© 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
Katalog / Publication:
Erscheinungsdatum / Date of delivery:
Medieninhaber und Herausgeber / Copyright:
URL / Internet:
E-Mail:
Wissenschaftliche Redaktion / Scientific editors: Daniel Desbruyeres, Michel Segonzac & Monika Bright
Redaktionelle Betreuung / Editorial assistance:
Layout, Druckorganisation /
Layout, printing organisation:
Umschlag-, Plakatgestaltung / Cover, placard:
Druck / Printing:
Bestellung / Ordering:
Umschlagfoto / Cover:
Ausstellung / Exhibition:
Ort / Address:
Zeitraum / Period:
Konzept, Organisation und Gestaltung /
Concept, organization, design:
Ausstellungstechnik, Mitarbeit /
Exhibition techniques, collaboration:
Museumspädagogik / Museum education:
Leihgeber / Lenders:
Impressum
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
6. April 2006
Land Oberösterrreich, Biologiezentrum der Oberösterreichische Landesmuseen, J.-W.-Klein-Str. 73, 4040
Linz, Austria
http://www.biologiezentrum.at
bio-linz@landesmuseum.at
Dr. Erna Aescht
Eva Rührnößl
Eva Rührnößl
Plöchl-Druck, Werndlstraße 2, 4240 Freistadt
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.
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.
Heiß und giftig – Oasen der Tiefsee / Hot and toxic – Oases in the deep sea
Biologiezentrum der Oberösterreichischen Landesmuseen, J.-W.-Klein-Str. 73, 4040 Linz, Austria
7. April bis 1. Oktober 2006
Mag. Stephan Weigl, Dr. Monika Bright, Johannes Rauch
Jürgen Plass, Roland Rupp, Franz Meindl, Bruno Tumfart, Erwin Kapl, Josef Schmidt, Roland Zarre
Mag. Sandra Kotschwar
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-
book of Deep-Sea Hydrothermal Vent Fauna“ gives on over-
view of our current knowledge on the animals living at hydro-
thermal vents. The discovery of hydrothermal vents and pro-
gresses made during almost 30 years are outlined. A brief intro-
duction is given on hydrothermal vent meiofauna and parasites.
Geographic maps and a table of mid-ocean ridges and back-arc
basins with the major known hydrothermal vent fields, their lo-
cation and depth range and the most prominent vent sites are
provided. Higher taxa are presented individually with informa-
tion on the current taxonomic and biogeographic status, the
number of species described, recommendations for fixation, and
Résumé
La seconde édition, considérablement étendue et améliorée
du « Handbook of Deep-Sea Hydrothermal Vent Fauna » don-
ne un panorama de notre connaissance sur les animaux qui vi-
vent autour des sources hydrothermales océaniques. Les avan-
cées faites depuis leurs découvertes il y a trente ans environ,
donnent lieu à une mise en perspective rédigée par des acteurs
des campagnes océanographiques. Une brève introduction est
proposée sur la méiofaune et les parasites, puis chaque taxon su-
périeur est présenté avec des rappels sur la taxinomie, des in-
formations sur sa distribution géographique, des représentations
schématiques de la morphologie permettant à des non-spécia-
listes de comprendre la nomenclature, des informations sur le
nombre d’espèces présentes, des conseils pour la fixation et la
conservation des échantillons. Des cartes de distribution des
Zusammenfassung
Die zweite, wesentlich erweiterte Auflage des „Handbook
of Deep-Sea Hydrothermal Vent Fauna“ gibt einen Überblick
über den momentanen Stand der Forschung. Die Entdeckung
der Hydrothermalquellen, Fortschritte der letzten beinahe 30
Jahre, sowie Meiofauna und Parasiten der Hydrothermalquellen
werden kurz umrissen. Alle ozeanischer Rücken und Back-Arc
Basins mit ihren bekannten Hydrothermalquellfeldern, ihre ge-
naue Lage und Tiefe, sowie ihre benannten Fundstellen werden
in geographischen Karten und einer Tabelle gezeigt. Höhere
Taxa werden individuell dargestellt mit Information über den
momentanen taxonomischen und biogeographischen Stand,
Key words: Hydrothermal vents, deep sea, mid-ocean ridge, back-arc basin, hydrothermal vent fauna.
schematic drawings, which aim to help non-specialists to iden-
tify the animals. 86 authors contributed with their expertise to
create a comprehensive database on animals living at hydro-
thermal vents, which contains information on the morphology,
biology, and geographic distribution of more than 500 current-
ly described species belonging to one protist and 12 animal
phyla. It comprises also the largest collection of more than
1000 pictures of hydrothermal vent animals taken in situ with
submersibles, in vivo after collection, and with various dissec-
tion, light, and scanning electron microscopes after fixation
and preparations.
champs hydrothermaux le long des rides océaniques et dans les
bassins arrière-arc sont proposées assorties d’un tableau qui
donne leur position géographique, leur profondeur et les sur-
noms des principaux sites. 86 auteurs ont contribué à la créa-
tion d’une base de données sur les animaux présents dans cet
écosystème surprenant, en donnant des informations sur la
morphologie, la biologie et la distribution géographique de plus
de 500 espèces appartenant à 12 phylums animaux. Cette base
de données, présentée sous forme de fiches individuelles, re-
groupe une série unique de plus de 1000 illustrations compor-
tant dessins au trait, photographies des organismes dans leur
milieu, vues d’animaux isolés après récoltes et micro-photogra-
phies prises au microscope électronique à balayage.
die Anzahl der beschriebenen Arten, Methoden zur Fixierung
und schematischen Zeichnungen, die es dem Nicht-Spezialis-
ten die Identifizierung erleichtern sollen. Über 80 Autoren ga-
ben ihren Beitrag zur Erstellung einer umfassenden Datenbank
mit mehr als 500 Tieren aus einem Protisten-Stamm und 12
Tierstämmen und Informationen zu deren Morphologie, Biolo-
gie und Verbreitung. Darüber hinaus werden die Tiere der
Hydrothermalquellen in mehr als 1000 Bildern, Unterwasser-
aufnahmen von bemannten und unbemannten U-Booten, in
vivo Aufnahmen und diversen licht- und elektronenmikrosko-
pischen Aufnahmen, dargestellt.
5
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Foreword
Deep-sea hydrothermal vents and their associated fauna
were discovered in 1977. The presence of these rich oases of li-
fe in the food-limited environment of basaltic Mid-Oceanic
Ridges fostered the interest of the marine biologists to explore
the deep sea using submersibles with a focus on fragmented ha-
bitats. During the intervening three decades, a large number of
cruises using deep submergence vehicles (HOV or ROV) were
devoted to the biological study of the hydrothermal communi-
ties on the Pacific, Atlantic, and Indian Ridges as well as in
Back-Arc Basins and Volcanic Arcs of the Pacific Rim of Fire.
These series of cruises were mainly devoted to the understan-
ding (1) the distribution, the phylogeny, and the dispersal of
this specialized fauna which is restricted to this fragmented “ex-
treme” habitat along the world’s oceanic ridges and (2) to the
niche characterization and the study of the physiological adap-
tations of the organisms (or symbiotic associations) living in
the mixing zone between the superheated fluid and the cold
deep-sea water. This endeavour remains unique in its extent
and intensity in the history of biological exploration of the
world’s oceans since the early studies of the deep-sea by the pio-
neering expeditions of the nineteenth century. However, be-
cause of logistic constrains, this accomplishment is still far from
providing an exhaustive overview of the composition and dis-
tribution of hydrothermal vent fauna; some Ridges (South At-
lantic and the three Indian Ridges) still deserve obviously mo-
re attention and the study of remote targets as high latitude rid-
ges (Arctic and Antarctic Seas) or isolated basins (South Sand-
wich and Andaman Sea) would bring valuable information on
the biogeography of the vent fauna. Multiyear surveys on the
East Pacific Rise and the Juan de Fuca Ridge have also demon-
strated that the hydrothermal environment is submitted to
temporal instability due to volcanic and tectonic events; this
instability greatly affects the patterns of the ecosystem deve-
lopment and produces a variation in specific composition of
communities only detected by temporal series of sampling.
Reliable morphological identification of collected speci-
mens presently remains the inescapable base of most of the bi-
ological work conducted at vents on animals, including the mo-
lecular approach. Because of the taxonomic novelty of deep-sea
hydrothermal vent communities, an impressive descriptive
6
« 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.“
work has been conducted for three decades producing more
than 500 fully identified species of which descriptions or
records are dispersed throughout the scientific literature. Nev-
ertheless the list of animal species living at vents is still evolv-
ing every week; new species are described from additional sam-
ples in known locations and from new locations, descriptions
are revisited, sibling species are detected based on molecular
taxonomy, and new combinations are proposed based on stud-
ies of large series and molecular information. The technology of
imaging in situ greatly improved in the last few years giving
considerable information on the shape, colors, behavior, and
micro-distribution of living animals. The dialogue between
classical and molecular taxonomy brought new insights and
clarifies the biogeographic patterns of vent communities as well
as their relationships with other deep-sea reduced habitats
(cold seeps, food falls, and minimum oxygen zones).
In the foreword of the first edition of the “Handbook of
Deep-Sea Hydrothermal Vent Fauna” (Ifremer Editions, 1997),
we proposed to publish periodic addenda to the first edition
based on new published observations and we invited contribu-
tors to help us by sending information and illustration. The re-
sults far exceeded our hopes and we realized soon that a new is-
sue of the handbook would be fully justified. At the same time,
the Biologiezentrum der Oberösterreichischen Landesmuseen
(Biology Center of the Upper Austrian Museums) was prepar-
ing an exhibition on deep-sea hydrothermal vent life planned
for April 2006 and was looking for authors willing to publish a
review on vent biology. It was the perfect opportunity for us to
restart the collection of contributions from the worldwide net-
work of taxonomists involved in the study of hydrothermal
vent samples. We want to thank here all the contributors who
participated in the wording of this new issue.
This book, published under the auspices of InterRidge and
Census of Marine Life/ChEss, is an overview of the present tax-
onomic knowledge of the deep-sea hydrothermal vent fauna,
intended for scientists and submersible pilots working at sea,
but also for students and the general public curious about the
life in the ocean.
D. Desbruyères, M. Segonzac & M. Bright
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Remarks
Collection of animals as well as in situ photographs and vi-
deos of animals from a deep-sea environment are mainly achie-
ved through the use of manned and unmanned submersibles.
Valuable sampled animals are often a by-product of large bulk
samples, fixation and storage sometimes are less than optimal
for all taxa, and the time for taking pictures while on board is
often scarce. Also, it is often due to pure luck that in situ
photographs and videos are taken at the decision of chief scien-
tists, pilots, and researchers on board as they must interrupt
their ongoing tasks and use valuable bottom time to document
animals in situ. Good quality pictures taken on the bottom pro-
vide a unique set of information on the micro-distribution, be-
havior, and biological interactions. Even those photographs,
which are in low resolution or out of focus, are highly valuable
because they give us an impression of where these animals oc-
cur and how they behave naturally.
We stress our belief that our successes in understanding the
fauna living at hydrothermal vents will be highly influenced by
the way we approach solving these issues. So we recommend for
taxonomy, ecology and outreach purposes, that researchers ta-
ke the time on the bottom to video and photograph, that scien-
tists onboard research vessels take high quality photographs of
animals before preservation as well as follow instructions for
proper fixation and storage, and that they get into contact with
taxonomists for identification and appropriate procedures for
voucher specimens. Moreover, we hope that more taxonomists
will be invited to research cruises to better support our goal of
learning more about hydrothermal vents and their inhabitants.
Recent experiences demonstrated that the presence of taxono-
mists on cruises improved by far our knowledge of the faunal
composition of vent communities and fostered interactions
with other disciplines.
D. Desbruyères, M. Segonzac & M. Bright
7
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Acknowledgements
We would like to thank all the contributors whose know-
ledge of the fauna and taxonomic expertise allowed us to revi-
se and supplement the second edition of this book. We are gra-
teful to Gerhard Aubrecht, the head/director of the Biologie-
zentrum der Oberösterreichischen Landesmuseen (Biology
Center of the Upper Austrian Museums), for making this pu-
blication possible and for his continuous support throughout
the entire process. Special thanks to Erna Aescht (Biology
Center) for the editing, Eva Rührnößl (Linz) for the page-ma-
king, Heidemarie Grillitsch (University Vienna) for the sche-
matic drawings, Ingrid Kolar (University Vienna) for checking
the numerous references, and Patrick Briand (Ifremer) for his
help with the pictures, Violaine Martin (Ifremer) for many dra-
wings, and Philippe Crassous (Ifremer) for his technical exper-
tise with the SEM preparations.
8
We express our heartful thanks to the chief scientists Fern-
ando Barriga, Nadine Le Bris, Charles R. Fisher, Francoise
Gaill, Anne Godfroy, Jun Hashimoto, S. Kim Juniper, François
Lallier, Marvin D. Lilley, Pierre-Marie Sarradin, Tim Shank,
Verena Tunnicliffe, Karen Von Damm, Cindy L. Van Dover
and Robert C. Vriejenhoek, who allowed us to have access to
samples, data, and in situ pictures. We are also grateful to the
many contributors who shared their pictures with us.
This book was supported by the Federal Government of
Upper Austria, the Biology Center of the Upper Austrian Mu-
seums, Ifremer, ChEss (especially the coordinators of the ChEss
project Eva Ramirez-Llodra and Maria Baker), InterRidge, the
Austrian Science Foundation, and the Faculty of Life Sciences,
University of Vienna.
D. Desbruyères, M. Segonzac & M. Bright
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Photo credits
Auger, V. – Canadian Scientific Submersible Facility, Canada
Biscoito, M. – Museu Municipal do Funchal, Portugal
Blake, J. – ENSR Marine & Coastal Center, USA
Blaston, P. – University of Otago, New Zealand
Briand, P. – Ifremer, Brest, France
Bright, M. – University of Vienna, Austria
Burreson, E. – College of William and Mary, Williamsburg,
USA
Chan, T. Y. – National Taïwan Ocean University, Taïwan
Cosel, R. von – Muséum National d’Histoire Naturelle, Paris,
France
Cunha, M.R. – Centro das Zonas Costeiras e do Mar, Universi-
dad, Portugal
Fautin, D. – Natural History Museum and Biodiversity Rese-
arch Center, Kansas, USA
Dejouannet, J.-F. – IRD (Institut Recherche Développement)
Paris, France
Drukker-Brammall, P. – Smith Institute of Ichthyology, Gra-
hamstown, South Africa
Dugornay O. – Ifremer, Brest, France
Encarnação, H. – Museu Municipal do Funchal, Portugal
Fifis, A. – Ifremer, Brest, France
Fisher, C. R. – The Pennsylvania State University, USA
Glover, A. – The Pennsylvania State University, USA
Guinot, D. – Muséum National d’Histoire Naturelle, Paris,
France
Harrison-Nelson, E. – National Museum of Natural History,
Smithsonian Institution, USA
Hashimoto, J. – JAMSTEC, Nagasaki, Japan
Heemstra, E. – Smith Institute of Ichthyology, Grahamstown,
South Africa
Hessler, R.R. – Scripps Oceanographic Institution, USA
Hourdez, S. – Station Biologique Roscoff, France
Ivanenko, V. N. – Moscow State University, Russia
Juniper, K. S. – Université du Québec, Montréal, Canada
Klitz, K. – National Museum of Natural History, Smithsonian
Institution, USA
Kornicker, L.S. – National Museum of Natural History, Smith-
sonian Institution, 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
Lozouet, P. – National Muséum d’Histoire Naturelle, Paris,
France
Lutz, R. A. – Rutgers University, USA
Maddocks, R. M. – University of Houston, USA
Maestrati, P. – Muséum National d’Histoire Naturelle, Paris,
France
Marcus, J. – University of Victoria, Canada
Martin, V. – Ifremer, Brest, France
McLean, J. – Museum of Natural History, Los Angeles, USA
Mills, S.W. – Woods Hole Oceanographic Institution, USA
Miura, T. – JAMSTEC, Nagasaki, Japan
Mullineaux, L. – Woods Hole Oceanographic Institution, USA
Ng, N. K. – National University of Singapore, Republic of Sin-
gapore
Okata, Y. – JAMSTEC, Nagasaki, Japan
Pleijel, F. – University of Göteborg, Sweden
Rouse, G. – University of Adelaide, Australia
Roux, M. – Université de Reims, France
Sagalevitch, A.M. – Shirshov Institute of Oceanology, Mos-
cow, Russia
Saldanha, L. † – LMG, Lisbonne, Portugal
Shagin, A. – PMGE (Mining Intitute, St Petersburg), Russia
Smith Jr, K. – Scripps Institution of Oceanography, USA
Southward, E.C. – Plymouth Marine Laboratory, United King-
dom
Stöhr, S. – Swedish Museum of Natural History, Sweden
Takeda, M. – JAMSTEC, Japan
Tivey, M. – Woods Hole Oceanographic Institution, USA
Tsuchida, S. – JAMSTEC, Japan
Tunnicliffe, V. – University of Victoria, Canada
Van Dover, C.L. – College of William and Mary, Williamsburg,
USA
Verschelde, D. – Universiteit Gent, Belgium
Vrijenhoek, R. – Monterey Bay Aquarium Research Institute,
USA
Warén, A. – Swedish Museum of Natural History, Sweden
Wilson, G.D.F. – Australian Museum, Syndney, Austrialia
Young, C.M. – Origon Institute of Marine Biology, USA
Zierenberg, R. – University of California, Davis, USA
9
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Contributors and addresses
Almeida, Armando J. – Laboratório Marítimo da Guia, IMAR,
Faculdade de Ciências da Universidade de Lisboa, Estrada
do Guincho, 2750-642 Cascais, Portugal aalmeida@fc.ul.pt
Baba, Keiji – Kumamoto University, Faculty of Education, 2-40-
1 Kurokami, Kumamoto 860-8555, Japan
keiji5962@yahoo.co.jp
Bamber, Roger – Environment: Coastal & Marine, The Natural
History Museum, Cromwell Road, London SW7 5BD, Uni-
ted Kingdom r.bamber@nhm.ac.uk
Barthélémy, Roxane – Université de Provence, Biologie Ani-
male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé-
dex 3, France bioplank@newsup.univ-mrs.fr
Bartsch, Ilse – Biologische Anstalt Helgoland, Zentrale Ham-
burg, Notkestrae 31, D-22607 Hamburg, Germany
bartsch@meeresforschung.de
Bellan-Santini, Denise – Université d’Aix-Marseille 2, Station
Marine d’Endoume-Luminy, Rue de la Batterie des Lions,
13007 Marseille, France bellan@com.univ-mrs.fr
Biscoito, Manuel – Museu Municipal do Funchal, Rua da Mou-
raria, 31, 9000 Funchal, Madeira, Portugal manuel.biscoi-
to@mail.cm-funchal.pt
Blake, James A. – ENSR Marine & Coastal Center, 89 Water
Street, Woods Hole, MA 02543, USA jablake@ix.net-
com.com
Böggemann, Markus – Habsburgerallee 37, 60385 Frankfurt am
Main, Germany mboeggem@aol.com
Bouchet, Philippe – Département Systématique et Évolution,
Muséum National d’Histoire Naturelle, Taxonomie-Collec-
tion Mollusques, 55 rue Buffon, 75231 Paris, France pbou-
chet@mnhn.fr
Boury-Esnault, Nicole – Université d’Aix-Marseille 2, Centre
d’Océanologie de Marseille, Station Marine d’Endoume -
Luminy, Rue de la Batterie des Lions, 13007 Marseille, Fran-
ce esnault@com.univ-mrs.fr
Briand, Patrick – Département «Etude des écosystèmes pro-
fonds», Ifremer, Centre de Brest, BP 70, 29280 Plouzané,
France patrick.briand@ifremer.fr
Bright, Monika – Department of Marine Biology, University of
Vienna, Althanstr. 14, A-1090 Vienna, Austria
monika.bright@univie.ac.at
Buron, de Isaure – Division of Natural Sciences and Enginee-
ring, University of South Carolina Spartanburg, Spartan-
burg, SC 29303, USA deburoni@cofc.edu
10
Burreson, Eugene M. – Department of Environmental and
Aquatic Animal Health, Virginia Institute of Marine Scien-
ce, College of William and Mary, PO. Box 1346, Gloucester
Point, VA 23062 USA gene@vims.edu
Casanova, Bernadette – Université de Provence, Biologie Ani-
male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé-
dex 3, France bioplank@newsup.univ-mrs.fr
Casanova, Jean-Paul – Université de Provence, Biologie Ani-
male (Plancton), 3, place Victor Hugo, 13331 Marseille Cé-
dex 3, France bioplank@newsup.univ-mrs.fr
Causse, Romain – Muséum National d’Histoire Naturelle, La-
boratoire d’Ichthyologie, 43, rue Cuvier, 75231 Paris cedex
05, France causse@mnhn.fr
Cohen, Daniel M. – California Academy of Sciences, Golden
Gate Park, San Francisco, CA 95521, USA dmco3@com-
cast.net
Corbera, Jordi – Carrer Gran, 90, 08310 Argentona, Spain cor-
bera@sct.ictnet.es
Cosel, von Rudo – Département Systématique et Évolution,
Muséum National d’Histoire Naturelle, Taxonomie-Collec-
tion Mollusques, 55 rue Buffon, 75231 Paris, France co-
sel@cimrs1.mnhn.fr
Cunha, Marina R. – Centro das Zonas Costeiras e do Mar, Uni-
versidad, Departamento de Biologia, 3810-193 Aveiro, Por-
tugal mcunha@bio.ua.pt
Daly, Marymegan – Dept. Evolution, Ecology and Organismal
Biology, Ohio State University, 315 Kinnear RD, Columbus
OH 43212, USA daly.66@osu.edu
Defaye, Danielle – Département Milieux et Peuplements Aqua-
tiques, Muséum National d’Histoire Naturelle, 61, rue de
Buffon, 75005 Paris, France ddefaye@cimrs1.mnhn.fr
Desbruyères, Daniel – Département «Etude des écosystèmes pro-
fonds», Ifremer, Centre de Brest, BP 70, 29280 Plouzané,
France Daniel.Desbruyeres@ifremer.fr
Gebruk, Andrey V. – Laboratory of Ocean Benthic Fauna, P.P.
Shirshov Institute of Oceanology, Nakhimovsky Pr., 36,
Moscow 117997, Russia agebruk@sio.rssi.ru
Geistdoerfer Patrick – Laboratoire d’Ichthyologie, 43, rue Cu-
vier, 75231 Paris, France Patrick.Geistdoerfer@
cimrs1.mnhn.fr
Gonzalez, Angel – Ecología y Biodiversidad Marina (ECOBIO-
MAR), Instituto de Investigaciones Marinas (CSIC), C/
Eduardo Cabello 6, 36208 Vigo, Spain afg@iim.csic.es
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Guerra, Angel – Ecología y Biodiversidad Marina (ECOBIO-
MAR), Instituto de Investigaciones Marinas (CSIC), C/
Eduardo Cabello 6, 36208 Vigo, Spain brc1@iim.csic.es
Guinot, Danièle – Département Milieux et Peuplements Aqua-
tiques, Muséum National d’Histoire Naturelle, 61, rue de
Buffon, 75005 Paris, France guinot@mnhn.fr
Haney, Todd – Department of Organismic Biology, Ecology, and
Evolution, University of California Los Angeles, 621 Char-
les E. Young Drive South, Box 1606, Los Angeles, Califor-
nia 90095-1606, USA haney@ucla.edu
Harrison-Nelson, Elisabeth – Department of Invertebrate Zoo-
logy, National Museum of Natural History, Smithsonian In-
stitution, MRC 163, Washington, D. C. 20013-7012, USA
nelsone@si.edu
Hashimoto, Jun – Japan Marine Science and Technology Cen-
ter (JAMSTEC) 2-15, Natsushima-Cho, Yokosuka, 237, Ja-
pan junh@net.nagasaki-u.ac.jp
Holzmann, Maria – Department of Paleontology, University of
Vienna, Althanstr. 14, A-1090 Vienna, Austria maria.holz-
mann@univie.ac.at
Hourdez, Stephan – Station Biologique, 29680 Roscoff, France
shourdez@sj2.seanet.in
Hurtado, Luis – University of Arizona, 310 Biosciences West,
P.O. Box 210088, Tucson, AZ, 85721-0088, USA lhurta-
do@u.arizona.edu
Ivanenko, Viatcheslav N. – Department of Invertebrate Zoolo-
gy, Biological Faculty, Moscow State University, Moscow
119899, Russia ivanenko.slava@mail.ru
Jaume, Damian – IMEDEA (CSIC-UIB), Instituto Mediterra-
neo de Estudios Avanzados, C/ Miquel Marques 21, 07190-
Esporles (Mallorca, Illes Balears), Spain d.jaume@uib.es
Jones, Diana S. – Western Australian Museum, Locked Bag 49.
WELSHPOOL DC, Western Australia 6986, Australia dia-
na.jones@museum.wa.gov.au
Justine, Jean-Lou – Institut Développement Recherche (IRD),
Nouméa, Nouvelle-Calédonie, France jean-
lou.justine@noumea.ird.nc
Komai, Tomoyuki – Natural History Museum and Institute Chi-
ba, 955-2, Aoba-cho, Chuo-ku, Chiba 260-8682, Japan ko-
mai@chiba-muse.or.jp
Kornicker, Louis S. – Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smithsonian Institution,
MRC 163, Washington, D. C. 20013-7012, USA kor-
nickl@si.edu
Kudenov, Jerry D. – Department Biological Sciences, Universi-
ty Alaska Anchorage, 3211 Providence Drive, Anchorage,
Alaska 99508, USA afjdk@uaa.alaska.edu
Larsen, Kim – Department of Invertebrates, Zoological Mu-
seum, Universitetsparken 15, 2100 KBH, Denmark ta-
naids@hotmail.com
Lemaitre, Rafael – Smithsonian Institution, Department of Sys-
tematic Biology, NMHN, MRC 163, PO Box 37012, Was-
hington, DC 20013-7012, USA lemaitre.rafael@
nmnh.si.edu
Lopez-Gonzalez, Pablo – Departamento de Fisiologia y Biologia
Animal, Facultad de Biologia, Universidad de Sevilla, Rei-
na Mercedes, 6, 41012 Sevilla , Spain pjlopez@us.es
Lutz, Richard A. – Center for Deep-Sea Biology & Biotechno-
logy, Rutgers University, Dudley Road, New Brunswick, NJ
08903-0231, USA rlutz@imcs.rutgers.edu
Macpherson, Enrique – CSIC/CEAB, Camí Santa Bàrbara, s/n,
17300 Blanes, Girona, Spain macpherson@ceab.csic.es
Maddocks, Rosalie F. – Department of Geosciences, Room 312
Science & Research Bldg. 1, University of Houston, Hous-
ton, TX 77204-5007, USA RMaddocks@uh.edu
Mah, Christopher – Department. of Invertebrate Zoology, Na-
tional Museum of Natural History, MRC-163, PO Box
37012, Smithsonian Institution, Washington DC 20013,
USA brisinga@gmail.com
Miura, Tomoyuki – Faculty of Fisheries, Kagoshima University,
4-50-20, Shimoarata, Kagoshima, 890 Japan
miura@ugs.agri.kagoshima-u.ac.jp
Möller, Peter R. – Zoological Museum, University of Copenha-
gen Universitetsparken 15, 2100 KBH, Denmark PDRMol-
ler@zmuc.ku.dk
Moreau, Xavier – Université de Provence, Biologie Animale
(Plancton), 3, place Victor Hugo, 13331 Marseille, Cédex 3,
France bioplank@newsup.univ-mrs.fr
Myers, Alan A. – Department of Zoology, Ecology and Plant
Science, National University of Ireland, Cork, Republic of
Ireland alanmyers@crustacea.net
Newman, William A. – Scripps Institution of Oceanography, La
Jolla, CA 92093-0202, USA wnewman@ucsd.edu
Ng, Ngan Kee – Department of Biological Sciences, National
University of Singapore, 14, Science Drive 4, Singapore
117543, Republic of Singapore ngankee@nus.edu.sg
Nielsen, Jorgen G. – Department of Invertebrates, Zoological
Museum, Universitetsparken 15, 2100 KBH, Denmark
JGNielsen@zmuc.ku.dk
Okutani, Takashi – Tokyo University of Fisheries, Konan 4-5-7,
Minato-ku, Tokyo 108, Japan okutani@jamstec.go.jp
11
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

12
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Milestones in the discovery of hydrothermal-vent faunas
Seafloor Spreading and Hot Springs
Any account of the discovery of hydrothermal-vent faunas
must begin with the geology of seafloor spreading centers. The
symmetry of magnetic anomalies on either side of the mid-ocean
ridges that girdle the globe and the correspondence of the anom-
aly patterns with the pattern of magnetic reversals on Earth con-
firmed the process of seafloor spreading and led to general ac-
ceptance of plate tectonic theory in the early 1960s (VINE &
MATTHEWS 1963). The bathymetric relief of mid-ocean ridges
was understood to be a consequence of the thermal buoyancy of
hot rock in volcanic systems. Conductive heat loss was expect-
ed to be greatest at the axis of these linear volcanoes and to di-
minish along transects away from the ridge crest, but heat-flow
measurements collected by placing a vertical array of thermistors
into seafloor sediments consistently documented a heat-deficit
near the ridge axis. This heat deficit suggested that conductive
heat loss was not the only operative mode of cooling (STEIN et
al. 1995). Geologists hypothesized that the convective heat
driven by cells of seawater percolating into the crust and subse-
quent heating and buoyancy-driven flux of fluids out of the crust
would account for the missing heat. Thus the presence of hot
springs on the seafloor with temperatures as great as 300°C was
anticipated (TALWANI et al. 1971; LISTER 1972; WOLERY & SLEEP
1976), and the first unequivocal evidence of warm-water, buoy-
ant plumes was collected by May 1976 using the Scripps Institu-
tion of Oceanography Deep-Tow vehicle (WEISS et al. 1977).
While systematic geophysical studies predicted the existence of
hydrothermal vents on the seafloor, the discovery of their atten-
dant chemosynthetic ecosystems was unimagined.
Strange Animals at Hydrothermal Vents on
the Galapagos Spreading Center
The study of chemosynthetic ecosystems in the deep sea
dates back to a Black and white photograph of large white clam
shells lying within cracks in a pavement of Black basalt that was
captured by the Deep-Tow camera system at 13:20:39 (GMT) on
29 May 1976 at 0°47‘84‘‘N, 86°09‘18‘‘W (Galapagos Spreading
Center, eastern Pacific Ocean; LONSDALE 1977). Given that bi-
valves are generally suspension feeders, the diet of the clams was
inferred to have been organic particulates concentrated by bot-
tom-water currents (convection cells) induced by hydrothermal
activity (LONSDALE 1977; ENRIGHT et al. 1981).
C.L. VAN DOVER, M. BISCOITO, A. GEBRUK, J. HASHIMOTO, V. TUNNICLIFFE,
P. TYLER & D. DESBRUYÈRES
During Alvin dives at the Galapagos Spreading Center in
1977, geologist Jack Corliss first described vent mussels (al-
though he erred in his initial identification). His difficulty in
finding the words to describe what was before him, a vista no
one had ever seen before, is evident:
“They are abalone shells. They are shells. They are big
shells. They are living. … attached shellfish….”
The wonder in his voice is captured on the audio record of
the dive.
Further into the dive series, geochemist John Edmond first
sounds incredulous as he provided the first description of giant
worms, and then frustrated as his observations were limited by
the green light of the thallium iodide bulb on Alvin:
“There are big ones [worms] out there. Looks like an Indian
[Native American] headdress. There are four of them in a row,
right outside my view port. I wonder what color they are. You
can’t tell. They may be red.”
Vent animals were useful as flux indicators of hydrothermal
activity for geologists prospecting for vents, but geologists also
appreciated the need for an explanation of the tremendous bio-
mass of animals and of the means by which species could be
maintained at vents in the face of inevitable local extinctions.
Following the first Alvin dive series to Galapagos vents, micro-
biologist John Baross postulated that the millimolar concentra-
tions of hydrogen sulfide in vent fluids were a source of reduced
sulfur for free-living chemolithoautotrophic, sulfur-oxidizing
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
ATP is then used to fix inorganic carbon (CO2) into organic
carbon. This contrasts with photosynthetic processes, where it
is light energy that is harvested to produce ATP. Subsequent
biochemical steps in the production of organic carbon can be
identical in chemoautotrophs and photoautotrophs. Discovery
of vents thus led us to understand for the first time that com-
plex food webs could be dependent on microbial chemosyn-
thetic primary production. The initial dives to deep-sea vents
also sparked the hypothesis that life on Earth may have origi-
nated at submarine hot springs (CORLISS et al. 1980).
The triptych of charismatic vent organisms – clams (Calyp-
togena magnifica), mussels (Bathymodiolus thermophilus), and gi-
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-

14
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
cal changes in the environment and to biological interactions
(e.g., SARRAZIN & JUNIPER 1999). The proximity of NE Pacific
vent sites to coastal ports has fostered studies that examine tem-
poral change and community comparisons.
Discoveries in the Atlantic
Discovery of shrimp- (Rimicaris exoculata) dominated vent
fields at TAG (36°N; RONA et al. 1986) and Snake Pit in 1985
(23°N; ODP Leg 106 Scientific Party 1986; MEVEL et al. 1989)
on the Mid-Atlantic Ridge (MAR) highlighted the fact that
similar types of organisms might be found at hydrothermal vents
throughout the world’s oceans (e.g., alvinocarid shrimp, bathy-
modioliform mussels, siphonostome copepods), but that the
species, and even many genera, are different from one ocean
basin to another. Major taxonomic groups familiar from EPR
vents (e.g., vestimentiferan tubeworms, alvinellid polychaetes,
stalked barnacles) are so far unknown at the MAR vents. In
contrast to the dynamic succession of macrofaunal species ob-
served in some NE Pacific vent habitats (e.g., SARRAZIN & JU-
NIPER 1999) and of megafaunal invertebrates at EPR vents (e.g.,
SHANK et al. 1998), communities at MAR vent fields are re-
markably stable on decadal time scales (COPLEY et al. 1997,
1999).
The visual impact of vent megafauna is different between
MAR and eastern Pacific vents: motile shrimp at MAR vents
were described by John Edmond as looking “like maggots
swarming on a hunk of rotten meat”; sessile worms and bivalves
at EPR and NE Pacific vents lend the sites a more garden-like
serenity, as evoked by names like Rose Garden and Garden of
Eden. There are differences as well in the details of the bacter-
ial-invertebrate symbioses implicated in the nourishment of the
dominant megafauna between the two ocean basins. Episym-
bionts likely contribute to the nutrition of rimicarid shrimp on
the MAR (VAN DOVER et al. 1988; GAL’CHENKO 1989; GEBRUK
et al. 1992; SEGONZAC et al. 1993), whereas tubeworms, clams,
and mussels that dominate EPR vents rely on endosymbionts.
MAR mussels have dual endosymbionts (methanotrophs and
thiotrophs), while EPR mussels have only thiotrophic en-
dosymbionts in their gills (FIALA-MEDIONI 1984; LE PENNEC &
HILY 1984). Discovery of shrimp swarms at Mid-Atlantic Ridge
vents ultimately led to the description of modified eyes in Rim-
icaris exoculata and other alvinocarid shrimp, eyes that are
adapted for detecting dim sources of light (VAN DOVER et al.
1989; WHITE et al. 2002).
Western Pacific Explorations
Hydrothermal vents in Manus Basin were discovered in
1985 (BOTH et al. 1986), the same year that vents on the Mid-
Atlantic Ridge were reported (RONA et al. 1986). Further at-
tention was brought to the southwestern Pacific in 1987 with
the first description of vent faunas from the Mariana back-arc
spreading center (HESSLER et al. 1988, HESSLER & LONSDALE
1991). Back-arc spreading centers are zones of extension of
oceanic crust between active and remnant volcanic arcs associ-
ated with subduction zones. Because back-arc systems tend to
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
similar niches was postulated (HESSLER & LONSDALE 1988).
Dense clusters of large, “hairy gastropods” (Alviniconcha hessleri)
with chemoautotrophic endosymbionts in their gills (STEIN et
al. 1988) proved to be one of the most distinctive species, be-
longing to a new genus and having trophic attributes not previ-
ously known within the Gastropoda.
Although Mariana vent organisms belonged to undescribed
species, more than half of the genera found there were already
known from vent fields in the eastern Pacific. A mussel in the
genus Bathymodiolus dominated the biomass; other familiar an-
imal types included polychaete worms in the genus Paralvinella
and limpets in the genus Lepetodrilus. Affinities of some Mari-
ana taxa were shared with species known from Atlantic hy-
drothermal vents – for example, shrimp in the genus Chorocaris.
There were no surprises in terms of fidelity to a particular niche
or microhabitat in genera shared between Mariana and other
mid-ocean ridge vents; the implication is that speciation has
largely been passive within these genera, a consequence of iso-
lation by distance and barriers to dispersal, rather than of adap-
tive radiations into new niches. The Mariana studies provided
the first incontestable evidence for faunal interchange and for
barriers or filters to dispersal of species between Pacific back-arc
basins and the mid-ocean ridge system (HESSLER & LONSDALE
1988). They also provided compelling evidence for the poten-
tial for discovery of unanticipated taxa (in this case, the hairy
gastropods) as new geographic regions are explored.
Subsequent explorations of other southwestern Pacific
back-arc systems [Manus (AUZENDE et al. 1997; HASHIMOTO et
al. 1999); North Fiji and Lau Basins (HASHIMOTO et al. 1989;
DESBRUYÈRES et al. 1994); New Ireland Basin (HERZIG et al.
1994)] demonstrated that some species have restricted distribu-
tions while other species, such as provannid gastropods, are
shared among basins. Molecular techniques have now distin-
guished four species of alviniconchid snails: Alviniconcha hessleri
at Mariana vents, two Alviniconcha species in the Manus and
North Fiji Basins, and a fourth Alviniconcha species at Indian
Ocean vents (KOJIMA et al. 2003; OKUTANI et al. 2004). In con-
trast to the basin-scale differentiation of alviniconchid gas-
tropods in the southwestern Pacific, Ifremeria nautilei (synony-
mous with Olgaconcha tufari) so far is only known to inhabit
vent sites in the Manus, North Fiji, and Lau Basin region (DES-
BRUYÈRES et al. 1994; HASHIMOTO et al. 1999). The relatively
narrow distributional range of Ifremeria nautilei might be attrib-
uted its lower dispersal ability compared to that of Alviniconcha
species (KOJIMA et al. 2000, 2001). Ifremeria nautilei is also the
only known gastropod that supports dual symbioses, with
methane- and sulfur-oxidizing bacteria in gill bacteriocytes
(GAL’CHENKO et al. 1992).
15
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Japanese biological explorations of deep-sea hydrothermal
systems in the northwestern Pacific began in 1988 with the dis-
covery of active hydrothermal vents at Izena Hole (Okinawa
Trough; TANAKA et al. 1990) by scientists using Shinkai 2000.
Subsequent work discovered venting in yet another setting on
the arc volcanos of the Izu-Bonin seamount chain south of
Japan (KOJIMA 2002). Recent exploration of the Mariana Arc
reveals variability in venting conditions and associated com-
munities (EMBLEY et al. 2004). Together, the southwestern and
northwestern Pacific hydrothermal fields represent a fertile re-
gion for the study of population genetics and the biogeography
and evolution of vent taxa (DESBRUYÈRES et al. 2006).
Discovery of Seeps and Other
Chemosynthetic Settings
The 1980s included other milestones: Seep communities
discovered first in 1984 at the base of the Florida Escarpment in
the Gulf of Mexico (PAULL et al. 1984; HECKER 1985) and in
Sagami Bay (OKUTANI & EGAWA 1985; HASHIMOTO et al.
1989), and then in association with subduction zone settings off
Oregon (SUESS et al. 1985), in the northwestern Pacific
(LAUBIER et al. 1986; OHTA & LAUBIER 1987), and off Barbados
(FAUGERES et al. 1987), taught us that vent-like taxa and troph-
ically complex, chemosynthetically based ecosystems are not
restricted to hydrothermal settings. Discovery of chemosyn-
thetic ecosystems and of vent- and seep-like taxa associated
with whale skeletons on the seafloor (SMITH et al. 1989; re-
viewed in SMITH & BACO 2003) underscored the need to adopt
a broad view of chemosynthetic faunas. Vent taxa were also dis-
covered in diverse volcanic and hydrothermal settings, includ-
ing alvinocarid shrimp at a mid-plate, hot-spot volcano (Loihi
seamount; KARL et al. 1989; WILLIAMS & DOBBS 1995), alvinel-
lids, mussels, crabs, and alvinocarid shrimp at Pito Seamount
(NAAR et al. 2004), and relatively shallow (489 m) populations
of vesicomyid clams at Piips volcano (SELIVERSTOV et al. 1986;
SAGALEVITCH et al. 1992). Thus we learn that vent and vent-
like taxa and trophically complex, chemosynthetically based
ecosystems are not restricted to mid-ocean ridge hydrothermal
settings, and that an understanding of the evolution and bio-
geography of vent taxa can only be understood in the broader
context of the evolution and biogeography of allied taxa at oth-
er reducing environments (SIBUET & OLU 1998; VAN DOVER et
al. 2002; TUNNICLIFFE et al. 2003a).
Other shallow hydrothermal settings were found to lack en-
demic invertebrate species and do not support invertebrates
with endosymbiotic chemoautotrophic bacteria [e.g., subtidal
vents off the coast of Southern California (KLEINSCHMIDT &
TSCHAUDER 1985); vents of the volcanic arc in the Mediter-
ranean (reviewed in DANDO et al. 1999); vents on the Kolbein-
sy (Olafsson et al. 1989) and Jan Mayen ridges (FRICKE et al.
1989) north of Iceland; vents in the crater of Ushishir volcano
of the Kamchatka region (TARASOV et al. 1990, TARASOV
2006)].
16
Hydrothermal vents also occur in fresh-water lakes. Fresh-
water hydrothermal systems, similar to those in the ocean, are
related either to rift zones, like lakes Baikal (CRANE et al. 1991)
and Tanganyika (TIERCELIN et al. 1993), or volcanic activity in
subduction zones, e.g. Crater Lake in Oregon (Anonymous
1989). No examples of fresh-water, vent-specific metazoa are
known. Bacterial mats are common at all known fresh-water
vents. Mineral structures analogous to Black smoker chimneys,
reaching up to 10 m in height, can also develop (e.g., in Crater
Lake). The Russian-operated Pisces submersible studies in Lake
Baikal documented aggregations of non-vent fauna in associa-
tion with bacterial mats; carbon derived from bacterial
methane oxidation is incorporated into local food web of the
benthic fauna (GEBRUK et al. 1993).
Eruptions and the Hydrothermal Cycle
1991 was the year of the eruption at 9°50’N on the East Pa-
cific Rise (HAYMON et al. 1993) and the beginning of a long-
term study of hydrothermal cycles in this region (SHANK et al.
1998). The bloom of bacteria that marked the commencement
of the cycle (NELSON et al. 1991; HAYMON et al. 1993) and oth-
er biological indications of recent volcanic activity were subse-
quently observed at other locales, including the NE Pacific
Ridges (e.g., TUNNICLIFFE et al. 1997; JUNIPER et al. 1998) and at
17°S on the East Pacific Rise (EMBLEY et al. 1998). Repeated vis-
its to developing vent communities documented the rapidity
with which colonization of new sites of venting takes place (e.g.,
well-established colonies of tubeworms within one year; TUNNI-
CLIFFE et al. 1997; SHANK et al. 1998) and confirmed earlier re-
ports of rapid growth rates in vent species (LUTZ et al. 1994).
Careful re-sampling of evolving vents allows insights into troph-
ic strategies and the role of biological interactions such as com-
petition (LEVESQUE et al. 2003).
The 9°50N vent field became a favored field site for eco-
logical studies for several reasons: (1) the eruption took place
there, (2) its relatively simple topography has been thoroughly
mapped, (3) its vents support a variety of foundation species
with different habitat preferences (e.g., tubeworm clumps, mus-
sel beds, alvinellid galleries, serpulid and anemone fields), (4)
there is a long record of contemporaneous and complementary
geological and geochemical studies, and (5) the transit time
from major ports in Mexico is relatively short. The 15+ years of
research at 9°50‘N since the eruption have resulted in impor-
tant new insights into successional sequences (SHANK et al.
1998), physiological adaptations to the vent environment (e.g.,
GOFFREDI et al. 1997; SHILLITO et al. 2001), reproductive at-
tributes of vent invertebrates (e.g., MARSH et al. 2000; PRADIL-
LON et al. 2001), biological interactions among species (e.g.,
MULLINEAUX et al. 2000; MICHELI et al. 2002; MULLINEAUX et
al. 2003), animal-chemical relationships (e.g., LUTHER et al.
2001; LE BRIS et al. 2006), population genetics (reviewed in
VRIJENHOEK 1997), and community structure (e.g., VAN DOVER
2003; GOVENAR et al. 2004).
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
One of the longest records of time-series studies at vents is
that for 13°N on the East Pacific Rise, where French and Amer-
ican scientists have made intermittent observations since 1982
(reviewed by DESBRUYÈRES 1995). At this site, a previously
“dead” vent area with relic populations resumed activity in
1987, and followed the successional pattern described for 9°N by
SHANK et al. (1998). The presumption is that there was a sub-
surface dyking event (i.e., an upward migration of molten rock
through a fissure, but without overflow onto the seabed) in 1987
that delivered heat to the system and restored the hydrothermal
circulation at this site.
More Atlantic Discoveries
Attention returned to the Mid-Atlantic Ridge in 1992 with
the chance collection of vent animals in a rock-dredge haul, and
in 1993 during dives at the site that came to be known as Lucky
Strike at 37°N (VAN DOVER et al. 1996; LANGMUIR et al. 1997).
Although Lucky Strike shares a few species with the TAG and
Snake Pit vent sites, most species at Lucky Strike were new to
science. This site and the nearby Menez Gwen vents explored in
1993 (FOUQUET et al. 1995; DESBRUYÈRES et al. 2001) appear to
belong to a biogeographic province different from that of the
more southerly TAG and Snake Pit faunas. Variables associated
with the depth differential between the two provinces (<1700 m
at Lucky Strike and Menez Gwen vs. >3000 m at TAG, Snake
Pit) have been implicated in the biogeographic differentiation
of the two vent faunas (VAN DOVER et al. 1996; DESBRUYÈRES et
al. 2000, 2001).
Continuing geological and geochemical exploration for hy-
drothermal activity on the Mid-Atlantic Ridge between the Ve-
ma transform fault at 11°N and the Azores hotspot (38°N) re-
sulted in the first submersible dives to hydrothermal sites at Lo-
gatchev in 1993 (BOGDANOV et al. 1995; GEBRUK et al. 2000),
Broken Spur in 1993 (MURTON et al. 1995), and Rainbow in
1997 (FOUQUET et al. 1997). The Logatchev site at 14°45’N was
the first vent discovery where the host rock was ultramafic (i.e.,
of mantle rather than crustal origin; BOGDANOV et al. 1997;
FOUQUET et al. 2006). Logatchev represents the current south-
ern limit of published information on vent faunas along the
Mid-Atlantic Ridge, although a new active vent field known as
Ashadze at 12°38’N has been reported (BELTENEV et al. 2004), as
have vent fields south of the equator (C. German, pers. comm).
Logatchev is notable as the only known site on the Mid-At-
lantic Ridge that supports large populations of vesicomyid clams;
in other aspects, it has a species list similar to that found at
Snake Pit (e.g., TURNIPSEED et al. 2004). Broken Spur (29°10’N)
is a hybrid zone for mussels, where two species of mussels (Ba-
thymodiolus puteoserpentis, dominant at southern locales, and B.
azoricus, known from the northern vent sites) co-occur and in-
ter-breed (WON et al. 2003a). Rainbow (FOUQUET et al. 1997),
Saldanha (BARRIGA et al. 1998), and Lost City (KELLEY et al.
2001) vent fields, like Logatchev, are hosted by ultramafic rocks.
Their megafaunal and macrofaunal communities are generally
meager [except for dense shrimp Rimicaris exoculata populations
at Rainbow], possibly due to the distinctive fluid chemistries as-
sociated with these sites (KELLEY et al. 2001).
The Indian Ocean
By 2000, a basic understanding of vent faunas of the north-
ern Mid-Atlantic Ridge, the East Pacific Rise, the Northeast
Pacific, and back-arc basins in the western Pacific was estab-
lished. One of the biggest missing pieces of the biogeographic
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,
and galatheid squat lobsters, but specimens were not collected
(JUNIPER et al. 1990). Japanese scientists brought attention to
the Indian Ocean in 2000, using the ROV Kaiko to explore the
Kairei hydrothermal field on the Central Indian Ridge, just
north of the Rodriguez Triple Junction (HASHIMOTO et al.
2001). Shrimp closely related to Rimicaris exoculata of Atlantic
vents dominate the biomass of the Kairei field so that the site
resembles TAG and Snake Pit, but other taxa, including hairy
gastropods and mussels, are more closely allied with Pacific vent
faunas. These observations supported the hypothesis that the
Indian Ocean ridges serve as a link between Atlantic and Pa-
cific vent faunas (HASHIMOTO et al. 2001). Further explorations
of the Kairei Field (VAN DOVER et al. 2001) led to the discov-
ery of a remarkable scaly-footed gastropod with no close al-
liance to any known Pacific or Atlantic vent genera (Waren et
al. 2003). Unlike other mollusks at vents that house autotroph-
ic endosymbionts in epithelial bacteriocytes of their gills, the
scaly-footed gastropod hosts its symbionts in a hypertrophied
esophageal gland (GOFFREDI et al. 2004).
Fossil Records
The first accounts of fossil vent invertebrates comes from
studies of Cretaceous Oman and Cyprus ophiolites that yielded
well-preserved tubes of worms, including what might be a ves-
timentiferan (HAYMON et al. 1984; OUDIN & CONSTANTINOU
1984; HAYMON & KOSKI 1985). Invertebrates have occupied
vents (KUZNETSOV 1993; LITTLE et al. 1997) and seeps (BARBI-
ERI et al. 2004) at least since the Silurian, but the ancient (old-
er than the early Cretaceous) assemblages described so far were
dominated by now-extinct families of brachiopods, monopla-
cophorans, bivalves, and gastropods (reviewed in CAMPBELL
2006). There is little evidence to support the view that vents
(or seeps) have served as a refuge during global extinction
events for most taxa (reviewed in WARÉN & BOUCHET 2001;
LITTLE & VRIJENHOEK 2003), although there may be exceptions
(YAMAGUCHI et al. 2004). As in extant chemosynthetic com-
munities, fossil records include indications of successional re-
sponses to changing conditions at a given site (CAMPBELL et al.
2002), and paleobiogeographic patterns that are likely to reflect
past plate tectonic configurations (CAMPBELL 2006).
17
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Prospects for Exploration and Discovery
There remain a number of oceanic regions where we can
anticipate discovery of entire communities of new species at hy-
drothermal vents (VAN DOVER et al. 2002; TYLER et al. 2003).
Of these, the Arctic Ocean has long been of great interest: its
relative youth and degree of bathymetric isolation from the rest
of the deep waters of the world’s oceans suggest that the vent
fauna of the region might provide insight into the invasibility
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-
mented and sampled. Shells of vesicomyid clams (Archivesica
sp.), indicative of reducing environments, occur in dredge sam-
ples from depths of ~2000 m near the Gakkel Ridge and seem
likely to have been associated with a hydrothermal site when
the animals were alive (SIRENKO et al. 1995).
Other sites of considerable biogeographic and taxonomic
interest are isolated ridge systems, such as the Scotia Ridge in
the southwest Atlantic, the Cayman Rise in the Caribbean, and
the Andaman back-arc ridge in the northeast Indian Ocean.
Even on the mid-ocean ridge system, there remain large stretch-
es of unexplored ridge axis. One of the most compelling ridge
segments is the Mid-Atlantic Ridge north of the Azores – here
the ridge axis dives from shallowest depths back down to more
than 2500 m – has the Azores hot spot served as a faunal break
to north and south? The deep, greatly offset fracture zones of
the Mid-Atlantic Ridge north of the Azores and south of the
equator (i.e., south of the Romanche Fracture Zone) beg for
studies of gene flow and the role of these fracture zones as bar-
riers or filters to species dispersal. The Chile Rise in the south-
east Pacific and the Antarctic ridges are also certain to be of in-
terest in comparative studies of taxonomy, diversity, and gene
flow. Phylogeographic studies of the vent fauna of the triple
ridge system of the Indian Ocean, in comparison to faunas of
the Atlantic and western Pacific, should allow us to determine
the direction of dispersal of species between the Atlantic and
Pacific.
Even within extremely well-explored vent fields, the po-
tential for discovery of previously unknown adaptations exists,
as in the recent documentation of a blood-sucking polychaete
(Galapagomystides aristata; JENKINS et al. 2002) that had been
described from some of the earliest collections at the Galapagos
Spreading Center (BLAKE 1985). Little is known of the biology
of most vent species. There is great potential in careful studies
of populations, life history traits and behavior of individual
species to learn more of the range of adaptations present in
these unusual habitats. Investigation of relationships of bio-
mass-dominant metazoans to microbial communities and chem-
ical settings will yield better insight into the complex processes
that control vent community structure and function (TUNNI-
CLIFFE et al. 2003b).
The ChEss program (a component of the Census of Marine
18
Life) was conceived to promote international efforts to improve
our understanding of the identity and biogeography of the fau-
na of these reducing environments (TYLER et al. 2003). ChEss
also promotes tests of hypotheses regarding potential barriers
and filters to dispersal of species whose adults are restricted to
these environments. Through ChEss and other ocean explo-
ration activities, we look forward to the next decade of discov-
ery that will undoubtedly add many new species, genera, and
families to the compendium of taxa so carefully compiled here.
Acknowledgements
We thank our many shipboard colleagues – Captains, pilots,
ships’ crews, scientists, technicians, and students – who have
contributed to the discovery of deep-sea hot springs and of the
strange animals that live at them. There are too many individ-
uals to cite by name; we honor their efforts with images of some
of the many vehicles that have given us such remarkable access
to the seafloor (Fig. 1, 2).
Bibliography
In addition to scientific reports, there have been many ex-
cellent reviews of the biology of vent and other chemosynthet-
ic communities that provide entrée into the primary literature
and deliver a historical perspective on the exploration of hy-
drothermal-vent biota. A selection of these reviews is listed
here.
CHILDRESS J.J. & C.R. FISHER (1992): The biology of hydrothermal vent ani-
mals: Physiology, biochemistry, and autotrophic symbioses. —
Oceanogr. Mar. Biol. 30: 337-441.
DESBRUYÈRES D. & M. SEGONZAC (1997): Handbook of Deep-sea Hydrother-
mal Vent Fauna. — Ifremer éd., Plouzané, France: 1-279.
GEBRUK A.V. (Ed.; 2002): Biology of Hydrothermal Systems. — Moscow,
KMK Press: 1-543.
GEBRUK A.V., GALKIN S.V., VERESHCHAKA A.L., MOSKALEV L.I. & A.J. SOUTHWARD
(1997): Ecology and biogeography of the hydrothermal vent fauna
of the Mid-Atlantic Ridge. — Adv. Mar. Biol. 32: 93-144.
GRASSLE J.F. (1985): Hydrothermal vent animals: Distribution and biology.
— Science 229: 713-717.
JOLLIVET D. (1996): Specific and genetic diversity at deep-sea hydrother-
mal vents: An overview. — Biodiv. Conserv. 5: 1619-1653.
JUNIPER S.K. & V. TUNNICLIFFE (1997): Crustal accretion and the hot ecosys-
tem. — Philos. Trans. R. Soc. Lond. A 355: 450-474.
TUNNICLIFFE V. (1991): The biology of hydrothermal vents: ecology and
evolution. — Oceanogr. Mar. Biol. 29: 319-407.
TUNNICLIFFE V. (1992): Hydrothermal vent communities of the deep sea. —
Amer. Sci. 80: 336-349.
TUNNICLIFFE V., FOWLER C.M.R. & A.G. MCARTHUR (1996): Plate tectonic his-
tory and hot vent biogeography. — In: MACLEOD C.J., TYLER P.A. &
C.L. WALKER (Eds.): Tectonic, Magmatic, Hydrothermal and Biological
Segmentation of Mid-Ocean Ridges. J. Geol. Soc. Lond. Spec. Publ.
118: 225-238.
TUNNICLIFFE V., MACARTHUR A.G. & D. MCHUGH (1998): A biogeographical
perspective of the deep-sea hydrothermal vent fauna. — Adv. Mar.
Biol. 34: 353-442.
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
TUNNICLIFFE V., JUNIPER S.K. & M. SIBUET (2003): Reducing environments of
the deep-sea floor. — In: TYLER P.A. (Ed.): Ecosystems of the Deep
Oceans. Ecosystems of the World 28, Elsevier, Amsterdam: 81-110.
VAN DOVER C.L. (1995): Ecology of Mid-Atlantic Ridge hydrothermal
vents. — J. Geol. Soc. Lond. Spec. Publ. 87: 257-294.
VAN DOVER C.L. (2000): The Ecology of Deep-Sea Hydrothermal Vents. —
Princeton University Press, Princeton, NJ: 1-424.
References
Anonymous (1989): Hydrothermal vent communities found in lake. —
Mar. Pollut. Bull. 20: 540-541.
ARP A.J. & J.J. CHILDRESS (1983): Sulfide binding by the blood of the hy-
drothermal vent tube worm Riftia pachyptila. — Science 219: 295-
297.
ARP A.J., CHILDRESS J.J. & C.R. FISHER (1984): Metabolic and blood gas trans-
port characteristics of the hydrothermal vent bivalve Calyptogena
magnifica. — Physiol. Zool. 57: 648-662.
AUZENDE J.-M., HASHIMOTO J., FIALA-MEDIONI A. & S. OHTA (1997): Étude
géologique et biologique in situ de deux zones hydrothermales du
bassin de Manus (Papouasie Nouvelle-Guinée). — C. R. Acad. Sci.
(Ser. 2a) 325: 585-591.
BALLARD R.D. (1977): Notes on a major oceanographic find. — Oceanus
20: 35-44.
BARBIERI R., ORI G.G. & CAVALAZZI B. (2004): A Silurian cold-seep ecosystem
from the Middle Atlas, Morocco. — Palaios 19: 527-542.
BARRIGA F.J.A.S., FOUQUET Y., ALMEIDA A., BISCOITO M.J., CHARLOU J.L., COSTA
R.L.P., DIAS A., MARQUES A.M.S.F., MIRANDA J.M.A., OLU K., PORTEIRO F. &
M.G.P.S. QUEIROZ (1998): Discovery of the Saldanha Hydrothermal
Field on the FAMOUS segment of the MAR (36°30’N). — Eos, Trans.
Amer. Geophys. Union 79: F67.
BELTENEV V.E., NESCHERETOV A.V., IVANOV V.N., V SHILOV.V., ROZHDESTVENSKAYA
I.I., SHAGIN A.A., STEPANOVA T.V., ANDREEVA I.A., SEMENOV Y.P., SERGEEV
M.B., CHERKASHEV G.A., BATUEV B.N., SAMOVAROV M.L., KROTOV A.G. &
V.F. MARKOV (2004): New hydrothermal ore field in the axial zone of
the Mid-Atlantic Ridge. — Dokl. Akad. Nauk 397: 78-82.
BLACK M.B., LUTZ R.A. & R.C. VRIJENHOEK (1994): Gene flow among vesti-
mentiferan tube worm (Riftia pachyptila) populations from hy-
drothermal vents in the eastern Pacific. — Mar. Biol. 120: 33-39.
BLAKE J.A. (1985): Polychaeta from the vicinity of deep-sea geothermal
vents in the Eastern Pacific. I. Euphrosinidae, Phylodocidae, Hesion-
idae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae and Mal-
danidae. — Proc. Biol. Soc. Wash. 6: 67-102.
BOGDANOV Y.A., SAGALEVITCH A.M., CHERNYAEV E.C., ASHADZE A.M., GURVITCH
E.G., LUKASHIN V.N., IVANOV G.V. & V.N. PERESYPKIN (1995): Hydrother-
mal field at 14°45’N on the Mid-Atlantic Ridge. — Dokl. Akad. Nauk
343: 353-357.
BOGDANOV Y.A., BORTNIKOV N.S., VIKENTEV I.V., GURVITCH E.G. & A.M. SAGALE-
VITCH (1997): A new type of modern mineral forming system: Black
smokers of the hydrothermal filed at 14°45’N on the Mid-Atlantic
Ridge. — Geol. Ore Deposits 39: 68-90.
BOTH R., CROOK K., TAYLOR B., BROGAN S., CHAPPELL B.W., FRANKEL E., LIU L.,
SINTON J. & D. TIFFIN (1986): Hydrothermal chimneys and associated
fauna in the Manus back-arc basin, Papua New Guinea. — Eos,
Trans. Amer. Geophys. Union 67: 489-490.
CAMPBELL K.A. (in press): Hydrocarbon seep and hydrothermal vent
palaeoenvironments and palaeontology: Past developments and
future research directions. — Palaeogeogr. Palaeoclimatol.
Palaeoecol.
CAMPBELL K.A., FARMER J.D. & D. DES MARAIS (2002): Ancient hydrocarbon
seeps from the Mesozoic convergent margin of California: Carbon-
ate geochemistry, fluids, and palaeoenvironments. — Geofluids 2:
63-94.
CASM (Canadian American Seamount Expedition; 1985): Hydrothermal
vents on an axis seamount of the Juan de Fuca Ridge. — Nature
313: 212-214.
CAVANAUGH C.M., GARDINER S.L., JONES M.L., JANNASCH H.W. & J.B. WATERBURY
(1981): Prokaryotic cells in the hydrothermal vent tube worm Riftia
pachyptila: Possible chemoautotrophic symbionts. — Science 213:
340-342.
COPLEY J., TYLER P.A., MURTON B.J. & C.L. VAN DOVER (1997): Spatial and in-
terannual variation in the faunal distribution at Broken Spur vent
field (29°N, Mid-Atlantic Ridge). — Mar. Biol. 129: 723-733.
COPLEY J., TYLER P.A., VAN DOVER C.L., SCHULZ A., SULANOWSKA M., SINGH S. &
P. DICKSON (1999): Effects of drilling on the TAG hydrothermal vent
community, 26°N Mid-Atlantic Ridge. — P.S.Z.N. Mar. Ecol. 20: 291-
306.
CORLISS J.B., BAROSS J.A. & S.E. HOFFMAN (1980): An hypothesis concerning
the relationship between submarine hot springs and the origin of
life on Earth. — Eos, Trans. Amer. Geophys. Union 61: 996.
CORLISS J.B., DYMOND J., GORDON L.I., EDMOND J.M., VON HERZEN R.P., BALLARD
R.D., GREEN K., WILLIAMS D., BAINBRIDGE A., CRANE K. & T.H. VAN ANDEL
(1979): Submarine thermal springs on the Galapagos Rift. — Science
203: 1073-1083.
CRANE K., HECKER B. & V. GOLUBEV (1991): Heat flow and hydrothermal
vents in Lake Baikal, USSR. — Eos, Trans. Amer. Geophys. Union 72:
585, 588-589.
DANDO P.R., STUEBEN D. & S.P. VARNAVAS (1999): Hydrothermalism in the
Mediterranean Sea. — Progr. Oceanogr. 44: 333-367.
DESBRUYÈRES D. (1995): Temporal variations of deep-sea hydrothermal
communities at 13°N/EPR. — InterRidge News 4: 6-10.
DESBRUYÈRES D. & L. LAUBIER (1980): Alvinella pompejana gen. sp. nov.,
Ampharetidae aberrant des sources hydrothermales de la ride East-
Pacifique. — Oceanologica Acta 3: 267-274.
DESBRUYÈRES D., CRASSOUS P., GRASSLE J., KHRIPOUNOFF A., REYSS D., RIO M. &
M. VAN PRAET (1982): Données écologiques sur un nouveau site d’hy-
drothermalisme actif de la ride du Pacifique oriental. — C. R. Acad.
Sci. 295: 489-494.
DESBRUYÈRES D., HASHIMOTO J. & M.-C. FABRI (in press): Composition and
biogeography of hydrothermal vent communities in Western Pacif-
ic back arc basins, in Back-Arc Spreading Systems: Geological, Bio-
logical, Chemical and Physical Interactions. — Geophys. Monogr.
DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA & the Scientific Parties of Bi-
olau and Starmer Cruises (1994): Deep-sea hydrothermal communi-
ties in Southwestern Pacific back-arc basins (the North Fiji and Lau
Basins): Composition, microdistribution and food web. — Mar. Ge-
ol. 116: 227-242.
DESBRUYÈRES D., ALMEIDA A., BISCOITO M., COMTET T., KHRIPOUNOFF A., LE BRIS
N., SARRADIN P.M. & M. SEGONZAC (2000): A review of the distribution
of hydrothermal vent communities along the northern Mid-Atlantic
Ridge: dispersal vs. environmental controls. — Hydrobiologia 440:
201-216.
DESBRUYÈRES D., BISCOITO M., CAPRAIS J.-C., COLAÇO A., COMTET T., CRASSOUS P.,
FOUQUET Y., KHRIPOUNOFF A., LE BRIS N., OLU K., RISO R., SARRADIN P.-M.,
SEGONZAC M. & A. VANGRIESHEIM (2001): Variations in deep-sea hy-
drothermal vent communities on the Mid-Atlantic Ridge near the
Azores plateau. — Deep-Sea Res. I 48: 1325-1346
EDMONDS H.N., MICHAEL P.J., BAKER E.T., CONNELLY D.P., SNOW J.E., LANGMUIR
C.H., DICK H.J.B., MUEHE R., GERMAN C.R. & D.W. GRAHAM (2003): Dis-
covery of abundant hydrothermal venting on the ultraslow-spread-
ing Gakkel ridge in the Arctic Ocean. — Nature 421: 252-256.
19
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
EINSELE G., GIESKES J.M., CURRAY J., MOORE D.M., E AGUAYO., AUBRY M.-P.,
FORNARI D., GUERRERO J., KASTNER M., KEITS K., LYLE M., MATOBA Y., MOLI-
NA-CRUZ A., NIEMITZ J., RUEDA J., SAUNDERS A., SCHRADER H., SIMONEIT B. &
V. VACQUIER (1980): Intrusion of basaltic sills into highly porous sedi-
ments, and resulting hydrothermal activity. — Nature 283: 441-445.
EMBLEY R.W., LUPTON J.E., MASSOTH G., URABE T., TUNNICLIFFE V., BUTTERFIELD
D.A., SHIBATA T., OKANO O., KINOSHITA M. & K. FUJIOKA (1998): Geolog-
ic, chemical, and biological evidence for Recent volcanism at 17.5°S;
East Pacific Rise. — Earth Planetary Sci. Lett. 163: 131-147.
EMBLEY R.W., BAKER E.T., CHADWICK W.W. Jr., LUPTON J.E., RESING K.A., MAS-
SOTH G.J. & K. NAKAMURA (2004): Explorations of Mariana Arc volca-
noes reveal new hydrothermal systems — Eos, Trans. Amer. Geo-
phys. Union 85: 37.
ENRIGHT J.T., NEWMAN W.A., HESSLER R.R. & J.A. MCGOWAN (1981): Deep-
ocean hydrothermal vent communities. — Nature 289: 219-221.
FAUGERES J.C., DESBRUYÈRES D., GONTHIER E., GRIBOULARD R., POUTIERS J., DE
RESSEGUIER A. & G. VERNETTE (1987): Sedimentary and biological wit-
ness of the present tectonic activity of the Barbados accretion
wedge. — C. R. Acad. Sci. (Ser. 2) 305: 115-119.
FELBECK H. (1981): Chemoautotrophic potential of the hydrothermal vent
tube worm Riftia pachyptila JONES (Vestimentifera). — Science 213:
336-338.
FIALA-MÉDIONI A. (1984): Mise en évidence par microscopie électronique
à transmission de l’abondance de bactéries symbiotiques dans la
branchie de mollusques bivalves de sources hydrothermales pro-
fondes. — C. R. Acad. Sci. 298: 487-492.
FOUQUET Y., ONDREAS H., CHARLOU J.-L., DONVAL J.P., RADFORD-KNOERY J., COS-
TA I., LOURENCO N. & M.K. TIVEY (1995): Atlantic lava lakes and hot
vents. — Nature 377: 201.
FOUQUET Y., CHARLOU J.L., ONDREAS H., RADFORD-KNOERY H., DONVAL J., DOU-
VILLE J.P., APPRIOU E., CAMBON R., PELL P., LANDURE H.J.Y., NORMAND A.,
PONSEVERA E., GERMAN C.R., PARSON L. M., BARRIGA F.J.A.S., COSTA I.M.A.,
RELVAS J.M.R.S. & A. RIBEIRO (1997): Discovery and first submersible in-
vestigations on the Rainbow hydrothermal field on the MAR
(36°14’N). — Eos, Trans. Amer. Geophys. Union 78: 832.
FOUQUET Y., CAMBON P., CHARLOU J.L., BARRIGA F.J.A.S., ONDREAS H., BAYON G.,
HENRY K., MURPHY P., ROUXEL O., POROSHINA I., DONVAL J.P. & J. ETOUBLEAU
(in press). Geodiversity of hydrothermal processes along the Mid-
Atlantic Ridge – Ultramafic-hosted mineralization: a new type of
oceanic Cu-Zn-Co-Au VMS deposit. — Economic Geol.
FRICKE H., GIERE O., STETTER K., ALFREDSSON G.A., KRISTJANSSON J.K., STOFFERS P.
& J. SVAVARSSON (1989): Hydrothermal vent communities at the shal-
low subpolar Mid-Atlantic Ridge. — Mar. Biol. 102: 425-429.
FUSTEC A., DESBRUYÈRES D. & S.K. JUNIPER (1987): Deep-sea hydrothermal
vent communities at 13°N on the East Pacific Rise: Microdistribution
and temporal variations. — Biol. Oceanogr. 4: 121-164.
GAL’CHENKO V.F., PIMENOV N.V., LEIN A.Y., GALKIN S.V., MOSKALEV L.I. & M.V.
IVANOV (1989): Autotrophic CO2-assimilation in tissues of shrimp
Rimicaris exoculata from a hydrothermal area on the Mid-Atlantic
Ridge. — Dokl. Akad. Nauk 308: 1478-1481.
GAL’CHENKO V.F., PIMENOV N.V., LEIN A.Y., GALKIN S.V., MILLER Y.M. & M.V.
IVANOV (1992): Mixotrophic type of feeding of Olgaconcha tufari
BECK (Gastropoda: Prosobranchia) from the active hydrothermal
field of the Manus Basin (Bismarck Sea). — Dokl. Akad. Nauk 323:
776-781.
GEBRUK A.V., PIMENOV N.V. & A.S. SAVVICHEV (1992): Bacterial “farm“ on the
mouthparts of shrimp. — Priroda 6: 37-39.
GEBRUK A.V., KUZNETSOV A.P., NAMSARAEV B.B. & Y.M. MILLER (1993): Role of
bacterial component in feeding of deep-water benthic fauna in
Froliha Bay, Lake Baikal. — Izvestija Acad. Nauk Ser. Biol. 6: 903-908.
20
GEBRUK A.V., CHEVALDONNÉ P., SHANK T., LUTZ R.A. & R.C. VRIJENHOEK (2000):
Deep-sea hydrothermal vent communities of the Logatchev area
(14°45’N, Mid-Atlantic Ridge): Diverse biotopes and high biomass.
— J. Mar. Biol. Ass. U.K. 80: 383-394.
GEISTDOERFER P., AUZENDE J.-M., BATIZA R., BIDEAU D., CORMIER M.-H., FOUQUET
Y., LAGABRIELLE Y., SINTON J. & P. SPADEA (1995): Hydrothermalisme et
communautés animales associés sur la dorsale du Pacifique oriental
entre 17°S and 19°S (Campagne NAUDUR, Décembre 1993). — C. R.
Acad. Sci. 320: 47-54.
GOFFREDI S.K., CHILDRESS J.J., DESAULNIERS N.T. & F.J. LALLIER (1997): Sulfide ac-
quisition by the vent worm Riftia pachyptila appears to be via up-
take of HS^, rather than H2S. — J. Exp. Biol. 200: 2609-2616.
GOFFREDI S.K., WARÉN A., ORPHAN V.J., VAN DOVER C.L. & R.C. VRIJENHOEK
(2004): Novel forms of structural integration between microbes. —
Appl. Environm. Microbiol. 70: 3082-3090.
GOVENAR B., FREEMAN M., BERGQUIST D.C., JOHNSON G.A. & C.R. FISHER (2004):
Composition of a one-year-old Riftia pachyptila community follow-
ing a clearance experiment: Insight to succession patterns at deep-
sea hydrothermal vents. — Biol. Bull. 207: 177-182.
GRASSLE J.F., BERG C.J., CHILDRESS J.J., GRASSLE J.P., HESSLER R.R., JANNASCH H.,
KARL D.M., LUTZ R.A., MICKEL T.J., RHOADS D.C., SANDERS H.L., SMITH K.L.,
SOMERO G.N., TURNER R.D., TUTTLE J.H., WALSH P.J. & A.J. WILLIAMS
(1979): Galapagos ’79: Initial findings of a deep-sea biological
quest. — Oceanus 22: 2-10.
GUINOT D. & L.A. HURTADO (2003): Two new species of hydrothermal vent
crabs of the genus Bythograea from the southern East Pacific Rise
and from the Galapagos Rift (Crustacea, Decapoda, Brachyura,
Bythograeidae). — C. R. Acad. Sci. 326: 423-439.
HASHIMOTO J., JOLLIVET D. & the KAIYO 88 Shipboard Party (1989): The hy-
drothermal vent communities in the North Fiji Basin: Results of
Japan-France cooperative research on board KAIYO 88. — La Mer
27: 62-71.
HASHIMOTO J., OHTA S., TANAKA T., HOTTA H., MATSUZAWA S. & H. SAKAI (1989):
Deep-sea communities dominated by the giant clam, Calyptogena
soyoae, along the slope foot of Hatsushima Island, Sagami Bay, cen-
tral Japan. — Palaeogeogr. Palaeoclimatol. Palaeoecol. 71: 179-192.
HASHIMOTO J., OHTA S., FIALA-MEDIONI A., AUZENDE J.-M., KOJIMA S., SEGONZAC
M., FUJIWARA Y., HUNT J. C., GENA K., MIURA T., KIKUCHI T., YAMAGUCHI T.,
TODA T., CHIBA H., S TSUCHIDA., ISHIBASHI J., HENRY K., ZBINDEN M., PRUSKI
A., INOUE A., KOBAYASHI H., BIRRIEN J.-L., NAKA J., YAMANAKA T., LAPORTE
C., NISHIMURA K., YEATS C., MALAGUN S., KIA P., OYAIZU M. & T. KATAYAMA
(1999): Hydrothermal vent communities in the Manus Basin, Papua
New Guinea: Results of the BIOACCESS cruise ’96 and ’98. — Inter-
Ridge News 8: 12-18.
HASHIMOTO J., OHTA S., GAMO T., CHIBA H., YAMAGUCHI T., TSUCHIDA S., OKU-
DAIRA T., WATABE H., YAMANAKA T. & M. KITAZAWA (2001): First hy-
drothermal vent communities from the Indian Ocean discovered. —
Zool. Sci. 18: 717-721.
HAYMON R.M. & R.A. KOSKI (1985): Evidence of an ancient hydrothermal
vent community: Fossil worm tubes in Cretaceous sulfide deposit of
the Samail Ophiolite, Oman. — Proc. Biol. Soc. Wash. 6: 57-65.
HAYMON R.M., KOSKI R.A. & C. SINCLAIR (1984): Fossils of hydrothermal vent
worms from Cretaceous sulfide ores of the Samail Ophiolite, Oman.
— Science 223: 1407-1409.
HAYMON R.M., FORNARI D.J., VON DAMM K.L., LILLEY M.D., PERFIT M.R., ED-
MOND J.M, SHANKS W.C. III, LUTZ R.A., GREBMEIER J.M., CARBOTTE S.,
WRIGHT C.D., MCLAUGHLIN E., SMITH M., BEEDLE N. & E. OLSON (1993):
Volcanic eruption of the mid-ocean ridge along the East Pacific Rise
crest at 9°45-52’N: Direct submersible observations of seafloor phe-
nomena associated with an eruption event in April 1991. — Earth
Planetary Sci. Lett. 119: 85-101.
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
HECKER B. (1985): Fauna from a cold sulfur-seep in the Gulf of Mexico:
Comparison with hydrothermal vent communities and evolutionary
implications. — Proc. Biol. Soc. Wash. 6: 465-473.
HERZIG P., HANNINGTON M., MCINNES B., STOFFERS P., VILLINGER H.W., SEIFERT R.,
BINNS R. & T. LIEBE (1994): Submarine volcanism and hydrothermal
venting studied in Papua, New Guinea. — Eos, Trans. Amer. Geo-
phys. Union 75: 513-, 515-516.
HESSLER R.R. & P.F. LONSDALE (1991): Biogeography of Mariana Trough hy-
drothermal vent communities. — Deep-Sea Res. A 38: 185-199.
HESSLER R.R. & W.M. SMITHEY (1983): The distribution and community
structure of megafauna at the Galapagos Rift hydrothermal vents.
— In: RONA P.A., BOSTRÖM K., LAUBIER L. & K.L. Jr. SMITH (Eds.): Hy-
drothermal Processes at Seafloor Spreading Centers, Plenum Press,
NY: 735-770.
HESSLER R.R., SMITHEY W.M. Jr. & C.H. KELLER (1985): Spatial and temporal
variation of giant clams, tube worms and mussels at deep-sea hy-
drothermal vents. — Bull. Biol. Soc. Wash. 6: 411-428.
HESSLER R.R., LONSDALE P. & J. HAWKINS (1988): Patterns on the ocean floor.
— New Scientist 1605: 47-51.
HURTADO L.A., LUTZ R.A. & R.C. VRIJENHOEK (2004): Distinct patterns of ge-
netic differentiation among annelids of eastern Pacific hydrother-
mal vents. — Mol. Ecol. 13: 2603-15
JENKINS C.D., WARD M., TURNIPSEED M., OSTERBERG J. & C.L. VAN DOVER (2002):
The digestive system of the hydrothermal vent polychaete Gala-
pagomystides aristata (Phyllodocidae): Evidence for hematophagy?
— Invertebr. Biol. 121: 243-254.
JONES, M.L. (1981): Riftia pachyptila JONES: Observations on the vesti-
mentiferan worm from the Galapagos Rift. — Science 213: 333-336.
JONES M.L. & C.F. BRIGHT (1985): Appendix 1. Bibliography of hydrother-
mal vents and related areas. Their biotas, ecological parameters
and ancillary data. — Proc. Biol. Soc. Wash. 6: 495-538.
JUNIPER S.K., TUNNICLIFFE V. & D. DESBRUYÈRES (1990): Regional-scale features
of Northeast Pacific, East Pacific Rise, and Gulf of Aden vent com-
munities. — In: MCMURRAY G.R. (Ed.): Gorda Ridge: A Seafloor
Spreading Center in the United States Exclusive Economic Zone.
Springer, NY: 265-278.
JUNIPER S.K., TUNNICLIFFE V. & E.C. SOUTHWARD (1992): Hydrothermal vents in
turbidite sediments on a Northeast Pacific spreading centre: Or-
ganisms and substratum at an ocean drilling site. — Canad.J. Zool.
70: 1792-1809.
JUNIPER S.K., BIRD D.F., SUMMIT M., PONG VONG M. & E.T. BAKER (1998): Bac-
terial and viral abundances in hydrothermal event plumes over
northern Gorda Ridge. — Deep-Sea Res. II 45: 2739-2749.
KARL D.M., BRITTAIN A.M. & B.D. TILBROOK (1989): Hydrothermal and mi-
crobial processes at Loihi Seamount, a mid-plate hot-spot volcano.
— Deep-Sea Res. A 36: 1655-1673.
KELLEY D.S., KARSON J.A., BLACKMAN D.K., FRUEH-GREEN G.L., BUTTERFIELD D.A.,
LILLEY M.D., OLSON E.J., SCHRENK M.O., ROE K.K., LEBON G.T. & P. RIV-
IZZIGNO (2001): An off-axis hydrothermal vent field near the Mid-At-
lantic Ridge at 30°N. — Nature 412: 145-149.
KLEINSCHMIDT M. & R. TSCHAUDER (1985): Shallow-water hydrothermal vent
systems off the Palos Verdes Peninsula, Los Angeles County, Califor-
nia. — Proc. Biol. Soc. Wash. 6: 485-488.
KOJIMA S. (2002): Deep-sea chemoautosynthesis-based communities in
the Northwestern Pacific. — J. Oceanogr. 58: 343-363.
KOJIMA S., SEGAWA R., FUJIWARA Y., HASHIMOTO J. & S. OHTA (2000): Genetic
differentiation of populations of a hydrothermal vent-endemic gas-
tropod, Ifremeria nautilei, between the North Fiji Basin and the
Manus Basin revealed by nucleotide sequences of mitochondrial
DNA. — Zool. Sci. 17: 1167-1174.
KOJIMA S., SEGAWA R., FUJIWARA Y., FUJIKURA K., OHTA S. & J. HASHIMOTO
(2001): Phylogeny of hydrothermal-vent-endemic gastropods
Alviniconcha spp. from the western Pacific revealed by mitochon-
drial DNA sequences. — Biol. Bull. Mar. Biol.Lab. Woods Hole 200:
298-304.
KOJIMA S., FUJIKURA K., OKUTANI T. & J. HASHIMOTO (2003): Phylogenetic re-
lationship of Alviniconcha gastropods from the Indian Ocean to
those from the Pacific Ocean (Mollusca: Provannidae) revealed by
nucleotide sequences of Mitochondrial DNA. — Venus 63: 65-68.
KUZNETSOV A.P., MASLENNIKOV V.V. & V.V. ZAIKOV (1993): Hydrothermal fau-
na of the Silurian paleoocean in South Ural. — Izvestiya Akademii
Nauk, Ser. Biol. 4: 525-534.
LANGMUIR C., HUMPHRIS S., FORNARI D., VAN DOVER C., VON DAMM K., TIVEY
M.K., COLODNER D., CHARLOU J.-L., DESONIE D., WILSON C., FOUQUET Y.,
KLINKHAMMER G. & H. BOUGAULT (1997): Hydrothermal vents near a
mantle hot spot: The Lucky Strike vent field at 37°N on the Mid-At-
lantic Ridge. — Earth Planetary Sci. Lett. 148: 69-91.
LAUBIER L. & D. DESBRUYÈRES (1985): Oases at the bottom of the ocean. —
Endeavour 9: 67-76.
LAUBIER L., OHTA S. & M. SIBUET (1986): Decouverte de communautes ani-
mals profondes durant la campagne franco-japonaise KAIKO de
plongees dans les fosses de subduction autour du Japon. — C. R.
Acad. Sci. (Ser. 3) 303: 25-29.
LE PENNEC M. & A. HILY (1984): Anatomie, structure et ultra structure de
la branchie d’un Mytilidae des sites hydrothermaux du Pacifique
Oriental. — Oceanol. Acta 7: 517-523.
LE BRIS N., GOVENAR B., LE GALL C. & C.R. FISHER (2006): Variability of physi-
co-chemical conditions in 9°50’N EPR diffuse flow vent habitats. —
Mar. Chem. 98: 167-182.
LEVESQUE C., JUNIPER S.K. & J. MARCUS (2003): Food resource partitioning
and competition among alvinellid polychaetes of Juan de Fuca
Ridge hydrothermal vents. — Mar. Ecol. Progr. Ser. 246: 173-182.
LITTLE C.T.S. & R.C. VRIJENHOEK (2003): Are hydrothermal-vent animals liv-
ing fossils? — Trends Ecol. Evol. 18: 582-588.
LITTLE C.T.S., HERRINGTON R.J., MASLENNIKOV V.V., MORRIS N.J. & V.V. ZAYKOV
(1997): Silurian hydrothermal-vent community from the southern
Urals, Russia. — Nature 385: 146-148.
LISTER C.R.B. (1972): On the thermal balance of a mid-ocean ridge. —
Geophys. J. Roy. Astronom. Soc. 26: 515-535.
LONSDALE P. (1977): Clustering of suspension-feeding macrobenthos near
abyssal hydrothermal vents at oceanic spreading centers. — Deep-
Sea Res. 24: 857-863.
LUTHER G.W. III, ROZAN T.F., TAILLEFERT M., NUZZIO D.B., DI MEO C., SHANK T.M.,
LUTZ R.A. & S.C. CARY (2001): Chemical speciation drives hydrother-
mal vent ecology. — Nature 410: 813-816.
LUTZ R.A., SHANK T.M., FORNARI D.J., HAYMON R.M., LILLEY M.D., VON DAMM
K.L. & D. DESBRUYÈRES (1994): Rapid growth at deep-sea vents. — Na-
ture 371: 663-664.
MARSH A.G., MULLINEAUX L.S., YOUNG C.M. & D.T. MANAHAN (2001): Larval
dispersal potential of the tubeworm Riftia pachyptila at deep-sea
hydrothermal vents. — Nature 411: 77-80
MEVEL C., AUZENDE J.-M., CANNAT M., DONVAL J.P., DUBOIS J., FOUQUET Y., GENTE
P., GRIMAUD D., KARSON J.A., SEGONZAC M. & M. STIEVENARD (1989): La
ride du Snake Pit (dorsale médio-Atlantique, 23°22’N); résultats
préliminaires de la campagne Hydrosnake. — C. R. Acad. Sci. (Ser. 2)
308: 545-552.
MICHELI F., PETERSON C.H., JOHNSON G.A., MULLINEAUX L.S., MILLS S.W., SANCHO
G., FISHER C.R. & H.S. LENIHAN (2002): Predation structures communi-
ties at deep-sea hydrothermal vents. — Ecol. Monogr. 72: 365-382.
21
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
MULLINEAUX L.S., FISHER C.R., PETERSON C.H. & S.W. SCHAEFFER (2000): Tube-
worm succession at hydrothermal vents: use of biogenic cues to re-
duce habitat selection error? — Oecologia 123: 275-284.
MULLINEAUX L.S., PETERSON C.H., MICHELI F. & S.W. MILLS (2003): Successional
mechanism varies along a gradient in hydrothermal fluid flux at
deep-sea vents. — Ecol. Monogr. 73: 523-542.
MURTON B.J., KLINKHAMMER G., BECKER K., BRIAIS A., EDGE D., HAYWARD N., MIL-
LARD N., MITCHELL I., ROUSE I., RUDNICKI M., SAYANAGI K., SLOAN H. & L.
PARSON (1994): Direct evidence for the distribution and occurrence
of hydrothermal activity between 27°N-30°N on the Mid-Atlantic
Ridge. — Earth Planetary Sci. Lett. 125: 119-128.
NAAR D.F., HEKINIAN R., SEGONZAC M., FRANCHETEAU J., & the Pito Dive Team
(2004): Vigorous venting and biology at Pito Seamount, Easter Mi-
croplate. — In: GERMAN C.R., LIN J., L.M. PARSON (Eds.): Mid-Ocean
Ridges: Hydrothermal Interactions Between the Lithosphere and
Oceans. Amer. Geophys. Union, Washington, D.C.: 305-318.
NELSON D.C., HAYMON R.M., LILLEY M. & R. LUTZ (1991): Rapid growth of un-
usual hydrothermal bacteria observed at new vents during the AD-
VENTURE dive program on the EPR crest at 9°45’-52’N. — Eos, Trans.
Amer. Geophys. Union 72: 481.
ODP Leg 106 Scientific Party (1986): Drilling the Snake Pit hydrothermal
sulfide deposit on the Mid-Atlantic Ridge, 23°22’N. — Geology 14:
3739-3762.
OHTA S. & L. LAUBIER (1987): Deep biological communities in the subduc-
tion zone of Japan observed by bottom photographs taken during
Nautile dives in the KAIKO project. — Earth Planetary Sci. Lett. 83:
329-342.
OKUTANI T. & K. EGAWA (1985): The first underwater observation on living
habitat and thanatocoenoses of Calyptogena soyoae in bathyal
depth of Sagami Bay. — Venus 49: 83-91.
OKUTANI T., HASHIMOTO J. & T. SASAKI (2004): New gastropod taxa from hy-
drothermal vent (Kairei Field) in the central Indian Ocean. — Venus
63: 1-12.
OUDIN E. & G. CONSTANTINOU (1984): Black smoker chimney fragments in
Cyprus sulphide deposits. — Nature 308: 349-353.
PAULL C.K., HECKER B., COMMEAU R., FREEMAN-LYNDE R.P., NEUMANN C., CORSO
W.P., GOLUBIC S., HOOK J.E., SIKES E. & J. CURRAY (1984): Biological com-
munities at the Florida Escarpment resemble hydrothermal vent
taxa. — Science 226: 965-967.
POWELL M.A. & G.N. SOMERO (1983): Blood components prevent sulfide
poisoning of respiration of the hydrothermal vent tube worm Rif-
tia pachyptila. — Science 219: 297-299.
POWELL M.A. & G.N. SOMERO (1986): Adaptations to sulfide by hydrother-
mal vent animals: Sites and mechanisms of detoxification and me-
tabolism. — Biol. Bull. Mar. Biol.Lab. Woods Hole 171: 274-290.
PRADILLON F., SHILLITO B., YOUNG C.M. & F. GAILL (2001): Developmental ar-
rest in vent worm embryos. — Nature 413: 698-699.
Rise Project Group (1980): East Pacific Rise: Hot springs and geophysical
experiments. — Science 207: 1421-1433.
RONA P.A., KLINKHAMMER G., NELSON T.A., TREFRY J.H. & H. ELDERFIELD (1986):
Black smokers, massive sulfides and vent biota at the Mid-Atlantic
Ridge. — Nature 321: 33-37.
SAGALEVITCH A.M., TOROKHOV P.V., MATVEENKOV V.V., GALKIN S.V. & L.I.
MOSKALEV (1992): Hydrothermal manifestations of the submarine Pi-
ip’s Volcano. — Izvestiya RAN, Ser. Geol. 9: 104-114.
SARRAZIN J. & S.K. JUNIPER (1999): Biological characteristics of a hydrother-
mal edifice mosaic community. — Mar. Ecol. Progr. Ser. 185: 1-19.
SARRAZIN J., ROBIGOU V., JUNIPER S.K. & J.R. DELANEY (1997): Biological and
geological dynamics over four years on a high-temperature sulfide
structure at the Juan de Fuca Ridge hydrothermal observatory. —
Mar. Ecol. Progr. Ser. 153: 5–24.
22
SEGONZAC M., DE SAINT LAURENT M. & B. CASANOVA (1993): L’énigme du com-
portement trophique des crevettes Alvinocarididae des sites hy-
drothermaux de la dorsale médio-atlantique. — Cah. Biol. Mar. 34:
535-571.
SELIVERSTOV N.I., AVDEIKO G.P., IVANENKO A.N., SHKIRA V.A. & S.A. HUBUNAIA
(1986): New submarine volcano in the western part of the Aleutian
Island Arc. — Volcanol. Seismol. 4: 3-16.
SHANK T.M., FORNARI D.J., VON DAMM K.L., LILLEY M.D., HAYMON R.M. & R.A.
LUTZ (1998): Temporal and spatial patterns of biological community
development at nascent deep-sea hydrothermal vents (9°50’N, East
Pacific Rise). — Deep-Sea Res. II 45: 465-515.
SHILLITO B., JOLLIVET D., SARRADIN P.-M., RODIER P., LALLIER F., DESBRUYÈRES D. &
F. GAILL (2001): Temperature resistance of Hesiolyra bergi, a poly-
chaetous annelid living on deep-sea vent smoker walls. — Mar. Ecol.
Progr. Ser. 216: 141-149.
SIBUET M. & K. OLU (1998): Biogeography, biodiversity and fluid depend-
ence of deep-sea cold-seep communities at active and passive mar-
gins. — Deep-Sea Res. II 45: 517-567.
SIRENKO B.I., PETRYASHOV V.V., RACHOR E. & K. HINZ (1995): Bottom bio-
cenoses of the Laptev Sea and adjacent areas. — Ber. Polar-
forsch./Rep. Polar Res. 176: 211-221.
SMITH C.R. & A.R. BACO (2003): Ecology of whale falls at the deep-sea
floor. — Oceanogr. Mar. Biol. 41: 311–354.
SMITH C.R., KUKERT H., WHEATCROFT R.A., JUMARS P.A. & J.W. DEMING (1989):
Vent fauna on whale remains. — Nature 323: 251-253.
STEIN J.L., CARY S.C., HESSLER R.R., OHTA S., VETTER R.D., CHILDRESS J.J. & H. FEL-
BECK (1988): Chemoautotrophic symbiosis in a hydrothermal vent
gastropod. — Biol. Bull. Mar. Biol. Lab. Woods Hole 174: 373-378.
STEIN C.A., STEIN S. & A.M. PELAYO (1995): Heat flow and hydrothermal cir-
culation. — In: HUMPHRIS S.E., ZIERENBERG R.A., MULLINEAUX L.S. & R.E.
THOMSON (Eds.): Seafloor Hydrothermal Systems: Physical, Chemical,
and Geochemical Interactions. Geophys. Monogr. 91, Amer. Geo-
phys. Union, Washington, DC: 425-445.
SUESS E., CARSON B., RITGER S.D., MOORE J.C., JONES M.L., KULM L.D. & G.R.
COCHRANE (1985): Biological communities at vent sites along the sub-
duction zone off Oregon. — Proc. Biol. Soc. Wash. 6: 475-484.
TALWANI M., WINDISCH C.C. & M.L. LANGSETH (1971): Reykjanes ridge crest:
A detailed geophysical study. — J. Geophys. Res. 76: 473-517.
TANAKA T., HOTA H., SAKAI H., ISHIBASHI J., OOMORI T., IZAWA E. & N. ODA
(1990): Occurrence and distribution of the hydrothermal deposits in
the Izena Hole, central Okinawa Trough. — JAMSTEC Deep Sea Res.
6: 11-26.
TARASOV V.G. (in press): Effects of shallow-water hydrothermal venting
on biological communities of coastal marine ecosystems of the
western Pacific. — Adv. Mar. Biol.
TARASOV V.G., PROPP M.V., PROPP L.N., ZHIRMUNSKY A.V., NAMSARAEV B.B., GOR-
LENKO V.M. & D.A. STARYNIN (1990): Shallow-water gasohydrothermal
vents of Ushishir Volcano and the ecosystem of Kraternaya Bight
(the Kurile Islands). — P.S.Z.N. Mar. Ecol. 11: 1-23.
TIERCELIN J.-J., PFLUMIO C., CASTREC M., BOULEGUE J., GENTE P., ROLET J., COUSSE-
MENT C., STETTER K.O., HUBER R., BUKU S. & W. MIFUNDI (1993): Hy-
drothermal vents in Lake Tanganyika, East African Rift system. —
Geology 21: 499-502.
TSURAMI M. & V. TUNNICLIFFE (2001): Characteristics of a hydrothermal vent
assemblage on a volcanically active segment of Juan de Fuca Ridge,
northeast Pacific. — Canad. J. Fisheries Aquat. Sci. 58: 530-542.
TUNNICLIFFE V. (1988): Biogeography and evolution of hydrothermal-vent
fauna in the eastern Pacific Ocean. — Proc. Roy. Soc. Lond. B 233:
347-366.
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

TUNNICLIFFE V., JUNIPER S.K. & M.E. DE BURGH (1985): The hydrothermal vent WARÉN A. & P. BOUCHET (2001): Gastropoda and Monoplacophora from
community on Axial Seamount, Juan de Fuca Ridge. — Proc. Biol. hydrothermal vents and seeps; new taxa and records. — The Veliger
Soc. Wash. 6: 453-464. 44: 116-231.

TUNNICLIFFE V., JUNIPER S.K. & M. SIBUET (2003a): Reducing environments of
the deep-sea floor. — In: P. TYLER (Ed.): Ecosystems of the Deep
Oceans. Elsevier, Amsterdam: 81-110.
TUNNICLIFFE V., M BOTROS., DE BURUGH M.E., DONET A., JONSON H.P., JUNIPER S.K.
& R.E. MCDUFF (1986): Hydrothermal vents of Explorer Ridge, north-
east Pacific. — Deep-Sea Res. A 33: 401-412.
TUNNICLIFFE V., EMBLEY R.W., HOLDEN J.F., BUTTERFIELD D.A., MASSOTH G.J. &
S.K. JUNIPER (1997): Biological colonization of new hydrothermal
vents following an eruption on Juan de Fuca Ridge. — Deep-Sea
Res. I 44: 1627-1644.
WARÉN A., BENGTSON S., GOFFREDI S.K. & C.L. VAN DOVER (2003): A hot-vent
gastropod with iron-sulphide dermal sclerites. — Science 302: 1007.
WHITE S.N., CHAVE A.D., REYNOLDS G.T. & C.L. VAN DOVER (2002): Ambient
light emission from hydrothermal vents on the Mid-Atlantic Ridge.
— Geophys. Res. Lett. 29: 10.1029/2002GL014977.
WILLIAMS A.G. & F.C. DOBBS (1995): A new genus and species of caridean
shrimp (Crustacea: Decapoda: Bresiliidae) from hydrothermal vents
on Loihi Seamount, Hawaii. — Proc. Biol. Soc. Wash. 108: 228-237.
WOLERY T.J. & N.H. SLEEP (1976): Hydrothermal circulation and geother-
mal flux at mid-ocean ridges. — J. Geol. 84: 249-275.

TUNNICLIFFE V., BAROSS J.A., GEBRUK A.V., GIERE O., HOLLAND M.E., KOSCHINSKY
A., REYSENBACH A.-L. & T.M. SHANK (2003b): Group Report: What are
the interactions between biotic processes at vents and physical,
chemical and geological conditions? — In: HALBACK P.E., TUNNICLIFFE V.
& J.R. HEIN (Eds.): Energy and Mass Transfer in Marine Hydrothermal
Systems. Dahlem Press, Berlin: 251-270.
TURNIPSEED M., JENKINS C.D. & C.L. VAN DOVER (2004): Community structure
in Florida Escarpment seep and Snake Pit (Mid-Atlantic Ridge) vent
mussel beds. — Mar. Biol. 145: 121-132.
TYLER P.A., GERMAN C.R., RAMIREZ-LLODRA E. & C.L. VAN DOVER (2003): Un-
derstanding the biogeography of chemosynthetic ecosystems. —
Oceanologica Acta 25: 227-241.
WON Y., HALLAM S.J., O’MULLAN G.D. & R.C.VRIJENHOEK (2003a): Cytonuclear
disequilibrium in a hybrid zone involving deep-sea hydrothermal
vent mussels of the genus Bathymodiolus. — Mol. Ecol. 12: 3185-
3190.
WON Y., YOUNG C.R., LUTZ R.A. & R.C. VRIJENHOEK (2003b): Dispersal barri-
ers and isolation among deep-sea mussel populations (Mytilidae:
Bathymodiolus) from eastern Pacific hydrothermal vents. — Mol.
Ecol. 12: 169-184.
YAMAGUCHI T., NEWMAN W.A. & J. HASHIMOTO (2004): A cold seep barnacle
(Cirripedia: Neolepadinae) from Japan and the age of the vent/seep
fauna. — J. Mar. Biol. Ass. U.K. 84: 111-120.

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

drothermal vent mussel beds of the East Pacific Rise. — Mar. Ecol.
Progr. Ser. 253: 55–66.

VAN DOVER C.L., GRASSLE J.F. & M. BOUDRIAS (1990): Hydrothermal vent fau-
na of Escanaba Trough (Gorda Ridge). — In: MCMURRAY G.R. (Ed.):
Gorda Ridge: A Seafloor Spreading Center in the United States Ex-
clusive Economic Zone. Springer-Verlag, NY: 253-264.

VAN DOVER C.L., FRY B., GRASSLE J.F., HUMPHRIS S. & P.A. RONA (1988): Feed-
ing biology of the shrimp Rimicaris exoculata at hydrothermal vents
on the Mid-Atlantic Ridge. — Mar. Biol. 98: 209-216.

VAN DOVER C.L., SZUTS E.Z., CHAMBERLAIN S.C. & J.R. CANN (1989): A novel eye
in ‘eyeless’ shrimp from hydrothermal vents of the Mid-Atlantic
Ridge. — Nature 337: 458-460.

VAN DOVER C.L., DESBRUYÈRES D., SEGONZAC M., COMTET T., SALDANHA L., FIALA-
MEDIONI A. & C. LANGMUIR (1996): Biology of the Lucky Strike hy-
drothermal field. — Deep-Sea Res. I 43: 1509-1529.

VAN DOVER C.L., HUMPHRIS S.E., FORNARI D., CAVANAUGH C.M., COLLIER R., GOF-
FREDI S.K., HASHIMOTO J., LILLEY M.D., REYSENBACH A.L., T SHANK.M., VON
DAMM K.L., BANTA A., GALLANT R.M. & R.C. VRIJENHOEK (2001): Bio-
geography and ecological setting of Indian Ocean hydrothermal
vents. — Science 294: 818-823.

VAN DOVER C.L., GERMAN C.R., SPEER K.G., PARSON L.M. & R.C. VRIJENHOEK
(2002): Evolution and biogeography of deep-sea vent and seep in-
vertebrates. — Science 295: 1253-1257.

VINE F.J. & D.H. MATTHEWS (1963): Magnetic anomalies over oceanic
ridges. — Nature 199: 947-949.

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

hadal (ultra-abyssal) fauna in relation to vertical zonation of the
ocean. — Sarsia 64: 41-50.

VRIJENHOEK R.C. (1997): Gene flow and genetic diversity in naturally frag-
mented metapopulations of deep-sea hydrothermal vent animals.
— J. Heredity 88: 285-293.

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

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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Hydrothermal vent meiofauna
Meiofauna, small animals and protists, which pass through
a net with 1 mm mesh size and are retained on a net with 63
µm (or smaller) mesh size (GIERE 1993), is part of the hy-
drothermal vent community. In general, very little is known
about the diversity of vent meiofauna. So far, only a few phyla
such as the Arthropoda (Acariformes, Copepoda, Cumacea,
Isopoda, Ostracoda, Tanaidacea), the foraminiferan Granu-
loreticulosa, and the Nematoda have been described from
vents. In addition, Plathelminthes and three protist phyla in-
cluding amoebae, colonial chrysophytes and ciliates are known
but have yet to be described (SMALL & GROSS 1985; M. B.,
pers. obs.). In contrast, other marine habitats, such as sedi-
ments from shallow waters to the deep sea harbor far more di-
verse meiofauna. For example gnathostomulids, gastrotriches,
loriciferans, tardigrades, and small-sized representatives of hy-
droids, bryozoans, nemertines, rotifers, or gastropods have not
yet been discovered (HIGGINS & THIEL 1988; GIERE 1993).
All together, two foraminiferan, seven nematode, nine ost-
racod, two acari, two cumacean, six tanaidacean, two isopod,
and 78 copepod species are included in this edition. Only the
Copepoda are known to a greater extent (IVANENKO & DEFAYE,
this volume). Currently, meiofauna species contribute to about
20% of the total diversity at vents. However, at large, the diver-
sity of meiofauna is unknown. There are entire mid-ocean rid-
ges or back-arc basins known for a relatively long time, in which
not a single species of ostracod, tanaidacean, mite, or nematode
is described. Most known species are reported from a single site.
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
known. The few ostracod species described exclusively come
from the eastern Pacific (DEEVEY 1969; KORNICKER 1969, 1991,
KORNICKER & HARRISON-NELSON 2005; MADDOCKS 2006). The
few tanaidaceans and the two isopod species were collected at
the site Lucky Strike at the Mid-Atlantic Ridge (CUNHA &
WILSON, in press; LARSEN et al., in press); tanaidaceans were al-
so observed at the site Rainbow (M. Segonzac, pers. comm.)
Only one mite is described from Lau and North Fiji Back-Arc
Basins (KRANTZ 1982) and the second species comes from the
Mid-Atlantic Ridge (BARTSCH 1990). Similarily, one cumacean
and one foraminiferan species each are decribed from the East
Pacific Rise (BRÖNNIMANN et al. 1998; CORBERA this volume),
the second species each come from the Mid-Atlantic Ridge
(KAMENSKAYA et al. 2002; CORBERA this volume). The few des-
cribed free-living nematode species exclusively were collected at
sedimented vent sites from the East Pacific Rise, the Guaymas
Basin, and the Lau and the North Fiji Back-Arc Basins (DE-
CRAEMER & GOURBAULT 1998; VERSCHELDE et al. 1998).
To date there is not a single site for which the entire meio-
fauna community structure is known. The picture emerging
from the few meiofauna studies points to a vent meiofauna,
which is low in abundance and diversity (DINET et al. 1988;
SHIRAYAMA 1992; VANREUSEL et al. 1997; TSURUMI et al. 2003).
Comprehensive studies are urgently needed not only to under-
stand the community structure of meiofauna but also to provi-
de insight in the biodiversity of the entire vent community and
their trophic interactions.
Denisia 18 (2006): 27–28
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

28
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Hydrothermal vent parasites
Currently, very few macroparasites are known from hydro-
thermal deep-sea vents. Only four copepods, Genesis vulcanoc-
topusi (East Pacific Rise, LOPEZ-GONZALEZ et al. 2000), Cholidya
polypi (Endeavour Segment, HUMES & VOIGHT 1997), Ceuthoe-
cetes aliger and Rimitantalus hirsutus (Galapagos Spreading Cen-
ter, HUMES & DOJIRI 1980), one leech Bathybdella sawyeri (East
Pacific Rise, BURRESON 1981; BURRESON & SEGONZAC, in
press), one acanthocephalan Hypoechinorhynchus thermaceri
(East Pacific Rise, BURON 1988), and one nematode, Moravec-
nema segonzaci (Mid-Atlantic Ridge, JUSTINE et al. 2002) have
been reported from these sites. Significantly, each of these spe-
cies was new to science. At least one monogenean and two di-
geneans are currently being described by various specialists
from material collected on crabs and fishes collected from sou-
thern East Pacific Rise sites and each of these also appear to be
previously unreported species.
Because the concentration of potential hosts in a vent eco-
system could possibly increase parasite transmission and diversi-
ty, BURON & MORAND (2004) hypothesized that the low diversi-
ty of macroparasites presently reported from these sites is likely a
reflection of these organisms having been overlooked by vent
biologists due to their inconspicuous nature. In support of this
idea, the use of molecular tools has shown the presence of a high
diversity of parasitic protists genetically linked to apicomplexans,
perkinsozoans, syndiniales, and kinetoplastids (MOREIRA & LO-
PEZ-GARCIA 2003). Additionally, Rickettsia-like inclusions have
also been found in numerous limpets from various vents (TERLIZ-
ZI et al. 2004), and in mussels from Snake Pit and Logatchev si-
tes (WARD et al. 2004) are further indications that parasite di-
versity is vastly underreported from these important sites.
1: Hirudinea Bathybdella sawyeri from East Pacific Rise: 17°S; cruise
Biospeedo (Ifremer, CNRS).
I. DE BURON & M. SEGONZAC
In brief, most often parasites must carefully looked for in or-
der to be noticed. Hosts are various and diverse and range from
the smallest of copepods to the largest of fish.
Since numerous parasites have a complex life cycle invol-
ving more than one host, a better knowledge of the parasite
vent fauna would help researchers to understand the trophic
structure present at each vent. Because trophically transmitted
parasites are in many cases known to alter intermediate host
behavior to increase capture by definitive hosts (LAFFERTY &
MORRIS 1996) these organisms may play significant roles in in-
creasing or directing the flow of energy into higher trophic le-
vels at the vents. For example, acanthocephalans at the East
Pacific Rise use fish as definitive hosts and appear to be trans-
mitted via amphipod ingestion (I. B., pers. obs.): evidence from
other systems indicates that such infected amphipods exhibit
altered behavior (BAUER et al. 2005). Significantly, at the At-
lantic sites these crustaceans were found to contain nematode
larvae of the ascarid genus Hysterothylacium although the adults
were not found (A. Petter, unpublished data).
Furthermore, non-trophically transmitted parasites present
at vents may also pose interesting problems for researchers. For
instance, the leech Batybdella sawyeri, living among Riftia pachyp-
tila and mussel Bathymodiolus thermophilus at the Galapagos Spre-
ading Center, raises an interesting specificity problem: although
a parasite of fish, it has never been found on such hosts but was
observed on vent crabs and bivalves from the southern East Pa-
cific Rise (BURRESON & SEGONZAC, in press). Because of the well
known specificity of many parasite-host relationships the recog-
nition and identification of vent parasites may play an important
role in deciphering vent colonization. In this regard it is impor-
2: Nemertea Carcinonemertes sp., semi-parasite on
bythograeid crabs from Lau Basin; cuise TUIMO6MV
(MBARI); by G. Rouse.
Denisia 18 (2006): 29–30
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

tant to note the presence of carcinonemertids (nemertean
worms) at vents. These worms are known to feed on the eggs of
shallow water decapods and were discovered on several vent crab
species (SHIELDS 2001). Given the depth (1800-2400 m) and iso-
lated nature of vent communities, the origin of vent colonization
by carcinonemertids has been raised and it has been speculated
that infections were acquired from a more shallow-water host
(see SHIELDS 2001 for review). Such egg predators were recently
collected on several bythograeid crab species from western back-
arc basins, and northern and southern East Pacific Rise (J.
Shields & M. Segonzac, unpublished data). Moreover, because of
the isolated nature of the vent ecosystems these sites will likely
provide excellent sources of material for the study of the proces-
ses of co-evolution in a relatively simple trophic system.
In summary, the study of parasitism in isolated vent com-
munities should be emphasized not just because it would furt-
her increase our understanding of the unique parasite fauna
present at these sites, but also because it would allow us to ad-
dress and answer important questions regarding the coloniza-
tion of vents and vent community structure as well as allow us
to identify and recognize new strategies for parasite survival.
Parasites may be located almost anywhere on or in a host.
Ideally, parasites should be isolated from their host while still
fresh and then prepared in a specific manner dependent on the
taxon collected (see, for example, PRITCHARD & KRUZE 1982
for macroparasite fixation techniques). Importantly, parasites
collected from formaldehyde fixed hosts, such as museum spe-
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.

30
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Granuloreticulosa, Foraminifera

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

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

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

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

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

Porifera

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

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

J. VACELET Denisia 18 (2006): 35

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

The cladorhizids are generally small sponges with a mor-
phology rather unusual in sponges. They are erect, with a pin-
nate or symmetrical shape, and their aquiferous system, long
considered as a diagnostic characteristic of the phylum Porifera,
has been discarded in most genera. A special mention, howev-
er, should be made for the genus Chondrocladia, in which a
canal system and choanocyte chambers are maintained, but
serve to inflate large spheres on which the prey is captured. The
prey, mostly small crustaceans, are passively captured by means
of hook-like cheloid microscleres lining the surface of the
sponge and catching the setae or appendages of various inver-
tebrates, acting as a Velcro cover. In the absence of a digestive
cavity, cells act individually to digest the prey.
The classification of the carnivorous sponges is presently
rather problematic (HAJDU & VACELET 2002). They all belong
to the order Poecilosclerida, and most are classified in the fam-
ily Cladorhizidae, with several genera dinstinguished mainly by
the microsclere spicules. These microscleres, however, belong
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.
These deep-sea sponges are highly fragile. The hexactinel-
lids, although supported by a highly developed siliceous skele-
ton, have very thin, partly syncytial living tissue which is very
difficult to preserve properly. They must be preserved in good
histological or cytological fixatives as soon as possible after col-
lection. The carnivorous sponges have very fragile appendages
that are generally only partially preserved during collection.
Observation of their morphology from manned submersibles or
ROVs is important. For instance, a long time was needed to re-
alize that a mysterious organism with pedunculate, translucent
spheres actually corresponds to the genus Chondrocladia as it
was known from preserved specimens, in which the spheres
were contracted. Their histology and reproduction is poorly
known and could be studied only from specimens preserved in
formalin or fixatives for electron microscopy. The present un-
certainties in the distinction of evolutionary lines in carnivo-
rous sponges require more information on their molecular char-
acters, and the preservation of fragments in alcohol is highly
desirable.

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

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

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

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

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

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

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

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:

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae
Chondrocladia lampadiglobus VACELET, in press
Size: Up to approximately 50 cm high, with inflated spheres 3-
5 cm in diameter.
Color: White in life, yellowish white to clear brown in alcohol.
Morphology: Large stalked sponge, composed of a rhizoid fixa-
tion system, a cylindrical stalk ending in an enlarged, ovoid
body from which radiate in all directions secondary branches,
each ending in translucent sphere in the living animal, in an ir-
regular swelling including crustacean debris on the preserved
specimen. Aquiferous system present, with large choanocyte
chambers and canals ending in the inflatable spheres. Skeleton:
stalk and branches made of large fusiform styles longitudinally
arranged, covered with a feltwork of rugose tylostyles; terminal
swellings made of smaller styles with an outer cover of numer-
ous microscleres. Spicules: fusiform styles, 700-4750 x 15-75 µm
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.
Vrijenhoek © MBARI.
J. VACELET & M. SEGONZAC
140 µm, alae 25 µm long, and 20-32 µm, alae 10-11 µm long;
sigmas 45-120 x 2-3 µm.
Biology: The collected specimen was rooted in sediment be-
tween pillow lava, near active hydrothermal sites, but in an
area still with low density of animal life. Sponges of similar
morphology have been often observed on various sites of the
East Pacific Rise, either rooted in sediment or attached to pil-
low lava, always at some distance from the rich animal com-
munities of the active hydrothermal sites. Their identification
to C. lampadiglobus cannot be ascertained from external mor-
phology alone, and several species may be present in this large
geographic zone. Carnivorous mode of feeding.
Distribution: East Pacific Rise: collected at 17°S. Sponges of
similar morphology observed in various sites extending from
23°S to 13°N, 2600-3000 m deep.
2A: Holotype; scale bar 17 mm; B: Tylostyle of the
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).
Denisia 18 (2006): 42–43
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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

Porifera, Demospongiae, Poecilosclerida, Guitarridae

Euchelipluma pristina TOPSENT, 1909

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

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Cnidaria, Hydrozoa, Leptolida (= Hydroida)
The leptolid fauna of hydrothermal vents of oceanic ridge
sites is still poorly investigated and understood. Fragmentary
colonies of leptolids from oceanic ridges in the Pacific have so
far made it possible to identify Zygophylax cervicornis (NUTTING,
1905), Zygophylax sp. and Halecium tenellum HINCKS, 1861.
While from vents on the Mid-Atlantic Ridge only a single an-
thoathecate leptolid, Candelabrum serpentarii SEGONZAC &
VERVOORT, 1995 can be considered to be a true companion of
the vent community. A second species, Candelabum phrygium
(FABRICIUS, 1780) is a North Atlantic deep-sea inhabitant oc-
casionally found associated with the hydroythermal vent fauna.
Another anthoathecate, Bouillonia sp. was found on the
Mid-Atlantic Ridge in the Gulf of Guinea area; it probably rep-
resents the deep-water leptolid community without direct
affinities to hydrothermal vents (SVOBODA, STEPANJANTS &
LJUBENKOV, in press). Photographs taken during surveys of Mid-
Atlantic vent areas show a third species of anthoathecate, oc-
casionally in great numbers; it may represent a species of Tubu-
laria or Ectopleura. There is presently no material available to
substantiate that guess. A probably undescribed species of Hy-
dractinia has also been observed.
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
Collecting activities at or near Mid-Atlantic vents areas
have currently provided material for proper identification of 3
anthoathecates and 23 leptothecates; 4 more leptothecates
could only be identified to the genus (CALDER & VERVOORT
1998). One more species (Hydrallmania falcata) was dead when
collected and accidentally introduced; all remaining leptothe-
cates belong to the deep-water fauna and are only party re-
stricted to the Atlantic.
Because of the technique used to collect animals at ridge
sites, only rarely do complete animals or colonies become avail-
able for scientific study. Anthoathecates in particular are hard
to collect and they must be narcotized prior preservation. So
far, only a single complete animal of Candelabrum serpentarii has
been obtained, in addition to many incomplete individuals,
which have been studied. Anthoathecates are best preserved in
6% formalin (after prior relaxation by adding MgSO4 solution
to the seawater), but Leptothecates get very brittle in formalin
and are best preserved in 70% ethanol. These preservation
techniques only apply to material for routine taxonomic inves-
tigations; more sophisticated methods are necessary for modern
DNA techniques.
Denisia 18 (2006): 48
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Cnidaria, Hydrozoa, Leptolida, Leptothecata, Halopterididae
Polyplumaria flabellata G.O. SARS, 1874
Size: Height of colony up to 350 mm, spread almost equal.
Morphology: Stolonal mass of fine tubules supporting a rigid,
pinnately branched colony of very regular structure. Stem poly-
siphonic and repeatedly branched, branches in opposite pairs;
branches originating from secondary tubes; primary axis visible
in polysiphonic parts in front of bundle of secondary tubules,
divided by oblique nodes into internodes of varied length, each
internode with one to five apophyses, alternately directed
obliquely upwards and to left and right. Primary and secondary
tubules with nematothecae, each apophysis with a ‘mamelon’
and three nematothecae; in addition two to four nematothecae
between two successive apophyses. Hydrocladia about 3 mm
long, supported by apophyses, slightly curved, first internode
athecate, with one nematothecae; all following internodes the-
cate, with one proximal, a pair of flanking, one distal and sev-
eral additional and variously distributed nematothecae on dis-
tal part of internode. All nematothecae bithalamic with deeply
scooped adcauline wall. Hydrotheca cylindrical with almost
parallel walls, adcauline wall adnate for half its length, length
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
axis of hydrotheca at 45∞ to length axis of internode; hy-
drothecal rim circular and smooth, perpendicular to hydrothe-
cal length axis; lateral (flanking) nematothecae on short
apophyses near hydrothecal axil. Hydrocladia occasionally
forked, secondary branch springing from base of first hydrothe-
ca. Depth of hydrotheca 140-190 µm, diameter at rim 135-160
µm. Female and male gonothecae of same shape, curved ovoid,
attached to apophyses by means of short pedicel; basal part of
gonothecae with 6-9 nematothecae. Aperture circular, laterally
disposed, turned towards hydrocladium, closed by circular lid.
Length of gonothecae 610-750 µm; diameter 340-400 µm. Ne-
matocysts: Not studied in detail.
Biology: Usually on soft bottoms (off vent areas). Serves as host
for many epizoic leptolid species.
Distribution: Mid-Atlantic Ridge: near Menez Gwen, 997-696
m. Moderately deep to deep waters of tropical, temperate, and
northern parts of the eastern Atlantic.
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).
Denisia 18 (2006): 53
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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

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

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

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

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

1 left: Distal part of stem with hydrothecae, nematothecae and insertion of two hydrocladia, scale bar 0.25 mm; top row left:
Axillary hydrotheca with nematotheca, scale bar 0.1 mm; middle: Hydrotheca and nematotheca, scale bar 0.1 mm; right: End 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

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

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

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

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

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

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

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

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

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

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.
Morphology: Erect colony, rising from creeping stolon. Proxi-
mal parts of stem and branches polysiphonic; branching irregu-
lar, basal part of stem ahydrothecate. Monosiphonic parts of
colony composed of long, slender, geniculate internodes sepa-
rated by nodes sloping in alternate directions. Hydrothecae
950-1.200 µm deep, greatest diameter about 500 µm, placed at
end of internode, one-fifth of abcauline wall adnate, curved
outwards, slender, abcauline wall convex, abcauline wall
bulging in proximal third. Hydrothecal rim not everted, occa-
sionally renovated, with three prominent cusps, one abcauline,
two lateral. Three triangular opercular plates attached in em-
bayments between marginal cusps, no renovations of opercu-
ameter at two-thirds of distance from top, with about seven
transverse ribs, petering our proximally. Orifice at end of a short
tube in middle of apical ‘field’ formed by distalmost rib. Pedicel
quite short, attaching gonothecae to internode at hydrothecal
base. Nematocysts: Not studies in detail.
Biology: On solid substrates in deep ocean water.
Distribution: Mid-Atlantic Ridge: Lucky Strike, Sintra site.
Deep water of Atlantic (Bay of Biscay) and Pacific (off Chile
and SE of New Caledonia).

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

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

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

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

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

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

Cnidaria, Scyphozoa, Stauromedusae, Eleuthrocarpidae

Lucernaria janetae COLLINS & DALY, 2005 “stalked jellyfish”

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae
Actinostola VERRILL, 1883
Size: Diameter of the pedal disc, 90 mm; mouth, 70 mm; col-
umn height, 60 mm; length of contracted tentacles, 11 mm.
Color: Column, tentacles, pharynx, and mesenteries yellowish
white in situ.
Morphology: Column cylindrical, with conspicuous mesogloeal
tubercles arranged regularly in longitudinal rows; tentacles
arranged in four or five cycles, with an apical orifice, aboral side
of tentacles often thickened at the base; sphincter weak,
mesogloeal, thus tentacles can not be completely retracted;
longitudinal musculature of tentacles mesogloeal; mesenteries
arranged according to Actinostola-rule; oldest cycles of mesen-
teries sterile.
Remarks: DOUMENC & VAN PRAËT (1986) discussed the possi-
ble specific identity of the single specimen examined without a
Reference:
DOUMENC D. & M. VAN PRAËT (1986) Oceanol. Acta 8: 61-68.
P. LÓPEZ-GONZÁLEZ & M. SEGONZAC
definitive conclusion. Although in the figure 1 of that paper,
the name A. callosa is included, it was probably a mistake dur-
ing the editorial process of the manuscript. DOUMENC & VAN
PRAËT (1986: 63) also reported the presence of four cycles of
tentacles, this number of cycles corresponding to 48 tentacles
(6+6+12+24). However, the number of tentacles seen in the in
situ photographs suggest five cycles. Additional material is
needed to study the specific pertinence and other anatomical
details.
Biology: As other sea anemones, probably feeds on caught prey,
with a relative wide prey size-range. All other biological aspects
are unknown. Often observed within the gravel, attached at
the top of lava pillars.
Distribution: East Pacific Rise: 13°N.
2: In situ specimen; cruise
Hot 96 © Ifremer.
1: In situ specimen;
cruise Hope 99 © Ifremer.
Denisia 18 (2006): 66
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Actinostolidae
Cyananthea hydrothermala DOUMENC & VAN PRAËT, 1986
Size: Diameter of pedal disc, 10 mm, height, 35 mm; tentacles,
ca. 25 mm.
Color: Whitish in situ, column and tentacles dark brownish,
mesenteries pale beige.
Morphology: Column smooth, with distal margin distinctly
marked; tentacles occupying most oral disc, reaching near to
the mouth; more mesenteries distally than proximally; sphinc-
ter mesogloeal, strong, on the endodermic face of the
mesogloeal layer; retractor musculature diffuse and weak; longi-
tudinal musculature of tentacles ectodermic; radial musculature
of the oral disc ecto-mesogleal.
1: Two specimens taken onboard from the East Pacific Rise: 13°N, cruise Phare
© Ifremer.
Reference:
DOUMENC D. & M. VAN PRAËT (1986) Oceanol. Acta 8: 61-68.
P. LÓPEZ-GONZÁLEZ & M. SEGONZAC
Remarks: This species was insufficiently described to be cor-
rectly diagnosed or compared with other actinostolid genera
and species. The familial pertinence was tentative after the sup-
posed absence of acontia in the single fragment available to
study by DOUMENC & VAN PRAËT (1986). Material collected in
the type locality should be re-described in order to precise the
taxonomic status of this genus and species.
Biology: As other sea anemones, probably feeds on caught prey,
with a relative wide prey size-range. All other biological aspects
are unknown. The association with bacterial symbionts has not
been demonstrated. Lives on the walls of the black smokers at
temperatures of 12-20°C and on sulphide rocks.
Distribution: East Pacific Rise: 13°N.
2: In situ specimen taken from the East Pacific
Rise: 13°N, among siboglinids Riftia pachyptila
and limpets Lepetodrilus elevatus, cruise Phare
© Ifremer.
Denisia 18 (2006): 67
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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.

P. LÓPEZ-GONZÁLEZ & M. SEGONZAC Denisia 18 (2006): 69

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

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

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

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.

P. LÓPEZ-GONZÁLEZ & E. RODRÍGUEZ Denisia 18 (2006): 71

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

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

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

Cnidaria, Anthozoa, Hexacorallia, Actiniaria, Hormathiidae

Chondrophellia cf. coronata (VERRILL, 1883)

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

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

Mollusca, Solenogastres

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

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

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

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

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

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

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
2: H. cf. acredema, in vivo specimen from East Pacific Rise:
9°N, site Tica; by M. Bright.
4: Helicoradomenia sp. 2, body length 2 mm, in ventral
view,
w anterior end at left showing slit-like dorsofrontal sen-
sory pit and atrio-buccal area, pedal groove (foot) wide;
from East Pacific Rise: 18°S; cruise Biospeedo © Ifremer.
6: Helicoradomenia sp. 3, body length 2.3 mm, lateral view
(anterior at left), mantle with incrustation by yellow-orange
deposits; from East Pacific Rise: 21°S; cruise Biospeedo
© Ifremer.
Reference:
SCHELTEMA
L A.H. & A.M. KUZIRIAN (1991) Veliger 34: 195-203.
3: Helicoradomenia sp. 1, in vivo specimen from East Pacific Rise:
9°N, site Tica; by M. Bright.
5: Simrothiellidae gen. et sp. 1, body length 3 mm, lateral view,
anterior end at left; mantle in part with dorsal incrustation of or-
ange iron oxide; from East Pacific Rise: 13°N, site PP Hot 3 (Elsa);
cruise Phare © Ifremer.
7: Two specimens of Simrothiellidae sp. in vivo from North Fiji
Back-Arc Basin, site White Lady; cruise TUIM06MV (June 2005,
MBARI) © G. Rouse.
79
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

Mollusca, Polyplacophora, Neoloricata, Leptochitonidae

Leptochiton tenuidontus SAITO & OKUTANI, 1990

portion of each valve, sutural laminae roughly sub-triangular

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Mollusca, Gastropoda
Currently about 60 genera and more than 100 species of gas-
tropods have been recognized in hydrothermal communities in
the Indo-Pacific, Indian and Atlantic Oceans. That makes the
gastropods the most species-rich group of vent animals thanks to
all researchers who have sent specimens to taxonomists for iden-
tification. The gastropods are probably also the group where
knowledge on distribution and zoogeography is the best.
Although the present handbook aims to present a fairly
complete overview with diagnostic information and illustra-
tion, it is important to stress several points:
(1) The identification of several genera of small gastropods
is difficult
f and is a matter of specialist work.
(2) The limpets can be assigned to genus based on shell and
external morphology only, but many can be confusingly similar
on superficial examination by an untrained eye. It is sometimes
even easier to identify limpets to species than to genus.
(3) For any work deemed to have results of more than tem-
porary importance and where species identification is involved,
1: Ifremeria nautilei from Kilo Moana, Lau Back-Arc Basin, TU-
IM 07 cruise © C.R. Fisher.
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
it is recommended that voucher specimens are deposited in a
recognized scientific museum collection. Many natural history
museums readily accept such deposits. For morphological work
formalin-fixed specimens are preferred, although also ethanol-
preserved specimens can be used. Frozen specimens are usually
too macerated to show details in tentacles and epithelia. As al-
ways when calcareous shells are involved, do not forget to
buffer the formalin, for example with a tea spoon of borax per
litre 4% formalin.
(4) Beginning genetic work has revealed existence of isolat-
ed genotypes or populations within some of the previously rec-
ognized species. In some species this is accompanied by morpho-
logical differences and/or geographic separation (e.g. Lepetodrilus
fucensis JOHNSON et al., in press). In others (e.g. Alviniconcha hess-
leri) there are no noticeable differences in morphology and their
geographic distribution is not easily explicable. It should be re-
membered that distribution, dispersal and speciation of deep-sea
animals still is “terra incognita“ and should be explored.
(5) This manual may give the impression that the fauna is ad-
equately known. However, many vent inhabitants are still 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-
tent be host specific since their appearance differs
f between host
species. They have only been observed in vent environments.
Denisia 18 (2006): 82
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

Mollusca, Gastropoda, Patelligastropoda, Lepetopsoidea, Neolepetopsidae

Neolepetopsis MCLEAN, 1990 “symmetrical limpets”

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 94

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

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

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

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

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.

A. WARÉN & P. BOUCHET & R. VON COSEL Denisia 18 (2006): 96

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

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

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

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

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

Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Sutilizonidae

Temnocinclis euripes MCLEAN, 1989 “slit-limpet”

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

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

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

Mollusca, Gastropoda, Vetigastropoda, Lepetodriloidea, Sutilizonidae

Temnozaga parilis MCLEAN, 1989

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

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.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 100

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

Mollusca, Gastropoda, Vetigastropoda, Trochoidea, Chilodontidae

Bathymargarites symplector WARÉN & BOUCHET, 1989

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

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

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

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

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

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.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 103

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

Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Cyathermia naticoides WARÉN & BOUCHET, 1989

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

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

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

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

Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Melanodrymia HICKMAN, 1984

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

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

Mollusca, Gastropoda, Neomphalina, Neomphaloidea, Neomphalidae

Neomphalus fretterae MCLEAN, 1981

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

Peltospira MCLEAN, 1989 “tapersnout limpets”

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

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

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

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

Mollusca, Gastropoda, Neomphalina, Peltospiroidea, Peltospiridae

“scaly foot gastropod”

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

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

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

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.

A. WARÉN, P. BOUCHET & R. VON COSEL Denisia 18 (2006): 121

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

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

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

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

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

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

1: E. desbruyeresi; top left to right: Abapertural, lateral and apertural view; bottom: Lateral views; by R. von Cosel & P. Lozouet.

P. BOUCHET Denisia 18 (2006): 124–125

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

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

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

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

Mollusca, Gastropoda, Caenogastropoda, Elachisinidae

Laeviphitus desbruyeresi WARÉN & BOUCHET, 2001

Size: Shell height up to 1.8 mm.
Morphology: Shell very small, not especially fragile, about four
whorls, with shallow suture, narrow umbilical crevice and an-
teriorly bluntly rounded and posteriorly slightly pointed aper-
ture, smooth teleoconch and distinctly demarcated and cancel-
late protoconch. Shell surface usually with strong ferrugineous
deposits. Operculum thin, yellowish brownish, paucispiral.
Biology: Among Bathymodiolus and in sediment. Genus known
from vents and driftwood (unpubl.). Development with plank-
totrophic larvae.
Distribution: Mid-Atlantic Ridge: Menez Gwen to Rainbow.
Genus also known from Japan (Laeviphitus japonicus OKUTANI,
FUJIKURA & SASAKI, 1993) and the Marianas (unpubl.). Larvae
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

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

Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae

Alviniconcha hessleri OKUTANI & OHTA, 1988

pertrophied and specialized gill cells contain endosymbiotic

Size: Shell height up to 85 mm.
Color: Yellowish.
Morphology: Shell globose, rather elastic, spectacularly orna-
mented with regularly spirally arranged periostracal hairs.
Aperture with a shallow sinus in front. Operculum horny and
ovo-quadrate.
Biology: Genus endemic in vents. Usually in stacks around
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-
chemoautotrophic bacteria of the sulphur cycle. Reduced di-
gestive tract as compared to related species of the same family.
Distribution: A single morphospecies in the Western Pacific:
Mariana, North-Fiji and Lau Back-Arc Basins; Indian Ocean:
Rodriguez Triple Junction, Kairei hydrothermal field. Never-
theless, A. hessleri from the type location is genetically different
from the three Alviniconcha populations collected at other
places.

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

A. WARÉN Denisia 18 (2006): 128

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

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

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

Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae

Ifremeria nautilei BOUCHET & WARÉN, 1991

Alviniconcha, the gill and circulatory system are hypertrophied

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

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

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

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

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

Mollusca, Gastropoda, Caenogastropoda, Loxonematoidea, Provannidae

Provanna DALL, 1918

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

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

Mollusca, Gastropoda, Caenogastropoda, Rissoidae

Alvania cf. stenolopha BOUCHET & WARÉN, 1993

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

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

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

Mollusca, Gastropoda, Caenogastropoda, Vitrinellidae

Neusas marshalli SYKES, 1925

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

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

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

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

Mollusca, Gastropoda, Heterobranchia, Orbitestellidae

Lurifax vitreus WARÉN & BOUCHET, 2001

cylindrical tentacles. Foot with well-demarcated propodium, no

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

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

Mollusca, Gastropoda, Heterobranchia, Xylodisculidae

Xylodiscula analoga WARÉN & BOUCHET, 2001

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

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

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

Mollusca, Gastropoda, Prosobranchia, Conoidea, Turridae

Phymorhynchus DALL, 1908

Species
P carinatus WARÉN & BOUCHET, 2001
P.
P. hyfifluxi BECK, 1996
P. major WARÉN & BOUCHET, 2001
P moskalevi SYSOEV & KANTOR, 1995
P.
P. ovatus WARÉN & BOUCHET, 2001
P. starmeri OKUTANI & OHTA, 1993
P wareni SYSOEV & KANTOR, 1995
P.
Distribution
Mid-Atlantic Ridge: 23-15°N
North Fiji Basin
East Pacific Rise: 13-9°N
Mid-Atlantic Ridge: 26-23°N
Mid-Atlantic Ridge: 37-15°N
North Fiji and Manus Back-Arc Basins; may be
common in both vent and non-vent areas in up-
per abyssal bottom
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

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Mollusca, Bivalvia
The Solemyidae are represented at hydrothermal vent or
cold seep biotopes by the genus Acharax. Eight named species
as well as several still unnamed species are known, but only a
single species has been collected at hydrothermal vents (Lau
Back-Arc Basin). Representatives of this genus can grow to rel-
atively large sizes between 10 and 22 cm. They live deeply
buried in soft sediment. At least some species are characterized
by the absence of a digestive tract, and their nourishment relies
exclusively on their chemosynthetic symbiotic bacteria.
Of the large mussels living at hydrothermal vents or cold
seeps, 20 species have been currently described, 18 of them in
the genus Bathymodiolus. Of these, 10 are known from hy-
drothermal vents and are covered herein. Several other large
mussels are still awaiting description. All known Bathymodiolus
have a larval shell of about 0.5 mm or smaller with set-off pro-
toconch 1; in general, the protoconch has a distinct rose color.
The Vesicomyidae are a rather diverse family in size, shape
and species number: at the moment 84 recently named species
are known, but there are still numerous undescribed species
currently under study by several authors. The family comprises
on one hand small species (genus Vesicomya sensu stricto),
which occur in the deep sea but are not necessarily associated
with cold seep and/or hot vent habitats and on the other hand,
most of the numerous medium-sized or large to very large
species are confined to reducing sediments, cold seeps, hydro-
1: Bathymodiolus azoricus from Rainbow, Mid-Atlantic Ridge, Atos cruise © Ifremer.
R. VON COSEL
carbon seeps or hydrothermal vents. In the past, species of Vesi-
comyidae have been assigned to different genera or subgenera
and many of the large species are now commonly placed in the
genus Calyptogena (sensu lato). Recently however, some au-
thors have revised tentatively all Vesicomyidae in the sole
genus Vesicomya (in a broad sense) pending future supra-specif-
ic revisions based mainly on molecular research. Anyway, shell
and soft part morphology remain important and key characters
are among others hinge dentition, shell size and shape and pres-
ence or absence of a well-developed pallial sinus. Herein, the
genus Calyptogena is maintained. The assignment of a vesi-
comyid species to Calyptogena differs with authors, and the
supra-specific systematics of the Vesicomyidae is still far from
settled. Most of the larger species have been described from
cold seep biotopes, but of the 28 named species currently treat-
ed as Calyptogena, seven were found inhabiting also (or exclu-
sively) hydrothermal vents. Not very much is known of the lar-
val development of vesicomyid clams, but many are known to
have lecithotrophic development.
The Pectinidae are represented by two species, Bathypecten
vulcani, present at the Galapagos Spreading Center, the north-
ern East Pacific Rise 13°N and the southern East Pacific Rise
38°S and Sinepecten segonzaci, from the Manus Back-arc Basin.
More sampling need of these small bivalves, difficult to see in
situ, to understand the processes of their distribution.
Denisia 18 (2006): 141
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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.

R. VON COSEL Denisia 18 (2006): 143

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Mollusca, Bivalvia, Heterodonta, Veneroida, Vesicomyidae
Calyptogena magnifica BOSS & TURNER, 1980
Size: Shell length up to 263 mm.
Morphology: Shell very large, variable, oval to elongate-oval in
outline, inequilateral and equivalve, usually without gape;
valves white, entirely aragonitic; periostracum present, but fre-
quently eroded away on older portions of shell; umbos prosogy-
rous, sometimes partially enrolled; lunule and escutcheon vari-
able, present or absent; ligament external, opisthodetic, and
parivincular; foot strong and rugose, with byssal gland.
Biology: At hot vents only. The life habit of the adult is semi-
epibiotic on bare basalt, living in cracks and crevices 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.
References:
ARP A. J., CHILDRESS J.J. & C.J. FISHER (1984) Physiol. l Zool. 57: 648-662.
BOSS K. & R. TURNER (1980) Malacologia 20: 161-194.
CHILDRESS J.J., FISHER C.R., FAVUZZI J.A. & N.K. SANDERS (1991) Physiol. Zool.
64: 1444-1470.
FATTON E. & M. ROUX (1981) C. R. Acad. Sc. Paris, Série III 293: 63-68.
FIALA-MÉDIONI A. & C. METIVIER (1986) Mar. Biol. 90: 215-222.
FISHER C. et al. (1988) Deep-Sea Res. 35: 1811-1831.
R. LUTZ & R.C. VRIJENHOEK
often encountered living within basaltic rubble associated with
such warm cracks and crevices. The presence of this species is
correlated with elevated levels of hydrogen sulphide. The soft
tissue is coloured dark red when retrieved living due to the
presence of intracellular hemoglobin. The thick and large gills
contain sulfur oxidizing chemoautotrophic symbionts. The
large rugose foot is often seen protruding when the clams are
viewed in life position.
Distribution: Entire Northern East Pacific Rise and Southern
East Pacific Rise: 21°N to 22°S; Galapagos Spreading Center.
3: Exterior, interior and dorsal view of left
valve, specimen from 21°N;
by R. von Cosel & P. Lozouet.
HURTADO L.A., MATEOS M., LUTZ R.A. & R.C. VRIJENHOEK (2003) Appl. Envi-
ronm. Microbiol. 69(4): 2058-2064.
KENNISH M. & R. LUTZ (1992) Rev. Aquat. Sci. 6: 29-66.
KOJIMA S., FUJIKURA K. & T. OKUTANI (2004) Mol. Phylogen. Evol. 32: 396-406
[400].
LUTZ R.A., FRITZ L.W. & R.M. CERRATO (1988) Deep-Sea Res. I 35: 1793-1810.
RIO M. & M. ROUX (1984) C. R.Acad Sci. Paris, Série II 299: 167-172.
VRIJENHOEK R. et al. (1994) Deep-Sea Res. 41: 1171-1189.
Denisia 18 (2006): 145
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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.

R. VON COSEL Denisia 18 (2006): 146

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus japonicus HASHIMOTO & OKUTANI, 1994

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

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

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

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.

R. VON COSEL Denisia 18 (2006): 156

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

Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus platifrons HASHIMOTO & OKUTANI, 1994

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

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

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

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

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.

J. HASHIMOTO Denisia 18 (2006): 159

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

Mollusca, Bivalvia, Pteriomorphia, Isofilibranchia, Mytilidae

Bathymodiolus thermophilus KENK & WILSON, 1985

Size: Shell length up to 18.4 cm.
Morphology: Shell smooth, modioliform, with subterminal um-
bones; periostracum present and straw-yellow to brown in
colour; external surface lacks sculpture and is dull white be-
neath periostracum; hinge edentulous; ligament opisthodetic,
paricincular, strong, extending most of the length of the dorsal
margin; well developed byssus. Valvular siphonal membrane
long and stretching towards ventrally; on anterior part fusion of
inner mantle fold reaching ventrally to 1/3 shell length, to-
gether with siphonal membrane leaving rather short byssus
opening in the middle.
Biology: The life habit of the adult is epibiotic on bare basalt
and other hard substrates (e.g. tubes of vestimentiferans) asso-
ciated with deep-sea hydrothermal vents. The presence of this
species is correlated with elevated levels of hydrogen sulfide.
Endemic to vents. Protoconch 0.4 mm in length, indicative of
a planktotrophic larval stage with a high dispersal capability.
Paired ctenidia consist of inner and outer demibranchs, each
with descending and ascending lamellae. The gills contain sul-
phur-oxidizing chemoautotrophic symbionts. A commensal
polychaete (Branchipolynoe symmytilida) is frequently found
within the mantle cavity of the mussel.
Distribution: Galapagos Spreading Center, East Pacific Rise:
13°N to 22°S. At the Pacific-Antarctic Ridge (31°S and 38°S)
another species of Bathymodiolus was collected in 2005 (R. 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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

Mollusca, Bivalvia, Pteriomorphia, Eupteriomorphia, Pectinidae

Bathypecten vulcani SCHEIN-FATTON, 1985

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

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

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

Mollusca, Bivalvia, Pteriomorphia, Eupteriomorphia, Pectinidae

Sinepecten segonzaci SCHEIN, in press

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

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

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

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

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

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

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

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

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

Mollusca, Cephalopoda, Octopoda, Grimpoteuthidae

Grimpoteuthis ROBSON, 1932

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

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

Mollusca, Cephalopoda, Octopoda, Octopodidae

Vulcanoctopus hydrothermalis GONZÁLEZ, GUERRA, PASCUAL & BRIAND, 1998

Size: Up to 56 mm mantle length, 235 mm total length and 45
g body weight.
Color: In situ, white, eyes black.
Morphology: Body semi-translucent with a muscular consisten-
cy; mantle pear-shaped and posteriorly mitre-like; presence of a
large white body (which covers the eye, the optic nerves and
the optic lobe), an inflated black bulb (dark swelling), in prox-
imal third of the intestine, a crop, and a multilobulate digestive
gland; absence of an ink sac. Arms 1.5-4.3 times mantle length.
Two rows of suckers on each arm. Arm formula typically 1.2.4.3
or 2.1.4.3. No enlarged suckers. Maximum depth of the largest
sector of the web about 22% of the longest arm. Gills with 7-8
lamellae per demibranch. Right arm III hectocotylized (HA
1.5-2.1 times mantle length) in males. Ligula short (8-10% of
HA), lance-shaped and without transverse ridges. Calamus rep-
resents 30-50% of the ligula length in fully mature specimens.
Spermatophore length 70-125% of mantle length. Only one fe-
male of 35 mm mantle length was collected; external morphol-
ogy similar to male. Finger-like oocytes (80) ranging from 0.15-
4 mm maximum length.
Biology: This benthic species has characters that represent ei-
ther adaptations to the deep-sea (absence of ink sac, loss of the
anal flaps, eye without iris and optic chiasma) or to a hy-
drothermal vent habitat (eyes are covered by a thin semi-
translucent skin, high concentrations of metals and presence of
amoebocytes clots in the venous system and in the renal sacs).
It inhabits an isolated extreme environment among aggrega-
tions of tubeworms Riftia pachyptila, Alvinellidae polychaetes or
mussels, very close to the chimneys (2-10°C); also observed on
the pillow lava at several meters from the active areas. No pre-
dation over these species was observed. Octopuses forage on ba-
thypelagic amphipods, apparently targeting their attacks based
on contact with the swarming amphipods (e.g. Halice hes-
monectes). Some male specimens were parasitised by Genesis
vulcanoctopusi, a species of cholidynid harpacticoid (copepod).
Distribution: East Pacific Rise: collected at 13°N (Fig. 2) and
21°S (Fig. 1); observed at 23°S (Fig. 3).

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

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

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.

J.R. VOIGHT Denisia 18 (2006): 171

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

Mollusca, Cephalopoda, Octopoda, Octopodidae

Graneledone JOUBIN, 1918

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

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

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

Nematoda

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

173
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Annelida, Polychaeta1
According to different countings, the polychaete worms, with
111 presently described species, constitute 18-20% of the total
number of species fully identified from vent samples. 30% of these
bristle worms are belonging to scale worms (Polynoidae), 10% to
alvinellids, 9% to siboglinids, 6% to spionids, 5% to hesionids and
dorvilleids. Other families account for less than 5% of described
annelid species. The present recorded species are largely (94%)
considered as ‘endemic’ of the deep-sea hydrothermal vent envi-
ronment (or according to Andrey Gebruk, vent ‘obligate’), less
than 4% are shared with other deep-sea reduced environments and
ca. 2% are ‘regular’ deep-sea species. However this high ratio of en-
demism will probably be revised downward when the sampling of
the peripheral areas will make progress accounting for opportunis-
tic groups as dorvilleids or spionids which could be thriving in ad-
jacent disturbed habitats.
Because of their odd biology and ecology, two species of an-
nelids Riftia pachyptila (‘the giant tube worm’) and Alvinella pompe-
jana (‘the Pompeii worm’) became emblematic of the vent research
and are still considered as ‘biological models’. Numerous research
projects dealing with the functioning of endosymbiosis in Riftia
pachyptila and the physiological adaptation of Alvinella pompejana
to its extreme habitat are presently underway.
Nevertheless, the basic taxonomy of vent annelids is far to be
completed. Recent cruises which occurred in the Western Pacific
(Lau Basin, Kermadec and Mariana Arcs) and on southern East Pa-
cific Rise brought back a lot of new interesting specimens present-
ly under study. For example, five new species of scale-worms 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.
The molecular identification of sibling species among wide-
spread morphotypes (e.g. Amphisamytha galapagensis or Archinome
rosacea) lead ‘classical’ taxonomists to dig back throughout former
collections looking for new diagnostic characters. For example, a
joint work between morphologists and molecular taxonomists al-
References:
HALANYCH K.M. (2005) Hydrobiologia 535/536: 297-307.
ROUSE G.W. & K. FAUCHALD (1997) Zool. Scr. 26: 139-204.
SOUTHWARD E.C, SCHULZE A. & S.L. GARDINER (2005) Hydrobiologia 535/536: 225-249.
D. DESBRUYÈRES
lows recognizing three different new species of Archinome and one
new genus of Amphinomidae among specimens previously identi-
fied as A. rosacea (J. Kudenov, pers. comm.) Conversely, new
records of previously described species from new locations (e.g. Hes-
iolyra bergi on Mid-Atlantic Ridge vents or Hesiospina vestimentifera
from Lau Basin) question the dispersal and allopatric speciation in
response to major vicariance events. Because of the influence of
joint occurrences in biogeographic analyses, the solution of these
taxonomic riddle must be seriously considered and we urge scien-
tists working at sea to focus on these widespread morphotypes and
to build up parallel samples for classical and molecular taxonomy.
Polychaetes are delicate and fragile animals, and special care
should be taken when handling them. To be described, the speci-
mens must be unfragmented and as intact as possible and damaged
specimens may be misidentified. Thus a gentle sorting is desirable as
soon as possible after recovery of the samples, using soft pliers for
macroscopic individuals and gentle sieving for others (avoid stack
of sieves and split the samples before sieving). Specimens may be re-
laxed prior to preservation (7.5% of magnesium chloride) and tubes
of tubicolous annelids must be opened; fixation is best in buffered
formalin 5-10%. After a suitable time of fixation, depending of the
size (in general < 24 hours), the worms may be transferred to 80%
ethyl or isopropyl alcohol (formaldehyde even buffered, is a very
poor preservative).
1: Laminatubus alvini; by courtesy of R. A. Lutz.
1
Even if the question is still strongly debated and the molecular information is
not consistent, we chose herein, following ROUSE & FAUCHALD (1997) to in-
clude the Pogonophora (Perviata and Vestimentifera) within the family Si-
boglinidae (Polychaeta: Sabellida). Contrarily SOUTHWARD et al. (2005) chose
a more conservative standpoint and retain the class ‘Pogonophora’ within An-
nelida, waiting for more conclusive information (HALANYCH 2005).
DENISIA 18 (2006): 185
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

1: Top left, proboscis and 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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta, Phyllodocida, Hesionidae

Amphiduropsis axialensis (BLAKE & HILBIG, 1990)

rophores. Neuropodial lobes and neurochaetae present from

Size: All known specimens incomplete, anterior end fragments
up to 8 mm long for 21 segments.
Color: Preserved animals yellow-white, pigmentation absent or
with dark brown-black pigmentation on lips, laterally on pros-
tomium, palpophores and nuchal organs, distally on proboscis,
and on venter.
Morphology: Prostomium rounded rectangular, wider than
long, posteriorly with small median incision. Palpophores very
short, palpostyles of equal width or with slightly enlarged
rounded tips. Paired antennae as long as palps, tapered. Both
palpostyles and paired antennae very long and slender. Median
antenna much shorter than paired antennae, with tapering tip,
inserted medio-dorsally on prostomium. Eyes absent. Proboscis
smooth and unarmed, ending anteriorly in ring of cilia but
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-
segment 5, notopodial lobes and notochaetae from segment 6.
Notopodial lobes conical, pointed. Notochaetae include many
fine capillaries with two rows of small teeth, three to four ven-
trally situated curved chaetae with distal serration on ventral
side, and two (rarely one) medially situated stout emerging aci-
cular chaetae. Neuropodial lobes conical, much larger than no-
topodial lobes. Neurochaetae all compound. Ventral cirri dis-
tally inserted on neuropodium.
Remarks: The new record from East Pacific Rise: 9°N is based
on a single specimen from the expedition Hope’99 (Nautile
dive 1372, 2500 m, 09°46’17’’N, 104°21’70’’W); although not
in good condition it agrees well with A. axialensis.
Distribution: Juan de Fuca Ridge: Axial Seamount and Middle
Valley; East Pacific Rise: 9°N (new record; see Remarks).

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Hesiolyra bergi BLAKE, 1985 “caterpillar worm“
Size: Up to 80 mm for 100 segments.
Morphology: Body anteriorly truncate and posteriorly tapered,
dorso-ventrally flattened. Prostomium rounded rectangular,
slightly wider than long. Palpophores short, stout; palpostyles
much longer and thinner than palpophores, cylindrical with
rounded tips. Paired antenna similar to palpostyles; median an-
tenna absent. Eyes absent. Outside of everted proboscis with
pointed papillae, increasing in size distally. Terminal ring of
proboscis with large number of pointed papillae. Inside of pro-
boscis with large number of denticles in band on ventral side;
dorsal and ventral teeth present, dorsal much larger. Dorsal cir-
ri and cirrophores on segment 1–5 and ventral cirri and cir-
rophores on segment 1–4 larger and longer than on following
segments. Neuropodial lobes and neurochaetae from segment 5,
notopodial lobes and notochaetae from segment 6. No-
tochaetae surrounded by large pre- and postchaetal lamellae,
smaller lamellae also surrounding neurochaetae. Notochaetae
of four kinds: one-two serrated flattened (difficult to detect),
1: Habitus in vivo; by P. Briand
 Ifremer.
F. PLEIJEL
Annelida, Polychaeta, Phyllodocida, Hesionidae
large number of lyriform, tapered spinulose, and subdistally en-
larged (and often twisted) spinulose. Neurochaetae all com-
pound, dark to black in color (possibly due to some inclu-
sions?), with unidentate tips. Ventral cirri inserted medially on
neuropodium, with distinct cirrophores.
Biology: On smoker walls, often visiting tubes of Alvinella pom-
pejana. Can occur in large numbers. Epibiotic filamentous bac-
teria on parapodia and chaetae. Egg diameter about 125 µm.
Distribution: East Pacific Rise: 9°N and 21°N, and a closely re-
lated species on Mid-Atlantic Ridge: at the Azores Triple
Junction (expeditions Diva 1 and 2, Marvel) and 22°N
(Microsmoke). Newly collected specimens from Mid-Atlantic
Ridge agree well with H. bergi. Whether they should be referred
to a new species or not cannot be stated at present; they are
currently best referred to as “Hesiolyra sp.”.
2: Population on a smoker wall, visit-
ing Alvinella’s tubes (East Pacific Rise
13°N, 2630 m). Phare cruise  Ifremer.
3: Proboscis, frontal view (SEM)
 Ifremer.
Denisia 18 (2006): 200–201
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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.

F. PLEIJEL Denisia 18 (2006): 204

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

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

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

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

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

Annelida, Polychaeta, Phyllodocida, Nautiliniellidae

Mytilidiphila enseiensis MIURA & HASHIMOTO, 1993

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

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

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

Annelida, Polychaeta, Phyllodocida, Nautiliniellidae

Mytilidiphila okinawaensis MIURA & HASHIMOTO, 1993

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

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

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

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

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

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

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

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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Nereis piscesae BLAKE & HILBIG, 1990
Size: A large, robust species, some fragments up to 5 cm long;
largest complete specimens 75 mm long, 7 mm wide, for 100
segments.
Morphology: Prostomium triangular, narrowing anteriorly to
rounded tip bearing two short antennae; eyes absent, but two
pairs of sunken depressions representing positions of missing
eyes present; palps thickened, fleshy, each bearing two finger-
like palpostyles. Peristomium enlarged, inflated, surrounding
prostomial structures; four pairs of tentacular cirri. Jaws golden,
pointed, each with 78 teeth. Paragnaths all conical, arranged as
follows: I – two large cones in vertical row; II – 12 large cones
in nearly circular arc; III – 21-22 small and large cones in clus-
ter; IV – 19-20 small and large in three distinct clusters
arranged in three transverse rows; V – 0; VI – 46 large pointed
cones; VII – two small cones; VIII – two groups of two small
cones. Parapodia of middle and posterior segments with upper
notopodial ligule becoming enlarged and flattened, with dis-
tinct dorsal curvature and tapering triangular tip, with thick-
ened glandular border, bearing thin dorsal cirrus; lower notopo-
dial ligule tapering, not enlarged; neuropodial ligule thickened
1: Anterior end, ventral view (SEM) © Ifremer.
J.A. BLAKE & D. DESBRUYÈRES
Annelida, Polychaeta, Phyllodocida, Nereididae
basally, tapering distally; ventral cirrus narrow, cirriform. No-
topodial acicular lobes small in both anterior and posterior seg-
ments. Neuropodial acicular lobes conical throughout. Four
types of chaetae present. Anterior notochaetae all homogomph
spinigers replaced from segments 30-35 by 1-2 heavy ho-
mogomph falcigers with elongated blades bearing 6-8 blunt
denticles [not seen in specimens from type locality studied by
D. Desbruyères, probably worn falcigers]; upper neurochaetae
including heterogomph spinigers and heterogomph falcigers;
lower neurosetae include homogomph spinigers and hetero-
gomph falcigers; blades of all spinigers with thin denticles on
cutting edge; falcigers with thin, pointed denticles; shafts of all
chaetae smooth; internally ornamented with dark brown to
black camerations.
Biology: An epifaunal species associated with siboglinids. The
species is probably an omnivore, feeding on filamentous bacte-
ria.
Distribution: Juan de Fuca Ridge.
2: Anterior end, lateral view (SEM) © Ifremer.
Denisia 18 (2006): 212–213
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

Annelida, Polychaeta, Phyllodocida, Phyllodocidae

Eulalia papillosa (BLAKE, 1985)

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

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.

J.A. BLAKE Denisia 18 (2006): 215

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

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.

J.A. BLAKE Denisia 18 (2006): 216

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

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.

J.A. BLAKE Denisia 18 (2006): 217

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

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

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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Annelida, Polychaeta, Phyllodocida, Polynoidae
Branchinotogluma sandersi PETTIBONE, 1985
Synonym: B. grasslei PETTIBONE, 1985.
Size: Up to 60 mm in length.
Color: Pink to light red in living specimens. Golden-colored
chaetae.
Morphology: Body rectangular flattened, tapering slightly an-
teriorly and posteriorly. Dorsum with transverse ciliated bands.
Elytra 10 pairs, large covering dorsum, round to oval, stiff,
opaque. Each segment from 4rd chaetiger with two groups of del-
icate arborescent compact branchiae, with terminal filaments.
A single group of branchiae on 3rd segment. Prostomium
bilobed, anterior lobes cylindrical with a terminal filament.
Median antenna with a short style. Palps stout tapered smooth.
Tentaculophores with two pairs of tentacular cirri, dorsal ones
as long as palps. Muscular pharynx showing five papillae around
opening (3 dorsal and two ventral). Jaws denticled in their in-
ner border.
In males, parapodia of segments 2-18 biramous. Notopodia con-
ical with projecting acicular processes, without bracts except on
segment 2. Neuropodia with long conical prechaetal acicular
lobes, postchaetal lobes rounded and shorter. Notochaetae
1: Dorsal view; by P. Briand © Ifremer.
Reference:
PETTIBONE M.H. (1985) Proc. Biol. Soc. Wash. 98: 447-469.
D. DESBRUYÈRES & S. HOURDEZ
short and acicular, superior neurochaetae with two rows of
prominent spines and a slightly hooked tip, lower neurochaetae
with flattened spinous hooked hood. Parapodia of the posterior
three segments modified directed posteriorly and enclosed in
parapodia of segment 18, all with ventral cirri but branchiae
lacking. Parapodia of elytrigerous segment 19 biramous with ra-
mi similar in length; notopodia with thickened dorsal bract.
Parapodia of segment 20 much smaller and enclosed in parapo-
dia of segment 19. Notopodial acicular lobe fused to cirrophore
of dorsal cirrus. Notochaetae few, short, stout, curved with two
rows of distal spines. Upper neurochaetae with scattered long
spines and rounded tips. Segment 21 similar to 20th but lacking
notochaetae. Four pairs of segmental ventral papillae on seg-
ment 12-15. Three pairs of rounded ventral lamellae on seg-
ments 16-18.
In females (described as B. grasslei) segments 19-21 not trans-
formed. With or without five pairs of small squarish segmental
ventral papillae near ventral bases of neuropodia 11-15.
Distribution: Galapagos Spreading Center, East Pacific Rise:
21°N to 17°S, Guaymas Basin.
2: Ventral view of the posterior segments, female;
by P. Briand © Ifremer.
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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Annelida, Polychaeta, Phyllodocida, Polynoidae
Branchipolynoe seepensis PETTIBONE, 1986
Size: Up to 31 mm in length and 13 mm in width.
Morphology: Body short, spindle-shaped, tapered anteriorly
and posteriorly, flattened ventrally and arched dorsally. 21 seg-
ments with 10 pairs of elytra attached on segments 2, 4, 5, 7, 9,
11, 13, 15, 17 and 19. Elytra moderatly large, oval, covering lat-
eral thirds of body on larger specimens and nearly covering
body of juveniles. Branchiae well developped, arborescent.
Prostomium oval, bilobed, with rounded anterior lobes lacking
frontal filaments. Median antenna lacking distinct cer-
atophore. Palp rather short, thick, smooth and tapered. Thick
muscular pharynx, five pairs of small delicate oval papillae
around opening and two pairs of small jaws lacking denticulate
base. Lower neurochaetae slender and having hooked tips with
some longer lateral denticles.
1: Proboscis, frontal view (SEM) © Ifremer.
D. DESBRUYÈRES
Remark: According to CHEVALDONNÉ et al. (1998) B. seepensis
from the Gulf of Mexico and B. seepensis from Mid-Atlantic
Ridge appear to have recently diverged and must be considered
as two isolated phylogenetic species, although no morphologi-
cal differences have been found on large series of specimens.
Biology: Commensal within the mantle cavity of mussels. Two
individuals (male and female) are frequently observed within a
single mussel. Sexual dimorphism with females larger than
males; females having two pairs of ventral papillae on segments
11 and 12. Sex ratio 0.5-0.7: 1. Females contain mature sperm
and there is evidence of internal fertilization. Inferred mode of
development lecitotrophic or direct. Asynchronous gametoge-
nesis, rapid oogenesis.
Distribution: Hypersaline seeps at the base of Florida Escarp-
ment, 26°02’24’’N and 84°54’48’’W; Mid-Atlantic Ridge:
Lucky Strike.
2: Tip of upper neurochaetae
(SEM) © Ifremer.
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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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

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

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

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

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

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
ventroposterior sides of the
neuropodia. © Briand/Ifremer.
5: Globular micropapillae
at the surface of elytron (SEM) 7: Two pairs of dorsal and ventral jaws minutely
© 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

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

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

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

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

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.
Morphology: Body flattened, with 30 or 31 segments. Elytra 14
pairs on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26 and
27; elytra on segment 2 oval, with fringes of short papillae cov-
ered with filamentous bacteria; other elytra elongated subreni-
form, lateral borders with fringes of short papillae; elytral sur-
face covered with hexagonal or polygonal areas with secondary
areolae. Prostomium partially fused to tentacular segment, and
withdrawn in anterior segments. Prostomium bilobed forming
separate rounded lobes with anterolateral extensions. Tentacu-
lar segment with long cylindrical tentaculophores lateral to
prostomium; each with a single aciculum. Nodular papillae aris-
ing from dorsum of anterior segments; more than 20 papillae on
anterior segments; arranged in two or more rows before segment
Elytrophores bulbous, transversaly elongated; places of attach-
ment with latero-posterior extensions. Dorsal tubercles on cir-
rigerous segments bulbous, prominent, transversally elongated,
continuous with enlarged cirrophores of dorsal cirri. Dorsal and
ventral cirri with short clavate papillae. Branchiae absent. No-
topodia subconical, with projecting acicular lobes hidden by
numerous notochaetae. Neuropodia large truncate. Hooked
neurochaetae occurring on segment 3.
Biology: Free living scale-worm. Epibiotic filamentous bacteria
on parapodia.
Distribution: North Fiji and Lau Back-Arc Basins.

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

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

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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Thermopolynoe branchiata MIURA, 1994
Size: Up to 53 mm in length. Up to 20 mm in width.
Morphology: Body flattened, with 27 segments (first
achaetous). Elytra 11 pairs on segments 2, 4, 5, 7, 9, 11, 13, 15,
17, 19 and 21, large, covering dorsum, stiff, rough with numer-
ous brownish papillae. Branchiae well developed, arborescent
on segment 3-26; separated into two groups on anterior and
posterior sides of notopodia on segment 3 and on some less de-
veloped posterior segments; forming single large branchial areas
encircling central parts of notopodia on other fully developed
parapodia. Prostomium bilobed. Anterior lobes prominent,
cylindrical, with small frontal filaments. Mouth opening situat-
ed between segments 1 and 2. Muscular pharynx encircled by
1: Habitus in vivo; by courtesy of C. Rouse (cruise TUIM06).
2: Elytron surface (SEM) © Ifremer.
3: Elytron surface detail (SEM) © Ifremer.
T. MIURA & D. DESBRUYÈRES
Annelida, Polychaeta, Phyllodocida, Polynoidae
seven pairs of bulbous papillae. Dorsal and ventral pairs of jaws
fused medially each with up to 15 teeth on basal sides. No-
topodia subconical with projecting acicular lobes and large flar-
ing bracts. Neuropodia diagonally truncate, deeply notched on
upper part; distal margins fimbriated with slender papillae cov-
ered with numerous filamentous bacteria. Ventral segmental
papillae long; three pairs present on segments 12, 13 and 14 and
lacking in half of the specimens.
Biology: Free living. Found in washings of Ifremeria and mus-
sels. Epibiotic bacteria on parapodia.
Distribution: North Fiji, Lau Basin, and Manus Back-Arc
Basins.
4: Prostomium and buccal region, frontal view (SEM)
© Ifremer.
5: Inferior neurochaetae (SEM) © Ifremer.
6: Superior notochaetae (SEM) © Ifremer.
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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Annelida, Polychaeta, Sabellida, Siboglinidae

Ridgeia piscesae JONES, 1985

Size: Tube length max. 1900 mm; anterior diameter 2-13 mm.
Morphology: Tube very variable, straight or sinuous, stiff or
soft, tapering to 1mm posterior end; anterior funnel and exter-
nal flanges present. Color white, grey, gold, brown; commonly
translucent. Top of obturaculum carries 1-15 brown saucers on
a central axial rod; if these are shed, traces of the axial structure
remain. Red branchial filaments in concentric lamellae parallel
to the obturaculum; most bear pinnules, but there are a few
thin, smooth filaments at the ends of each row.
Remark: R. piscesae was amended by SOUTHWARD et al. (1995)
to include R. phaeophiale JONES 1985
Biology: Grows gregariously in clusters at sites of diffuse or
warm venting, temperature range 5-60°C (?). Uses internal
symbiotic sulphide-oxidizing bacteria. This species can even
grow and survive in areas of low diffuse vent flow with very low
plume level exposure to sulfide.
Distribution: Explorer Ridge, Juan de Fuca Ridge, Gorda
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

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

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

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
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.
ARP A.J., DOYLE M.L., DI CERA E. & S.J. GILL (1990) Resp. Physiol. 80: 323-
334.
BELKIN S., NELSON D.C. & H.W. JANNASCH (1986) Biol. Bull. 170: 110-121.
BLACK M.B., LUTZ R.A. & R.C. VRIJENHOEK (1994) Mar. Biol. 120: 33-39.
BRIGHT M., KECKEIS H. & C R. FISHER (2000) Mar. Biol. 136: 621-632.
CHILDRESS J.J., ARP A.J. & C.R. FISHER (1984) Mar. Biol. 83: 109-124.
CHILDRESS J.J., FISHER C.R., FAVUZZI J.A., KOCHEVAR R.E., SANDERS N.K. & A.M.
ALAYSE (1991) Bio. Bull. 180: 135-153.
HAAS F. de, ZAL F., LALLIER F., TOULMOND A. & J.N. LAMY (1996) Proteins 26:
241-256.
FELBECK H. (1985) Physiol. Zool. 53: 272-281.
GAILL F., HERBAGE D. & L. LEPESCHEUX (1988) Oceanologica Acta Numéro Spé-
cial 8: 155-160.
GAILL F., SHILLITO B., MÉNARD F., GOFFINET G. & J.J. CHILDRESS (1997) Mar. Ecol.
Progr. Ser. 148: 135-143.
GARDINER S.L. & M.L. JONES (1985) Trans. Amer. Microsc. Soc. 104: 19-44.
GIRGUIS P.R., LEE R.W., DESAULNIERS N., CHILDRESS J.J., POSPESEL M., FELBECK H. &
F. ZAL (2000) Appl. Environ. Microbiol. 66: 2783-2790.
GOFFREDI S. K., CHILDRESS J.J., DESAULNIERS N.T. & F.H. Lallier (1997) J. Exp. Bi-
ol. 200: 2609-2616.
JONES M.L. (1981) Proc. Biol. Soc. Wash. 93: 1295-1313.
MARSH A.G., MULLINEAUX L.S., YOUNG C.M. & D.T. MANAHAN (2001) Nature
411: 77-80.
POWELL M.A. & G.N. SOMERO (1983) Science 219: 297-299.
SCOTT K.M., FISHER C.R., VODENICHAR J.S., NIX E.R. & E. MINNICH (1994) Physi-
ol. Zool. 67: 617-638.
THIEBAUT E., HUTHER X., SHILLITO B., JOLLIVET D. & F. GAILL (2002) Mar. Ecol.
Progr. Ser. 234: 147-157.
ZAL F., LALLIER F.H., WALL J.S. & S.N. VINOGRADOV (1996) J. Biol. Chem. 271:
8869-8874.
263
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

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.

E.C. SOUTHWARD Denisia 18 (2006): 265

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

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
chaetigers.
Morphology: Color in alcohol light to dark brown, with gold
pigment on anterior end and dark, rusty pigment posteriorly;
some EPR specimens with glandular areas appearing dark green
or black. Prostomium broadly rounded anteriorly, merging pos-
teriorly with arched cephalic keel; cephalic plate lacking,
nuchal grooves curved. Segments longer than wide, flaring an-
teriorly, but not forming collars. Notochaetae capillaries; long
filamentous capillary notochaetae absent. Neurochaetae of
from chaetiger 4 increasing to 9-10 per fascicle middle seg-
ments; hooks with four apical teeth and tuft of bristles below
main fang. Anus surrounded by funnel bearing about 25 papil-
lae.
Biology: An epifaunal species, found in association with mus-
sels, clams, siboglinids, and other associations. Maldanids are
known as bulk-ingestors of particles.
Distribution: Galapagos Spreading Center; East Pacific Rise:
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

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

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

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

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
chaetigers.
Color: In alcohol tan.
Morphology: Prostomium conical, smoothly rounded along an-
terior margin, with small nuchal organs on lateral boundary of
prostomium and peristomium. Peristomium with one or two
achaetous rings depending on size and state of preservation;
both rings apparent in smaller specimens, vaguely apparent in
larger specimens. Thorax with nine similar chaetigers. Notopo-
dia with thin, cirriform postchaetal lobes; neuropodia, with
short triangular shaped postchaetal lobes, developing thick-
ened base in abdominal region; neuropodial lobe of abdominal
segments becoming thicker, blunted, then developing weak
bilobed appearance in far abdominal segments. Notochaetae
camerated capillaries; furcate chaetae absent. Thoracic neu-
cluding 2-3 capillaries and 1-2 thin acicular spines. Branchiae
from chaetiger 13-15; each thick in cross section, appearing
glandular; anterior and posterior abdominal branchiae short,
stubby in appearance, with broad base and narrow apex;
branchiae from middle abdominal segments, longer, but still
with broad base. Pygidial segment broadly rounded, with two
lateral cirri.
Biology: Exact habitat unknown, collected near an active vent.
The species probably burrows into mud and is likely a deposit
feeder. Two large eggs measuring approximately 100 mm in di-
ameter were observed in one specimen. They were enclosed in
a chamber that was located on the dorsum between chaetigers
6 and 7.
Distribution: Juan de Fuca Ridge.

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

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

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.

J.A. BLAKE Denisia 18 (2006): 269

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

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

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

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

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

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.

D. DESBRUYÈRES Denisia 18 (2006): 272

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

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

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

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

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

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.

J.A. BLAKE Denisia 18 (2006): 275

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

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

Annelida, Polychaeta, Spionida, Spionidae

Prionospio sandersi MACIOLEK, 1981

Size: Small, up to 3.2 mm long, 0.5 mm wide, with about 35
segments.
Morphology: Prostomium broadly rounded anteriorly, ending
posteriorly at chaetiger 1. Peristomium encompassing prostomi-
um laterally and ventrally, not developing lateral wings; fused
dorsally to chaetiger 1. Branchiae from chaetiger 2, nine pairs;
each gill broad, robust, appearing wrinkled, but lacking pin-
nules; branchiae longest anteriorly, becoming shorter, stubby
posteriorly. Dorsal crests absent. Anterior chaetae capillaries;
multidentate hooded hooks from chaetiger 15 in neuropodia,
chaetiger 29 in notopodia; ventral sabre chaetae from chaetiger
14. Pygidium cup-shaped, with deeply rounded ventral edge
and small dorsal elongation.
Remark: A closely related form of Prionospio sp. has been com-
monly sampled on the Northern East Pacific Rise at 13°N and
9°N. It differs from Prionospio sandersi by the number of
branchiae (12-14 pairs instead nine pairs) by the occurrence of
the sabre chaetae on neuropodia from chaetiger 15 instead of
14 and the occurrence of the multidentate hooks on neuropo-
dia from chaetiger 17 instead 15 and in notopodia from
chaetiger 23 instead of 29.
Biology: The known specimens were collected from washings
of Riftia pachyptila and might no be a fully developed. The
species is likely a surface deposit feeder, scavenging particles
with its short palps.
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
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Prionospio unilamellata SIGVALDADÓTTIR & DESBRUYÈRES, 2003
Size: Up to 17 mm long and 0.9 mm width for 114 chaetigers.
Morphology: Prostomium broadly triangular, widest at anterior
margin, narrowing into a raising caruncle, reaching to posteri-
or delineation of first chaetiger. Anterior margin sometimes
with 3-4 ciliated tubercles. Peristomium partly surrounding
prostomium, forming a low collar, not fused to chaetiger one.
Pair of indistinct ciliated nuchal organs situated on each side of
caruncle. Postchaetal notopodial lamella of chaetiger one re-
duced, dorsally pointed, free from prechaetal lamella. On
chaetigers 2 to 16-17, a single large notopodial lamella is en-
closing chaetae. Postchaetal lamellae on postbranchial seg-
ments becoming smaller, separated from prechaetal lamellae,
round in form. On anterior chaetigers, postchaetal lamellae
with dorsal tip. On posteriormost chaetigers lamellae dorsally
pointed. Prechaetal lamellae small on postbranchial chaetigers.
Dorsal crests absent. Neuropodial postchaetal lamellae reduced
on first chaetiger, elliptical in shape. Lamellae largest on
branchial chaetigers, successively becoming smaller. On
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
part of notopodial hooded hook (SEM)  Ifremer.
Reference:
SIGVALDADOTTIR E. & D. DESBRUYÈRES (2003) Cah. Biol. Mar. 44: 219-225.
D. DESBRUYÈRES
Annelida, Polychaeta, Spionida, Spionidae
chaetigers lamellae leaflike, dorsally pointed. Prechaetal lamel-
lae separated from postchaetal lamellae. Branchiae from
chaetiger two, 15-16 in number. Anterior branchiae slightly
longer than notopodial lamellae, on subsequent chaetigers suc-
cessively longer and more slender. Branchiae apinate with
dense lateral ciliation. Interparapodial pouches missing. Small,
papilla-like, structures can sometimes be observed laterally be-
tween neuropodia. These structures start on 2-3 posteriormost,
or posterior to, branchial segments and occur on 7-8 segments.
Chaetae on anterior chaetigers slender, arranged in dense dou-
ble rows. Neuropodial hooded hooks from chaetiger 19-27. In-
troduction of hooks moving backwards with increasing size.
Hooks with 4-6 secondary teeth over main fang. Notopodial
hooks from chaetiger 25-46. Notopodial hooks with longer
shaft than neuropodial hooks. Sabre chaeta from chaetiger 12-
20, one or two per rami. Sabre chaeta distally granulated. Py-
gidium with long median cirrus and two rounded lateral flaps.
Distribution: Mid-Atlantic Ridge: Lucky Strike, Rainbow and
Snake Pit.
2: Anterior part (SEM)  Ifremer.
Denisia 18 (2006): 280
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

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

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

Annelida, Polychaeta, Terebellida, Alvinellidae

Alvinella caudata DESBRUYÈRES & LAUBIER, 1986

Size: Up to 140 mm.
Morphology: Four pairs of lamellate gills; two transformed
chaetigerous segments bearing stout spines; body in two parts,
the posterior one with enlarged notopods bearing distal digita-
tions; bacterial epibiosis.
Biology: Dwelling inside organic tubes in active chimney walls.
Temperature ranging from 10-50°C. Seldom observed in situ.
Feeding on free bacteria, associated with filamentous bacteria
epibiosis. Gut functional; retractile buccal tentacles; gonochor-
ic, sexual dimorphism.
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

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

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

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

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).
Color: Dark red when preserved.
Morphology: Alvinellid worm with four pairs of pinnate
branchiae. Prostomium very reduced, similar to that of P. bac-
tericola with a median lobe having a median incision and two
very well developed and bilobed lateral expansions. Numerous
grooved buccal tentacles inserted on two lateral pads. As in P.
grasslei, a median hemispherical body is observed ventrally in
the buccal cavity, bordered by two lateral pads. Two long point-
ed and grooved peribuccal tentacles are present in males, each
with a deep longitudinal groove devoid of ciliation. Four stout
Uncinigerous neuropodial tori from chaetiger 48-52 to the end
of the body. No notopodial or anal cirrus.
Remarks: The species is a sister species of P. bactericola from
Guaymas basin and P. hessleri from Western Pacific Back-Arc
Basins which were grouped in the subgenus Miralvinella.
Biology: This worm is usually found in tube worm clusters.
Distribution: Juan de Fuca Ridge: Axial Seamount, sites Ashes
and Casm.

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

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

Annelida, Polychaeta, Terebellida, Alvinellidae

Paralvinella (Miralvinella) hessleri DESBRUYÈRES & LAUBIER, 1989

digitiform lobe bearing two groups of capillary chaetae, one

Size: Up to 22 mm.
Morphology: Alvinellid worm with four pairs of pinnate
branchiae. Up to 61 segments. Prostomium medially reduced
with a median incision and two well developed lateral lobes
ventrally enclosing peristomium. Male buccal apparatus com-
prising a ventral globular bulky organ, two lateral strong and
pointed tentacles bearing a deep groove without ciliation, and
many grooved and ciliated smaller tentacles inserted in two
groups on a quadrilobed upper lip. First two segments (II and
III) achaetous and fused to the first three chaetigerous. The first
15-20 chaetigerous segments with notopodia only. First three
notopodia smaller than others and dorsally elevated. Chaetiger
4 with a median dorsal expansion which protrude forward. No-
topodia from 4 to 13-17 (7 excepted) cylindrical with a dorsal
with short and the other with long. Chaetigerous segment 7
strongly modifies lacking cylindrical notopodia but bearing on
each side 4-5 strongly modified acicular hooks directed posteri-
orly. Segment 8 with cylindrical notopodium and very strong
digitiform lobe directed forward. Uncinigerous neuropodial tori
on each segment from segment 15-20 to the end of the body.
Biology: All specimens found in tubes on rocks directly ex-
posed to venting water whose temperature was recorded up to
25°C. Tubes whitish and corneous in aspect, amoeba like in
shape with long anchor filaments. Tube walls thick and multi-
layered. Inner surface bearing important bacterial mats.
Distribution: Mariana and Manus Back-Arc Basins.

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

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

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

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

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

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

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

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.

D. DESBRUYÈRES Denisia 18 (2006): 292

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

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.

D. DESBRUYÈRES Denisia 18 (2006): 293

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

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.
Color: Bright red in life and chocolate brown with brown
chaetae after preservation in formalin.
Morphology: 54-68 chaetigerous segments. Body regular, taper-
ing abruptly in terminal 10 segments. Buccal apparatus com-
prising small grooved and ciliated tentacles, one pair of large
basal tubercles and in males two large lateral tentacles and ter-
minating in a curved margin. First 24-30 chaetigerous segments
with notopodia only. Four pairs of branchiae present, all similar
strong, abruptly attenuated. Branchial stem relatively short,
with a large number of slender filaments arising in two opposite
ranks and bearing ciliated longitudinal line. First 22 notopodia
strongly modified with 4-5 very large stout notopodial hooks di-
rected posteriorly. Tubes cylindrical translucent and thin.
Biology: Temperatures of 20-80°C have been measured on sur-
faces colonized by sulfide worms and its most likely that the
worms regularly experience temperatures within the lower part
of this range. All the collections containing this species are
coming from active sulfide edifices where the animal occupies
a distinct microhabitat.
Distribution: Explorer Ridge, Juan de Fuca Ridge, Gorda
Ridge.

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.

D. DESBRUYÈRES Denisia 18 (2006): 294

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

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.

D. DESBRUYÈRES Denisia 18 (2006): 295

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

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.

D. DESBRUYÈRES Denisia 18 (2006): 296

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

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.

E.M. BURRESON & M. SEGONZAC Denisia 18 (2006): 297

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

Arthropoda, Chelicerata, Arachnida, Acariformes, Halacaridae

Copidognathus papillatus KRANTZ, 1982

Size: 500-570 µm (from tip of rostrum to end of anal papillae).
Morphology: Marine mite with ovate idiosoma, slender
gnathosoma with almost parallel-sided rostrum, two pairs of
legs directed forward and two pairs backward. Dorsal plates of
idiosoma with areolate-reticulate sculpturing. Anal papillae en-
larged, extending beyond anal aperture. Legs with aciculate
reticulation; they lack long spine-like setae.
Biology: Present within detritus among colonies of vestimen-
tiferan tubeworms, barnacles, and mussels.
Distribution: Galapagos Spreading Center, East Pacific Rise:
13°N, Juan de Fuca Ridge: Main Endeavour; North Fiji and Lau
Back-Arc Basins.

1: Habitus; by I. Bartsch.

References:
BARTSCH I. (1991) Zool. Sci. 8: 789-792.
KRANTZ G.W. (1982) Can. J. Zool. 6: 1728-1731.

I. BARTSCH Denisia 18 (2006): 299

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

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.

I. BARTSCH Denisia 18 (2006): 300

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

Arthropoda, Pycnogonida, Ammotheidae

Ammothea verenae CHILD, 1987

auxiliary claws. Male cement gland opening a tiny dorsodistal

Synonym: Scipiolus thermophilus TURPAEVA, 1988.
Size: Male leg span about 43 mm. Female slightly larger.
Morphology: Trunk slender with segment posteriors flared into
cowling. Ocular tubercle a low, blind, truncate cone at neck
mid length. Lateral processes separated by less than half their
diameters. Proboscis long, with median and distal constrictions.
Abdomen long, down curved. Chelifores tiny, short, chelae at-
rophied bumps. Palps nine-segmented, 5 distal segments very
short, heavily setose ventrally. Oviger segment 4 and 5 sub
equal in length, proximally setose, strigilis segments 6, 7 and 8
with many long lateral setae, segments 8, 9 and 10 with 1-2
small endal denticulate spines. Eggs tiny, about 0.25 diameter of
main oviger segments, carried in large round clusters. Legs long,
slender, extremely setose. Third coxae with many long ventral
setae. Distal leg segments alike, tarsus very short, propodus
slender, well curved, sole spines alike. Main claws with long
pore. Female oviger strigilis with segments lacking dense setae,
with 1-2 denticulate spines on distal segments. Female legs with
far fewer setae, without large fields of setae.
Remarks: This species differs in basic morphology from any
other known member of the genus. It is blind, lacks the con-
spicuous dorsomedian tubercles common on the trunk cowls of
most species, lacks differentiated heel spines, and does not have
differences in propodi shape between the anterior and posteri-
or legs.
Biology: Found commonly in close proximity to hydrothermal
vents and sometimes encrusted with polymetallic sulfides.
Distribution: Juan de Fuca Ridge: Axial Seamount, Endeavour
Segment; Explorer Ridge.

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.

R. BAMBER Denisia 18 (2006): 301

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

Arthropoda, Pycnogonida, Ammotheidae

Sericosura cochleifovea CHILD, 1989

ae and propodus. No thick fields of setae. Tarsus short, propo-

Synonym: Sericosura bifurcata STOCK, 1991.
Size: Small for the genus, male leg span about 13 mm.
Morphology: Trunk slightly ovoid, lateral processes separated
by about half their diameters, armed with one or more dor-
sodistal and laterodistal spines. Trunk segments with swollen
posterior cowls. Ocular tubercle only as tall as its base, blind.
Proboscis ovoid, originating on slender stalk, greatly constrict-
ed distally. Abdomen long, downcurved, with two pairs of long
dorsal spines, pair of short distal spines. Chelifores typical;
short, with atrophied chelae. Palps nine-segmented, second
segment longest, five distal segments little longer than wide,
heavily setose. Oviger fourth segment longest, fifth slightly
shorter. Strigilis sixth segment with field of long lateral setae,
with 1-2 tiny denticulate spines on distal three segments. Legs
moderately slender, very setose. Male leg major segments with
rows of many setae, some longer than segment diameter. Female
leg major segments with few very long femoral setae, lateral
rows of extremely long setae and few long dorsal setae on tibi-
dus slender, almost straight, claws moderately long. Eggs large,
almost as wide as oviger segment diameters around which they
are wound. Cement gland at extreme proximal end of femur
with laterally pointing tube slightly longer than segment diam-
eter.
Remarks: One of the major differences between this species
and the others in this genus is the sexual dimorphism displayed
in leg setae. The very long lateral leg setae of females differen-
tiate them conspicuously from males. The shorter leg setae and
very long cement gland tube of the males sets them off from fe-
males of this species and males of other known species which
have much shorter tubes.
Biology: Taken in the proximity of a vent site with other fau-
na: snails, crabs, shrimps and anemones.
Distribution: Only known from the Mariana Islands Back-Arc
Basin hydrothermal vent fields.

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.

R. BAMBER Denisia 18 (2006): 302

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

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.

R. BAMBER Denisia 18 (2006): 303

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

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].

R. BAMBER Denisia 18 (2006): 304

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

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

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

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

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

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 incisure small; posterior edge of each
that the species is a scavenger or predator on small inverte-
valve with small glandular process dorsal to midheight; without brates. The collecting site suggests that the species, which is an
posterodorsal spines. First antenna with proximal black spots. active swimmer, dwells in deep-water close to the substrate;
Second antenna: exopod with short medial setae on article 1; collected within Riftia pachyptila aggregations.
endopod article 1 without processus mamillaris; ventral margin
Distribution: East Pacific Rise: 9°N, site Tica.
with minute digitations and spines. Basal endite of mandible
with row of triangular teeth. Bellonci Organ elongate, reaching

3: Left first antenna and

1: Complete specimen from left side. Bellonci Organ.

4: Right
mandible,
distal coxa and
basis.

2: Complete specimen, dorsal view, anterior to right (black spots

are on first antenna; BO – tip of Bellonci Organ).

Reference:
KORNICKER L.S. & E. HARRISON-NELSON (2005) Zootaxa 1071: 19-38.

L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 307

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

Arthropoda, Crustacea, Ostracoda, Myodocopa, Halocyprida, Halocypridae

Bathyconchoecia deeveyae KORNICKER, 1969

Size (length and height excluding spines): Adult or A-1 female Biology: Collections indicate that the species, a swimmer, lives
collected off Surinam: length 1.49 mm, height 1.10 mm. Juve- close to the bottom at bathyal and abyssal depths.
nile collected off Peru: length 1.12 mm, height 0.89 mm.
Distribution: Guaymas Basin, Southern Trough (2000 m), col-
A-4 instar collected in Guaymas Basin: length 0.66 mm, height
lected in plankton net 3-4 m above bottom. Also collected off
0.46 mm.
Peru in the Peru-Chile Trench system (520 m), and in western
Morphology: Carapace with seven spines (each valve with an- Atlantic off Surinam (508–523 m).
terior spine, dorsal spine near midlength, and lateral spine be-
low middle; right valve with posterior spine). Surface with or
without punctae and arcuate ridges between punctae. Pos-
terodorsal corner of each valve with glandular process.

2: Juvenile carapace, lateral view (length 0.87 mm);

by Kornicker.

3: Endopod of second antenna

(adult or A-1 instar);
by Kornicker.

1: Juvenile carapace, dorsal, lateral, and ventral views

(length 0.87 mm); by Kornicker.

References:
KORNICKER L.S. (1969) Proc. Biol. Soc. Wash. 82: 403–408.
KORNICKER L.S. (1981) Proc. Biol. Soc. Wash. 89(4): 1237–1243.
KORNICKER L.S. (1991) Smithson. Contr. Sci. 516: 1–46.

L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 308

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

Arthropoda, Crustacea, Ostracoda, Myodocopa, Halocyprida, Halocypridae

Bathyconchoecia paulula DEEVEY, 1968

on each lamella and long unpaired bristle; small oval area prox-
Size: Adult male length 0.95 mm, height 0.60–0.61 mm (Guay-
imal to claw 2. Copulatory organ inner rod with pointed re-
mas Basin); length 0.95 mm, height 0.70 mm (Gulf of Mexico).
curved tip. Adult female carapace and appendages, except en-
Morphology: Adult male carapace rostrum with prolonged dopod of second antenna, fairly similar to those of adult male.
downward pointed tip; surface with crescent-like reticulations
Biology: Collections indicate that the species, a swimmer, lives
containing minute pits; height at least 70% length; posterodor-
close to the bottom at bathyal and abyssal depths. Specimens
sal corner of each valve with glandular opening; list along pos-
from the Gulf of Mexico were collected in the intestines of bot-
terior infold with four transparent lamella with four triangular
tom fish.
flagella-like cusps. First antenna terminal segments with more
than 200 sensory filaments. Second antenna exopod with un- Distribution: Guaymas Basin, Southern Trough (collected in
usually long first article (71–72% length of protopod); endopod plankton net 3-4 m above bottom). Also collected in Gulf of
first article with two bristles (distal bristle about twice length of Mexico (from intestines of bottom fish), 1000 m and southwest
proximal bristle). Fifth limb terminal article of endopod with of Sao Miquel, Azores (620–800 m) in plankton net open be-
two claws and one shorter ringed bristle. Sixth limb terminal tween 680 and 780 m.
article of endopod with three bristles (one very long, two
short). Seventh limb with two bristles. Furca with eight claws

2: Adult male
endopod of second
antenna; by Kornicker.

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

3: Adult male sixth limb; by Kornicker.

References:
DEEVEY G.B. (1968) Proc. Biol. Soc. Wash. 81: 539–570.
KORNICKER L.S. (1991) Smithson. Contr. Sci. 516: 1–46.
POULSEN E.M. (1972) Tethys 4(2): 445–456.

L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 309

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

Arthropoda, Crustacea, Ostracoda, Myodocopa, Myodocopida, Philomedidae

Euphilomedes climax KORNICKER, 1991

Size: Adult female carapace length 2.84 mm, height 2.02 mm. Biology: Benthic swimmer and burrower. One female with four
eggs in marsupium. Detritus feeder; guts generally with uniden-
Morphology of adult female (adult male unknown): Carapace
tified particles, but one specimen with two copepods in gut.
oval in lateral view with broad rostrum and deep incisure; sur-
The collecting site suggests that the species may be restricted to
face with many short pointed bristles, and long widely separat-
deep water.
ed bristles more numerous along valve edge. Mandible exopod
about half length of dorsal margin of article 1 of endopod. Sev- Distribution: Explorer Ridge: Pogo Peaks Vent, Gulati Gusher
enth limb with 17–19 bristles; comb with 13 alate teeth with Vent, Lunch Hour Vent, Crab Vent, Upper Magic Mountain
two small teeth on each side of base, side opposite comb with Vent, Busted Thruster Vent; Juan de Fuca Ridge: Long Term
two or three pegs. Furca with 11–14 claws on each lamella; claw Observatory Vent, Axial Seamount, Hamond’s Hell Vent.
3 small, secondary, remaining claws decreasing in length along
lamella. Bellonci Organ elongate with suture at midlength and
pointed or narrowly rounded tip. Lateral eyes absent in females
at all stages, present in males at all stages.

2: Right lamella of furca; by Kornicker.

1: Complete carapace showing four eggs; by Kornicker. 4: Anterior of body

from right side show-
ing Bellonci Organ;
by Kornicker.

3: Seventh limb; by Kornicker.

References:
KORNICKER L.S. (1991) Smithson. Contr. Zool. 516: 1–46.
TSURUMI M. & V. TUNNICLIFFE (2003) Deep-Sea Res. I 50: 611–629.
TUNNICLIFFE V. (1988) Proc. R. Soc. Lond. B 233: 347–366.

L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 310

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

Arthropoda, Crustacea, Ostracoda, Myodocopa, Myodocopida, Polycopidae

Polycopetta pax KORNICKER & HARRISON-NELSON, 2005

ing distal end of endopod, and with eight long terminal bristles.
Size: Adult female carapace length 0.54 mm, height 0.47 mm.
Fifth limb exopod well-developed, and with four terminal spin-
Morphology of female (male unknown): Carapace: anterior ous bristles (three long, one short). Bellonci Organ consists of
margin with concavity, but without rostrum or anterodorsal spinous bristle divided at midlength.
tooth; ventral margin with about 25 small teeth and minute
Biology: Unique specimen with several eggs in marsupium, and
spines (or serrations) between teeth. First antenna dorsal mar-
filled internal sperm sac near base of furca. Gut with brown par-
gin without processes, but with long proximal bristle.
ticles, some round or disc-like, and with few fragments of tubu-
Mandible: coxa endite bifurcate distally; exopod interpreted to
lar structures. Species is a bottom dweller; found within Riftia
have two articles, but without suture separating articles: article
pachyptila aggregations, maximal water temperature 23°C.
1 broad with terminal bare bristle; article 2 with proximal
spines and funnel-like tip. Maxilla: proximal dorsal half of ba- Distribution: East Pacific Rise: 9°N, site Riftia Field.
sis with long backward projection; distal end of exopod reach-

2: First antennae and Bellonci Organ (not all bristles shown

on first antennae).

1: Complete specimen from right side.

4: Right maxilla (not

all bristles shown;
precoxa endite not
shown).
3: Right mandible.

Reference:
KORNICKER L.S. & E. HARRISON-NELSON (2005) Zootaxa 1071: 19-38.

L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 311

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

Arthropoda, Crustacea, Ostracoda, Myodocopa, Myodocopida, Cylindroleberidae

Prionotoleberis styx KORNICKER, 1991

Size: Adult female carapace length 1.89 mm, height 1.09 mm. Biology: Benthic swimmer and burrower. Filter feeder; gut slen-
der, containing fine-grained material. The collecting site sug-
Morphology of adult female (adult male unknown): Carapace
gests that the species may be restricted to deep water.
elongate with deep incisure. First antenna sensory bristle with
fairly long proximal bristle and six long terminal filaments. Distribution: East Pacific Rise: National Geographic Site,
Sixth limb anterior margin with 18–20 short bristles. Seventh Clam Acres.
limb with 24 or 25 bristles. Furca with 10 claws on each lamel-
la. Lateral eyes absent.

1: Complete carapace showing central adductor

muscles; by Kornicker. 2: Sixth limb; by Kornicker.

3: Seventh limb;
by Kornicker.
4: Mandible; by Kornicker.

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

L.S. KORNICKER & E. HARRISON-NELSON Denisia 18 (2006): 312

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

Arthropoda, Crustacea, Ostracoda, Podocopida, Pontocyprididae

Thomontocypris brightae MADDOCKS, 2006

and reefal sand and plants (Madagascar, Australia), and exper-

Size: Carapace length 0.53-0.57 mm, height 0.30-0.33 mm, no
imental wood falls in the deep sea (MADDOCKS & STEINECK
carapace dimorphism.
1987; MADDOCKS 1991).
Morphology: Carapace compressed, lateral outline rounded-
Biology: Living specimens were collected in washings of Riftia
subtriangular with very broadly rounded anterior margin and
pachyptila. Appendages and genitalia are normal for the genus,
three-segmented dorsal margin, distinct ventral indentation;
and gut contents include no recognizable objects. Well-devel-
conspicuous anteroventral hornlike process on upper lip; palp
oped swimming setae suggest active demersal swimming, while
of male fifth limb broad, subtriangular, with thumb-like ven-
the rather weak mouthparts of this family indicate a soft detri-
trodistal projection; terminal setae of seventh limb smooth;
tus or bacterial diet. The function of the hornlike process is un-
Zenker’s organ with three bulbous swellings.
known.
Remarks: Other species of Thomontocypris have been reported
Distribution: East Pacific Rise: 9°N, site Riftia Field and Tica.
living in anchialine pools and caverns (Bermuda), sublittoral

1: Left side of entire animal. 4: Left side of entire female animal.

5: Dorsal view of entire male animal.

2: Exterior of right valve, male specimen.

6: Right side of anterior body, showing anteri-

oventral horn on upper lip and palps of male fifth
3: Exterior of left valve, male specimen. limb. 1-6; from MADDOCKS (2006).

References:
MADDOCKS R.F. (1991) Zool. J. Linn. Soc. 103: 309-333.
MADDOCKS R.F. (2006) Micropaleontology 51: 345-372.
MADDOCKS R.F. & P.L. STEINECK (1987) Micropaleontology 33: 318-355.

R.F. MADDOCKS Denisia 18 (2006): 313

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

Arthropoda, Crustacea, Ostracoda, Podocopa, Podocopida, Pontocyprididae

Thomontocypris gollnerae MADDOCKS, 2006

imental wood falls in the deep sea (MADDOCKS & STEINECK

Size: Carapace length 0.65 mm, height 0.32 mm, no carapace
1987; MADDOCKS 1991).
dimorphism.
Biology: Living specimens were collected in washings of Riftia
Morphology: Carapace egg-shaped, slightly flexible, swollen
pachyptila. Appendages and genitalia are normal for the genus,
ventrolaterally, with rounded-subtrapezoidal lateral outline,
and gut contents include no recognizable objects. Well-devel-
weakly convex ventral margin, and slightly produced anterior
oped swimming setae suggest active demersal swimming, while
end; Y-aesthetasc of antenna small, elongate; terminal setae of
the rather weak mouthparts of this family indicate a soft detri-
seventh limb smooth; furcal claws with opposing setules form-
tus or bacterial diet. The unusually weak calcification of the
ing terminal thorn.
carapace may be either a response or a preadaptation to low pH.
Remarks: Other species of Thomontocypris have been reported
Distribution: East Pacific Rise: 9°N, site Riftia Field.
living in anchialine pools and caverns (Bermuda), sublittoral
and reefal sand and plants (Madagascar, Australia), and exper-

3: Left valve exterior with ovaries.

4: Furca.

1: Ventral view of entire female.

5: Detail of left valve exterior, showing adductor

muscle scars (asterisk) and normal pore canals.
2: Right valve exterior with ovaries. 1-6; from MADDOCKS (2006).

References:
MADDOCKS R.F. (1991) Zool. J. Linn. Soc. 103: 309-333.
MADDOCKS R.F. (2006) Micropaleontology 51: 345-372.
MADDOCKS R.F. & P.L. STEINECK (1987) Micropaleontology 33: 318-355.

R.F. MADDOCKS Denisia 18 (2006): 314

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

Arthropoda, Crustacea, Ostracoda, Podocopida, Cytheruridae

Xylocythere vanharteni MADDOCKS, 2006

(Atlantic and Pacific, Upper Oligocene to Holocene; MAD-

Syn.: Xylocythere sp. A in VAN HARTEN (1993).
DOCKS & STEINECK 1987; STEINECK et al. 1990).
Size: Carapace length 0.56-0.66 mm, height 0.27-0.32 mm,
Biology: Living specimens were collected in washings of Riftia
males smaller than females.
pachyptila. The mouthparts and legs are normal for the family,
Morphology: Lateral outline elongate-subquadrate to subreni- and the gut contents contains no recognizable objects. A crawl-
form, with broad ventral indentation; reticular network of low, ing habit and bacterial or detrital diet are probable. The func-
narrow muri and flat, polygonal sola uniformly developed over tion of the pore clusters is unknown but might be related to res-
entire lateral surface, except for nearly smooth submarginal piration (VAN HARTEN, 1992, 1993).
fields; 20-45 pores in pore clusters; prominent ventrolateral
Distribution: East Pacific Rise: 13°N and 9°N, sites Tica and
spine and single anterior and posterior marginal spines on each
Riftia Field.
valve.
Remarks: Other living and fossil species of Xylocythere are re-
ported from experimental wood falls and deep-sea sediments

1: Entire animal, right side.

4: Left valve exterior (SEM).

2: Right valve exterior.

5: Right valve interior showing hinge and pore clusters (SEM).

3: Left valve exterior. 1-5: from MADDOCKS (2006).

References:
MADDOCKS R.F. (2006) Micropaleontology 51: 345-372.
MADDOCKS R.F., & P.L. STEINECK (1987) Micropaleontology 33: 318-355.
STEINECK P.L., MADDOCKS R.F., COLES G. & R.C. WHATLEY (1990) in WHATLEY R. & C. MAYBURY (Eds.) Ostracoda and Global Events,
Chapman & Hall: 307-319.
VAN HARTEN D. (1992) Deep-Sea Res. 39: 1067-1070.
VAN HARTEN D. (1993) in MCKENZIE K.G. & P.J. JONES (Eds.) Ostracoda in the Earth and Life Sciences. A.A. Balkema: 571-580.

R.F. MADDOCKS Denisia 18 (2006): 315

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Copepoda
Almost 80 species of copepods are described from hydro-
thermal vents. More than half of the copepods species recorded
represent copepods of the family Dirivultidae (order Siphonos-
tomatoida) which were exclusively found at different vent sites
with exception for the type species of Dirivultus dentaneus (HU-
MES & DOJIRI 1980; IVANENKO & FERRARI 2003). Additionally,
three dirivultid species descriptions from animals of the East
Pacific Rise, the West Pacific, and the Mid-Atlantic Ridge are
in preparation; and at least two dozens of new species, which
are in our disposition, are waiting for thorough descriptions.
These are mostly representatives of calanoids, cyclopoids and
harpacticoids from different localities of the East Pacific Rise
1: Oncaea preclara from East Pacific Rise: 9°N, Tica;
by M. Bright.
V.N. IVANENKO
and the Mid-Atlantic Ridge, which came mainly from the sam-
ples obtained during in situ colonization experiments and sedi-
ment traps. Some of these copepods represent common genera
(such as the harpacticoid genus Tisbe and the cyclopoid genus
Heptnerina close to Cyclopina) and even common families (the
harpacticoid family Tegastidae) known till now only from shal-
low waters (IVANENKO & DEFAYE 2004a, b; V. Ivanenko & D.
Defaye, unpublished).
Certainly new methods of meiofauna sampling and explo-
ration of new sites will reveal many new copepods representing
different taxonomical and ecological groups. Further ecological
studies of copepods from different microhabitats, distinct loca-
lities, environments surrounding deep-sea hydrothermal vents
and cold seeps (see HEPTNER & IVANENKO 2002; V. Ivanenko,
D. Defaye & Cuoc, unpublished), as well as meiofauna associa-
ted with whale remnants (to date unknown) will let us better
understand the role of the remarkably diverse copepods in
structuring and functioning of deep-sea chemosynthetic com-
munities.
For more information about taxonomic composition, distri-
bution, morphological, and biological traits, and the presumed
habitat specificity of the copepods from deep-sea chemosynthe-
tic environments we refer to the paper of HEPTNER & IVANEN-
KO (2002) as an attempt to summarize all data available for that
moment.
Denisia 18 (2006): 316–317
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

2: Aphotopontius sp. from East Pacific Rise: 13°N;

by M. Bright.

References:
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta 11(2): 117-134.
HUMES A.G. & M. DOJIRI (1980) Pac. Sci. 34: 143-151.
IVANENKO V.N. & D. DEFAYE (2004a) Cah. Biol. Mar. 45(3): 255-268.
IVANENKO V.N. & D. DEFAYE (2004b) Zoosystema 26(1): 49-64.
IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11(3): 177-185.

317
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Copepoda, Harpacticoida, Aegisthidae

Andromastax CONROY-DALTON & HUYS, 1999

Species Distribution Body length in mm

A. cephaloceratus LEE & HUYS, 2000 Okinawa Trough Female 3.05, male 2.85
A. muricatus CONROY-DALTON & HUYS, 1999 Galapagos Spreding Center Female 3.49, male 3.24

Morphology: Body with reticulated integument. Cephalic
shield, coxae of legs 2-4 with spinous processes. Caudal rami
slightly longer than rest of body. Antennule of female seven-
segmented, second segment with two spine-like processes. An-
tennule of male nine-segmented, segment 8 elongate. Feeding
appendages of male reduced.
Biology: Large benthopelagic copepods with non-feeding
males. A. muricatus has been found in “washing of mussel sam-
ple”. A. cephaloceratus has been collected 0.5-3 m above the
bottom.

1: A. cephaloceratus; A: Male habitus,

dorsal; B: Female cephalosome, dorsal;
from LEE & HUYS (2000).

References:
CONROY-DALTON S. & R. HUYS (1999) J. Crustac. Biol. 19(2): 408-431.
LEE W. & R. HUYS (2000) Zool. J. Linn. Soc. 129: 1-71.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 318

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Ancorabolidae

Uptionyx verenae CONROY-DALTON & HUYS, 2000

Size: Body length of female 1.11 mm. Male unknown. Biology: Two females were found associated with macroinver-
tebrates. Most ancorabolids inhabit mud sediment.
Morphology: Body with dorsal and lateral bulbous processes on
all somites except last two somites. Rostrum subtriangular Distribution: Juan de Fuca Ridge: Middle Valley Segment,
prominence. Antennule three-segmented, with aesthetasc on Peanut Vent.
segments 2 and 3. Antenna three-segmented, without exopod.
Legs 1-4 biramous; endopods of legs 1-4 and exopod of leg 1
two-segmented, exopod of legs 2-4 three-segmented. Leg 5:
long endopod fused with protopod, with four setae; exopod
one-segmented, with five setae.

1: Female; A: Habitus, dorsal; B: Habitus, lateral;

from CONROY-DALTON & HUYS (2000).

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

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 319

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Argestidae

Argestoides prehensilis HUYS & CONROY-DALTON, 1997

Size: Body length of male 0.34 mm. Female unknown. Biology: One male has been found in a “fraction of bucket mus-
sel washings”, bearing, attached to its shield of cephalothorax
Morphology: Prosome four-segmented; urosome six-segmented.
and anal somite, two stages of a tantulocarid, an ectoparasitic
Antennule eleven-segmented, geniculation between segments
crustacean.
8 and 9. Antennary exopod two-segmented, with three setae.
Maxilliped subchelate, three-segmented. Legs 1-4 biramous, Distribution: Galapagos Spreading Center.
with three-segmented rami. Endopod of leg 1 prehensile, first
segment elongate. Leg 5 two-segmented, baseoendopod with
endopodal lobe bearing two spines, exopod with six setae.

1: Male; A: Habitus, dorsal; B: Urosome, ventral; from HUYS & CONROY-DALTON (1997).

Reference:
HUYS R. & CONROY-DALTON S. (1997) Cah. Biol. Mar. 38: 235-249.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 320

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Aegisthidae

Jamstecia terazakii LEE & HUYS, 2000

Size: Body length of female 3.38 mm. Male unknown. Biology: Benthopelagic copepod sampled 0.5-3 m above the
bottom.
Morphology: Body with reticulated integument and paired
spinous processes. Caudal rami 1.5 longer than rest of body. Distribution: Okinawa Trough.
Antennule seven-segmented, first segment elongate. Antenna
with two-segmented exopod, one-segmented endopod shorter
than allobasis. Leg 5 large, one-segmented.

1: Female habitus, lateral;

from LEE & HUYS (2000).

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

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 321

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Aegisthidae

Scabrantenna yooi LEE & HUYS, 2000

Size: Body length of female 3.53 mm; male 3.32 mm. Biology: Benthopelagic copepod sampled 0.5-3 m above the
bottom. Male non-feeding.
Morphology: Body with reticulated integument. Caudal rami
slightly longer than rest of body. Rostrum spiniform, small in fe- Distribution: Okinawa Trough.
male and strongly developed in male. Antennule seven-seg-
mented in female, nine-segmented in male. Antenna with
three-segmented exopod, allobasis and endopod sexually di-
morphic. Mandible and maxillule reduced in male. Leg 1-4 bi-
ramous, with three-segmented rami, excepting one-segmented
endopod of leg 1. Leg 5 almost reaching posterior edge of anal
somite, one-segmented in female and three-segmented in male.

1A: Female habitus, dorsal; B: Male antenna; C: Antenna; from LEE & HUYS (2000).

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

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 322

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Asterocheridae

Cheramomyzon abyssale HUMES, 1989

Size: Body length of female 2.09-2.27 mm. Male unknown. Remark: Has been found in three samples.
Morphology: Prosome four-segmented, tergites of last two Distribution: East Pacific Rise: 13°N.
somites pointed posteriorly. Urosome five-segmented; first
somite with leg 5. Genital double-somite and caudal ramus
elongate. Oral cone elongate. Antennule nineteen-segmented;
segment 17 with aesthetasc. Antenna with short one-segment-
ed exopod. Mandible with stylet-like gnathobase and two-seg-
mented palp bearing two terminal setae. Maxilla two-segment-
ed, distal segment claw-like. Legs 1-4 biramous, with three-seg-
mented rami. Leg 5 two-segmented, distal segment with four se-
tae.

1: Female; A: Habitus, dorsal; B: Habitus, lateral; C: Leg 5 and genital double-somite, ventral; D: Antennule; E: Antenna;
F: Mandible. ae – aesthetasc; gn – gnathobase of mandible; l5 – leg 5; pl – palp of mandible; from HUMES (1989).

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

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 323

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Asterocheridae

Collocherides brychius HUMES, 1999

segment of endopod of legs 1-3 with two inner setae. Leg 5 two-
Size: Body length of female 0.55-0.58 mm; male 0.50-0.51 mm.
segmented, first segment with outer seta, second segment with
Morphology: Prosome four-segmented. Urosome five-segment- four setae in female and five setae in male.
ed in female and six-segmented in male; first somite with leg 5.
Biology: Females and males have been found in washings of
Longest seta of caudal ramus slightly longer (1.18:1) than ra-
tube worms. All congeners are endosymbionts living in the
mus. Oral cone short. Antennule of female twenty-segmented,
stomach of shallow-water ophiuroids (HUMES 1993).
segment 18 with aesthetasc. Antenna with exopod represented
by one short seta. Mandible with stylet-like gnathobase and Distribution: Juan de Fuca Ridge: Cleft Segment, Marker M of
one-segmented palp with one terminal seta. Maxilla three-seg- North Field, 2253 m.
mented. Legs 1-4 biramous, with three-segmented rami. Second

1: Female; A: Antennule; B: Antenna; C: Mandible; D: Maxilla; E: Maxilliped; F: Leg 5; G: Male leg 5.

ex – exopod of antenna represented by seta; gn – gnathobase of mandible; pl – palp of mandible; from HUMES (1999).

References:
HUMES A.G.(1993) Bijdr. Dierkd. 63: 121-127.
HUMES A.G. (1999) Proc. Biol. Soc. Wash. 112(1): 181-188.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 324

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

Arthropoda, Crustacea, Copepoda, Cyclopoida, Clausidiidae

Hyphalion captans HUMES, 1987

mented, second segment with long terminal seta; maxilliped of

Size: Body length of female 1.94-2.17 mm; male 1.71-1.82 mm.
male subchelate, four-segmented, second segment triangular
Morphology: Body elongate, flattened. Prosome four-segment- with two rows of spinules, third segment not well defined,
ed. Urosome six-segmented in female, five-segmented in male; fourth segment claw-like. Legs 1-4 biramous with three-seg-
first somite with leg 5. Rostrum broad, projecting anterior. An- mented rami. Leg 5 two-segmented, first segment with one se-
tennule six-segmented in both sexes; male antennules non- ta; second segment with three spines and one seta.
geniculate. Antenna three-segmented, without exopod; distal
Biology: Symbiont of bivalves; type of feeding unclear, may
segment with three stout recurved claws. Mandible flexed,
feed on tissues, mucus or utilize host’s food.
gnathobase with three spines and one seta; palp absent. Lobe of
maxillule with five setae. Maxilla two-segmented; first segment Distribution: Guaymas Basin.
with one seta; second segment with one seta and three spines.
Maxilliped sexually dimorphic; maxilliped of female two-seg-

1: Female; A: Habitus, dorsal; B: Antennule; C: Antenna; D: Maxilliped; E: Male habitus, dorsal; F: Maxilliped; from HUMES (1987).

References:
DEFAYE D. & T. TODA (1994) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 16, section A (1): 87-94.
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 325

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

Arthropoda, Crustacea, Copepoda, Cyclopoida, Cyclopinidae

Barathricola rimensis HUMES, 1999

1-4 biramous, with three-segmented rami. Leg 1 with one seta

Size: Body length of female 0.67-0.79 mm; male 0.47-0.57 mm.
on inner segment. Leg 5 three-segmented with intercoxal scle-
Morphology: Prosome slender, five-segmented. Urosome five- rite: armature 0-0; 1-0; II, 1, I in female and 0-0; 1-0; II, 1, I, 1
segmented in female, six-segmented in male, first somite with in male.
leg 5. Caudal ramus long, of length/width ratio 11:1. Anten-
Distribution: Juan de Fuca Ridge: Coaxial Segment, Flow vent
nule thirteen-segmented; geniculation of male antennules be-
site (Vent HDV).
tween segments 11 and 12. Antenna four-segmented, without
exopod. Mandible with biramous palp: endopod two-segment-
ed; exopod small, one-segmented, with two terminal setae. Legs

1: Female; A: Habitus, dorsal; B: Urosome, dorsal; C: Antenna; D: Mandible; E: Maxilliped; F: Leg 5; from HUMES (1999).

Reference:
HUMES A.G. (1999) J. Nat. Hist. 33: 961-978.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 326

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

Arthropoda, Crustacea, Copepoda, Cyclopoida, Cyclopinidae

Heptnerina confusa IVANENKO & DEFAYE, 2004

Size: Body length of female 0.82 mm; male 0.65 mm. Biology: Many adult and subadult copepodid stages of H. con-
fusa have been collected during in situ colonization experi-
Morphology: Prosome five-segmented; tergite of somite bearing
ments at the base of the active chimney Eiffel Tower covered by
leg 1 covered laterally by shield of cephalosome. Urosome five-
a layer of Bathymodiolus azoricus (Bivalvia, Mytilidae).
segmented in female, six-segmented in male; first somite with
leg 5. Antennule ten-segmented in female and seventeen-seg- Distribution: Mid-Atlantic Ridge: Lucky Strike, Chimney Eif-
mented in male. Antenna four-segmented; exopod represented fel Tower.
by two setae on first segment. Mandible with biramous palp.
Legs 1-4 biramous, with three-segmented rami. Endopod of leg
4 with three robust setae. Leg 5 two-segmented with armature
1-0; I, 1, I in female and 1-0; I, 1, I, 2 in male. First segment of
female leg 5 can have stout endopodal element.

1: Female; A: Habitus, dorsal; B: Antennule; C: Antenna; D: Leg 5, endopodal element arrowed; E: Male habitus, dorsal;
from IVANENKO & DEFAYE (2004).

Reference:
IVANENKO V.N. & D. DEFAYE (2004) Zoosystema 26 (1): 49-64.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 327

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Aphotopontius HUMES, 1987

Species 1 2 3 4 5 6 7 8 9 10 11
A. acanthinus HUMES & LUTZ, 1994 +
A. arcuatus HUMES, 1987 + + +
A. atlanteus HUMES, 1996 + +
A. baculigerus HUMES, 1987 + +
A. flexispina HUMES, 1987 +
A. forcipatus HUMES, 1987 + + + +
A. hydronauticus HUMES, 1989 +
A. limatulus HUMES, 1987 + + +
A. mammillatus HUMES, 1987 + + + +
A. probolus HUMES, 1990 +

Size: Body length 0.6-1.2 mm.
Morphology: Prosome four-segmented. Urosome five-segment-
ed in female, six-segmented in male; first somite with leg 5.
Oral cone short. Antenna with one-segmented exopod.
Mandible with stylet-like gnathobase, without palp. First seg-
ment of maxilla with inner sinuous seta. Exopods of legs 1-4
and endopods of legs 1-3 three-segmented. Endopod of leg 4
two-segmented; first segment with one inner seta; second seg-
ment with two (terminal and inner) setae.
Remarks: A. rapunculus HUMES & SEGONZAC, 1998 transferred
to Rhogobius HUMES, 1987; A. temperatus HUMES, 1997 was syn-
onymized with A. atlanteus. Only females are known for A. flex-
ispina and A. hydronauticus.
Biology: All species of Aphotopontius have been found free-liv-
ing or associated with invertebrates. A. arcuatus, A. atlanteus,
A. flexispina and A. probolus found in association with bivalves;
A. arcuatus, A. flexispina and A. mammillatus with vestimen-
tiferans; A. acanthinus and A. forcipatus with crustaceans. Eco-
logical studies at the Juan de Fuca Ridge showed that A. forci-
patus is more abundant at new vents appearing after eruption
and has female to male ratio 10.6:1 (TSURUMI et al. 2003).
Functional analysis suggests that copepods of the genus Aphoto-
pontius are swimming and moving over substratum, they can
feed on bacterial films and flakes on substratum and in near-
bottom water column (HEPTNER & IVANENKO 2002).
Distribution: See the table after Ivanenko & Ferrari, 2003: 1 –
Galapagos Rift; 2 – East Pacific Rise at 10°N; 3 – East Pacific
Rise at 13°N; 4 – East Pacific Rise at 21°N; 5 – Guaymas Basin,
27°N; 6 – Gorda Ridge, 41°N; 7 – Juan de Fuca Ridge, 46°N; 8
– Explorer Ridge, 49°N; 9 – Mid-Atlantic Ridge at 37°N
(Lucky Strike); 10 – Mid-Atlantic Ridge at 37°N (Menez
Gwen); 11 – Mid-Atlantic Ridge at 23°N (Snake Pit).

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 328-329

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

1: Female of A. mammillatus; A: Habitus,

dorsal; B: Habitus, lateral; C: Antenna;
D: Mandible; E: Maxilla; F: Leg 4.
b – basis; c – coxa; en1-en2 – endopodal
segments; ex1-ex3 – exopodal segments 1-3;
gn – gnathobase of mandible;
p1-p4 – segments of prosome; ss – sinuous
seta of maxilla; u1-u5 – segments of urosome;
from IVANENKO & HEPTNER (1998).

2: Female of A. atlanteus (SEM);

A: Habitus, dorsal; B: Habitus, ventral;
C: Distal part of oral cone; D: Urosome, dorsal.
es – egg sac; lb – distal part of labrum
of oral cone, oral cone, distal part of labrum;
lm – distal part of labium of oral cone
ornamented with setiform elements;
md – mandible, tip of gnathobase; original.

References:
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta 11(2): 117-134.
HUMES A.G. (1987) Bull. Mar. Sci. 41(3): 645-788.
HUMES A.G.(1990) Sci. Mar. 54(2): 145-154.
HUMES A.G. & R.A. LUTZ (1994) Jour. Crustac. Biol. 14(2): 337-345.
HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.
IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11(3): 177-185.
IVANENKO V.N & M.V. HEPTNER (1998) J. Mar. Syst. 15: 243-254.
TSURUMI M., DE GRAAF R.C. & V. TUNNICLIFFE (2003) J. Mar. Biol. Ass. U.K. 83(3): 469-478.

329
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Benthoxynus HUMES, 1984

Species Distribution Body length in mm
B. spiculifer HUMES, 1984 Juan de Fuca Ridge: Axial Seamount, Heineken Hollow, Female 1.61-1.79; male 0.97-1.09
Megaplume Site; Gorda Ridge; Explorer Ridge
B. tumidiseta HUMES, 1989 East Pacific Rise: 13°N, sites Genesis, Elsa, Totem Female 1.53-1.76; male unknown

Morphology: Prosome four-segmented. Urosome five-segment-
ed in female, six-segmented in male; first somite with leg 5.
Caudal ramus elongate; innermost terminal seta minute. Oral
cone short. Antenna with one-segmented exopod. Mandible
with stylet-like gnathobase, without palp. First segment of max-
illa with inner sinuous seta. Exopods of legs 1-4 and endopods
of legs 1-3 three-segmented. First exopodal segment of leg 3
lacks inner seta. Leg 4 with two-segmented endopod; first seg-
ment unarmed; second segment armed with terminal spine. Leg
5: small lobe with 3 setae in female and two setae in male.
Biology: B. spiculifer has been found in debris from vestimen-
tiferan tubes. Ecological studies at the Juan de Fuca Ridge re-
vealed that it is more common at older vents (TSURUMI et al.
2003). The blood of B. spiculifer has haemoglobin supporting
aerobic respiration in low-oxygen conditions (HOURDEZ et al.
2000).

1: Female of B. tumidiseta; A: Habitus, dorsal; B: Anal somite and elongate caudal ramus with reduced innermost terminal seta,
dorsal; C: Antennule; D: Maxilla; female of B. spiculifer; E: Antennule; F: Leg 4. is – innermost terminal seta of caudal ramus;
from HUMES (1989) (A-D) and HUMES (1984) (E, F).

References:
HOURDEZ S., LAMONTAGNE J., PETERSON P., WEBER R.E. & C.R. FISHER (2000) Biol. Bull. 199: 95-99.
HUMES A.G. (1984) Can. J. Zool. 62: 2594-2599.
HUMES A.G. (1989) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 11, section A, no. 4: 829-849.
TSURUMI M., DE GRAAF R.C. & V. TUNNICLIFFE (2003) J. Mar. Biol. Ass. U.K. 83(3): 469-478.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 330

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Ceuthoecetes HUMES & DOJIRI, 1980

two-segmented endopod; first segment unarmed; second seg-

Type species: C. aliger HUMES & DOJIRI, 1980. Other included
ment armed with terminal spine and inner seta. Maxilliped of
species: C. acanthothrix HUMES, 1987; C. cristatus HUMES, 1987;
male with stout second segment.
C. introversus HUMES, 1987.
Remarks: Only female was described for C. aliger; only males
Size: Body length 0.91-1.50 mm.
are known for three other species.
Morphology: Prosome four-segmented. Urosome five-segment-
Biology: Abundant in washings of vestimentiferans and Calyp-
ed in female, six-segmented in male; first somite with leg 5.
togena. Type of feeding unclear, may feed on tissues or associat-
Oral cone short, with distal disk formed by labium. Antenna
ed mucus of their hosts. Ceuthoecetes aliger was found in wash-
with small one-segmented exopod; endopod with curved termi-
ing of tentacular crowns of vestimentiferans.
nal claw. First segment of maxilla without inner sinuous seta;
second segment elongate, with distal setiform part. Exopods of Distribution: East Pacific Rise, Galapagos Spreading Center.
legs 1-4 and endopods of legs 1-3 three-segmented. Leg 4 with

1: Female of C. aliger; A: Habitus, dorsal; B: Antenna;
C: Maxilla; D: Leg 4; E: Male of C. cristatus, habitus, dorsal;
F: Male of C. introversus, maxilla.
ex – exopod; tc – terminal claw of endopod;
from HUMES & DOJIRI (1980) (A-D) and HUMES (1987) (F).
2: Male of C. introversus (SEM). A: Habitus, dorsal;
B: Habitus, ventral; C: Oral cone, ventral.
a1 – antennule; cr – caudal ramus; dd – distal disk of
oral cone formed by labium; l1-l3 – swimming legs
1-3. lb – labium; m1 – maxillule; m2 – maxilla;
mp – maxilliped; p1-p4 – segments of prosome;
u1-u6 – somites of urosome; original.

References:
HUMES A.G. (1987) Bull. Mar. Sci. 41(3): 645-788.
HUMES A.G. & M. DOJIRI (1980) Proc. Biol. Soc. Wash. 93(3): 697-707.
HUMES A.G. & M. SEGONZAC (1998) Cah.Biol. Mar. 39: 51-62.
IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11(3): 177-185.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 331

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Chasmatopontius thescalus HUMES, 1990

Size: Body length of female 1.14-1.39 mm; male 1.06-1.28 mm. Biology: Abundant. Found in washings of tubes of Paralvinella
hessleri DESBRUYÈRES & LAUBIER, 1989 in the Mariana Back-
Morphology: Prosome four-segmented. Urosome four-segment-
Arc Basin.
ed in female, five-segmented in male; first somite with leg 5.
Copulatory pores near posterior border of female genital dou- Distribution: Mariana Back-Arc Basin: Alice Spring, Illium;
ble-somite. Oral cone short. Antenna with one-segmented ex- Lau Back-Arc Basin: Vailili.
opod. Mandible with stylet-like gnathobase, without palp. First
segment of maxilla with inner sinuous seta. Exopods of legs 1-4
and endopods of legs 1-3 three-segmented. Leg 4 with two-seg-
mented endopod; first segment unarmed; second segment with
three inner setae. Second segment of male maxilliped expand-
ed, with knob on inner edge.

1: Female; A: Habitus, dorsal; B: Urosome, dorsal; C: Maxilla; D: Male leg 4, anterior; E: Maxilliped, posterior.
cp – copulatory pore; kb – knob on inner edge of male maxilliped; from HUMES (1990).

References:
HUMES A.G. (1990) J. Nat. Hist. 24: 289-304.
HUMES A.G. (1991) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 13, section A, nos. 1-2: 121-134.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 332

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Dirivultus HUMES & DOJIRI, 1980

dopods of legs 2-3 three-segmented; endopods of legs 1 two- or

Type species: D. dentaneus HUMES & DOJIRI, 1980; one further
species D. spinigulatus HUMES, 1999.
Size: Body length of female 1.00-1.16 mm; male 0.92-1.09 mm.
Morphology: Prosome four-segmented. Urosome five-segment-
ed in female, six-segmented in male; first somite with leg 5.
Rostral area projecting anteriorly. Oral cone with distal disk
formed by labium; labrum with dentiform spines directed pos-
teriorly. Antenna with one-segmented exopod; endopod with
curved terminal claw. Mandible with stylet-like gnathobase,
without palp. First segment of maxilla without inner sinuous se-
ta. Second segment of maxilla armed with long slender claw
and ornamented with long setules. Exopods of legs 1-4 and en-
three-segmented. Leg 4 with two-segmented endopod; first seg-
ment unarmed; second segment armed with terminal spine and
inner seta.
Biology: May “feed on tentacular lamellae or associated mucus”
of siboglinids (HUMES & DOJIRI 1980). The type species D. den-
taneus HUMES & DOJIRI, 1980 has been found in tentacular
crown of Lamellibrachia barhami WEBB, 1969 off Southern Cali-
fornia.
Distribution: D. dentaneus from Southern California
(32°19’6’’N, 117°19’08’’W; 1125 m); D. spinigulatus from
Tabar-Feni Volcanic Fore-Arc: Edison Seamount.

1A: Female habitus, dorsal; B: Habitus lateral; C: Antenna, posterior; D: Maxilla; E: Leg 4; F: Male habitus, dorsal; G: Female
habitus, dorsal; H: Egg sac. ec – egg sac; ex – exopod; s2 – second segment of maxilla ornamented with setules; tc – terminal
claw of endopod; from HUMES & DOJIRI (1980) (A-F D. dentaneus) and HUMES (1999) (G-H D. spinigulatus, female).

References:
HUMES A.G. (1999) J. Mar. Biol. Ass. U.K. 79: 1053-1060.
HUMES A.G. & M. DOJIRI (1980) Pac. Sci. 34: 143-151.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 333

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Exrima HUMES, 1987

Species Distribution Body length in mm

E. dolichopus HUMES, 1987 East Pacific Rise: 13°N, Parigo Female 1.12-1.15; male unknown
E. singular HUMES, 1987 East Pacific Rise: 21°N, Clam Acres Female 0.97-1.07; male unknown

Morphology: Prosome four-segmented. Urosome five-segment- Biology: E. singula has been found in washing of vestimentifer-
ed; first somite with leg 5. Oral cone short. Antenna with one- ans and Calyptogena.
segmented exopod; distal segment of endopod with two long
terminal setae. Mandible with stylet-like gnathobase, without
palp. First segment of maxilla with inner sinuous seta. Exopods
of legs 1-4 and endopods of legs 1-3 three-segmented. Leg 4
with two-segmented endopod; first segment unarmed; second
segment armed with terminal spine. Leg 5 two-segmented.

1: Female of E. singula; A: Habitus, dorsal; B: Antenna; C: Maxilla; D: Leg 4; E: Female of E. dolichopus, habitus, dorsal;
from HUMES (1987).

References:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
HUMES A.G. (1989) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 11, section A, no. 4: 829-849.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 334

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Fissuricola caritus HUMES, 1987

Size: Body length of female 1.19-1.34 mm. Male unknown. Remark: The only dirivultid with maxilla reduced to vestige.
Morphology: Prosome four-segmented. Urosome five-segment- Distribution: East Pacific Rise: 21°N, Clam Acres.
ed; first somite with leg 5. Oral cone short. Antenna with one-
segmented exopod. Mandible with stylet-like gnathobase, with-
out palp. Maxilla reduced to small vestige. Exopods of legs 1-4
and endopods of legs 1-3 three-segmented. Leg 4 with two-seg-
mented endopod; first segment with inner seta; second segment
with one terminal spine and three inner setae.

1: Female; A: Habitus, dorsal; B: Antenna; C: Maxilla; D: Leg 4; E: Leg 5; from HUMES (1987).

References:
HUMES A.G. (1987) Bull. Mar. Sc. 41: 645-788.
IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11 (3): 177-185.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 335

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Humesipontius arthuri IVANENKO & FERRARI, 2003

Remark: The only dirivultid missing endopod of leg 4.

Size: Body length of female 1.8 mm. Male unknown.
Distribution: Juan de Fuca Ridge.
Morphology: Body elongate. Prosome four-segmented. Uro-
some five-segmented; first somite with leg 5. Oral cone short.
Antenna with one-segmented exopod; second segment of en-
dopod with elongate terminal seta. Mandible with stylet-like
gnathobase, without palp. First segment of maxilla with inner
sinuous seta. First segment of maxillipedal endopod with elon-
gate seta. Exopods of legs 1-4 and endopods of legs 1-3 three-
segmented. Leg 4 without endopod.

1: Female; A: Habitus, dorsal; B: Antenna; C: Maxilla; D: Maxilliped, posterior; E: Leg 4, anterior; after IVANENKO & FERRARI (2003).

Reference:
IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11(3): 177-185.

V.N. IVANENKO & D. DEFAYE Denisia 18: 336

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Nilva torifera HUMES, 1987

Size: Body length of female 0.67-0.88 mm, male 0.79-0.96 mm. Biology: Common in washings of vestimentiferans and Calyp-
togena; type of feeding unclear, may feed on tissues or mucus of
Morphology: Prosome four-segmented. Urosome five-segment-
hosts.
ed in female, six-segmented in male; first somite with leg 5; last
two somites indistinctly separated. Genital double-somite of fe- Distribution: Galapagos Spreading Center; East Pacific Rise:
male and genital somite of male with dorsal hump. Oral cone 21°N, Clam Acres: 13°N, Parigo, Totem, Genesis.
short, with distal disk formed by labium. Antenna with small
one-segmented exopod; endopod with curved terminal claw.
First segment of maxilla without inner sinuous seta; second seg-
ment elongate with setiform distal part. Exopods of legs 1-4 and
endopods of legs 1-3 three-segmented. Leg 4 with two-seg-
mented endopod; first segment unarmed; second segment
armed with inner seta and terminal spine.

1: Female; A: Habitus, dorsal; B: Urosome, lateral; C: Antenna; D: Maxilla; E: Male habitus, dorsal; F: Urosome, lateral.
dh – dorsal hump; tc – terminal claw of antennal endopod; from HUMES (1987).

Reference:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 337

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Rhogobius HUMES, 1987

Species Distribution Body length in mm
R. contractus HUMES, 1987 Galapagos Spreading Center; East Pacific Rise: 13°N, 21° Female 0.7-0.94, male 0.6-0.71
R. pressulus HUMES, 1989 Galapagos Spreading Center Female 0.67-0.78; male unknown
R. rapunculus (HUMES & SEGONZAC, 1998) East Pacific Rise: 9°N Female 0.89-0.91; male unknown

segmented endopod; first segment with inner seta; second seg-

Morphology: Body with broad prosome. Prosome four-segment-
ment with terminal spine and inner seta.
ed. Urosome five-segmented in female, six-segmented in male;
first somite with leg 5. Last abdominal somite with terminal Remark: R. rapunculus (HUMES & SEGONZAC, 1998) transferred
paired lobes. Oral cone short. Antenna with one-segmented from Aphotopontius.
exopod; second segment of endopod elongate and slender.
Biology: R. contractus has been found in number of samples
Mandible with stylet-like gnathobase, without palp. First seg-
(HUMES & SEGONZAC 1998). Only two females of R. rapunculus
ment of maxilla without inner sinuous seta. Exopods of legs 1-
and 10 females of R. pressulus have been found until now.
4 and endopods of legs 1-3 three-segmented. Leg 4 with two-

1: Female of R. contractus.;
A: Habitus, dorsal; B: Genital double-
somite, abdominal somites
and caudal rami; C: Antenna;
D: Maxilla; E: Leg 5, male; F: Habitus,
dorsal; G: R. rapunculus. Female
leg 5; H: Habitus, dorsal; I: Leg 4;
J: Female of R. pressulus, leg 5;
K: Genital double-somite and
abdominal somites.
s2 – second segment of endopod;
tl – terminal lobe on anal somite;
from HUMES (1987) (A-G), HUMES &
SEGONZAC (1998) (H-J) and HUMES
(1989) (K).

References:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
HUMES A.G. (1989) Pac. Sci. 43: 27-31.
HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 338

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Rimipontius mediospinifer HUMES, 1996

Size: Body length of female 0.84-1.01 mm; male 0.57-0.69 mm. Biology: Found in washing of the decapod Rimicaris (HUMES
1996) and in plankton 80-300 m over the hydrothermal field
Morphology: Prosome four-segmented. Urosome five-segment-
among shrimps and other copepods (IVANENKO 1998). It can
ed in female, six-segmented in male; first somite with leg 5.
“feed on the bacteria on the mouthparts of the shrimps, but al-
Genital double-somite of female with dorsal longitudinal crest.
so, when they are free, perhaps on the bacteria free in the wa-
Caudal rami with 5 setae: innermost terminal seta reduced.
ter” (HUMES 1996).
Oral cone short. Antenna with one-segmented exopod.
Mandible with stylet-like gnathobase, without palp. First seg- Distribution: Mid-Atlantic Ridge: Logatchev, Irina-2; Snake
ment of maxilla with inner sinuous seta. Exopods of legs 1-4 Pit, Broken Spur (80-300 m over Saracen’s Head).
and endopods of legs 1-3 three-segmented. Leg 4 with two-seg-
mented endopod, first segment unarmed; second segment with
terminal spine.

1: Female; A: Habitus, dorsal; B: Habitus, lateral; C: Anal somite and caudal ramus, dorsal; D: Maxilla; E: Leg 4;
F: Male habitus, dorsal. cr – caudal ramus; ds – dorsal spiniform process of genital double somite; en – endopod of leg 4;
ss – sinuous seta of maxilla; from HUMES (1996).

References:
HUMES A.G. (1996) Bull. Mar. Sci. 58: 609-653.
HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.
IVANENKO V.N. (1998) Zool. Zh. 77(11): 1249-1256.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 339

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Scotoecetes introrsus HUMES, 1987

Size: Body length of female 1.65-1.79 mm; length of male 1.30- Biology: Abundant, associated with siboglinids, feeding on
1.42 mm. bacterial flakes; different copepodid stages have been found in
washings of vestimentiferans collected at 9°N (unpublished da-
Morphology: Prosome four-segmented. Urosome five-segment-
ta). The blood of S. introrsus as well as that of Benthoxynus spi-
ed in female, six-segmented in male; first somite with leg 5.
culifer has haemoglobin supporting aerobic respiration in low-
Oral cone short. Antenna with one-segmented exopod.
oxygen conditions (SELL 2000).
Mandible with stylet-like gnathobase, without palp. First seg-
ment of maxilla with inner sinuous seta. Exopods of legs 1-4 Distribution: East Pacific Rise: 9°N, 13°N.
and endopods of legs 2-3 three-segmented. Endopod of leg 1
two-segmented in female and three-segmented in male; distal
segment of endopod with outer terminal seta directed inward.
Leg 4 with two-segmented endopod; first segment unarmed,
second segment with terminal spine and inner seta.

1: Female; A: Habitus, dorsal; B: Maxilla; C: Leg 1; D: Leg 4; E: Male habitus, dorsal.

lb – labrum of oral cone; a2 – antenna; m1 – maxillule; m2 – maxilla; from HUMES (1987).

References:
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta, 11(2): 117-134.
HUMES A.G.(1987) Bull. Mar. Sci. 41: 645-788.
HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.
SELL A.F. (2000) Proc. R. Soc. London 267: 2323-2326.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 340

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Dirivultidae

Stygiopontius HUMES, 1987

1 2 3 4 5 6 7 8 9 10 11 12 13
S. appositus + +
S. brevispina +
S. cinctiger + + +
S. cladarus + +
S. flexus + +
S. hispidulus + + +
S. latulus +
S. lauensis +
S. mirus + +
S. mucroniferus + +
S. paxillifer + +
S. pectinatus + + + +
S. quadrispinosus + + +
S. regius +
S. rimivagus +
S. sentifer + + +
S. serratus +
S. stabilitus + +
S. teres +
S. verruculatus + +

Size: Body length 0.6-1.6 mm.
Morphology: Prosome four-segmented. Urosome five-segment-
ed in female, six-segmented in male; first somite with leg 5.
Oral cone short. Antenna with one-segmented exopod.
Mandible with stylet-like gnathobase, without palp. First seg-
ment of maxilla with inner sinuous seta. Exopods of legs 1-4
and endopods of legs 1-3 three-segmented. Leg 4 with two-seg-
mented endopod; first segment unarmed; second segment
armed with terminal spine and inner seta.
Remarks: S. lumiger HUMES, 1989 and S. bulbisetiger HUMES,
1996 were synonymized with S. sentifer and S. pectinatus, re-
spectively. Only females are known for eight species (S.
cinctiger, S. flexus, S. hispidulus, S. mucroniferus, S. pectinatus, S.
sentifer, S. stabilitus, and S. teres), and only males are known for
six species (S. appositus, S. latulus, S. mirus, S. paxillifer, S. rimi-
vagus, and S. verruculatus).
Biology: All species of the genus Stygiopontius have been found
free-living or associated with invertebrates. S. hispidulus and S.
rimivagus have been found in association with bivalves; S.
hispidulus, S. sentifer, and S. stabilitus with polychaetes; S. flexus
with siboglinids; S. cladarus and S. pectinatus with crustaceans.
S. sentifer lives in tubes of polychaete Alvinella pompejana DES-
BRUYÈRES & LAUBIER, 1980. S. pectinatus has been found in gill
cavity of the shrimps Rimicaris exoculata WILLIAMS & RONA,
1986 and Chorocaris chacei (WILLIAMS & RONA, 1986) and free-
living in plankton over hydrothermal field (IVANENKO 1998). S.
quadrispinosus is more abundant at high temperature at the Juan
de Fuca Ridge and has the ratio of females to males 7.6:1 (TSU-
RUMI et al. 2003).
Distribution: See the table after Ivanenko & Ferrari (2003): 1
– East Pacific Rise at 10°N; 2 – East Pacific Rise at 13°N; 3 –
East Pacific Rise at 21°N; 4 – Guaymas Basin, 27°N; 5 – Gor-
da Ridge, 41°N; 6 – Juan de Fuca Ridge, 46°N; 7 – Explorer
Ridge, 49°N. WP – West Pacific: 8 – Mariana Back-Arc Basin,
18°N; 9 – Lau Back-Arc Basin, 23°S; 10 – Mid-Atlantic Ridge
at 37°N (Lucky Strike); 11 – Mid-Atlantic Ridge at 29°N (Bro-
ken Spur); 12 – Mid-Atlantic Ridge at 26°N (TAG); 13 – Mid-
Atlantic Ridge at 23°N (Snake Pit).

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 341–342

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

1: Female of S. quadrispinosus;
A: Habitus, dorsal; B: Maxilla;
C: Leg 4; D: Male habitus,
dorsal; from HUMES (1987).

2: S. pectinatus ventral view (left)

and dorsal view (right); original.

References:
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta 11(2):117-134.
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
IVANENKO V.N. (1998) Zool. Zh. 77 (11): 1249-1256.
IVANENKO V.N. & F.D. FERRARI (2003) Arthropoda Selecta 11 (3): 177-185.
TSURUMI M., DE GRAAF R.C. & V. TUNNICLIFFE (2003) J. Mar. Biol. Assoc. U.K. 83(3): 469-478

342
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Ecbathyriontidae

Ecbathyrion prolixicauda HUMES, 1987

mous, with three-segmented rami. Basis of leg 1 without inner

Size: Body length of female 1.01-1.11 mm; male 0.77-0.84 mm.
spine or seta. Leg 5: first segment fused with somite and repre-
Morphology: Prosome four-segmented. Urosome four-segment- sented with one outer seta; second segment bearing three setae
ed in female and five-segmented in male; first somite with leg in female, and five setae in male.
5. Caudal ramus elongate. Oral cone short. Antennule eight-
Remarks: Found in many samples.
een-segmented in female and seventeen-segmented in male;
aesthetasc on segments 15 and 17 in female and male, respec- Distribution: Galapagos Rift; East Pacific Rise: 9°N, 13°N,
tively. Antenna with one-segmented exopod having pointed 21°N.
projection. Mandible with stylet-like gnathobase, without palp.
First segment of maxilla with inner sinuous seta. Legs 1-4 bira-

1: Female; A: Habitus, dorsal; B: Urosome, dorsal; C: Antennule; D: Antenna; E: Maxilla; F: Leg 4; G: Male habitus, dorsal.
ae – aesthetasc; cr – caudal ramus; ds – genital double-somite; en – endopod; ex – exopod; ss – sinuous seta; from HUMES (1987).

References:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
HUMES A.G. & M. SEGONZAC (1998) Cah. Biol. Mar. 39: 51-62.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 343

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

Arthropoda, Crustacea, Copepoda, Cyclopoida, Erebonasteridae

Ambilimbus IVANENKO, DEFAYE & HUYS, 2005

Species Distribution Body length in mm

A. altalis (HUMES & HUYS, 1992) Juan de Fuca Ridge Female 2.41-2.46; male 1.98-2.13
A. arcuscelestis IVANENKO, DEFAYE & HUYS, 2005 Mid-Atlantic Ridge: Rainbow Female 2.35; male 1.65
A. tuerkayi (MARTINEZ-ARBIZU, 1999) Fiji Basin Female 1.51; male 1.2

Maxilla two-segmented, distal segment claw-like, with inner

Synonym: Amphicrossus HUYS, 1991
process. Maxilliped four-segmented, sexually dimorphic, distal
Morphology: Body elongate and flattened, ornamented with segment claw-like, with inner process. Legs 1-4 biramous with
spinules. Prosome five-segmented, epimera of somites bearing three-segmented rami. Leg 5 two-segmented.
legs 1-4 rounded. Urosome five-segmented in both sexes; first
Biology: Deep-sea poecilostomatoid cyclopoids found in sedi-
somite with leg 5; copulatory pore on ventral side of genital
ment. Most likely associated with bivalves. Type of feeding un-
double-somite. Rostrum prominent with row of dentiform spin-
clear; can feed on tissues, mucus or utilize hosts food. Ere-
ules and long setules. Antennule six-segmented in both sexes.
bonaster protentipes HUMES, 1987 has been found in mantle cav-
Antenna four-segmented, without exopod. Mandibular
ity and in washings of a Nuculana-like protobranch bivalve
gnathobase with four spines, mandibular palp one-segmented
from cold seeps at Guaymas Basin.
with four setae. Maxillule represented by lobe with five setae.

1: A. arcuscelestis, female; A: Habitus, dorsal; B: Mandible; C: Maxilla; D: Maxilliped;

E: Male habitus, dorsal; F: Maxilliped; from IVANENKO et al. (2005).

References:
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
HUMES A.G. & R. HUYS (1992) Can. J. Zool. 70: 1369-1380.
IVANENKO V.N., DEFAYE D. & R. HUYS (2005) Marine Biology Research 19: 93-105.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 344

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Laophontidae

Bathylaophonte LEE & HUYS, 1999

Species Distribution Body length in mm
B. azorica LEE & HUYS, 1999 Mid-Atlantic Ridge: Menez Gwen, Lucky Strike Female 0.69; male 0.66
B. pacifica LEE & HUYS, 1999 East Pacific Rise:17°S north of Easter Island Female 0.81; male 0.74

hensile, first segment longer than exopod, first segment un-

Morphology: Body elongate, sub-cylindrical, with reticulated
armed, second segment with claw and one seta. Leg 5 two-seg-
integument and spinules. Prosome four-segmented; urosome
mented; endopodal lobe of female well developed, with four se-
five-segmented in female and six-segmented in male; genital
tae; endopodal lobe of male reduced, with one seta; exopod
double-somite of female subdivided dorsally. Rostrum bell-
with five setae in female and four setae in male.
shaped. Anal operculum well-developed. Antennule seven-seg-
mented in female, eight-segmented in male. Antenna with Biology: Both species were “extracted from the invertebrates
one-segmented exopod bearing four setae. Maxilliped sub- living in the hydrothermal fluids”.
chelate, three-segmented. Legs 1-4 biramous, with three-seg-
Distribution: East Pacific Rise: 17°S north of Easter Island.
mented exopods and two-segmented endopods, except male
with three-segmented endopod of leg 3. Endopod of leg 1 pre-

1: B. azorica; A: Female habitus, lateral; B: Leg 5, anterior; C: Male leg 5, anterior; D: Female of B. pacifica, leg 1,
anterior; en1 – first endopodal segment; from LEE & HUYS (1999).

Reference:
LEE W. & R. HUYS (1999) Cah. Biol. Mar. 40: 293-328.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 345

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

Arthropoda, Crustacea, Copepoda, Cyclopoida, Lubbockiidae

Laitmatobius crinitus HUMES, 1987

Size: Body length of male 1.29-1.38 mm. Female unknown. Biology: More than 60 males of L. crinitus have been found in
two samples. Other family members occur in plankton at dif-
Morphology: Body slender. Prosome five-segmented. Urosome
ferent depths and probably associated with pelagic inverte-
six-segmented, first somite with leg 5, second somite elongate.
brates (HUYS & BÖTTGER-SCHNACK 1997 cited in BOXSHALL &
Rostrum rounded lobe in lateral view. Caudal ramus elongate,
HALSEY 2004).
length/width ratio 2.7:1. Antennule five-segmented. Antenna
four-segmented with four terminal setiform claws and two sub- Distribution: Guaymas Basin.
terminal setae. Mandible elongate, without palp; basal part
with two spines and one seta, distal part forming long lash.
Maxillule bilobed. Maxilla two-segmented; second segment
with one seta and two spines. Maxilliped subchelate, four-seg-
mented. Legs 1-4 biramous, with three-segmented rami; basis of
leg 1 with inner spine. Leg 5 represented by small segment bear-
ing two setae and adjacent seta.

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

References:
BOXSHALL G.A & S.H. HALSEY (2004) An Introduction to Copepod Diversity. Ray Society, London: i-xv, 1-966.
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 346

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

Arthropoda, Crustacea, Copepoda, Siphonostomatoida, Megapontiidae

Hyalopontius boxshalli HUMES, 1988

Maxilliped subchelate, three-segmented with long terminal

Size: Body length of male 4.75-4.86 mm. Female unknown.
claw. Legs 1-4 biramous, with three-segmented rami. Leg 5 one-
Morphology: Body elongate, 5.7 times as long as wide. Prosome segmented, armed with three setae; one outer seta near seg-
four-segmented, epimera pointed posteriorly. Urosome six-seg- ment.
mented, first somite with leg 5. Oral cone elongate. Antennule
Biology: Two males have been found only. Functional analysis
eleven-segmented. Antenna with short one-segmented exopod
suggests that megapontiids suck out blood or soft tissues of un-
bearing three long setae and two-segmented endopod bearing
known hosts (HEPTNER 1968).
two unequal terminal setae. Mandible with stylet-like
gnathobase, without palp. Maxillule with lobe bearing three se- Distribution: Galapagos Spreading Center.
tae and one adjusted to lobe seta. Maxilla two-segmented.

1: Male; A: Habitus, dorsal; B: Oral cone, lateral;

C: Antenna; from HUMES (1988).

References:
HEPTNER M.V. (1968) Zool. Zh. 47(11):1628-1638.
HUMES A.G. (1988) Proc. Biol. Soc. Wash. 101(4): 825-831.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 347

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Miraciidae

Stenhelia gundulae WILLEN, 2003

baseoendopod with one outer and four or two inner setae in fe-
Size: Body length of female 0.89 mm; male 0.81 mm.
male and male, respectively; exopod with five setae in female
Morphology: Body with distinct separation between prosome and four setae in male.
and urosome. Prosome four-segmented; urosome five-segment-
Biology: Found only in “the North and South Clam fields, with
ed in female and six-segmented in male. Rostrum separated
a relatively high abundance”. The samples with the copepods
from cephalothorax, triangular, with bifid tip. Anal operculum
“contained basaltic pepperite with numerous small xenoliths as
large and prominent. Caudal rami 2.6 times as long as wide.
well as a great number of vesicomyiid clams and a rich associ-
Antennule short, eight-segmented in female and ten-segment-
ated fauna indicating hydrothermal activity” (WILLEN 2003).
ed in male. Antennary exopod three-segmented, with one, one,
four setae. Maxilliped three-segmented. Legs 1-4 biramous, Distribution: Tabar-Feni Volcanic Fore-Arc: Edison Seamount,
with three-segmented rami, except two-segmented endopod of Southern Clam field, North Clam field.
leg 1 in female and of legs 1 and 2 in male. Leg 5 with exopod
articulated with baseoendopod in female and fused in male;

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

Reference:
WILLEN E. (2003) J. Nat. Hist. 37(14): 1691-1711.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 348

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

Arthropoda, Crustacea, Copepoda, Misophrioida, Misophriidae

Misophriopsis longicaudata HUMES, 1999

Size: Body length of female 1.80-1.94 mm; male 1.28-1.45 mm. Remarks: Misophriid copepods inhabit shallow waters, deep-
sea hyperbenthos, deep-sea plankton, anchialine caves, and la-
Morphology: Prosome slender, five-segmented; tergite of
va tubes. Bathypelagic misophriids of the genus Benthomisophria
somite bearing leg 1 covered by shield of cephalosome. Uro-
SARS, 1909 “are opportunistic gorgers, feeding on variety of
some ornamented with lamellae and setules, five-segmented in
other animals, including copepods and cnidarians” (BOXSHALL
female, six-segmented in male; first somite with leg 5, genital
1982). Cave misophriids of the genus Speleophriopsis JAUME &
double-somite of female subdivided dorsally. Rostrum beak-like
BOXSHALL, 1996 (Speleophriidae) have been found in baited
in lateral view. Caudal rami longer than wide. Antennule
traps and can be scavengers (BOXSHALL & HALSEY 2004).
eighteen-segmented in female, thirteen-segmented in male.
Antenna with three-segmented endopod and six-segmented Distribution: Juan de Fuca Ridge: Coaxial Segment, Flow site
exopod. Legs 1-4 biramous, with three-segmented rami. Second (Vent HDV).
endopodal segment of leg 2 with two inner setae. Leg 5 with
one-segmented endopod and two- or three-segmented exopod
in female and male, respectively.

1A: Female habitus, dorsal; B: Habitus, lateral; C: Antenna; D: Leg 5; E: Male habitus, dorsal; F: Urosome, dorsal;
G: Leg 5; from HUMES (1999).

References:
BOXSHALL G.A. (1982) Philos. Trans. R. Soc. Lond. B 297: 125-181.
BOXSHALL G.A. & S.H. HALSEY (2004) An Introduction to Copepod Diversity. Ray Society, London: i-xv, 1-966.
HUMES A.G. (1999) J. Nat. Hist. 33: 961-978.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 349

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

Arthropoda, Crustacea, Copepoda, Cyclopoida, Oncaeidae

Oncaea praeclara HUMES, 1988

Size: Body length of female 1.01-1.30 mm; male 0.91-0.98 mm. Biology: Common in the “samples taken in the vicinity of the
vents by means of box corers and slurp guns” (HUMES 1988).
Morphology: Body elongate. Prosome five-segmented. Uro-
Oncaea sp. has been found in plankton over the Mid-Atlantic
some five-segmented in female and six-segmented in male, first
Ridge among dirivultids and subadult calanoids (IVANENKO
somite with leg 5. Genital double-somite of female and genital
1998). Most of more than 70 species of Oncaea occur in the
somite of male, anteriorly dilated and elongate. Caudal ramus
epipelagic zone, several species have been found in the deep ba-
long, length/width ratio 6.7:1 in female and 3.2:1 in male. An-
thypelagic zone. GO et al. (1998) recently reported that Oncaea
tennule six-segmented in both sexes. Antenna three-segment-
feeds on the integuments of various planktonic animals and in-
ed, without exopod; distal segment with two groups of setae.
flicts especially heavy injuries to chaetognaths; the copepods of
Mandible with three setae and two broad spines; palp absent.
this group “must gnaw or cut out pieces of host integument with
Maxillule lobe with seven setae. Maxilla two-segmented, sec-
their mandibles. To do this, they adhere tightly to the host by
ond segment with one spine and one seta. Maxilliped sexually
means of their antennae and maxillipeds” (HEPTNER & IVA-
dimorphic, four-segmented in female and three-segmented in
NENKO 2002).
male. Legs 1-4 biramous, with three-segmented rami. Female
leg 5 as a small segment bearing two setae and adjacent dorsal Distribution: Galapagos Rift; East Pacific Rise: 9°N, 13°N,
seta; male leg 5 represented by three setae. 21°N; Guaymas Basin.

1A: Female habitus, lateral; B: Urosome, dorsal; C: Antenna; D: Maxilliped; E: Male habitus, dorsal; F: Maxilliped; from HUMES (1987).

References:
GO Y.-B., OH B.-C. & M.J. TERAZAKI (1998) J. Mar. Sys. 15: 474-482.
HEPTNER M.V. & V.N. IVANENKO (2002) Arthropoda Selecta 11(2): 117-134.
HUMES A.G. (1987) Bull. Mar. Sci. 41: 645-788.
HUMES A.G. (1988) J. Plankton Res. 10: 475-485.
HUMES A.G. & M. SEGONZAC (1998) Zool. Zh. 77(11): 1249-1256.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 350

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

Arthropoda, Crustacea, Copepoda, Calanoida, Scolecitrichidae

Grievella shanki FERRARI & MARKHASEVA, 2000

Size: Body length of female 2.1-2.2 mm. Male unknown. Biology: One female has been collected in “less than a meter
above the surface on the side of a small diffusing vent chim-
Morphology: Prosome four-segmented; last segment of prosome
ney”. The copepods of the family Scolecitrichidae “usually are
with leg 5. Urosome three-segmented; first somite is genital
collected from pelagic and benthopelagic habitats of marine
double-somite with small integumental bumps. Rostrum bear-
waters below 200 m” (FERRARI & MARKHASEVA 2000).
ing two filaments, each on robust base. Antennule twenty four-
segmented; segment 22 with ear-like extension. Distal segment Distribution: East Pacific Rise: 21°S, Droopy Vent.
of maxilla with nine sensory setae. Exopods of legs 1-4 and en-
dopods of legs 3-4 three-segmented; endopods of leg 1 and leg
2 one- and two-segmented, respectively. Distal exopodal seg-
ment of leg 2 with two outer spines. Leg 5 two-segmented, dis-
tal segment with three setae.

1: Female; A: Habitus, lateral; B: Rostrum; C: Genital double-somite, lateral; D: Distal segment of maxilla; E: Leg 5;
from FERRARI & MARKHASEVA (2000).

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

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 351

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

Arthropoda, Crustacea, Copepoda, Calanoida, Spinocalanidae

Isaacsicalanus paucisetus FLEMINGER, 1983

Size: Body length of female 2.44-2.56 mm. Male unknown. Biology: The copepods have been sampled with the aid of a
slurp gun within 1 m of the sea floor. They “were swimming in
Morphology: Prosome five-segmented; tergites of two proximal
a tightly clustered swarm of several thousand individuals just
somites fused laterally. Urosome four-segmented; first somite is
about a small depression… formed in collapsed pillow lava”
genital double-somite. Rostrum present, not bifurcate. Caudal
(FLEMINGER 1983). Water temperatures in the depression range
rami slightly asymmetrical: left inner seta projecting ventrally,
from 5-15°C. The genus belongs to Spinocalanidae, “a com-
right inner seta directed dorsally. Antennule twenty three-seg-
mon group of small- to medium-sized calanoid copepods found
mented. Exopod of antenna longer than endopod. Exopods of
at meso- and bathypelagic depths of all oceans” (SCHULZ 1996).
legs 1-4 and endopods of legs 3-4 three-segmented; endopods of
legs 1-2, 1- and two-segmented, respectively. Leg 1: first seg- Distribution: East Pacific Rise: 21°N.
ment of exopod without inner seta. Leg 5 absent.

1: Female; A: Habitus, dorsal; B: Rostrum; C: Caudal rami; D: Antenna; from FLEMINGER (1983) (A, B, D) and SCHULZ (1989) (C).

References:
FLEMINGER A. (1983) Proc. Biol. Soc. Wash. 96 (4): 605-622.
SCHULZ K. (1989) Mitt. Hamb. Zool. Mus. Inst. 86: 185-208.
SCHULZ K. (1996) Polar Biol.16: 595-600.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 352

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Tegastidae

Smacigastes micheli IVANENKO & DEFAYE, 2004

Size: Body length of female 0.63 mm; male 0.57 mm. Biology: Numerous adult and subadult copepodid stages of S.
micheli have been collected during in situ colonization experi-
Morphology: Body strongly compressed laterally, shield of
ments at the base of the active chimney Eiffel Tower covered by
cephalothorax produced ventrolaterally, and male genital com-
a layer of Bathymodiolus azoricus (Bivalvia, Mytilidae).
plex produced ventrally. Legs 1-4 biramous, with three-seg-
mented rami. Elongate caudal rami and ten-segmented anten- Distribution: Mid-Atlantic Ridge: Lucky Strike.
nule of male are the distinctive features of Smacigastes.

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

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

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 353

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Tisbidae

Cholidya polypi FARRAN, 1914

with four setae and blunt terminal teeth. Leg 5 of male repre-
Size: Body length of female 0.74-0.83 mm; male 0.57-0.62 mm.
sented with lateral expansion bearing four setae.
Morphology: Body of female with flattened shield-like
Biology: Widespread ectoparasite living on arms and web of
cephalothorax and swollen metasome, urosome with slight in-
deep-sea octopuses in the Atlantic and Pacific Oceans. One of
dication of segmentation. Body of male with distinct somites
these octopuses, Graneledone sp., sampled at the sediment-
and six-segmented urosome. Rostrum produced anteriorly. An-
laden vent field was described by JUNIPER et al. (1992). The
tennule six-segmented in female and seven-segmented in male.
parasitic tisbids are found on arms, head, and gills of the deep-
Antenna: exopod one-segmented, endopod two-segmented.
sea octopuses associated to soft substrates. The redescription of
Mandibular palp three-segmented. Maxillule bilobed, with
Cholidya polypi and the list of species parasitizing octopuses is
three and five setae. Maxilla three-segmented, last two seg-
given in HUMES & VOIGHT (1997). Number of free-living tis-
ments forming claw. Maxilliped four-segmented, distal segment
bids (including Tisbe sp. nov.) have been found in the Mid-At-
claw-shaped. Leg 1 biramous with three segmented rami; distal
lantic Ridge (Lucky Strike) during in situ colonization experi-
segment of endopod with three claws; distal segment of exopod
ments (Ivanenko et al., in preparation).
armed with one spine and four setae ornamented with setules.
Leg 2 of female biramous, with two-segmented rami. Legs 3-4 of Distribution: Juan de Fuca Ridge: Middle Valley; none vent lo-
female represented by small segment with three and one setae, calities, Pacific and Atlantic Oceans (HUMES & VOIGHT 1997).
respectively. Legs 2-4 of male with three-segmented exopods
and two-segmented endopods. Leg 5 of female one-segmented,

1A: Female habitus, dorsal; B: Habitus ventral; C: Leg 1; D: Male habitus, dorsal. en – endopod; l5 – leg 5; from HUMES & VOIGHT (1997).

References:
HUMES A.G. & J.R. VOIGHT (1997) Ophelia 46(1): 65-81.
JUNIPER S.K., TUNNICLIFFE V. & E.C. SOUTHWARD (1992) Can. J. Zool. 70: 1792-1809.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 354

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

Arthropoda, Crustacea, Copepoda, Harpacticoida, Tisbidae

Genesis vulcanoctopusi LÓPEZ-GONZÁLEZ, BRESCIANI & HUYS, 2000

Size: Body length of female 2.5-2.8 mm. Male unknown. Biology: Females and subadults have been found embedded and
encapsulated in the integument of the head and of the mantle
Morphology: Body indistinctly segmented; prosome laterally
of the deep-sea octopus Vulcanoctopus hydrothermalis GONZÁLEZ
expanded with protruding margins. Rostrum produced beyond
& GUERRA, 1998. It was suggested that the life cycle of Genesis
anterior margin of cephalothorax. Antennule four-segmented.
vulcanoctopusi would consist of both endoparasitic and ectopar-
Antenna with one-segmented exopod and two-segmented en-
asitic phases.
dopod. Mandibular palp two-segmented. Maxillule bilobed,
with eight setae. Maxilla two-segmented, second segment claw- Distribution: East Pacific Rise: 13°N, near Genesis.
like. Maxilliped three-segmented, distal segment claw-shaped.
Leg 1-4 biramous, exopods of legs 1-4 and endopod of leg 2
three-segmented; endopods of legs 1, 3, and 4 two-segmented.
Leg 5 represented by lobe fused to body and bearing six setae.

1: Female; A: Habitus, dorsal;

B: Habitus ventral; from
LÓPEZ-GONZÁLEZ et al. (2000).

Reference:
LÓPEZ-GONZÁLEZ P.J., BRESCIANI J., HUYS R., GONZÁLEZ A.F., GUERRA A. & S. PASCUAL (2000) Cah. Biol. Mar. 41: 241-253.

V.N. IVANENKO & D. DEFAYE Denisia 18 (2006): 355

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Cirripedia
The cirripedes or barnacles are crustaceans found perma-
nently attached as adults. They occur in brackish as well as ma-
rine conditions, ranging from rigorous habitats such as estuaries
and the highest reaches of the tides to the relatively benign
depths of the oceans. There are three superorders, the Acrotho-
racica, Rhizocephala and Thoracica, and it is in the last that
the greatest diversity is found. While the barnacle suborders as-
sociated with hydrothermal vents and seeps are, like their deep-
sea counterparts, predominantly pedunculates belonging to the
Scalpellomorpha and the asymmetrical sessile Verrucomorpha,
their representatives, such as Ashinkailepas and Neoverruca, are
remarkably generalized. Surprisingly, vents also include repre-
sentatives of the two symmetrical sessile suborders, the
Brachylepadomorpha and Balanomorpha. The former were
thought to have gone extinct in the Miocene, until Neo-
brachylepas was discovered at Lau. The latter, represented by
Eochionelasmus, first appear in the Paleocene and include the
1: Barnacle Vulcanolepas “Lau A” sp. from Lau Back-Arc Basin; cruise TUIM07; by courtesy of C.R. Fisher.
W. NEWMAN, T. YAMAGUCHI, A.J. SOUTHWARD & M. SEGONZAC
dominant shallow-water barnacles today. All four of these gen-
era are judged to be the most primitive surviving members of
their suborders; e.g., relics of bygone ages.
In addition to being unusual in composition and primitive
character, as a group the vent and seep barnacles differ from all
other thoracicans in having their mandibles uniquely modified
for feeding on very fine particles such as bacteria and fine de-
bris. Some of the species have filamentous chemoautotrophic
bacteria growing profusely on modified cirral setae, which they
nurture by holding in prevailing vent currents and upon which
they presumably feed. Furthermore, while vent barnacle larvae
are non-feeding, they are relatively large and yolky and studies
to date indicate they can remain in the plankton for three
months or so before having to settle. This is five to six times
longer than any known shallow-water barnacle which, as one
would expect, indicates they are very good dispersers.
Denisia 18 (2006): 356–357
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
The relative abundance of vent barnacles appears to differ
greatly from place to place as well as between species. While
Neolepas zevinae is common along the Central East Pacific Rise,
dense stands are rarely encountered. However, an undescribed
neolepadine from the Central Indian Ocean Ridge, Vul-
canolepas osheai from the Kermadecs, Leucolepas longa from off
New Guinea, Neoverruca brachylepadoformis from the Mariana
Back-Arc Basin, and Eochionelasmus ohtai from Lau, may occur
in extremely dense populations around vents or seeps. But cu-
riously, the remains of vent barnacles have yet to be reported
from vent or seep deposits of any age. On the other hand, no
specimens of Neobrachylepas relica or Imbricaverruca yamaguchii
have been taken since first discovered, despite several subse-
quent expeditions to Lau, and to date Neoverruca spp. are
known only by a chance photograph from Lau and in a sample
from Manus, and both were among crowded Eochionelasmus. It
thus seems likely many new and exciting forms remain to be
discovered.
Vent-seep barnacles are now known to range pretty much
throughout the Indo-Pacific vent-seep systems, except in the
NE Pacific and the Galapagos where they are conspicuously ab-
sence. They are also absent from the Atlantic, but so are nu-
merous other ancient groups of marine invertebrates. The cen-
ter of distribution, where representatives of most genera are
presently found, is in the SW Pacific, specifically the Lau Basin,
Tonga. It is noteworthy that the centers of diversity at high tax-
onomic levels in numerous other relic groups of marine inverte-
brates are also found in the SW Pacific. Taxonomic diversity at-
tenuates to north and south, as well as to the east, and appar-
ently also to the relatively unexplored vents of the far west. Bar-
riers, such as disjunctions in active venting, the sill between the
Marianas vents and the vents of Japan, or the Eastern Pacific
and the Galapagos, have been proposed to explain these atten-
uations. However, considering the ephemerality and patchiness
of vent and seep fields, extinction rates associated with them
must be relatively high. It follows that the center of distribution
is where availability of the vent-seep habitat has been the least
interrupted in space and time, for otherwise such a high taxo-
nomic endemicity could not have accumulated. Like the high
taxonomic endemicity of the SW Pacific in general, that of the
vent barnacles appears to be the result of reliction due to habi-
tat restriction rather than to barriers to dispersal per se.
357
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Cirripedia, Balanomorpha, Chionelasmatoidae

Eochionelasmus ohtai YAMAGUCHI, 1990

brachylepadomorphs, the most parsimonious explanation was

Size: This species, measuring up to 25 mm in height, is sub-
that balanomorphs evolved from them rather than independ-
stantially larger than its bathyal and more highly derived coun-
ently from scalpellomorphs. However, forthcoming genetic ev-
terpart, Chionelasmus darwini, which is about 15 mm in height.
idence indicates the balanomorphs evolved from very different
Morphology: Chionelasmatines are balanomorphs having a scalpellomorph stocks than the neoverrucids whereby the no-
wall comprising two pairs of dedicated latera, in addition to the table similarities between neoverrucids and verrucids are con-
rostrum, carina and basal imbricating plates seen in vergent.
brachylepadomorphs. Eochionelasmus is distinguished from
Biology: Eochionelasmus apparently occupies the same micro-
Chionelasmus by the basal whorls of imbricating plates of sever-
habitat as Neoverruca, but when co-occurring it tends to be
al to numerous distinct whorls rather than being condensed in-
physically closer to the mussels then are sympatric ne-
to what appears to be a single whorl, and by its trophi being
olepadines. Equipped for feeding on the same material as all
much like those of other vent species. Two subspecies of Eo-
vent barnacles studied to date. Eggs are relatively large, but
chionelasmus were proposed by YAMAGUCH & NEWMAN (1997),
rarely recovered brooding whereby the nature of the larval
E. ohtai ohtai and E. ohtai manusensis from Lau, Fiji and Manus
stages is unknown.
Back-Arc Basins, respectively. They are distinguished from
each other by differences in the frequency of appearance of the Distribution: Lau, North Fiji and Manus Back-Arc Basins.
rl1 and cl1 plates of the imbricating whorls (see E. paquensis). Morphological differences between the Lau-Fiji and Manus
populations are considered sub-specific (see above) while dif-
Remarks: Eochionelasmus represents the most primitive living
ferences between these three and a population from near East-
balanomorph. While the order in which the whorls of imbri-
er Island were interpreted as being specific (E. paquensis).
cating plates are added during ontogeny is the reverse of that in

1: E. o. othai, collected at Lau Back-Arc Basin; by 2: Group of E. o. manusensis from N-Fiji Back-Arc Basin; cruise Starmer
courtesy of T. Yamaguchi. II; P. Briand © Ifremer.

References:
BUCKERIDGE J.S. & W.A. NEWMAN (1992) J. Paleontol. 66: 341-345.
GALKIN S.V. (1992) Zl. zh. Rss. Akad. Nauk. 71(11): 139-134.
KLEPAL W, NEWMAN W. A. & W. TUFAR (2006) in SCHRAM F.R. & J.C. VON VAUPEL KLEIN (Eds.) Crustaceans and Biodiversity. Brill, Leiden: 195-199.
PERÉZ-LOSADA M., JARA C.G., BOND-BUCKUP G., PORTER M.L. & K.A. CRANDALL (2002) J. Crust. Biol. 22: 661-669.
TUFAR W. & H. JULLMANN (1991) Spiegel der Forschung 8(1): 39-44.
YAMAGUCHI T. & W.A. NEWMAN (1990) Pacific Sc. 44(2): 135-155.

W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 358

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

Arthropoda, Crustacea, Cirripedia, Balanomorpha, Chionelasmatoidae

Eochionelasmus paquensis YAMAGUCHI & NEWMAN 1997

Size: 5 mm in diameter. Biology: Living with mussels (Bathymodiolus thermophilus).

Morphology: The shell is low conic, its orifice large and rhom- Distribution: East Pacific Rise: 17°S, site Rehu. Undescribed
boidal. Differs externally from E. ohtai in the alae of primary species of this genus are also known from Izu Ogasawara Arc,
plates being very indistinct and by the rostrum rising obliquely the Pacific-Antarctic Ridge: 38°S and the Central Indian
rather than nearly vertically to the orifice of the wall. Often Ocean.
coated with ferromanganese. The scutum is very distinct in
lacking an articular ridge and adductor muscle pit, a wider ar-
ticular furrow, and a tergal magin that is indented rather than
straight. The tergum has a produced articular ridge. The num-
ber of basal imbricating plates ist greatly reduced compared to
E. othai.

1A: Right side of the holotype; B: Above side of the same specimen; scale bar 1 mm; 2: Specimen, view from above; scale
from YAMAGUCHI & NEWMAN (1997). Capital letters indicate the principal wall plates: bar 1 mm; from East Pacific Rise:
R – rostrum; RL – rostrolateral; CL – carinolateral; C – carina; small letters indicate the 17°S, cruise Naudur © Ifremer; from
imbricating plates YAMAGUCHI & NEWMAN (1997).

Reference:
YAMAHUCHI T. & W.A. NEWMAN (1997) J. Crust. Biol. 17(3): 488-496.

W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 359

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

Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae

Ashinkailepas seepiophilia YAMAGUCHI, NEWMAN & HASHIMOTO, 2004

particles, but no significant differences noted compared too

Size: Up to 36 mm in height.
those of ordinary vent barnacles (with the exception of those
Morphology: Capitular plates ornamented by conspicuous lon- with cirri having exceptionally long setae generally festooned
gitudinal ribs or ridges, peduncular to capitular ratio of 1:1 or with filamentous bacteria). Peduncles not elongate, as in some
less and, if unaltered by bending over to one side, with whorls neolepadines, but rather relatively short and bilaterally asym-
of six large scales which are considerably wider than high. metrical peduncular plates when individuals are bent over too
one side or the other. Function of bending over unknown, but
Remarks: Ashinkailepas is considered as the most primitive of
it may be an advantage too extend the cirri closer too the sub-
the Neolepadinae. The subfamily Neolepadinae is considered
stratum, or perhaps too present a lower profile too marauding
as the most primitive of the living Scalpellomorpha as well as
predators. Hermaphroditic; eggs large (~300 x 500 µm) and rel-
closest relative of the ancestor of the Brachylepadomorpha of
atively few in number, as in N. zevinae NEWMAN, 1979, free
which the only living representative is Neobrachylepas relica
nauplii likely present and lecithotrophic (cf. JONES 1993; TUN-
known from Lau.
NICLIFFE & SOUTHWARD 2004; WATANABE et al. 2004).
Biology: This species was collected from a cold deep. Associ-
Distribution: Off Hatsushima Island, Sagami Bay (hydrother-
ates included a seep clam (Calyptogena sp.), and shrimps
mally driven cold seep). Undescribed species of this genus are
(Alvinocaris sp. and Lebbeus sp.). An ordinary deep-sea crab
known from Japan: 32°N and 27°N, Lihir and the Kermadec
(Paralomis multispina) inhabited crevices between the boulders.
Ridge.
Equipped with long cirri for setose feeding in gentle currents
and mouth parts uniquely modified for handling extremely fine

2: Group of Ashinkailepas sp. from Japan;

by courtesy of T. Yamaguchi.

1: Right side; from YAMAGUCHI et al. (2004).

References:
TUNNICLIFFE V. & A.J. SOUTHWARD (2004) J. Mar. Biol. Assoc. U.K. 84: 121-132.
WATANABE H., KADO R., TSUCHIDA S., MIYAKE H., KYO M. & S. KOJIMA (2004) J. Mar. Biol. Assoc. U.K. 84: 743-745.
YAMAGUCHI T., NEWMAN W.A. & J. HASHIMOTO (2004) Mar. Biol. Assoc. U.K. 84: 797-812.

W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 360

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

Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae

Leucolepas longa SOUTHWARD & JONES, 2003

long, often gently curved peduncle appears to be to keep the ca-

Size: Peduncular length commonly five but up to 12 times
pitulum above those of its peers rather than adjusting to changes
height of the capitulum, total length reaching 40 cm or more.
in prevailing currents, as seems to be the case in Vulcanolepas.
This is, proportionately as well as literally, the longest peduncle
Growth appears to be rapid with reproductive activity and re-
of any known staked barnacle.
cruitment continuous. Lecithotrophic nauplii, released upon re-
Morphology: Capitulum relatively narrow, tergal apex more trieval were cultivated in vitro for 45 days, metamorphosed into
acute than in other neolepadines; e.g., <65° rather than some- stage IV swimming nauplii containing lipid reserves deemed suf-
what <75° as in Neolepas and >80° as in Vulcanolepas; capitular ficient to insure wide dispersal. In a similar in vitro rearing of lar-
plates of mature individuals separated by a band of arthrodial vae of Neoverruca sp., it tooks 96 days to reach the cyprid stage.
membrane; basal angle of tergum elevated well above rather
Remarks: Leucolepas longa differs considerably in general from
than close to capitulo-peduncular junction; ratio of peduncular
Neolepas and Vulcanolepas. While what is apparently a new
length to capitular height generally >5:1 to as much as 12:1.
genus from the Izu Ogasawara Arc has the same general ap-
Leucolepas longa can be distinguished from other neolepadines
pearance, the basal angle of the tergum reaches the capitulo-
in the basal angle of the tergum being well separated from the
junction, the plates are contiguous, the median latus has a
capitulo-peduncular junction. This is true even in early juve-
height to width ratio is two rather than 1.5:1 or less, and the p-
nile stages in which the relatively contiguous plates and the
c ratio is in the order of eigth rather than 12:1.
more convex occludent margin of the capitulum, as well as
short, blunt peduncular scales, suggest affinities with Vul- Distribution: Tabar-Feni Volcanic Fore-Arc: Edison Seamount.
canolepas. In adult L. longa the capitular plates are well separat- While presently monotypic, undescribed populations range as
ed rather than contiguous with each other. far East as the Izu Ogasawara Arc (see remarks). An unde-
scribed species is known from North Fiji Back-Arc Basin, an-
Biology: It occurs in densities of over 1000 m-2, mostly associat-
other undescribed species from Sunda Trench, and perhaps a re-
ed with vesicomyid clams, but occasionally on rocks and vesti-
lated genus from Izu Ogasawara Arc.
mentiferans over sulfide-rich sediments. The function of the

3: In situ view of clumps of barnacles, some

1: Capitulum after removal of the
prosoma and cirri which empha-
sizes the uncalfified interspaces
between the shell plates; from
SOUTHWARD & JONES (2003).
2: Group of specimens; from SOUTHWARD &
JONES (2003).
with cirral net open, at the edge of a bed of
live vesicomyids; also visible, associated fauna
of Phymorhynchus gastropods and squat lob-
ster; scale bar 15 cm; from TUNNICLIFFE & SOUTH-
WARD (2004); by courtesy of P. Hertzig.

References:
SOUTHWARD A.J. & D.S. JONES (2003) Senckenberg. maritima 32 (1/2): 77-93.
TUNNICLIFFE V. & A.J. SOUTHWARD (2004) J. Mar. Biol. Ass. U.K. 84: 121-132.

W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 361

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

Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae

Neolepas rapanuii JONES, 1993

and all subsequently described vents and seep species, but the
Size: Up to approximately 8.0 cm in total length.
functional basis for the difference is unknown.
Morphology: Distinguished from N. zevinae by the apex of ros-
Remarks: Peduncles elongated as in most other neolepadine
trum generally being equal to rather than higher than the me-
genera, evidently to keep the capitulum and cirral net in cur-
dian latus, a peduncle often as much as fives times the height of
rents rather than to host chemoautotrophic endosymbionts.
the capitulum and clothed with less robust scales, and a
This is a different strategy than practiced by Ashinkailepas, N.
mandible without a strong spine near the superior margin of the
zevinae, and of course the sessile vent barnacles, Neoverruca,
second tooth and denticles of the second and third teeth not
Imbricaverruca, Neobrachylepas and Eochionelasmus.
rolled over toward the inside.
Distribution: East Pacific Rise: 23°S. An undescribed species of
Biology: While little known, its biology is likely much the same
this genus is known from Pacific-Antarctic Ridge: 32°S, anoth-
as that of N. zevinae, except for the development of a longer
er species from Izu Ogasawara Arc and another morphological-
and often curved peduncle, presumably to keep the cirral net in
ly similar from is known from Toto Caldera Marianos Arc.
prevailing currents, and the comb of denticles along the supe-
Three other populations from the Indian Ocean Rigde (25°S,
rior margin of the second and third mandibular teeth and the
17°S, 41°S) while genetically close to Neolepas, appear inter-
upper part of the inferior angle, that stand nearly erect rather
mediate between it and Leucolepas.
than being rolled over toward the inner surface of the
mandible. This is curious as they are rolled over in N. zevinae

1: In vivo specimen; scale bar 1 cm; from JONES (1993).

References:
JONES D.S. (1993) Bull. Mar. Sci. 52(3): 937-948
SOUTHWARD A.J. & D.S. JONES (2003) Senckenberg. maritima 32(1/2): 77-93

D. JONES, W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 362
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae

Neolepas zevinae NEWMAN, 1979

Size: This species is known to be up to 5.8 cm in total length, Biology: Neolepadines are hermaphroditic, like all vent barna-
whereas alone the peduncle of N. rapanuii can exceed that cles studied to date. They are gregarious but, like the cyprid lar-
length, and some neolepadines in the western Pacific have pe- vae of Ashinkailepas, those of Neolepas zevinae apparently prefer
duncles more than a decimeter in length. to settle on the substratum rather than on established individ-
uals, such as on the long peduncles of Neolepadines from else-
Morphology: Capitulum with relatively smooth, unornament-
where. Individuals tend to align along ridges and contours fac-
ed capitular plates and an adult peduncle with whorls of 16 or
ing the prevailing current. Neolepadines and Eochionelasmus
more scales which are about as wide as high. Distinguished from
occur sympatric with mussels, the latter tend to be closer to the
N. rapanuii by the apex of rostrum generally being higher than
vent. Equipped with long cirri for setose feeding in gentle cur-
that of the median latus, a peduncle clothed with proportion-
rents and equipped with mouth parts uniquely modified for
ately more robust scales and of no more than three rather than
handling extremely fine particles, as in all vent barnacles stud-
as much as five times the height of the capitulum, and a
ied to date. Peduncles become elongate in some forms, evi-
mandible having a strong spine near the superior margin of the
dently to keep the capitulum and cirral net in currents rather
second tooth and the denticles of the second and third teeth
than to host chemoautotrophic endosymbionts. Eggs large in
rolled over toward the inside.
size, egg nauplius present in Neolepas rapanuii, but there are
Remarks: Neolepadines are considered the most primitive of likely free lecithotrophic stages, as in other vent barnacles.
the living scalpellomorphs known, and are evidently early
Distribution: East Pacific Rise: 9-21°N.
Mesozoic in age. They apparently arose in the Triassic (BUCK-
ERIDGE & GRANT-MACKIE 1985) and are near the base of the
clade that gave rise to the first sessile barnacles, the
brachylepadomorphs.

1: Specimen in vivo collected from 2: Specimen in vivo collected from

East Pacific Rise: 21°N; East Pacific Rise: 21°N;
by A. Southward. by A. Southward.

References:
BUCKERIDGE J.S. & J.A. GRANT-MACKIE (1985) Géologie de la France 1: 77-80.
JONES D.S. (1993) Bull. Mar. Sci. 52: 937-948.
NEWMAN W.A. (1979) Trans. San Diego Soc. Nat. Hist. 19(11): 153-167.
NEWMAN W.A. (1985) Bull. Biol. Sc. Wash. 6: 231-242.

W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 363

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

Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae

Vulcanolepas osheai (BUCKERIDGE, 2000)

1998), an hypothesis compatible with the molecular phyloge-

Size: Capitulum up to approximately 20 mm, overall length to
netic and isotopic evidence for ectosymbiosis in V. osheai
at least 120 mm, so pedunclar to capitular 5:1 or 6.8:1.
(Suzuki et al., in prep.).
Morphology: Capitulum broad, tergal apex blunter than in oth-
Remarks: This species is very similar in external appearance to
er Neolepadines (>80 versus <75°); capitular plates approxi-
“Lau A“ of SOUTHWARD & NEWMAN (1998). In most individu-
mate, not separated by a band of arthrodial membrane; basal
als of Lau A, these setae were festooned with filamentous bac-
angle of tergum close to rather than elevated well above capit-
teria. Since the cirral rami as well as the exterior surfaces of the
ulo-peduncular junction; maximal observed ratio of pedunclar
barnacle in general were not covered with such bacteria,
length to capitular height 7:1 but usually less.
SOUTHWARD & NEWMAN (1998) concluded those on the setae
Biology: Found living on volcanic debris in the vicinity of a were being farmed for consumption.
new dome forming within the caldera. While no other mac-
Distribution: Kermadec Arc: Brothers Seamount.
robenthic animals were collected, the presence of shrimp was
observed. Cirri apparently equipped with exceptionally long,
relatively soft setae (BUCKERIDGE 2000, Fig. 4C, E), which sug-
gests bacterial ectosymbiosis (cf. SOUTHWARD & NEWMAN

1: Habitus; scale bar 1 cm

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

References:
BUCKERIDGE J.S. (2000) New Zealand J. Mar. Freshwater Res. 34: 409-418.
SOUTHWARD A.J. (2005) Senckenberg. maritima 35(2): 147-156.
SOUTHWARD A.J. & W.A. NEWMAN (1998) Cah. Biol. Mar. 39: 259-262.

W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 364

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Cirripedia, Scalpellomorpha, Eolepadidae
Vulcanolepas parensis SOUTHWARD, 2005
Size: Capitulum up to 15 mm in height and peduncular length
of 103 mm; peduncular to capitular ratio 6.86:1, while perhaps
a smaller species than V. osheai, peduncular to capitular ratio es-
sentially the same.
Morphology: Tergum shorter (basal angle well above capitulo-
peduncular margin) and median latus is taller than in V. osheai,
median latus forming an approximately equilateral triangle
whereas in V. osheai and Leucolepas longa it is curved along the
tergal margin.
1: Capitulum of a specimen with the capitular
plates; from SOUTHWARD (2005).
Reference:
SOUTHWARD A.J. (2005) Senckenberg. maritima 35(2): 147-156.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD
Remarks: Morphologically this species appears to be a Vul-
canolepas, and if so, it is relatively isolated since the genus is
otherwise only known from Lau, East to the Kermadecs. How-
ever, there could be a vent fauna along the Pacific-Antarctic
and Indian-Antarctic Ridges to be discovered that would help
explain the situation.
Biology: Posterior cirri with ctenopod rather than lasiopod cir-
ri as in Vulcanolepas sp.
Distribution: Pacific-Antarctic Ridge: 37°S to 38°S. A bacte-
ria-farming form from Lau Back-Arc Basin is representative of
this genus.
2: Cluster of specimens on a small fragment
of the substratum, peduncles of medium
length; from SOUTHWARD (2005).
Denisia 18 (2006): 365
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Cirripedia, Brachylepadomorpha, Neobrachylepadidae

Neobrachylepas relica NEWMAN & YAMAGUCHI, 1995

Size: Only a few possibly juvenile or protandric specimens have
been collected, suggesting they may have been waifs; that is,
the microhabitat of the adult populations likely has yet to be
sampled. The largest specimen was mature as a male, but there
were no eggs and it was only 6 mm high. It is nonetheless pos-
sible this is a small species compared to other vent barnacles.
Morphology: A living representative of the Brachylepadomor-
pha (U. Jurassic-Miocene), symmetrical sessile barnacles with
basal whorls of imbricating plates surrounding a wall consisting
of but two principal plates, the rostrum and carina. Neo-
brachylepas differs from †Brachylepas in the rl-l-cl tiers of imbri-
cating plates covering the gap between the principal wall plates
standing three rather than four plates high (compare to Imbri-
caverruca in this regard). Distinguished from neolepadines in
being sessile and operculate, from neoverrucids in being sym-
metrical, and from balanomorphs in the operculum including a
pair of median latera and the oldest whorl of imbricating plates
being basal rather than situated between the younger imbricat-
ing whorls and the wall.
Biology: Unfortunately a reproductive population of this
species has yet to be discovered, but one would expect its mem-
bers to occupy much the same situations occupied by Neoverru-
ca brachylepadoformis and Eochionelasmus ohtai because, while
perhaps a smaller species, it has essentially the same feeding
mechanism and is hermaphroditic. The first and often the sec-
ond pair of cirri in vent barnacles are antenniform and likely
serve in part in orienting the cirral net to currents. However, in
Neobrachylepas, the second pair is not only shorter than the first
but it apparently functions as maxillipeds. Another unique fea-
ture in Neobrachylepas is a median-dorsal appendage on the pro-
soma that may serve to hold the egg mass in place while brood-
ing, but ovigerous individuals have not been observed.
Remarks: Neobrachylepas relica is the only known surviving
member of the Brachylepadomorpha, fossil representatives of
which are known from the Jurassic to the Miocene.
Distribution: Lau Back-Arc Basin.

1: Viewed from above (A) and from the right (B). Me-
dian latera (L), scuta (S) and terga (T) comprise the op-
erculum; rostrum (R) and carina (C) are in contact at
their lateral margins; the other letters indicate the im-
bricating plates; from NEWMAN & YAMAGUCHI (1995).
2: Specimen viewed from the left side, almost com-
pletely buried beneath sulfide and oxide deposits
on a small block of basalt; cruise Biolau © Ifremer;
from NEWMAN & YAMAGUCHI (1995).

References:
NEWMAN W.A. & T. YAMAGUCHI (1995) Bull. Mus. natl. Hist. nat., Paris. 4e sér 17A (3-4): 211-243.
NEWMAN W.A. (1993) in TRUSEDALE J. (Ed.) The History of Carcinology. Crustacean Issues 8: 349-434.

W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 366

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

Arthropoda, Crustacea, Cirripedia, Verrucomorpha, Neoverrucidae

Imbricaverruca yamaguchii NEWMAN, 2000

Size: Apparently a small species, largest of several specimens Biology: No data.

having a rostro-carinal diameter of 7 mm and standing about
Distribution: Known only from the type locality; Lau Back-Arc
5.5 mm high.
Basin, site Hine Hina.
Morphology: Differs from Neoverruca in having 1) a more ver-
rucid-like operculum albeit including a well developed median
latus, 2) a fixed scutum and tergum much wider than high
rather than at least as high as wide, and 3) in retaining a com-
plete set of well-developed imbricating plates on the movable
side of the wall essentially for life whereas the imbricating sys-
tem in Neoverruca is incomplete (at least plates r4 -l4-c4 fail to
develop) and the older (marginal) imbricating plates tend to
fall off.

1A: Oblique view of movable side; B: Viewed from scutal end; C – carina, FS,
MS – scutum, FT, MT – tergum, R – rostrum, cl – carinolaterale, l – laterale, rl
rostrolaterale; from NEWMAN (2000).

Reference:
NEWMAN W.A. (2000) Zoosystema 22(1): 71-84.

W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD Denisia 18 (2006): 367

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Cirripedia, Verrucomorpha, Neoverrucidae
Neoverruca brachylepadoformis NEWMAN, 1989
Size: Moderately large sessile barnacle measuring up to 25 mm
in height.
Morphology: Neoverrucids are bilaterally asymmetrical sessile
barnacles built the same general plan as the brachylepado-
morphs. They are distinguished from verrucids in having basal
whorls of imbricating plates surrounding the wall and an oper-
culum including a median latus, which in N. brachylepadoformis
becomes vestigial with age and may even be lost, but in Imbri-
caverruca it remains relatively large and significant part of the
operculum. Imbricaverruca is further distinguished from Neover-
ruca in having the rl and cl tiers of imbricating plates as in
†Brachylepas; e.g., standing four rather then three or less plates
high, and in having the basal-most ones well attached rather
than easily shed.
Remarks: Neoverrucids appear not only to be the most primi-
tive living verrucomorphs but to represent the “missing link“
between the Brachylepadomorpha and the remainder of the
Verrucomorpha.
Biology: Neoverruca brachylepadoformis is hermaphroditic. It is
gregarious, and the cyprid larvae settle on established individu-
als as well as on the substratum. Settled juveniles first pass
through several pedunculate stages before undergoing meta-
morphosis into the first sessile stage. Cyprids metamorphosis in-
to juveniles with their cirral nets facing the prevailing current,
and those that settled on the right side of an established indi-
1: Large specimen
(2.1 cm high), right
side, with two juve-
niles lacking ferro-
manganese deposit;
Mariana Back-Arc
Basin; from NEWMAN
& HESSLER (1989)
References:
NEWMAN W.A. (1989) Bull. Mar. Sci. 45(2): 467-477.
NEWMAN W.A. & R.R. HESSLER (1989) Trans. San Diego Soc. Nat. Hist. 21(16): 259-273.
WATANABE H., KADO R., TSUCHIDA S., MIYAKE H., KYO M. & S.J. KOJIMA (2004) Mar. Biol. Assoc. U.K. 84(4): 743- 745.
W.A. NEWMAN, T. YAMAGUCHI & A.J. SOUTHWARD
vidual develop their operculum on the right side, while those
settling on the left develop it on the left side. Thus right and
left-sidedness is ecotypically determined, as is likely the case in
other verrucomorphans where right and left-sidedness occurs in
the same species. It appears that some species of the genus are
occasionally sympatric with Eochionelasmus, at Lau and Manus
Basins for example.
Equipped to feed on the same material as all vent barnacles
studied to date. Eggs are relatively large and held in place by
ovigerous frenae. Lecithotrophic nauplii in vitro pass through
six stages before metamorphosis after to the cyprid stage, and
this took 96 days in the deep-water form from the Okinawa
Trough reared in vitro. Unusual among sessile barnacles in pass-
ing through several pedunculate stages resembling neolepadids
during ontogeny, as likely do the brachylepadomorphs. While
the earliest pedunculate stages are essentially symmetrical,
asymmetry becomes apparent before an abrupt metamorphosis
into the first sessile juvenile, during which the peduncle be-
comes buried beneath an expanded membranous basis, and the
basal whorl of capitular plates becomes the oldest or basal-most
whorl of imbricating plates of the sessile form. Additional
whorls are added between it and the wall plates until at least
the median latera of the movable side stand four plates high.
Distribution: Mariana Back-Arc Basin. Undescribed species
are found in Marianas, Japan, Manus and Lau.
2: Closely
packed
aggregation;
Mariana Back-
Arc Basin;
by courtesy of
R.R. Hessler.
Denisia 18 (2006): 368
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Malacostraca, Leptostraca, Nebaliidae

Dahlella caldariensis HESSLER, 1984

visual elements. Scale of the first antenna with the anterior

Size: Up to 10 mm.
margin densely clothed with setae ranging from stout and den-
Morphology: Carapace ovoid, posterior edge emarginate, ex- tate to long and slender (see upper right figure).
posing at least part of pleonite one dorsally. Rostrum 0.4 times
Biology: In washings of mussels and vestimentiferan worms.
of carapace length, 3 time longer than wide. First thoracomere
Observed swimming above clumps of animals at vents.
seen from carapace even dorsally, postero-lateral angles of
pleonites 1-3 rounded, that of pleonite 4 pointed. Anal plates Distribution: Galapagos Spreading Center and East Pacific
like equilateral triangles. Eyestalk boomerang shaped (see up- Rise: 21°N to 18°S.
per centre figure) and denticulate articulated to head without

1: Habitus (critical point dried specimen)  Ifremer.

4: Lateral side of pleonites 3: Distal penduncular article of

(SEM)  Ifremer. the first antenna (SEM)  Ifremer.

2: Eyestalk (carapace removed) SEM  Ifremer. 5: Pleopods (SEM)  Ifremer. 6: Caudal ramus (SEM)  Ifremer.

References:
HESSLER R.R. (1984) J. Crustac. Biol. 4(4): 655-664.
LEDOYER M. (1995) Mar. Life 4(1): 25-29.

T. HANEY Denisia 18 (2006): 369

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

Arthropoda, Crustacea, Cumacea, Bodotriidae

Atlantocuma bidentatum LEDOYER, 1988

long as dactylus. Pereopod 2 basis shorter than the remaining ar-

Size: 3.9 mm.
Morphology: Subfamily: Mandibles not broad at base. Exopods
only on maxilliped 3 and pereopod 1 in female. Males usually
with five pairs of pleopods (sometimes two or three) but they
are absent in Atlantocuma that has been considered as an aber-
rant genus (JONES , 1984). Without free telson. Uropod endo-
pod one- or two-articulated.
Female: Carapace smooth, about twice as long as high and
slightly shorter than 1/3 of total length; pseudorostrum not very
long, pointed; antennal notch shallow, anterolateral angle not
prominent, marked by two teeth; eyelobe pointed at front, with-
out lenses. Five thoracic segments visible. Pleon slightly longer
than carapace and thoracic segments combined. Antenna 1 pe-
duncle three-articulated; main flagellum two-articulated with
two aesthetascs terminally, accessory flagellum rudimentary. Ex-
opods on maxilliped 3 and pereopod 1. Maxilliped 3 basis slen-
der, 1.5 as long as the rest of appendage. Pereopod 1 basis slight-
ly longer than rest of appendage with a small simple seta on dis-
tal corner; carpus longer than propodus, which is 1.5 times as
ticles together, ischium very short, carpus longer than dactylus.
Pereopod 3 basis longer than rest of appendage, carpus about 1/3
of basis. Pereopod 4-5 basis shorter than the rest of appendage,
carpus longer than half length of basis. Uropod peduncle longer
than pleonite 5 and twice length of endopod; exopod article 2
with a simple seta on distal third of inner margin and a long sim-
ple seta terminally; endopod one-articulated, with 10 setae on
inner margin and a longer one terminally.
Remark: Although the specimen described above agrees with
the LEDOYER’S (1988) description in most of the features, it dif-
fers by a greater number of setae on the endopod of uropod.
Biology: Collected in a sediment trap (12 m above the bottom)
located at East Pacific Rise: 13°N, 2 m north of the vent site
Parigo.
Distribution: East Pacific Rise: 13°N, site Parigo. Previously
known only from South-East of Glorieuses Islands, Mozam-
bique Channel, at 3716 m depth (LEDOYER 1988).

1: Preadult female; A: Habitus; B: Anterolatereal angle of carapace; C: Maxilliped 3; D: Pereopod 1; E: Pereo-

pod 2; F: Pereopod 3; G: Pereopod 4; H: Pereopod 5; I: Uropod; by J. Corbera.

References:
JONES N.S. (1984) Bull. Br. Mus. Nat. Hist. (Zool.) 46(3): 207-289.
LEDOYER M. (1988) Mésogée 48: 131-172.

J. CORBERA Denisia 18 (2006): 370

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

Arthropoda, Crustacea, Cumacea, Bodotriidae

Bathycuma brevirostre (NORMAN, 1879)

short. Pereopod 3 and 4 basis longer than rest of appendage,

Size: 15.5 mm.
Morphology: Subfamily (Vaunthompsoniinae): Mandibles not
broad at base. Exopods on at least first three pairs of pereopods.
Males usually with five pairs of pleopods (sometimes three).
Without free telson. Uropod endopod one- or two-articulated.
Species, adult male: Carapace with a mid-dorsal paired row of
teeth on anterior half; anterolateral angle acute and lateral
margin serrated. Antenna 1 peduncle three-articulated, article
1 slightly shorter than article 2 and 3 combined lengths; main
flagellum two-articulated with 2 aesthetascs terminally, acces-
sory flagellum rudimentary. Well developed exopods on maxil-
liped 3 and pereopods 1-4. Maxilliped 3 basis longer than rest
of appendage, distal outer corner produced reaching merus.
Pereopod 1 basis shorter than rest of appendage, with cuspidate
setae on ventral face and pappose setae on distal half of outer
and inner margins; carpus and dactylus of the same length.
Pereopod 2 basis as long as rest of appendage, ischium very
with pappose setae on margin. Pereopod 5 basis shorter than
the three following articles combined lengths. Uropod pedun-
cle longer than rami, with more than 20 setae on inner margin,
being largest the distal one; exopod article 2 with simple setae
on outer margin, plumose setae on the inner margin and two
long simple setae terminally; endopod two-articulated, article 1
twice long as article 2 with more than 20 setae on inner mar-
gin; article 2 with eight setae on inner margin and 2 terminal-
ly.
Biology: Collected in a sediment trap (2.5 m above the bot-
tom) located between two active vents at 1630 m depth.
Distribution: Previously known from south of Ireland (CAL-
MAN 1905), Bay of Biscay (JONES 1985) and Mediterranean Sea
(REYSS 1973) up to 5000 m depth. Mid-Atlantic Ridge: Lucky
Strike.

1: Adult male; A: Habitus; B: Antenna 1; C: Maxilliped 3; D: First pereopod; E: Second pereopod; F: Third pereo-
pod; G: Fourth pereopod; H: Fifth pereopod; I: Uropod; by J. Corbera.

References:
CALMAN W.T. (1905) Fish. Ireland, Sci. Invest.1904(1): 1-7.
JONES N.S. (1985) in LAUBIER L. & C. MONNIOT (Eds.) Peuplements Profonds du Golfe de Gascogne: 429-433.
NORMAN A.M. (1879) Ann. Mag. Nat. Hist. 3(5): 54-73.
REYSS D. (1972) Deep-Sea Res. 20: 1119-1123.

J. CORBERA Denisia 18 (2006): 371

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

Arthropoda, Crustacaea, Tanaidacea

In some deep-sea environments tanaids are the most di- All species known from hydrothermal vents belong to pre-
verse and abundant fauna but because of their small size (most viously known genera, even Gordotanais was found before al-
are 2-5mm long) they are frequently overlooked. Their distri- though not yet described indicating that the fauna at Lucky
bution in hydrothermal environments is virtually unknown. Up Strike is not dramatically different from the surrounding deep-
to now only two species (Leptognathia ventralis and Typhlotanais sea habitat. A similar observation was made in the study of
sp.) are known from Western Pacific Back-Arc Basins, seven tanaids from the Juan de Fuca vent system (LARSEN, in press).
species from the Mid-Atlantic Ridge, Lucky Strike (LARSEN et
Although the present handbook aims to present diagnostic
al., in press) and one (Typhlotanais sp.) from Rainbow (M.
information and illustration it is important to stress that the
Segonzac, pers. obs.).
identification of tanaid species is often difficult and usually re-
At Lucky Strike, the highest diversity and abundance of quires the dissection of the specimens.
tanaids occurred in peripheral mussel clumps. Besides the two
species, Agathotanais ingolfi and Pseudotanais vulsella, known
from other deep-sea environments, Mesotanais styxis was found
both inside and outside the vent field.

1: Armaturatanais atlanticus; by K. Larsen, M. Blazewicz-Praszkowycz & M. R. Cuhna.

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

M.R. CUNHA Denisia 18 (2006): 372

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

Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, fam. indet.

Armaturatanais atlanticus LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press

Pleon marginally longer than in female. Antennular article 3

Size: 2.0 mm.
Morphology: (Genus, female) Body slightly dorso-ventrally
flattened. Cuticle heavily calcified. Pleotelson acorn-shaped in
lateral view. Antennula with four articles. Antenna with six ar-
ticles and fusion line on article 4. Mandibles small; molar fair-
ly broad with flat distal crushing area surrounded by small den-
ticles. Maxilliped endites without denticles, processes or flat se-
tae. Pereopods I-III merus and carpus without spiniform setae.
Pereopods and pleopods attached on the inner side of the lat-
eral shield. Pereopods with coxa; dactylus and unguis not fused
to a claw; pereopods I-III unguis longer than dactylus; pere-
opods IV-VI dactylus longer than unguis. Pleopods present in
female, with simple or plumose setae. Uropods biramous, endo-
pod with two articles; exopod with one article. (Genus, male)
shorter than in female and without fusion line. Functional
mouthparts retained. All pleonites bearing pleopods with
plumose setae, pleopods larger than in female. (Species, fe-
male) Antennula article 3 more than half as long as article 2.
Right mandibular incisors smooth; left mandibular/lacinia mo-
bilis with few dorsal denticles. Chelipedal dactylus naked. Pere-
opods longer than pleon. Pereopods IV-VI dactylus naked; un-
guis of same with simple apex.
Biology: Collected from samples of volcanic rocks (hyaloclas-
tic) and hydrothermal slab in the proximity of active vents.
Distribution: Mid-Atlantic Ridge: Lucky Strike.

1: Female; A: Habitus, lateral view; B: Antennula; C: Antenna; D: Cheliped; E: Maxilliped; scale bars
1 mm (A), 0.1 mm (others); from LARSEN et al. (in press).

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

K. LARSEN & M.R. CUNHA Denisia 18 (2006): 373

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

Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Colleteidae

Leptognathiella fragilis LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press

setae only. Uropods biramous; rami with one or two articles.

Size: 1.3 mm.
(Species, female) Body fairly elongated (length/width ca. 9.5).
Morphology: (Family and Genus) Body cylindrical, relatively Cephalothorax as long as combined length of first two pere-
small (rarely over 2 mm in length). Pleon and pleotelson short onites. Pereopods I-III carpus and propodus without small ven-
(never longer than combined length of three last pereonites). tral spines. Uropods twice as long as pleotelson but shorter than
Antennule with four or five articles; article 2 frequently with combined pleon; basal article shorter than pleotelson and biar-
dorsal projection overlapping basal part of article 3. Antenna iculated exopod half as long as first endopod article.
with five or six articles. Mouthparts well developed and func-
Biology: Collected from samples of hydrothermal slabs in the
tional in both sexes. Molar process thin with few terminal
vicinity of active venting.
spines. Maxillule with seven or eight terminal spiniform setae.
Maxilliped endite not fused, distal edge often with medial, fre- Distribution: Mid-Atlantic Ridge: Lucky Strike.
quently triangular, process. Chelipeds attached via sclerite.
Pereopods I-III often stout, with or without coxa. Pereopods IV-
VI without coxa and not stouter than pereopods I-III; dactylus
and unguis not fused. Pleopods absent or present with simple

1: Female; A: Habitus lateral view; B: Antennula; C: Antenna; D: Maxilliped; E: Cheliped;

scale bars 1 mm (A), 0.1 mm (others); from LARSEN et al. (in press).

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

K. LARSEN & M.R. CUNHA Denisia 18 (2006): 374

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

Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Leptocheliidae

Mesotanais styxis LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press

out coxa; basis thicker than pereopods I-III dactylus/unguis in-

Size: 1.9 mm.
completely fused to an elongated claw. Pleopods present and
Morphology: (Family and Genus) Body cylindrical. Eye lobes well developed, with plumose setae. Uropods biramous; exopod
present but without visual pigment. Antennula with three arti- biarticulated; endopod with three or more articles. (Species, fe-
cles. Antenna with six articles; article 2 and 3 with spiniform male) Antennula without long (almost as long as antennule)
dorsal setae. Mandibles well developed with broad molar. Labi- setae. Maxilliped basis with only one distal setae. Uropodal en-
um with two pairs of lobes. Maxillule with nine spiniform ter- dopod with four articles.
minal setae. Maxilliped basis and endites not fused; endites
Biology: Collected from samples of hydrothermal slabs in the
with two or three short, flat, unequal-sized setae and none or
proximity of active chimneys but also on volcanic rocks outside
one simple seta; basis with one or two long simple setae near
the vent field (Lucky Strike segment).
palp insertion. Mouthparts reduced in males. Chelipeds at-
tached via sclerite. Pereopod I almost twice as long as following Distribution: Mid-Atlantic Ridge: Lucky Strike.
pereopods; dactylus/unguis longer than propodus. On other
pereopods dactylus/unguis shorter than propodus. Pereopods I-
III with coxa; dactylus/unguis not fused. Pereopods IV-VI with-

1: Female; A: Habitus, dorsal view; B: Habitus, lateral view; C: Antennula; D: Antenna; E: Cheliped;
F: Pleopod; scale bars 1 mm (A, B), 0.1 mm (others); from LARSEN et al. (in press).

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

K. LARSEN & M.R. CUNHA Denisia 18 (2006): 375

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

Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Nototanaidae

Obesutanais sigridi LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press

the length of endopod). Uropod endopod is incompletely fused

Size: 1.2 mm.
to one article; endo- and exopod with thick specialized termi-
Morphology: (Family and Genus, female) Body habitus short nal setae; exopod unarticulated, almost as long as endopod.
and stout (almost pseudotanais-like) with a wide head. Body ta- (Species, female) Pereopods long (longer than pleon). Pere-
pering off in posterior direction. Antennula with long setae on opods IV-VI dactylus and unguis partly fused but not into a
article 1. Mandibular molar with few distal spines. Maxilliped claw. Uropods longer than pleotelson; endopod with pseudoar-
endites with one distal tubercule. Mouthparts reduced in males. ticulation; exopod with one article, almost as long as endopod.
Cheliped merus and carpus with long seta (longer than fixed
Biology: Collected from samples of volcanic rocks (hyaloclas-
finger). Pereopods II and III merus with one seta as long as or
tic) and sulphide chimneys active and inactive. Often found in-
longer than carpus; carpus with a long seta (almost as long as
side tubes built with small particles of sediment.
propodus). Pereopod IV-VI without process with microspines
(clinging apparatus). Pleopods setation, except for endopod Distribution: Mid-Atlantic Ridge: Lucky Strike.
proximal seta, restricted to the distal end; large gap between
endopod proximal seta and other setae, long (more than half

2: Tubes on a piece of sulphide

(Lucky Strike), cruise Seahma 1
© FCIJL; by P. Briand © Ifremer.

1: Female; A: Habitus, dorsal view; B: Habitus, lateral view; 3: Specimen removed

C: Antennula; D: Antenna; E: Cheliped; scale bars 0.1 mm; from tube; by M. Cunha.
from LARSEN et al. (in press).

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

K. LARSEN & M.R. CUNHA Denisia 18 (2006): 376

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

Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Nototanaidae

Typhlotanais incognitus LARSEN, BLAZEWICZ-PASZKOWYCZ & CUNHA, in press

fused. Pleopods present, with simple or plumose setae. Uropods

Size: 1.5 mm.
Morphology: (Family and Genus, female) Body almost com-
pletely cylindrical. Cephalothorax wider than pereon. Anten-
nula with three articles. Antenna with five or six articles. Mo-
lar process broad with several terminal denticles. Maxillula
with seven or eight terminal spiniform setae. Maxilliped basis
partially fused, endites not fused; distal edge with lateral
process. Chelipeds ventrally attached via sclerite. Pereopods I-
III with coxae; merus and carpus with simple setae only. Pere-
opods IV-VI with or without coxae; basis stouter or not stouter
than pereopods I-III; merus and carpus with at least one spini-
form seta, usually with clinging apparatus; dactylus and unguis
biramous; endopod with two articles; exopod with one or two
articles. (Species, female) Pereopods long (longer than pleon).
Pereopods IV-VI not stouter than Pereopods I-III; dactylus and
unguis fused but not into a claw. Uropods longer than pleotel-
son; endopod with two articles; exopod with one article, almost
as long as endopod.
Biology: Collected from samples of hydrothermal slab in the
proximity of active vents.
Distribution: Mid-Atlantic Ridge: Lucky Strike.

1: Female; A: Habitus, lateral view; B: Habitus, dorsal view; C: Antennula; D: Antenna; E: Cheliped; scale bars 0.1 mm; from
LARSEN et al. (in press).

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

K. LARSEN & M.R. CUNHA Denisia 18 (2006): 377

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

Arthropoda, Crustacea, Tanaidacea, Tanaidomorpha, Pseudotanaidae

Pseudotanais vulsella BIRD & HOLDICH, 1989

broad. Pereopods II-VI carpal blade-like spine long. Pereopods

Size: 2.3 mm.
IV-VI similar dactylus and unguis fused. Pleopods well devel-
Morphology: (Family and Genus) Body habitus short and ro- oped.
bust. Eyes absent. Pereonites 1-2 reduced. Antennula with
Biology: On Lucky Strike area, this species was frequently col-
three articles, distal article bearing setae with complex tips.
lected from samples of volcanic rocks (hyaloclastic and lava)
Maxilliped endites fused. Cheliped attachment via sidepiece.
and sulphide rubble but also in areas of diffuse venting and near
Pereopods I-III coxae absent. Pereopod I different from II-III.
active chimneys active.
Pereopods II-VI carpus antero-inferior spine flattened, “blade-
like”. Uropod endopod with two articles; exopod with two arti- Distribution: North Atlantic: North Feni Ridge, Rockall
cles. One pair of oostegites only on pereopods IV. (Species) Trough, Hebridian slope, Porcupine Seabight and Celtic Slope,
pereonites 1-3 combined shorter than pereonite 4. Cheliped of 1028-1640 m; Mid-Atlantic Ridge: Lucky Strike.
forcipate type. Pereopod I slender, basis about 12 times longer
than broad. Pereopods II-III similar, basis 6 times longer than

1: A: Female habitus, dorsal view; B: Male habitus, dorsal view; C: Right cheliped, female; D: Antenulla, male; scale bar 1 mm;
from BIRD & HOLDICH (1989).

References:
BIRD G.J. & D.M. HOLDICH (1989) Zool. J. Linnean Soc. 97: 233-298.
LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.

M.R. CUNHA & M. SEGONZAC Denisia 18 (2006): 378

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

Arthropoda, Crustacaea, Isopoda

Recent expeditions to the Lucky Strike vent field at the
Mid-Atlantic Ridge provided an uncommon collection of per-
acarid crustaceans (CUNHA et al. 2001, CUNHA & WILSON
2003, in press) that accounted for about 50% of the species
richness in the samples and included about 20 isopod species.
Among these four asellotes are new species: the two Heterome-
sus presented herein, one undescribed Katianiridae and one un-
described Munnopsidae. These species were collected in areas
of diffuse low venting, in peripheral mussel (Bathymodiolus
azoricus) aggregations and in a zone of filter feeding organisms.
Nevertheless, we cannot infer that these isopods are vent en-
demic because they do not show special adaptations to the en-
vironment and our knowledge of their distribution is minimal.
The Lucky Strike isopod fauna also includes a number of
known deep-sea asellotes (mostly Haplomunnidae, Haplonisci-
dae, and Munnopsidae) and a few species of other isopods (An-
thurida, Cirolanidae, and Gnathiidae, unpublished data). Some
of these were already cited in the previous version of the Hand-
book (DESBRUYÈRES & SEGONZAC 1997) together with the epi-
carid Thermaloniscus cotylophorus from East Pacific Rise: 13ºN.
Isopods are among the most abundant and diverse taxa in
the deep sea (HESSLER & SANDERS 1967, HESSLER et al. 1979;
RAUPACH et al. 2004), but there are only very few records of
these crustaceans in hydrothermal communities. The reasons
for this are unclear because isopods, especially asellotes, are
known for their adaptative potential (HESSLER & THISTLE
1975) and at least in relatively shallow (<2000 m) hydrother-
mal vent fields, such as in Lucky Strike, the less toxic environ-
ment likely facilitates the immigration of background species
(VAN DOVER 1995).

References:
CUNHA M.R. & G.D.F. WILSON (2003) Zootaxa 323:1-16.
CUNHA M.R. & G.D.F. WILSON (in press) Zootaxa.
CUNHA M.R., HILARIO A.M. & I.G. TEIXEIRA (2001) IOC Workshop Report 175: 73-74.
DESBRUYÈRES D. & M. SEGONZAC (1997) Handbook of Deep-Sea Hydrothermal Vent Fauna. Ifremer Ed.: 1-279.
HESSLER R.R. & T.D. THISTLE (1975) Mar. Biol. 32: 155-165.
HESSLER R.R. & H.L. SANDERS (1967) Deep-Sea Research 14: 65-78.
HESSLER R.R., WILSON G.& D. THISTLE (1979) Sarsia 64: 67-76.
LARSEN K., BLAZEWICZ-PASZKOWYCZ M. & M.R. CUNHA (in press) Zootaxa.
RAUPACH M.J., HELD C. & J.-W. WÄGELE (2004) Deep-Sea Research II 51: 1787-1795.
VAN DOVER C.L. (1995) in PARSON L.M., WALKER C.L. & D.R. DIXON (Eds.) Hydrothermal vents and processes. Geol. Soc. Spec. Publ. 87: 257-294.

M.R. CUNHA Denisia 18 (2006): 379

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Isopoda, Asellota, Ischnomesidae
Heteromesus calcar CUNHA & WILSON, in press
Size: 4 mm.
Morphology: (Family) Body elongate, subcylindrical and nar-
row. Pereonites 4-5 elongate, pereonite 5 longest, pereonite 4
widest anteriorly, 5 widest posteriorly. Head fused and embed-
ded in pereonite 1. Pereonites 1 (posterior margin) to 4 free and
articulating. Anus separated from branchial chamber. Eyes ab-
sent. Antennula terminating with simple seta, article 1 squat
and globular, article 2 elongate, at least twice as long as article
1. Antenna length more than half body length, without squa-
ma. Maxilla inferior margin with two medial pectinate setae.
Pereopod I robust strongly subchelate; pereopods II-VII ambu-
latory. Uropod uniramous terminal. (Genus) Pereonite 5 freely
articulated with pereonite 6, articulations not expressed be-
tween pereonite 6, pereonite 7, pleonite 1 and pleotelson. Pere-
onite 4 produced posteriorly but always broader than long. An-
tennula article 1 globular, article 2 inserting dorsally; article 2
strongly curved anteriorly at proximal insertion; articles distal
to article 2 reduced to 1-3 articles, distal articles altogether
1: Female (left) head, lateral view and habitus, dorsal and lat-
eral views; scale bar 1 mm; by M. Cunha.
Reference:
CUNHA M.R. & G.D.F. WILSON (in press) Zootaxa.
M.R. CUNHA & G.D.F. WILSON
tiny, length less than 0.3 times of article 2 length. Pereopod I
carpus distally expanded, widest point distal to midpoint of car-
pus, with one elongate robust seta and a proximal shorter robust
seta. Pleopod II female operculum with narrow proximal neck,
almost circular posteriorly, with plumose setae. Pleopod III ex-
opod with plumose setae and fringe of fine setae. Pleopod V
absent. Uropod with single article, conical, tapering distally;
extending beyond posterior margin of pleotelson. (Species)
Pereonites 1-3 with anterolateral spines. Pereonite 2 in female
with no paired dorsal spines or tubercles. Pereonite 5 in female
length 2.0 times the width. Antennula with 3 articles altogeth-
er. Pereopods IV-V bases with pedestal spines; ischia with elon-
gate pedestal spines.
Biology: Collected in the vicinity of active venting sites on sul-
phide deposits, sulphide rubble and volcanic rocks.
Distribution: Mid-Atlantic Ridge: Lucky Strike.
2: Male (right) habitus, dorsal view, and pleotelson, dorsal and
ventral views; scale bar 1 mm; by M. Cunha.
Denisia 18 (2006): 380
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Isopoda, Asellota, Ischnomesidae

Heteromesus ctenobasius CUNHA & WILSON, in press

to article 2 reduced to 1-3 articles, distal articles altogether

Size: 4 mm.
Morphology: (Family) Body elongate, sub-cylindrical and nar-
row. Pereonites 4-5 elongate, pereonite 5 longest, pereonite 4
widest anteriorly, 5 widest posteriorly. Head fused and embed-
ded in pereonite 1. Pereonites 1 (posterior margin) to 4 free and
articulating. Anus separated from branchial chamber. Eyes ab-
sent. Antennula terminating with simple seta, article 1 squat
and globular, article 2 elongate, at least twice as long as article
1. Antenna length more than half body length, without squa-
ma. Maxilla inferior margin with two medial pectinate setae.
Pereopod I robust strongly subchelate; pereopods II-VII ambu-
latory. Uropod uniramous terminal. (Genus) Pereonite 5 freely
articulated with pereonite 6, articulations not expressed be-
tween pereonite 6, pereonite 7, pleonite 1 and pleotelson. Pere-
onite 4 produced posteriorly but always broader than long. An-
tennula article 1 globular, article 2 inserting dorsally; article 2
strongly curved anteriorly at proximal insertion; articles distal
tiny, length less than 0.3 times of article 2 length. Pereopod I
carpus distally expanded, widest point distal to midpoint of car-
pus, with one elongate robust seta and a proximal shorter robust
seta. Pleopod II female operculum with narrow proximal neck,
almost circular posteriorly, with plumose setae. Pleopod III ex-
opod with plumose setae and fringe of fine setae. Pleopod V
absent. Uropod with single article, conical, tapering distally;
extending beyond posterior margin of pleotelson. (Species)
Pereonite 1-3 median pedestal spines. Pereonites 4-5 in female
with lateral pedestal spines. Pleotelson terminal margin with
pedestal spines. Antennula with five articles altogether. Pereo-
pod II-VII bases and ischia with pedestal spines.
Biology: Collected in the vicinity of active venting sites on
volcanic rocks.
Distribution: Mid-Atlantic Ridge: Lucky Strike.

1: Female habitus, dorsal view; the color picture is a false color composite of three different SEM images,
by G.D.F. Wilson & S. Lindsay © Australian Museum.

Reference:
CUNHA M.R. & G.D.F. WILSON (in press) Zootaxa.

M.R. CUNHA & G.D.F. WILSON Denisia 18 (2006): 381

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

Arthropoda, Crustacea, Amphipoda

Currently, 24 species of Amphipoda are described from
hydrothermal vent samples. In addition, some other species we-
re collected in the surroundings of vents or at stations not
strictly linked with hydrothermal vent communities. A certain
number of new species are currently being described.
The following preliminary conclusions can be drawn from
the species so far identified (BELLAN-SANTINI 1998):
(1) The subfamily of Lysianassoidea (including Lysianassi-
dae sensu lato and Uristidae) is exclusively found in Pacific
samples with 10 species.
(2) Twelve other families are represented by one to three
species.
(3) The number of endemic species is very high. However,
the large repartition of species most likely is due to insufficient
identification and appears doubtful.
(4) Four species are abundant and build swarms: Ventiella
sulfuris and Halice hesmonectes in Pacific Ocean and Bouvierella
curtirama and Luckia striki in Atlantic Ocean (VAN DOVER et al.
1992; MARTIN et al. 1993; SHEADER et al. 2003). The other spe-
cies are known to occur only in small numbers per sample.
(5) Different techniques have been used in the past to sam-
ple amphipods. Trawls are not recommended because large sur-
faces are sampled without distinction of communities (in the
case of the sampling of the Guaymas area). Also, only large an-
imals are collected and the delicate amphipods are often bro-
ken. Slurp guns or grabs manipulated by the submarine are the
best methods for collecting fragile animals but for rare species
they are not very efficient. Washing of mytilids, tubeworms and
other species is a good method if rinsing of samples is conduc-
ted carefully. Sediment traps deployed for a certain period of ti-
me are also a good, but probably selective method.
Amphipods are delicate and fragile animals in particular
the representatives of the deep-sea families. They frequently
loose appendices and damaged individuals are difficult to de-
termine. Generally there are only one or two individuals of a
species per sample. Consequently, if they are damaged identifi-
cations and descriptions are impossible. Alcohol is the best fi-
xative, but colors will vanish after preservation.

1: Ventiella sulfuris from East Pacific Rise: 9°N; by M. Bright.

References:
BELLAN-SANTINI D. (1998) Cah. Biol. Mar. 39: 143-152.
BELLAN-SANTINI D. (2005) J. Nat. Hist. 39(39): 3435-3452.
MARTIN J.W., FRANCE S.C. & C.L.VAN DOVER (1993) Can. J. Zool. 71: 1724-1732.
SEGONZAC M. (1992). C.R. Acad. Sci. Paris. 314: 593-600.
SHEADER M. VAN DOVER C.L. & M.H. THURSTON (2004) Mar. Biol. 144: 503-514.
VAN DOVER C.L., KAARTVEDT S., BOLLENS S.M., WIEBE P.H., MARTIN J.W. & S.C. FRANCE (1992) Nature 358: 25-26.
VINOGRADOV G.M. (1993) Zool. Zh. 72(2): 40-53.

D. BELLAN-SANTINI Denisia 18 (2006): 382

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Ampeliscidae

Ampelisca romigi BARNARD, 1954

Epimeral plate posterior edge convex, lower posterior corner

Synonyms: A. isocornea: BARNARD, 1954, A. romigi ciego
quadrate. Urosomite 1, dorsal surface with a weakly seddle-
BARNARD, 1966.
shaped process. Telson rather broad. (Subspecies A. romigi
Size: Up to 16 mm. ciego) Eyes absent. Pereiopod 7 with propodus more strongly
narrowed in the proximal part while the propodus is wider.
Morphology: (Family) Mouthparts basic. Eyes composed of
corneal lenses or absent. Body without process except uro- Biology: Unknown. Dredged mainly from soft-bottom areas.
somite 1. Urosomites 2 and 3 coalesced. Antenna 1 without ac- Trawl in periphery of hydrothermal vents.
cessory flagellum. Gnathopods 1 and 2 slender subchelate or
Distribution: East Pacific Rise. Along the coast of southern
nearly simples. Pereopods 3-4 slender with merus elongate.
California and in the Channel Islands; Gulf of California at
(Genus) Pereopods 5-6 with basisvery broad. Pereopod 7 basis
Angel de la Guardia Island, Isla Partida; Angeles Bay, San Mar-
dilated with posterior lobe greatly expanded distally, bearing
cos Island, Tortuga Island and Tiburon Island; Isabel Island,
marginal plumose setae. Telson longer than broad and deeply
Mexico; Salinas Bay, Costa Rica; Secas Islands, Panama; Santa
cleft. (Species) Eyes present. Pereiopod 7 lower edge of basis
Elena Bay, Ecuador, 3-500 m. The subspecies A. romigi ciego was
reaching joint between merus and carpus, merus with a posteri-
found also at Guaymas vent site.
or setose lobe, carpus with distal anterior edge notched.

1: Female 9.5 mm;

after BARNARD (1966).

References:
BARNARD J.L. (1954) Allan Hancock Pac. Exped. 18(1): 1-137.
BARNARD J.L. (1966) Allan Hancock Pac. Expec. 27(5):1-166.
VINOGRADOV G.M. (1993) Zool. Zh. 72(2): 40-53 [in Russian].

D. BELLAN-SANTINI Denisia 18 (2006): 383

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Amphilochidae

Gitanopsis alvina BELLAN-SANTINI & THURSTON, 1996

Size: 2.5 mm in adult female. Biology: Collected in hydrothermal field.

Morphology: (Family) Coxa 4 immensely broadened, coxa 1 Distribution: Mid-Atlantic Ridge: Lucky Strike.
very small and hidden by coxa 2. Peduncle of uropode 3 elon-
gate. Telson entire elongate. (Genus) Mandibular molar large,
triturative. Dactylus of gnathopods simple. (Species) Head with
rostrum. Accessory flagellum one-articulated. Antenna 2
slightly longer than antenna 1. Gnathopods small and feeble.
Epimeral plate 3 posterodistal margin rounded. Uropod 3, pe-
duncle 1.5x longer than inner ramus, outer ramus shorter than
inner. Telson elongate, triangular, apex tridentate.

1A: Head incomplete of female; B: Head of juvenile, C: Antenna 1, D: Gnathopod 1, E: Epimeral plates, F: Uropod 3, G: Telson;
from BELLAN-SANTINI & THURSTON (1996).

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

D. BELLAN-SANTINI Denisia 18 (2006): 384

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

Arthropoda, Crustacea, Amphipoda, Corophiidea, Aoridae

Autonoe longicornis CHEVREUX, 1909

spine, about one third length of peduncle; inner ramus longer

Size: 8 mm.
than outer and subequal in length with peduncle. Uropod 2 pe-
Morphology: (Family) Head anteroventral margin weakly re- duncle with short inter-ramal spine about one sixth length of
cessed, moderately excavate. Pereopods 3-4 basis glandular, peduncle; inner ramus longer than outer and subequal with pe-
pereopod 7 very elongate. Gnathopod 1 enlarged in males and duncle. Uropod 3 peduncle longer than broad, rami subequal
females. Uropod 3 with slender and robust setae. Telson and only a little longer than peduncle, inner ramus with small
dorsoventrally thickened. Species, male: Antennae 1 and 2 second article. Telson with each dorsal crest bearing a fine seta.
subequal in length, less than half body length. Antenna 1 pe-
Biology: Collected from samples of volcanic rocks (pillow lava
duncle article 3, one quarter length of article 1; flagellum short-
and hyaloclastic rocks). The proximity of active venting was
er than peduncle; accessory flagellum composed of one rudi-
inferred from the occurrence of Bathymodiolus azoricus (living
mentary article. Mandible palp article 3 longer than article 2,
specimens and/or shell debris) in the samples. Previously
terminally falcate. Maxilla 1 inner plate with a single, long,
known only by the type specimens (only females) collected by
pectinate seta. Labium mandibular processes acute. Gnathopod
CHEVREUX (1909) from similar depths (1360 m) also in the
1 coxa produced anterodistally, subacute; basis robust; carpus
Azores region.
enlarged, longer than propodus, the posterodistal corner pro-
duced into a spine. Gnathopod 2 carpus and propodus elongate, Distribution: Mid-Atlantic Ridge: Lucky Strike.
subequal in length. Pereopods 5-7 basis only weakly expanded.
Epimera 1-3 rounded. Uropod 1 peduncle with inter-ramal

1: Male; by A.A. Myers. G1 – gnathopod 1; G2 – gnathopod 2; Lab – labium; Md – mandible; Mx1 – maxilla; Mxp –
maxilliped; T – telson; U1, U2, U3 – uropods 1-3.

References:
CHEVREUX E. (1909) Bull. Inst. Océanogr. Monaco 150: 1-7.
MYERS A.A. & M.R. CUNHA (2004) J. Mar. Biol. Ass. U.K. 84: 1019-1025.

A.A. MYERS & M.R. CUNHA Denisia 18 (2006): 385

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Calliopiidae

Oradarea longimana (BOECK, 1871)

Size: Up to 11 mm. Biology: Frequently associated to other invertebrates. SHOE-

MAKER (1930) describes a collection from whelk egg cases, per-
Morphology: (Family) Lateral cephalic lobes small. Coxae of
haps as a scavenger or predator of eggs. One of samples from the
medium length or short. Antennae long; accessory flagellum
Juan de Fuca vent sites was taken from the decapod Macrorego-
vestigial or absent. Mouthparts basic. Gnathopods powerful or
nia macrochira SAKAI. Oradarea is a genus attracted to baited
feeble, usually subchelate, occasionnally simple. Telson of
traps, and it seems probable that these amphipods are scav-
medium size, entire, acuminate, emarginate or notched.
engers that cue on exudates of egg capsules and possibly pre-
(Genus) Rostrum large. Accessory flagellum one-articulated.
moult decapods.
Antenna 2 longer than 1. Gnathopods diverse. Pereopods 3-7
ordinary. Epimeron 3 smooth. (Species) Accessory flagellum Distribution: Recorded from the North Atlantic (54 m depth)
very short. Both gnathopods very slender; gnathopod 2 much and the N Pacific coast of North America, and probably broad-
longer than 1. Pereopods 3-4 alike in structure. Urosomite 1-2 ly extended along the northern Hemisphere. Present also at hy-
dorsally produced backward. Epimeron 3 with lower posterior drothermal vent sites of the Juan de Fuca Ridge (Endeavour
angle very obtuse and postero-lateral margin evenly convex. Segment, Lieutenant Obo vent, Dual Smoker; Explorer Seg-
Uropod 3 peduncle one quarter of inner ramus and half of out- ment, Upper Magic Mountain, Magic Mountain.
er. Telson a little longer than wide, apex rounded with a very
shallow cental notch.

1A: Posterior part of the body; B: Head; C: Gnathopod 1; D: Gnathopod 2; E: Telson; from SHOEMAKER (1930).

References:
BOECK A. (1871) Forhandl. Vidensk.-Selskabet i Christiania 1870: 83-280.
SHAW P. (1989) Can. J. Zool. 67(8): 1882-1890.
SHOEMAKER C.R. (1930) Contr. Canad. Biol. Fisheries, new series 5(10): 221-359.

D. BELLAN-SANTINI & D. JAUME Denisia 18 (2006): 386

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

Arthropoda, Crustacea, Amphipoda, Caprellidea, Caprellidae

Caprella bathytatos MARTIN & PETTIT, 1998

Size: Largest male 14.9 mm; smallest juvenile 2.2 mm. Biology: It is the first caprellid found in association with the
crab Macroregonia macrochira, or in the vicinity of marine hy-
Morphology: (Family) Body slender, cylindrical. Eyes lateral.
drothermal vent. 30 individuals were sampled on a single crab.
Gnathopods often different. Number of pereopods variable.
(Genus) Mandible lacking palp. Antenna 2, flagellum two-ar- Distribution: Juan de Fuca Ridge.
ticulate. Pereopods 3-4 lacking. Pereopods 5-7 normal.
(Species) Head rounded, lacking spines or projectings. Eyes re-
duced. Pereon lacking any spines or projections. Gnathopod 2,
propodus bears one blunt, rectangular tooth, separated by a U-
shaped notch from a slightly more acute tooth, an acute proxi-
mal lobe, dactylus 2/3 length of propodus; gnathopod attached
to body at midlength of pereionite 2. Each pereopod with grasp-
ing spines on propodus forming with the dactylus a mechanism
for attachment to host setae.

1A: Habitus of male, attached to maxilliped 3 of the majid crab; B: Gnathopod 2; C: Pereopod 7; D: Abdomen, male;
E: Abdominal appendage; from MARTIN & PETTIT (1998).

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

D. BELLAN-SANTINI Denisia 18 (2006): 387

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Eusiridae

Bouvierella curtirama BELLAN-SANTINI & THURSTON, 1996

Size: 5 mm in adult female. Biology: B. curtirama is abundant. This species lives in Bathy-
modiolus azoricus beds. Females maturing at a minimum body
Morphology: (Family) Accessory flagellum 0-2-articulated.
length of 3.5 mm and males at 2.4 mm.
Gnathopods variable. (Genus) Body not toothed. Eyes absent.
Coxal plates 1-4 ordinary. Epimeron 3 smooth. (Species) Head, Distribution: Mid-Atlantic Ridge: Lucky Strike, sites Sintra
rostrum moderate, lateral cephalic lobe rounded. Antenna 1 and Tour Eiffel sites.
longer than antenna 2, flagellum with a number of articles vari-
able with the size of the body, 8-46, 22 in the holotype. An-
tenna 2 flagellum variable about 7-37, 12 in holotype.
Gnathopods weakly subchelate, dissimilar, second longer than
first. Pereopods 3-7 ordinary, dactylus simple not prehensile.
Epimeral plate 3 smooth. Telson rounded, without armament.

1: A: Habitus; B: Gnathopod 1; C: Epimeral plates; D: Uropod 1; E: Uropod 2; F: Uropods 3 and telson; from BELLAN-SANTINI &
THURSTON (1996).

References:
BELLAN-SANTINI D. & M.H. THURSTON (1996) J. Mar. Biol. Ass. U.K. 30: 685-702.
SHEADER M., VAN DOVER C.L. & M.H. THURSTON (2004) Mar. Biol. 144: 503-514.

D. BELLAN-SANTINI Denisia 18 (2006): 388

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Eusiridae

Luckia striki BELLAN-SANTINI & THURSTON, 1996

Size: 7 mm in adult female. Biology: Collected in vent community among shrimps, gas-
tropods, crabs, and limpets.
Morphology: (Family) Accessory flagellum 0-2-articulated.
Gnathopods variable. (Genus) Body normally compressed. Distribution: Mid-Atlantic Ridge: Lucky Strike.
Rostrum short. Antenna 1 longer than antenna 2. Accessory
flagellum one-articulated. Labrum entire, molar triturative.
Gnathopods different in size. Epimeral plate 3 smooth. Outer
rami of uropods 1-3 shorter than inner rami. Telson cleft with-
out armament. (Species) Body smooth. Eyes absent. Rostrum
short. Antenna 1 long as body length. Antenna 2 slightly
longer than half of antenna 1. Gnathopods slender, linear. Tel-
son triangular, cleft.

1: A: Habitus; B: Accessory flagellum; C: Gnathopod 2; D: Epimeral plates; E: Uropod 1; F: Uropod 2;

G: Uropods 3 and telson; from BELLAN-SANTINI & THURSTON (1996).

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

D. BELLAN-SANTINI Denisia 18 (2006): 389

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

Arthropoda, Crustacea, Amphipoda, Corophiidea, Ischyroceridae

Bonnierella compar MYERS & CUNHA, 2004

peduncle elongate, broad proximally, narrow distally, rami

Size: 3 mm.
much less than half length of peduncle. Telson without cusps or
Morphology: Family: Head anteroventral margin moderately to spines.
strongly recessed and moderately excavate. Gnathopod 2 en-
Biology: Collected in the vicinity of active venting sites to-
larged in males and females. Pereopods 3-4 basis glandular.
gether with filter-feeding organisms (small sponges, hydrozoans
Uropod 3 peduncle broad proximally, narrow distally, rami with
and cirripeds). Some of the specimens of B. compar were found
tiny apical setae. Telson dorsoventrally thickened. Species:
inside their tubes of fine pelagic sediments cemented with mu-
Head anteroventral margin strongly regressed. Eyes absent. An-
cus, in some cases built around hydrozoan stalks. A previous
tenna 1 and 2 subequal in length, setose; accessory flagellum
record of the genus in hydrothermal vents is given by SHAW
with one long and one rudimentary article. Maxilliped raptori-
(1989) who collected a single specimen of B. linearis from a sta-
al, with reduced plates and strong palp terminating in a dactyli-
tion at the Juan de Fuca Ridge.
form palp article 4. Gnathopod 1 small, unornamented.
Gnathopod 2 similar in both sexes, massive, with palmar exca- Distribution: Mid-Atlantic Ridge: Lucky Strike.
vation and with defining spine on posterior margin. Uropod 3

1: By A. Myers. FG1, FG2 – female gnathopods 1 and 2; MG2 – male gnatophod 2; Lbr – labrum; Md – mandible; Mx1 –
maxilla; Mxp – maxilliped; T – telson; U1, U2, U3 – uropods 1-3.

References:
MYERS A.A. & M.R. CUNHA (2004) J. Mar. Biol. Ass. U.K. 84: 1019-1025.
SHAW P. (1989) Can. J. Zool. 67: 1882-1890.

A.A. MYERS & M.R. CUNHA Denisia 18 (2006): 390

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Ischyroceridae

Bonnierella cf. linearis BARNARD, 1964

setation, and number of spines in the mandibular spine row. But

Size: Up to 4 mm.
virtually all characters used to distinguish the present Bon-
Morphology: (Family) Head anteroventral margin moderately nieriella species are only present on the males, which makes the
to strongly recessed and moderately excavate. Gnathopod 2 specific assignment of this specimen dubious.
enlarged in males and females. Pereopods 3-4 basis glandular.
Biology: Unknown. At hydrothermal vents, collected from
Uropod 3 peduncle broad proximally, narrow distally, rami with
washings of associated vent fauna (vestimentiferans and ar-
tiny apical setae. Telson dorsoventrally thickened. (Species)
chaeogastropods).
Eyes absent. Lateral lobes of head produced and acute. Anten-
nae nearly as long as body. Gnathopod 1, palm not ornamed. Distribution: Off Peru, 10°13’S 80°05’W, 6324 m. Reported al-
Gnathopod 2, armed with three sharp cusps in male; with two so from hydrothermal vents at Explorer Ridge, Juan de Fuca
blunt process in female defining cusps obsolete. Uropod 1, in- Ridge: site Gulati Gusher.
ner ramus lacking a marginal spine. Uropod 3 with outer ramus
bearing 5 to 6 minute fringe-like spinules and a distal setule.
Telson triangular , the narrow apex blunt. The single specimen
known from hydrothermal vents, a female, differs slightly from
the original description (BARNARD 1964) in the tuberosities of
the palmar margin of gnathopod 2, minor differences in telson

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

References:
BARNARD J.L. (1964) Bull. Am. Mus. Nat. Hist. 127(1): 1-46.
SHAW P. (1989) Can. J. Zool. 67(8): 1882-1890.

D. BELLAN-SANTINI & D. JAUME Denisia 18 (2006): 391

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Apotectonia heterostegos BARNARD & INGRAM, 1990

multitoothed; molar of both mandibles simple, pubescent. In-

Size: Up to 18.3 mm.
ner plate of either maxilla 1-2 fully setose medially. Inner plate
Morphology: (Family) Body compact, robust. Coxae generally of maxilliped with strongly oblique apex. Gnathopod 1 simple
large. Antenna 1, peduncle article 1 large, inflated, articles 2-3 and with reduced coxa. Basis of pereopods 5-7 rounded-attenu-
much shorter, often telescoped. Mouthparts very variable. ate. Oostegite present on coxa 1 of brooding females.
Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-
Biology: Largely unknown. Readily attracted to baited traps;
acteristic for the family, slender and microchelate. Pereopods 5-
some specimens sorted also from siboglinid washings.
7 basis usually broadly expanded. Telson variable. (Genus) Dor-
sal process of urosomite 1 complexly toothed. Gnathopod 1 Distribution: Galapagos Spreading Center, apparently endem-
simple. (Species) Antenna 1, primary flagellum with basal ic to the sites Garden of Eden and Rose Garden.
spines. Urosomite 1 with 2 sharp dorsal teeth. Mandibular right
lacinia mobilis tiny, bifid; left lacinia mobilis scarcely larger and

1: Female 18.3 mm. A: Head; B: Pleosome and urosome with attached uropods and telson; C: First gnathopod; D: Second
gnathopod; E: Proximal portion of pereopods 5-7; from BARNARD & INGRAM (1990).

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

D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 392

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Cyclocaris tahitensis STEBBING, 1888

lacking rostrum. Mandibular palp present. Coxa 1much wider

Size: Up to 15 mm.
at distal margin than at insertion. Epimeron 3 with posterior
Morphology: (Family) Body compact, robust. Coxae generally angle rounded. Rami of uropod 1 about equal in length. Pe-
large. Antenna 1 peduncle article 1 large, inflated, articles 2-3 duncle of uropod 2 shorter than rami.
much shorter, often telescoped. Mouthparts very variable.
Biology: Unknown. Originally described from volcanic mud
Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-
bottoms around Tahiti and trapped also around Cape Verde on
acteristic for the family, slender and microchelate. Pereopods 5-
sandy bottoms, the only report of the species from hydrother-
7 basis usually broadly expanded. Telson variable. (Genus)
mal vents could be accidental since the accompanying fauna
Head tall, horizontally short, weakly grotesque. Coxae 1-2
was also not typically hydrothermal.
small, strongly shortened and partly covered by coxa3. Coxa 4
large excavate. Gnathopod 1 long, simple. Uropod 3 elongate. Distribution: Off Tahiti, 17°30’26’’S, 149°33’45’’W (STEBBING
Telson elongate, deeply cleft. (Species) Base of primary flagel- 1888). Off Cape Verde Islands, 1477 m. East Pacific Rise,
lum of antenna 1 with callinophore. Body capable of coiling in- Guaymas Basin.
to a circle, bending its head round to the protection of the cox-
ae of the third and fourth pereopods. Head extremely short,

1: Habitus; after STEBBING (1888).

References:
CHEVREUX E. (1935) Résult. Camp. Sci. Prince Albert Ier de Monaco 90: 1-214.
STEBBING T.R.R. (1888) Rep. Sci. Results Voyage H. M. S. Challenger Years 1873-1876, Zoology 29: 1-1737.
VINOGRADOV G.M. (1993) Zool. Zh. 72(2): 40-53 [in Russian].

D. BELLAN-SANTINI & D. JAUME Denisia 18 (2006): 393

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Euonyx mytilus BARNARD & INGRAM, 1990

Gnathopod 1 with propodus about 1.2 times as long as carpus.

Size: Up to 20 mm.
Palm of gnathopod 2 long. Epimeron 2 with a strong tooth.
Morphology: (Family) Body compact, robust. Coxae generally Epimeron 3 with tiny posteroventral tooth.
large. Antenna 1 peduncle article 1 large, inflated, articles 2-3
Biology: Members of this genus come readily to baited traps
much shorter, often telescoped. Mouthparts very variable.
and some appear occasionally associated with echinoderms and
Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-
deep-sea corals. Euonyx mytilus was sorted from washings of
acteristic for the family, slender and microchelate. Pereopods 5-
vent clams, trapped also with baited traps, and also caught di-
7 basis usually broadly expanded. Telson variable. (Genus)
rectly with a slurp gun around vent fauna.
Mandible with palp attached strongly distal. Inner plate of
maxilla 1 weakly setose. Coxa 1 strongly shortened. Gnathopod Distribution: East Pacific Rise, 13°N; Galapagos Spreading
1 elongate. Inner ramus of uropod 2 without notch. Uropod 3 Center: Garden of Eden and Rose Garden.
ordinary. Telson elongate, deeply cleft. (Species) Body lacking
any prominent dorsal tooth. Lateral cephalic lobes rounded.

1: Female 20.04 mm; A: anterior portion of body, lateral; B: pleosome and urosome with uropods and telson attached, lateral;
C: First gnathopod; D: second gnathopod; E: fifth to seventh pereopods; from BARNARD & INGRAM (1990).

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

D. BELLAN-SANTINI & D. JAUME Denisia 18 (2006): 394

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Hirondellea glutonis BARNARD & INGRAM, 1990

shorter than carpus, propodus minutely chelate. Telson elon-

Size: Up to 12.6 mm.
gate, cleft. (Species) Dactylus of gnathopod 1 scarcely overlap-
Morphology: (Family) Body compact, robust. Coxae generally ping palm. Epimera 3 rounded behind. Inner ramus of uropod 2
large. Antenna 1 peduncle article 1 large, inflated, articles 2-3 constricted. Telson weakly cleft.
much shorter, often telescoped. Mouthparts very variable.
Biology: Attracted by baited traps, but caught also directly with
Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-
slurp gun on and around, or sorted from vent mussel and clam
acteristic for the family, slender and microchelate. Pereopods 5-
washings. Most specimens had the midgut densely packed with
7 basis usually broadly expanded. Telson variable. (Genus)
bait food, and with the sternites ventrally extended.
Mandible with incisor ordinary, molar simple, palp attached
opposite molar. Coxa 1 strongly shortened. Gnathopod 1 short, Distribution: Galapagos Spreading Center, East Pacific Rise.
subchelate, palm transverse. Gnathopod 2, propodus greatly

1: Female 12.45 mm; A: head; B: Detail of coxal plates of anterior pereopods; C: Pleosome and urosome with attached uropods
and telson; D: First gnathopod; E: Second gnathopod; F: Proximal portion of pereopods 5-7; from BARNARD & INGRAM (1990).

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

D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 395

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Orchomene (Abyssorchomene) abyssorum STEBBING, 1888

short and covering 100% of palmar edge, palm defined thin

Size: 6.1 mm in adult male.
Morphology: (Family) Body compact. Peduncle of antenna 1
short and stout, articles 2-3 much shorter than 1 and partly
telescoped. Article 3 of gnathopod 2 elongate. (Genus) Mouth-
parts forming quadrate bundle. Coxa 1 large and visible not ta-
pering. Gnathopod 1 subchelate. Inner ramus of uropod 2 with-
out notch. (Species) Lateral cephalic lobes broadly rounded.
Eyes long, lunate, colorless in alcohol, glandular, ommatidia
not evident. Peduncle of antenna 1 barely keeled. Carpus of
gnathopod 1 short, posterior lobe narrow, well exceeding pos-
terior tangential line between merus and propodus. Propodus of
gnathopod 2 with palm short, chelate, not lunate, dactylus
spinule and bearing weak inner setal basket. Article 2 of pere-
opods 5-7 with sparse, weak posterior setule-notches. Epimeral
plate 3 weakly sinuous posteriorly, produced into weak blunt
tooth posteroventrally, smooth. Dorsal process of urosomite 1
hood-shaped, weakly over-vaulting urosomite 2. Telson cleft
about 50% of its length.
Biology: Sampled in mussel and clam beds.
Distribution: Galapagos Spreading Center; worldwide abyssal
distribution to 9000 m.

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

References:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contrib. Zool. 499: 1-80.
THURSTON M.H. (1990) Prog. Oceanogr. 24: 257-274.

D. BELLAN-SANTINI Denisia 18 (2006): 396

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Orchomene (Abyssorchomene) distinctus BIRSTEIN & VINOGRADOV, 1960

covering only 60% of palmar edge, palm defined by cusp. Arti-

Size: 12 mm in adult male.
cle 2 of pereopods 5-7 with sparse, very weak posterior setule-
Morphology: (Family) Body compact. Peduncle of antenna 1 notches. Epimeral plate 3 broadly rounded posteroventrally,
short and stout, articles 2-3 much shorter than 1 and partly smooth. Dorsal process of urosomite 1 hood-shaped, weakly
telescoped. Article 3 of gnathopod 2 elongate. (Genus) Mouth- over-vaulting urosomite 2. Telson cleft about 53% of its length.
parts forming quadrate bundle. Coxa 1 large and visible not ta-
Biology: Sampled in sediments by slurp gun and in washings of
pering. Gnathopod 1 subchelate. Inner ramus of uropod 2 with-
siboglinid tubeworms.
out notch. (Species) Lateral cephalic lobes broadly rounded.
Eyes long, flask-shaped, very pale pink. Carpus of gnathopod 1 Distribution: East Pacific Rise: 13°N; worldwide abyssal distri-
short, posterior lobe narrow, scarcely exceeding posterior tan- bution to 4850 m.
gential line between merus and propodus. Propodus of gnatho-
pod 2 with palm elongate, transverse, sinuate, dactylus long but

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

References:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contrib. Zool. 499: 1-80.
THURSTON M.H. (1990) Prog. Oceanogr. 24: 257-274.

D. BELLAN-SANTINI Denisia 18 (2006): 397

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Tectovalopsis diabolus BARNARD & INGRAM, 1990

cal, densely setulose, tapering to tiny apical plaque with weak

Size: Up to 23.4 mm.
triturative surface. Inner plate of maxillae with medial setae.
Morphology: (Family) Body compact, robust. Coxae generally Gnathopod 1 not elongate and subchelate, with reduced coxa.
large. Antenna 1 peduncle article 1 large, inflated, articles 2-3 Palm of gnathopod 2 long, strongly oblique. Teeth on epimera
much shorter, often telescoped. Mouthparts very variable. 2-3 weak. Epimeron 2 lacking facial spine. Keel of urosomite 1
Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char- simple. Urosomite 3 with erect dorsal keel.
acteristic for the family, slender and microchelate. Pereopods 5-
Biology: Unknown. Collected with slurp gun or baited traps.
7 basis usually broadly expanded. Telson variable. (Genus)
Mandible with molar large, conical, setulose; palp attached Distribution: East Pacific Rise: 13°N. Apparently endemic to
strongly distal of molar. Coxa 1 strongly shortened. Gnathopod this vent site.
strongly subchelate. Urosomite 1 carinate. Telson elongate
deeply cleft. (Species) Mandibular incisor with two teeth; right
lacinia mobilis very small, flake-like, much broader than long;
left lacinia also broader than long, evenly serrate; molar coni-

1: Female 19.1 mm; A: Head; B: Pleosome and urosome with uropods and telson attached; C: Antennule; D: Second gnathopod;
from BARNARD & INGRAM (1990).

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

D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 398

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Tectovalopsis wegeneri BARNARD & INGRAM, 1990

adz-shaped coxa 2; (4) the slightly stouter lobe of coxa 4; (5)

Size: Up to 33.9 mm.
Morphology: (Family) Body compact, robust. Coxae generally
large. Antenna 1 peduncle article 1 large, inflated, articles 2-3
much shorter, often telescoped. Mouthparts very variable.
Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-
acteristic for the family, slender and microchelate. Pereopods 5-
7 basis usually broadly expanded. Telson variable. (Genus)
Mandible with molar large, conical, setulose; palp attached
strongly distal of molar. Coxa 1 strongly shortened. Gnathopod
1 strongly subchelate. Urosomite 1 carinate. Telson elongate
deeply cleft. (Species) Similar to the foregoing T. diabolus
BARNARD & INGRAM, 1990 except for: (1) the wider peduncle
of antenna 1; (2) the slightly less tapering coxa 1; (3) the less
the slightly weaker gnathopods with shorter carpus and propo-
dus on gnathopod 2; (6) the less oblique palm of gnathopod 2;
(7) the more numerous and more widely spread spines on
epimera 2-3; (8) the less stronger dorsal projection of urosomite
2 and dorsal keel of urosomite 3; and (9) the outer ramus of uro-
pod 2 a bit longer.
Biology: Largely unknown. Collected with slurp gun around
Alvinella polychaetes.
Distribution: East Pacific Rise: 13°N. Apparently endemic to
this vent site.

1: Female 33.9 mm long; from BARNARD & INGRAM (1990).

Reference:
BARNARD J.L. & C. INGRAM (1990) Smithon. Contrib. Zool. 499: 1-80.

D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 399

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Lysianassidae

Transtectonia torrentis BARNARD & INGRAM, 1990

strongly tapering. Urosomite 1 with a sharp dorsal tooth.

Size: Up to 21.2 mm.
Morphology: (Family) Body compact, robust. Coxae generally
large. Antenna 1 peduncle article 1 large, inflated, articles 2-3
much shorter, often telescoped. Mouthparts very variable.
Gnathopod 1 simple, subchelate or chelate. Gnathopod 2 char-
acteristic for the family, slender and microchelate. Pereopods 5-
7 basis usually broadly expanded. Telson variable. (Genus)
Mandible with incisor strongly toothed, laciniae mobilis longer
than broad, molar large, conical. Coxa 1 not shortened and no
strongly covered by coxa 2. Pereopods 5-7 elongate, basis
Telosn elongate, deeply cleft. (Species) Antennae especially
short. Inner plate of maxillae setose medially only along distal
half. Gnathopod 1, palm strongly oblique. Epimeron 3 rounded
behind with six spines narrowly spread.
Biology: Caught associated with worms and attracted to baited
traps.
Distribution: East Pacific Rise: 13°N. This monotypic genus is
apparently endemic to this vent site.

1: Female 10.6 mm; A: Head; B: Pleosome and urosome with uropods and telson attached; C: Coxal plates of first and second
gnathopods; D: First gnathopod; from BARNARD & INGRAM (1990).

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

D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 400

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Pardaliscidae

Halice hesmonectes MARTIN, FRANCE & VAN DOVER, 1993

Size: 5.3 mm in adult male. Biology: In large monospecific swarms in the immediate vicin-
ity of low temperature vent openings. 1000 ind. l-1 estimated.
Morphology: (Family) Body laterally compressed, coxae short.
Accessory flagellum well developed, molar absent. (Genus) Distribution: East Pacific Rise: 9°N.
Rostrum well developed. Eyes absent. Gnathopods simple, slen-
der. Pereopods simple. (Species) Teeth of urosomites 1-2
strong. Accessory flagellum biarticulate. Dactylus of pereopods
3-4 minute, constricted at mid length. Pereopods 5-7 extreme-
ly long. Telson cleft along 2/3 and terminating in acute tip.

1A: Habitus; B: Accessory flagellum; C: Lateral view of urosome; D: Gnathopod 1; E: Dorsal view of urosome and telson; from
MARTIN et al. (1993).

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

D. BELLAN-SANTINI Denisia 18 (2006): 401

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Pardaliscidae

Pardalisca endeavouri SHAW, 1989

3, posteroventral corner acute. Telson cleft to 80%, lobes wide-

Size: 11.5 mm.
ly separated, each apex tridentate, bearing paired marginal
Morphology: (Family) Body laterally compressed, coxae short. spines, in addition to paired penicillate setae on the upper face.
Accessory flagellum well developed. Molar absent. Inner plate
Biology: Unknown. Collected with slurp gun from the vicinity
of maxilliped short to evanescent. (Genus) Rostrum small. Eyes
of siboglinids and archaeogastropods.
absent. Mandible asymetrical, incisor on left smooth, weakly
toothed, on right strongly toothed. Palp of maxilla 1 apically Distribution: Explorer Ridge: site Gulati Gusher.
expanded. Coxae 1-4 subquadrate. Gnathopods simple. Telson
scarcely elongate, partly cleft. (Species) Body slender smooth.
Antenna 2 with peduncle segments 3 and 4 elongate. Epimeron

1: Male 11.5 mm; after SHAW (1989).

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

D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 402

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Sebidae

Seba profundus SHAW, 1989

gitudinal axis of limb. Gnathopod 2 elongate, propodus slender.

Size: Up to 3.5 mm.
Basis of pereopods 5-7 evenly expanded, with convex anterior
Morphology: (Family) Urosomites 2-3 free or coalescent. An- and posterior margins. Telson triangular, armed with two pairs
tenna 1, peduncle elongate, accessory flagellum two-articulate. of marginal setae.
Gnathopods 1-2 chelate or gnathopod 1 strongly subchelate.
Biology: Sorted from samples of associated vent fauna (the si-
Gnathopod 1 larger than gnathopod 2. Uropod 3 uniramous.
boglinid Ridgeia piscesiae and archaeogastropods). Other mem-
Telson entire. (Genus) Body slender. Urosomites 2-3 coales-
bers of the genus are typically associated either as a commensal
cent. Gnathopods diverse. Gnathopod 1 much the larger, sub-
or as inquiline with invertebrate hosts.
chelate or chelate. Gnathopod 2 strongly chelate. Telson lin-
guiform. (Species) Body stout. Antennae subequal. Epimeron 3 Distribution: Explorer Ridge: Crab Vent.
lacking large posterodistal tooth. Gnathopod 1, subchelate,
with posterodistal corner of propodus extending out from lon-

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

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

D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 403

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Stegocephalidae

Steleuthera ecoprophycea BELLAN-SANTINI & THURSTON, 1996

Size: 3 mm in adult female. Biology: Collected with shrimps Rimicaris exoculata and Choro-
caris chacei, and in mussel washing.
Morphology: (Family) Body appears stout and globular. Coxae
1-4 forms lateral shield. Antennae very short. Mandible with- Distribution: Mid Atlantic Ridge: Snake Pit.
out molar and palp. Gnathopods feeble. (Genus) Accessory fla-
gellum biarticulate. Mandibular incisor toothed. Pereopods 3-4
simple. Article 2 of pereopod 6 unexpanded. Pereopod 7 with 7
articles. (Species) Epimeron 3 with postero-distal corner
prominent and without seta. Gnathopods 1-2 with propodus ta-
pered distally. Basis of pereopod 7 serrulate. Telson broader
than long, scarcely cleft.

1: A: Habitus; B: Antenna 1; C: Mandible; D: Gnathopod 1; E: Pereopod 7; F: Epimeral plates; G: Uropod 3 and telson;
from BELLAN-SANTINI & THURSTON (1996).

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

D. BELLAN-SANTINI Denisia 18 (2006): 404

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Stenothoidae

Metopa (Prometopa) samsiluna BARNARD, 1966

Size: 4.5 mm. Biology: Unknown.

Morphology: (Family) Coxa 1 very small and partially covered Distribution: Originally described from the submarine canyons
by following coxae. Coxa 4 enlarged, shield-like, not posteri- of off southern California (San Clemente Rift Valley, 32°44’N,
odorsally excavate. Uropod 3 uniramous. Telson entire. 118°12’W, 1096-1620 m). Found also in the Gulf of California,
(Genus) Antenna 1 lacking nasiform process. Accessory flagel- in the Guaymas Basin.
lum absent or vestigial. Palp of mandible two- to three-articu-
late. Gnathopods 1-2 subchelate, different from each other in
size and shape. Pereopod 5 with basis rectolinear. Pereopod 6-7
with expanded lobate basis. (Species) Eyes absent Antennae
very long. Accessory flaggellum vestigial. Coxa 2 very broad.
Gnathopod 1 short. Gnathopod 2 with a large medial tooth,
defining corner with large tooth. Telson spinose.

1: Habitus; from BARNARD (1966).

References:
BARNARD J.L. (1966) Allan Hancock Pac. Exped. 27(5): 1-166.
VINOGRADOV G.M. (1993) Zool. Zh. 72(2): 40-53 [in Russian].

D. JAUME & D. BELLAN-SANTINI Denisia 18 (2006): 405

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Stenothoidae

Stenothoe menezgweni BELLAN-SANTINI, 2005

Telson ordinary, flat. (Species) Antenna 1 with accessory fla-

Size: 5 mm in adult female.
Morphology: (Family) Coxa 1 very small and partially covered
by following coxae, coxa 4 enlarged, shield-like, not posteri-
odorsally excavate. Uropod 3 uniramous. Telson entire.
(Genus) Antenna 1 lacking nasiform process on article 1, ac-
cessory flagellum absent or 1-articulated. Gnathopods 1-2 sub-
chelate, very different from each other in size and shape.
Gnathopod 1 small. Gnathopod 2 large. Pereopod 5 with rec-
tolinear basis. Pereopods 6-7 with expanded and lobate basis.
gellum reduced at a small scale. Eyes present. Gnathopod 1
small. Gnathopod 2 greatly enlarged, with the propodus long
and narrow, palm excavate at the distal part dactylus as long as
half propodus. Pereopods 6-7 slightly lobate. Telson entire.
Biology: Collected on hydrothermal site
Distribution: Mid-Atlantic Ridge: SW of Azores, Menez
Gwen.

1A: Habitus; B: Gnathopod 1; C: Gnathopod 2; D: Epimeral plates; E: Uropod 1; F: Uropod 2; G: Uropods 3; H: Telson; from BEL-
LAN-SANTINI (2005).

Reference:
BELLAN-SANTINI D. (2005) J. Nat. Hist. 39(15): 1101-1110.

D. BELLAN-SANTINI Denisia 18 (2006): 406

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

Arthropoda, Crustacea, Amphipoda, Gammaridea, Stenothoidae

Torometopa saldanhae BELLAN-SANTINI, 2005

strongly indented, limited at the distal corner by a tooth, pos-

Size: 4 mm in adult male.
terior margin fringed with six small spines, dactylus as long as
Morphology: (Family) Coxa 1 very small and partially covered the palm, margin smooth. Pereopods 6-7 with expanded and lo-
by following coxae, coxa 4 enlarged, shield-like, not posteri- bate basis, more expanded in pereopod 7. Telson entire, longer
odorsally excavate. Uropod 3 uniramous. Telson entire. than wide, distal triangular.
(Genus) Antenna 1 lacking nasiform process on article 1, ac-
Biology: Collected a few meters away from the fluid emission
cessory flagellum zero- to two-articulated. Gnathopods 1-2 dif-
among sponges or dead mussels, and out of the active hy-
ferent from each other in size and shape. Gnathopod 1 small,
drothermal areas amongst gorgonians. This species is present
almost simple or weakly subchelate. Gnathopod 2 large. Pereo-
within hydrothermal sites and in the bathyal environment, can
pod 5, basis not lobate in the middle of the article but with a
be regarded as an opportunist amongst hydrothermal communi-
poster-distal lobe expanded to the end of ischium. Pereopods 6-
ties.
7 with expanded and lobate article 2. Telson ordinary, flat.
(Species) Antenna 1 without accessory flagellum. Eyes present Distribution: Mid-Atlantic Ridge: Menez Gwen, Lucky Strike,
of moderate size. Gnathopod 1 subchelate, feeble, propodus Rainbow.
with a short and oblique palm, defined by a spine. Gnathopod
2 large, propodus distally expanded with palm transverse,

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

Reference:
BELLAN-SANTINI D. (2005) J. Nat. Hist. 39(15): 1101-1110.

D. BELLAN-SANTINI Denisia 18 (2006): 407

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

Arthropoda, Crustacea, Amphipoda, Lysianassoidea, Uristidae

Ventiella sulfuris BARNARD & INGRAM, 1990

Size: 6.4 mm in adult female. Biology: Collected from worms and clams washings as well as
crab trap wash. Very abundant. Ventiella sulfuris is the most fre-
Morphology: (Family) Body compact. Peduncle of antenna 1
quent species in Eastern Pacific and represents up to 98% of the
short and stout, articles 2-3 much shorter than 1 and partly
catches.
telescoped. Article 3 of gnathopod 2 elongate. (Genus) Mouth-
parts forming quadrate bundle, labrum dominant, molar tritu- Distribution: Galapagos Spreading Center; East Pacific Rise.
rative. Eyes absent. Coxa 1 strongly shortened partly covered
by coxa 2. Gnathopod 1 short, poorly subchelate. Inner ramus
of uropod 2 without notch. Telson short weakly cleft. (Species)
Monotypic.

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

References:
BARNARD J.L. & C. INGRAM (1990) Smithson. Contr. Zool. 499: 1-80.
VINOGRADOV G.M. 1993. Zool. Zh. 72(2): 40-53 [in Russian].

D. BELLAN-SANTINI Denisia 18 (2006): 408

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

Arthropoda, Crustacea, Euphausiacea, Euphausiidae

Thysanoessa parva HANSEN, 1905

Curiously, all the specimens were juveniles. Chemical analysis

Size: Max. length 9-10.5 mm.
Morphology: Rostrum acute, ending beyond the eyes. Eyes
large, with a transverse constriction between two lobes, the up-
per one being smaller than the lower. Carapace with a lateral
small denticle on its inferior margin, posterior to its mid-point.
Thoracopods almost uniform in structure, although the second
pair is elongated; their setae are provided with setules visible
only at high magnifications. The eighth pair is rudimentary.
The seventh pair has well developed exopodites; endopodites
are absent in males and short in females. Sixth abdominal seg-
ment shorter than the sum of lengths of the two preceding seg-
ments. Photophores are present on the eye stalks, at the bases
of the thoracopods 3 and 7 and on the mid ventral part of ab-
dominal segments 1-4.
Biology: It is interesting to note the great abundance of speci-
mens in almost all samples from the hydrothermal vent field
Rainbow (Mid-Atlantic Ridge). Indeed, a total of 700 speci-
mens were caught in the different traps (KHRIPOUNOFF et al.
2001). They were regularly present from September to January
and more markedly in October and November at different dis-
tances from the vents. They were always absent in traps from
February to June, except for traps laid far off vent sites (2 km).
of the composition of particles from the stomach content has
been done by EDS X-ray microanalysis. A comparison between
T. parva specimens and specimens of the related species T. gre-
garia indicate that probably the T. parva population is more or
less linked with the particle flux. Indeed, a peak of the Fe ele-
ment is obvious and might be related with the high concentra-
tion of iron (7%) in the vent particles (KHRIPOUNOFF et al.
2001) that does not exist in T. gregaria (unpublished data).
However, studies of T. parva specimens caught far from the
vents have yet to verify this hypothesis. Curiously, there were
no euphausiaceans in traps laid in the South Atlantic (Gulf of
Guinea) and in the Pacific (East Pacific Rise: 13°N) (M.
Segonzac, pers. comm.).
Distribution: It is a rare meso-bathypelagic species. In the At-
lantic, it has been recorded off Portugal and Africa, from the
latitude of Gibraltar to near Cap Town; on the western side, it
is mentioned near the Bermuda Islands. In the Pacific, it is
known in scattered areas: east of south Japan, off California and
east of New Zealand. In the Indian Ocean, it exists west of
Sumatra. Mid-Atlantic Ridge: Rainbow.

1: Adult male; from BAKER et al. (1990).

References:
BAKER A. DE C., BODEN B.P. & E. BRINTON (1990) A Practical Guide of the Euphausiids of the World. Nat. Hist. Mus. Publ., London: 1-96.
BRINTON E. (1962) Bull. Scripps Inst. Oceanogr. 8: 51-270.
KHRIPOUNOFF A., VANGRIESHEIM A., CRASSOUS P., SEGONZAC M., COLACO A., DESBRUYÈRES D. & R. BARTHELEMY (2001) J. Mar. Res. 59: 633-656.
MAUCHLINE J. & L.R. FISHER (1969) Adv. Mar. Biol. 7: 1-454.

R. BARTHELEMY & B. CASANOVA Denisia 18 (2006): 409

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Decapoda, Caridea
MARTIN & HANEY (2005) listed 52 species of nine families
of caridean shrimp from the vicinity of hydrothermal vents and
cold seeps, although several of them have not been identified
to species level. Of them species of the family Alvinocarididae
are endemic to such reduced environments, whereas the occur-
rence of most of other families is considered to be rather op-
portunistic. It is highly likely that pelagic or demersal species
(particularly species of Oplophoridae) will be further recorded
from waters around vent sites by future studies. Periclimenes
thermohydrophilus of the Palaemonidae is associated with the tu-
beworm Lamellibrachia satsuma, and such commensalism is rat-
her unique among the shrimp species known from the chemo-
synthetic communities.
This book may give an impression that inventory
of the caridean fauna of the hydrothermal vents or cold seeps is
nearly completed. Indeed, in particular, thanks to the recent
studies, progress on the taxonomy of the Alvinocarididae has
been made, but there are still some species to be described
(MARTIN & HANEY, 2005). For example, in spite of the revision
by KOMAI & SEGONZAC (2005) and subsequent work by KOMAI
et al. (2005), the taxonomic identity of some species of Alvino-
caris remain unclear. There is a tendency that oceanographic
researchers publish lists of vent or seep communities based on
identification made in a rather preliminary manner and that
the voucher material is stored at universities or personal free-
zers or other non-recognized repositories. This makes access to
the specimens, as well as accurate assessment of specific identi-
ties, difficult. It is advisable to send specimens for accurate
identifications to relevant specialists and to deposit voucher
specimens in museums or other appropriate institutions.
References:
KOMAI T. & M. SEGONZAC (2005) Journal of Natural History 39(15): 1111-1175.
KOMAI T. SHANK T.M. & C.L. VAN DOVER (2005) Zootaxa 1019: 27-42.
MARTIN J.W. & T. HANEY (2005) Zool. J. Linn. Soc. 145: 445-522.
T. KOMAI
Below, some important points are given:
1. Juveniles of congeneric species of Alvinocarididae are
very similar in morphology and sometimes it is impossible to
identify them without locality data. In case of the occurrence of
more than one congeneric species sympatrically, identification
should only be done to genus level.
2. Adults of alvinocaridids often exhibit a high degree of
polymorphism in the morphology of the chela and ambulatory
legs. Therefore careful observations of various characters is ne-
cessary for accurate identification.
3. Some species of alvinocaridids, for example, species of Ri-
micaris, shows dramatic ontogenetic change after postlarval sta-
ges. Therefore, it is strongly recommended that a formal des-
cription of new species is based on adult specimen(s).
4. For any work including results of identification of spe-
cies, it is recommended that voucher specimens are deposited
in a recognized museum collection. For morphological studies,
specimens preserved in 70–75% ethanol is recommended, alt-
hough formalin fixed specimens are also useful. However, 70–
75% ethanol or formalin may be not adequate for DNA extrac-
tion. Therefore, it is advisable to preserve specimens in two dif-
ferent concentration of ethanol according to further treatment.
5. Color in life is sometimes very useful in discriminating
caridean species, particularly hippolytids. It is advisable to take
color photographs just after capture of specimens.
6. There is little doubt that new species are likely to be dis-
covered by new expeditions even at “well-known“ or “well-col-
lected“ sites.
Denisia 18 (2006): 410
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Decapoda, Caridea, Nematocarcinidae
Nematocarcinus burukovskyi KOMAI & SEGONZAC 2005
Size: Carapace length 25.2 mm, total length ca. 74 mm.
Color: In life, body light orange, yellowish hepatopancreas vis-
ible through integument. Cornea of eye brown, but reflective;
eye-stalk dark orange. Pereopods nearly colorless.
Morphology: Body surface smooth. Rostrum slender, slightly
falling short of anterior margin of antennal scale, about 0.6 of
carapace length; dorsal margin armed with three fixed teeth;
ventral margin with four fixed teeth. Carapace relatively well
sculptured with distinct postorbital, cervical, post-cervical, he-
patic, branchiocardiac grooves; postrostral median ridge with
six teeth clearly separated from three teeth on rostrum proper,
at least some of them with basal suture. Abdomen dorsally
smooth; fifth somite with one or two acute posteroventral
teeth; inner pleural ridge on fifth somite low; sixth somite with
two rows of setae on ventral surface, but devoid of paired pos-
teroventral spots. Telson with six pairs of small dorsolateral
spines; all dorsolateral spines aligned; terminal margin with
three pairs of spines, second pair longest. Eye with well devel-
oped, distinctly faceted cornea. Second pereopod longest
1, 3-5: Holotype, subadult male carapace length 18.6 mm, from Wormwood vent site, Southern East Pacific Rise: 17°S
© J.-F. Dejouannet/IRD.
1: Habitus; scale bar 1 cm.
T. KOMAI & M. SEGONZAC
among pereopod. Third to fifth pereopods markedly elongate,
but still relatively short and stout for nematocarcinids; propodi
very short, subcylindrical. Exopods on third and fourth pere-
opods very short, particularly that on fourth pereopod rudi-
mentary.
Remarks: This species can be confused in situ with Nemato-
carcinus ovalis, whose a female ovigerous carapace length 27.5
mm, collected a few tens of meters above the hydrothermal
sites East Pacific Rise: 13°N, 2600 m, presents a similar color.
This species is observed on the active sites, but also in the
abyssal milieu at several hundred meters of sites.
Biology: Live around the active chimneys diffusing milky fluid
at low temperature between 2 and 7°C, above beds of the
mytilid Bathymodiolus spp., close to siboglinid tubeworm Riftia
pachyptila, crabs Bythograea spp., and galatheid crabs Munidopsis
spp. Necrophagous feeder.
Distribution: Southern East Pacific Rise: 17°S, 23°S and 31°S.
2: Individual in situ, taken at Fred’s Fortress
vent site, Southern East Pacific Rise: 31°S; cruise
PAR 5; on mussel bed of Bathymodiolus sp.,
with bythograeid crabs (Bythograea
vrijenhoeki); by R. Vrijenhoek © MBARI.
Denisia 18 (2006): 411-412
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

3: Rostrum, anterior part of carapace and cephalic appendages, lateral view;

scale bar 5 mm.

4: Posterior part of third abdominal somite to telson and left uropod, lateral 5: Telson, dorsal view; scale
view; scale bar 5 mm. bar 2 mm.

6: Holotype taken on board, just after collection; cruise
Biospeedo; P. Briand © Ifremer.
7: Paratype juvenile carapace length 13.8
mm, rostrum, anterior part of carapace and
cephalic appendages, taken on board, just
after collection; Southern East Pacific Rise:
31°S; cruise PAR 5; G. Rouse © Australia.

Reference:
KOMAI T. & M. SEGONZAC (2005) Zoosystema 27(2): 343-364.

412
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
Alvinocaris brevitelsonis KIKUCHI & HASHIMOTO, 2000
Size: Only the ovigerous female holotype is known: carapace
length 13.8 mm, total length ca. 57 mm.
Color: Ivory to dull ivory white.
Morphology: Rostrum directed forward, slightly curved dorsal-
ly, 0.55% of carapace length, reaching distal margin of second
segment of antennular peduncle; dorsal margin armed with 13
teeth, including eight teeth on rostrum proper and five rela-
tively large teeth on carapace posterior to orbital margin, pos-
teriormost tooth arising from 0.38% of carapace length; ventral
margin armed with seven small teeth on anterior 0.60%. Cara-
pace 0.7 times as wide as long; postrostral ridge relatively high,
dorsal angle about 150°; branchial region not notably inflated,
slightly convex; pterygostomian tooth relatively weak. Fourth
abdominal pleuron with three tiny teeth on posteroventral
margin. Fifth abdominal pleuron with strong posteroventral
tooth and one additional tooth on posterior margin. Telson
falling short of posterior margin of uropodal endopod, armed
with six or seven dorsolateral spines; posterior margin moder-
ately convex, armed with six pairs of spines (second pair longest
and third pair shortest) and two pairs of plumose setae subequal
in length to longest second spine. Eye with small spiniform tu-
2: Anterior part of carapace
and cephalic appendages,
dorsal.
References:
KIKUCHI T. & J. HASHIMOTO (2000) Species Diversity 5: 135-148.
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.
T. KOMAI & M. SEGONZAC
bercle on anterior surface. Antennular peduncle with second
segment 1.8 times longer than wide. Antennal scale half length
of carapace, 1.87 times longer than wide. Third to fifth pere-
opods moderately slender; dactyli each with single row of ac-
cessory spinules; meri armed with spines on ventrolateral sur-
faces; ischia also with spines at least in third and fourth pere-
opods. No epipods on third maxilliped to fourth pereopod. Sec-
ond to fourth pereopods each with slender, cincinnulate ap-
pendix interna.
Remarks: A. brevitelsonis is represented only by the holotype.
The possession of more than two pairs of spines on the posteri-
or margin of the telson links A. brevitelsonis to A. lusca and A.
stactophila. As KIKUCHI & HASHIMOTO (2000) correctly point-
ed out, the shorter telson, which falls short of the posterior end
of the uropod, distinguishes A. brevitelsonis from A. lusca. A.
dissimilis and Shinkaicaris leurokolos occurred sympatrically.
Biology: This species occurs at a hydrothermally influenced
area.
Distribution: Okinawa Trough: Minami-Ensei Knoll.
1: Body, lateral.
1-3: Holotype
ovigerous female
(carapace length
13.8 mm) from Mi-
nami Ensei Knoll,
Mid-Okinawa
Trough;
scale bars 1 mm;
by T. Kikuchi.
3: Telson and
left uropod,
dorsal.
Denisia 18 (2006) 413
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Alvinocaris dissimilis KOMAI & SEGONZAC, 2005

ners and 12-14 plumose setae all longer than mesial pair of lat-
Size: Largest male carapace length 8.4 mm, largest female cara-
eral spines.
pace length 9.8 mm, ovigerous females carapace length 7.3-9.8
mm. Maximal total length 41 mm. Remarks: Alvinocaris dissimilis, A. markensis, A. muricola and A.
longirostris are morphologically very similar to one another.
Morphology: Rostrum directed forward, straight or very slight-
These four species share the rostrum reaching at least to the
ly curved dorsally, usually reaching to second segment of an-
second segment of the antennular peduncle and armed usually
tennular peduncle; dorsal margin armed with 13-17 teeth, in-
with one or more ventral teeth, relatively high postrostral me-
cluding 8-10 teeth on rostrum proper and 5-8 relatively small
dian ridge on the carapace, and the posterior margin of the tel-
teeth on carapace posterior to orbital margin; ventral margin
son with long plumose setae and two lateral pairs of spines.
armed with one or two small subdistal teeth. Carapace width
These characters should be used with caution, as most of them
0.65-0.80 of length; dorsal angle about 155°. Fourth abdominal
are useful only for adult specimens.
pleuron with small posteroventral tooth and additional one or
two teeth on posterior margin. Fifth abdominal somite similar- Biology: This recently described species occurs in a hydrother-
ly armed with strong posteroventral tooth and additional 1-5 mally active area. Alvinocaris brevitelsonis and Shinkaicaris leu-
tiny teeth on posterior margin or ventral margin. Telson not rokolos occurred sympatrically with this species.
reaching posterior margin of uropodal endopod; armed with 5-
Distribution: Known only from Minami-Ensei Knoll, Mid-Ok-
8 dorsolateral spines; posterior margin convex (rarely shallowly
inawa Trough, 705 m.
notched medially), armed with two pairs of spines at lateral cor-

1: Carapace and cephalic appendages, lateral.

3: Anterior part of carapace and
cephalic appendages, dorsal.

1-4: Holotype
ovigerous female
(carapace length
11.9 mm) from
Minami-Ensei
Knoll, Mid-
Okinawa Trough;
by T. Komai.
4: Telson and left uropod, dorsal (marginal setae on
2: Third to sixth abdominal somites, lateral. uropod omitted).

References:
KIKUCHI T. & J. HASHIMOTO (2000) Species Diversity 5: 135-148.
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.

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

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Alvinocaris longirostris KIKUCHI & OHTA, 1995

14 plumose setae all longer than mesial pair of lateral spines.

Size: Largest female 16.9 mm. Maximal total length ca. 85 mm.
Eye with small spiniform tubercle on anterior surface. Anten-
Color: Ivory to dull ivory white. nular peduncle with second segment 1.58-1.69 times longer
than wide. Antennal scale about half lengh of carapace and
Morphology: Rostrum slightly descending or directed forward,
1.72-1.90 times longer than wide. Third to fifth pereopods
straight or weakly curved dorsally, 0.64-1.26 of carapace length
moderately slender; dactyli each with single row of accessory
in females, usually overreaching distal margin of antennular pe-
spinules; meri armed with spines on ventrolateral surfaces; is-
duncle, armed with 9-15 teeth including 5-10 relatively large
chia also with spines at least in third and fourth pereopods. No
teeth on carapace posterior to orbital margin, posteriormost
epipods on third maxilliped to fourth pereopod. Second to
tooth arising 0.38-0.48 of carapace length; ventral margin
fourth pereopods each with slender, cincinnulate appendix in-
armed with 4-9 small teeth on anterior 0.30-0.50. Carapace
terna.
width 0.65-0.72 of length; postrostral median ridge relatively
high, dorsal angle about 150°; branchial region not notably in- Biology: Usually associated with hydrothermal vents. FUJIKURA
flated, slightly convex; pterytostomial tooth relatively strong. ET AL. (1995) reported the occurrence of A. longirostris at cold
Fourth abdominal pleuron with 1-4 posterolateral teeth. Fifth seeps on off Hatsushima site, Sagami Bay.
abdominal pleuron similarly armed with strong posteroventral
Distribution: Okinawa Trough: Iheya Ridge and Hatoma
tooth and additional 2-5 small teeth. Telson not reaching to
Knoll; Japan: Sagami Bay, off Hatsushima site; Kermadec-Arc:
posterior margin of uropodal endopod, distinctly narrowed pos-
Brothers Seamount Caldera.
teriorly, armed with 5-7 dorsolateral spines; posterior margin
convex, armed with two pairs of spines at lateral angle and 12-

2: Posterior part of abdomen and uropods in ven-

1: Adult female, lateral.
tral view.

3: Anterior part of
carapace and
cephalic ap-
pendages, dorsal.

4: Telson and
uropod, dorsal.
1-4: from KIKUCHI &
OTHA (1995).

References:
FUJIKURA K., HASHIMOTO J., FUJIWARA Y. & T. OKUTANI (1995) JAMSTEC J. Deep-Sea Res. 11: 227-241.
KIKUCHI T. & S. OHTA (1995) J. Crustac. Biol. 15: 219-240.
WATABE H. & H. MIYAKE. (2000) JAMSTEC J. Deep-Sea Res. 17: 29-34.
WEBBER W.R. (2004) Zootaxa 444: 1-26.

T. KOMAI Denisia 18 (2006): 415

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Alvinocaris lusca WILLIAMS & CHACE, 1982

Size: Carapace length largest male 7.3 mm; largest female 13.5
mm. Maximal total length ca. 85 mm.
Morphology: Rostrum slightly descending or directed forward,
straight, 0.37-0.44 times of carapace length, usually reaching
second segment of antennular peduncle, armed with 10-14
teeth including 4-6 moderately large teeth on carapace posteri-
or to orbital margin, posterior tooth arising 0.32-0.39 of cara-
pace length; ventral margin armed with three or four small
teeth on anterior 0.20-0.40. Carapace width 0.55-0.67 of
length; postrostral median ridge relatively low, dorsal angle
about 170°; branchial region weakly convex; pterytostomial
tooth relatively weak. Fourth abdominal pleuron with 1-5
(most frequently four) posterolateral teeth. Fifth abdominal
pleuron similarly armed with strong posteroventral tooth and
additional 1-5 small teeth. Telson not reaching to slightly over-
reaching posterior margin of uropodal endopod, very slightly
narrowed posteriorly, armed with 7-9 dorsolateral spines; poste-
rior margin shallowly notched medially or slightly convex,
armed with 5-7 spines (mesial 3-5 spines unequal in length, but
third spine from lateral shorter than others) and one or two
short plumose setae on either side of median notch. Eyes with
small spiniform tubercle on anterior surface. Antennular pe-
duncle with second segment 2.00-2.20 times longer than wide.
Antennal scale about half lengh of carapace and 2.20-2.60
times longer than wide. Third to fifth pereopods moderately
slender; dactyli each with single row of accessory spinules; meri
armed with spines on ventrolateral surfaces; ischia also with
spines at least in third and fourth pereopods. No epipods on
third maxilliped to fourth pereopod. Second to fourth pere-
opods each with slender, cincinnulate appendix interna.
Biology: This species is associated with the vestimentiferan
worm Riftia pachyptila, and could be nourished from its biologi-
cal production. VAN DOVER et al. (1985) described plank-
totrophic larval develoment.
Distribution: Known with certainty only from the Galapagos
Spreading Center: Rose Garden area and East Pacific Rise:
9°N.

1: Body, lateral; enlargement x5.

2: Anterior part of carapace

and cephalic appendages; en-
largement x10.

3: Telson and
uropods, dorsal;
enlargement x10.
1-3: from WILLIAMS
& CHACE (1982).

References:
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.
VAN DOVER C.L., FACTOR J.R., WILLIAMS A.B. & C.J. BERG (1995) Bull. Biol. Soc. Wash. 6: 223-227.
WILLIAMS A.B. & F.A. CHACE (1982) J. Crustac. Biol. 2(1): 136-147.

T. KOMAI & M. SEGONZAC Denisia 18 (2006): 416

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
Alvinocaris markensis WILLIAMS, 1988
Size: Carapace length largest male 12.9 mm; largest female 22.3
mm, ovigerous female 13.0 mm. Maximal total length ca. 82
mm.
Color: In situ red-orange, abdomen paler.
Morphology: Rostrum directed forward or slightly descending,
nearly straight or slightly curved dorsally, armed with 14-19
teeth, including 8-12 on rostrum proper and 5-7 moderately
large teeth on carapace posterior to level of orbital margin, pos-
teriormost tooth arising from 0.24-0.31 of carapace length; ven-
tral margin armed with 6-9 small teeth. Carapace 0.52-0.65
times as wide as long; postrostral median ridge relatively high,
dorsal angle about 150°; pterygostomial tooth relatively strong;
branchial region not particularly inflated, slightly convex.
Fourth abdominal pleuron with 1-4 (most frequently two or 3)
teeth posteriorly. Fifth abdominal pleuron similarly armed with
one strong posterolateral tooth and 1-3 additional teeth on pos-
terior margin. Telson narrowed posteriorly, not reaching or
reaching posterior margin of uropodal endopod, armed with 6-
8 dorsolateral spines; posterior margin always convex, armed
with two pairs of lateral spines and 12-14 plumose setae all
1: Female (carapace length 16.7 mm) from site Les Ruches, Snake Pit, MAR; carapace
and cephalic appendages, lateral; scale bar 5 mm.
T. KOMAI & M. SEGONZAC
longer than mesial pair of lateral spines. Antennular peduncle
moderately stout, second segment 1.79-2.05 times longer than
wide. Eye with small spiniform tubercle on anterior surface.
Antennal scale 1.91-2.04 times longer than wide. Third to fifth
pereopods moderately slender; dactyli each with single row of
accessory spinules; meri armed with spines on ventrolateral sur-
faces; ischia also with spines at least in third and fourth pere-
opods. No epipods on third maxilliped to fourth pereopod. Sec-
ond to fourth pereopods each with slender, cincinnulate ap-
pendix interna.
Biology: This species always lives more or less solitary at the
base and on walls of active chimneys, close to aggregates of
Rimicaris exoculata, or on mussel beds. The trophic mode is
necrophagous, as captures of the shrimps by baited traps indi-
cate. However, the gut contents examined generally contained
mainly mineral particles. Ovigerous females collected in Au-
gust.
Distribution: The occurrence of this species is confirmed at
several locations at Mid-Atlantic Ridge: TAG, Snake Pit, Bro-
ken Spur, Logatchev, Rainbow, and Lucky Strike.
2: Anterior part of carapace and
cephalic appendages, dorsal;
scale bar 2 mm.
3: Telson and right uropod, dorsal; scale
bar 5 mm.
Denisia 18 (2006): 417–418
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

4: Fresh specimen taken on board; from Briand © Ifremer.

5: In situ at the base of hot vent on the site Les Ruches, Snake Pit. The shrimp is seen in pro-
file; one notes the camber cephalothorax/abdomen, characteristic of this species. Surround-
ed by Chorocaris chacei shrimps. Cruise Hydrosnake © Ifremer. 1-5: by T. Komai (from KOMAI
& SEGONZAC 2005).

References:
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.
SEGONZAC M., SAINT LAURENT M. DE & B. CASANOVA (1993) Cah. Biol. Mar. 34: 535-571.
SHANK T., BLACK M.B., HALANYCH K.M., LUTZ R.A. & R.C. VRIJENHOEK (1999) Mol. Phylogenet. Evol. 13: 244-255.
VAN DOVER C.L. (1995) in PARSON L., WALKER C.L. & D.R. DIXON (Eds) Hydrothermal Vents and Processes: 257-294.
WILLIAMS A.B. (1988) J. Crustac. Biol. 2(1): 136-147.
WILLIAMS A.B. (1988) Fish. Bull. 86(2): 263-287.

418
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Alvinocaris niwa WEBBER, 2004

pereopod without fine row of long submarginal setae on outer

Size: Male holotype carapace length 15.38 mm, total length ca.
surface along cutting edges of fingers. Third to fifth pereopods
57 mm.
moderately slender; dactyli armed with two or more rows of ac-
Morphology: Rostrum short, not reaching distal margin of first cessory spinules on ventral surface; meri usually unarmed; is-
segment of antennular peduncle, directed forward, weakly com- chia of third and fourth pereopods armed usually with two lat-
pressed laterally, terminating acutely, dorsal margin sharply eral spines. No strap-like epipods on third maxilliped and pere-
carinae, armed with 5-17 teeth including 4-12 on rostrum prop- opods. Appendices internae on second to fourth pereopods
er and 0-6 on anterior part of carapace posterior to level of or- slender, without coupling hooks.
bital margin; ventral margin usually unarmed or rarely with one
Remarks: The reduced dorsal rostral series, the reduced arma-
tiny subterminal tooth. Carapace somewhat compressed later-
ture of the meri of the third to fifth pereopods and the posses-
ally, with sharp postrostral ridge reaching anterior 0.2 of cara-
sion of two or more rows of accessory spinules on the ventral
pace length; antennal spine acuminate; no conspicuous lobe
surface of each dactylus of the third to fifth pereopods set this
mesial to antennal spine; pterygostomial angle weakly to some-
species apart from Alvinocaris, and link it to a group composed
what produced in adults, reaching or distinctly overreaching
of Shinkaicaris, Opaepele, Chorocaris and Rimicaris.
antennal spine, terminating in sharp spine. Abdomen smooth
dorsally; pleuron of third somite usually smooth, those of fourth Biology: This species is associated with hydrothermal springs
and fifth somites at least with posterolateral tooth and fre- on active submarine volcanoes, among mytilid bivalve Giganti-
quently with additional small teeth ventrally and/or posterior- das gladius, but also abundant bathyal fauna (coral, barnacles,
ly. Telson with six or seven dorsolateral spines arranged in sin- asteroid echinoderms).
uous row on either side; posterior margin convex, with 1-3
Distribution: Kermadec-Arc: Brothers Seamount Caldera and
small spines at each lateral angle and row of numerous long
Rumble V Seamount.
plumose setae. Eyestalks degenerated, broadly fused mesially,
cornea unfacetted; anterior surface unarmed. Chela of first

1: Lateral view; scale bar 5 mm.

3: Front, 2: Abdomen, ventrolateral

dorsal view. view; scale bar 5 mm.

4: Telson and uropods,

dorsal view.
1-4: Holotype, carapace
length 15.38 mm;
from WEBBER (2004).

References:
WEBBER W.R. (2004) Zootaxa 444: 1-26.
WEBBER W.R. & N.L. BRUCE (2002) Biodiversity Update 5: 6-7.

T. KOMAI & M. SEGONZAC Denisia 18 (2006): 419

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
Alvinocaris williamsi SHANK & MARTIN, 2003
Size: Largest male carapace length 6.1 mm; largest female 10.3
mm, ovigerous female 5.8 mm.
Morphology: Rostrum directed forward, straight or slightly
downward, not reaching or reaching distal margin of first seg-
ment of antennular peduncle, 0.27-0.37 of carapace length;
dorsal margin armed with 10-15 teeth, including 5-9 teeth on
rostrum proper and 4-8 small to moderately large teeth on cara-
pace posterior to orbital margin, posteriormost tooth arising
from 0.13-0.20 of carapace length; ventral margin usually un-
armed. Carapace about 0.70 times as wide as long, postrostral
median ridge relatively low, with dorsal angle about 150°;
branchial region not notably inflated, slightly convex; bran-
chiostegal tooth moderately strong. Fourth abdominal pleuron
rounded or with small posteroventral tooth. Fifth abdominal
pleuron armed with strong posteroventral tooth and 2-4 addi-
tional tiny teeth on posterior margin. Telson not reaching pos-
terior margin of uropodal endopod; armed with six or seven
dorsolateral spines arranged in straight or slightly convex line;
2: Anterior part of
carapace and cephalic
appendages, dorsal.
References:
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.
SHANK T.M. & J.W. MARTIN (2003) Proc. Biol. Soc. Wash. 116: 158-167.
T. KOMAI & M. SEGONZAC
posterior margin broadly rounded, armed with two pairs of
spines at posterolateral angles and 14-16 plumose setae all
longer than mesial pair of lateral spines. Eye with small tuber-
cle on anterior surface. Antennular peduncle very stout, with
second segment 1.05-1.11 times longer than wide. Antennal
scale about 0.4 of carapace length, 1.40-1.50 times longer than
wide. Third to fifth pereopods moderately slender; dactyli each
with single row of accessory spinules; meri usually armed with
spines on ventrolateral surfaces in third and fourth; ischia also
with spines at least in third and fourth pereopods. No epipods
on third maxilliped to fourth pereopod. Second to fourth pere-
opods each with slender, cincinnulate appendix interna.
Biology: The specimens were collected from the mussel beds of
the mytilid Bathymodiolus azoricus.
Distribution: Known only from Mid-Atlantic Ridge: Menez
Gwen.
1: Body, lateral.
3: Telson and left
uropod, dorsal.
1-3: Holotype,
ovigerous female;
scale bar 1 mm;
from SHANK & MAR-
TIN (2003).
Denisia 18 (2006): 420
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae
Chorocaris chacei (WILLIAMS & RONA, 1986)
Size: Female, max. carapace length 17.4 mm; total length 55
mm.
Color: Cephalothorax red orange, abdomen paler.
Morphology: Rostrum broadly rounded, flattened dorsoventral-
ly, reaching or overreaching antennal spines; no dorsal or ven-
tral teeth; ventral surface convex. Carapace somewhat com-
pressed laterally, rounded dorsally; antennal spine blunt; ptery-
gostomial angle strongly produced, terminating bluntly. Fourth
abdominal pleuron blunt angular posteroventrally; fifth pleu-
ron with subacutely pointed posterolateral angle. Telson nar-
rowed posteriorly, bearing row of 6-8 dorsolateral spines
arranged in sinuous row on either side; posterior margin broad-
ly convex, with 1-3 spines at each lateral angle and with row of
numerous plumose setae. Eye-stalks degenerated, broadly fused
mesially without trace of median separation; no distinct spine
or tubercle on anterior surface. Chela of first pereopod without
fine row of long submarginal setae on outer surface along cut-
ting edges of fingers. Third to fifth pereopods moderately slen-
der to stout; dactyli armed with two or more rows of accessory
1: Holotype, lateral view; enlargement x5.2;
from WILLIAMS & RONA (1986).
2: Specimen taken on board; cruise Exomar © Ifremer.
References:
RAMIREZ LLODRA E.,TYLER P.A. & J.T.P. COPLEY (2000) J. Mar. Biol. Ass. U.K. 80: 473-484.
SEGONZAC M., SAINT LAURENT M. DE & B. CASANOVA (1993) Cah. Biol. Mar. 39(15): 1111-1175.
WILLIAMS A.B. & P.A. RONA (1986) J. Crustac. Biol. 6: 446-462.
T. KOMAI & M. SEGONZAC
spinules on ventral surface; meri and ischia unarmed. Third
maxilliped and first to fourth pereopods without strap-like
epipods. Appendices internae on second to fourth pereopods
simple, slender, without cincinnuli. Protopod of endopod ta-
pering distally, terminating subcautely or acutely.
Remarks: A single female specimen collected at the site Eiffel
Tower on Lucky Strike differs from typical series of C. chacei in
having an acuminate rostrum and acuminate antennal and
pterygostomial spines on the carapace. We regard that the spec-
imen is an abnormal individual of C. chacei, because it was col-
lected along with individuals of typical C. chacei and we have
not encountered such a form in other collections from the
Lucky Strike.
Biology: This species is much less abundant (2-3 ind./dm2, by
places) than Rimicaris exoculata. It is present close to chimneys
and on sulfide blocks, mostly where fluid emissions are weak.
Distribution: Mid-Atlantic Ridge: TAG, Snake Pit, Logatchev,
Rainbow, and Lucky Strike.
3: Cephalothorax
of a single aber-
rant specimen
collected at Lucky
Strike in July
2001 (American
cruise DiversExpe-
dition) by P.
Briand © Ifremer.
4: Anterior part of carapace 5: Telson and uropods,
and cephalic appendages, dor- dorsal; cruise Exomar
sal; cruise Exomar © Ifremer. © Ifremer.
Denisia 18 (2006): 421
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Chorocaris paulexa MARTIN & SHANK, 2005

numerous plumose setae. Eye-stalks degenerated, broadly fused

Size: Female, carapace length 9.6 mm
mesially without trace of median separation; no distinct spine
Color: Live shrimp mostly pale translucent white to slightly or tubercle on anterior surface. Chela of first pereopod without
pink on posterior and dorsal surfaces of carapace and abdomen. fine row of long submarginal setae on outer surface along cut-
ting edges of fingers. Third to fifth pereopods moderately slen-
Morphology: Rostrum broadly triangular, flattened dorsoven-
der to stout; dactyli armed with two or more rows of accessory
trally, overreaching antennal spines; no dorsal or ventral teeth;
spinules on ventral surface; meri and ischia unarmed. Third
ventral surface convex. Carapace somewhat compressed later-
maxilliped and first to fourth pereopods without strap-like
ally, rounded dorsally; spawning female with very shallow de-
epipods. Appendices internae on second to fourth pereopods
pression on either side of midline in anterior half; antennal
simple, slender, without couppling hooks. Protopod of endopod
spine blunt; pterygostomial angle strongly produced as relative-
tapering distally, terminating in blunt tooth.
ly narrow triangular projection, terminating subacutely. Fourth
abdominal pleuron bluntly angular posteroventrally; fifth pleu- Biology: Living directly on the black smokers in numbers up to
ron with subacutely pointed posterolateral angle. Telson slight- 47 individuals per half meter square.
ly narrowed posteriorly, bearing row of 6-8 dorsolateral spines
Distribution: Southern East Pacific Rise: 17°S.
arranged in sinuous row on either side; posterior margin broad-
ly convex, with 1-3 spines at each lateral angle and with row of

1: Body, lateral; scale bar 10 mm.

2: Anterior part of carapace and cephalic appendages, dor- 3: Telson and left uropod, dorsal;
sal; scale bar 5 mm. scale bar 2 mm.
1-3: from MARTIN & SHANK (2005).

Reference:
MARTIN J. & T. SHANK (2005) Proc. Biol. Soc. Wash. 118(1): 183-198.

T. KOMAI & M. SEGONZAC Denisia 18 (2006): 422

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Chorocaris vandoverae (MARTIN & HESSLER, 1990)

or tubercle on anterior surface. Chela of first pereopod without

Size: Female carapace length 13.4 mm.
Morphology: Rostrum broadly rounded, flattened dorsoventral-
ly, overreaching antennal spines; no dorsal or ventral teeth;
ventral surface convex. Carapace somewhat compressed later-
ally, rounded dorsally; antennal spine blunt; pterygostomial an-
gle strongly produced, terminating bluntly. Fourth abdominal
pleuron bluntly angular posteroventrally; fifth pleuron with
subacutely pointed posterolateral angle. Telson slightly nar-
rowed posteriorly, bearing row of 6-8 dorsolateral spines
arranged in sinuous row on either side; posterior margin broad-
ly convex, with 1-3 spines at each lateral angle and with row of
numerous plumose setae. Eye-stalks degenerated, broadly fused
mesially without trace of median separation; no distinct spine
fine row of long submarginal setae on outer surface along cut-
ting edges of fingers. Third to fifth pereopods moderately slen-
der to stout; dactyli armed with two or more rows of accessory
spinules on ventral surface; meri and ischia unarmed. Third
maxilliped and first to fourth pereopods without strap-like
epipods. Appendices internae on second to fourth pereopods
simple, slender, without cincinnuli. Protopod of endopod
slightly tapering and rounded distally.
Biology: Living in profusion at vents or adjacent rocks.
Distribution: Mariana, Manus, North Fiji and Lau Back-Arc
Basins.

1: Habitus, lateral view; by J. Martin.

2: Dorsal view of carapace and bases

of antennae; by J. Martin.

4: In situ view of two specimens among gastropods

(Alviniconcha hessleri); Mariana Back-Arc Basin; by
3: Telson and uropods; by J. Martin. courtesy of R.R. Hessler.

References:
MARTIN J.W. & R.R. HESSLER (1990) Nat. Hist. Mus. Los Angeles Cty 417: 1-11.
Williams A.B. & P.A. Rona (1986) J. Crustac. Biol. 6: 446-462.

T. KOMAI & M. SEGONZAC Denisia 18 (2006): 423

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Mirocaris fortunata (MARTIN & CHRISTIANSEN, 1995)

ly fused mesially without trace of median separation; no distinct

Size: Females carapace length 3.3-8.7 mm, males carapace
length 3.8-7.2 mm.
Morphology: Rostrum triangular, terminating bluntly or suba-
cutely in dorsal view, flattened dorsoventrally, reaching to
slightly overreaching antennal spine; no dorsal or ventral teeth.
Carapace somewhat compressed laterally, with short transverse
(vertical) rows of minute to short setae on lateral parts; dorsal
surface rounded in males and non-ovigerous females, broadly
carinate in ovigerous females, general outline in lateral view
faintly sinuous to weakly convex; in ovigerous females, subme-
dian areas very shallowly depressed and ornamented with nu-
merous longitudinal striae; antennal spine acute; pterygostomi-
al angle weakly produced, terminating subacutely. Fourth ab-
dominal pleuron rounded posteriorly; fifth pleuron with suba-
cutely pointed posterolateral angle. Telson barely narrowed
posteriorly, bearing row of 7-9 dorsolateral spines arranged in
sinuous row; posterior margin broadly convex, occasionally
with shallow median emargination, bearing 12-19 spines in to-
tal; 1-3 spines at posterolateral angle shorter than mesial spines,
simple, while remaining mesial spines elongate, bearing minute
marginal setules. Eye-stalks rather large but degenerated, broad-
spine or tubercle on anterior surface. Chela of first pereopod
with fine row of long submarginal setae on outer surface along
cutting edges of fingers. Third to fifth pereopods moderately
slender to stout; dactyli armed with single row of accessory
spinules on ventral margin; meri unarmed; ischia with spines in
third, usually unarmed in fourth and fifth. Third maxilliped and
first to fourth pereopods with strap-like epipods. Appendices
internae on second to fourth pereopods rudimentary or absent.
Biology: Lives in close proximity to vent openings or in shim-
mering water from crevices with diffuse flow, among numerous
Bathymodiolus bivalves. Numerous ovigerous females in the
June-July samples. This species was found in the enteron of
Candelabrum phrygium (Cnidaria, Hydrozoa) which lives in the
same area.
Distribution: Known from hydrothermal vent sites along the
Mid-Atlantic Ridge between 38°N and 14°N: Menez Gwen,
Lucky Strike, Rainbow, Broken Spur, TAG, Snake Pit, Lo-
gatchev.

1: Body, lateral; by J.W. Martin & J.C. Christiansen.

2: Anterior part of carapace and

cephalic appendages, dorsal 3: Telson and left uro-
(setae partially omitted; right pod, dorsal (setae omit-
antenna removed); ted), scale bar 1 mm; 4: In situ aggregation on crevice of suphide edifice, Lucky Strike
scale bar 2 mm; by T. Komai. by T. Komai. site during Exomar cruise © Ifremer.

References:
KOMAI T. & M. SEGONZAC (2003) Cah. Biol. Mar. 44: 199-215.
MARTIN J.W. & J.C. CHRISTIANSEN (1995) Proc. Biol. Soc. Wash. 108(2): 220-227.
SEGONZAC M. & W. VERVOORT (1995) Bull. Mus. Natl. Hist. Nat., Paris, 4e sér. 17: 31-64.

T. KOMAI & M. SEGONZAC Denisia 18 (2006): 424

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Mirocaris indica KOMAI, MARTIN, ZALA, TSUCHIDA & HASHIMOTO, in press

terolateral angle shorter than mesial spines, simple, while re-

Size: Carapace length 9.2-13.8 mm (females), 4.6-10.2 mm
(males).
Color: In life, body and appendages generally white, but dorsal
surface of carapace and anterior two abdominal somites reddish;
eye yellowish.
Morphology: Rostrum triangular, terminating bluntly or suba-
cutely in dorsal view, flattened dorsoventrally, reaching to
slightly overreaching antennal spine; no dorsal or ventral teeth.
Carapace somewhat compressed laterally, with short transverse
(vertical) rows of minute to short setae on lateral parts; dorsal
surface rounded, general outline in lateral view slightly convex;
antennal spine acuminate; pterygostomial angle weakly pro-
duced, terminating subacutely. Fourth abdominal pleuron
rounded posteriorly; fifth pleuron with subacutely pointed pos-
terolateral angle. Telson barely narrowed posteriorly, bearing
row of 7-9 dorsolateral spines arranged in sinuous row; posteri-
or margin broadly convex, occasionally with shallow median
emargination, bearing 12-19 spines in total; 1-3 spines at pos-
maining mesial spines elongate, bearing minute marginal se-
tules. Eye-stalks rather large but degenerated, broadly fused; no
distinct spine or tubercle on anterior surface. Chela of first
pereopod without submarginal row of setae on outer surface
along cutting edges of fingers. Third to fifth pereopods moder-
ately slender to stout; dactyli armed with single row of accesso-
ry spinules on ventral margin; meri unarmed; ischia with spines
in third, usually unarmed in fourth and fifth. Third maxilliped
and first to fourth pereopods with strap-like epipods. Appen-
dices internae on second to fourth pereopods rudimentary or
absent.
Biology: The collection sites were an active vent field that in-
cluded black smoker complexes with dense patches of vent as-
sociated animals, including Rimicaris kairei, Bathymodiolus mus-
sel, Alviniconcha snail, Austinograea crab, Marianactis anemone.
Distribution: Central Indian Ridge, Kairei Field, Edmond Vent
Field.

2: Front, dorsal view.

1: Habitus holotype, lateral view; scale bar 5 mm.

1-3 from KOMAI et al. (2006) © Scientia Marina.

3: Telson and right uropod,

dorsal view (setae omitted).

References:
KOMAI. T., MARTIN J.W., ZALA K., TSUCHIDA S. & J. HASHIMOTO (in press) Scientia Marina 70(1).

T. KOMAI Denisia 18 (2006): 425

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Nautilocaris saintlaurentae KOMAI & SEGONZAC, 2005

distinct dorsolateral tooth. Chela of first pereopod with fine

Size: Largest male carapace length 5.6 mm, largest female cara-
pace length 7.8 mm. Maximal total length ca. 35 mm.
Color: Generally whitish; matured female gonads reddish, visi-
ble throughout cuticle.
Morphology: Rostrum carinate and dentate dorsally, reaching
distal margin of basal segment of antennular peduncle; ventral
surface unarmed. Carapace somewhat compressed laterally;
postrostral median carina low, blunt, restricted to anterior 0.15
of carapace; antennal tooth acuminate; pterygostomial angle
weakly produced anteriorly, extending as far as antennal spine,
terminating in sharp tooth. Third to fifth pleonal pleura den-
tate posteroventrally. Telson with 7-9 dorsolateral spines
arranged in slightly sinuous row; posterior margin convex, bear-
ing 12-19 spines in total, 1-3 spines at each posterolateral cor-
ner shorter than mesial spines, simple, while remaining mesial
spines elongate, bearing minute marginal setules. Eyes rather
large but degenerate, broadly fused mesially; anterior surface
smooth; no trace of pigment. Antennal scale broadly oval, with
row of long submarginal setae on outer surface along cutting
edges of fingers. Third to fifth pereopods moderately slender to
stout; each dactylus armed with single row of accessory spinules
on ventral margin; meri unarmed; ischia with spines in third,
usually unarmed in fourth and fifth. Third maxilliped to fourth
pereopods with strap-like, terminally hooked epipods, corre-
sponding to setobranchs above first to fifth pereopods; appen-
dices internae on second to fourth pereopods rudimentary.
Biology: This species lives around black smokers where the flu-
ids are percolating at temperatures between 5 and 33°C, among
molluscan beds. The similarity in the mouthparts and chela
morphology suggests that the shrimps ingest free bacteria, mol-
lusc pseudo-faeces, or other detritus, as most other species of
the family. Some ovigerous females were collected in May 2005
(American cruise PAR 5).
Distribution: North Fiji and Lau Back-Arc Basins.

1: Holotype female (carapace length 7.3

mm) from the North Fiji Basin; habitus,
lateral view (fourth and fifth pereopods
and second to fourth pleopods missing);
scale bar 2 mm; by T. Komai.

2: Anterior part of carapace 3: Telson, dorsal view

and cephalic appendages, (setae omitted);
dorsal view; scale bar 0.5 mm; scale bar 1 mm;
by T. Komai. by T. Komai. 4: Several individuals in situ on sulphide wall; cruise TUIM07.

References:
DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA (1994) Mar. Geol. 116: 227-242.
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.

T. KOMAI & M. SEGONZAC Denisia 18 (2006): 426

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Opaepele loihi WILLIAMS & DOBBS, 1995

ther side; posterior margin convex, with 1-3 small spines at

Size: Male carapace length 13.6 mm.
each lateral angle and row of long plumose setae. Eyestalks de-
Color: When collected, intensely orange, with orange to yel- generated, broadly fused mesially, cornea unfacetted, poorly or-
lowish deposit on abdomen (iron oxyhydroxide particles). ganized retinal pigment evident; anterior surface unarmed.
Chela of first pereopod without fine row of long submarginal se-
Morphology: Rostrum short, not reaching distal margin of first
tae on outer surface along cutting edges of fingers. Third to fifth
segment of antennular peduncle, directed forward or slightly
pereopods moderately slender; dactyli armed with two or more
downward, somewhat depressed dorsoventrally, terminating
rows of accessory spinules on ventral surface; meri and ischia
acutely, triangular in dorsal view; dorsal surface bluntly carinae
unarmed. No strap-like epipods on third maxilliped and pere-
in midline, unarmed or armed with minute denticles in adults,
opods. Appendices internae on second to fourth pereopods
armed with up to six very small teeth in juveniles; ventral sur-
slender, without coupling hooks.
face flat or slightly elevated in midline, unarmed. Carapace
somewhat compressd laterally, rounded dorsally; antennal spine Biology: Lives on the flank of small volcano, among numerous
acute; no conspicuous lobe mesial to antennal tooth; pterygos- vents which discharge fluids at 37°C (ambiant water, 4°C).
tomial angle somewhat produced in adults, distinctly exceeding The fluid is clear. At Loihi Seamount, no other animal live
antennal tooth, terminating in sharp tooth. Abdomen smooth there.
dorsally; pleura of third and fourth somites smooth or faintly
Distribution: So far known only from Hawaii, Loihi Seamount.
denticulate posteriorly, posterolateral angle of fourth pleuron
rounded or sharply pointed; fifth pleuron with marginal denti-
cles posteriorly and sharply pointed posteroventral angle. Tel-
son with 6-8 dorsolateral spines arranged in sinuous row on ei-

2: Female paratype, anterior re-

gion, dorsal view.

1: Female holotype, lateral view; scale bar 5 mm.

3: Female paratype,
telson and uropods,
dorsal view.
1-3: from WILLIAMS &
DOBBS (1995).

Reference:
WILLIAMS A.B. & F.C. DOBBS (1995) Proc. Biol. Soc. Wash. 108(2): 228-237.

T. KOMAI Denisia 18 (2006): 427

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Rimicaris exoculata WILLIAMS & RONA, 1986

markedly shorter toward posterior; dactyli armed with several

Size: Carapace length 22.0 mm; total length up to 55 mm.
Morphology: Rostrum strongly reduced to very low, broadly
rounded lobe. Carapace with branchial region very strongly in-
flated, distinctly wider than abdomen; dorsal surface rounded;
antennal spine modified as broadly rounded lobe; pterygosto-
mial angle somewhat produced anteriorly, broadly rounded; sur-
face with scattered puntae each bearing tuft of short setae.
Third abdominal pleuron with posterolateral margin forming
light angle. Fourth and fifth pleura with posterolateral angles
subacutely or blunly pointed. Telson barely narrowed posterior-
ly, bearing row of 6-8 dorsolateral spines arranged in sinuous
row on either side; posterior margin broadly convex, with 1-3
spines at each lateral angle and with row of numerous plumose
setae. Eye-stalks greatly reduced and modified into broad ocu-
lar plate. Chela of first pereopod always slender, without fine
row of long submarginal setae on outer surface along cutting
edges of fingers. Third to fifth pereopods stout, becoming
rows of accessory spinules on ventral surface; propodi of third
and fourth pereopods with covering of numerous spinules on
ventral surfaces; meri and ischia unarmed. No strap-like
epipods on third maxilliped to fourth pereopods. Appendices
internae on second and fourth pereopods simple, slender, with-
out coupling hooks, those of fourth and fifth pleopods normal-
ly developed, with coupling hooks.
Biology: Shrimp highly mobile occurring in swarms reaching
densities of about 2500 ind./m2 on active chimney walls where
temperatures range from 10-30°C. Primary consumer, eating
bacteria “cultivated” on the mouthparts, and on the mineral
particles ingested. Very few ovigerous females collected, be-
tween August and November.
Distribution: Mid-Atlantic Ridge: TAG, Snake Pit, Rainbow,
very few at Lucky Strike.

1: Specimen taken onboard, dorsal view; 2: In situ aggregation of adults and juveniles or subadults (red cephalothorax)
cruise Exomar by P. Briand © Ifremer. specimens; TAG; cruise Exomar © Ifremer.

References:
MARTIN J.W. & R.R. HESSLER (1990) Nat. Hist. Mus. Los Angeles C. 417: 1-11.
SEGONZAC M., SAINT LAURENT M. DE & B. CASANOVA (1993) Cah. Biol. Mar. 34: 535-571.
WILLIAMS A.B. & P.A. RONA (1986) J. Crustac. Biol. 6: 446-462.
ZBINDEN M. & M.-A. CAMBON-BONAVITA (2003) FEMS Microbiol. Ecol. 46: 23-30.

T. KOMAI & M. SEGONZAC Denisia 18 (2006): 428

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Rimicaris kairei WATABE & HASHIMOTO, 2002

Size: Up to 21.6 mm in carapace length. Remark: Comparing the sequence of gene coding for mito-
chondrial cytochrome oxydase subunit one, VAN DOVER et al.
Morphology: Integument completely spineless, almost mem-
(2001) found a small divergence (0.9%) from R. exoculata.
branous on branchiostegite. Carapace longitudinally oval in
dorsal view, sparsely pitted by shallow punctuations without Biology: Swarming with extremely high density on the surface
any setae. Rostrum and eyestalks completely absent in adult. of active chimneys and diffuse vents. Like R. exoculata, symbio-
Antennal and antennular flagella strongly developed. Anten- sis with chemoautotrophic bacteria.
nal scale enlarged, completely filling anterior margin of cara-
Distribution: Central Indian Ridge: Kairei and Edmond Vent
pace. First and second pereopods chelate, usually hidden be-
Fields, near the Rodriguez Triple Junction.
neath branchiostegite. Third to fifth pereopods well developed,
stout.

1: Two specimens taken onboard; scale bar 5 mm; 2: In situ aggregation of adult specimens at the Kairei
by courtesy of J. Hashimoto. Vent Field; below right, sea anemone Marianactis cf.
bythios, and mytilid bivalves Bathymodiolus sp.; cruise
ROV Kaiko © JAMSTEC.

References:
HASHIMOTO J., OHTA S., GAMO T., CHIBA H., YAMAGUCHI T., TSUCHIDA S., OKUDAIRA T., WATABE H., YAMANAKA T. & M. KITAZAWA (2001) Zool. Sci. 18(5): 717-721.
VAN DOVER C.L., HUMPHRIS S.E., FORNARI D., CAVANAUGH C.M., COLLIER R., GOFFREDI S.K., HASHIMOTO J., LILLEY M.D., REYSENBACH A.L., SHANK T.M., VON DAMM
K.L., BANTA A., GALLANT R.M., GÖTZ D., GREEN D., HALL J., HARMER T.L., HURTADO L.A., JOHNSON P., MCKINESS Z.P., MEREDITH C., OLSON E., PAN I.L.,
TURNIPSEED M., WON Y., YOUNG III C.R. & R.C. VRIJENHOEK (2001) Science 294: 818-823.
WATABE H. & J. HASHIMOTO (2002) Zool. Sci. 19(10): 1167-1174.

J. HASHIMOTO Denisia 18 (2006): 429

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

Arthropoda, Crustacea, Decapoda, Caridea, Alvinocarididae

Shinkaicaris leurokolos (KIKUCHI & HASHIMOTO, 2000)

teeth including posteroventral tooth. Telson elongate subrec-

Size: An ovigerous female carapace length 12.5 mm.
tangular, with 5-7 dorsolateral spines arranged in a sinuous row;
Color: Ivory to dull ivory white. posterior margin convex, with two pairs of lateral spines (mesial
pair longer than lateral) and 22-24 long plumose setae. Eyes on
Morphology: Integument of body thin, but not membranous,
basally separated, movable stalks broadly fused mesially; divi-
surface shining. Rostrum compressed laterally, reaching distal
sion of corneal region and stalk unclear; corneal region, un-
margin of first segment to midlength of second segment of an-
faceted, with irregular, scattered pigment-like masses within
tennular peduncle; dorsal margin armed with 7-10 teeth slight-
stalk; anteromedial face flat, sloping to anterolateral face, with-
ly diminishing in size anteriorly, including 6-9 on rostrum prop-
out spiniform tubercle on anterior surface. Dorsolateral flagel-
er and 1-3 on carapace. Carapace somewhat compressed later-
lum longer than carapace; ventromesial flagellum somewhat
ally; postrostral median ridge relatively low, rather blunt; an-
longer. Antennal flagellum longer than body. Uropod with ra-
tennal tooth sharp, distinctly buttressed, margin inferior to an-
mi subequal in length, exopod with small movable spine mesial
tennal tooth forming small rounded lobe. Strong median ster-
to smaller distolateral tooth and sinuous diaeresis.
nal spine between coxae of fifth pereopods. Abdomen smooth
dorsally; pleural margin of anterior two somites broadly round- Biology: This species occurs in a thermally influenced area.
ed, that of third somite broadly rounded or with one small pos-
Distribution: Known only Okinawa Trough: Minami-Ensei
teroventral tooth; pleuron of fourth somite with acute postero-
Knoll, 705 m.
lateral tooth and occasionally with additional 1-3 acute teeth
on posterior margin; pleuron of fifth somite with 2-4 posterior

1: Holotype female in lateral view.

2: Anterior region,
dorsal view.

3: Telson and uropods,

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

References:
KIKUCHI T. & J. HASHIMOTO (2000) Species Diversity 5: 135-148.
KOMAI T. & M. SEGONZAC (2005) J. Nat. Hist. 39(15): 1111-1175.

T. KOMAI & M. SEGONZAC Denisia 18 (2006): 430

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Decapoda, Caridea, Hippolytidae
Lebbeus carinatus ZARENKOV, 1976
Size: Maximum carapace length 17 mm.
Morphology: Rostrum short, not reaching distal margin of first
segment of antennular peduncle; dorsal margin armed with 4-6
teeth including 2-4 on rostrum proper and two on carapace pos-
terior to orbital margin, posteriormost tooth arising about 0.15
length of carapace; ventral blade poorly developed; ventral
margin with 1-3 small teeth subdistally. Carapace with small
supraorbital spine; broad V-shaped notch just below supraor-
bital spine; orbital margin depressed, forming shallow concavi-
ty accommodating eye-stalk; suborbital lobe acutely triangular,
somewhat upturned, slightly exceeding antennal spine; antero-
lateral margin between antennal spine and pterygostomial an-
gle strongly sinuous; pterygostomial spine small; dorsal surface
strongly convex in spawning females, not inflated in males. Ab-
domen rounded dorsally; fourth pleuron broadly rounded, fifth
pleuron with strong posteroventral tooth. Telson with 4-6 dor-
solateral spines on either side; posterior margin slightly convex
or truncate, with two pairs of spines at lateral angle and one or
two pairs of plumose setae. Eye small, corneal diameter
0.09–1.00 of carapace length; cornea semispherical, darkly pig-
mented and clearly faceted. Antennule with stylocerite not
reaching distal margin of first segment of peduncle; first pedun-
cular segment armed with two or three spines on dorsodistal
margin laterally; second and third segments each with one dor-
sodistal spine; flagella sexually dimorphic, those of males elon-
gate, particularly ventral flagellum of male exceeding 1.5
T. KOMAI
length of carapace. Antennal scale moderately broad; distolat-
eral tooth falling short of or reaching rounded distal lamella.
Third maxilliped overreaching antennal scale by half length of
ultimate segment. First pereopod chelate. Second pereopod al-
so chelate, carpus divided into seven articles. Third to fifth
pereopods slender, third overreaching antennal scale by full
length of propodus; merus of third and fourth pereopods with
four or five spines on lateral surface; that of fifth unarmed or
armed with one spine; dactyli short, 0.10-0.13 length of propo-
di, each with four or five accessory spinules on ventral margin.
Third maxilliped and first and second pereopods each with
strap-like epipod.
Remark: This species is easily distinguished from L. washingto-
nianus of KIKUCHI & OHTA (1995), the other congeneric
species occurring at hydrothermal vents, by the absence of a
strap-like epipod on the third pereopod and the more slender,
elongate third to fifth pereopods. Another different species was
described by SAINT LAURENT (1984) under the same name. It
differs mainly from the other caridean species associated with
hydrothermal vents by the well developed eyes and the carpus
of the second pereopod divided in seven articles.
Biology: This species live among the tubeworm Riftia pachypti-
la and mussel beds of Bathymodiolus spp.
Distribution: Known with certainty only from the East Pacific
Rise: 13°N.
Denisia 18 (2006): 431–432
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

2: Cephalothorax, lateral view; from

East Pacific Rise 13°N, cruise Hope 99;
by P. Briand © Ifremer.

3: Specimen in situ, among tubeworms

(Riftia pachyptila), from East Pacific Rise
1: Body, lateral view; from N. ZARENKOV (1976). 13°N, cruise Hope 99 © Ifremer.

References:
KIKUCHI T. & S. OHTA (1995) J. Crustac. Biol. 15: 219-240.
SAINT LAURENT M. DE (1984) C. R. Acad. Sc., Paris 299(9), sér. III: 355-360.
ZARENKOV N.A. (1976) J. Mar. Biol. 5: 277-285.

432
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Decapoda, Astacidea, Nephropidae

Thymopides laurentae SEGONZAC & MACPHERSON, 2004

Size: Carapace length 45 mm; total length 97 mm. Biology: Lives on the slope of the Snake Pit vent area, near the
black smoker edifice Beehive, among galatheid crab Munidopsis
Morphology: Carapace smooth, rostrum slender, nearly 0.3 of
exuta, and chaetopterid polychaetes.
carapace length, overreaching antennular and antennal pedun-
cles. Abdomen without longitudinal median carina and palm of Distribution: Mid-Atlantic Ridge: near the Snake Pit hy-
first chela distinctly longer than wide. Telson 1.7 times longer drothermal area.
than wide and longer than abdominal somite 6. Eyes small, un-
pigmented, cornea as wide as ocular peduncle.

1–4: Scale bar 1 cm; drawings by M. de Saint Laurent.

1: Body, lateral view. 2: Carapace, anterior part and 3: Fifth and sixth abdominal somites 4: Carapace dorsal
cephalic appendages, lateral view. and tail fan with telson, dorsal view. view.

6: In situ of one lobster (probably T. laurentae)

taken from the submersible at Snake Pit field;
5: Photograph taken on board, after sampling © Ifremer cruise Hydrosnake © Ifremer

References:
MÈVEL C., AUZENDE J.-M., CANNAT M., DONVAL J.-P., DUBOIS J., FOUQUET Y., GENTE P., GRIMAUD D., KARSON J.A., SEGONZAC M. & M. STIEVENARD (1989) C. R. Acad.
Sci., Paris 308(II): 545-552.
SEGONZAC M. (1992) C. R. Acad. Sci., Paris 314(III): 593-600.
SEGONZAC M. & E. MACPHERSON (2004) Cah. Biol. Mar. 44: 361-367.

M. SEGONZAC & E. MACPHERSON Denisia 18 (2006): 433

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda: Decapoda, Anomura
Worldwide, there are over 2500 species of anomouran
crabs, which comprise ca 5% of all crustacean species. The In-
fraorder Anomura represents a paraphyletic group that includes
the superfamilies Lomisoidea, Hippoidea, and the much more
diverse Galatheoidea and Paguroidea. Species of these taxa are
commonly found living from the intertidal zone to the abyssal
plain >2000 m, including one terrestrial representative. Mor-
phologically they have little in common, some are like crabs
(e.g. Lithodidae) and others are like hermit crabs (e.g. Paguri-
dae). They only share one character: the small fifth pereiopod.
Molecular studies have shown that Galatheoidea and Paguroi-
dea are more related to each other than to Hippoidea, although
more work is needed to completely resolve these relationships.
Munidopsis sp. (probably M. lauensis); cruise TUIM 07;
by C.R. Fisher.
References:
MARTIN J.W. & G.E. DAVIS (2001) Nat. Hist. Mus. Los Angeles Cty, Sci. Ser. 39: 1-124.
MARTIN J.W. & T.A. HANEY (2005) Zool. J. Linn. Soc. 145: 445-522.
MORRISON C.L., HARVEY A.W., LAVERY S., TIEU K., HUANG Y. & C.W. CUNNINGHAM (2002) Proc. R. Soc. Lond., B 269: 345-350.
PERÉZ-LOSADA M., JARA C.G., BOND-BUCKUP G., PORTER M.L. & K.A. CRANDALL (2002) J. Crustac. Biol. 22: 661-669.
E. MACPHERSON & K. BABA
The vent fauna contains representatives of the superfami-
lies Galatheoidea and Paguroidea, including species of four fa-
milies and a recent new family (Kiwaidae). Despite their ecolo-
gical importance and high diversity, many aspects of their sys-
tematics and distribution are still poorly known.
The anomurans exhibit a considerable diversity of repro-
duction modes, life cycles and capacities for dispersal. The vast
majority of species have relatively small pelagic eggs, with the
exception of some representatives of the families Galatheidae
and Chirostylidae, and a pelagic larval phase, which enhances
their capacity for dispersal. There is evidence for prolonged
brooding periods. Usually they produce only a few large eggs,
probably related to an abbreviated or direct larval develop-
ment. Most species are not restricted to hydrothermal vents
and cold seeps. They usually occur in low densities, although
there are some interesting exceptions (e.g. Munidopsis lentigo,
Shinkaia crosnieri). The abundance of anomurans in vent and
seep environments probably reflects the benefit derived from
the organic matter produced by the chemosynthetic communi-
ty inhabiting these zones. In general, their abundance increased
in the vicinity of active hydrothermal sites. In the center of
hydrothermal activity, however, abundance decreases.
Information on the geographic distribution and taxonomy
of the different species is still limited and incomplete. Most spe-
cies have a broad geographic distribution, as has been shown for
a number of deep-sea taxa. There appears to be some inter-
change among the different communities separated by long dis-
tances, associated with high dispersal capabilities, although a
high degree of local endemism can also exist. Unfortunately,
mechanisms for colonization by vent organisms remain largely
unknown for most of the mid-ocean ridge systems. Therefore,
additional studies on molecular data and larval dispersal pro-
cesses are desirable.
Denisia 18 (2006): 434
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Decapoda, Anomura, Parapaguridae

Paragiopagurus ventilatus LEMAITRE, 2004

pereopods) with bifid spine. Telson asymmetrical; posterior

Size: Shield length 5.8 mm max. (male); 4.9 mm max. (oviger-
ous female).
Color: In life, shield and cephalic appendages cream with some
faded pink areas. Chelipeds and ambulatory legs red or pink.
Morphology: Gills biserial. Ocular acicles subtriangular, termi-
nating in strong, simple spine (rarely bifid) with transverse stri-
ae. Outer faces of basis and ischium of third maxillipeds, che-
lipeds, and second and third pereopods, with plumose, bacte-
riophore setae. Chelipeds markedly dissimilar; proportions and
armature of right strongly affected by size and sex, with that of
males considerably longer and more elongate than in females.
Dactyls of second and third pereopods with irregular rows of
spines on ventromesial margins. Propodal rasp of fourth pereo-
pod with one row of scales. Anterior lobe of sternite XII (third
margin separated by broad, shallow cleft, into rounded projec-
tions armed with few short, corneous spines often ventrally
curved. Males lacking first pleopods; with unpaired rudimenta-
ry left pleopod.
Biology: Specimens collected by beam-trawl among other
species newly described as mytilid and lucinid bivalves
(BOUCHET & COSEL 2004), and numerous grapsid crabs (NG et
al. 2000). At this area, diffuse gas seepage from the bottom.
The specimens live in gastropod shells of the buccinid Siphona-
lia. Probably not part of the “obligate” vent fauna. Ovigerous fe-
males found in May 2001.
Distribution: Okinawa Arc: Northeastern Taiwan, Tashi fish-
ing grounds.

1: Male in vivo; by courtesy of T. Yam Chan. 2: Female in vivo; by courtesy of T. Yam Chan.

References:
BOUCHET P. & R. VON COSEL (2004) Zool. Stud. 43: 704-711.
LEMAITRE R. (2004) Cah. Biol. Mar. 45: 325-334.
NG N.K., HUANG J.F. & P.H. HO (2000) Nat. Taiwan Mus., Spec. Publ. Ser. 10: 191-199.

R. LEMAITRE Denisia 18 (2006): 435

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

Arthropoda, Crustacea, Decapoda, Anomura, Lithodidae

Paralomis hirtella DE SAINT LAURENT & MACPHERSON, 1997

along ventral margin. Scattered tuft of setae on posterior sur-

Size: Up to 59 mm carapace length.
face, dorsal and ventral margins of articles.
Morphology: Carapace more or less hexagonal. Dorsal surface
Biology: Lives in few numbers around active chimneys and on
smooth, with few scattered small granules. Numerous tuft of se-
mussel beds.
tae of different length scattered on carapace surface. Rostrum
with basal spine and two small dorsal spines. Chelipeds bearing Distribution: North Fiji Back-Arc Basin: White Lady vent
thick spines and acute granules. Walking legs moderately long, field; Lau Back-Arc Basin: Hine Hina vent field.
with some thick spines on dorsal border of merus, carpus and
propodus, dactylus slightly curved, with corneous spinelets

1: Holotype male, 32 x 35 mm, habi- 2: Holotype, abdomen; scale bar 10 mm; by 3: Other specimen 44 x 45 mm, abdomen
tus, dorsal view; scale 10 mm; by M. M. de Saint Laurent & E. Macpherson. of ovigerous female; scale bar 10 mm; by
de Saint Laurent & E. Macpherson. M. de Saint Laurent & E. Macpherson.

4: Specimen collected at Lau Back-Arc Basin: Hine Hina site; 5: Two specimens in situ on gastropod bed of Ifremeria
by P. Briand © Ifremer. nautilei at North Fiji Back-Arc Basin: White Lady site;
cruise Starmer 2 © Ifremer.

References:
DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA (1994) Mar. Geol. 116: 227-242.
SAINT LAURENT M. DE & E. MACPHERSON (1997) Zoosystema 19: 721-727.

E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 436

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

Arthropoda, Crustacea, Decapoda, Anomura, Lithodidae

Paralomis verrilli (BENEDICT, 1895)

vergent dorsal spines, upwardly directed, extending well past

Size: Up to 110 mm carapace length.
Morphology: Carapace more or less hexagonal, as long as wide.
Dorsal carapace surface covered with small granules of various
size. Granules usually with several setae. Regions well-defined.
Gastric region rather more prominent than other regions, with
a thick spine on apex. Cardiac region clearly less prominent
than branchial regions, with four thick granules in square pat-
tern. Each branchial region with one median spiniform granule
and two smaller thick granules near intestinal region. Basal
spine of rostrum more or less horizontal, slightly overreaching
end of cornea, with spiniform tubercles on ventral side; two di-
end of cornea. External orbital spine slightly shorter than eyes.
Anterolateral spine slightly shorter than external orbital. Che-
lipeds bearing thick spines on mesial border of merus, carpus
and hand. Walking legs moderately long, with well-developed
spines along dorsal and ventral margins of meri and propodi,
dactyli as long or longer than propodi.
Distribution: North Pacific, from Japan, Bering Sea to Califor-
nia, 850-2379 m. Collected on vent sites of Juan de Fuca Ridge:
Axial Seamount.

1: Specimen among siboglinid tubeworms and microbial mats on side of sulphide edifice; from R/V Thomas G. Thompson, cruise
TN149 (Zooarium vent field, Explorer Ridge; 31 July 2002) © NOAA.

References:
DAWSON E.W. (1989) N.Z.O.I Misc. Publ. 101: 1-338.
IKEDA H. (1998) The Deep-sea Crabs of Sagami Bay, Imamoto Printing: 1-180.
MACPHERSON E. (2001) Zoosystema 23: 797-805.
ZAKLAN S.D. (2002) Alsk. Sea Grant Coll. Progr.: 751-845.

E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 437

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munida magniantennulata BABA & TÜRKAY, 1992

Distomesial spine of first antennal article very small, never

Size: Up to 12.8 mm carapace length.
reaching midlength of second article. Merus of third maxilliped
Morphology: Dorsal carapace surface smooth, with epigastric with only one spine on flexor margin. Chelipeds moderately
spines. Lateral branchial margins with five small spines. First massive, about three times carapace length. Walking legs slen-
lateral spine of carapace near anterolateral angle, second spine der.
clearly smaller than first. Second abdominal segment with
Biology: No data. Collected by grab TV from an area close to
transverse row of small spines, other segments unarmed.
active sites.
Corneae small, width equal to or less than distance between
than distance between sinus formed by supraocular spines and Distribution: Lau Back-Arc Basin: Hine Hina area.
rostrum. Basal segment of antennular peduncle unusually large.

1: Holotype male, carapace

length 7.2 mm; dorsal view;
scale bar 1 mm;
from BABA & TÜRKAY (1992).

References:
BABA K. (1994) Mem. Qld. Mus. 35: 1-21.
BABA K. (2005) Galathea Report 20: 1-317.
BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.
BABA K. & M. TÜRKAY (1992) Senckenb. Marit. 22: 203-210.

E. MACPHERSON & K. BABA Denisia 18 (2006): 438

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis acutispina BENEDICT, 1902

Size: Up to 8.0 mm carapace length.
Morphology: Carapace with numerous pointed tubercles. Ros-
trum narrow triangular, horizontal and dorsally carinated. Front
margin convex, antennal spine small. Abdomen unarmed.
Cornea well exposed, visible in dorsal view, eye spine mesial
and small. Fixed finger of chelipeds without denticulate carina
on distolateral margin. Walking legs not reaching end of che-
lipeds. Propodi of walking legs of uniform width, with distinct
spines. Epipods absent from pereopods.
Biology: Occurs in both hydrothermal and cold seep areas.
Distribution: Mid-Atlantic Ridge: Lost City vent field, and
Eastern Mediterranean Sea, Kazan mud volcano, cold seep,
2030 m. Known as well off Western North Africa and the
Azores Islands, between 698 and 845 m, and from the Mediter-
ranean Sea (W Tyrhenian Sea, Sardinia) between 374 and
1036 m.

2: Carapace and abdomen, lateral view; scale bar 5 mm;

male; from FROGLIA et al. (2002).

1: Carapace, dorsal view; male, carapace

length 8.0 mm; from FROGLIA et al. (2002).

3: Telson; scale bar 5 mm; from FROGLIA et 4: Specimen from Lost City; cruise Exomar; by P. Briand
al. (2002). © Ifremer.

References:
FROGLIA C., MURA, M. & A. BONFITTO (2002) Crustac. Int. J. Crustac. Res. 75: 375-382.
GEBRUK A.V., GALKIN S.V., KRYLOVA E.M., VERESHCHAKA A.L. & G.M. VINOGRADOV (2002) InterRidge News 11(2): 18-19.
MACPHERSON E. & M. SEGONZAC (2005) Zootaxa 1095: 1-60.
OLU-LE ROY K., SIBUET M., FIALA-MÉDIONI A., GOFAS S., SALAS C., MARIOTTI A., FOUCHER J.-P. & J. WOODSIDE (2004) Deep-Sea Res. I 51: 1915-1936.

E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 439

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis alvisca WILLIAMS, 1988

First walking leg reaching nearly tip of cheliped, dorsal crest of

Size: Up to 27.9 mm carapace length.
merus and carpus with row of spines along dorsal border; dacty-
Morphology: Dorsal carapace surface without spines, covered lus smoothly narrowed distally, flexor margin slightly curving,
by scale-like rugosities. Rostrum unarmed, broad at base, distal- bearing low spines. Epipods absent from pereopods.
ly narrowed and upturned, weakly carinated dorsally. Front
Biology: Living around hydrothermal vent sites.
margin oblique, small antennal spine present. Abdomen un-
armed. Eyes not movable, eyespine well developed, cornea rel- Distribution: Explorer Ridge: Magic Mountain; Juan de Fuca
atively large, clearly wider than eyespine. Chelipeds with sev- Ridge; Guaymas Basin.
eral spines on merus and carpus, palm spineless, fixed finger of
chelipeds without denticulate carina on distolateral margin.

1: Carapace and abdomen, dorsal

view; holotype carapace length
23.7 mm (including rostrum);
scale bar 5 mm; by A.B. Williams.

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

References:
BABA K. (2005) Galathea Report 20: 1-317.
KHODKINA I.V. (1991) Zool. Zh. 70: 71-76.
WILLIAMS A.B. (1988) Fish. Bull. 86: 263-287.

E. MACPHERSON & K. BABA Denisia 18 (2006): 440

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis diomedeae (FAXON, 1893)

slender, with numerous granules, subcylindrical. First walking

Size: Up to 30 mm (with rostrum).
leg overreaching chelipeds. Dactyli compressed laterally, flexor
Morphology: Carapace covered with tubercles, dorsally armed border moderately curving, bearing proximally diminishing low
with two well-developed epigastric, one extremely strong later- spines. Epipods present on chelipeds and first and second walk-
ally compressed mesogastric, and one moderately large cardiac ing legs.
spine, with two lateral spines. Dorsal carapace surface smooth,
Distribution: East Pacific Rise. The species has been sampled
without spines. Rostral spine upturned distally. Second to
in Eastern Pacific, from the Gulf of California to Chile, be-
fourth abdominal tergites each with prominent median spine.
tween 768 and 2026 m.
Eyes movable and spineless, corneae subglobular. Chelipeds
covered with granules, nearly spineless. Walking legs long,

1: Male, carapace length ca. 30 mm

(with rostrum), habitus, dorsal view;
from FAXON (1895).

References:
BABA K. (2005) Galathea Report 20: 1-317.
FAXON W. (1895) Mem. Mus. Comp. Zool. Harvard Coll. 18: 1-292.
HAIG J. & M. WICKSTEN (1975) Bull. South Calif. Acad. Sci. 74: 100-105.
KHODKINA I.V. (1991) Zool. Zh. 70: 71-76.

E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 441

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis exuta MACPHERSON & SEGONZAC, 2005

Size: Up to 39.5 mm carapace length. Biology: This species occurs in both abyssal (Gulf of Biscay,
2860 m) and hydrothermal vent environments (Mid-Atlantic
Morphology: Carapace with pair of epigastric spines, some-
Ridge: Snake Pit, Elan). It has been collected by baited trap to-
times 2-3 additional small spines, covered by scale-like rugosi-
gether with a nephropid lobster Thymopides laurentae SEGON-
ties. Rostrum broad at base, distally narrowed and upturned,
ZAC & MACPHERSON, and a macrurid fish Coryphaenoides arma-
weakly carinated dorsally. Front margin oblique, antennal spine
tus (HECTOR), at the base of the active edifice, on sulphide
absent. Abdomen unarmed. Eyes not movable, cornea relative-
rocks, among scattered sea anemones, chaetopterid tubeworms
ly large, clearly wider than eyespine. Chelipeds with several
and alvinocaridid shrimps. The species has also been collected
spines on merus and carpus, palm spineless, fixed finger moder-
at the base of inactive chimney complex, among dead mytilid
ately ridged along distal third of lateral margin. First walking
bivalves. An additional specimen (unfortunately lost) was col-
leg slightly exceeding cheliped, dorsal crest of merus and carpus
lected during the cruise BRIDGE 1993 at the Mid-Atlantic
with row of spines along dorsal border; dactylus smoothly nar-
Ridge hydrothermal vent site Broken Spur site (dive Alvin
rowed distally, flexor margin slightly curving, bearing low
2625, 27.06.1993, E. Southward, personal communication).
spines. Epipods on chelipeds, not on walking legs. The species
was identified as M. subsquamosa by SAINT LAURENT (1985) Distribution: Mid-Atlantic Ridge: Snake Pit, TAG, probably
and M. crassa by SEGONZAC (1992). Broken Spur, and Logatchev; Gulf of Biscay.

2: Carapace, lateral view; holo-

type, carapace length 34.7
mm; scale bar 5 mm; from
MACPHERSON & SEGONZAC (2005).

1: Carapace, dorsal view; holotype, carapace length

34.7 mm; scale bar 5 mm; from MACPHERSON & SEGON- 3: Specimen from Snake Pit; cruise Hydrosnake; by P. Briand
ZAC (2005). © Ifremer.

References:
MACPHERSON E. & M. SEGONZAC (2005) Zootaxa 1095: 1-60.
SAINT LAURENT M. DE (1985) in Ifremer Brest (Ed.) Peuplements Profonds du Golfe de Gascogne: 469-478.
SEGONZAC M. (1992) C. R. Acad. Sci. Paris III 314: 593-600.

E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 442

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis lauensis BABA & DE SAINT LAURENT, 1992

overreaching chelipeds. Fixed finger of chelipeds without den-

Size: Up to 12.4 mm carapace length.
ticulate carina on distolateral margin. Chelipeds and walking
Color: Generally white, but some rare specimens are blackish, legs, with few small spines. Dactylus of walking legs with spines
due to a manganese deposit. along entire flexor border.
Morphology: Carapace without dorsal spines or tubercles. Ros- Biology: On active hydrothermal sites, often in high density.
trum spiniform, horizontal and unarmed. Eye with small mesial
Distribution: North Fiji Back-Arc Basin: White Lady vent
spine, cornea large. Abdomen unarmed. Sixth abdominal seg-
field; Lau Back-Arc Basin: Valu Fa Ridge, Hine Hina vent
ment having posteromedian margin transverse, exceeded by
fields.
lateral lobes. Epipods absent from pereopods. First walking legs

2: Carapace, lateral view; holotype male, carapace length 10.8 mm; scale bar 1
mm; by K. Baba & M. de Saint Laurent.

1: Carapace and abdomen, dorsal view; holo-

type male, carapace length 10.8 mm; scale 3: Munidopsis sp. (probably M. lauensis); cruise TUIM 07;
bar 1 mm; by K. Baba & M. de Saint Laurent. by courtesy of C.R. Fisher.

References:
BABA K. (2005) Galathea Report 20: 1-317.
BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.
DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA (1994) Mar. Geol. 116: 227-242.

E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 443

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis lentigo WILLIAMS & VAN DOVER, 1983

tened light brown spot on ventral surface of chela. First walk-

Size: Up to 12.8 mm carapace length.
ing leg reaching nearly tip of cheliped, dorsal crest of merus
Morphology: Dorsal carapace surface smooth, without spines. with row of small spines; dactylus short, flexor margin bearing
Rostrum unarmed, broad triangular and horizontal, not cari- low spines. Epipods absent from pereopods.
nated dorsally. Front margin transverse, antennal spine present.
Biology: Lives around the hydrothermal vents.
Abdomen unarmed. Flattened eyes, depressed cornea, large me-
dian eyespine present. Chelipeds with several spines on merus Distribution: East Pacific Rise: 21°N; south of Baja California.
and carpus, and palm, fixed finger of chelipeds without dentic-
ulate carina on distolateral margin. Lenticular-shapped, flat-

2: Carapace, lateral view; female

paratype, carapace length 11.9 mm;
from WILLIAMS & VAN DOVER (1983).

1: Habitus, dorsal view; scale bar 1 mm; female paratype, carapace length 11.9 mm;
from WILLIAMS & VAN DOVER (1983).

References:
BABA K. (2005) Galathea Report 20: 1-317.
WILLIAMS A.B. & C. VAN DOVER (1983) Proc. Biol. Soc. Wash. 96: 481-488.

E. MACPHERSON & K. BABA Denisia 18 (2006): 444

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis marianica WILLIAMS & BABA, 1989

chelipeds without denticulate carina on distolateral margin.

Size: Up to 38.0 mm carapace length.
Morphology: Dorsal carapace surface without spines, covered
by scale-like rugosities. Rostrum unarmed, broad triangular and
horizontal, weakly carinated dorsally. Front margin oblique,
small antennal spine present. Abdomen unarmed. Eyes mov-
able, large mesial eyespine small lateral eyespine, cornea rela-
tively large, wider than mesial eyespine. Chelipeds with sever-
al spines on merus and carpus, palm spineless, fixed finger of
First walking leg reaching nearly tip of cheliped, dorsal crest of
merus and carpus with row of spines along dorsal border; dacty-
lus smoothly narrowed distally, flexor margin nearly stright,
bearing low spines. Epipods present on chelipeds.
Biology: Lives at both vent and non vent areas.
Distribution: Western Pacific Ocean, Mariana Back-Arc Basin.

1-4: Female holotype, carapace length 38.0 mm; scale bar 10 mm; by A.B. Williams & K. Baba.

3: Anterior part of carapace, lateral view.

2: Rostrum and anterior part of 4: Telson, no scale.

1: Carapace and abdomen, dorsal view. carapace, dorsal view.

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

E. MACPHERSON & K. BABA Denisia 18 (2006): 445

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis sonne BABA, 1995

tubercular processes and blunt short spines, longer than second

Size: Up to 12.8 mm carapace length.
walking legs; fixed finger without denticulate carina on disto-
Morphology: Dorsal carapace surface covered with numerous lateral margin. Walking legs diminishing in length posteriorly;
tubercle-like setiferous processes, gastric region with pair of epi- dactyli rather straight in first and second legs, more curving on
gastric processes. Rostrum nearly triangular, nearly horizontal, third; setae along distal two-thirds of flexor border. Epipods
lateral margins serrate. Front margin with antennal acute present on chelipeds, first and second walking legs.
process. Anterolateral angle produced. Abdomen unarmed;
Biology: Lives around hydrothermal vents.
sixth abdominal segment with weak posterolateral lobe, poste-
rior margin not produced. Eyes immovable, cornea narrowed Distribution: North Fiji Back-Arc Basin.
distally. Antennular basal segment cristate dorsolaterally, with
row of several spines. Chelipeds subcylindrical, covered with

1-3: Ovigerous female holotype, carapace length 10.5 mm; scale bar 1 mm; by K. Baba.

2: Carapace and abdomen, lateral view.

3: Telson.

1: Carapace and abdomen, dorsal view.

References:
BABA K. (1995) Crustac. Res. 24: 188-193.
BABA K. (2005) Galathea Report 20: 1-317.

E. MACPHERSON & K. BABA Denisia 18 (2006): 446

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis starmer BABA & DE SAINT LAURENT, 1992

chelipeds without denticulate carina on distolateral margin.

Size: Up to 33.4 mm carapace length.
Chelipeds and walking legs, with few small spines. Dactylus of
Morphology: Carapace with two epigastric spines, dorsal sur- walking legs with spines along entire flexor border.
face smooth. Rostrum spiniform, upwards directed, unarmed.
Biology: Lives on active hydrothermal vents.
Eye with small mesial spine, cornea large. Abdomen unarmed.
Sixth abdominal segment having posteromedian margin trans- Distribution: North Fiji Back-Arc Basin: site White Lady.
verse, exceeded by lateral lobes. Epipods absent from pere-
opods. First walking legs overreaching chelipeds. Fixed finger of

1-3: Female holotype, carapace length 31.4 mm; by K. Baba & M. de Saint Laurent.

2: Carapace and abdomen, lateral view; scale bar 10 mm.

1: Carapace and abdomen, dorsal view; scale bar 10 mm. 3: Telson; scale bar 1 mm.

References:
BABA K. (2005) Galathea Report 20: 1-317.
BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.
DESBRUYÈRES D., ALAYSE-DANET A.M., OHTA S. & the Scientific Parties of Biolau and Starmer Cruises (1994) Mar. Geol. 116: 227-242.

E. MACPHERSON & M. SEGONZAC Denisia 18 (2006): 447

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Munidopsis WHITEAVES, 1874

fixed finger, bearing a few spines along mesial margin; fingers

Size: Up to 45 mm carapace length.
spooned at tip. Dactyli of walking legs smoothly narrowed dis-
Morphology: Carapace with gastric region having group of tally, flexor margin bearing ultimate denticle equidistant be-
spines including two epigastric spines and some additional tween penultimate denticle and end of article. Epipods present
small spines or tubercular spines on scale-like ridges. Lateral on chelipeds.
margin having first spine (anterolateral) distinctly larger than
Remarks. This species has been reported from the hydrother-
antennal spine, directed anterolateral, second spine stronger
mal vents of the Galapagos Spreading Center under the name
than first. Rostrum subtriangular in proximal half, distally nar-
of M. subsquamosa HENDERSON, 1885, but it belongs to a differ-
rowed, slightly upcurved, dorsally carinated. Front margin
ent species, probably undescribed. A more careful examination,
oblique, antennal spine small. Abdominal segments unarmed,
including molecular analyses, of the specimens from different
sixth segment having posteromedian margin slightly convex,
localities will clarify the systematic status of this species.
not produced. Telson composed of 10 plates, midlateral plate
produced on anterolateral margin. Ocular peduncles broad at Biology: Occurrences around the hydrothermal vents may be
base, distomesially with eye-spine distinctly longer than revised.
cornea; cornea relatively small, as broad as eye-spine. Fixed fin-
Distribution: Galapagos Spreading Center, East Pacific Rise.
ger of chelipeds without denticulate carina on distolateral mar-
gin. First walking leg exceeding chelipeds; palm shorter than

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

1: Carapace, dorsal view; scale bar 5 mm;

from BABA (2005).

Reference:
BABA K. (2005) Galathea Report 20: 1-317.

E. MACPHERSON & K. BABA Denisia 18 (2006): 448

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

Arthropoda, Crustacea, Decapoda, Anomura, Galatheidae

Shinkaia crosnieri BABA & WILLIAMS, 1998

dorsally flattened projection. Chelipeds strong, broad, de-

Size: Up to 43.7 mm carapace length (Taiwanese specimen 45
pressed, with numerous short spines. Walking legs stout and
mm, 58.9 mm with rostrum).
moderately flattened. Epipods present on chelipeds, first and
Morphology: Dorsal carapace surface smooth, lateral margins second walking legs.
bearing many small spines. Rostrum prominent, flattened dor-
Distribution: Western Pacific: Edison Seamount; Okinawa
sally, triangular, not carinated dorsally. Orbital margin concave,
Trough, Iheya Ridge, North Iheya Knoll; off NE Taiwan, 1200-
antennal spine present. Pterygostomian flap anteriorly pro-
1500 m.
duced, covering greater part of antennal peduncle. Dense long
plumose setae on sternum, pterygostomian flap and ventral sur-
face of pereopods. Abdomen unarmed. Flattened eyes, with

1: Male carapace
length 45 mm,
dorsal view;
by T.-Y. Chan.

References:
BABA K. (2005) Galathea Report 20: 1-317.
BABA K. & A.W. WILLIAMS (1998) Zoosystema 20: 143-156.
CHAN T.Y., LEE D.-A. & C.S. LEE (2000) Bull. Mar. Sci. 67(2): 799-804.
FUJIKURA K. & T. OKUTANI (2002) Benthos Research 57: 21-30.
WATABE H. (2000) JAMSTEC J. Deep-Sea Res. 17: 29-34.

E. MACPHERSON & K. BABA Denisia 18 (2006): 449

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Arthropoda, Crustacea, Decapoda, Anomura, Kiwaidae
Kiwa hirsuta MACPHERSON, JONES & SEGONZAC, 2005 “yeti crab”
Size: Carapace length 51.5 mm (58.6 mm with rostrum), total
length 88.4 mm.
Color: Uniformly white, with yellow corneous tip at the ex-
tremity of the chelipeds 1.
Morphology: Carapace, excluding rostrum, 1.3 times longer
than broad, dorsal surface smooth. Front margin slightly
oblique, with small tooth near rostrum; anterolateral angle
rounded. Rostrum broadly triangular, horizontal, slightly con-
cave dorsally, lateral borders granulated, with long uniramous
setae; ventral side slightly carinated. Insertion of fifth pereopod
not visible and situated below sternal plastron. Eyes strongly re-
duced, membranous remains, without pigment. Antennal scale
absent. Antennal peduncle without scaphocerite. Flagellum as
long as carapace without rostrum. Chelipeds and walking legs
with dense mat of setae. Telson as wide as long, median trans-
verse suture separating anterior and posterior portions. Che-
lipeds and walking legs with numerous rows of spines, each
spine with yellow corneal tip and tuft of long and dense
plumose setae, only absent in cheliped fingers, setae denser and
longer in mesial and ventral sides than in lateral and dorsal
sides. Chelipeds nearly symmetric, slightly more than twice as
long as carapace including rostrum. Fingers somewhat triangu-
lar, without setae, having numerous spines decreasing in size
distally, distal areas of fingers unarmed; slightly gapping, and
1: Carapace and abdomen, lateral view; scale bar 10 mm;
by J.-F. Dejouannet © IRD.
2: Sixth segment of abdomen, telson and uropods, dorsal view;
scale bar 5 mm; by J.-F. Dejouannet © IRD.
M. SEGONZAC
distally spoon-shaped; movable finger with proximal large den-
ticulate tooth followed by cutting margin bordered with
smooth, low, corneous scales, ending in acute corneous point;
fixed finger with some proximal small teeth, followed by cut-
ting edge similar to movable finger and ending in acute cor-
neous point, additional row of mesial granules ending in acute
corneous point; fingers distally crossing, corneous tip of mov-
able finger crossing between two corneous tips of fixed finger.
Paired pleopods present.
Setae: the pereopods, and in particular the chelipeds, are dense-
ly covered with flexible setae (ca. 15 mm) having clusters of fil-
amentous bacteria, mainly at distal part. Other rigid chitinous
setae (ca. 13 mm) are barbed in the distal part, ending in a rigid
spine, and they are regularly inserted in pairs mainly on the
merus of the chelipeds. They are deprived of bacteria.
Biology: Occurs at densities of one to two individuals per 10
m2, more or less regularly spaced on the zone of pillow basalt
surrounding active hydrothermal vents, and at the base of
chimneys among vent mussels Bathymodiolus sp., crabs
bythograeid spp. and ophidiid fish. Omnivorous.
Distribution: Pacific-Antarctic Rise: German Flats, 38°S. This
animal was first observed (and noted as “type Shinkaiinae”, but
not collected), in 2001 by the German cruise Sonne SO-157
(STECHER et al. 2002).
3: Two specimens on mussel bed, among
bythograeid crabs on vent site Annie’s Anthill
(Southern East Pacific Rise); cruise PAR 5; by courtesy
of R. Vrijenhoek © MBARI.
Denisia 18 (2006): 450–451
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

4: Male holotype, dorsal view; by A. Fifis © Ifremer.

5: One specimen on pillow lava, taken on the site Pâle Etoile (Southern East Pacific Rise); cruise, PAR 5;
by courtesy of R. Vrijenhoek © MBARI.

References:
MACPHERSON E., JONES W.J. & M. SEGONZAC (2005) Zoosystema 27(4): 709-723.
STECHER J., TÜRKAY M. & C. BOROWSKI (2002) Cah. Biol. Mar. 43: 271-274.

451
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Decapoda, Anomura, Chirostylidae

Uroptychus bicavus BABA & DE SAINT LAURENT, 1992

Size: Up to 7.5 mm carapace length. Biology: Living in low number among vent community.
Morphology: Carapace smooth, lateral margin without distinct Distribution: North Fiji Back-Arc Basin: site Mussel Valley.
spine other than anterolateral spine. Epigastric spines on dorsal
surface. Rostrum moderately broad triangular, slightly upturned
apically. Fourth sternite rounded on anterolateral corner. An-
tennal scale barely reaching midlength of fifth antennal article.
Propodus of first walking legs slightly longer than carpus.
Dactylus of walking legs with flexor marginal spines not con-
tiguous to flexor margin.

2: Habitus, lateral view; holotype male, carapace length 7.5 mm;

scale bar 1 mm; by K. Baba & M. de Saint Laurent.

1: Habitus, dorsal view; holotype male,

carapace length 7.5 mm; scale bar 1 mm;
by K. Baba & M. de Saint Laurent.

References:
BABA K. (2005) Galathea Report 20: 1-317.
BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.
DESBRUYÈRES D., ALAYSE-DANET A.-M. & S. OHTA (1994) Mar. Geol. 116: 227-242.

E. MACPHERSON & K. BABA Denisia 18 (2006): 452

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

Arthropoda, Crustacea, Decapoda, Anomura, Chirostylidae

Uroptychus edisonicus BABA & WILLIAMS, 1998

Size: Up to 8.4 mm carapace length. Distribution: Western Pacific: Bismarck Archipelago, Edison
Seamount, near Lihi Island.
Morphology: Carapace smooth, lateral margin without distinct
spine other than anterolateral spine, not laterally serrate. Epi-
gastric spines absent. Rostrum narrow triangular, weakly curv-
ing dorsad distally. Fourth sternite relatively narrow on antero-
lateral corner. Antennal scale overreaching end of penultimate
segment. Propodus of walking legs unarmed on dorsal crest,
with spines on distal portion of flexor margin, distal-most re-
motely separated from distal second, and situated near juncture
with dactylus; propodus with convex flexor distal margin.
Dactylus of second walking leg with two distal spines remotely
separated from proximal group of spines.

2: Ovigerous female holotype, carapace length 6.2 mm; lateral view; scale
bar 3 mm; by K. Baba & A.B. Willians.

1: Ovigerous female holotype, carapace length

6.2 mm; dorsal view; scale bar 3 mm; by K. Baba
& A.B. Willians.

References:
BABA K. (2005) Galathea Report 20: 1-317.
BABA K. & A.B. WILLIAMS (1998) Zoosystema 20: 143-156.

E. MACPHERSON & K. BABA Denisia 18 (2006): 453

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

Arthropoda, Crustacea, Decapoda, Anomura, Chirostylidae

Uroptychus thermalis BABA & DE SAINT LAURENT, 1992

Size: Up to 8.4 mm carapace length. Biology: Living in few number among vent community.
Morphology: Carapace smooth, lateral margin without distinct Distribution: North Fiji Back-Arc Basin: White Lady. Known
spine other than anterolateral spine. Epigastric spines absent. also from Queensland, at 1497 m.
Rostrum narrow triangular, weakly curving dorsad distally.
Fourth sternite relatively narrow on anterolateral corner. An-
tennal scale overreaching end of penultimate segment. Propo-
dus of walking legs with spines on distal portion of flexor mar-
gin, distal-most remotely separated from distal second, and sit-
uated near juncture with dactylus; propodus without convex
flexor distal margin. Dactylus of second walking leg with two
distal spines remotely separated from proximal group of spines.

2: Habitus, lateral view; male holotype, carapace length 8.4 mm;

scale bar 1 mm; from BABA & SAINT LAURENT (1992).

1: Habitus, dorsal view; male holotype, carapace

length 8.4 mm; scale bar 1 mm; from BABA &
SAINT LAURENT (1992).

References:
AHYONG S. & G.C.B. POORE (2004) Zootaxa 436: 1-88.
BABA K. (2005) Galathea Report 20: 1-317.
BABA K. & M. DE SAINT LAURENT (1992) Sci. Mar. 56: 321-332.

E. MACPHERSON & K. BABA Denisia 18 (2006): 454

 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Chordata, Chondrichthyes & Osteichthyes
Since the discovery of animal communities in oceanic hy-
drothermal vents in 1977, fishes have been regularly observed
in association with these chemosynthetically driven communi-
ties, but in most cases they are difficult to catch and therefore
species identification can often only rely on images captured by
the diving vehicles.
Ichthyologic information pertaining to species inhabiting
the deep-sea hydrothermal vents is mentioned in over 30 pa-
pers and even in the more detailed and updated lists (BISCOITO
et al. 2002; GEISTDOERFER 1996, 1998; TUNNICLIFFE 1991) there
are species missing. The situation for bathyal species inhabiting
the periphery of active vent fields is even less clear since the
vast majority of identifications have been based on video
records or photographs.
Species living inside active fields
Vent fishes were found at only 20 of some 50 active vent
fields discovered to date. The specific diversity found is low and
the degree of endemism is high, which seems to be an overall
characteristic of the hydrothermal vent fauna (TUNNICLIFFE
1991).
Hydrolagus pallidus from Lucky Strike, Mid-Atlantic Ridge, Atos cruise © Ifremer.
M. BISCOITO
The family Zoarcidae dominates in terms of the number of
species and biomass. Eelpouts form a highly diverse family with
over 220 known, mostly benthic species (WEITZMAN 1997) and
are one of the more successful fish families to occupy continen-
tal slopes down to 5000 m (ANDERSON 1994; WEITZMAN 1997).
Nonetheless, it is remarkable that they have been able to adapt
(and evolve) to the vent environments. As pointed out by
GEISTDOERFER (1996) these adaptations are neither anatomical,
nor trophic. In order to cope with the chemical conditions of
vents and seeps, these species must have biochemical adapta-
tions. These adaptations, as well as the factors conditioning the
distribution of the species need further investigation. At the
present level of our knowledge on the taxonomy and distribu-
tion of the species, zoogeographical considerations cannot be
produced.
Bathyal species living in
the vicinity of the vents and seeps
As for the peripheral bathyal fish fauna, data available are
far from being satisfactory. The five best-represented families
sensu lato (Centrophoridae, Somniosidae and Etmopteridae),
Denisia 18 (2006): 489-490
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Macrouridae, Ophidiidae, Squalidae, Moridae and Synapho-
branchidae correspond to the most common families in the
deep sea (WEITZMAN 1997). However, the variations in num-
bers of recorded species from field to field are enormous and
definitely reflect insufficient research.
Although there are twice as many known active sites in the
Pacific as in the Atlantic, the number of species recorded in the
latter is higher (38 versus 43). Depth may well be the main rea-
son for this, as vent fields in the Atlantic range from 850 m
down to 3650 m and the ones in the East Pacific are all around
2500 m of depth. As pointed out by MERRETT & HAEDRICH
(1997) for the Porcupine Seabight, demersal fish abundance
tends to increase with depth attaining a maximum around 1000
m and then decreases sharply. This is in accordance with avail-
able data, wherein the highest number of records is from Menez
Gwen (850 m) and Lucky Strike (1700 m) on the Mid-Atlantic
Ridge (DESBRUYÈRES et al. 2001).
The fact that the fields near the Azores Triple Junction
(Mid-Atlantic Ridge) have been more intensively studied by
professional ichthyologists can also have an influence in the re-
sults obtained. This bias precludes us from drawing conclusions
on abundance, diversity and zoogeography of this fauna. To
overcome it, scientific teams working on hydrothermal vent bi-
ology should be reinforced with fish biologists as well as making
more ship-time available for this kind of research.
The species treated herein are not only those who live in-
side the active fields, which are commonly considered as “vent
endemic”, but also some of the more commonly seen in the pe-
riphery of the vents, in some cases feeding even on vent inver-
tebrates and thus contributing for the export of energy from the
chemosynthetically-driven environment to the photosyntheti-
cally-dependent bathyal environment.
New species
Amongst the so-called “vent endemic” species, at least six
new to science are being described while this contribution is
being published: three Zoarcidae from 9°N East Pacific Rise,
the Kermadec Arc and the Rodriguez Triple Junction in the In-
dian Ocean respectively, one Ophidiidae from 17°S southern
Pacific Pacific Rise, one Myxinidae from 38°S Pacific-Antarc-
tic Ridge and one Cynoglossidae from seamounts at the Mari-
ana Arc and the Kermadec Arc (E. Anderson, J. Hashimoto, J.
Nielsen & C. Roberts, pers. comm.).

References:
ANDERSON M.E. (1994) Ichthyol. Bull. J.L.B. Smith. Inst. Ichtyol. 60: 1-120.
BISCOITO M., SEGONZAC M., ALMEIDA A.J., DESBRUYÈRES D., GEISTDOERFER P., TURNIPSEED M. & C. VAN DOVER (2002) Cah. Biol. Mar. 43: 359-362.
DESBRUYÈRES D., BISCOITO M., CAPRAIS J.-C., COLAÇO A., COMTET T., CRASSOUS P., FOUQUET Y., KHRIPOUNOFF A., LE BRIS N., OLU K., RISO R., SARRADIN P.-M., SEGONZAC
M. & A. VANGRIESHEIM (2001) Deep-Sea Res. Part I 48: 1325-1346.
GEISTDOERFER P. (1996) Oceanol. Acta 19(5): 539-548.
GEISTDOERFER P. (1998) Ann. Inst. Océanogr. 74(2): 201-215.
MERRETT N.R. & R.L. HAEDRICH (1997) Deep-sea Demersal Fish and Fisheries. Chapman & Hall, London: 1-282.
TUNNICLIFFE V. (1991) Oceanogr. Mar. Biol. Annu. Rev. 29: 319-407.
WEITZMAN S.H. (1997) in RANDALL J.E. & A.P. FARRELL (Eds.) Deep-Sea Fishes: 43-78.

490
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Chordata, Vertebrata, Chondrichthyes, Chimaeriformes, Chimaeridae

Hydrolagus affinis (CAPELLO, 1868) “small-eyed rabbitfish”

Size: To about 1300 mm total length.
Color: Uniform dark violet/brown.
Morphology: Body greatly tapering from a massive head and
trunk to a pointed caudal fin with a short filament at its tip.
Snout short, somewhat conical, overhanging mouth. First dor-
sal fin short-based, triangular and high, with a strong spine in
front. Second dorsal fin long and low. Pectoral fins not reach-
ing to pelvic fin base when laid back. Anal fin continuous with
caudal fin.
Biology: Benthopelagic on continental slopes and down to
abyssal plains. At Lucky Strike, specimens were observed swim-
ming gently above the bottom and off the hydrothermal vent
field. Carnivorous. At Lucky Strike, preys included the vent
endemic mussel, Bathymodiolus azoricus. Oviparous.
Distribution: General: Western Atlantic: from Davis Strait
and off Newfoundland to Cape Cod. Eastern Atlantic: Den-
mark Strait, Rockall Trough, northern Bay of Biscay and off the
coast of Portugal. Possibly a much wider Distribution than
records show. Mid-Atlantic Ridge vents: Lucky Strike, Mount
Saldanha, Famous Segment, Rainbow. Depth range: 300 to
2400 m.

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

References:
HARDY G.S. & M. STEHMANN (1990) Arch. Fischereiwiss. 40(3): 229-248.
MARQUES A. & F. PORTEIRO (2000) Copeia 2000(3): 806-807.
MØLLER P.R., KULLBERG T. & O.A. JØRGENSEN (2004) Cybium 28(1): 55-60.
STEHMANN M. & D.L. BÜRKEL (1984) in WHITEHEAD P.J.P., BAUCHOT M.-L., HUREAU J.-C., NIELSEN J. & E. TORTONESE (Eds.) Fishes of the North-eastern Atlantic
and the Mediterranean 1: 212-215.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 491

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

Chordata, Vertebrata, Chondrichthyes, Chimaeriformes, Chimaeridae

Hydrolagus pallidus HARDY & STEHMANN, 1990 “pallid ghost shark”

Size: To about 1300 mm total length.
Color: Uniform creamy to light greyish.
Morphology: Body greatly tapering from a massive head and
trunk to a pointed caudal fin with a short filament at its tip.
Snout short, somewhat conical, overhanging mouth. First dor-
sal fin short-based, triangular and high, with a strong spine in
front. Second dorsal fin long and low. Pectoral fins not reach-
ing to pelvic fin base when laid back. Anal fin continuous with
caudal fin.
Biology: Benthopelagic on continental slopes and down to
abyssal plains. At Lucky Strike, specimens were observed swim-
ming gently above the bottom and off the hydrothermal vent
field. Carnivorous, possibly feeding on small fishes and inverte-
brates. Oviparous.
Distribution: General: Western Atlantic: Davis Strait, Cana-
da, Bear Seamount, New England; Eastern Atlantic: Green-
land, off Iceland, from the southern Bay of Biscay to off west-
ern Scotland and Mid-Atlantic Ridge: Lucky Strike, Menez
Hom, Mount Saldanha, Rainbow. Depth range: 1200-2075 m.

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

2: At Rainbow, Mid-Atlantic Ridge; cruise Atos © Ifremer.

References:
HARDY G.S. & M. STEHMANN (1990) Arch. Fischereiwiss. 40(3): 229-248.
MØLLER P.R., KULLBERG T. & O.A. JØRGENSEN (2004) Cybium 28(1): 55-60.
SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 492

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

Chordata, Vertebrata, Osteichthyes, Anguilliformes, Synaphobranchidae

Dysommina rugosa GINSBURG, 1951

field with substantial coverage of microbial mat. Not found in

Size: Up to 352 mm total length.
non-venting habitats at the same depth nearby. Swims short
Morphology: Naked ilyophine eel with small broad-based pec- distances into the water column, probably to feed.
toral fins, their tips not sharply pointed. Dorsal-fin origin for-
Distribution: General: Widespread at tropical latitudes. West-
ward, about one head length behind a vertical through the gill
ern Atlantic Ocean, from off the Carolinas to the Caribbean.
slits. Fleshy, papillose snout slightly overhangs tip of lower jaw.
Southwestern Indian Ocean, Canal of Mozambique. Pacific
Gill slits separate, small (about equal to diameter of eye), cres-
Ocean, Hawaii. At vents, this species has been reported only
centic, oriented nearly vertically on the ventrolateral surface.
from warm-water vents at the peak of a volcanic cone, Nafan-
Vertebrae 127-134.
ua, at Vailulu’u Seamount, Samoan Archipelago. Depth range:
Biology: Benthopelagic; carnivorous, feeding on shrimps and 260-775 m.
possibly other invertebrates. Oviparous. At Vailulu’u, living in
crevices and holes in a low-temperature (10°C) diffuse vent

1: Habitus; from ROBIN & ROBIN (1989).

2: In situ view at Vailulu’u Seamount; by C.

Young © OIMB.

Reference:
ROBINS C.H. & C.R. ROBINS (1989) in: BÖHLKE E. (Ed.) Fishes of the Western North Atlantic. Part 9(1): 207-253.

C. YOUNG & M. BISCOITO Denisia 18 (2006): 493

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

Chordata, Vertebrata, Osteichthyes, Anguilliformes, Synaphobranchidae

Ilyophis saldanhai KARMOVSKAYA & PARIN, 1999

straight, its rictus situated behind a vertical through posterior

Size: Total length 410 mm (Atlantic specimens), and 335 mm
margin of orbit. Mouth cavity covered with longitudinal folds
(Pacific specimen).
bearing numerous papillae that differ in size and shape. Teeth
Color: Uniformly brownish. in jaws conical, pointed, slightly curved, bent inside, and close-
ly set in rows forming a band on each jaw. Teeth increasing in
Morphology: Body elongate, compressed and naked. Origin of
size in inner rows and medially on each jaw. Teeth on palate
dorsal fin above the 11th pore of the lateral line and behind
larger. Teeth on vomer forming two irregular longitudinal rows
end of pectoral fin, origin of anal fin after the 32nd pore of the
(Fig. 2). Eyes small, round, covered with a thin membrane,
lateral line, both fused with caudal fin (fig. 1). Head well dif-
their diameter 6.7 times in head and 2.5 times in snout. Ante-
ferentiated from the rest of the body, its profile slightly convex
rior nostrils tubular, directed forward, bearing small flaps.
at level of posterior margin of orbit. Head 3.3 times in pre-anal
Openings of posterior nostrils simple and round, their indented
length. Snout length moderate (2.65 times in head). Fleshy tip
margins slightly turned out. Gill slits semicircular, situated hor-
of snout with two pairs of well-developed plicae (fig. 2). Two
izontally at lower part of head before bases of pectoral fins
sharply pronounced medial folds descending from tip of snout;
(CAUSSE et al. 2005; KARMOVSKAYA & PARIN 1999).
at its lower edge, then bending horizontally to the right and left
of median line of snout. Two other shorter lyre-shaped folds sit- Biology: This species is benthopelagic, associated with vent
uated laterally to two median folds. Supra-orbital canal with communities. Carnivorous, feeding on hydrothermal vent in-
three pores, postorbital canal with two pores, infra-orbital canal vertebrates (small shrimps and polychaetes).
with five pores, preoperculo-mandibular canal with eight pores,
Distribution: East Pacific Rise: 21°S, hydrothermal vent site
six before the commisure, the seventh at the level of the com-
Gromit, and MAR: Broken Spur vent field.
misure and the eigth on the preopercle. Mouth opening

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

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

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

2: In vivo, after collection from East Pacific Rise: 21°S, Gromit site; cruise Biospeedo, by Briand © Ifremer.

3: In situ, cruise Biospeedo © Ifremer.

References:
CAUSSE R., BISCOITO M. & P. BRIAND (2006) Cybium 29(4): 413-416.
KARMOVSKAYA E.S. & PARIN N.V. (1999) J. Ichthyol. 39(5): 353-362.
NIELSEN J. & D.C. COHEN (2005) Cybium 29(4): 395-398.
PARIN N.V. (1995) Vopr. Ihtiol. 35(5): 698-701.

495
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Chordata, Vertebrata, Osteichthyes, Anguilliformes, Synaphobranchidae

Thermobiotes mytilogeiton GEISTDOERFER, 1991

Size: Up to 247 mm. Biology: Dwelling among mussels.

Color: In formalin pale brownish, beige in vivo with abdomen Distribution: Lau Back-Arc Basin: Valu Fa Rise.
darker.
Morphology: Eel-shaped, no scales, no pectoral fins, lateral line
present. Dorsal fin origin behind level of anus. Teeth on both
jaws, small and needle-like. Vertebrae 132.

1: Total view; by P. Briand © Ifremer. 2: Anterior part in lateral view; by P. Briand © Ifremer.

4: In situ view of a specimen on a mussel bed of Bathymodio- 3: Anterior end in ventral view; by P. Briand © Ifremer.
lus brevior, at Lau Basin; cruise Biolau © Ifremer.

References:
GEISTDOERFER P. (1991) C. R. Acad. Sci. Paris, Sér. III 312: 91-97.
MEUNIER F.J. & P. GEISTDOERFER (1991) Cybium 15: 83-87.

P. BRIAND Denisia 18 (2006): 496

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

Chordata, Vertebrata, Osteichthyes, Gadiformes, Lotidae

Gaidropsarus RAFINESQUE, 1810

Size: To about 360 mm standard length. Remark: A specimen was collected with the “Nautile” sub-
mersible during Diva 2 cruise and its description is in press.
Color: Generally reddish pink with a distinct white mark on
Other specimens caught in the Bay of Biscay and the Rockall
cheeks.
Trough, currently under study, may eventually prove to belong
Morphology: Elongate gadoid fish with two dorsal fins, the first to the same yet undescribed species.
with a moderately elongate first finray, followed by fine short
Biology: Very few data. Benthic. The specimens observed at
rays. A single anal fin. Second dorsal and anal fins long and
Lucky Strike were usually inside crevices, on the mussel beds,
moderately high. Three barbels, one on chin and one on each
well inside the active hydrothermal vent field. Carnivorous,
anterior nostril. Pre-anal length longer than post-anal (exclud-
feeding on hydrothermal vent crustaceans (alvinocaridid
ing the caudal fin). Snout longer than twice the eye diameter.
shrimps).
Forty seven vertebrae (16+31).
Distribution: Mid-Atlantic Ridge: Lucky Strike.

2: At Lucky Strike; cruise Seahma

1: By Luiz Saldanha © LMG, 1986. © FCT & Ifremer.

3: At Lucky Strike; cruise Diva © FCT & Ifremer. 4: Anterior part in dorsal view; by P. Briand © Ifremer.

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

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 497

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

Chordata, Vertebrata, Osteichthyes, Gadiformes, Moridae

Lepidion schmidti SVETOVIDOV, 1936

Size: Up to 1230 mm standard length. Biology: Probably benthopelagic on continental slope. At

Rainbow, Lucky Strike and Menez Gwen, specimens were
Morphology: Head conical and robust. Posterior nostril ahead
found lying on the bottom, amongst rocks or in crevices, in the
of erye. Chin barbel present, as long as snout. Orbit 3.5-5.8 in
inner periphery of the active fields (over brown sulphide de-
head length. Second ray of first dorsal fin very elongate. Later-
posits).
al line well marked until near the origin of caudal peduncle.
Distribution: General Eastern North Atlantic: in the Bay of
Remark: One specimen caught with the “Nautile“ at Rainbow
Biscay and West of Ireland; Pacific Ocean: Sagami Bay, Japan;
and others at Lucky Strike with bottom long lines.
Mid-Atlantic Ridge: Menez Gwen. Depth range: 900-2300 m.

1: At Rainbow vent field; cruise Atos © Ifremer.

3: At Lucky Strike vent field; cruise Marvel © Ifremer.

2: At Lucky Strike vent field; cruise Victor Première

© Ifremer.

References:
COHEN D.M. (1984) in WHITEHEAD P.J.P., BAUCHOT M.L., HUREAU J.-C., NIELSEN J. & E. TORTONESE (Eds.) Fishes of the North-eastern Atlantic and the Mediter-
ranean 2: 713-723.
SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 498

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

Chordata, Vertebrata, Osteichthyes, Ophidiiformes, Bythitidae

Cataetyx laticeps KOEFOED, 1927

Size: To about 650 mm standard length.
Color: Greyish brown.
Morphology: Body elongate. Head dorsoventrally flattened,
with a strong opercular spine. Dorsal and anal fins confluent
with caudal. Dorsal finrays 91-107; anal finrays 74-87; pectoral
finrays 22-29; pelvic finrays 1.
Remarks: Specimens were collected with the submersible at
Lucky Strike. A second species of this genus was recorded from
Rainbow (a single specimen caught in a fish trap) and so far
cannot be assigned to any of the previously known species of
Cataetyx.
Biology: Benthic species. At Lucky Strike, specimens occurring
often in pairs on sulphide deposits amongst shells of dead mus-
sels, on the inner border of the hydrothermal field. Carnivo-
rous. At Lucky Strike, feeding on hydrothermal vent crus-
taceans (crab Segonzacia mesatlantica, alvinocaridid shrimps).
Viviparous.
Distribution: Generally known from a few localities in the
Northeast Atlantic and western Mediterranean. Found also in
the Azores and along West Africa to the Cape of Good Hope.
Possibly also in the Gulf of Mexico. Mid-Atlantic Ridge:
Menez Gwen, Lucky Strike, Mount Saldanha, Menez Hom, and
Rainbow. Depth range: 900-2830 m.

1: At Lucky Strike, Tower Eiffel vent field; cruise Exomar © 4: At Lucky Strike, Tower Eiffel vent field; cruise
Ifremer. Exomar © Ifremer.

References:
NIELSEN J.G. (1984) in WHITEHEAD P.J.P., BAUCHOT M.L., HUREAU J.-C., NIELSEN J. & E. TORTONESE (Eds.) Fishes of the North-eastern Atlantic and the Mediter-
ranean 3: 1153-1157.
SALDANHA L. (1994) Cybium 18(4): 460-462.
SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 499

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

Chordata, Vertebrata, Osteichthyes, Ophidiiformes, Bythitidae

Thermichthys hollisi (COHEN, ROSENBLATT & MOSER 1990)

(Thermarces-like) with half of its length, during cruise

Size: 218-304 mm standard length.
Biospeedo. In the video images, only the posterior half of the
Morphology: A bythitid genus with joined vertical fins, short prey-fish has been swallowed and the anterior part is slowly
body and blunt snout. Body with imbricate scales and head moving.
naked. Mouth terminal, upper jaw ending well behind eye. Eye
Distribution: Known from two specimens from the Galapagos
shorter than snout. Opercular spine strong. Palatine with teeth.
Spreading Center and Southern East Pacific Rise: 17°S, site
Developed rakers on anterior arch 3. Rays in dorsal fin 122-
Hobbs.
124, anal fin 88, causal fin 12, pectoral fin 36-37 and ventral fin
1. Vertebrae 76, precaudal 20-21.
Biology: One specimen was caught associated with hydrother-
mal vents. A specimen of T. hollisi was seen eating a zoarcid fish

1 top: Specimen taken in vivo, onboard, dorsal view; bottom: dorsolateral view; East Pacific Rise: 17°S,
site Hobbs, cruise Biospeedo © Ifremer.

References:
COHEN D.M., ROSENBLATT R.H. & H.G. MOSER (1990) Deep-Sea Res. A 37: 267-283.
NIELSEN J.G. & D.M. COHEN (2005) Cybium 29(4): 395-398.

J.G. NIELSEN & D.M. COHEN Denisia 18 (2006): 500

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

Chordata, Vertebrata, Osteichthyes, Ophidiiformes, Ophidiidae

Ventichthys biospeedoi NIELSEN, MOLLER & SEGONZAC, in press

Size: Maximum standard length 282 mm. Biology: Species abundant, living in shimmering vent fluids
with temperatures between 2 and 7°C, among hydrothermal
Morphology: Body robust, very small, overlapping scales on
community: mytilid and clam bivalves, stalked barnacles.
head and body, thick skin, and four indistinct lateral lines; dor-
Necrophagous.
sal fin origin above tip of pectorals, basis of pelvic fins below
hind margin of opercle; head broad with blunt snout; strong op- Distribution: East Pacific Rise: 17°S, site Oasis, but probably
ercular spine covered by thick skin; upper jaw ends just behind the same species occurs on northern and southern sites.
eye; teeth granular, one median basibranchial tooth patch; an-
terior gill arch with 10-11 long rakers; number of rays in dorsal
fin 80-89, caudal fin 8, anal fin 64-72, pelvic fin 2, pectoral fin
24-25; number of vertebrae 16-17+36.

1: Couple of specimens caught at East Pacific Rise: 17°S, site Oasis; cruise Biospeedo, by P. Briand © Ifremer.

2: In situ view of several speci-

mens on shimmering water, on
mussel bed Bathymodiolus
thermophilus, with stalked
barnacle (Neolepas n. sp.) and
sea anemaone (Chondrophellia
cf. coronata); East Pacific Rise:
17°S, site Oasis; cruise
Biospeedo © Ifremer.

Reference:
NIELSEN J.G. & P.R. MOLLER & M. SEGONZAC (in press) Zootaxa.

J.G. NIELSEN, MOLLER P.R. & M. SEGONZAC Denisia 18 (2006): 501

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

Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae

Pachycara gymninium ANDERSON & PEDEN, 1988

Size: Up to about 420 mm standard length. Biology: Benthic over brown and green mud bottoms. Carniv-
orous, eating amphipods, isopods and polychaetes. Oviparous, a
Color: In life dark brown, head and pectoral fin darker, almost
gravid female caught in February.
black. Margins of vertical fins and peritoneum black.
Distribution: General: Northeast Pacific Ocean, off the Queen
Morphology: Body short, deep, broader in cross section, when
Charlotte Islands, British Columbia, south of Guadalupe Is-
compared with its congeners. Head large, ovoid. Pelvic fins
land, Mexico and Gulf of California. Juan de Fuca Ridge: Cryp-
present (4.8-11.3% of head length). Mediolateral branch of lat-
to Vent Field, Axial Seamount; Endeavour Segment; Ham-
eral line originating in pectoral axil just posterior to vertical
mond’s Hell vent. Depth range: 1575-3219 m.
through pectoral base. Scales absent on nape, or, if present, not
extending anterior to line connecting anterodorsal edges of gill
slits. Vertebrae 102-109. Anal fin origin associated with verte-
brae 27-31.

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

References:
ANDERSON M.E. & A.E. PEDEN (1988) Proc. Calif. Acad. Sci. 46(3): 83-94.
TUNNICLIFFE V., MCARTHUR A.G. & D. MCHUGH (1998) Adv. Mar. Biol. 34: 353-451.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 502

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

Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae

Pachycara rimae ANDERSON, 1989

Size: Up to 403 mm standard length. Remark: Data available are not sufficient to determine
whether this species is endemic to hydrothermal vent environ-
Color: Uniformly light brown, eye and abdomen bluish.
ments or not.
Morphology: Head deep, rounded, somewhat shorter than sim-
Distribution: Galapagos Spreading Center. Known only from
ilarly sized congeners. Pelvic fins nublike, of two soft rays. Ver-
the holotype.
tebrae 93. Dorsal-fin rays 86, anal-fin rays 70. Lateral line of
mediolateral branch only. Dorsal fin origin associated with ver-
tebra 8. Pseudobranchs absent.

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

References:
ANDERSON M.E. (1989) Proc. Calif. Acad. Sci. 46(10): 221-242.
COHEN D.M. & R.L. HAEDRICH (1983) Deep-Sea Res. 30(4A): 371-379.
COHEN D.M., ROSENBLATT R.H. & R.L. HAEDRICH (1985) Biol. Soc. Wash. Bull. 6: 229-230.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 503

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

Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae

Pachycara saldanhai BISCOITO & ALMEIDA, 2004

Size: Up to 256 mm total length. Biology: Benthic, not very numerous over sulphide deposits
and among mussels inside the active field. Food: hydrothermal
Color: In life, light brownish grey, head, dorsal and anal fins
vent crustaceans.
darker. When preserved light brown, body with conspicuous
whitish scale pockets. Distribution: Up to present restricted to Mid-Atlantic Ridge:
Rainbow.
Morphology: Body elongate and compressed, scaled. Mouth
subterminal, no crests on chin. Dorsal fin origin over pectoral
fins. Pelvic fins present. Lateral line with two branches. Dorsal
fin rays 108-115, anal fin rays 90-95, vertebrae 117-123.

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

2: At Rainbow vent field; cruise

Marvel © Ifremer.

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

Reference:
BISCOITO M. & A.J. ALMEIDA (2004) Copeia 3: 562-568.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 504

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

Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae

Pachycara thermophilum GEISTDOERFER, 1994

Size: Up to 388 mm total length. Biology: Benthic, in areas with active smokers, swimming in
sea water 5-20°C, away from rocks and along smokers near
Color: Light brown with pinkish hues. Border of fins darker.
crowds of shrimps Rimicaris exoculata and Chorocaris chacei.
Morphology: Eelpout shaped. Pelvic fin rays 2. Mediolateral Carnivorous, feeding on hydrothermal vent shrimps.
lateral line only, originating just behind posterior-most postor-
Distribution: Mid-Atlantic Ridge: Snake Pit and TAG. Depth
bital pore. Scales absent on nape. Dorsal fin rays 105-107. Anal
range: 3500-3700 m.
fin rays 86-89. Total vertebrae 113-114. Dorsal fin origin asso-
ciated with vertebrae 7-8.

1: Habitus; by E. Heemstra © JLB Smith Inst. Ichth., 1996.

2: In situ at Snake Pit © Ifremer. 3: Habitus; by P. Briand © Ifremer.

References:
ANDERSON M.E. & H. BLUHM (1996) Trans. R. Soc. Afr. 51: 219-227.
GEISTDOERFER P. (1994) Cybium 18(2): 109-115.
PARIN N.V. (1995) J. Ichthyol. 35(9): 328-332.

P. GEISTDOERFER Denisia 18 (2006): 505

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

Chordata, Vertebrata, Osteichthyes, Perciformes, Zoarcidae

Pyrolycus manusanus MACHIDA & HASHIMOTO, 2002

Size: Up to 170 mm total length. Biology: Seen in areas close to active smokers and/or in and
around diffuse vent fluids. Carnivorous, alvinocaridid shrimps
Color: Light brown or beige.
were found in stomach contents of some specimens.
Morphology: Eeelpout shaped. Suborbital bones 6, canal with
Distribution: Manus Back-Arc Basin: Pacmanus and Desmos
6-7 pores; flesh gelatinous; dermal papillae absent from head;
sites.
gill slit large, extending ventrally beyond pectoral fin base;
pelvic fins present, each with 2-3 rays; scales, lateral line and
pseudobranch absent; pyloric caeca present; oral valve weak;
interorbital pore absent; postorbital pores 3; occipital pores 1;
palatopterygoid series weak; vomerine and palatine teeth pres-
ent; pectoral fin rays 16-17; caudal fin rays 8-10; vertebrae 22-
23+56-59=78-81.

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

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

References:
HASHIMOTO J., OHTA S., FIALA-MÉDIONI A., AUZENDE J.-M., KOJIMA S., SEGONZAC M., FUJIWARA Y., HUNT J.-C., GENA K., MIURA T., KIKUCHI T., YAMAGUCHI T., TODA T.,
CHIBA H., TSUCHIDA S., ISHIBASHI J., HENRI K., ZBINDEN M., PRUSKI A., INOUE A., KOBAYASHI H., BIRRIEN J.-L., NAKA J., YAMANAKA T., LAPORTE C., NISHIMURA K.,
YEATS C., MALAGUN S., KIA P., OYAIZU M. & T. KATAYAMA (1999) InterRidge News 8: 12-18.
MACHIDA Y. & J. HASHIMOTO (2002) Ichthyol. Res. 49: 1-6.

J. HASHIMOTO Denisia 18 (2006): 506

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

Chordata, Osteichthyes, Perciformes, Zoarcidae

Thermarces cerberus ROSENBLATT & COHEN, 1986

Size: Up to 370 mm total length.
Color: Whitish or pinkish.
Morphology: Eelpout shaped, head and body laterally com-
pressed, naked, covered with mucous, without scales. Lips dis-
tinct, thick and fleshy, continuous and smooth; oral valves ob-
solete. Head pores large and conspicuous. Occipital pores ab-
sent. Teeth in both jaws stout, conical and pointed. No lateral
line. Pelvic fins absent. Pectoral fins small, rounded, with rays
covered by thick skin. Vertebrae: 93-97.
Remarks: 1. Another close species, T. andersoni ROSENBLATT &
COHEN, 1986, was described from Galapagos Rift, but its syn-
onymy with T. cerberus is debated. 2. A new species of Ther-
marces was found at East Pacific Rise: 9°N, showing strong mor-
phological and color differences. It is being described elsewhere
(M. Biscoito & M. Segonzac, unpublished data).
Biology: Observed in areas of active smokers, associated with
Riftia pachyptila or on smoker walls. Very often stays in diffuse
venting areas, amongst rubble. Carnivorous, feeding on small
invertebrates, mainly amphipods and limpets, but also poly-
chaetes. Occasionally they have been seen biting the gills of R.
pachyptila.
Distribution: Galapagos Spreading Center; East Pacific Rise:
9°N, 13°N and 21°N; probably also at South East Pacific vent
sites.

1: Fresh specimen collected at East Pacific Rise: 13°N; by P. Briand © Ifremer.

M. BISCOITO & M. SEGONZAC Denisia 18 (2006): 507-508

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

2: In situ specimens from East Pacific Rise: 13°N, among vestimentiferan (Riftia pachyptila) and mussel bed of Bathymodiolus
thermophilus; cruise Phare © Ifremer.

References:
GEISTDOERFER P. (1986) Bull. Mus. Natl. Hist. Nat. Paris, Sér. 4A 8: 969-980.
GEISTDOERFER P. (1996) Oceanol. Acta 19(5): 539-548.
ROSENBLATT R.H. & D.M. COHEN (1986) Trans. San Diego Soc. Nat. Hist. 21: 71-79.

508
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Chordata, Vertebrata, Osteichthyes, Scorpaeniformes, Liparidae

Careproctus hyaleius GEISTDOERFER, 1994

Size: Up to 112 mm standard length. Biology: Frequently observed at the boundary of active vent
fields and occasionally among Riftia or within diffuse venting
Color: Whitish to pinkish; translucent.
areas.
Morphology: Small-sized fish, tadpole-shaped, with a round
Distribution: East Pacific Rise: 9° N to 13° N.
and globular head accounting for 22% of total length.
Branchial aperture small (32% of head length) and located in
the upper part of the body. Teeth simple, hooked and all simi-
lar. Body flaccid, covered by thick layer of mucous. Bare and
fragile skin. Pelvic fins modified into a sucking disc.

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

2: Ventral view; by P. Briand © Ifremer. 3: View in situ of a specimen (righ), with a zoarcid Thermarces
cerberus, tubeworms Riftia pachyptila and serpulids Lami-
natubus alvini; East Pacific Rise: 13°N. Cruise Hot 96 © Ifremer.

References:
BISCOITO M., SEGONZAC M., ALMEIDA A.J., GEISTDOERFER P., TURNSIPSEED M. & C. VAN DOVER (2002) Cah. Biol. Mar. 43: 359-362.
GEISTDOERFER P. (1994) Cybium 18(3): 325-333.

P. BRIAND Denisia 18 (2006): 509

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

Chordata, Vertebrata, Osteichthyes, Scorpaeniformes, Sebastidae

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

Size: Up to 500 mm standard length. Biology: Benthic. At Menez Gwen on pillow lavas, off the ac-
tive sites. Carnivorous. One anguilliform fish found inside the
Color: Reddish, dorsal fin with bluish pigment.
stomach of a specimen collected at Menez Gwen. Oviparous.
Morphology: Head large. Orbit large, much wider than snout
Distribution: General: Eastern Atlantic, from Ireland south-
length. Well developed spines on head. Dorsal fin ray with 12
ward to Mauritania. Mid-Atlantic Ridge: Menez Gwen.
spines and 8-9 rays. Pectoral fin with characteristic shape
(longest rays near upper part of fin) and with 20-23 rays.

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

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

References:
HUREAU J.-C. & N.I. LITVINENKO, (1984) in WHITEHEAD P.J.P., BAUCHOT M.-L., HUREAU J.-C., NIELSEN J. & E. TORTONESE (Eds.) Fishes of the North-eastern Atlantic
and the Mediterranean 3: 1211-1229.
SALDANHA L. & M. BISCOITO (1997) Bol. Mus. Munici. Funchal 49(283): 189-206.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 510

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

Chordata, Vertebrata, Osteichthyes, Macrouridae

Coryphaenoides armatus (HECTOR, 1875) “armed grenadier”

Size: To at least 800 mm total length. Biology: Benthopelagic. Carnivorous, feeding on benthic ani-
mals (amphipods, isopods, cumaceans), also pelagic animals
Color: Generally brownish to reddish brown, fin membranes
(mysids, euphausids and other crustaceans, echinoderms,
brownish, mouth and gill cavity blackish.
cephalopods and fishes). Two specimens were collected with
Morphology: Ventral parts of head mostly naked, including bottom long lines at Lucky Strike and one at Snake Pit, the lat-
snout, most ventral surfaces of suborbital space, ventral preop- ter had Rimicaris exoculata in its stomach. Oviparous.
ercular margin and anterior part of mandible. Premaxillary
Distribution: General: Worldwide. Marginal to the Southern
teeth in one or two rows, one row on mandible. Inner gill rak-
Ocean. Mid-Atlantic Ridge: Lucky Strike, Snake Pit, Rainbow;
ers on first arch 11 to 14. First dorsal fin with two spines and 8-
Galapagos Spreading Center; East Pacific Rise: 9°N. Depth
10 rays, pectoral fins rays I+17-21. Pelvic fins 10. Anus close to
range: 282-4700 m.
anal fin origin. No light organ.

1: Habitus; from GÜNTHER (1887).

2: In situ from
Mid-Atlantic Ridge;
cruise Atos © Ifremer.

References:
COHEN D.M. (1990) in QUÉRO J.-C., HUREAU J.-C., KARRER C., POST A. & L. SALDANHA (Eds.) Check-list of the Fishes of the Eastern Tropical Atlantic 2: 541-
563.
COHEN D.M., T. INADA, T. IWAMOTO & N. SCIALABBA (1990) FAO Species Catalogue. FAO Fish. Synop. 125(10): 1-442.
GEISTDOERFER P. (1988) Oceanol. Acta, 8 n° sp.: 125-130.
GEISTDOERFER P. (1991) C. R. Acad. Sci. Paris, Sér. III 312: 91-97.
GÜNTHER A. (1987) Challenger Reports, Zool. 22: 1-268.
MARQUES A & A.J. ALMEIDA (2000) InterRidge News 9(2): 16-17.
MERRETT N.R. & N.B. MARSHALL (1981) Progr. Oceanogr. 9: 185-244.

M. BISCOITO & A.J. ALMEIDA Denisia 18 (2006): 511

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

Major known deep-sea hydrothermal vent fields

Ocean Ridge / BAB Field Lat. Long. Depth (m) Sites
Atlantic Jan-Mayen Ridge Kolbeinsey 67°05’N 18°43’W 90
Mid-Atlantic Ridge Menez Gwen 37°51’N 31°31’W 840-865
Lucky Strike 37°17’N 32°16’W 1620-1730 Eiffel Tower, Statue of Liberty,
Sintra, Bairro-Alto, Isabel, Petit
chimiste, Fantôme, Montségur,
Hélène, Pico, Nuno, Elisabeth
Mount Saldanha 36°33’N 33°28’W 2300
Rainbow 36°13’N 33°54’W 2260-2350
Lost City 30°07’N 42°07’W 750-900
Broken Spur 29°10’N 43°10’W 3050-3875 Saracen’s Head, Wasp’s nest,
Spire
TAG 26°08’N 44°49’W 3635-3670 Mir zone, Kremlin
Snake Pit 23°23’N 44°58’W 3480 Moose, Beehive, The Nail
Logatchev 14°45’N 44°58’W 2930-3020 Irina Mound, Main Mound,
Anna-Louise
Ashadze-2 12°59’N 44°54’W 3200-3300
Ashadze-1 12°58’N 44°52’W 4100-4200
Ascension 4°48’S 12°22’W 3100
Mediterranean Gulf of Naples Capo Misseno 40°50’N 14°05’E coastal
Tyrrhenian Sea Capo Palinuro 40°01’N 15°16’E 9
Aegian Sea Milos 36°40’N 24°23’E 0-115
Indian Central Indian Ridge Edmond Vent Field 23°52’S 69°35’E 3290-3320
Kairei Vent Field 25°19’S 70°02’E 2415-2460
Gulf of Aden 11°57’N 43°40’E 1400-1600
North East Pacific Southern Explorer Magic Mountain 49°46’N 130°15’W 1797 Zooarium, Dead Chimney,
Ridge Zoo Chimney Field Recordbreaker, Zeus
Magic Mountain 49°46’N 130°15’W 1778 Stump, Tubeworm, Beercan,
Merlin Mound Limpet, Anhydrite
Magic Mountain 49°46’N 130°16’W 1778 Einstein, Eastern Island,
Mystic Mound Ridge Top
Magic Mountain 49°46’N 130°15’W 1785 Anhydrite 1-4, Magestic,
Magestic Chimney Obelisk, Digit Field
Juan de Fuca Ridge Middle Valley 48°27’N 128°42’W 2400
Endeavour Segment 48°N 129°04’W 2200
Sasquatch Field
Endeavour Segment 47°58’N 129°04’N 2200 Grendl
Salty Dawg Field
Endeavour Segment 47°57’N 129°06’W 2200 S & M, Peanut, Puffer, Hulk
Main Endeavour
Endeavour Segment 47°57’N 129°07’W 2200-2220 Godzilla
High Rise Field
Endeavour segment 47°55’N 129°08’W 2200 Faulty Towers Complex
Mothra Vent Field
CoAxial Segment 46°19’N 129°42’W 2291 Source site, Floc site , Flow site,
Mongo vent, Beard vent, Church
vent
Axial Volcano 45°59’N 130°02’W 1580 Shepherd vent
CASM Vent Field
Axial Volcano 45°55’N 129°59’W 1530-1544 Magnesia, Oxide, Blue nose, Easy,
Lava Flow 98 45°57’N Milky, Old worms, Minisnow,
nSRZ Roof, The Pit, Old flow, Nascent,
Cloud, Snail, Snow, Circ, Castle,
Village, Pillar, Joystick, Iron City,
Bag City, Crevice, Coquille vent
field, Dying

D. DESBRUYÈRES Denisia 18 (2006) 513–517

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

Ocean Ridge / BAB Field Lat. Long. Depth (m) Sites

Axial Volcano 45°56’N 130°00W 1547 White, Virgins Daughter, Virgin,
Ashes Vent Field Marshmallow, Mushroom, Gol-
lum, Inferno, Daves, Hairdo,
Medusa, Ropos, Phoenix, Styx,
Crack, Steve Mound
Cleft Segment 44°39’N 130°15’W 2270 Fountain
(Southern Juan
de Fuca)
Gorda Ridge Sea Cliff Vent Field, 42°15’N 126°42’W 2700
Northern Gorda Ridge
Escanaba Trough 41°00’N 127°29’W 3250
East Pacific Southern California Palos Verdes 33°42’N 118°19’W 1-10
White Point
Gulf of California Guaymas Basin 27°00’N 111°24’W 2000
East Pacific Rise 21°N 20°50’N 109°06’W 2615 Clam Acres
(Northern) 13°N 12°48’N 103°56’W 2635 Pogosud, Pogonord, Totem,
12°50’N 103°57’W Genesis, Elsa
9°N (Venture Hydro- 9°48’N 104°17’W 2500 Biovent, M-vent, Riftia Field,
thermal Field) 9°51’N Mussel Bed, East Wall, Tube-
worm Pilar, Tevnia Hole, Brasou-
cade, Tica
Galapagos Ridge Galapagos Spreading 00°47’N 86°07’W 2450-2490 Rose Garden, Mussel bed, Small
Center 00°49’N 86°13’W Fry, Garden of Eden
East Pacific Rise 7°S 7°21’S 107°47’W 2735-2752 White Christmas, Sarah’spring,
(Southern) 7°25’S Petite cheminée, Last hope
14°S 13°58’S 112°28’W 2623-2635 Pagoda, Smokin’Shank,
13°59’S Alvinella chimney, Lucky Eric
17°24’S 17°24’S 113°12’W 2575-2590 Stanley, Robbie Roost, S-vent,
17°25’S Nadir, Oasis, Rehu Marka
17°34’S 17°34’S 113°14’W 2590-2600 Calvin, Hobbs, Suzie, Miss
17°35’S Wormwood
18°S 18°24’S 113°24’W 2636-2680 Fromveur, Cathedral, Sojourn,
18°36’S Animal Farm
21°S 21°25’S 114°17’W 2800-2850 Les trois diffuseurs, Bordreaux,
21°34’S Grande cheminée, Krasnov, Gro-
mit, Brandon, Preston.
23°S 23°32’S 115°34’W 2598 Needles
Pacific-Antarctic 31°S 31°09’S 111°55’W 2330 Fred’s Fortress, Nolan’s Nook
Ridge
Saguaro Field 31°51’S 112°02’W 2235
Foundation 37°35’S 110°35’W 2220
German Flats - 38°S 37°47’S 110°55’W 2216 Sebastian’s Steamer, Pâle Etoile,
Annie’s Anthill, Serpulid City
West Pacific Bering Sea Piyp Submarine 55°41’N 167°27’E 352-450
Volcano
Kuril Islands Kraternaya Bight 47°31’N 152°49’E 49
Japan Kagoshima Bay 31°39’N 130°48’E 82-110
Izu Ogasawara Arc Myojin Knoll 32°07’N 139°51’E 1300-1400
(Bonin)
Sumisu Caldera 31°28’N 140°04’E 670-690
Suiyo Seamount 28°34’N 140°39’E 1367
Kaikata Seamount 26°42’N 141°04’E 460-910
Nikko Seamount 23°06’N 142°40’E 430-600
Okinawa Trough Minami-Ensei Knoll 28°24’N 127°38’E 640-720
127°39’E
North Iheya Knoll 27°47’N 126°54’E 990-1070
Iheya Ridge 27°33’N 126°59’E 1400-1430
Izena Cauldron 27°16’N 127°04’E 1310-1580
127°05’E
Hatoma Knoll 28°24’N 123°50’E 1470-1520
Okinawa Arc Northeastern Taiwan 24°50’N 121°60’E 20-180 Tashi fishing grounds
Mariana Mariana Trough 18°12’N 144°42’E 3595
Back-Arc Basin
Anemone Heaven 18°02’N 144°45’E 3676

514
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Ocean Ridge / BAB Field Lat. Long. Depth (m) Sites

Snails Pit
Alice Spring Field 13°23’N 143°55’E 1450 Forecast vent field
Mariana Volcanic Arc Daikoku Seamount 21°19’N 144°12’E 378 Fish Spa, AA, Bluff, Bottomless
pit, Bubbles, Cactus flower, Cra-
ter tuff luck, Fish crack
Kagusa 2 Seamount 21°37’N 143°38’E 387-393 Barnacle Boulders, Mat Ridge,
Crabe slope, Cracked vent, Flat
Bottom, Hairy mat, Hairy rock,
Pinnacle, SW eels, Whale rock,
Yellow overlord
North West Eifuku 21°29’N 144°02’E 1573-1604 More Mussels, Top Towers,
Seamount Champagne, Near Fouling, Fou-
ling Heights, Sulphur spicules,
Bacterial balls, Cliff house, Floc,
Ice cream, Ice Fall, Ice sheet,
Mussel mound, Walter grenade
East Diamante 15°56’N 145°40’E 179-457 Barnacle Beach, Diamante Fo-
Seamount rest, Central Cone (Boulder
Vent), Central Cone (Aquarium),
Fe-Mn Crust, Intense Diffuse,
Floc Storm, Basket case, Five to-
wers, Limpets, Snail flange, Softy
North West Rota 14°36’N 144°46’E 516-599 Brimstone Pit, Cnidaria area,
Seamount Dark sands, Eastern Fault, Fault
Shrimp, Flank vent, Gastros,
High Flow, Iceberg, Loose sands,
Pit Edge, Scarp top, Shimmering
sands, Shimmering vent, Shrim-
p’s peak, Snowcone, Sulfurs
and mats, White mat, White
wall, Yellow granules, Yellow
top
Tabar-Feni Volcanic Edison Seamount 03°01’S 152°03’E 1450
Fore-Arc
Manus Back-Arc Vienna Woods 03°09’S 150°16’E 2500
Basin
Desmos Cauldron 03°41’S 151°52’E 1860-2000 Onsen, Genge-ba
Pacmanus Complex 03°43’S 151°40’E 1700 Field D, Mont Blanc, Field F,
Field E, Black smoker, Chimney
Forest, Field G
Intra-plate Loihi Seamount 18°57’N 155°16’W 969
seamounts
Vailulu’u Seamount 14°13’S 169°04’W 600-1000
North Fiji White Lady 16°59’S 173°55’E 2000 LHOS, White Lady,
Back-Arc Basin Kaiyo Chimney
Mussel Valley 18°49’S 173°29’E 2700
Lau Back-Arc Basin Kilo Moana 20°03’S 176°08’W 2618
Tow Cam 20°06’S 176°34’W 2703
ABE 20°46’S 176°11’W 2148
Tu’i Malila 21°59’S 176°34’W 1887
Vaï Lili 22°13’S 176°37’W 1764-1707
Hine Hina 22°32’S 176°43’W 1832-1887
Kermadec-Arc Macauley Caldera 30°12’S 178°27’E 200-504
Giggenbach 30°02’S 178°43’E 144-175
Seamount
Brothers Seamount 34°52’S 179°04’E 1197
Rumble III Seamount 35°44’S 178°28’E 200
Rumble V Seamount 36°08’S 178.11’E 367-755
Bay of Plenty Calypso Vents 37°39S 177°07’E 159-200

515
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Fig. 1: Atlantic-East
Pacific regions.

Fig. 2: Atlantic-East
Pacific sites.

516
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Fig. 3: Indian-
West Pacific
regions.

Fig. 4: Indian-
West Pacific sites.

517
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Abbreviations, ridges, back-arc basins and vent fields

Abbreviation Ridge, Back-Arc Basin Abbreviation Field

Azores Azores Plateau
CIR Central Indian Ridge EVF Edmond Vent Field
KVF Kairi Vent Field
EPR East Pacific Rise 21°N-23°S
Explorer Southern Explorer Ridge MM Magic Mountain
Galapagos Galapagos Ridge Galapagos Spreading Center
Guaymas Gulf of California Guaymas Basin
Gorda Gorda Ridge
Izu Izu Ogasawara Arc KS Kaikata Seamount
MK Myojin Knoll
NS Nikko Seamount
SS Suijo Seamount
JFR Juan de Fuca Ridge AV-AVF Axial Volcano, Ashes Vent Field
AV-CASM Axial Volcano, CASM Vent Field
CAS CoAxial Segment
ES Endeavour Segment
MV Middle Valley
S-JFR Southern Juan de Fuca Ridge (Cleft Segment)
Kagoshima Japan KB Kagoshima Bay
Kermadec Kermadec Arc BS Brothers Seamount
RIIIS Rumble III Seamount
RVS Rumble V Seamount
Lau Lau Back-Arc Basin VF Valu Fa Ridge
HH Hine Hina
MAR Mid-Atlantic Ridge A1 Ashadze-1
BS Broken Spur
LC Lost City
LO Logatschev
LS Lucky Strike
MG Menez Gwen
MS Mount Saldanha
RB Rainbow
SP Snake Pit
TAG TAG
Mariana Mariana Back-Arc-Basin
Manus Manus Back-Arc Basin
NFiji North Fiji Back-Arc Basin WL White Lady
MV Mussel Valley
Okinawa A Okinawa Arc NET North Eastern Taiwan
Okinawa T Okinawa Trough HK Hatoma Knoll
IR Iheya Ridge
MEK Minami-Ensei Knoll
PAR Pacific-Antarctic Ridge 31°S-38°S
Tabar-Feni Tabar-Feni Volcanic Fore-Arc ES Edison Seamount
Vailulu Intra-plate Seamounts VS Vailulu’u Seamount

M. BRIGHT Denisia 18 (2006): 518

 Species list

Phylum, Class, Order, Family Species Distribution Page

Granuloreticulosa, Foraminifera, Allogromida, Arboramminidae Luffammina atlantica KAMENSKAYA, BAGIROV & SIMDIANOV, 2002 MAR: RB 32
Granuloreticulosa, Foraminifera, Textulariida, Remaneicidae Abyssotherma pacifica BRÖNNIMANN, VAN DOVER & WHITTAKER, 1989 EPR: 21°N, 9°N 33
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae Abyssocladia dominalba VACELET, in press NFiji: WL 37
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae Abyssocladia naudur VACELET, in press EPR: 17°S 38
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae Asbestopluma agglutinans VACELET, in press EPR: 14°S, 18°S 39
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae Asbestopluma (Helophloeina) formosa VACELET, in press NFiji 40
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae Asbestopluma pennatula (SCHMIDT, 1875) MAR: LS; non-vent locations 41
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae Chondrocladia lampadiglobus VACELET, in press EPR: 17°S 42
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae Cladorhiza abyssicola G.O. SARS, 1872 MAR: LO non-vent locations 44
Porifera, Demospongiae, Poecilosclerida, Cladorhizidae Cladorhiza segonzaci VACELET, in press EPR: 17°S 45
Porifera, Demospongiae, Poecilosclerida, Guitarridae Euchelipluma pristina TOPSENT, 1909 MAR: LS, RB 46
Porifera, Hexactinella, Lyssacinosida, Caulophacidae Caulophacus cyanae BOURY-ESNAULT & DE VOS, 1988 EPR: 13°N 47
Cnidaria, Hydrozoa, Anthoathecata, Candelabridae Candelabrum phrygium FABRICIUS, 1780 MAR: LS, RB; non-vent locations 49
Cnidaria, Hydrozoa, Anthoathecata, Candelabridae Candelabrum serpentarii SEGONZAC & VERVOORT, 1995 MAR: SP 50
Cnidaria, Hydrozoa, Anthoathecata, Eudendriidae Eudendrium planum BONNEVIE, 1898 MAR: LO, LS; non-vent locations 51
Cnidaria, Hydrozoa, Anthoathecata, Tubulariidae Ectopleura larynx ELLIS & SOLANDER, 1786 MAR: MG; non-vent locations 52
Cnidaria, Hydrozoa, Leptothecata, Halopterididae Polyplumaria flabellata G.O. SARS, 1874 MAR: MG; non-vent locations 53
Cnidaria, Hydrozoa, Leptothecata, Hebellidae Halisiphonia arctica KRAMP, 1932 MAR: LO 54
Cnidaria, Hydrozoa, Leptothecata, Lafoeidae Grammaria abietina SARS M., 1850 MAR: MG; non-vent locations 55
Cnidaria, Hydrozoa, Leptothecata, Lafoeidae Lafoea dumosa FLEMING, 1820 MAR: RB; non-vent locations 56
Cnidaria, Hydrozoa, Leptothecata, Lafoeidae Zygophylax echinata CALDER & VERVOORT, 1998 MAR: MG 57
Cnidaria, Hydrozoa, Leptothecata, Lafoeidae Zygophylax leloupi RAMIL & VERVOORT, 1992 MAR: RB; non-vent locations 58
Cnidaria, Hydrozoa, Leptothecata, Sertulariidae Hydrallmania falcata LINNAEUS, 1758 MAR: LS; non-vent locations 59
Cnidaria, Hydrozoa, Leptothecata, Sertulariidae Sertularella tenella ALDER, 1856 MAR: LS; non-vent locations 60
Cnidaria, Hydrozoa, Leptothecata, Sertulariidae Symplectoscyphus bathyalis VERVOORT, 1972 MAR: LS; non-vent locations 61
Cnidaria, Hydrozoa, Leptothecata, Tiarannidae Stegolaria geniculata ALLMAN, 1888 MAR: LS, RB; non-vent locations 62
Cnidaria, Hydrozoa, Siphonophora, Rhodaliidae Thermopalia taraxaca PUGH, 1983 EPR, Galapagos 63
Cnidaria, Scyphozoa, Stauromedusae, Eleuthrocarpidae Lucernaria janetae COLLINS & DALY, 2005 EPR: 21°N-7°S 64
Cnidaria, Anthozoa, Actiniaria, Actinostolidae Actinostola VERRILL, 1883 EPR: 13°N 66
Cnidaria, Anthozoa, Actiniaria, Actinostolidae Cyananthea hydrothermala DOUMENC & VAN PRAET, 1986 EPR: 13°N 67
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Cnidaria, Anthozoa, Actiniaria, Actinostolidae Maractis rimicarivora FAUTIN & BARBER, 1999 MAR: A1, SP, TAG 68
Cnidaria, Anthozoa, Actiniaria, Actinostolidae Marianactis bythios FAUTIN & HESSLER, 1989 Mariana 69
Cnidaria, Anthozoa, Actiniaria, Actinostolidae Pacmanactis hashimotoi LÓPEZ-GONZÁLES, RODRIGUEZ & SEGONZAC, 2005 Manus 70
Cnidaria, Anthozoa, Actiniaria, Actinostolidae Paranthosactis denhartogi LÓPEZ-GONZÁLES, RODRIGUEZ & SEGONZAC, 2003 Guaymas 71
Cnidaria, Anthozoa, Actiniaria, Boloceroididae Boloceroides daphneae DALY, in press EPR: 9°N-23°S; PAR: 32°S 72
Cnidaria, Anthozoa, Actiniaria, Hormathiidae Chondrophellia cf. coronata (VERRIL, 1883) EPR: 13°N, 7°S-23°S; PAR: 32°S 73

519
 Phylum, Class, Order, Family Species Distribution Page

520
Mollusca, Solenogastres, Cavibelonia, Simrothiellidae Helicoradomenia acredema SCHELTEMA, 2000 EPR: 21°N; 17°S; Galapagos 76
Mollusca, Solenogastres, Cavibelonia, Simrothiellidae Helicoradomenia bisquama SCHELTEMA, 2000 EPR: 21°N 77
Mollusca, Solenogastres, Cavibelonia, Simrothiellidae Helicoradomenia juani SCHELTEMA & KUZIRIAN, 1991 Exporer, Gorda, JFR: ES 78
Mollusca, Polyplacophora, Neoloricata, Ischnochitonidae Thermochiton undocostatus SAITO & OKUTANI, 1990 Okinawa T: IR 80
Mollusca, Polyplacophora, Neoloricata, Leptochitonidae Leptochiton tenuidontus SAITO & OKUTANI, 1990 Okinawa T: IR 81
Mollusca, Gastropoda, Patelligastropoda, Neolepetopsidae Eulepedopsis vitrea MCLEAN, 1990 EPR: 21°N-17°S; Galapagos 83
Mollusca, Gastropoda, Patelligastropoda, Neolepetopsidae Neolepetopsis densata MCLEAN, 1990 EPR: 13°N; Galapagos 84
Mollusca, Gastropoda, Patelligastropoda, Neolepetopsidae Neolepetopsis gordensis MCLEAN, 1990 EPR: 20°N, 5°S; Gorda, 84
Mollusca, Gastropoda, Patelligastropoda, Neolepetopsidae Neolepetopsis occulta MCLEAN, 1990 EPR: 21°N 84
Mollusca, Gastropoda, Patelligastropoda, Neolepetopsidae Neolepetopsis verruca MCLEAN, 1990 EPR: 21°N 84
Mollusca, Gastropoda, Patelligastropoda, Neolepetopsidae Paralepetopsis ferrugivora WARÉN & BOUCHET, 2001 MAR: LS 85
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Clypeosectus curvus MCLEAN, 1989 Explorer, JFR 86
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Clypeosectus delectus MCLEAN, 1989 EPR: 21°N, 13°N; Galapagos 86
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Gorgoleptis emarginatus MCLEAN, 1988 EPR: 21°N-9°N 87
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Gorgoleptis patulus MCLEAN, 1988 EPR: 13°N; Galapagos 87
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Gorgoleptis spiralis MCLEAN, 1988 EPR: 13°N 87
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus atlanticus WARÉN & BOUCHET, 2001 MAR: 38°N-23°N 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus corrugatus MCLEAN, 1993 JFR 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus cristatus MCLEAN, 1988 EPR: 21°N, 13°N; Galapagos 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus elevatus MCLEAN, 1988 EPR: 21°N-17°S; Galapagos; Lau; Mariana; NFiji 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus fucensis MCLEAN, 1988 JFR 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus galriftensis MCLEAN, 1988 EPR: 9°N; Galapagos 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus guaymasensis MCLEAN, 1988 Guaymas 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus japonicus OKUTANI FUJIKURA & SASAKI, 1993 Okinawa T 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus nux OKUTANI FUJIKURA & SASAKI, 1993 Okinawa T 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus ovalis MCLEAN, 1988 EPR: 21°N-17°S; Galapagos 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus pustulosus MCLEAN, 1988 EPR: 21°N-17°S; Galapagos 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus schrolli BECK, 1993 Manus 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Lepetodrilus tevnianus MCLEAN, 1991 EPR: 11°N 88
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Pseudorimula marianae MCLEAN, 1989 Mariana 91
Mollusca, Gastropoda, Vetigastropoda, Lepetodrilidae Pseudorimula midatlantica MCLEAN, 1992 MAR: 37°N-14°N 91
Mollusca, Gastropoda, Vetigastropoda, Pyropeltidae Pyropelta corymba MCLEAN & HASZPRUNAR, 1987 Guaymas 93
Mollusca, Gastropoda, Vetigastropoda, Pyropeltidae Pyropelta musaica MCLEAN & HASZPRUNAR, 1987 JFR 93
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Mollusca, Gastropoda, Vetigastropoda, Pyropeltidae Pyropelta yamato SASAKI OKUTANI & FUJIKURA, 2003 Izu 93
Mollusca, Gastropoda, Vetigastropoda, Pyropeltidae Pyropelta bohlei BECK, 1996 Tabar-Feni: ES 93
Mollusca, Gastropoda, Vetigastropoda, Skeneidae Bruceiella globulus WARÉN & BOUCHET, 1993 Lau, NFiji 94
Mollusca, Gastropoda, Vetigastropoda, Skeneidae Fucaria mystax WARÉN & BOUCHET, 2001 Tabar-Feni: ES 95
Mollusca, Gastropoda, Vetigastropoda, Skeneidae Fucaria striata WARÉN & BOUCHET, 1993 JFR 95
Mollusca, Gastropoda, Vetigastropoda, Skeneidae Leptogyra inflata WARÉN & BOUCHET, 1993 Lau 96
 Mollusca, Gastropoda, Vetigastropoda, Skeneidae Protolira thorvaldssoni WARÉN, 1996 MAR: SP; non-vent locations 97
Mollusca, Gastropoda, Vetigastropoda, Skeneidae Protolira valvatoides WARÉN & BOUCHET, 1993 MAR: LS, MG 97
Mollusca, Gastropoda, Vetigastropoda, Sutilizonidae Sutilizona pterodon WARÉN & BOUCHET, 2001 MAR: SP 98
Mollusca, Gastropoda, Vetigastropoda, Sutilizonidae Sutilizona theca MCLEAN, 1989 EPR: 13°N 98
Mollusca, Gastropoda, Vetigastropoda, Sutilizonidae Sutilizona tunnicliffae WARÉN & BOUCHET, 2001 JFR: ES 98
Mollusca, Gastropoda, Vetigastropoda, Sutilizonidae Temnocinclis euripes MCLEAN, 1989 JFR 99
Mollusca, Gastropoda, Vetigastropoda, Sutilizonidae Temnozaga parilis MCLEAN, 1989 EPR: 21°N 100
Mollusca, Gastropoda, Vetigastropoda, Trochidae Bathymargarites symplector WARÉN & BOUCHET, 1989 EPR: 13°N, 21°S 101
Mollusca, Gastropoda, Vetigastropoda, Trochidae Helicrenion reticulatum WARÉN & BOUCHET, 1993 Lau: HH 102
Mollusca, Gastropoda, Vetigastropoda, Trochidae Vetulonia phalcata WARÉN & BOUCHET, 1993 NFiji 103
Mollusca, Gastropoda, Neomphalina, Neomphalidae Cyathermia naticoides WARÉN & BOUCHET, 1989 EPR: 21°N-9°N 104
Mollusca, Gastropoda, Neomphalina, Neomphalidae Lacunoides exquisitus WARÉN & BOUCHET, 1989 Galapagos 105
Mollusca, Gastropoda, Neomphalina, Neomphalidae Lacunoides vitreus WARÉN & BOUCHET, 2001 JFR: AV-AVF 105
Mollusca, Gastropoda, Neomphalina, Neomphalidae Melanodrymia aurantiaca HICKMAN, 1984 EPR: 13°N, 7°S-23°S; PAR: 32°S 106
Mollusca, Gastropoda, Neomphalina, Neomphalidae Melanodrymia brightae WARÉN & BOUCHET, 1993 JFR: ES 106
Mollusca, Gastropoda, Neomphalina, Neomphalidae Melanodrymia galeronae WARÉN & BOUCHET, 1993 EPR: 13°N 106
Mollusca, Gastropoda, Neomphalina, Neomphalidae Neomphalus fretterae MCLEAN, 1981 EPR: 21°N-9°N; Galapagos 107
Mollusca, Gastropoda, Neomphalina, Neomphalidae Pachydermia laevis WARÉN & BOUCHET, 1989 EPR: 21°N-17°S 108
Mollusca, Gastropoda, Neomphalina, Neomphalidae Pachydermia sculpta WARÉN & BOUCHET, 1993 Lau, NFiji 108
Mollusca, Gastropoda, Neomphalina, Neomphalidae Planorbidella depressa WARÉN & BOUCHET, 1993 Lau: HH 109
Mollusca, Gastropoda, Neomphalina, Neomphalidae Planorbidella planispira WARÉN & BOUCHET, 1989 EPR: 21°N-17°S 109
Mollusca, Gastropoda, Neomphalina, Neomphalidae Symmetromphalus hageni BECK, 1992 Manus 110
Mollusca, Gastropoda, Neomphalina, Neomphalidae Symmetromphalus regularis MCLEAN, 1990 Mariana 110
Mollusca, Gastropoda, Neomphalina, Peltospiridae Ctenopelta proifera WARÉN & BOUCHET, 1993 EPR: 13°N 111
Mollusca, Gastropoda, Neomphalina, Peltospiridae Depressigyra globulus WARÉN & BOUCHET, 1989 JFR 112
Mollusca, Gastropoda, Neomphalina, Peltospiridae Echinopelta fistulosa MCLEAN, 1989 EPR: 13°N, 21°S 113
Mollusca, Gastropoda, Neomphalina, Peltospiridae Hirtopelta hirta MCLEAN, 1989 EPR: 13°N-21°S 114
Mollusca, Gastropoda, Neomphalina, Peltospiridae Hirtopelta tufari BECK, 2002 EPR: 21°S 114
Mollusca, Gastropoda, Neomphalina, Peltospiridae Lirapex costellata WARÉN & BOUCHET, 2001 MAR: LS 115
Mollusca, Gastropoda, Neomphalina, Peltospiridae Lirapex granularis WARÉN & BOUCHET, 1989 EPR: 21°N-9°N 115
Mollusca, Gastropoda, Neomphalina, Peltospiridae Lirapex humata WARÉN & BOUCHET, 2001 EPR: 21°N 115
Mollusca, Gastropoda, Neomphalina, Peltospiridae Nodopelta heminoda MCLEAN, 1989 EPR: 21°N, 13°N 116
Mollusca, Gastropoda, Neomphalina, Peltospiridae Nodopelta rigneae WARÉN & BOUCHET, 2001 EPR: 13°N 116
Mollusca, Gastropoda, Neomphalina, Peltospiridae Nodopelta subnoda MCLEAN, 1989 EPR: 13°N 116
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Mollusca, Gastropoda, Neomphalina, Peltospiridae Peltospira delicata MCLEAN, 1989 EPR: 13°N-9°N 118
Mollusca, Gastropoda, Neomphalina, Peltospiridae Peltospira lamellifera WARÉN & BOUCHET, 1989 EPR: 13°N 118
Mollusca, Gastropoda, Neomphalina, Peltospiridae Peltospira operculata MCLEAN, 1989 EPR: 21°N-9°N, 17°S 118
Mollusca, Gastropoda, Neomphalina, Peltospiridae Peltospira smaragdina WARÉN & BOUCHET, 2001 MAR: 38°N-15°N 118
Mollusca, Gastropoda, Neomphalina, Peltospiridae Rhynchopelta concentrica MCLEAN, 1989 EPR: 21°N-17°S 119
Mollusca, Gastropoda, Neomphalina, Peltospiridae „scaly foot gastropod“ CIR 120

521
 Phylum, Class, Order, Family Species Distribution Page

522
Mollusca, Gastropoda, Neritimorpha, Phenacolepadidae Olgasolaris tollmanni BECK, 1992 Manus 121
Mollusca, Gastropoda, Neritimorpha, Phenacolepadidae Shinkailepas briandi WARÉN & BOUCHET, 2001 MAR: LO, LS, MG 122
Mollusca, Gastropoda, Neritimorpha, Phenacolepadidae Shinkailepas kaikatensis OKUTANI SAITO & HASHIMOTO, 1989 Izu: KS 122
Mollusca, Gastropoda, Neritimorpha, Phenacolepadidae Shinkailepas mojinensis SASAKI OKUTANI & FUJIKURA, 2003 Izu 122
Mollusca, Gastropoda, Neritimorpha, Phenacolepadidae Shinkailepas tufari BECK, 1992 NFiji 122
Mollusca, Gastropoda, Caenogastropoda, Buccinidae Eosipho auzendei WARÉN & BOUCHET, 2001 EPR: 17°S 124
Mollusca, Gastropoda, Caenogastropoda, Buccinidae Eosipho desbruyeresi nipponensis OKUTANI & OHTA, 1993 Okinawa T 124
Mollusca, Gastropoda, Caenogastropoda, Buccinidae Eosipho desbruyeresi NIPPONENSIS OKUTANI & FUJIWARA, 2000 Lau, Mariana, NFiji 124
Mollusca, Gastropoda, Caenogastropoda, Cerithiopsidae Speculator cariosus WARÉN & BOUCHET, 2001 Exporer: MM 126
Mollusca, Gastropoda, Caenogastropoda, Elachisinidae Laeviphitus desbruyeresi WARÉN & BOUCHET, 2001 MAR: MG-RB 127
Mollusca, Gastropoda, Caenogastropoda, Provannidae Alvinoconcha hessleri OKUTANI & OHTA, 1988 CIR: KVF; Lau; Mariana; NFiji 128
Mollusca, Gastropoda, Caenogastropoda, Provannidae Desbruyeresia cancellata WARÉN & BOUCHET, 1993 Lau, NFiji 129
Mollusca, Gastropoda, Caenogastropoda, Provannidae Desbruyeresia marianaensis OKUTANI, 1990 Manus 129
Mollusca, Gastropoda, Caenogastropoda, Provannidae Desbruyeresia marisindica OKUTANI HASHIMOTO & SASAKI, 2004 CIR 129
Mollusca, Gastropoda, Caenogastropoda, Provannidae Desbruyeresia melanioides WARÉN & BOUCHET, 1993 NFiji 129
Mollusca, Gastropoda, Caenogastropoda, Provannidae Desbruyeresia spinosa WARÉN & BOUCHET, 1993 Lau, Manus 129
Mollusca, Gastropoda, Caenogastropoda, Provannidae Ifremeria nautilei BOUCHET & WARÉN, 1991 Lau, Manus, NFiji 130
Mollusca, Gastropoda, Caenogastropoda, Provannidae Provanna buccinoides WARÉN & BOUCHET, 1993 Lau, NFiji 132
Mollusca, Gastropoda, Caenogastropoda, Provannidae Provanna glabra OKUTANI TSUCHIDA & FUJIKURA, 1992 Okinawa T; non-vent locations 132
Mollusca, Gastropoda, Caenogastropoda, Provannidae Provanna ios WARÉN & BOUCHET, 1986 EPR: 21°N-17°N,13°N; Galapagos 132
Mollusca, Gastropoda, Caenogastropoda, Provannidae Provanna laevis WARÉN & PONDE, 1991 Guaymas; JFR; non-vent locations 132
Mollusca, Gastropoda, Caenogastropoda, Provannidae Provanna muricata WARÉN & BOUCHET, 1986 EPR: 21°N; Galapagos; Lau; NFiji 132
Mollusca, Gastropoda, Caenogastropoda, Provannidae Provanna nassariaeformis OKUTANI, 1990 Mariana, Manus 132
Mollusca, Gastropoda, Caenogastropoda, Provannidae Provanna segonzaci WARÉN & PONDER, 1991 CIR 132
Mollusca, Gastropoda, Caenogastropoda, Provannidae Provanna variabilis WARÉN & BOUCHET, 1986 JFR; non-vent locations 132
Mollusca, Gastropoda, Caenogastropoda, Rissoidae Alvania stenolopha BOUCHET & WARN, 1993 MAR: LS, MG 133
Mollusca, Gastropoda, Caenogastropoda, Vitrinellidae Neusas marshalli SYKES, 1925 MAR: MG 134
Mollusca, Gastropoda, Heterobranchia, Hyalogyrinidae Hyalogyrina globularis WARÉN & BOUCHET, 2001 JFR: ES 135
Mollusca, Gastropoda, Heterobranchia, Hyalogyrinidae Hyalogyrina grasslei WARÉN & BOUCHET, 1993 Guaymas 135
Mollusca, Gastropoda, Heterobranchia, Orbitestellidae Lurifax vitreus WARÉN & BOUCHET, 2001 MAR: MG, LS 136
Mollusca, Gastropoda, Heterobranchia, Xylodisculidae Xylodiscula analoga WARÉN & BOUCHET, 2001 MAR: LS, MG 137
Mollusca, Gastropoda, Heterobranchia, Xylodisculidae Xylodiscula major WARÉN & BOUCHET, 2001 NFiji 137
Mollusca, Gastropoda, Prosobranchia, Turridae Phymorhynchus carinatus WARÉN & BOUCHET, 2001 MAR: 23°N-15°N 138
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Mollusca, Gastropoda, Prosobranchia, Turridae Phymorhynchus hyfifluxi BECK, 1996 NFiji, non-vent locations 138
Mollusca, Gastropoda, Prosobranchia, Turridae Phymorhynchus major WARÉN & BOUCHET, 2001 EPR: 13°N-9°N 138
Mollusca, Gastropoda, Prosobranchia, Turridae Phymorhynchus moskalevi SYSOEV & KANTOR, 1995 MAR: 26°N-23°N 138
Mollusca, Gastropoda, Prosobranchia, Turridae Phymorhynchus ovatus WARÉN & BOUCHET, 2001 MAR: 37°N-15°N 138
Mollusca, Gastropoda, Prosobranchia, Turridae Phymorhynchus starmeri OKUTANI & OHTA, 1993 Manus, NFiji 138
Mollusca, Gastropoda, Prosobranchia, Turridae Phymorhynchus wareni SYSOEV & KANTOR, 1995 Tabar-Feni: ES 138
 Mollusca, Gastropoda, Nudibranchia, Dendronotidae Dendronotus comteti VALDES & BOUCHET, 1998 MAR: LS 140
Mollusca, Bivalvia, Heterodonta, Vesicomyidae Calyptogena edisonensis OKUTANI, KOJIMA & KIM, 2003 Tabar-Feni: ES 142
Mollusca, Bivalvia, Heterodonta, Vesicomyidae Calyptogena extenta KRYLOVA & MOSKALEV, 1996 Gorda; non-vent locations 143
Mollusca, Bivalvia, Heterodonta, Vesicomyidae Calyptogena gigas DALL, 1896 Guaymas, JFR; non-vent locations 144
Mollusca, Bivalvia, Heterodonta, Vesicomyidae Calyptogena magnifica BOSS & TURNER, 1980 EPR: 21°N-23°S; Galapagos 145
Mollusca, Bivalvia, Heterodonta, Vesicomyidae Calyptogena nankaiensis OKUTANI, KOJIMA & ASHI, 1996 Okinawa T: IR; non-vent locations 146
Mollusca, Bivalvia, Heterodonta, Vesicomyidae Calyptogena okutanii KOJIMA & OHTA, 1997 Okinawa T: IR; non-vent locations 147
Mollusca, Bivalvia, Heterodonta, Vesicomyidae Calyptogena solidissima OKUTANI, HASHIMOTO & FUJIKURA, 1992 Okinawa T: MEK 148
Mollusca, Bivalvia, Protobranchia, Solemyidae Acharx alinae MÉTIVIER & COSEL, 1993 Lau: HH 149
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus aduloides HASHIMOTO & OKTUANI, 1994 Okinawa T: MEK, IR; non-vent locations 150
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus azoricus COSEL & COMTET, 1998 MAR: LS, MG, RB 151
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus brevior COSEL, MÉTIVIER & HASHIMOTO, 1994 Lau, NFiji 153
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus elongatus COSEL, MÉTIVIER & HASHIMOTO, 1994 NFiji 154
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus japonicus HASHIMOTO & OKTUANI, 1994 Okinawa T: MEK, IR; non-vent locations 155
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus marisindicus HASHIMOTO, 2001 CIR: KVF 156
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus platifrons HASHIMOTO & OKTUANI, 1994 Okinawa T: IR, Izena Caldron; 157
non-vent locations
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus puteoserpentis COSEL, MÉTIVIER & HASHIMOTO, 1994 MAR: LO, SP 158
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus septemdierum HASHIMOTO & OKTUANI, 1994 Izu: MK, SS 159
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Bathymodiolus thermophilus KENK & WILSON, 1985 EPR: 13°N-21°S 160
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Gigantidas gladius COSEL & MARSHALL, 2003 Kermadec: RIIIS, RVS 162
Mollusca, Bivalvia, Pteriomorphia, Mytilidae Gigantidas horikoshii HASHIMOTO & YAMANE, 2005 Izu: KS 163
Mollusca, Bivalvia, Pteriomorphia, Pectinidae Bathypecten vulcani SCHEIN-FATTON, 1985 EPR: 13°N-9°N; PAR: 32°S, 38°S; Galapagos 164
Mollusca, Bivalvia, Pteromorphia, Pectinidae Sinepecten segonzaci SCHEIN, in press Manus 165
Mollusca, Cephalopoda, Octopoda, Cirroteuthidae Cirrothauma murrayi CHUN, 1911 EPR, non-vent locations 167
Mollusca, Cephalopoda, Octopoda, Cirroteuthidae Cirroteuthis magna HOYLE, 1885 MAR: LO; EPR: 17°S; non-vent locations 168
Mollusca, Cephalopoda, Octopoda, Grimpoteuthidae Grimpoteuthis ROBSON, 1932 EPR, MAR, non-vent locations 169
Mollusca, Cephalopoda, Octopoda, Octopodidae Vulcanoctopus hydrothermalis GONZÁLES, GUERRA, ROCHA & BRIAND, 2002 EPR: 13°N, 23°S 170
Mollusca, Cephalopoda, Octopoda, Octopodidae Benthoctopus GRIMPE, 1921 Gorda, JFR, non-vent locations 171
Mollusca, Cephalopoda, Octopoda, Octopodidae Graneledone JOUBIN, 1918 Explorer, Galapagos, JFR, Kermadec, MAR 172
Nematoda, Adenophorea, Desmodorida, Desmodoridae Desmodora alberti VERSCHELDE, GOURBAULT & VINCX, 1998 Guaymas 174
Nematoda, Adenophorea, Desmodorida, Desmodoridae Desmodora marci VERSCHELDE, GOURBAULT & VINCX, 1998 Lau: HH 175
Nematoda, Adenophorea, Desmodorida, Desmodoridae Desmodorella balteata DECRAEMER & GOURBAULT, 1998 Guaymas 176
Nematoda, Adenophorea, Desmodorida, Desmodoridae Desmodorella spineacaudata DECRAEMER & GOURBAULT, 1998 Guaymas 177
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Nematoda, Adenophorea, Chromadorida, Draconematidae Cephalochaetosoma pacificum notium DECRAEMER & GOURBAULT, 1997 Lau 178
Nematoda, Adenophorea, Chromadorida, Draconematidae Dinetia nycterobia DECRAEMER & GOURBAULT, 1997 EPR: 21°N 179
Nematoda, Secernentea, Spirurida, Cystidicolidae Moravecnema segonzaci JUSTINE, CASSONE & PETTER, 2002 MAR: LO, SP 180
Acanthocephala, Palaeacanthocephala, Echinorhynchida, Hypoechinorhynchus thermaceri BURON, 1988 EPR: 13°N 181
Hypoechinorhynchidae
Nemertini, Hoplonemertini, Monostilifera, family unknown Thermonemertes valens ROGERS, GIBSON & TUNNICLIFFE, 1996 JFR 183

523
 Phylum, Class, Order, Family Species Distribution Page

524
Annelida, Polychaeta, Eunicida, Amphinomidae Archinome rosacea BLAKE, 1985 Galapagos 186
Annelida, Polychaeta, Eunicida, Dorvilleidae Exallopus jumarsi BLAKE, 1985 Guaymas 187
Annelida, Polychaeta, Eunicida, Dorvilleidae Ophryotrocha akessoni BLAKE, 1985 EPR, Galapagos, Guaymas 188
Annelida, Polychaeta, Eunicida, Dorvilleidae Ophryotrocha globopalpata BLAKE & HILBIG, 1990 JFR 189
Annelida, Polychaeta, Eunicida, Dorvilleidae Ophryotrocha platykephale BLAKE, 1985 Guaymas 190
Annelida, Polychaeta, Eunicida, Dorvilleidae Parougia wolfi BLAKE & HILBIG, 1990 JFR 192
Annelida, Polychaeta, Eunicida, Eunicidae Eunice pulvinopalpata FAUCHALD, 1982 EPR: 21°N, 13°N 193
Annelida, Polychaeta, Phyllodocida, Crysopetalidae Thraumastos dieteri WATSON, 2001 Lau, Nfiji; non-vent locations 194
Annelida, Polychaeta, Phyllodocida, Glyceridae Glycera branchiopoda MOORE, 1911 Guaymas; non-vent locations 195
Annelida, Polychaeta, Phyllodocida, Glyceridae Glycera tesselata GRUBE, 1863 MAR: LS; non-vent locations 196
Annelida, Polychaeta, Phyllodocida, Hesionidae Amphiduropsis axialensis (BLAKE & HILBIG, 1990) EPR: 9°N; JFR: AV-AVF, MV 198
Annelida, Polychaeta, Phyllodocida, Hesionidae Hesiodeiria glabra BLAKE & HILBIG, 1990 JFR: AV-AVF, AV-CASM 199
Annelida, Polychaeta, Phyllodocida, Hesionidae Hesiolyra bergi BLAKE, 1985 EPR: 21°N, 9°N 200
Annelida, Polychaeta, Phyllodocida, Hesionidae Hesiospina vestimentifera BLAKE, 1985 EPR; Explorer; JFR; Galapagos 202
Annelida, Polychaeta, Phyllodocida, Hesionidae Nereimyra alvinae BLAKE, 1985 Galapagos; Guaymas 204
Annelida, Polychaeta, Phyllodocida, Hesionidae Sirsoe grasslei (BLAKE, 1985) Guaymas 205
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae Iheyomytilidicola tridentatus MIURA & HASHIMOTO, 1996 Okinawa T 206
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae Mytilidiphila enseiensis MIURA & HASHIMOTO, 1993 Okinawa T 207
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae Mytilidiphila okinawaensis MIURA & HASHIMOTO, 1993 Okinawa T 208
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae Shinkai longipedata MIURA & OHTA, 1991 Okinawa T 209
Annelida, Polychaeta, Phyllodocida, Nautiliniellidae Shinkai semilonga MIURA & HASHIMOTO, 1996 Okinawa T: MEK 210
Annelida, Polychaeta, Phyllodocida, Nereidae Nereis piscesae BLAKE & HILBIG, 1990 JFR 212
Annelida, Polychaeta, Phyllodocida, Nereidae Nereis sandersi BLAKE, 1985 EPR; Galapagos; Guaymas 214
Annelida, Polychaeta, Phyllodocida, Phyllodocidae Eulalia papillosa (BLAKE, 1985) EPR: 21°N 215
Annelida, Polychaeta, Phyllodocida, Phyllodocidae Galapagomystides aristata BLAKE, 1985 EPR: 13°N, 9°N; Galapagos 216
Annelida, Polychaeta, Phyllodocida, Phyllodocidae Protomystides verenae BLAKE & HILBIG, 1990 Explorer, JFR 217
Annelida, Polychaeta, Phyllodocida, Polynoidae Bathykurila guaymasensis PETTIBONE, 1898 Guaymas 218
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchinotogluma hessleri PETTIBONE, 1985 EPR: 9°N; Galapagos; Guaymas 219
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchinotogluma marianus (PETTIBONE, 1989) Manus 221
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchinotogluma sandersi PETTIBONE, 1985 EPR: 21°N-17°S; Galapagos; Guaymas 223
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchinotogluma segonzaci (MIURA & DESBRUYÈRES, 1995) Lau, Manus, NFiji 225
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchinotogluma trifurcus (MIURA & DESBRUYÈRES, 1995) Lau, NFiji 227
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchinotogluma tunnicliffae PETTIBONE, 1988 Explorer, JFR: AV, ES, S-JFR 228
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchiplicatus cupreus PETTIBONE, 1985 EPR: 21°N-9°N; Guaymas 230
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta, Phyllodocida, Polynoidae Branchipolynoe pettibonae MIURA & HASHIMOTO, 1991 Izu, Okinawa T; NFiji, Lau 231
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchipolynoe seepensis PETTIBONE, 1986 MAR: LS; non-vent locations 232
Annelida, Polychaeta, Phyllodocida, Polynoidae Branchipolynoe symmytilida PETTIBONE, 1984 EPR; Galapagos 234
Annelida, Polychaeta, Phyllodocida, Polynoidae Iphionella risensis PETTIBONE, 1986 EPR: 21°N, 9°N 235
Annelida, Polychaeta, Phyllodocida, Polynoidae Lepidonotopodium atalantae DESBRUYÈRES & HOURDEZ, 2000 EPR: 13°N-17°S 236
Annelida, Polychaeta, Phyllodocida, Polynoidae Lepidonotopodium fimbriatum PETTIBONE, 1983 EPR: 21°N-9°N 237
 Annelida, Polychaeta, Phyllodocida, Polynoidae Lepidonotopodium jouinae DESBRUYÈRES & HOURDEZ, 2000 MAR: LS, MG 238
Annelida, Polychaeta, Phyllodocida, Polynoidae Lepidonotopodium piscesae PETTIBONE, 1986 Explorer, Gorda, JFR: AV, ES, S-JFR 239
Annelida, Polychaeta, Phyllodocida, Polynoidae Lepidonotopodium riftense PETTIBONE, 1984 EPR: 21°N-9°N; Galapagos 241
Annelida, Polychaeta, Phyllodocida, Polynoidae Lepidonotopodium williamsae PETTIBONE, 1984 EPR: 21°N-9°N; Galapagos 242
Annelida, Polychaeta, Phyllodocida, Polynoidae Levensteiniella iris HOURDEZ & DESBRUYÈRES, 2003 MAR: LS, RB 243
Annelida, Polychaeta, Phyllodocida, Polynoidae Levensteiniella plicata HOURDEZ & DESBRUYÈRES, 2000 EPR: 9°N 244
Annelida, Polychaeta, Phyllodocida, Polynoidae Thermiphione fijiensis MIURA, 1994 Lau, NFiji 246
Annelida, Polychaeta, Phyllodocida, Polynoidae Thermiphione tufari HARTMANN-SCHRÖDER, 1992 EPR: 7°S-21°S 247
Annelida, Polychaeta, Phyllodocida, Polynoidae Thermopolynoe branchiata MIURA, 1994 Lau, Manus, NFiji 249
Annelida, Polychaeta, Phyllodocida, Polynoidae Vampiropolynoe embleyi MARCUS & HOURDEZ, 2002 JFR: AV 251
Annelida, Polychaeta, Phyllodocida, Syllidae Sphaerosyllis ridgiensis BLAKE & HILBIG, 1990 Explorer, JFR 252
Annelida, Polychaeta, Sabellida, Serpulidae Laminatubus alvini TEN HOVE & ZIBROWIUS, 1986 EPR: 21°N-23°S; Galapagos 253
Annelida, Polychaeta, Sabellida, Serpulidae Protis hydrothermica TEN HOVE & ZIBROWIUS, 1986 EPR: 21°N, 13°N, 17°S; Galapago; PAR: 38°S 254
Annelida, Polychaeta, Sabellida, Siboglinidae Alaysia spiralis SOUTHWARD, 1991 Lau: VF 255
Annelida, Polychaeta, Sabellida, Siboglinidae Arcovestia ivanovi SOUTHWARD & GALGIN, 1997 Lau, Manus 256
Annelida, Polychaeta, Sabellida, Siboglinidae Lamellibrachia barhami WEBB, 1969 JFR: MV; non-vent locations 257
Annelida, Polychaeta, Sabellida, Siboglinidae Lamellibrachia columna SOUTHWARD, 1991 Lau: VF 258
Annelida, Polychaeta, Sabellida, Siboglinidae Lamellibrachia satsuma MIURA, TSUKAHARA & HASHIMOTO, 1997 Japan: Kagoshima; Nankai Trough 259
Annelida, Polychaeta, Sabellida, Siboglinidae Oasisia alvinae JONES, 1985 EPR: 21°N-9°N 260
Annelida, Polychaeta, Sabellida, Siboglinidae Ridgeia piscesae JONES, 1985 Explorer; JFR; Guaymas 261
Annelida, Polychaeta, Sabellida, Siboglinidae Riftia pachyptila JONES, 1981 EPR, Galapagos, Guaymas 262
Annelida, Polychaeta, Sabellida, Siboglinidae Siphonobrachia lauensis SOUTHWARD, 1991 Lau: VF 264
Annelida, Polychaeta, Sabellida, Siboglinidae Tevnia jerichonana JONES, 1985 EPR: 13°N-21°S 265
Annelida, Polychaeta, Scolecida, Maldanidae Nicomache arwidssoni BLAKE, 1985 EPR: 21°N,13°N; Galapagos; Mariana? 266
Annelida, Polychaeta, Scolecida, Maldanidae Nicomache venticola BLAKE & HILBIG, 1990 Guaymas; Explorer; JFR: AV-AVF, S-JFR 267
Annelida, Polychaeta, Scolecida, Orbiniidae Leitoscoloplos pachybranchiatus BLAKE & HILBIG, 1990 JFR 268
Annelida, Polychaeta, Scolecida, Orbiniidae Orbiniella aciculata BLAKE, 1985 Galapagos 269
Annelida, Polychaeta, Scolecida, Orbiniidae Orbiniella hobsonae BLAKE & HILBIG, 1990 MAR: RB, TAG 270
Annelida, Polychaeta, Scolecida, Orbiniidae Scoloplos ehlersi BLAKE, 1985 Galapagos 271
Annelida, Polychaeta, Spionida, Chaetopteridae Spiochaetopterus SARS, 1853 MAR: RB, TAG 272
Annelida, Polychaeta, Spionida, Spionidae Laonice athecata SIGVALDADÓTTIR & DESBRUYÈRES, 2003 MAR: LS, LO 273
Annelida, Polychaeta, Spionida, Spionidae Laubieriellus grasslei MACIOLEK, 1981 Galapagos 274
Annelida, Polychaeta, Spionida, Spionidae Lindaspio dibranchiata BLAKE & MACIOLEK, 1992 Guaymas 275
Annelida, Polychaeta, Spionida, Spionidae Lindaspio southwardorum BLAKE & MACIOLEK, 1992 JFR: MV 276
Annelida, Polychaeta, Spionida, Spionidae Prionospio sandersi MACIOLEK, 1981 Galapagos 278
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta, Spionida, Spionidae Prionospio unilamellata SIGVALDADOTTIR & DESBRUYÈRES, 2003 MAR: LS, RB, SP 280
Annelida, Polychaeta, Spionida, Spionidae Xandaros acanthodes MACIOLEK, 1981 Galapagos 281
Annelida, Polychaeta, Terebellida, Alvinellidae Alvinella caudata DESBRUYÈRES & LAUBIER, 1986 EPR: 21°N-17°S 282
Annelida, Polychaeta, Terebellida, Alvinellidae Alvinella pompejana DESBRUYÈRES & LAUBIER, 1980 EPR: 21°N-23°S 283
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Miralvinella) bacteriocola DESBRUYÈRES & LAUBIER, 1991 Guaymas 285
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Miralvinella) dela DETINOVA, 1988 JFR: AV, ES 286

525
 Phylum, Class, Order, Family Species Distribution Page

526
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Miralvinella) hessleri DESBRUYÈRES & LAUBIER, 1989 Mariana, Manus 287
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Nautalvinella) pandorae DESBRUYÈRES & LAUBIER, 1986 JFR 288
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Nautalvinella) unidentata DESBRUYÈRES & LAUBIER, 1993 Lau, NFiji 289
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Paralvinella) fijiensis DESBRUYÈRES & LAUBIER, 1993 Lau, NFiji 291
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Paralvinella) grasslei DESBRUYÈRES & LAUBIER, 1982 EPR; Galapagos; Guaymas 292
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Paralvinella) palmiformis DESBRUYÈRES & LAUBIER, 1986 Exporer, Gorda, JFR 293
Annelida, Polychaeta, Terebellida, Alvinellidae Paralvinella (Paralvinella) sulfincola DESBRUYÈRES & LAUBIER, 1993 Explorer; JFR; Guaymas 294
Annelida, Polychaeta, Terebellida, Ampharetidae Amathys lutzi DESBRUYÈRES & LAUBIER, 1996 MAR: BS, LS, SP 295
Annelida, Polychaeta, Terebellida, Ampharetidae Amphisamytha galapagensis ZOTTOLI, 1983 Manus, Mariana, NFiji, Okinawa T 296
Annelida, Oligochaeta, Hirudinea, Piscicolidae Bathybdella sawyeri BURRESON, 1981 EPR: 14°S, 17°S; Galapagos 297
Arthropoda, Arachnida, Acariformes, Halacaridae Copidognathus papillatus KRANTZ, 1982 Lau, NFiji 299
Arthropoda, Arachnida, Acariformes, Halacaridae Halacarellus auzendei (BARTSCH, 1990) MAR: BS, LS 300
Arthropoda, Pycnogonida, , Ammotheidae Ammothea verenae CHILD, 1987 Explorer; JFR: AV, ES 301
Arthropoda, Pycnogonida, , Ammotheidae Sericosura cochleifovea CHILD, 1989 Mariana 302
Arthropoda, Pycnogonida, , Ammotheidae Sericosura cyrtoma CHILD & SEGONZAC, 1996 EPR: 13°N 303
Arthropoda, Pycnogonida, , Ammotheidae Sericosura heteroscela CHILD & SEGONZAC, 1996 MAR: LS, LO; MG, RB, SP 304
Arthropoda, Pycnogonida, , Ammotheidae Sericosura mitrata GORDON, 1944 MAR: SP; non-vent locations 305
Arthropoda, Pycnogonida, , Ammotheidae Sericosura venticola CHILD, 1987 JFR: ES 306
Arthropoda, Crustacea, Ostracoda, Halocypridae Archiconchoecia (Archiconchoecia) chavturi KORNICKER & HARRISON-NELSON, 2005 EPR: 9°N 307
Arthropoda, Crustacea, Ostracoda, Halocypridae Bathyconchoecia deeveyae KORNICKER, 1969 Guaymas, non-vent locations 308
Arthropoda, Crustacea, Ostracoda, Halocypridae Bathyconchoecia paulula DEEVEY, 1968 Guaymas, non-vent locations 309
Arthropoda, Crustacea, Ostracoda, Philomedidae Euphilomedes climax KORNICKER, 1991 Explorer; JFR 310
Arthropoda, Crustacea, Ostracoda, Polycopidae Polycopetta pax KORNICKER & HARRISON-NELSON, 2005 EPR: 9°N 311
Arthropoda, Crustacea, Ostracoda, Cylindroleberidae Prionotoleberis styx KORNICKER, 1991 EPR: 21°N 312
Arthropoda, Crustacea, Ostracoda, Pontocyprididae Thomontocypris brightae MADDOCKS, 2006 EPR: 9°N 313
Arthropoda, Crustacea, Ostracoda, Pontocyprididae Thomontocypris gollnerae MADDOCKS, 2006 EPR: 9°N 314
Arthropoda, Crustacea, Ostracoda, Cytheruridae Xylocythere vanharteni MADDOCKS, 2006 EPR: 13°N, 9°N 315
Arthropoda, Crustacea, Copepoda, Aegistidae Andromastax muricatus CONROY-DALTON & HUYS, 1999 Galapagos 318
Arthropoda, Crustacea, Copepoda, Aegistidae Andromastax cephaloceratus LEE & HUYS, 2000 Okinawa T 318
Arthropoda, Crustacea, Copepoda, Ancorabolidae Uptionyx verenae CONROY-DALTON & HUYS, 2000 JFR: MV 319
Arthropoda, Crustacea, Copepoda, Argestidae Argestoides prehensilis HUYS & CONROY-DALTON, 1997 Galapagos 320
Arthropoda, Crustacea, Copepoda, Argestidae Jamstecia terazakii LEE & HUYS, 2000 Okinawa T 321
Arthropoda, Crustacea, Copepoda, Argestidae Scabrantenna yooi LEE & HUYS, 2000 Okinawa T 322
Arthropoda, Crustacea, Copepoda, Asterocheridae Cheramomyzon abyssale HUMES, 1989 EPR: 13°N 323
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Copepoda, Asterocheridae Collocherides brychius HUMES, 1999 JFR: S-JFR 324
Arthropoda, Crustacea, Copepoda, Clausidiidae Hyphalion captans HUMES, 1987 Guaymas 325
Arthropoda, Crustacea, Copepoda, Cyclopinidae Barathricola rimensis HUMES, 1999 JFR: CAS 326
Arthropoda, Crustacea, Copepoda, Cyclopinidae Heptnerina confusa IVANENKO & DEFAY, 2004 MAR: LS 327
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius acanthinus HUMES & LUTZ, 1994 Galapagos 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius arcuatus HUMES, 1987 EPR: 9°N; Galapagos 328
 Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius atlanteus HUMES, 1996 MAR: LS, MG 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius baculigerus HUMES, 1987 EPR: 21°N; Galapagos 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius flexispina HUMES, 1987 EPR: 13°N 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius forcipatus HUMES, 1987 Gorda; JFR; Guaymas 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius hydronauticus HUMES, 1989 EPR: 13°N 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius limatulus HUMES, 1987 EPR: 21°N, 9°N; Galapagos 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius mammillatus HUMES, 1987 EPR: 13°N, 21°N, 13°N; Galapagos; Guaymas 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Aphotopontius probolus HUMES, 1990 Galapagos 328
Arthropoda, Crustacea, Copepoda, Dirivultidae Benthoxynus spiculifer HUMES, 1984 Exporer, Gorda, JFR: AV 330
Arthropoda, Crustacea, Copepoda, Dirivultidae Benthoxynus tumidiseta HUMES, 1989 EPR: 13°N 330
Arthropoda, Crustacea, Copepoda, Dirivultidae Ceuthoecetes acanthothrix HUMES, 1987 EPR; Galapagos 331
Arthropoda, Crustacea, Copepoda, Dirivultidae Ceuthoecetes aliger HUMES & DOJIRI, 1980 EPR; Galapagos 331
Arthropoda, Crustacea, Copepoda, Dirivultidae Ceuthoecetes cristatus HUMES, 1987 EPR; Galapagos 331
Arthropoda, Crustacea, Copepoda, Dirivultidae Ceuthoecetes introversus HUMES, 1987 EPR; Galapagos 331
Arthropoda, Crustacea, Copepoda, Dirivultidae Chasmatopontius thescalus HUMES, 1990 Lau, Mariana 332
Arthropoda, Crustacea, Copepoda, Dirivultidae Dirivultus spinigulatus HUMES, 1999 Tabar-Feni: ES 333
Arthropoda, Crustacea, Copepoda, Dirivultidae Exrima dolichopus HUMES, 1987 EPR: 13°N 334
Arthropoda, Crustacea, Copepoda, Dirivultidae Exrima singular HUMES, 1987 EPR: 21°N 334
Arthropoda, Crustacea, Copepoda, Dirivultidae Fissuricola caritus HUMES, 1987 EPR: 21°N 335
Arthropoda, Crustacea, Copepoda, Dirivultidae Humesipontius arthuri IVANENKO & FERRARI, 2003 JFR 336
Arthropoda, Crustacea, Copepoda, Dirivultidae Nilva torifera HUMES, 1987 EPR: 21°N, 13°N; Galapagos 337
Arthropoda, Crustacea, Copepoda, Dirivultidae Rhogobius contractus HUMES, 1987 EPR: 21°N, 13°N; Galapagos 338
Arthropoda, Crustacea, Copepoda, Dirivultidae Rhogobius pressulus HUMES, 1989 Galapagos 338
Arthropoda, Crustacea, Copepoda, Dirivultidae Rhogobius rapunculus (HUMES & SEGONZAC, 1998) EPR: 9°N 338
Arthropoda, Crustacea, Copepoda, Dirivultidae Rimipontius mediospinifer HUMES, 1996 MAR: BS, LO, SP 339
Arthropoda, Crustacea, Copepoda, Dirivultidae Scotoecetes introrsus HUMES, 1987 EPR: 13°N, 9°N 340
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius appositus HUMES, 1989 EPR: 21°N, 13°N 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius brevispina HUMES, 1991 Lau 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius cinctiger HUMES, 1987 EPR: 21°N, 13°N, 9°N 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius cladarus HUMES, 1996 MAR: BS, TAG 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius flexus HUMES, 1987 EPR: 9°N; Guaymas 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius hispidulus HUMES, 1987 EPR: 21°N, 13°N, 9°N 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius latulus HUMES, 1996 MAR: SP 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius lauensis HUMES, 1991 Lau 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius mirus HUMES, 1996 EPR: 9°N; MAR: SP 341
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius mucroniferus HUMES, 1987 EPR: 9°N; Guaymas 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius paxillifer HUMES, 1989 EPR: 21°N, 9°N 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius pectinatus HUMES, 1987 Mariana, MAR: BS, SP, TAG 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius quadrispinosus HUMES, 1987 Gorda, Explorer, JFR 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius regius HUMES, 1996 MAR: SP 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius rimivagus HUMES, 1997 MAR: LS 341

527
 Phylum, Class, Order, Family Species Distribution Page

528
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius sentifer HUMES, 1987 EPR: 21°N, 13°N, 9°N 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius serratus HUMES, 1996 MAR: SP 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius stabilitus HUMES, 1990 EPR: 13°N, Mariana 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius teres HUMES, 1996 MAR: SP 341
Arthropoda, Crustacea, Copepoda, Dirivultidae Stygiopontius verruculatus HUMES, 1987 EPR: 21°N, 9°N 341
Arthropoda, Crustacea, Copepoda, Ecbathyriontidae Ecbathyrion prolixicauda HUMES, 1987 EPR: 21°N, 13°N, 9°N; Galapagos 343
Arthropoda, Crustacea, Copepoda, Erebonasteridae Ambilimbus altalis (HUMES & HUYS, 1992) JFR 344
Arthropoda, Crustacea, Copepoda, Erebonasteridae Ambilimbus arcuscelestis IVANENKO DEFAYE & HUYS, 2005 MAR: RB 344
Arthropoda, Crustacea, Copepoda, Erebonasteridae Ambilimbus tuerkayi (MARTINEZ-ARBIZU, 1999) NFiji 344
Arthropoda, Crustacea, Copepoda, Laophontidae Bathylaophonte azorica LEE & HUYS, 1999 MAR: LS, MG 345
Arthropoda, Crustacea, Copepoda, Laophontidae Bathylaophonte pacifica LEE & HUYS, 1999 EPR: 17°S 345
Arthropoda, Crustacea, Copepoda, Lubbockiidae Laitmatobius crinitus HUMES, 1987 Guaymas 346
Arthropoda, Crustacea, Copepoda, Megapontiidae Hyalopontius boxshalli HUMES, 1988 Galapagos 347
Arthropoda, Crustacea, Copepoda, Miraciidae Stenhelia gundulae WILLEN, 2003 Tabar-Feni: ES 348
Arthropoda, Crustacea, Copepoda, Misophriidae Misophriopsis longicaudata BOXSHALL, 1983 JFR: CAS 349
Arthropoda, Crustacea, Copepoda, Oncaeidae Oncaea praeclara HUMES, 1988 EPR: 21°N, 13°N, 9°N; Galapagos; Guaymas 350
Arthropoda, Crustacea, Copepoda, Scolecitrichidae Grievella shanki FERRARI & MARKHASEVA, 2000 EPR: 21°S 351
Arthropoda, Crustacea, Copepoda, Spinocalanidae Isaacsicalanus paucisetus FLEMINGER, 1983 EPR: 21°N 352
Arthropoda, Crustacea, Copepoda, Tegastidae Smacigastes micheli IVANENKO & DEFAYE, 2004 MAR: LS 353
Arthropoda, Crustacea, Copepoda, Tisbidae Cholidya polypi FARRAN, 1914 JFR: MV; non-vent locations 354
Arthropoda, Crustacea, Copepoda, Tisbidae Genesis vulcanoctopusi LÓPEZ-GONZALEZ, BRESCIANI & HUYS, 2000 EPR: 13°N 355
Arthropoda, Crustacea, Cirripedia, Chionelasmatoidae Eochionelasmus ohtai YAMAGUCHI, 1990 NFiji, Lau, Manus 358
Arthropoda, Crustacea, Cirripedia, Chionelasmatoidae Eochionelasmus paquensis YAMAGUCHI & NEWMAN, 1997 EPR: 17°S 359
Arthropoda, Crustacea, Cirripedia, Eolepadidae Ashinkailepas seepiophilia YAMAGUCHI, NEWMAN & HASHIMOTO, 2004 Sagami Bay (vent part) 360
Arthropoda, Crustacea, Cirripedia, Eolepadidae Leucolepas longa SOUTHWARD & JONES, 2003 Tabar-Feni: ES 361
Arthropoda, Crustacea, Cirripedia, Eolepadidae Neolepas rapanuii JONES, 1993 EPR: 23°S 362
Arthropoda, Crustacea, Cirripedia, Eolepadidae Neolepas zevinae NEWMAN, 1979 EPR: 21°N-9°N 363
Arthropoda, Crustacea, Cirripedia, Eolepadidae Vulcanolepas osheai (BUCKERIDGE, 2000) Kermadec: BS 364
Arthropoda, Crustacea, Cirripedia, Eolepadidae Vulcanolepas parensis SOUTHWARD, 2005 PAR: 37°S-38°S 365
Arthropoda, Crustacea, Cirripedia, Neobrachylepadidae Neobrachylepas relica NEWMAN & YAMAGUCHI, 1995 Lau 366
Arthropoda, Crustacea, Cirripedia, Neoverrucidae Imbricaverruca yamaguchii NEWMAN, 2000 Lau: HH 367
Arthropoda, Crustacea, Cirripedia, Neoverrucidae Neoverruca brachylepadoformis NEWMAN, 1989 Mariana 368
Arthropoda, Crustacea, Leptostraca, Nebaliidae Dahlella caldariensis HESSLER, 1984 EPR: 21°N-18°S; Galapagos 369
Arthropoda, Crustacea, Cumacea, Bodotriidae Atlantocuma bidentatum LEDOYER, 1988 EPR: 13°N; non-vent locations 370
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Cumacea, Bodotriidae Bathycuma breviostre (NORMAN, 1879) MAR: LS 371
Arthropoda, Crustacea, Tanaidacea, Fam. indet. Armaturatanais atlanticus LARSEN, BLAZEWICZ-PASZKOWYCZ & CUHNA, in press MAR: LS 373
Arthropoda, Crustacea, Tanaidacea, Colleteidae Leptognathiella fragilis LARSEN, BLAZEWICZ-PASZKOWYCZ & CUHNA, in press MAR: LS 374
Arthropoda, Crustacea, Tanaidacea, Leptocheliidae Mesotanais styxis LARSEN, BLAZEWICZ-PASZKOWYCZ & CUHNA, in press MAR: LS 375
Arthropoda, Crustacea, Tanaidacea, Nototanaidae Obesutanais sigridi LARSEN, BLAZEWICZ-PASZKOWYCZ & CUHNA, in press MAR: LS 376
Arthropoda, Crustacea, Tanaidacea, Nototanaidae Typhlotanais incognitus LARSEN, BLAZEWICZ-PASZKOWYCZ & CUHNA, in press MAR: LS 377
 Arthropoda, Crustacea, Tanaidacea, Pseudotanaidae Pseudotanais vulsella BIRD & HOLDICH, 1989 MAR: LS; non-vent locations 378
Arthropoda, Crustacea, Isopoda, Ischnomesidae Heteromesus calcar CUNHA & WILSON, in press MAR: LS 381
Arthropoda, Crustacea, Isopoda, Ischnomesidae Heteromesus ctenobasius CUNHA & WILSON, in press MAR: LS 380
Arthropoda, Crustacea, Amphipoda, Ampeliscidae Ampelisca romigi BARNARD, 1954 EPR, Guaymas, non-vent locations 383
Arthropoda, Crustacea, Amphipoda, Amphilochidae Gitanopsis alvina BELLAN-SANTINI & THURSTON, 1996 MAR: LS 384
Arthropoda, Crustacea, Amphipoda, Aoridae Autonoe longicornis CHEVREUX, 1909 MAR: LS 385
Arthropoda, Crustacea, Amphipoda, Calliopiidae Oradarea longimana (BOECK, 1871) JFR; non-vent locations 386
Arthropoda, Crustacea, Amphipoda, Caprellidae Caprella bathytatos MARTIN & PETTIT, 1998 JFR 387
Arthropoda, Crustacea, Amphipoda, Eusiridae Bouvierella curtirama BELLAN-SANTINI & THURSTON, 1996 MAR: LS 388
Arthropoda, Crustacea, Amphipoda, Eusiridae Luckia striki BELLAN-SANTINI & THURSTON, 1996 MAR: LS 389
Arthropoda, Crustacea, Amphipoda, Ischyroceridae Bonnierella compar MYERS & CUNHA, 2004 MAR: LS 390
Arthropoda, Crustacea, Amphipoda, Ischyroceridae Bonnierella linearis BARNARD, 1964 JFR, Explorer, non-vent locations 391
Arthropoda, Crustacea, Amphipoda, Lysianassidae Apotectonia heterostegos BARNARD & INGRAM, 1990 Galapagos 392
Arthropoda, Crustacea, Amphipoda, Lysianassidae Cyclocaris tahitensis STEBBING, 1888 Guaymas, non-vent locations 393
Arthropoda, Crustacea, Amphipoda, Lysianassidae Euonyx mytilus BARNARD & INGRAM, 1990 EPR:13°N, Galapagos 394
Arthropoda, Crustacea, Amphipoda, Lysianassidae Hirondellea glutonis BARNARD & INGRAM, 1990 Galapagos, EPR 395
Arthropoda, Crustacea, Amphipoda, Lysianassidae Orchomene (Abyssorchomene) abyssorum STEBBING, 1888 Galapagos, non-vent locations 396
Arthropoda, Crustacea, Amphipoda, Lysianassidae Orchomene (Abyssorchomene) distinctus BIRSTEIN & VINOGRADOV, 1960 EPR: 13°N, non-vent locations 397
Arthropoda, Crustacea, Amphipoda, Lysianassidae Tectovalopsis diabolus BARNARD & INGRAM, 1990 EPR: 13°N 398
Arthropoda, Crustacea, Amphipoda, Lysianassidae Tectovalopsis wegeneri BARNARD & INGRAM, 1990 EPR: 13°N 399
Arthropoda, Crustacea, Amphipoda, Lysianassidae Transtectonia torrentis BARNARD & INGRAM, 1990 EPR:13°N 400
Arthropoda, Crustacea, Amphipoda, Oedicerotidae Bathymedon curtipalpus VINOGRADOV, 1993 EPR
Arthropoda, Crustacea, Amphipoda, Pardaliscidae Halice hesmonectes MARTIN, FRANCE & VAN DOVER, 1993 EPR: 9°N 401
Arthropoda, Crustacea, Amphipoda, Pardaliscidae Pardalisca endeavouri SHAW, 1989 Explorer 402
Arthropoda, Crustacea, Amphipoda, Phoxocephalidae Harpiniopsis fulgens BARNARD, 1960 EPR, Guaymas
Arthropoda, Crustacea, Amphipoda, Sebidae Seba profundus SHAW, 1989 Explorer 403
Arthropoda, Crustacea, Amphipoda, Stegocephalidae Steleuthera ecoprophycea BELLAN-SANTINI & THURSTON, 1996 MAR: LS 404
Arthropoda, Crustacea, Amphipoda, Stenothoidae Metopa (Prometopa) samsiluna BARNARD, 1966 Guaymas, non-vent locations 405
Arthropoda, Crustacea, Amphipoda, Stenothoidae Stenothoe menezgweni BELLAN-SANTINI, 2005 MAR: MG 406
Arthropoda, Crustacea, Amphipoda, Stenothoidae Torometopa saldanhae BELLAN-SANTINI, 2005 MAR: MG; LS; RB 407
Arthropoda, Crustacea, Amphipoda, Uristidae Ventiella sulfuris BARNARD & INGRAM, 1990 Galapagos; EPR 408
Arthropoda, Crustacea, Euphausiacea, Euphausiidae Thysanoessa parva HANSEN, 1905 MAR: RB; non-vent locations 409
Arthropoda, Crustacea, Decapoda, Nematocarcinidae Nematocarcinus burukovskyi KOMAI & SEGONZAC, 2005 EPR: 17°S, 23°S; PAR: 31°S 411
Arthropoda, Crustacea, Decapoda, Alvinocarididae Alvinocaris brevitelsonis KIKUCHI & HASHIMOTO, 2000 Okinawa T: MEK 413
Arthropoda, Crustacea, Decapoda, Alvinocarididae Alvinocaris dissimilis KOMAI & SEGONZAC, 2005 Okinawa T: MEK 414
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Decapoda, Alvinocarididae Alvinocaris longirostris KIKUCHI & OHTA, 1995 Kermadec: BS; Okinawa T: IR, HK; 415
non-vent locations
Arthropoda, Crustacea, Decapoda, Alvinocarididae Alvinocaris lusca WILLIAMS & CHACE, 1982 EPR: 9°N; Galapagos 416
Arthropoda, Crustacea, Decapoda, Alvinocarididae Alvinocaris markensis WILLIAMS, 1988 MAR: BS, LO, LS, RB, SP, TAG 417
Arthropoda, Crustacea, Decapoda, Alvinocarididae Alvinocaris niwa WEBBER, 2004 Kermadec: BS, RVS 419
Arthropoda, Crustacea, Decapoda, Alvinocarididae Alvinocaris williamsi SHANK & MARTIN, 2003 MAR: MG 420

529
 Phylum, Class, Order, Family Species Distribution Page

530
Arthropoda, Crustacea, Decapoda, Alvinocarididae Chorocaris chacei (WILLIAMS & RONA, 1986) MAR: LO, LS, RB, SP, TAG 421
Arthropoda, Crustacea, Decapoda, Alvinocarididae Chorocaris paulexa MARTIN & SHANK, 2005 EPR: 17°S 422
Arthropoda, Crustacea, Decapoda, Alvinocarididae Chorocaris vandoverae (MARTIN & HESSLER, 1990) Lau, Mariana, Manus, NFiji 423
Arthropoda, Crustacea, Decapoda, Alvinocarididae Mirocaris fortunata (MARTIN & CHRISTIANSEN, 1995) MAR: BS, LO, LS, MG, RB, SP, TAG 424
Arthropoda, Crustacea, Decapoda, Alvinocarididae Mirocaris indica KOMAI, MARTIN, ZALA, TSUCHIDA & HASHIMOTO, in press CIR: KVF, EVF 425
Arthropoda, Crustacea, Decapoda, Alvinocarididae Nautilocaris saintlaurentae KOMAI & SEGONZAC, 2005 Lau, NFiji 426
Arthropoda, Crustacea, Decapoda, Alvinocarididae Opaepele loihi WILLIAMS & DOBBS, 1995 Hawaii: Loihi Seamount 427
Arthropoda, Crustacea, Decapoda, Alvinocarididae Rimicaris exoculata WILLIAMS & RONA, 1986 MAR: LS, RB, SP, TAG 428
Arthropoda, Crustacea, Decapoda, Alvinocarididae Rimicaris kairei WATABE & HASHIMOTO, 2002 CIR: KVF, EVF 429
Arthropoda, Crustacea, Decapoda, Alvinocarididae Shinkaicaris leurokolos (KIKUCHI & HASHIMOTO, 2000) Okinawa T: MEK 430
Arthropoda, Crustacea, Decapoda, Hippolytidae Lebbeus carinatus DE SAINT LAURENT, 1984 EPR: 13°N 431
Arthropoda, Crustacea, Decapoda, Nephropidae Thymopides laurentae SEGONZAC & MACPHERSON, 2004 MAR: SP 433
Arthropoda, Crustacea, Decapoda, Parapaguridae Paragiopagurus ventilatus LEMAITRE, 2004 Okinawa A: NET 435
Arthropoda, Crustacea, Decapoda, Lithodidae Paralomis hirtella DE SAIN LAURENT & MACPHERSON, 1997 Lau: HH; NFiji: WL 436
Arthropoda, Crustacea, Decapoda, Lithodidae Paralomis verrilli (BENEDICT, 1895) JFR: AV; non-vent locations 437
Arthropoda, Crustacea, Decapoda, Galatheidae Munida magniantennulata BABA & TÜRKAY, 1992 Lau: HH 438
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis acutispina BENEDICT, 1902 MAR: LC; non-vent locations 439
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis alvisca WILLIAMS, 1988 Explorer: MM; Guaymas; JFR 440
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis diomedeae (FAXON, 1893) EPR 441
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis exuta MACPHERSON & SEGONZAC, 2005 MAR: SP, TAG 442
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis lauensis BABA & DE SAINT LAURENT, 1992 Lau: VF, HH; NFiji: WL 443
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis lentigo WILLIAMS & VAN DOVER, 1983 EPR: 21°N 444
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis marianica WILLIAMS & BABA, 1989 Manus 445
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis sonne BABA, 1995 NFiji 446
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis starmer BABA & DE SAINT LAURENT, 1992 NFiji: WL 447
Arthropoda, Crustacea, Decapoda, Galatheidae Munidopsis WHITEAVES, 1874 Galapagos; EPR 448
Arthropoda, Crustacea, Decapoda, Galatheidae Shinkaia crosnieri BABA & WILLIAMS, 1998 Okinawa A: NET; Okinawa T: IR, NIK; Tabar-Feni: ES; 449
Ryuku Islands
Arthropoda, Crustacea, Decapoda, Kiwaidae Kiwa hirsuta MACPHERSON, JONES & SEGONZAC, 2005 PAR: 38°S 450
Arthropoda, Crustacea, Decapoda, Chirostylidae Uroptychus bicavus BABA & DE SAINT LAURENT, 1992 NFiji: MV 452
Arthropoda, Crustacea, Decapoda, Chirostylidae Uroptychus edisonicus BABA & WILLIAMS, 1998 Tabar-Feni: ES 453
Arthropoda, Crustacea, Decapoda, Chirostylidae Uroptychus thermalis BABA & DE SAINT LAURENT, 1992 NFiji: WL; non-vent locations 454
Arthropoda, Crustacea, Decapoda, Majidae Macroregonia macrochira SAKAI, 1978 Explorer, JFR, non-vent locations 456
Arthropoda, Crustacea, Decapoda, Portunidae Bathynectes maravigna (PRESTANDREA, 1839) MAR: LS, MG; non-vent locations 457
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Crustacea, Decapoda, Geryonidae Chaceon affinis MILNE-EDWARDS & BOUVIER, 1894 MAR: MG, LS; non-vent locations 458
Arthropoda, Crustacea, Decapoda, Bythograeidae Allograea tomentosa GUINOT, HURTADO & VRIJENHOEK, 2002 PAR: 31°S 459
Arthropoda, Crustacea, Decapoda, Bythograeidae Austinograea alayseae GUINOT, 1990 Lau: HH, VL; Manus; NFiji: WL 460
Arthropoda, Crustacea, Decapoda, Bythograeidae Austinograea rodriguezensis TSUCHIDA & HASHIMOTO, 2002 CIR: KVF, EVF 462
Arthropoda, Crustacea, Decapoda, Bythograeidae Austinograea williamsi HESSLER & MARTIN, 1989 Mariana 463
Arthropoda, Crustacea, Decapoda, Bythograeidae Austinograea yunohana TAKEDA, HASHIMOTO & OHTA, 2000 Izu 464
 Arthropoda, Crustacea, Decapoda, Bythograeidae Bythograea galapagensis GUINOT & HURTADO, 2003 Galapagos 465
Arthropoda, Crustacea, Decapoda, Bythograeidae Bythograea intermedia DE SAINT LAURENT, 1988 Galapagos
Arthropoda, Crustacea, Decapoda, Bythograeidae Bythograea laubieri GUINOT & SEGONZAC, 1997 EPR: 11°S-23°S; PAR: 31°S 466
Arthropoda, Crustacea, Decapoda, Bythograeidae Bythograea microps DE SAINT LAURENT, 1989 EPR: 21°N-23°S; PAR: 31°S-38°S; Galapagos 468
Arthropoda, Crustacea, Decapoda, Bythograeidae Bythograea thermydron WILLIAMS, 1980 EPR: 21°N-18°S; Galapagos 469
Arthropoda, Crustacea, Decapoda, Bythograeidae Bythograea vrijenhoeki GUINOT & HURTADO, 2003 PAR: 31°S-38°S 470
Arthropoda, Crustacea, Decapoda, Bythograeidae Cyanagraea preadator DE SAINT LAURENT, 1984 EPR: 21°N, 13°N, 14°S, 18°S 472
Arthropoda, Crustacea, Decapoda, Bythograeidae Segonzacia mesatlantica (WILLIAMS, 1988) MAR: LO, LS, MG, RB, SP, TAG 473
Arthropoda, Crustacea, Decapoda, Varunidae Xenograpsus testudinatus NG, HUANG & HO, 2000 Okinawa A: NET 474
Echinodermata, Crinoidea, Cyrtocrinida, Hyocrinidae Hyocrinus biscoitoi ROUX, 2004 EPR: 13°N; non-vent locations 475
Echinodermata, Crinoidea, Cyrtocrinida, Hyocrinidae Laubiericrinus pentagonalis ROUX, 2004 NFiji; non-vent locations 476
Echinodermata, Asteroidea, Brisingida, Brisingidae Brisinga endecacnemos ASBJØRNSEN, 1856 MAR: LO; non-vent locations 477
Echinodermata, Asteroidea, Brisingida, Freyellidae Freyella PERRIER, 1885 EPR: 13°N; PAR; Lau, NFiji 478
Echinodermata, Echinoidea, Echinidae Echinus alexandri DANIELSSEN & KOREN, 1883 MAR: LS; non-vent locations 479
Echinodermata, Holothuroidea, Chiridotidae Chiridota hydrothermica SMIRNOV & GEBRUK, 2000 EPR: 17°S-21°S; Manus, NFiji 480
Echinodermata, Ophiuroidea, Ophiurida, Ophiactidae Ophiactis tyleri STÖHR & SEGONZAC, 2005 MAR: MG, 38°N 482
Echinodermata, Ophiuroidea, Ophiurida, Ophiuridae Ophioctenella acies TYLER et al., 1995 MAR; non-vent locations 483
Echinodermata, Ophiuroidea, Ophiurida, Ophiuridae Spinophiura jolliveti STÖHR & SEGONZAC, 2006 EPR: 13°N-9°N, 17°S-18°S; PAR: 38°S 484
Echinodermata, Ophiuroidea, Ophiurida, Ophiacanthidae Ophiolamina eprae STÖHR & SEGONZAC, 2006 EPR: 13°N, 9°N, 17°S-18°S 485
Chaetognatha, Spadellidae Calispadella alata CSANOVA & MOREAU, 2005 MAR: LS 486
Hemichordata, Enteropneusta, , Saxipendiidae Saxipendium coronatum WOODWICK & SENSENBAUGH, 1985 EPR: 21°N-17°S; PAR: 38°S; Galapagos 487
Chordata, Chondrichthyes, Chimaeriformes, Chimaeridae Hydrolagus affinis (CAPELLO, 1868) MAR: LS, MS, RB; non-vent locations 491
Chordata, Chondrichthyes, Chimaeriformes, Chimaeridae Hydrolagus pallidus HARDY & STEHMANN, 1990 MAR: LS, Menez Hom, MS; RB; non-vent locations 492
Chordata, Osteichthyes, Anguilliformes, Synaphobranchidae Dysommina rugosa GINSBURG, 1951 Intra-plate Seamount: Vailulu 493
Chordata, Osteichthyes, Anguilliformes, Synaphobranchidae Ilyophis saldanhai KARMOVSKAYA & PARIN, 1999 EPR: 21°S; MAR: BS 494
Chordata, Osteichthyes, Gadiformes, Synaphobranchidae Thermobiotes mytilogeiton GEISTDORFER, 1991 Lau: VV 496
Chordata, Osteichthyes, Gadiformes, Lotidae Gaidropsarus RAFINESQUE, 1810 MAR: LS 497
Chordata, Osteichthyes, Gadiformes, Moridae Lepidion schmidti SVETOVIDOV, 1936 MAR: MG; non-vent locations 498
Chordata, Osteichthyes, Ophidiiformes, Bythitidae Cataetyx laticeps KOEFOED, 1927 MAR: MG, MS, Menez Hom, LS, RB; non-vent locations 499
Chordata, Osteichthyes, Ophidiiformes, Bythitidae Thermichthys hollisi (COHEN, ROSENBLATT & MOSER, 1990) EPR: 17°S, 21°S; Galapagos 500
Chordata, Osteichthyes, Ophidiiformes, Ophidiidae Ventichthys biospeedoi NIELSEN, MOLLER & SEGONZAC, in press EPR: 17°S 501
Chordata, Osteichthyes, Perciformes, Zoarcidae Pachycara gymninium ANDERSON & PEDEN, 1988 JFR: AV, ES; non-vent locations 502
Chordata, Osteichthyes, Perciformes, Zoarcidae Pachycara rimae ANDERSON, 1989 Galapagos 503
Chordata, Osteichthyes, Perciformes, Zoarcidae Pachycara saldani BISCOITO & ALMEIDA, 2004 MAR: RB 504
Chordata, Osteichthyes, Perciformes, Zoarcidae Pachycara thermophilum GEISTDORFER, 1994 MAR: SP, TAG 505
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Chordata, Osteichthyes, Perciformes, Zoarcidae Pyrolycus manusanus MACHIDA & HASHIMOTOT, 2002 Manus 506
Chordata, Osteichthyes, Perciformes, Zoarcidae Thermarces cerberus ROSENBLATT & COHEN, 1986 EPR: 21°N,13°N, 9°N; Galapagos 507
Chordata, Osteichthyes, Scorpaeniformes, Liparidae Careproctus hyaleius GEISTDORFER, 1994 EPR: 13°N-9°N 509
Chordata, Osteichthyes, Scorpaeniformes, Sebastidae Trachyscorpia cristulata echinata (KOEHLER, 1896) MAR: MG; non-vent locations 510
Chordata, Osteichthyes, Gadiformes, Macrouridae Coryphaenoides armatus HECTOR, 1875 EPR: 9°N; Galapagos; MAR: LS, RB, SP; 511
non vent locations

531
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Cnidaria, Anthozoa

532
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Mollusca, Gastropoda

533
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Mollusca, Bivalvia

534
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Mollusca, Cephalopoda

535
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta

536
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Annelida, Polychaeta

537
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Amphipoda

Arthropoda, Decapoda, Carididae

Total length (TL): from the tip of the rostrum to the tip of the telson. Cephalothorax length (CL): from the level of the posterior
margin of the orbit to the midpoint of the posterodorsal margin of the carapace.

538
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Ostracoda

Arthropoda, Copepoda

539
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Arthropoda, Decapoda, Brachyura

540
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Vertebrata, Osteichthyes

541
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Genus index
Abyssocladia, 35, 37f., 519 Boloceroides, 65, 72, 519 Dysommina, 493, 531
Abyssorchomene, 396f. Bonnierella, 390f., 529 Ecbathyrion, 343, 528
Abyssotherma, 31, 33, 519 Bouillonia, 48 Echinopelta, 113, 521
Acharax, 149, 523 Bouvierella, 382, 388, 529 Echinus, 479, 531
Achelia, 305 Brachylepas, 366 Ectopleura, 48, 52, 519
Actinostola, 66, 71, 519 Branchinotogluma, 183, 219-229, 301, 524 Eochionelasmus, 356-359, 362f., 366, 480,
Adula, 207 Branchiplicatus, 230, 524 528
Agathotanais, 372 Branchipolynoe, 14, 160, 162, 231-234, 524 Eosipho, 124f., 459, 522
Alaysia, 255, 525 Brisinga, 477, 531 Erebonaster, 344
Allograea, 455, 459, 470, 530 Bruceiella, 94, 520 Euchelipluma, 46, 519
Alvania, 133, 522 Bythograea, 411, 455, 463, 465-472, 530f. Eudendrium, 51, 519
Alvinella, 14, 27, 104, 106, 108f., 116, 118, Calispadella, 486, 531 Eulalia, 215, 524
182, 200, 282ff., 341, 399, 525 Callinectes, 297 Eulepedopsis, 83, 520
Alvinocaris, 360, 410, 413-421, 529 Calocarcinus, 455 Eunice, 193, 524
Alvinoconcha, 15, 82, 128, 130, 422, 425, Calyptogena, 13, 77, 80f., 101, 106, 141-148, Euonyx, 394, 529
480, 522 209f., 215, 219, 230, 297, 334, 337, Euphilomedes, 310, 526
Amathys, 295, 526 360, 480, 484, 523 Exallopus, 187, 524
Ambilimbus, 344, 528 Candelabrum, 46, 49f., 140, 424, 519 Exrima, 334, 527
Ammothea, 301, 306, 526 Caprella, 387, 529 Fissuricola, 335, 527
Ampelisca, 383, 529 Carcinonemertes, 29 Freyella, 478, 531
Amphicrossus, 344 Careproctus, 509, 531 Fucaria, 95, 520
Amphiduropsis, 198, 524 Cataetyx, 499, 531 Gaidropsarus, 497, 531
Amphisamytha, 130, 183, 190, 296, 526 Caulophacus, 47, 519 Galapagomystides, 18, 216, 524
Andromastax, 318, 526 Cephalochaetosoma, 178, 523 Genesis, 29, 170, 355, 528
Anisopes, 304 Cerianthus, 72 Gigantidas, 162f., 419, 523
Aphotopontius, 316, 328f., 338, 526f. Ceuthoecetes, 29, 331, 527 Gitanopsis, 384, 529
Apotectonia, 392, 529 Chaceon, 455, 458, 530 Glycera, 195ff., 524
Arborammina, 32 Chasmatopontius, 332, 527 Gordotanais, 372
Archiconchoecia, 307, 526 Cheramomyzon, 323, 526 Gorgoleptis, 87, 520
Archinome, 183, 185, 524 Chionelasmus, 358 Grammaria, 55, 519
Arcovestia, 18, 70, 256, 525 Chionoecetes, 297 Graneledone, 166, 172, 523
Argestoides, 320, 526 Chiridota, 480, 531 Grievella, 351, 528
Armaturatanais, 372f., 528 Cholidya, 29, 354, 528 Grimpoteuthis, 166, 169, 523
Asbestopluma, 39ff., 519 Chondrocladia, 36, 42f., 519 Halacarellus, 300, 526
Ashinkailepas, 356, 360, 362f., 528 Chondrophellia, 73, 164, 501, 519 Halecium, 48
Atlantocuma, 370, 528 Chorocaris, 15, 341, 404, 419-423, 466, 505, Halice, 170, 382, 401, 529
Austinograea, 35, 425, 455, 460-466, 530 530 Halisiphonia, 54, 519
Autonoe, 385, 529 Cirroteuthis, 166, 168, 523 Harmothoe, 183
Barathricola, 326, 526 Cirrothauma, 167, 523 Harpiniopsis, 529
Bathybdella, 29, 297, 526 Cladorhiza, 44f., 519 Helicoradomenia, 75-79, 520
Bathyconchoecia, 308f., 526 Clypeosectus, 86, 520 Helicrenion, 102, 521
Bathycuma, 371, 528 Collocherides, 324, 526 Heptnerina, 316, 327, 526
Bathykurila, 218, 524 Copidognathus, 299, 526 Hesiodeiria, 199, 524
Bathylaophonte, 345, 528 Coryphaenoides, 442, 511, 531 Hesiolyra, 183, 200f., 524
Bathymargarites, 101, 521 Ctenopelta, 111, 521 Hesiospina, 183, 202f., 524
Bathymedon, 529 Cyanagraea, 14, 297, 455, 472, 531 Heteromesus, 379ff., 529
Bathymodiolus, 13, 15, 29, 68ff., 80f., 83, 85, Cyananthea, 65, 67, 519 Hirondellea, 395, 529
91, 97, 115, 125, 137, 141, 150-161, Cyathermia, 104, 521 Hirtopelta, 114, 521
194, 206, 208, 231, 243f., 297, 327, Cyclocaris, 393, 529 Humesipontius, 336, 527
353, 359, 379, 385, 411, 421, 424f., Cyclopina, 316 Hyalogyrina, 135f., 522
429, 430, 450, 458, 461, 469, 473, Dahlella, 369, 528 Hyalopontius, 347, 528
483ff., 491, 496, 501, 508, 523 Dendronotus, 140, 523 Hydractinia, 48
Bathynectes, 457, 530 Depressigyra, 82, 112, 521 Hydrallmania, 59, 519
Bathypecten, 141, 164, 523 Desbruyeresia, 129, 522 Hydrolagus, 489, 491f., 531
Beggiatoa, 188, 205 Desmodora, 174f., 523 Hyocrinus, 475, 531
Benthoctopus, 166, 171, 523 Desmodorella, 176f., 523 Hyphalion, 325, 526
Benthomisophria, 349 Dinetia, 179, 523 Hypoechinorhynchus, 29, 181, 523
Benthoxynus, 330, 340, 527 Dirivultus, 316, 333, 527 Hysterothylacium, 29

542
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Ifremeria, 15, 82, 110, 121, 130f., 227, 249, Nodopelta, 116f., 521 Sericosura, 302-306, 526
436, 460, 480, 522 Nuculana, 344 Sertularella, 60, 519
Iheyomytilidicola, 206, 524 Oasisia, 260, 288, 525 Shinkai, 209ff., 524
Ilyophis, 494f., 531 Obesutanais, 376, 528 Shinkaia, 434, 449, 530
Imbricaverruca, 357, 362, 367, 528 Olgaconcha, 15, 130 Shinkaicaris, 413f., 419, 430, 530
Iphionella, 235, 524 Olgasolaris, 121, 131, 522 Shinkailepas, 122f., 522
Isaacsicalanus, 352, 528 Oncaea, 316, 350, 528 Sinepecten, 141, 165, 523
Jamstecia, 321, 526 Opaepele, 419, 427, 530 Siphonalia, 435
Johanssonia, 297 Ophiactis, 482, 531 Siphonobrachia, 264, 525
Kiwa, 450f., 530 Ophioctenella, 483, 531 Sirsoe, 205, 524
Lacunoides, 105, 521 Ophiolamina, 481, 484f., 531 Smacigastes, 353, 528
Laeviphitus, 127, 522 Ophryotrocha, 188-191, 524 Speculator, 126, 522
Lafoea, 56, 519 Opisthotrochopodus, 219, 221, 228 Speleophriopsis, 349
Laitmatobius, 346, 528 Oradarea, 386, 529 Sphaerosyllis, 252, 525
Lamellibrachia, 257ff., 333, 410, 525 Orbiniella, 269f., 525 Spinophiura, 484f., 531
Laminatubus, 125, 164, 183, 253, 509, 525 Orchomene, 396f., 529 Spiochaetopterus, 272, 525
Laonice, 273, 525 Pachycara, 68, 158, 180, 502-505, 531 Stegolaria, 62, 519
Laubiericrinus, 476, 531 Pachydermia, 108, 521 Steleuthera, 404, 529
Laubieriellus, 274, 525 Pacmanactis, 70, 519 Stenhelia, 348, 528
Lebbeus, 360, 431f., 461, 530 Paragiopagurus, 435, 530 Stenothoe, 406, 529
Leitoscoloplos, 268, 525 Paralepetopsis, 84f., 520 Stygiopontius, 341f., 527f.
Lepetodrilus, 15, 67, 82, 88ff., 152, 520 Paralomis, 360, 436f., 530 Sutilizona, 98, 521
Lepidion, 498, 531 Paralvinella, 15, 183, 285-294, 332, 525f. Symmetromphalus, 110, 521
Lepidonotopodium, 183, 236-242, 524f. Paranthosactis, 71, 519 Symplectoscyphus, 61, 519
Leptochiton, 80f., 520 Parasicyonis, 68 Tectovalopsis, 398f., 529
Leptognathia, 372 Pardalisca, 402, 529 Temnocinclis, 99, 521
Leptognathiella, 374, 528 Parougia, 183, 192, 524 Temnozaga, 100, 521
Leptogyra, 96, 520 Periclimenes, 410 Tevnia, 65, 88, 265, 288, 525
Leucolepas, 357, 361f., 365, 528 Peltospira, 118, 521 Thalassomonohystera, 173
Levensteiniella, 183, 243ff., 525 Phymorhynchus, 68, 138f., 156, 361, 522 Thermaloniscus, 379
Lindaspio, 275ff., 525 Planorbidella, 109, 521 Thermarces, 181, 500, 507ff., 531
Lirapex, 115, 521 Polycirrus, 183 Thermichthys, 500, 531
Lucernaria, 64, 519 Polycopetta, 311, 526 Thermiphione, 246ff., 525
Luckia, 382, 389, 529 Polyplumaria, 53, 519 Thermobiotes, 496, 531
Luffammina, 32, 519 Prionospio, 278ff., 525 Thermochiton, 80f., 520
Lurifax, 136, 522 Prionotoleberis, 312, 526 Thermonemertes, 183, 523
Macroregonia, 386f. 456, 530 Prometopa, 405 Thermopalia, 14, 63, 519
Maractis, 68, 462, 519 Protis, 254, 525 Thermopolynoe, 249f., 525
Marianactis, 69, 156, 158, 426, 429, 519 Protolira, 97, 152, 521 Thomontocypris, 313f., 526
Melanodrymia, 106, 521 Protomystides, 217, 524 Thraumastos, 194, 524
Mesotanais, 375, 528 Provanna, 93, 132, 522 Thymopides, 433, 442, 530
Methanotrix, 32 Pseudorimula, 91f., 520 Thysanoessa, 409, 529
Metopa, 405, 529 Pseudotanais, 372, 378, 529 Tisbe, 354
Miralvinella, 285ff. Pyrolycus, 506, 531 Torometopa, 407, 529
Mirocaris, 424f., 530 Pyropelta, 93, 520 Trachyscorpia, 510, 531
Misophriopsis, 349, 528 Rhogobius, 338, 527 Transtectonia, 400, 529
Moravecnema, 29, 180, 523 Rhynchopelta, 119, 521 Tubularia, 48, 53
Munida, 438, 530 Rickettsia, 29 Typhlotanais, 372, 377, 528
Munidopsis, 70, 411, 434, 439-448, 530 Ridgeia, 89, 106, 126, 172, 261, 293, 455, Uptionyx, 319, 526
Mytilidiphila, 207f., 524 525 Uroptychus, 452ff., 530
Myzobdella, 297 Riftia, 14, 29, 33, 67, 71, 88, 90, 101, 104, Vampiropolynoe, 251, 525
Nautalvinella, 288ff. 106, 111, 119, 170, 182f., 202, 215, Ventichthys, 501, 531
Nautilocaris, 426, 460, 530 219, 230, 262f., 278, 297, 307, 311, Ventiella, 382, 408, 529
Nautilus, 166 313ff., 403, 411, 416, 424, 431f., 464, Vesicomya, 141
Nematocarcinus, 411f., 466, 529 468f., 484f., 508f., 525 Vetulonia, 103, 521
Neobrachylepas, 356f., 362, 366, 528 Rimicaris, 17, 68, 156, 339, 341, 404, 417, Vulcanoctopus, 166, 170, 355, 523
Neolepas, 73, 357, 361ff., 528 419, 428f., 462, 505, 511, 530 Vulcanolepas, 145, 355f., 361, 364f., 528
Neolepetopsis, 84f., 520 Rimipontius, 339, 527 Xandaros, 281, 525
Neomphalus, 107, 521 Rimitantalus, 29 Xenograpsus, 455, 474, 531
Neoverruca, 356ff., 362, 366, 368, 528 Saxipendium, 14, 487, 531 Xylocythere, 315, 526
Nereimyra, 204, 524 Scabrantenna, 322, 526 Xylodiscula, 137, 522
Nereis, 212ff., 524 Scoloplos, 271, 525 Zooarium, 458
Neusas, 134, 522 Scotoecetes, 340, 527 Zygophylax, 48, 57f., 519
Nicomache, 266f., 525 Seba, 403, 529
Nilva, 337, 527 Segonzacia, 455, 463, 473, 499, 531

543
 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Author index
Almeida, 182, 491f., 497ff., 502ff., 510f. Larsen, 373-377
Baba, 434f., 438, 440, 444ff., 448f., 452ff. Lemaitre, 435
Bamber, 301-306 Lopez-Gonzalez, 65-73
Barthélémy, 409 Lutz, 145, 160f.
Bartsch, 299f. Macpherson, 433-449, 452ff., 456
Bellan-Santini, 382-408 Maddocks, 313ff.
Biscoito, 13-25, 457f., 489-495, 497ff., 502ff., 507f., 510f. Mah, 477f.
Blake, 187-193, 212f., 215ff., 252, 266-271, 274-279, 281 Miura, 206-211, 225f., 231, 246, 249f., 259
Böggemann, 195 Möller, 501
Bouchet, 83f., 94f., 99-104, 107, 111ff., 119f., 124f., 130, 133ff., Moreau, 486
137, 140 Myers, 385, 390
Boury-Esnault, 47 Newman, 356-368
Briand, 457, 494, 496, 509 Ng, 474
Bright, 6-9, 27f., 174-179, 518 Nielsen, 500f.
Buron de, 29f., 181 Okutani, 138f.
Burreson, 297 Pleijel, 198-205
Casanova B., 409 Pugh, 63
Casanova J.-P., 486 Ramirez-Llodra, 479
Causse, 494f. Rodríguez, 70f.
Cohen, 500 Roux, 475f.
Corbera, 370f. Salvini-Plawen, 75-79
Cosel, 80f., 83, 85, 94, 96, 99-104, 107, 111ff., 119f., 126f., 130, Schein, 164f.
134f., 137, 141-148, 151-154, 156, 158, 162
Segonzac, 6-9, 29f., 37-46, 49-69, 73, 140, 167-170, 253f., 297,
Cunha, 372-381, 385, 390 356ff., 378, 411-414, 416-424, 426, 428, 430, 433, 436f., 439,
Daly, 64, 72 441ff., 447, 450f., 455, 460f., 463, 466-472, 474, 477, 478, 482,
Defaye, 319-327, 332, 335ff., 339f., 343, 346-355 485, 501, 507
Desbruyeres, 6-9, 13-25, 149, 184f., 190-194, 196f., 212ff., 218- Senz, 183
224, 227-230, 232-245, 247-250, 252f., 256, 260, 262f., 266, Southward A.J., 356-368
268, 272f., 280, 282-296, 487, 513-517 Southward E.C., 255, 257f., 260f., 264f.
Gebruck, 13-25, 480 Stöhr, 481ff., 485
Geistdoerfer, 484, 505 Tsuchida, 462
Gonzalez, 170 Tunnicliffe, 13-25
Guerra, 166, 168, 170 Tyler, 13-25, 479, 483
Guinot, 455, 459ff., 465-473 Vacelet, 35-46
Haney, 369 Van Dover, 13-25,
Harrison-Nelson, 307-312 Verschelde, 174-177
Hashimoto, 13-25, 148, 150, 155, 157, 159, 163, 429, 464, 506 Vervoort, 48-62
Holzmann, 31ff. Villanueva, 166f., 169
Hourdez, 218f., 223, 227, 235ff., 242ff., 247, 251 Voight, 166, 171f., 339
Hurtado, 459, 465 Vrijenhoek, 145, 160, 459
Ivanenko, 316-355 Warén, 82-123, 126-137
Jaume, 386, 391-395, 398ff., 402-405 Wilson, 380f.
Jones, 362 Yamaguchi, 356-368
Justine, 180 Young, 493
Komai, 410-428, 430ff. Zekely, 174-179
Kornicker, 307-312 Zibrovius, 254
Kudenov, 186

544