followed by a se-
ventral fold. The branchial plume including the obturaculum is
ries of collars (flanges) diminishing in size toward the posterior
slightly longer than the vestimental region. A pair of genital
end. The tube surface is slightly ridged, but its general appear-
grooves runs from the posterior end of the vestimental region to
ance is shiny and smooth. The plume is made of slender fila-
the base of the anterior collar in mature males.
ments that surround a slender obturaculum with a spoon-
shaped top. The obturaculum stalk has a sharp ventral ridge, Distribution: Manus and Lau Back-Arc Basins.
extending from the base to the apex and a dorsal groove sepa-

1: Co-dominant species at the DESMOS site (Manus Back-Arc Basin) are 2: Anterior end of the tube, obturaculum and
galatheids (Munidopsis lauensis), vestimentiferans (Arcovestia ivanovi) and plume; specimen from Lau Back-Arc Basin;
zoarcids (Pyrolycus manusanus); by courtesy of J. Hashimoto © JAMSTEC. cruise TUIM 06; by courtesy of Greg Rouse.

3: Habitus of two animals in their tube. Specimens from Lau Basin; cruise TUIM 06; by courtesy of Greg Rouse.

References:
SOUTHWARD E.C. & S.V. GALKIN (1997) J. Nat. Hist. 31: 43-45.
KOJIMA S., OHTA S., YAMAMOTO T., MIURA T., FUJIWARA Y., FUJIKURA K. & J. HASHIMOTO (2002) Mar. Biol. 141: 57-64.

D. DESBRUYÈRES Denisia 18 (2006): 256


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Annelida, Polychaeta, Sabellida, Siboglinidae

Lamellibrachia barhami WEBB, 1969

Size: Tube length max. ca. 1500 mm; anterior diameter 7-12 Biology: Sprawling tangles of tubes lie on muddy surface, with
mm. posterior ends burried in sediment or under rocks. Cold seep
animals live at ambient sea temperature; Middle Valley animals
Color: Grey/white, opaque.
at fringes of diffuse venting regions with sulphide seepage but
Morphology: Tube rigid, thick walled, sinuous, tapering to <1 no apparent temperature anomaly. Use internal symbiotic sul-
mm at posterior end; short external flanges anteriorly, eroded phide-oxidizing bacteria.
and indistinct over much of the tube. Top of obteraculum cup-
Distribution: Subduction zone cold seeps on the North Amer-
shaped, smooth interior surface, stalk elliptical in section. Ob-
ica continental margin and a sedimented hydrothermal region
turaculum short (max. 12 mm), about 1/3 length of vestimen-
at Middle Valley on the Juan de Fuca Ridge.
tal region. Branchial filaments are parallel to the obturaculum,
forming lamellae of two types: pale outer sheath lamellae (2-5
pairs) made of adherent filaments without pinnules and red in-
ner lamellae of pinnulate filaments.

2: Branchial region
in lateral view; by
E. Southward.

1: Population on the bottom at Middle valley on 3: Population on the bottom at Middle val-
4: Anterior region in vivo; by
the Juan de Fuca Ridge; by courtesy of V. Tunni- ley on the Juan de Fuca Ridge; by courtesy
courtesy of V. Tunnicliffe.
cliffe. of V. Tunnicliffe.

References:
JONES M.L. (1985) Bull. Biol. Soc. Wash. 6:117-158.
JUNIPER S.K., TUNNICLIFFE V. & E.C. SOUTHWARD (1992) Can. J. Zool. 70: 1792-1809.
SOUTHWARD E.C., TUNNICLIFFE V., BLACK M.B., DIXON D.R.& L.R.J. DIXON (1996) J. Geol. Soc., Spec. Publ. 118: 211-224.
SUESS E., CARSON B., RITGER S.D., MOORE J.C., JONES M.L., KULM L.D.& G.R. COCHRANE (1985) Bull. Biol. Soc. Wash. 6: 475-484.
WEBB M. (1969) Bull. Mar. Sci. 19: 18-47.

E.C. SOUTHWARD Denisia 18 (2006): 257


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Annelida, Polychaeta, Sabellida, Siboglinidae

Lamellibrachia columna SOUTHWARD, 1991

Size: Tube length max. 820 mm; anterior diameter 14-20 mm. Biology: The tubes are attached within rock crevices, and proj-
ect vertically, near sources of diffuse venting, but probably
Color: Tube white/grey, opaque.
there is no temperature anomaly in the surrounding water; in-
Morphology: Tubes smooth surfaced, straight, hick walled, ternal bacteria present probably sulphide oxidizers.
rigid, tapering to sinuous posterior region 10 to <5 mm diame-
Distribution: Valu Fa Ridge in the Lau Back-Arc Basin.
ter. The white obturaculum (15-42 mm long) has a funnel-
shaped top, with thin anterior rim and smooth inner surface, on
a slender stalk elliptical in section. Branchial filaments are par-
allel to the obturaculum forming lamellae of two types: pale
outer sheath lamellae (8-16 pairs) made of adherent filaments
without pinnules and red inner lamellae composed of pinnulate
filaments.

1: Left, anterior end of the 2: Anterior end of the animal showing obturaculum and 3: A tube projecting vertically,
tube and anterior end of the branchial filaments; cruise TUIM 06; by courtesy of G. Rouse. Valu Fa Ridge; cruise Biolau ©
animal, dorsal view; from Ifremer.
SOUTHWARD (1991).

References:
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.
SOUTHWARD E.C. (1991) J. Nat. Hist. 25: 859-881.

E.C. SOUTHWARD Denisia 18 (2006): 258


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Annelida, Polychaeta, Sabellida, Siboglinidae


Lamellibrachia satsuma MIURA, TSUKAHARA & HASHIMOTO, 1997

Size: Tube length max. 1400 mm; anterior diameter max. 9 Biology: This tubeworm species lives in shallow water at depths
mm. of 82-110 m in Kagoshima Bay and of about 300 m in Nankai
Trough. The worms make a cluster on sediment bottom. Most
Color: Tube grey/white.
of clusters are less than 5 m in length with about 1 m height,
Morphology: Tube rigid, thick walled, tapering to less than 1 but reaches more than 10 m in length and 3 m in height. Am-
mm at posterior end; short external flanges anteriorly, eroded bient temperature is constant at about 16°C throughout the
and indistinct in posterior half; sinuous but anterior parts more year. Internal bacteria present, sulfide oxidizers, may chiefly use
straight than posterior: Obturaculum cup- or funnel- shape, biogenic hydrogen sulfide rather than volcanic gas.
with smooth interior surface; stalk elliptical in transverse sec-
Distribution: West Pacific: Kagoshima Bay; Nankai Trough.
tion, with up to 19 pairs of branchial lamellae hidden by up to
four pairs of peripheral lamellar sheets. Vestimentum of ale
with paired ciliated grooves, diverging at anterior ends. Vesti-
mental cuticular plaques 35-63 mm in diameter, trophosomal
ones 51-82 mm.

3: Living specimens; by T. Miura.

1: Anterior end, ventral view, body fixed


after removal from the tube; by T. Miura.

2: Uncini of the opisthosome (SEM); 4: A clump of vestimentiferan tube worms, 82 m deep,


by T. Miura. Kagoshima Bay; by T. Miura © Jamstec.

References:
MIURA T., TSUKAHARA J. & J. HASHIMOTO (1997) Proc. Biol. Soc. Wash. 110: 447-456.
MIURA T., NEDACHI M. & J. HASHIMOTO (2002) J. Mar. Biol. Ass. U. K. 82: 537-540.

T. MIURA Denisia 18 (2006): 259


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Annelida, Polychaeta, Sabellida, Siboglinidae

Oasisia alvinae JONES, 1985

Size: Tube length max. 123 mm; anterior diameter 1.5-2.5 mm. Biology: Found on sites of diffuse venting; occurs with Tevnia
jerichonana; endosymbiotic sulfide oxidizing bacteria.
Morphology: Tube transparent, tapering, flexible, with widely
spaced flanges. Top of obturaculum with conspicuous central Distribution: East Pacific Rise: 21°N to 9°N.
axial rod carrying 2-3 translucent white saucer-like structures
(small specimens found at East Pacific Rise 9°N having 5-6
saucers); broad dorsal groove. Branchial filaments in concentric
lamellae, most filaments pinnulate, a few without pinnules on
dorsal end of basal lamellae. Paired excretory pores opening
dorsally at the base of obturaculum. Anterodorsal wings of ves-
timentum become sheath at base of obturaculum, continuing
ventrally, free from obturaculum and vestimentum proper.
Opisthosome with as many as 37 segments.

1: Right lateral view of the anterior 2 left: Tube; right: Habitus; from East Pacific Rise: 13°N; cruise Biocyarise;
region; by E. Southward. by P. Briand © Ifremer.

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


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Annelida, Polychaeta, Sabellida, Siboglinidae

Ridgeia piscesae JONES, 1985

Size: Tube length max. 1900 mm; anterior diameter 2-13 mm. Remark: R. piscesae was amended by SOUTHWARD et al. (1995)
to include R. phaeophiale JONES 1985
Morphology: Tube very variable, straight or sinuous, stiff or
soft, tapering to 1mm posterior end; anterior funnel and exter- Biology: Grows gregariously in clusters at sites of diffuse or
nal flanges present. Color white, grey, gold, brown; commonly warm venting, temperature range 5-60°C (?). Uses internal
translucent. Top of obturaculum carries 1-15 brown saucers on symbiotic sulphide-oxidizing bacteria. This species can even
a central axial rod; if these are shed, traces of the axial structure grow and survive in areas of low diffuse vent flow with very low
remain. Red branchial filaments in concentric lamellae parallel plume level exposure to sulfide.
to the obturaculum; most bear pinnules, but there are a few
Distribution: Explorer Ridge, Juan de Fuca Ridge, Gorda
thin, smooth filaments at the ends of each row.
Ridge.

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

References:
BLACK M. B. (1991) M.Sc., University of Victoria.
JONES M. L. & S.L. GARDINER (1989) Biol. Bull. 177: 254-276.
SARRAZIN J., ROBIGOU V., JUNIPER S.K. & J.R. DELANEY (1997) Mar. Ecol. Progr. Ser. 153: 5-24.
SOUTHWARD E.C., TUNNICLIFFE V. & M. BLAKE (1995) Can. J. Zool. 73: 282-285.
TUNNICLIFFE V. & S.K. JUNIPER (1990) Progr. Oceanogr. 24: 1-13.
TUNNICLIFFE V. (1988) Proc. R. Soc. London B 233: 347-366.
URCUYO I., MASSOTH G., JULIAN D. & C.R. FISHER (2003) Deep Sea Res. I 50: 763-780.

E.C. SOUTHWARD Denisia 18 (2006): 261


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Annelida, Polychaeta, Sabellida, Siboglinidae

Riftia pachyptila JONES, 1981 “giant tube worm“

Size: Individual tubes up to 1.5 m.


Morphology: Four body regions (1) anterior tentacular plume Biology: Forms clusters on rocks in zone of diffuse venting.
on obturaculum; winged vestimentum; trunk; segmented poste- Feeds only on internal symbiotic sulfide oxidizing bacteria.
rior opisthosome. Plume red, with tentacular lamellae perpen-
Distribution: Galapagos Spreading Center, East Pacific Rise
dicular to axis; paired halves of apical split end forming oper-
(including Guaymas Basin).
culum upon withdrawal into tube. Vestimentum with dorsolat-
eral flaps overlapping one another ventrally. Opisthosome with
variable number of segments, ending in rounded posterior tip.
Anterior segments completely encircled with paired single rows
of chaetae, becoming incomplete posteriorly. Tube white,
smooth, flexible, extremely sturdy, essentially cylindrical, basal-
ly blind ending. Tube made of a chitin-protein system.

1: Population in situ from East Pacific Rise: 13°N; cruise Phare  Ifremer.

D. DESBRUYÈRES Denisia 18 (2006): 262–263


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2: Specimen, total view (left) and detail of anterior body regions, dorsal view (middle), ventral view (right) at East
Pacific Rise: 9°N; by M. Bright.

References:

ARP A.J., CHILDRESS J.J. & R.D. VETTER (1987) J. Exp. Biol. 128: 139-158. GARDINER S.L. & M.L. JONES (1985) Trans. Amer. Microsc. Soc. 104: 19-44.
ARP A.J., DOYLE M.L., DI CERA E. & S.J. GILL (1990) Resp. Physiol. 80: 323- GIRGUIS P.R., LEE R.W., DESAULNIERS N., CHILDRESS J.J., POSPESEL M., FELBECK H. &
334. F. ZAL (2000) Appl. Environ. Microbiol. 66: 2783-2790.
BELKIN S., NELSON D.C. & H.W. JANNASCH (1986) Biol. Bull. 170: 110-121. GOFFREDI S. K., CHILDRESS J.J., DESAULNIERS N.T. & F.H. Lallier (1997) J. Exp. Bi-
BLACK M.B., LUTZ R.A. & R.C. VRIJENHOEK (1994) Mar. Biol. 120: 33-39. ol. 200: 2609-2616.
BRIGHT M., KECKEIS H. & C R. FISHER (2000) Mar. Biol. 136: 621-632. JONES M.L. (1981) Proc. Biol. Soc. Wash. 93: 1295-1313.
CHILDRESS J.J., ARP A.J. & C.R. FISHER (1984) Mar. Biol. 83: 109-124. MARSH A.G., MULLINEAUX L.S., YOUNG C.M. & D.T. MANAHAN (2001) Nature
CHILDRESS J.J., FISHER C.R., FAVUZZI J.A., KOCHEVAR R.E., SANDERS N.K. & A.M. 411: 77-80.
ALAYSE (1991) Bio. Bull. 180: 135-153. POWELL M.A. & G.N. SOMERO (1983) Science 219: 297-299.
HAAS F. de, ZAL F., LALLIER F., TOULMOND A. & J.N. LAMY (1996) Proteins 26: SCOTT K.M., FISHER C.R., VODENICHAR J.S., NIX E.R. & E. MINNICH (1994) Physi-
241-256. ol. Zool. 67: 617-638.
FELBECK H. (1985) Physiol. Zool. 53: 272-281. THIEBAUT E., HUTHER X., SHILLITO B., JOLLIVET D. & F. GAILL (2002) Mar. Ecol.
GAILL F., HERBAGE D. & L. LEPESCHEUX (1988) Oceanologica Acta Numéro Spé- Progr. Ser. 234: 147-157.
cial 8: 155-160. ZAL F., LALLIER F.H., WALL J.S. & S.N. VINOGRADOV (1996) J. Biol. Chem. 271:
GAILL F., SHILLITO B., MÉNARD F., GOFFINET G. & J.J. CHILDRESS (1997) Mar. Ecol. 8869-8874.
Progr. Ser. 148: 135-143.

263
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Annelida, Polychaeta, Sabellida, Siboglinidae

Siphonobrachia lauensis SOUTHWARD, 1991

lowed by two rows of dark crescentic plaques on the anterior


Size: Tube length max. 290 mm; anterior diameter 0.8-1.2 mm.
trunk papillae.
Color: Tube, transparent yellow anteriorly, then orange brown
Biology: The tubes are partly burried in sediment, projecting
darkening to opaque brown posteriorly.
vertically, between rocks covered with oxides and bacterial
Morphology: The tube is straight and stiff, with a funnel at the mats. Temperature “not elevated“. Internal bacteria present,
top and external collars at irregular intervals, tapering to < 0.7 probably sulphur-oxidizers.
mm posteriorly. No obturaculum. Tentacles (10-21) stuck to-
Distribution: Lau Back-Arc Basin: Valu Fa Ridge.
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-

1 left: Anterior end of the 2: Tubes in hydrothermal sediment, close to bacterial mats. Stalked barnacles and
tube; right: Anterior end of galatheids in background. Biolau cruise © Ifremer.
animal; by E. Southward.

References:
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.
SOUTHWARD E.C. (1991) J. Nat. Hist. 25: 859-881.

E.C. SOUTHWARD Denisia 18 (2006): 264


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Annelida, Polychaeta, Sabellida, Siboglinidae

Tevnia jerichonana JONES, 1985 ”Jericho worm“

Size: Tube length max. 350 mm; anterior diameter 3-8 mm. Biology: Uses internal symbiotic sulphide-oxidizing bacteria.
Forms cluster on rocks in zone of diffuse venting, temperature
Morphology: Tube sinuous, rigid, tapering to <1 mm posterior-
5-30°C; an early colonizer of new vent sites.
ly; closely spaced external flanges. Color white or gold/brown,
opaque. Top of obturaculum has yellow/brown crust and small Distribution: East Pacific Rise: 13°N to 21°S.
axial lamina; obturaculum is short (length < 2 x diameter);
conspicuous dorsal groove. Short red branchial filaments in
concentric lamellae parallel to opteraculum, most bear pin-
nules, but a few without pinnules at dorsal ends of rows.

2: Plume;
by courtesy of P. Baston.

1: Population in situ from East Pacific Rise: 13°N; cruise Hydronaut  Ifremer.
Upper left: Anterior region; drawing by E. Southward. Box: Tubes.

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.

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Annelida, Polychaeta, Scolecida, Maldanidae

Nicomache arwidssoni BLAKE, 1985 “bamboo worm“

chaetigers 1-3 with 1-2 blunt-tipped spines; rostrate chaetae


Size: A large species, up to 84 mm long, 3 mm wide for 21
from chaetiger 4 increasing to 9-10 per fascicle middle seg-
chaetigers.
ments; hooks with four apical teeth and tuft of bristles below
Morphology: Color in alcohol light to dark brown, with gold main fang. Anus surrounded by funnel bearing about 25 papil-
pigment on anterior end and dark, rusty pigment posteriorly; lae.
some EPR specimens with glandular areas appearing dark green
Biology: An epifaunal species, found in association with mus-
or black. Prostomium broadly rounded anteriorly, merging pos-
sels, clams, siboglinids, and other associations. Maldanids are
teriorly with arched cephalic keel; cephalic plate lacking,
known as bulk-ingestors of particles.
nuchal grooves curved. Segments longer than wide, flaring an-
teriorly, but not forming collars. Notochaetae capillaries; long Distribution: Galapagos Spreading Center; East Pacific Rise:
filamentous capillary notochaetae absent. Neurochaetae of 13°N, 21°N; Mariana Back-Arc Basin (questionable).

4: Tube  Ifremer.

2: Prostomium, lateral view


(preserved specimen)
 Ifremer.

1: Habitus, dosal
view; by V. Martin
 Ifremer.

3: Funnel with papillae sur- 5: Hooks with tuffs of bristles


rounding anus  Ifremer. (SEM)  Ifremer.

References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 6: 67-101.
BLAKE J.A. (1991) Proc. Biol. Soc. Wash. 104: 175-180.
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.
VAN DOVER C.L. (2002) Mar. Ecol. Prog. Ser. 230: 137-158.

J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 266


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Annelida, Polychaeta, Scolecida, Maldanidae

Nicomache venticola BLAKE & HILBIG, 1990

spines. Following neuropodia with 6-8 rostrate hooks in single


Size: A large species, up to 130 mm long, 2.5 mm wide, for 27-
rows, increasing to 9-10 in middle segments; hoods with main
32 segments.
fang surmounted by 2-3 teeth and with numerous fibrils below
Color: In alcohol greenish gray or light brown, with dark pig- main fang; unworn hooks with distinct tuft of fibrils emerging
mented areas sometimes apparent on neuropodia. below main fang; worn spines lacking tuft of fibrils; rostrate
hooks with prominent manubrium on shaft. Anus terminal,
Morphology: Prostomium rounded on anterior margin, contin-
surrounded by irregular funnel bearing about 22 papillae.
uing posteriorly as arched cephalic keel; cephalic plate lacking;
nuchal grooves curved. Anterior segments except first one or 2, Biology: An epifaunal species, collected from washings of larg-
longer than wide, flared anteriorly, overlapping preceding seg- er biota.
ments but not forming collars. Few posterior segments crowded,
Distribution: Juan de Fuca Ridge: Axial Seamount, Southern
with swollen parapodial lobes. Parapodia biramous. No-
Juan de Fuca Ridge; Explorer Ridge; Guaymas Basin.
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

1: Specimen in vivo; by courtesy of V. Tunnicliffe. 2: Prostomium, lateral view showing arched cephalic
keels (SEM) © Ifremer.

3: Neuropodium of chaetiger III with four acicular spines 4: Rostrate hooks of middle segments, with main fang
(SEM) © Ifremer. surmounted by 2-3 teeth and with numerous fibrils below
main fang (SEM) © Ifremer.

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

J.A. BLAKE Denisia 18 (2006): 267


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Annelida, Polychaeta, Scolecida, Orbiniidae

Leitoscoloplos pachybranchiatus BLAKE & HILBIG, 1990

rochaetae camerated capillaries; abdominal neurochaetae in-


Size: Small species, up to 9.0 mm long and 0.64 mm wide for 54
cluding 2-3 capillaries and 1-2 thin acicular spines. Branchiae
chaetigers.
from chaetiger 13-15; each thick in cross section, appearing
Color: In alcohol tan. glandular; anterior and posterior abdominal branchiae short,
stubby in appearance, with broad base and narrow apex;
Morphology: Prostomium conical, smoothly rounded along an-
branchiae from middle abdominal segments, longer, but still
terior margin, with small nuchal organs on lateral boundary of
with broad base. Pygidial segment broadly rounded, with two
prostomium and peristomium. Peristomium with one or two
lateral cirri.
achaetous rings depending on size and state of preservation;
both rings apparent in smaller specimens, vaguely apparent in Biology: Exact habitat unknown, collected near an active vent.
larger specimens. Thorax with nine similar chaetigers. Notopo- The species probably burrows into mud and is likely a deposit
dia with thin, cirriform postchaetal lobes; neuropodia, with feeder. Two large eggs measuring approximately 100 mm in di-
short triangular shaped postchaetal lobes, developing thick- ameter were observed in one specimen. They were enclosed in
ened base in abdominal region; neuropodial lobe of abdominal a chamber that was located on the dorsum between chaetigers
segments becoming thicker, blunted, then developing weak 6 and 7.
bilobed appearance in far abdominal segments. Notochaetae
Distribution: Juan de Fuca Ridge.
camerated capillaries; furcate chaetae absent. Thoracic neu-

1: Prostomium, lateral view (SEM) 2: Nucchal organ (SEM) 3: Camerated capillaries (SEM)
© Ifremer. © Ifremer. © Ifremer.

4: Lateral view showing transition 5: Lateral view showing abdominal 6: Far abdominal segments, lateral
between thoracic and abdominal paramodia with thick glandular view (SEM) © Ifremer.
regions (SEM) © Ifremer. branchiae (SEM) © Ifremer.

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

J.A. BLAKE & D. DESBRUYÈRES Denisia 18 (2006): 268


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Annelida, Polychaeta, Scolecida, Orbiniidae

Orbiniella aciculata BLAKE, 1985

neurochaetae include 1-4 acicular spines and 2-10 long barbed


Size: Small, probably no more than 4 mm long, 0.6 mm wide,
capillaries; number of chaetae size dependent, smaller speci-
for 25 segments.
mens with fewest chaetae. Pygidium with two short cirri.
Color: In alcohol opaque white.
Biology: Infaunal, in mud; exact habitat unknown, may not be
Morphology: Body small, wide, not divided into thorax and ab- a true vent species; a deposit feeder.
domen. Prostomium broadly rounded along anterior margin;
Distribution: Galapagos Spreading Center hydrothermal vent
eyes and nuchal organs absent. Peristomium consisting of two
fields.
achaetous rings with first narrower than second. Noto- and

1A: Dorsal view; B: Anterior end, dorsal view; C: Acicular spine from notopodium; D: Basal part
of capillary notochaetae; from BLAKE (1985).

Reference:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.

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Annelida, Polychaeta, Scolecida, Orbiniidae

Orbiniella hobsonae BLAKE & HILBIG, 1990

again. Pygidium simple, without lobes or cirri. Parapodia re-


Size: Small, threadlike, up to 4.5 mm long, 0.2 mm wide, for 25
duced, without distinct podial or postchaetal lobes; branchiae
chaetigers.
lacking. Chaetae include 1-2 barbed acicular spines and 2-5
Color: In alcohol opaque white. bristled capillaries in both noto- and neuropodia; bristles of
capillaries arranged in uniform transverse rows.
Morphology: Prostomium nearly circular in outline, rounded
along anterior margin, with pair of lateral nuchal organs. First Biology: Unknown, but probably living in crevices containing
achaetous peristomial ring reduced, compressed between pros- pockets of soft sediments; a deposit feeder.
tomium and larger second ring. All chaetigers similar, without
Distribution: Juan de Fuca Ridge.
apparent abdominal or thoracic regions. Chaetigers 1-4 gener-
ally narrower, shorter, more compressed; middle chaetigers larg-
er, more elongate; posterior chaetigers becoming compressed

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.

J.A. BLAKE Denisia 18 (2006): 270


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Annelida, Polychaeta, Scolecida, Orbiniidae

Scoloplos ehlersi BLAKE, 1985

lowing segments; abdominal neuropodia elongate, flattened,


Size: Up to 10 mm long, 0.5 mm wide, for 50+ segments.
with ventral lobe in middle chaetigers. Branchiae from
Color: In alcohol opaque white, with golden flocculent materi- chaetiger 21, continuing to posterior end [late occurrence of
al observed in gut. branchiae unusual]. Thoracic notopodia with fascicles contain-
ing numerous camerated capillaries; neuropodia with dense fas-
Morphology: Body elongate, cylindrical in cross section; ante-
cicles of capillaries and short, ribbed spines.
rior nine chaetigers swollen. Thorax consisting of 11
chaetigers; transition to abdomen abrupt, denoted by appear- Biology: Infaunal, in mud; may not be a true vent species; a de-
ance of elongate neuropodial cirrus and disappearance of tho- posit feeder.
racic neuropodial uncini. Prostomium elongate, pointed anteri-
Distribution: Galapagos Spreading Center, hydrothermal vent
orly; eyes and nuchal organs absent. Parapodia of chaetigers 1-
fields.
3 without postchaetal lamellae; from chaetiger 4, notopodia
with fingerlike postchaetal lamellae, becoming longer over fol-

1A: Anterior end dorsal view; B: Chaetigers 10-13, dorsal view; C: Left parapodium from abdominal segment, posterior
view; D: Thoracic capillary notochaetae; E: Abdominal flail chaetae; F: Thoracic acicular neurochaetae; from BLAKE (1985).

References:
BLAKE J.A. (1985) Bull. Biol. Soc. Wash. 8: 67-101.
VAN DOVER C.L. (2002) Mar. Ecol. Progr. Ser. 230: 137-158.

J.A. BLAKE Denisia 18 (2006): 271


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Annelida, Polychaeta, Spionida, Chaetopteridae

Spiochaetopterus SARS, 1853

topodia of the posterior region are erected, slightly inflated pos-


Size: Unknown, only fragmented specimens.
teriorly.
Color: Dark tan in alcohol with ventral white glandular shield
Remarks: Likely two different species of Spiochaetopterus are
extending from segment 5-8.
present at Mid-Atlantic Ridge vent fields. The poor conserva-
Morphology: Body long, slender, soft; tubes ringed, horny. Pros- tion state of the collections does not allow a specific identifica-
tomium small, and enfolded by the truncate buccal segment tion; however, the shape of the specialized chaetae is unique.
which bears long grooved palps but no tentacular cirri. Fourth
Biology: All chaetopterids have highly modified body regions
chaetiger with a fan of lanceolate chaetae, a large and long
that enable them to strain water through mucous bags. All
spine with a truncated top, slightly cordate and a small aristate
chaetopterids secrete the tube from glands located on the ven-
spine. Middle region (very damaged) with numerous segments
tral side of the anterior body region.
(>15), with two rami, all similar with a bilobed Y-shaped inner
part and a small unilobed foliaceous outer part. The neuropodia Distribution: Mid-Atlantic Ridge: TAG and Rainbow.
are unilobed on B1 and bilobed on following segments. The no-

2: View of the extremity and the


1: Chaetopterid on the bottom at middle part of the tube, by P. 3: Lateral view, anterior part of a specimen collected at
TAG; cruise Exomar © Ifremer. Briand © Ifremer. the Rainbow vent field; by P. Briand © Ifremer.

4: Long and thick


4: Dorsal view of the posterior part of a specimen 5: Short and thick aristate chaeta from segment slightly cordate spine
collected at the Rainbow vent field; by P. Briand © four (SEM) © Ifremer. from chaetiger four
Ifremer. (SEM) © Ifremer.

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.

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Annelida, Polychaeta, Spionida, Spionidae

Laonice athecata SIGVALDADÓTTIR & DESBRUYÈRES, 2003

one reduced, round ventrally, with dorsal projection. On subse-


Size: Up to 51 mm long.
quent chaetigers lamella larger, dorsally and ventrally evenly
Color: Greenish in vivo, white when preserved in ethanol. projected. On posterior chaetigers lamella becoming smaller
and less projecting. Interparapodial pouches missing. Branchi-
Morphology: Prostomium broadly triangular with straight an-
ae from chaetiger two, present on about 30 chaetigers. Branchi-
terior margin, laterally tapering backwards into a caruncle
ae smooth, little longer than notopodial lamellae, heavily cili-
reaching to posterior margin of chaetiger four. Small occipital
ated laterally. Chaetae of anterior chaetigers arranged in simple
antennae on level of chaetiger one. Eyes absent. Nuchal organ
row, not particularly long, becoming longer and reduced in
arranged in two rows enclosing caruncle. Peristomium sur-
number on posterior chaetigers. Neuropodial hooded hooks
rounding prostomium almost completely, laterally forming
from chaetiger 30-32. Hooks with one pair of teeth over main
prominent lateral wings, partly fused with chaetiger one.
fang which is slightly grooved longitudinally. Notopodial hooks
Postchaetal notopodial lamellae of chaetiger one reduced, tri-
absent. Sabre chaeta from chaetiger 20-23. Chaetae long, thin,
angular, with dorsal projection. Lamellae on following
with a curved delicate tip. Pygidium with 6-8 short cirri.
chaetigers large, foliate, with a dorsal protrusion, extending al-
most over dorsum. Lamellae successively becoming smaller, on Distribution: Mid-Atlantic Ridge: Lucky Strike and Loga-
posteriormost chaetigers ventrally round with dorsal tip. tchev.
Prechaetal lamellae small. Neuropodial lamellae on chaetiger

1: Anterior part (preserved specimen)  Briand/Ifremer.


2: Anterior part (SEM)  Ifremer.

3: Neuropodial lamella with hooded hooks, and sabre chaeta 4: Hooded hook showing pair of teeth over main fang (SEM)
(SEM)  Ifremer.  Ifremer.

Reference:
SIGVALDADOTTIR E. & D. DESBRUYÈRES (2003) Cah. Biol. Mar. 44: 219-225.

D. DESBRUYÈRES Denisia 18 (2006): 273


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Annelida, Polychaeta, Spionida, Spionidae

Laubieriellus grasslei MACIOLEK, 1981

sabre chaetae from chaetiger 10-11. Pygidium with two short,


Size: Small, up to 11.5 mm long, 0.75 mm wide, with about 45
rounded lobes and one long cirrus (D: E).
segments.
Biology: A cryptic species, found among mussels, from experi-
Morphology: Prostomium rounded anteriorly, continuing pos-
mental fouling panels, and present with other invertebrates
teriorly as caruncle to chaetiger 2-3 (A-C). Peristomium sepa-
washed from various collections. The species is likely a surface
rated from chaetiger 1, without lateral wings. Branchiae
deposit feeder that forms temporary tubes.
smooth, apinnate, four pairs, from chaetiger 2. Neuropodia of
anterior segments connected by ventral crests from chaetiger 2; Distribution: Galapagos Spreading Center, hydrothermal vent
dorsal crests present on several postbranchial chaetigers (A). fields.
Notochaetae all capillaries; neurochaetae include capillaries
and multidentate hooded hooks from chaetiger 10; ventral

1A: Anterior end, dorsolateral view; B: Anterior end, lateral view; C: Ante-
rior end, dorsal view; D, E: Pygidium, ventral view; from MACIOLEK (1981).

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

J.A. BLAKE Denisia 18 (2006): 274


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Annelida, Polychaeta, Spionida, Spionidae

Lindaspio dibranchiata BLAKE & MACIOLEK, 1992

veloped by chaetiger 30 and fully developed by chaetiger 40;


Size: A large species, holotype 31 mm long, 4 mm wide, with
ventral branchiae broader than dorsal branchiae, not meeting
165 segments.
at ventral midline, continuing to posterior end. Notochaetae of
Color: Brown in alcohol. chaetigers 2-4 modified into unique rosette of 9-10 heavy
spines; subsequent notochaetae consisting of capillaries until
Morphology: Anterior end broad, widest between chaetigers
about chaetiger 40 where 3-5 unidentate hooded hooks begin;
12-20, dorsoventrally flattened throughout. Body terminating
hooks sharply curved with closely adhering hood. Neu-
in conical pygidial cone, lacking appendages. Prostomium bifid
rochaetae of chaetigers 1-5 capillaries; with distinct fascicles of
anteriorly, with two thick rounded lobes, extending posteriorly
15-20 heavy spines and thin capillaries from chaetigers 6-28;
as short, mounded caruncle to chaetiger 1. Palps short, thick.
spines replaced by multidentate hooded hooks by about
Chaetiger 1 reduced, lacking notochaetae; following segments
chaetiger 45; neuropodial spines distinctly tapered distally,
with well-developed noto- and neuropodia bearing elaborate
when worn, appearing acicular; neuropodial hooks thinner,
pre- and postchaetal lamellae encompassing chaetal fascicles.
more delicate than notopodial hooks.
Notopodia of middle and posterior chaetigers with anterior and
posterior lamellae, but these shorter, more triangular and only Biology: A large benthic infaunal species, probably a surface
partly encompassing chaetae. Dorsal branchiae present from deposit feeder.
chaetiger 2, each anterior branchia thickened, extending across
Distribution: Guaymas Basin, Southern Trough, in sediments
dorsal midline; after chaetiger 20, becoming very thin, contin-
of hydrothermal mounds.
uing to posterior end. Ventral branchiae from about chaetiger
20 as swollen protuberance of neuropodium, becoming well de-

1A: Anterior end, dorsal view; B: Posterior parapodium, anterior view; C: Modified neuropo-
dial spines from anterior chaetiger; D: Neuropodial hooded hooks; E: Notopodial hooded
hooks; from BLAKE & MACIOLEK (1992).

Reference:
BLAKE J.A. & N.J. MACIOLEK (1992) Proc. Biol. Soc. Wash. 105 (4): 723-732.

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Annelida, Polychaeta, Spionida, Spionidae

Lindaspio southwardorum BLAKE & MACIOLEK, 1992

cylindrical; in far posterior segments, nearly reaching ventral


Size: A large species, recorded up to about 160 mm long, 6-7
midline, but never as long as dorsal branchiae. Notochaetae of
mm wide, for more than 340 segments.
chaetigers 2-4 modified into cluster of about 20 heavy spines;
Color: In life pinkish white with pale violet streak down dor- subsequent notochaetae numerous, thin capillaries; 5-8 hooded
sum; in alcohol light brown. hooks from chaetiger 75; capillaries become heavier and more
limbate in far posterior segments; individual notopodial hooks
Morphology: Anterior half of body dorsoventrally flattened,
strongly curved, with pointed main fang surmounted by several
with dorsum becoming rounded posteriorly. Pygidium not ob-
minute teeth. Anterior neurochaetae include row of 25-30
served. Prostomium narrow, pear-shaped, flaring anteriorly,
heavy spines, thin companion capillaries, and ventral bundle of
forming two broadly swollen lobes, continuing posteriorly as
thin capillaries; neuropodial spines each with smooth shaft that
narrow, folded caruncle to anterior margin of chaetiger 2. Palps
tapers abruptly, then continues as fine, pointed tip with fine
short, thick. Chaetiger 1 reduced, lacking notochaetae. No-
serrations or bristles sometimes visible along edge; spines pres-
topodia of chaetigers 2-4 modified, dorsally elevated, with pre-
ent until about chaetiger 40, then replaced by thin capillaries;
and postchaetal lamellae forming cup enclosing cluster of mod-
neuropodial hooded hooks from about chaetiger 75; each hook
ified spines; notopodia from chaetiger 5 and neuropodia from
smaller, more delicate than notopodial hook; each hook with
chaetiger 2 with well-developed, elaborate pre- and postchaetal
several minute teeth above main fang.
lamellae enclosing chaetal fascicles; notopodial lamellae even-
tually becoming more elongate, somewhat triangular; neuropo- Biology: A benthic infaunal species living in sediments near
dial lamellae remaining broadly rounded throughout. Dorsal hydrothermal vents; the species is likely a surface deposit feed-
branchiae first present from chaetiger 2, each relatively short er.
for first 45-50 chaetigers, extending only half of distance to
Distribution: Juan de Fuca Ridge: Middle Valley Segment, in
midline, thereafter branchiae becoming thinner, longer, ex-
sediment from high heat areas.
tending to midline. Ventral branchiae from about chaetiger 55
as extensions of postsetal lamellae, remaining relatively short,
until about chaetiger 100-125, then becoming longer, more

J.A. BLAKE Denisia 18 (2006): 276–277


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

1A: Anterior end, dorsal view; B: Anterior parapodium anterior view; C: Middle parapodium, anterior
view; D: Posterior parapodium, anterior view; E: Modified anterior neurochetae and accompanying
capillaries; F-G: Notopoal hooked hooks; H: Neuropodial hooked hooks; from BLAKE & MACIOLEK (1992).

Reference:
BLAKE J.A. & N.J. MACIOLEK (1992) Proc. Biol. Soc. Wash. 105(4): 723-732.

277
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Annelida, Polychaeta, Spionida, Spionidae

Prionospio sandersi MACIOLEK, 1981

Size: Small, up to 3.2 mm long, 0.5 mm wide, with about 35 Remark: A closely related form of Prionospio sp. has been com-
segments. monly sampled on the Northern East Pacific Rise at 13°N and
9°N. It differs from Prionospio sandersi by the number of
Morphology: Prostomium broadly rounded anteriorly, ending
branchiae (12-14 pairs instead nine pairs) by the occurrence of
posteriorly at chaetiger 1. Peristomium encompassing prostomi-
the sabre chaetae on neuropodia from chaetiger 15 instead of
um laterally and ventrally, not developing lateral wings; fused
14 and the occurrence of the multidentate hooks on neuropo-
dorsally to chaetiger 1. Branchiae from chaetiger 2, nine pairs;
dia from chaetiger 17 instead 15 and in notopodia from
each gill broad, robust, appearing wrinkled, but lacking pin-
chaetiger 23 instead of 29.
nules; branchiae longest anteriorly, becoming shorter, stubby
posteriorly. Dorsal crests absent. Anterior chaetae capillaries; Biology: The known specimens were collected from washings
multidentate hooded hooks from chaetiger 15 in neuropodia, of Riftia pachyptila and might no be a fully developed. The
chaetiger 29 in notopodia; ventral sabre chaetae from chaetiger species is likely a surface deposit feeder, scavenging particles
14. Pygidium cup-shaped, with deeply rounded ventral edge with its short palps.
and small dorsal elongation.
Distribution: Galapagos Spreading Center, hydrothermal vent
fields.

2: Living specimen from East Pacific Rise: 13°N; by P.


Briand © Ifremer.

1: Drawings from MACIOLEK (1981). A: Anterior end, dorsolateral view; B:


Prostomium, dorsal view; C: Branchia, dorsal view, showing details of 3: Two neuropodia of the middle part of the body in
the wringled surface; D: Ventral sabre chaetae; E, F: Hooded hook; G: lateral view showing sabre chaetae and multidentate
Pygidium, lateral view; H. Pygidium, ventral view. hooks (SEM); East Pacific Rise: 13°N © Ifremer.

J.A. BLAKE Denisia 18 (2006): 278–279


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

4: A microphotograph mosaic, lateral 5: A microphotograph mosaic, dorsal


view (SEM); East Pacific Rise: 13°N view (SEM); East Pacific Rise: 13°N
© Ifremer. © Ifremer.

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

279
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Annelida, Polychaeta, Spionida, Spionidae

Prionospio unilamellata SIGVALDADÓTTIR & DESBRUYÈRES, 2003

chaetigers lamellae leaflike, dorsally pointed. Prechaetal lamel-


Size: Up to 17 mm long and 0.9 mm width for 114 chaetigers.
lae separated from postchaetal lamellae. Branchiae from
Morphology: Prostomium broadly triangular, widest at anterior chaetiger two, 15-16 in number. Anterior branchiae slightly
margin, narrowing into a raising caruncle, reaching to posteri- longer than notopodial lamellae, on subsequent chaetigers suc-
or delineation of first chaetiger. Anterior margin sometimes cessively longer and more slender. Branchiae apinate with
with 3-4 ciliated tubercles. Peristomium partly surrounding dense lateral ciliation. Interparapodial pouches missing. Small,
prostomium, forming a low collar, not fused to chaetiger one. papilla-like, structures can sometimes be observed laterally be-
Pair of indistinct ciliated nuchal organs situated on each side of tween neuropodia. These structures start on 2-3 posteriormost,
caruncle. Postchaetal notopodial lamella of chaetiger one re- or posterior to, branchial segments and occur on 7-8 segments.
duced, dorsally pointed, free from prechaetal lamella. On Chaetae on anterior chaetigers slender, arranged in dense dou-
chaetigers 2 to 16-17, a single large notopodial lamella is en- ble rows. Neuropodial hooded hooks from chaetiger 19-27. In-
closing chaetae. Postchaetal lamellae on postbranchial seg- troduction of hooks moving backwards with increasing size.
ments becoming smaller, separated from prechaetal lamellae, Hooks with 4-6 secondary teeth over main fang. Notopodial
round in form. On anterior chaetigers, postchaetal lamellae hooks from chaetiger 25-46. Notopodial hooks with longer
with dorsal tip. On posteriormost chaetigers lamellae dorsally shaft than neuropodial hooks. Sabre chaeta from chaetiger 12-
pointed. Prechaetal lamellae small on postbranchial chaetigers. 20, one or two per rami. Sabre chaeta distally granulated. Py-
Dorsal crests absent. Neuropodial postchaetal lamellae reduced gidium with long median cirrus and two rounded lateral flaps.
on first chaetiger, elliptical in shape. Lamellae largest on
Distribution: Mid-Atlantic Ridge: Lucky Strike, Rainbow and
branchial chaetigers, successively becoming smaller. On
Snake Pit.
chaetigers 2-3 lamellae ventrally round, dorsally with a tip, on
subsequent chaetigers becoming more round. On posteriormost

1: Habitus (preserved specimen)  Ifremer. Box: Distal 2: Anterior part (SEM)  Ifremer.
part of notopodial hooded hook (SEM)  Ifremer.

Reference:
SIGVALDADOTTIR E. & D. DESBRUYÈRES (2003) Cah. Biol. Mar. 44: 219-225.

D. DESBRUYÈRES Denisia 18 (2006): 280


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Annelida, Polychaeta, Spionida, Spionidae

Xandaros acanthodes MACIOLEK, 1981

straighter, then replaced by bidentate hooks with half hoods


Size: A small species, 6 mm long, 0.3 mm wide, with about 55
from chaetiger 16 continuing to posterior end; spines and hood-
segments.
ed hooks accompanied by 4-5 large and 2-3 thin capillaries
Morphology: A unique spionid. Prostomium rounded anterior- throughout; ventral sabre chaetae absent. Pygidium with two
ly, without posterior keel or caruncle. Peristomium well devel- dorsal and two larger ventral lobes.
oped, but not producing lateral wings. Notochaetae absent on
Biology: Epifaunal, collected among siboglinids and mussels.
chaetiger 1. Branchiae from chaetiger 4, continuing to
The species is likely a surface deposit feeder.
chaetiger 10-13; each gill elongate, cylindrical, wrinkled, lack-
ing distinct pinnules. Notochaetae capillaries; neurochaetae in- Distribution: Galapagos Spreading Center, hydrothermal vent
clude capillaries on chaetigers 1-2, replaced with unhooded aci- fields.
cular spines on chaetigers 3-10, these in turn becoming thinner,

1A: Anterior end, dorsal view, palps and several branchiae removed; B: Anterior end, lateral view; C: Distal tip of branchia in
lateral view; D: Entire branchia dorsal view; E: Pygidium and last three chaetigers, lateral view; from MACIOLEK (1981).

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

J.A. BLAKE Denisia 18 (2006): 281


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Annelida, Polychaeta, Terebellida, Alvinellidae

Alvinella caudata DESBRUYÈRES & LAUBIER, 1986

Size: Up to 140 mm. Biology: Dwelling inside organic tubes in active chimney walls.
Temperature ranging from 10-50°C. Seldom observed in situ.
Morphology: Four pairs of lamellate gills; two transformed
Feeding on free bacteria, associated with filamentous bacteria
chaetigerous segments bearing stout spines; body in two parts,
epibiosis. Gut functional; retractile buccal tentacles; gonochor-
the posterior one with enlarged notopods bearing distal digita-
ic, sexual dimorphism.
tions; bacterial epibiosis.
Distribution: East Pacific Rise: 21°N to 17°S (not observed in
Guaymas Basin and Galapagos Spreading Center).

2: Habitus, ventral view;


by V. Martin  Ifremer.

1: Population in situ (East Pacific Rise: 13°N, 2630 m);


Phare cruise  Ifremer.

3: “Caudal“ part in dorsal view; 4: “Caudal“ part in ventral view;


by P. Briand  Ifremer. by P. Briand  Ifremer.

References:
ALAYSE-DANET A.M., DESBRUYÈRES D. & F. GAILL (1987) Symbiosis 4: 51-62.
CHEVALDONNÉ P. & D. JOLLIVET (1993) Mar. Ecol. Prog. Ser. 95: 251-262.
DESBRUYÈRES D. & L. LAUBIER (1980) Oceanol. Acta 3: 267-274.
DESBRUYÈRES D., GAILL F., LAUBIER L. & Y. FOUQUET (1985) Bull. Biol. Soc. Wash. 6: 103-116.
DESBRUYÈRES D. & L. LAUBIER (1986) Can. J. Zool. 64: 2227-2245.
GAILL F. & S. HUNT (1991) Rev. Aquat. Sci. 4: 107-137.
JOLLIVET D. (1993) Thèse de doctorat de l’Université de Bretagne Occidentale.

D. DESBRUYÈRES Denisia 18 (2006): 282


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Annelida, Polychaeta, Terebellida, Alvinellidae

Alvinella pompejana DESBRUYÈRES & LAUBIER, 1980 “Pompei worm“

species, although no significant morphological differences have


Size: Up to 150 mm.
been detected.
Morphology: Tube dwelling polychaetes with four pairs of
Biology: Dwelling inside organic tubes in active chimney walls.
lamellate branchiae; two transformed chaetigerous segments;
Temperature ranging from 10-80°C (debated?). Feeding on free
body tappering posteriorly, no obvious abdominal part; bacteri-
bacteria, associated with filamentous bacteria epibiosis. Func-
al epibiosis on the dorsum. Retractile buccal tentacles and two
tional gut, retractile buccal tentacles, gonochoric, sexual di-
specialized sexual ones.
morphism.
Remark: HURTADO et al. (2004) found a very significant mito-
Distribution: East Pacific Rise: 21°N to 23°S (not observed in
chondrial divergence between north and south EPR popula-
Guaymas Basin and Galapagos Spreading Center).
tions, leading to hypothesize the presence of a southern cryptic

2: Cluster of
filamentous bacteria
from dorsal part of
the worm (SEM)
 Ifremer.

3: Transformed
notosetae on two
anterior segments
 Ifremer.

1: Population in situ; East Pacific Rise: 13°N;


Cruise Phare  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.

D. DESBRUYÈRES Denisia 18 (2006): 283–284


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

4: Specimen in vivo; by Dugornay © Ifremer.

284
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Miralvinella) bactericola DESBRUYÈRES & LAUBIER, 1991

covered with minute spines. Uncinigerous neuropodial tori


Size: Up to 42 mm for 130 segments.
present from chaetiger 35-37 to the end of the body. Unciniger-
Morphology: Alvinellid worm with four pairs of pinnate ous tori increasing slightly in length posteriorly. Uncini bre-
branchiae. Animals brownish in colour after preservation. Pros- viacicular, stout as in other alvinellids, arranged in a single ver-
tomium reduced, with a dorsal lobe, having a dorsal incision tical row and oriented with the teeth pointing backwards. Py-
and two well developed lateral expansions. Dorsal part of the gidium rounded without appendages.
dorsal cavity with several rows of smooth oral tentacles with a
Biology: The animal seems to live freely in the sediment with-
ciliated groove, surrounding dorsal organ dorsally. Dorsal organ
out conspicuous tubes. On living animals, the tentacles extend
with a deep median hollow ending ventrally in two stout lobes
far over the bacterial mats and move gently in the moiré water,
from which arise two large, long and strong pointed tentacles
while the tips of the branchiae reach the surface of the sedi-
(male), each with a deep longitudinal groove without any cili-
ment.
ation. Tentacles are often rolled in a spiral. Notopodia of 7th
chaetigerous segment modified with 4-6 stout and slightly Distribution: Guaymas basin.
curved acicular hooks instead of capillary chaetae. Notopodia
of chaetigers 8-32 with digitiform lobes. Capillary notochaetae

2: Anterior end, ventro-lateral view (pre-


served specimen) © Ifremer.

1: Habitus, lateral view of a male specimen by V. Mar-


tin © Ifremer. Upper box, prostomium, dorsal view; 3: Buccal part of a male specimen showing
center, buccal segment ventral view; bottom, the sexual grooved tentacles and dorsal
chaetigers 7 and 8, showing digitiform lobes (SEM) view (SEM) © Ifremer.
© Ifremer.

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

D. DESBRUYÈRES Denisia 18 (2006): 285


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Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Miralvinella) dela DETINOVA, 1988

and slightly curved acicular spines in the 7th chaetiger.


Size: Up to 110 mm (170 chaetigerous segments).
Uncinigerous neuropodial tori from chaetiger 48-52 to the end
Color: Dark red when preserved. of the body. No notopodial or anal cirrus.
Morphology: Alvinellid worm with four pairs of pinnate Remarks: The species is a sister species of P. bactericola from
branchiae. Prostomium very reduced, similar to that of P. bac- Guaymas basin and P. hessleri from Western Pacific Back-Arc
tericola with a median lobe having a median incision and two Basins which were grouped in the subgenus Miralvinella.
very well developed and bilobed lateral expansions. Numerous
Biology: This worm is usually found in tube worm clusters.
grooved buccal tentacles inserted on two lateral pads. As in P.
grasslei, a median hemispherical body is observed ventrally in Distribution: Juan de Fuca Ridge: Axial Seamount, sites Ashes
the buccal cavity, bordered by two lateral pads. Two long point- and Casm.
ed and grooved peribuccal tentacles are present in males, each
with a deep longitudinal groove devoid of ciliation. Four stout

1: Female specimen from Axial Seamount, frontal view 2: Living specimens in: “A Bestiary of
© Ifremer. the Endeavour Hot Vents”; by cour-
tesy of V. Tunnicliffe.

References:
DESBRUYÈRES D. & L. LAUBIER (1989) Proc. Biol. Soc. Wash. 102: 761-767.
DESBRUYÈRES D. & L. LAUBIER (1993) Proc. Biol. Soc. Wash 106: 225-236.
DETINOVA N.N. (1988) Zool. Zh. 67: 858-864.
DETINOVA N.N. (1989) Trans. Shirshov Instit. Oceanol. 123: 71-80.

D. DESBRUYÈRES Denisia 18 (2006): 286


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Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Miralvinella) hessleri DESBRUYÈRES & LAUBIER, 1989

digitiform lobe bearing two groups of capillary chaetae, one


Size: Up to 22 mm.
with short and the other with long. Chaetigerous segment 7
Morphology: Alvinellid worm with four pairs of pinnate strongly modifies lacking cylindrical notopodia but bearing on
branchiae. Up to 61 segments. Prostomium medially reduced each side 4-5 strongly modified acicular hooks directed posteri-
with a median incision and two well developed lateral lobes orly. Segment 8 with cylindrical notopodium and very strong
ventrally enclosing peristomium. Male buccal apparatus com- digitiform lobe directed forward. Uncinigerous neuropodial tori
prising a ventral globular bulky organ, two lateral strong and on each segment from segment 15-20 to the end of the body.
pointed tentacles bearing a deep groove without ciliation, and
Biology: All specimens found in tubes on rocks directly ex-
many grooved and ciliated smaller tentacles inserted in two
posed to venting water whose temperature was recorded up to
groups on a quadrilobed upper lip. First two segments (II and
25°C. Tubes whitish and corneous in aspect, amoeba like in
III) achaetous and fused to the first three chaetigerous. The first
shape with long anchor filaments. Tube walls thick and multi-
15-20 chaetigerous segments with notopodia only. First three
layered. Inner surface bearing important bacterial mats.
notopodia smaller than others and dorsally elevated. Chaetiger
4 with a median dorsal expansion which protrude forward. No- Distribution: Mariana and Manus Back-Arc Basins.
topodia from 4 to 13-17 (7 excepted) cylindrical with a dorsal

2: Male buccal apparatus (SEM)


1: Habitus;  Ifremer.
by V. Martin.
Box: Anterior part,
dorsal view (SEM)
 Ifremer.

3: Cross cut of a tube wall, show-


ing epibiotic bacteria on the inner
part (SEM)  Ifremer.

Reference:
DESBRUYÈRES D. & L. LAUBIER (1989) Proc. Biol. Soc. Wash. 102: 761-767.

D. DESBRUYÈRES Denisia 18 (2006): 287


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Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Nautalvinella) pandorae DESBRUYÈRES & LAUBIER, 1986

two adjacent lines. Uncinigerous tori from fifth chaetigerous


Size: Up to 20 mm.
segment (P. p. p.) or sixth chaetigerous segment (P.p.i.). No
Color: Light brown to pinkish in ethanol. digitiform lobes.
Morphology: Body tapering posteriorly comprising about 60 Biology: Associated with early colonizer siboglinids (e.g. Oa-
chaetigerous segments. Seventh chaetigerous segment with no- sisia, Tevnia). Continuous or semi-continuous recruitment (sin-
topodial chaetae deeply modified forming stout hooks directed gle mode size-frequency distribution). MCHUGH (1989) hy-
backward. Prostomium reduced. Buccal apparatus retractile pothesized a brooding of larvae by adults.
made of a median groove with a pointed end. Numerous buccal
Distribution: P. p. pandorae is found on North East Pacific:
tentacles grooved and smooth inserted in two groups. The first
Juan de Fuca Ridge; P. p. irlandei is found at the East Pacific
three chaetigerous segments bearing only notopodia with
Rise: 21°N to 20°S.
chaetae. The branchial region comprises the first achaetous
segment plus the three following chaetigerous segments. Four
pairs of pinnate gills with lanceolated secondary filaments in

1: Drawing of
P. p. pandorae;
by V. Martin
 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 the 4: P. p. irlandei (East Pacific Rise: 13°N), ventral view; Box:
sixth chaetigerous segment  Briand/Ifremer. Median groove (SEM)  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.

D. DESBRUYÈRES Denisia 18 (2006): 288


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Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Nautalvinella) unidentata DESBRUYÈRES & LAUBIER, 1993

cepted 7th) cylindrical bearing two groups of capillary chaetae.


Size: 4.8-11 mm in length for 77-88 chaetigerous segments.
Notopodia without digitiform lobes. Chaetiger 7 strongly mod-
Color: Pale grey-pinkish in ethanol. ified bearing two to three straight short acicular notopodial
chaetae on each side. Uncinigerous neuropodia from 26-29
Morphology: Prostomium well developed, with oviform shield
chaetiger; uncini numerous (20-50 per torus) in single rows,
shape, clearly separated from buccal segment, with anterior me-
with teeth directed anteriorly. Uncini with only a single main
dian incision on two third of the length. Buccal apparatus with
tooth, lacking a secondary tooth. Pygidium rounded with five
many grooved buccal tentacles. Paired sexual tentacles or ven-
conspicuous rounded papillae.
tral organ not observed in the type series. First 25-30 chaetiger-
ous segments with notopodia only. Branchiae four pairs all sim- Biology: In anhydrite close to vent opening and on rocks under
ilar arranged as funnel-like structure, with strong basal stem Alviniconcha beds in active areas. Tubes horny with apical digi-
bearing small secondary filaments and a thin terminal tip de- tations.
void of secondary filaments as long as the basal stem. Secondary
Distribution: North Fiji Back-Arc Basin; Lau Back-Arc Basin:
filaments inserted along stem on two opposite areas; each leaf-
Vaï Lili vent field.
shaped, strongly flattened with median ciliated area and point-
ed tip. Notopodia (from chaetiger 1 to the end of the body - ex-

2: Anterior portion (SEM);


by V. Martin © Ifremer.

1: Habitus; by V. Martin © Ifremer.

References:
DESBRUYÈRES D. & L. LAUBIER (1993) Proc. Biol. Soc. Wash. 106: 225-236.
DESBRUYÈRES D., ALAYSE-DANET A. M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.

D. DESBRUYÈRES Denisia 18 (2006): 289–290


© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

3: in vivo; by courtesy of Greg Rouse.

4: Anterior part of a preserved speci-


men showing the funnel-like structure
of gills. Photo by P. Briand © Ifremer. 5: Tubes of P. unidentata on a sulphide spire © Ifremer.

7: 7th and 8th parapods, left side of the body (SEM)


6: Funnel like structure of gills (SEM) © Ifremer. © Ifremer.

8: Prostomium (SEM) © Ifremer. 9: Uncinigerous torus (SEM) © Ifremer.

290
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Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Paralvinella) fijiensis DESBRUYÈRES & LAUBIER, 1993

bearing dorsal and ventral rounded lobes. Chaetigerous seg-


Size: Up to 25 mm long.
ment 7 bearing three to four stout transformed chaetae.
Color: In vivo reddish and yellow gills. Uncinigerous neuropodial tori present from 12-19 to the end of
the body. Each uncinus with a main tooth surmounted by
Morphology: Body maggot-shaped in large specimens (50-
smaller secondary tooth. Pygidium blunt.
65 cs). Prostomium reduced medially, with two anterior lobes.
Buccal apparatus comprising numerous exertile grooved tenta- Biology: Most specimens were collected on active edifice walls.
cles and two paired sexual tentacles in males ending with three Tubes are cylindrical and isolated.
unequally developed rounded lobes. Notopodia of chaetigerous
Distribution: North Fiji and Lau Back-Arc Basins.
segments 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

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

2: Habitus in vivo; Lau Basin, cruise TUIM06; by courtesy of


F. Pleijel.

3: Notopodial 4: Posterior 5: Sexual trilobed tenta-


chaeta (SEM) uncinigerous tori cles (SEM) © Ifremer.
© Ifremer. (SEM) © Ifremer.

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

D. DESBRUYÈRES Denisia 18 (2006): 291


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Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Paralvinella) grasslei DESBRUYÈRES & LAUBIER, 1982

Size: Up to 80 mm. Biology: Worm dwelling both in warm part of active chimneys
where mineral load is low and on Riftia tubes. Deposit feeder
Color: Brownish when preserved, reddish in vivo.
feeding mainly on bacterial mats.
Morphology: Alvinellid worm with four pairs of pinnate
Distribution: East Pacific Rise, Guaymas Basin, Galapagos
branchiae. One transformed notopod wearing stout acicular
Spreading Center.
spines. Uncinigerous segment from 13-17th chaetigerous seg-
ment to the posterior part. Chaetigerous segments up to 110.
Gonochoric species; males with two robust ventral peribuccal
tentacles ending in three rounded lobes. No bacterial epibiosis.
Mucous tubes unconspicuous.

1: Habitus; by V. Martin  2: Population in situ on a white 3: Anterior part  Ifremer.


Ifremer. smoker (East Pacific Rise: 13°N);
cruise Phare  Ifremer.

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.

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Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Paralvinella) palmiformis DESBRUYÈRES & LAUBIER, 1986

Size: Up to 80 mm. Biology: Most commonly found with its caudal end coiled
around the distal portion of siboglinid tubes or on the surface of
Color: Red or pinkish after preservation in ethanol.
sulfide mineral deposits covered in a sheath of inorganic par-
Morphology: Body gradually tapering. Holotype with 118 seg- ticulates accumulated by mucus secretion. Deposit feeder.
ments, paratypes 100-118. Prostomium medially reduced. Ven-
Distribution: Gorda Ridge, Explorer Ridge, Juan de Fuca
trally on the peristomium of males, two blind cavities. Buccal
Ridge.
apparatus 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. The
first 20-31 chaetigerous segments with notopodia only.
Chaetigerous segment 7 strongly modified.

1: Endeavour segment,
P. palmiformis associated
with Ridgeia piscesae; by
courtesy of S.K. Juniper.
Bottom right: Habitus; by
V. Martin  Ifremer. Top
left: Comparison between
the edges of peribuccal
tentacles (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.

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Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Paralvinella) sulfincola DESBRUYÈRES & LAUBIER, 1993 “sulfide worm“

(excepted 7) bearing a dorsal digitiform lobe. Chaetigerous 7


Size: Up to 80 mm.
strongly modified with 4-5 very large stout notopodial hooks di-
Color: Bright red in life and chocolate brown with brown rected posteriorly. Tubes cylindrical translucent and thin.
chaetae after preservation in formalin.
Biology: Temperatures of 20-80°C have been measured on sur-
Morphology: 54-68 chaetigerous segments. Body regular, taper- faces colonized by sulfide worms and its most likely that the
ing abruptly in terminal 10 segments. Buccal apparatus com- worms regularly experience temperatures within the lower part
prising small grooved and ciliated tentacles, one pair of large of this range. All the collections containing this species are
basal tubercles and in males two large lateral tentacles and ter- coming from active sulfide edifices where the animal occupies
minating in a curved margin. First 24-30 chaetigerous segments a distinct microhabitat.
with notopodia only. Four pairs of branchiae present, all similar
Distribution: Explorer Ridge, Juan de Fuca Ridge, Gorda
strong, abruptly attenuated. Branchial stem relatively short,
Ridge.
with a large number of slender filaments arising in two opposite
ranks and bearing ciliated longitudinal line. First 22 notopodia

1: Anterior part (preserved specimen)


 Ifremer.

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

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.

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Annelida, Polychaeta, Terebellida, Ampharetidae

Amathys lutzi DESBRUYÈRES & LAUBIER, 1996

shape as thoracic ones. Abdominal notopodia absent. Pygidium


Size: 1.25-15.7 mm in length, 2.1-6 mm in width.
terminal without circle of papillae or anal cirri. Mucus lined
Color: No color pattern observed on preserved material. tubes covered with rusty colored mineral particles, mussel pe-
riostracum pieces and fragments of byssal threats; sometimes,
Morphology: 38 chaetigerous segments of which 20 thoracic
tube simple covered with gray mud.
and 18 abdominal; 17 thoracic uncinigerous segments. Pros-
tomium lacking glandular ridge or eyes; no pallae. Buccal ten- Biology: Sorted from mussel washings, or found in tubes at-
tacles smooth, grooved and inserted on a buccal membrane. tached to mussel shell hinges and to sulfide or basaltic rocks.
Four pairs of branchiae, all smooth and similar, regularly atten-
Distribution: Mid-Atlantic Ridge: Lucky-Strike, Broken Spur,
uated. Thoracic parapodia without cirri. Each thoracic uncinus
Snake Pit.
with a single row of four teeth, abdominal uncini with the same

2: Anterior part, lateral view; 3: Tube from Mid-Atlantic Ridge:


by P. Briand  Ifremer. Rainbow; by P. Briand  Ifremer.

1: Habitus; by V. Martin
 Ifremer.

4: Prostomium, frontal view; 5: Thoracic uncini (SEM)


by P. Briand  Ifremer.  Ifremer.

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.

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Annelida, Polychaeta, Terebellida, Ampharetidae, Ampharetinae

Amphisamytha galapagensis ZOTTOLI, 1983

abdominal segment. Tube mucus-lined covered with small


Size: Maximum size in Guaymas individuals 18 mm, smaller
chips of volcanic glass or mud.
elsewhere.
Biology: Ubiquitous, often found in other invertebrate shells or
Color: Whitish or grey when preserved. Pinkish to green in vi-
carapaces. Deposit feeder.
vo.
Distribution: Galapagos Spreading Center, East Pacific Rise,
Morphology: Ampharetinae with four pairs of smooth gills, no
Guaymas Basin, North East Pacific, Lau Back-Arc Basin, North
pallae, first three chaetigerous segments reduced, uncini from
Fiji Back-Arc Basin, Manus Back-Arc Basin, Okinawa Trough,
fourth chaetigerous segment continuing posteriorly on 14 tho-
Mariana Back-Arc Basin. Likely a group of cryptic species (see
racic segments. Uncini avicular with one row of teeth. Buccal
CHEVALDONNÉ et al. 2002).
tentacles smooth, inserted on a “buccal membrane”, 12-15

5: Buccal membrane and tentacles (SEM)  Ifremer.

3: Anterior part, lateral view


(SEM)  Ifremer.
1: Habitus; by V. Martin
 Ifremer.

6: Abdominal segments, lateral view (SEM)  Ifremer.

2: Specimen in vivo 4: Posterior end (SEM)


 Briand/Ifremer.  Ifremer. 7: Thoracic unici (SEM)  Ifremer.

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.

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Annelida, Oligochaeta, Hirudinida, Piscicolidae, Platybdellinae

Bathybdella sawyeri BURRESON, 1981

Size: Maximal 13 mm. Biology: Found free-living in washings of bivalves vesicomyid


Calyptogena magnifica and mytilid Bathymodiolus thermophilus,
Morphology: The morphology of most specimens corresponds
bythograeid crabs Bythograea thermydron and Cyanagraea
with the original described morphology of B. sawyeri, with
praedator, tubeworm Riftia pachyptila, and bythitid fish. Little is
suckers of approximately equal size, but all specimens collected
known on the biology of Bathybdella sawyeri. No specimens
from Cyanagraea praedator have unusually large oral suckers
were collected from fish hosts, but the presence of nucleated
that were distinctly larger than the caudal sucker. However,
red blood cells in the gut suggest that this leech feeds on the
based on the internal morphology using serial sections, also the
blood of fishes, as do all other members of the family Piscicoli-
specimens from C. praedator could be identified as Bathybdella
dae. The high abundance of leeches found among vent inver-
sawyeri. The leech is unpigmented and lacks eyespots and ocel-
tebrates suggests that the leech leaves the fish host after each
li, but often appears red in color because of the blood in the gut.
blood meal and seeks refuge among various invertebrates until
The body is smooth and lacks papillae, tubercles or lateral pul-
digestion is complete and the leech is ready to feed again. Al-
satile vesicles. Internally B. sawyeri is characterized by six pairs
though cocoons (egg cases) have not been found, B. sawyeri un-
of testisacs and a very unusual reproductive system that in-
doubtedly deposits cocoons on available hard substrate includ-
cludes paired lateral invaginations in segment 12 connected via
ing shells or carapaces of invertebrates.
vector tissue to a large bilobed spermatheca.
Distribution: Galapagos Spreading Center; Southern East Pa-
cific Rise: 14°S, 17°S.

1: Habitus,
reconstruc-
tion of diges-
tive system,
reproductive
system, body
shape and
sucker shape
from dorsal
aspect; 3: Leech cocoons of Johanssonia arc-
A – anus; tica on the walking leg of the tan-
C – crop ner crab, Chionoecetes bairdi, off
caecum; the coast of Oregon (USA); scale bar
I – intestine; 6 mm; by E. Burreson © VIMS.
LI – lateral
invagination;
M – mouth-
pore;
P – proboscis;
PC – post-
caeca;
R – rectum;
T – testisac;
from BURRESON
(1981).
4: Leech cocoons of Myzobdella
lugubris on the carapace of the blue
2: Specimen taken onboard, after collection crab, Callinectes sapidus, in Chesa-
from Southern East Pacific Rise: 17°S, Oasis site; peake Bay, Virginia (USA); scale bar
cruise Biospeedo; by S. Hourdez © CNRS. 6 mm; by E. Burreson © VIMS.

References:
BURRESON E.M. (1981) Proc. Biol. Soc. Wash. 94(2): 483-491.
BURRESON E.M. & M. SEGONZAC (submitted) Zootaxa.

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Arthropoda, Chelicerata, Arachnida, Acariformes, Halacaridae

Copidognathus papillatus KRANTZ, 1982

Size: 500-570 µm (from tip of rostrum to end of anal papillae). Biology: Present within detritus among colonies of vestimen-
tiferan tubeworms, barnacles, and mussels.
Morphology: Marine mite with ovate idiosoma, slender
gnathosoma with almost parallel-sided rostrum, two pairs of Distribution: Galapagos Spreading Center, East Pacific Rise:
legs directed forward and two pairs backward. Dorsal plates of 13°N, Juan de Fuca Ridge: Main Endeavour; North Fiji and Lau
idiosoma with areolate-reticulate sculpturing. Anal papillae en- Back-Arc Basins.
larged, extending beyond anal aperture. Legs with aciculate
reticulation; they lack long spine-like setae.

1: Habitus; by I. Bartsch.

References:
BARTSCH I. (1991) Zool. Sci. 8: 789-792.
KRANTZ G.W. (1982) Can. J. Zool. 6: 1728-1731.

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Arthropoda, Chelicerata, Arachnida, Acariformes, Halacaridae

Halacarellus auzendei (BARTSCH, 1990)

Size: 650-700 µm (from tip to rostrum to end of anal papilla). Biology: Several mites have suctorians (Ciliophora) fixed on
the idiosoma, gnathosoma or legs.
Morphology: Marine mite with ovate idiosoma and two pairs of
legs directed forward and two pairs backward. Gnathosoma Distribution: Mid-Atlantic Ridge: Snake Pit and Broken Spur.
about 1/3 as long as the idiosoma; rostrum slender. Dorsal idio-
somatic plates reticulate, membranous integument between
plates 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.

1: Habitus; by I. Bartsch.

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.

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Arthropoda, Pycnogonida, Ammotheidae

Ammothea verenae CHILD, 1987

auxiliary claws. Male cement gland opening a tiny dorsodistal


Synonym: Scipiolus thermophilus TURPAEVA, 1988.
pore. Female oviger strigilis with segments lacking dense setae,
Size: Male leg span about 43 mm. Female slightly larger. with 1-2 denticulate spines on distal segments. Female legs with
far fewer setae, without large fields of setae.
Morphology: Trunk slender with segment posteriors flared into
cowling. Ocular tubercle a low, blind, truncate cone at neck Remarks: This species differs in basic morphology from any
mid length. Lateral processes separated by less than half their other known member of the genus. It is blind, lacks the con-
diameters. Proboscis long, with median and distal constrictions. spicuous dorsomedian tubercles common on the trunk cowls of
Abdomen long, down curved. Chelifores tiny, short, chelae at- most species, lacks differentiated heel spines, and does not have
rophied bumps. Palps nine-segmented, 5 distal segments very differences in propodi shape between the anterior and posteri-
short, heavily setose ventrally. Oviger segment 4 and 5 sub or legs.
equal in length, proximally setose, strigilis segments 6, 7 and 8
Biology: Found commonly in close proximity to hydrothermal
with many long lateral setae, segments 8, 9 and 10 with 1-2
vents and sometimes encrusted with polymetallic sulfides.
small endal denticulate spines. Eggs tiny, about 0.25 diameter of
main oviger segments, carried in large round clusters. Legs long, Distribution: Juan de Fuca Ridge: Axial Seamount, Endeavour
slender, extremely setose. Third coxae with many long ventral Segment; Explorer Ridge.
setae. Distal leg segments alike, tarsus very short, propodus
slender, well curved, sole spines alike. Main claws with long

1: Male holotype. A: Trunk, dorsal view; B: Trunk, lateral 2: In situ view showing numerous specimens and a scale
view; C: Oviger; female paratype: D: Third leg with ova; worm (Branchinotogluma?) © NOAA Ocean Exploration.
from CHILD (1987).

References:
CHILD C.A. (1987) Proc. Biol. Soc. Wash. 100(4): 892-896.
TURPAEVA E.P. (1988) Zool. Zh. 67(6): 950-953.

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Arthropoda, Pycnogonida, Ammotheidae

Sericosura cochleifovea CHILD, 1989

ae and propodus. No thick fields of setae. Tarsus short, propo-


Synonym: Sericosura bifurcata STOCK, 1991.
dus slender, almost straight, claws moderately long. Eggs large,
Size: Small for the genus, male leg span about 13 mm. almost as wide as oviger segment diameters around which they
are wound. Cement gland at extreme proximal end of femur
Morphology: Trunk slightly ovoid, lateral processes separated
with laterally pointing tube slightly longer than segment diam-
by about half their diameters, armed with one or more dor-
eter.
sodistal and laterodistal spines. Trunk segments with swollen
posterior cowls. Ocular tubercle only as tall as its base, blind. Remarks: One of the major differences between this species
Proboscis ovoid, originating on slender stalk, greatly constrict- and the others in this genus is the sexual dimorphism displayed
ed distally. Abdomen long, downcurved, with two pairs of long in leg setae. The very long lateral leg setae of females differen-
dorsal spines, pair of short distal spines. Chelifores typical; tiate them conspicuously from males. The shorter leg setae and
short, with atrophied chelae. Palps nine-segmented, second very long cement gland tube of the males sets them off from fe-
segment longest, five distal segments little longer than wide, males of this species and males of other known species which
heavily setose. Oviger fourth segment longest, fifth slightly have much shorter tubes.
shorter. Strigilis sixth segment with field of long lateral setae,
Biology: Taken in the proximity of a vent site with other fau-
with 1-2 tiny denticulate spines on distal three segments. Legs
na: snails, crabs, shrimps and anemones.
moderately slender, very setose. Male leg major segments with
rows of many setae, some longer than segment diameter. Female Distribution: Only known from the Mariana Islands Back-Arc
leg major segments with few very long femoral setae, lateral Basin hydrothermal vent fields.
rows of extremely long setae and few long dorsal setae on tibi-

1: Holotype male. A: Trunk, dorsal view; B: Trunk, lateral view; C: Palp; D: Third leg, with cement gland tube enlarged; E: Oviger
with several eggs attached; from CHILD (1989).

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.

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Arthropoda, Pycnogonida, Ammotheidae

Sericosura cyrtoma CHILD & SEGONZAC, 1996

segment. Legs with dorsal, ventral and lateral rows of short


Size: Leg span 17 mm.
spines in sockets, tibiae with 1-2 longer dorsal setae. Tarsus very
Morphology: Trunk robust, very broad anteriorly, armed with short, propodus typical, with few sole spines. Claw long, auxil-
many short spines on sockets on trunk anterior, distally on lat- iaries half main claw length. Male features unknown.
eral processes and chelifores, and on palp second segment. Oc-
Remark: The tapered and bent proboscis is a unique character
ular tubercle broad, not as tall as basal width, blind. Proboscis
among the known species of Sericosura.
tapering from broad base, downcurved distally, lips flat. Ab-
domen broad, long, with dorsal spines. Chelifore scapes twice as Biology: Found associated with siboglinid worms in a hy-
long as wide, chelae atrophied to rounded nubs. Palp seven-seg- drothermal vent area.
mented, fourth segment slightly longer than second, fifth
Distribution: East Pacific Rise: 13°N.
longer than sixth and seventh combined, few setae. Oviger (fe-
male) small, plain, strigilis with 1-2 tiny denticulate spines per

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

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

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Arthropoda, Pycnogonida, Ammotheidae

Sericosura heteroscela CHILD & SEGONZAC, 1996

cate conical tube directed anterolaterally. All female legs with


Size: Leg span 26.7 mm.
normal sized propodi and fewer short setae than those of male.
Morphology: Trunk and lateral processes typical. Lateral Neither sex with fields of leg setae.
processes with many short dorso- and latero-distal spines. Ocu-
Remarks: The grossly inflated posterior four propodi of males
lar tubercle slender, twice as long as diameter, blind, bifurcate
serve to set this species off from any other known in the genus.
at tip. Proboscis and abdomen typical. Chelifore scapes hardly
TURPAEVA (1998) allocates this species to a new genus
longer than wide, chelae rounded atrophied nubs. Palps seven-
Anisopes: A. heterocella (sic). Nevertheless, we suggest that we
segmented, only slightly longer than proboscis, moderately se-
need more material to check the probable morphological vari-
tose. Oviger typical, field of long lateral setae on sixth and sev-
ations.
enth segments, distal three segments with tiny denticulate
spines. Male legs of two kinds. Anterior two pairs with normal- Biology: Specimens of this species were found in association
ly proportioned distal segments, posterior two pairs with gross- with hydrothermal vents in several different collecting locali-
ly enlarged and inflated propodi twice diameter and slightly ties.
longer than anterior propodi. Claw of inflated propodi larger
Distribution: Mid-Atlantic Ridge: Snake Pit, Logatchev, Rain-
and longer than normal propodi and auxiliary claws shorter in
bow, Lucky Strike, and Menez Gwen.
relation to main claw. Cement gland of all male legs a small
bulbous dorsolateral inflation on proximal femur with a trun-

3: Specimen collected at Rainbow


site; cruise Atos; note the strong
1: Holotype male. A: Trunk, dorsal view; B: Trunk, lateral view; C: sulphide deposit on the entire
Palp; D: Oviger; E: Third leg; from CHILD & SEGONZAC (1996). body; by P. Briand © Ifremer.

2: Specimen collected at Lucky Strike site 4: Specimen with eggs, and colonized by fila-
(cruise Atos); by P. Briand © Ifremer. 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].

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Arthropoda, Pycnogonida, Ammotheidae

Sericosura mitrata (GORDON, 1944)

eter. Female legs with dorsal, lateral and ventral rows of short
Synonym: Achelia mitrata GORDON, 1944.
sharp spines pointing distally, lateral spines slightly longer.
Morphology: Trunk typical with lateral processes spaced at dis- Some specimens with very long lateral setae. Propodus with
tances of half their diameters, armed with 2-3 short dorsodistal row of many short sole spines.
and laterodistal setae. Ocular tubercle more than twice as long
Remarks: The very long leg setae of Gordon’s female specimen
than its basal diameter, blind, bifurcate at tip with prominent
may be an infestation of something growing on the integument
sensory papillae. Proboscis ovoid with basal and distal constric-
and not setae. One of the easily identified characters in this
tions. Abdomen long with 2-3 short distal setae. Chelifores
genus is the number of palp segments. The genus has species
very short, scapes broad, chelae tiny, atrophied. Palps seven-
with either seven or nine segments with two of the above hav-
segmented, segments 2 and 4 subequal in length, seventh seg-
ing seven and two having nine.
ment as long as fifth and sixth coined lengths, moderately se-
tose. Oviger second segment longest, fifth slightly longer than Biology: This species was described before the discovery of hy-
fourth. Sixth segment with field of lateral setae longer than seg- drothermal vent areas. It is an opportunistic species in the At-
ment diameter. Strigilis segments 8, 9, 10 with pairs of tiny den- lantic vent communities where it has been collected since.
ticulate spines. Male legs dimorphic in setae lengths and
Distribution: Eastern margin of Antarctica at 219 m, in the
arrangement. Tibiae with sparse setae longer than segment di-
Ross Sea at 106 m, on the Walvis Ridge off South Africa at
ameters, with 1-2 ventral spines. Propodus slender, slightly
2100 m; Mid-Atlantic Ridge: Snake Pit.
curved, 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-

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

References:
CHILD C.A. (1982) Smithon. Contrib. Zool. 349: 19-21, Fig. 6.
FRY W.G. & J.W. HEDGPETH (1969) Mem. New. Zeal. Oceano. Inst. 49: 112-113.
GORDON I. (1944) Brit., Austr., New Zeal. Ant. Res. Exped. ser. B 5(1): 54-57.
SEGONZAC M. (1992) C. R. Acad. Sci., Paris 314(III): 593-600.

R. BAMBER Denisia 18 (2006): 305


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Arthropoda, Pycnogonida, Ammotheidae

Sericosura venticola CHILD, 1987

Size: Male leg span about 28 mm. Female size unknown. Remarks: The main difference between specimens of this
genus and those in the genus Ammothea is the placement of the
Morphology: Trunk moderately slender, first three segments
male cement gland and its orifice. They are otherwise difficult
with flaring posterior cowls. Lateral processes separated by less
to separate.
than half their diameters, armed with 3-6 lateral spines. Ocular
tubercle taller than basal diameter, blind. Proboscis massive, Biology: The only known specimens were found associated
with proximal and distal constrictions. Abdomen long, down- with siboglinids at a hydrothermal vent on the Endeavour Seg-
curved. Chelifores small, short, chelae atrophied to nubs. Palps ment.
seven-segmented, segments 2 and 4 subequal in length, distal
Distribution: NE-Pacific: Endeavour Segment, Juan de Fuca
three short segments heavily setose. Oviger second segment
Ridge; a juvenile belonging probably to the same species was
longest, fourth and fifth subequal in length. Strigilis segments
collected at Southern East Pacific Rise: 17°S, Rehu-Marka site.
6, 7 and 8 with many long lateral setae, segments 8, 9, 10 with
1-2 small denticulate spines. Eggs very tiny, about one fifth as
wide 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, propodus
slender, well curved, claws of moderate length. Male cement
gland a swelling at proximal end of femur, with a short tube
pointing laterally. Female characters unknown.

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).

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

R. BAMBER Denisia 18 (2006): 306


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Arthropoda, Crustacea, Ostracoda, Myodocopa, Halocyprida, Halocypridae

Archiconchoecia (Archiconchoecia) chavturi KORNICKER & HARRISON-NELSON, 2005

bend at tip of first antenna, with rounded tip bearing two


Size: Adult female carapace length 0.60 mm, height 0.40 mm.
minute spines.
Morphology of adult female (male unknown): Carapace: oval
Biology: Unique female with 15 eggs in marsupium. Gut with
in lateral view with greatest height just posterior to midlength;
appendage fragments, bristles, and bristle-like claws indicating
anterior rostrum and i