Axioidea of the World and a Reconsideration of the
Callianassoidea (Decapoda, Thalassinidea,
Callianassida)
CRUSTACEANA MONOGRAPHS constitutes a series of books on carcinology in its widest sense. Contributions are handled by the Series Editor(s) and may be submitted through the office of KONINKLIJKE
BRILL Academic Publishers N.V., P.O. Box 9000, NL-2300 PA Leiden, The Netherlands.
Series Editors for the current volume: J.C. VON VAUPEL K LEIN, P.O. Box 30, NL-3720 AA Bilthoven, The
Netherlands; e-mail: jcvvk@xs4all.nl and C.H.J.M. F RANSEN, c/o National Museum of Natural History,
P.O. Box 9517, NL-2300 RA Leiden, The Netherlands; e-mail: Charles.Fransen@ncbnaturalis.nl
Editorial Committee: N.L. B RUCE, Wellington, New Zealand; Mrs. M. C HARMANTIER -DAURES, Montpellier, France; Mrs. D. D EFAYE, Paris, France; H. D IRCKSEN, Stockholm, Sweden; J. F OREST, Paris,
France; R.C. G UIA ŞU, Toronto, Ontario, Canada; R.G. H ARTNOLL, Port St. Mary, Isle of Man;
E. M ACPHERSON, Blanes, Spain; P.K.L. N G, Singapore, Rep. of Singapore; H.-K. S CHMINKE, Oldenburg, Germany; F.R. S CHRAM, Langley, WA, U.S.A.; G. VAN DER V ELDE, Nijmegen, Netherlands;
H.P. WAGNER, Leiden, Netherlands; D.I. W ILLIAMSON, Port Erin, Isle of Man.
Published in this series:
CRM 001 - Stephan G. Bullard
CRM 002 - Spyros Sfenthourakis et al.
CRM 003 - Tomislav Karanovic
CRM 004 - Katsushi Sakai
CRM 005 - Kim Larsen
CRM 006 - Katsushi Sakai
CRM 007 - Ivana Karanovic
CRM 008 - Frank D. Ferrari & Hans-Uwe Dahms
CRM 009 - Tomislav Karanovic
CRM 010 - Carrie E. Schweitzer et al.
CRM 011 - Pedro Castro et al. (eds.)
CRM 012 - Patricio R. De los Ríos-Escalante
CRM 013 - Katsushi Sakai
In preparation (provisional titles):
CRM 014 - Charles H.J.M. Fransen et al. (eds.)
CRM 015 - Akira Asakura et al. (eds.)
CRM 016 - Danielle Defaye et al. (eds.)
CRM 00x - Chang-tai Shih
Larvae of anomuran and brachyuran crabs of
North Carolina
The biology of terrestrial isopods, V
Subterranean Copepoda from arid Western Australia
Callianassoidea of the world (Decapoda,
Thalassinidea)
Deep-sea Tanaidacea from the Gulf of Mexico
Upogebiidae of the world (Decapoda, Thalassinidea)
Candoninae (Ostracoda) from the Pilbara region in
Western Australia
Post-embryonic development of the Copepoda
Marine interstitial Poecilostomatoida and Cyclopoida
(Copepoda) of Australia
Systematic list of fossil decapod crustacean species
Studies on Brachyura: a homage to Danièle Guinot
Crustacean zooplankton communities in Chilean
inland waters
Axioidea of the world and a reconsideration of
the Callianassoidea (Decapoda, Thalassinidea,
Callianassida)
Studies on Malacostraca: Lipke Bijdeley Holthuis
Memorial Volume
New frontiers in crustacean biology – Proceedings
of the TCS Summer Meeting, Tokyo,
20-24 September 2009
Studies on freshwater Copepoda: a volume in honour
of Bernard Dussart
Marine Calanoida of the China seas
Author’s address:
Prof. Dr. K. S AKAI, 1570 Nishiyama, Kamihachiman-cho, Tokushima 770-8041, Japan;
e-mail: ksakai@mf.pikara.ne.jp
Manuscript first received 9 October 2008; final version accepted 5 April 2010.
Cover: Gyanacaris hirsutimana (Boesch & Smalley, 1972), male; see p. 122, fig. 22A.
Axioidea of the World and a
Reconsideration of the
Callianassoidea (Decapoda,
Thalassinidea, Callianassida)
By
Katsushi Sakai
C RUSTACEANA M ONOGRAPHS , 13
LEIDEN • BOSTON
This book is printed on acid-free paper.
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The Library of Congress Cataloging-in-Publication Data is available from the Publisher.
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CONTENTS
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Infraorder classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Superfamily Axioidea Huxley, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Axiidae Huxley, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Acanthaxius Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . .
Genus Albatrossaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Alienaxiopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Allaxiopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Allaxius Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . . . . . .
Genus Amakusaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Anophthalmaxius De Man, 1905 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Australocaris Poore & Collins, 2009 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Axiopsis Borradaile, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Axiorygma Kensley & Simmons, 1988 . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Axius Leach, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Balssaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Boasaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Bouvieraxius Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . .
Genus Bruceaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Calaxius Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . . . . .
Genus Calocarides Wollebæk, 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Coelhocalaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Colemanaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Dorphinaxius Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . .
Genus Eutrichocheles Wood-Mason, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Guyanacaris gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Heterocalaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Leonardsaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Levantocaris Galil & Clark, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Litoraxius Komai & Tachikawa, 2007 . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Manaxius Kensley, 2003 (sensu nov.) . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Marianaxius Kensley, 1996 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Michelaxiopsis Poore & Collins, 2009 . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Newzealandaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Oxyrhynchaxius Parisi, 1917 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Parascytoleptus Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . .
Genus Paratrichocheles gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Paraxiopsis De Man, 1905 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Paraxius Bate, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
xi
1
9
13
15
19
29
32
36
40
44
45
48
50
63
64
73
76
79
82
84
86
98
103
106
109
119
123
124
129
130
131
142
143
148
150
152
155
156
164
vi
CRM 013 – Katsushi Sakai
Genus Pilbaraxius Poore & Collins, 2009 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pillsburyaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Planaxius Komai & Tachikawa, 2008 . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Platyaxiopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Platyaxius Sakai, 1994 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pseudoaxiopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Ralumcaris gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Scytoleptus Gerstaecker, 1856 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Spongiaxius Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . .
Family Callianideidae Kossmann, 1880 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Callianidea H. Milne Edwards, 1837 . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Paracallianidea Sakai, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Calocarididae Ortmann, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Ambiaxius Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . . . .
Genus Briancaris Sakai & Ohta, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Calastacus Faxon, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Calaxiopsis Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . . .
Genus Calocaris Bell, 1846 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Calocarisopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Eucalastacus Sakai, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Lophaxiopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Lophaxius Kensley, 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Nipponcalaxiopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Paracalocaris Sakai, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Coralaxiidae Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Coralaxius Kensley & Gore, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Meinertaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Eiconaxiidae Sakai & Ohta, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Eiconaxius Bate, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Eiconaxiopsididae fam. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Eiconaxiopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Meticonaxiidae Sakai, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Marcusiaxius De Carvalho & Rodrigues, 1973 . . . . . . . . . . . . . . . . . . .
Genus Meticonaxius De Man, 1905 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Tethisea Poore, 1994 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Micheleidae Sakai, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Michelea Kensley & Heard, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Strahlaxiidae Poore, 1994 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Neaxiopsis Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . . .
Genus Neaxius Borradaile, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Strahlaxius Sakai & De Saint Laurent, 1989 . . . . . . . . . . . . . . . . . . . . .
Superfamily Callianassoidea Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Anacalliacidae Manning & Felder, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Anacalliaopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Anacalliax De Saint Laurent, 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Capecalliax gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Bathycalliacidae Sakai & Türkay, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Subfamily Bathycalliacinae Sakai & Türkay, 1999 . . . . . . . . . . . . . . . . . . . . . . . .
167
169
171
172
174
177
182
185
193
197
198
205
209
211
213
214
223
228
242
244
245
246
248
250
253
254
260
267
268
289
290
297
301
305
309
311
311
319
320
324
332
335
341
342
343
345
347
347
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Genus Bathycalliax Sakai & Türkay, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Subfamily Vulcanocalliacinae Dworschak & Cunha, 2007 . . . . . . . . . . . . . . . . .
Genus Vulcanocalliax Dworschak & Cunha, 2007 . . . . . . . . . . . . . . . . . . . . . .
Family Callianassidae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Subfamily Callianassinae Dana, 1852 (sensu nov.) . . . . . . . . . . . . . . . . . . . . . . . .
Genus Callianassa Leach, 1814 (sensu nov.) . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Cheramoides gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Cheramus Bate, 1888 (sensu nov.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Gilvossius Manning & Felder, 1992 (sensu nov.) . . . . . . . . . . . . . . . . .
Genus Notiax Manning & Felder, 1991 (sensu nov.) . . . . . . . . . . . . . . . . . . . .
Genus Trypaea Dana, 1852 (sensu nov.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Subfamily Callichirinae Manning & Felder, 1991 (sensu nov.) . . . . . . . . . . . . . .
Genus Balsscallichirus gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Barnardcallichirus gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Callichirus Stimpson, 1866 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Corallichirus Manning, 1992 (sensu nov.) . . . . . . . . . . . . . . . . . . . . . . .
Genus Forestcallichirus gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Glypturoides gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Glypturus Stimpson, 1866 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Grynaminna Poore, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Lepidophthalmoides gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Lepidophthalmus Holmes, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Michaelcallianassa Sakai, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Neocallichirus Sakai, 1988 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Podocallichirus Sakai, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Sergio Manning & Lemaitre, 1994 (sensu nov.) . . . . . . . . . . . . . . . . . .
Genus Thailandcallichirus gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Tirmizicallichirus gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Callianopsidae Manning & Felder, 1991 (sensu nov.) . . . . . . . . . . . . . . . . . .
Subfamily Callianopsinae Manning & Felder, 1991 . . . . . . . . . . . . . . . . . . . . . . . .
Genus Callianopsis De Saint Laurent, 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pleurocalliax gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Subfamily Neocallianopsinae subfam. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Neocallianopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Ctenochelidae Manning & Felder, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Ctenocheles Kishinouye, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Eucalliacidae Manning & Felder, 1991 (sensu nov.) . . . . . . . . . . . . . . . . . . .
Subfamily Calliapaguropinae Sakai, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Calliapagurops De Saint Laurent, 1973 . . . . . . . . . . . . . . . . . . . . . . . . .
Subfamily Eucalliacinae Manning & Felder, 1991 (sensu nov.) . . . . . . . . . . . . .
Genus Andamancalliax gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Calliax De Saint Laurent, 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Calliaxina Ngoc-Ho, 2003 (sensu nov.) . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Eucalliax Manning & Felder, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Eucalliaxiopsis gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Paraglypturus Türkay & Sakai, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pseudocalliax gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Gourretiidae Sakai, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
vii
349
350
350
353
357
359
362
363
372
381
385
412
414
416
418
424
426
428
429
438
440
446
450
451
466
467
473
474
477
477
478
480
482
482
485
486
491
491
492
493
494
495
497
502
503
504
505
507
viii
CRM 013 – Katsushi Sakai
Genus Dawsonius Manning & Felder, 1991 sensu Sakai, 2005 . . . . . . . . . . .
Genus Gourretia De Saint Laurent, 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Laurentgourretia Sakai, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Paracalliax De Saint Laurent, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Paragourretia Sakai, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Lipkecallianassidae Sakai, 2005 (new status) . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Lipkecallianassa Sakai, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Pseudogourretiidae Sakai, 2005 (new status) . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pseudogourretia Sakai, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Thomassiniidae De Saint Laurent, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Crosniera Kensley & Heard, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Garyia gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Heardaxius gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Mictaxius Kensley & Heard, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Thomassinia De Saint Laurent, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Data on biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Addendum I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Addendum II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Errata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Taxonomic index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
508
510
514
515
516
521
522
523
523
525
527
530
533
534
535
539
549
551
593
595
597
599
PREFACE
With this book on the superfamily Axioidea I conclude my worldwide
overview of the second major clade of the infraorder Thalassinidea, started
in 2005 with a first treatise on the Callianassoidea (cf. Sakai, 2005b). The
present text treats the Axioidea and reconsiders the classification of the Callianassoidea, thus resulting in a complete review of the Callianassida sectio
nov., now conceived as a distinct section within the infraorder Thalassinidea
(obviously in addition to the nominate section Thalassinida sectio nov.). In
addition, the Upogebiidae have also been reviewed (Sakai, 2006), a family
contained in the superfamily Thalassinoidea. Together these volumes are intended to present both an inventory of the currently known species of those
taxa of the infraorder Thalassinidea, as well as my views on their classification.
Descriptions of the taxa included have been made as accurate as required,
both in text and in figures. Nomenclature and literature have been screened
to comply with international standards and to be as exhaustive as possible.
The principles of classification I have used primarily refer to the classical
school in taxonomic practice, also known as “Evolutionary Systematics”,
that was established by J. Huxley, G.G. Simpson, and E. Mayr, largely
during the years 1940-1960. I fully realize that in the future other criteria of
classification may increasingly be employed, mainly those of “Phylogenetic
Systematics” based on the principles explained by W. Hennig, most effectively
in 1966. Also, classifications will progressively be founded on data other than
those derived from morphology and anatomy, as, e.g., molecular and genetic
evidence. Nevertheless, I trust that my present compilation may serve as a basis
from which to work on a more and more fine-tuned and well-corroborated
classification of this interesting and important group of animals. I also hope
that the distributional data presented may likewise be of value in detailing the
biogeography of the Callianassida (as well as, of course, of the Upogebiidae
in the Thalassinida).
Having remarked this, I all the same point to the fact that, where during
recent years cladistic analyses have become available, I have carefully compared those with my own data and, where I deemed such appropriate, have
x
CRM 013 – Katsushi Sakai
implemented some of those results into my own classification, i.e., as far as
I judged relevant.
While acknowledging the above, I am fully aware of the fact that new
thalassinidean taxa are continuously being described as more material and
more sophisticated techniques become available. Yet, as with every review
work, the present text can only give a momentaneous picture of what is known
to date — though I have made every attempt to ensure that this report on the
current status quo will be as complete as humanly possible∗ ).
Tokushima, April 2010
K. S AKAI
∗ ) Meanwhile, some new results have indeed been published: I refer to the Addenda following
the main text for recent additions to thalassinidean systematics.
Tokushima, July 2010
K. S AKAI
INTRODUCTION
The infraorder Thalassinidea Latreille, 1831 as a whole was reviewed by
Borradaile (1903), Poore (1994, 1997), and Tudge et al. (2000). The families
Axiidae and Callianassidae were reviewed by De Man (1925, 1928); the family
Axiidae was reviewed by Kensley (1981a, 1989, 1994, 1996a-e, 2003) and by
Sakai & De Saint Laurent (1989); the Callianassidae and Ctenochelidae were
reviewed by Manning & Felder (1991); the Callianassidae were reviewed by
Sakai (1999c); the families contained in the Callianassoidea were reviewed by
Sakai (2005b); and the Upogebiidae were reviewed by Sakai (2006).
The recent morphological phylogeny of the families Callianassidae and
Ctenochelidae was established by Manning & Felder (1991) on the basis
of features of the carapace, the third maxilliped, the larger cheliped, the
abdomen, the tail-fan, and the appendix interna. The families included in
the three superfamilies (Thalassinoidea Dana, 1852a in the Thalassinida, and
Callianassoidea Dana, 1852a and Axioidea Huxley, 1879 in the Callianassida)
were further reviewed by Poore (1994) using cladograms resulting from
cladistic analysis. Sakai (2005a: 1117) and Sakai & Sawada (2006: 1353)
proposed that the Thalassinida [as Thalassinidea] should be distinguished from
the Callianassida [as Callianassidea] by morphological features of the second
pereiopod, the gastric mill, and the uropodal exopod.
A dichotomous scheme for the relationships among 14 species of thalassinidean shrimp (Callianassidae, Laomediidae, Strahlaxiidae, Thalassinidae,
and Upogebiidae) has been drawn up by using partial sequences of the 18S
nuclear and 16S mitochondrial ribosomal genes (Tudge & Cunningham, 2002:
842). According to that analysis, there appear to be two strongly supported
major clades in the Thalassinidea: the first clade includes a monophyletic Callianassidae Dana, 1852 as the sister-group of a representative of the axioid
family, Strahlaxiidae Poore, 1994; the second clade includes three families,
Laomediidae Borradaile, 1903, Thalassinidae Latreille, 1831, and Upogebiidae Borradaile, 1903.
By employing partial segments of two nuclear protein-coding genes, phosphoenolpyruvate carboxykinase (PEPCK, 570 bp) and sodium-potassium
2
CRM 013 – Katsushi Sakai
ATPase a-subunit (NaK, 534 bp), Chu et al. (2008) reconstructed the phylogeny of six species of the superfamily Axioidea Huxley, 1879, i.e., Calastacus crosnieri Kensley & Chan, 1998, Calaxius acutirostris Sakai & De Saint
Laurent, 1989 (syn. Calaxius manningi Kensley et al., 2000), Paracalocaris
sagamiensis Sakai, 1991, Eiconaxius indicus (De Man, 1907), and Calocarides chani Kensley, Lin & Yu, 2000, which are separated from Upogebia
edulis Ngoc-Ho & Chan, 1992 (= Austinogebia edulis) and Thalassina anomala (Herbst, 1804).
Those results mentioned above are systematically in accordance with the results of the classification proposed on the basis of the morphology of the gastric mill (Sakai, 2005a; Sakai & Sawada, 2006): the first sectio Callianassida
Dana, 1852 (new status), was supposed to contain the superfamily Callianassoidea Dana, 1852 with five families [as then conceived, see Sakai 2005a]:
Axiidae Huxley, 1879, Callianassidae Dana, 1852, Callianideidae Kossmann,
1880, Ctenochelidae Manning & Felder, 1991, and Gourretiidae Sakai, 1999;
and the second sectio Thalassinida Latreille, 1831 (new status) with a single
superfamily, Thalassinoidea Latreille, 1831, was presumed to include three
families: Laomediidae Borradaile, 1903, Thalassinidae Latreille, 1831, and
Upogebiidae Borradaile, 1903.
The present taxa, then, comprise two sections, Thalassinida including one
superfamily, Thalassinoidea Dana, 1852a, and Callianassida including two
superfamilies, Axioidea Huxley, 1879 and Callianassoidea Dana, 1852a.
Those two superfamilies are characterized by the presence or absence of
a rostral carina, (a) cardiac sulcus(i), cardiac prominence, dorsal oval of the
carapace, linea thalassinica, setal rows on carapace, abdomen, tail-fan, and
pereiopods, a posterior whip on the maxilla 2 scaphognathite, a dorsal plate or
lateral notch on the uropodal exopod, male pleopods 1-2, and the median tooth
of the urocardiac ossicle in the gastric mill.
In the present, new classification the section Callianassida consists of two
superfamilies (Axioidea and Callianassoidea), 19 families including 3 new
(or newly ranked) families, 119 genera including 38 new genera and 9 sensu
novo, and 432 species including 22 uncertain species [= species inquirendae],
as outlined below; the classification at family level of the whole infraorder
Thalassinidea is here given for convenience:
Infraorder Thalassinidea Latreille, 1831
Section Thalassinida Latreille, 1831 [sectio nov.]
Superfamily Thalassinoidea Latreille, 1831
Family Laomediidae Borradaile, 1903
Family Thalassinidae Latreille, 1831
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
3
Family Upogebiidae Borradaile, 1903
Section Callianassida Dana, 1852 [sectio nov.]
Superfamily Axioidea Huxley, 1879
Family Axiidae Huxley, 1879
Family Callianideidae Kossmann, 1880
Family Calocarididae Ortmann, 1891
Family Coralaxiidae Sakai & De Saint Laurent, 1989
Family Eiconaxiidae Sakai & Ohta, 2005
Family Eiconaxiopsididae fam. nov.
Family Meticonaxiidae Sakai, 1992
Family Micheleidae Sakai, 1992c
Family Strahlaxiidae Poore, 1994
Superfamily Callianassoidea Dana, 1852
Family Anacalliacidae Manning & Felder, 1991
Family Bathycalliacidae Sakai & Türkay, 1999
Family Callianassidae Dana, 1852
Family Callianopsidae Manning & Felder, 1991
Family Ctenochelidae Manning & Felder, 1991
Family Eucalliacidae Manning & Felder, 1991
Family Gourretiidae Sakai, 1999
Family Lipkecallianassidae Sakai, 2005 (new status)
Family Pseudogourretiidae Sakai, 2005 (new status)
Family Thomassiniidae De Saint Laurent, 1979
TABLE I
List of families, subfamilies, and genera in the Axioidea and Callianassoidea as conceived herein
Families
Subfamilies
Genera
Species
Acanthaxius
Albatrossaxius gen. nov.
7
1+1
sp. inq.
2
3
1
1
1
1
3+1
sp. inq.
1
4
1
1
5
1
1+1
sp. inq.
Superfamily A XIOIDEA Huxley, 1879
A XIIDAE
Alienaxiopsis gen. nov.
Allaxiopsis gen. nov.
Allaxius
Amakusaxius gen. nov.
Anophthalmaxius
Australocaris
Axiopsis
Axiorygma
Axius
Balssaxius gen. nov.
Boasaxius gen. nov.
Bouvieraxius
Bruceaxius gen. nov.
Calaxius
4
CRM 013 – Katsushi Sakai
TABLE I
(Continued)
Families
Subfamilies
Genera
Species
Calocarides
Coelhocalaxius gen. nov.
Colemanaxius gen. nov.
Dorphinaxius
Eutrichocheles
Guyanacaris gen. nov.
Heterocalaxius gen. nov.
Leonardsaxius gen. nov.
14
1
1
1
7
2
1
3+1
sp. inq.
1
1
12
1
1
1
3
1
1
9+1
sp. inq.
1
1
1
1
1
2
1
1
2
1
3
1
4+1
sp. inq.
3
8
1+1
sp. inq.
4+1
sp. inq.
1
1
1
2
1
1
3
1
29
Levantocaris
Litoraxius
Manaxius (sensu nov.)
Marianaxius
Michelaxiopsis
Newzealandaxius gen. nov.
Oxyrhynchaxius
Parascytoleptus
Paratrichocheles gen. nov.
Paraxiopsis
C ALLIANIDEIDAE
C ALOCARIDIDAE
Paraxius
Pilbaraxius
Pillsburyaxius gen. nov.
Planaxius
Platyaxiopsis gen. nov.
Platyaxius
Pseudoaxiopsis gen. nov.
Ralumcaris gen. nov.
Scytoleptus
Spongiaxius
Callianidea
Paracallianidea
Ambiaxius
Briancaris
Calastacus
Calaxiopsis
Calocaris
C ORALAXIIDAE
E ICONAXIIDAE
Calocarisopsis gen. nov.
Eucalastacus
Lophaxiopsis gen. nov.
Lophaxius
Nipponcalaxiopsis gen. nov.
Paracalocaris
Coralaxius
Meinertaxius gen. nov.
Eiconaxius
5
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
TABLE I
(Continued)
Families
Subfamilies
E ICONAXIOPSIDIDAE
fam. nov.
M ETICONAXIIDAE
Genera
Eiconaxiopsis gen. nov.
Marcusiaxius
Meticonaxius
Tethisea
Michelea
M ICHELEIDAE
S TRAHLAXIIDAE
Neaxiopsis
Neaxius
Strahlaxius
Species
2
7+1
sp. inq.
7+1
sp. inq.
2
12 + 1
sp. inq.
2
4
2
Superfamily C ALLIANASSOIDEA Dana, 1852
A NACALLIACIDAE
BATHYCALLIACIDAE
C ALLIANASSIDAE
Bathycalliacinae
Vulcanocalliacinae
Callianassinae
(sensu nov.)
Anacalliaopsis gen. nov.
Anacalliax
Capecalliax gen. nov.
Bathycalliax
Vulcanocalliax
Callianassa (sensu nov.)
Cheramoides gen. nov.
Cheramus (sensu nov.)
Gilvossius (sensu nov.)
Notiax (sensu nov.)
Trypaea (sensu nov.)
Callichirinae
(sensu nov.)
C ALLIANOPSIDAE
Callianopsinae
Neocallianopsinae
subfam. nov.
Balsscallichirus gen. nov.
Barnardcallichirus gen. nov.
Callichirus
Corallichirus (sensu nov.)
Forestcallichirus gen. nov.
Glypturoides gen. nov.
Glypturus
Grynaminna
Lepidophthalmoides gen. nov.
Lepidophthalmus
Michaelcallianassa
Neocallichirus
Podocallichirus
Sergio (sensu nov.)
Thailandcallichirus gen. nov.
Tirmizicallichirus gen. nov.
Callianopsis
Pleurocalliax gen. nov.
Neocallianopsis gen. nov.
1
1
1
1
1
1
2
15 + 1
sp. inq.
12 + 1
sp. inq.
6
52 + 1
sp. inq.
2
2
5
3
1
1
11
1
8
5
1
28 + 1
sp. inq.
1
7
1
1
1
1
1
6
CRM 013 – Katsushi Sakai
TABLE I
(Continued)
Families
Subfamilies
C TENOCHELIDAE
E UCALLIACIDAE
Calliapaguropinae
Eucalliacinae
G OURRETIIDAE
L IPKECALLIANASSIDAE
(new status)
P SEUDOGOURRETIIDAE
(new status)
T HOMASSINIIDAE
Genera
Species
Ctenocheles
Laurentgourretia
Paracalliax
Paragourretia
Lipkecallianassa
6+4
spp. inq.
2
1
2
8
1
1
1
1
1
6+1
sp. inq.
1
1
6
1
Pseudogourretia
1
Calliapagurops
Andamancalliax gen. nov.
Calliax
Calliaxina (sensu nov.)
Eucalliax
Eucalliaxiopsis gen. nov.
Paraglypturus
Pseudocalliax gen. nov.
Dawsonius
Gourretia
Crosniera
Garyia gen. nov.
Heardaxius gen. nov.
Mictaxius
Thomassinia
19 families (incl. 3 fam.
nov. or new status)
8 subfamilies (incl.
1 subfam. nov.
and 2 sensu nov.)
119 genera (incl. 38 gen.
nov. and 9 sensu nov.)
4+2
spp. inq.
1
1
1
2+1
sp. inq.
432 + 22
spp. inq.
Legend. — sp(p). inq. = species inquirenda(e).
Abbreviations
Museums and institutes referred to for scientific material examined. —
AM, Australian Museum, Sydney; CBM, Natural History Museum and Institute, Chiba; DOUFPe, Departamento de Oceanografia, Universidade Federal de Pernambuco, Pernambuco; MCZ, Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts; MNHN, Muséum national
d’Histoire naturelle, Paris; NHM, Natural History Museum, London; NSMT,
National Science Museum, Tokyo; NTOU, National Taiwan Ocean University, Keelung; QM, Queensland Museum, Brisbane; RMNH, Rijksmuseum
van Natuurlijke Historie [now National Museum of Natural History], Leiden;
SMF, Forschungsinstitut Senckenberg, Frankfurt am Main; SMNH, Swedish
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
7
Museum of Natural History (Natur Historiske Riksmuseet), Stockholm; UF,
Florida Museum of Natural History, University of Florida, Gainesville, Florida;
USNM, United States National Museum, Smithsonian Institution, Washington, D.C.; ZIASP, Zoological Institute of the Academy of Science in St. Petersburg, St. Petersburg; ZMA, Zoölogisch Museum, Universiteit van Amsterdam, Amsterdam; ZMB, Zoologisches Museum, Humboldt-Universität zu
Berlin, Berlin; ZMM, Zoological Museum of Moscow University, Moscow;
ZMUC, Zoological Museum, Copenhagen; ZRC, Zoological Reference Collection, University of Singapore, Singapore.
Abbreviations in morphological terminology. — A1, antennule or antenna
1; A2, antenna or antenna 2; CL, carapace length; Mxp, maxilliped; P,
pereiopod; Plp, pleopod.
INFRAORDER CLASSIFICATION
Infraorder T HALASSINIDEA Latreille, 1831
Section T HALASSINIDA Latreille, 1831
Superfamily T HALASSINOIDEA Latreille, 1831
Diagnosis. — Urocardiac∗ ) ossicle triangular, anteriorly protruded downward, and posteriorly concave. Its surface shows a longitudinal median carina
with a row of fine transverse septa on each side. P2 simple (in the Laomediidae
and Upogebiidae) or subchelate (in the Thalassinidae). [After Sakai & Sawada,
2006: 1355.]
Families included. — Laomediidae Borradaile, 1903; Thalassinidae Latreille, 1831; Upogebiidae Borradaile, 1903.
Section C ALLIANASSIDA Dana, 1852
Diagnosis. — Median tooth of urocardiac ossicle simple, with a smooth
posterior surface. P2 chelate. [After Sakai & Sawada, 2006: 1357.]
Superfamilies included. — Axioidea Huxley, 1879; Callianassoidea, Dana,
1852a.
Remarks. — In this revision, the section Callianassida is divided into two
superfamilies, Callianassoidea and Axioidea, by the presence or absence of
the linea thalassinica. In Callianassoidea the linea thalassinica is present,
but in Axioidea the linea thalassinica is absent, although in each of those
two superfamilies there is a family that includes genera in which the linea
thalassinica is morphologically intermediate. In Crosniera and Thomassinia
included in the Thomassiniidae under Callianassoidea, the linea thalassinica
is present, though it is incomplete, and in Paracallianidea included in the
Callianideidae under Axioidea, the linia thalassinica is present, though very
short.
∗ ) As pointed out by A. Brösing (in litt.), the ossicle we have previously referred to as
“prepyloric ossicle” in reality is the urocardiac ossicle.
10
CRM 013 – Katsushi Sakai
Much attention is paid also to the male Plps1-2, though it is difficult
to define their features, because they are sometimes either undeveloped, or
underdeveloped in juvenile individuals. Furthermore, only a few descriptions
of them have been made until recently, or if any, mostly of female Plps1-2,
because little attention was paid to male Plps1-2. All the genera included in the
families belonging to the Axioidea and Callianassoidea have been reviewed
in this respect. In consequence, almost all of the families and genera are
classified by the morphological features of the male Plps1-2 of the species
included, as shown in the following sections. The only exceptions involve
the genera Bouvieraxius and Eiconaxius in the Axioidea, as well as the
genera Callichirus, Lepidophthalmoides gen. nov. and Sergio sensu nov. in
the Callianassoidea, in which male Plp2 appendices interna and masculina are
not consistent in characteristics, since 2 or 3 different forms occur within one
genus.
1. Families and genera lacking male Plps1-2; male Plps3-5 endopods with appendix interna
Family Callianassidae Dana, 1852a: subfam. Callianassinae; Gilvossius (= Pestarella;
Paratrypaea).
2. Families and genera lacking male Plp1, but bearing Plp2; male Plps3-5 endopods with or
without appendix interna
a. Male Plp2 with appendices interna and masculina.
Family Axiidae: Acanthaxius; Albatrossaxius gen. nov.; Alienaxiopsis gen. nov.; Allaxiopsis gen. nov.; Allaxius; Anophthalmaxius; Axiopsis; Calaxius; Calocarides; Dorphinaxius; Manaxius (sensu nov.); Newzealandaxius gen. nov.; Platyaxiopsis gen. nov.;
Platyaxius; Pseudoaxiopsis gen. nov.
Family Eiconaxiidae: Eiconaxius∗ ).
Family Bathycalliacidae: subfam. Bathycalliacinae; Bathycalliax.
Family Thomassiniidae: Garyia gen. nov.; Mictaxius; Thomassinia.
b. Male Plp2 with appendix interna, but without appendix masculina.
Family Axiidae: Axiorygma; Heterocalaxius gen. nov.; Michelaxiopsis; Paratrichocheles
gen. nov. (Plps3-5 endopods lacking appendix interna); Pilbaraxius.
Family Eiconaxiidae: Eiconaxius∗ ); Pseudocalliax gen. nov.
Family Strahlaxiidae: Neaxiopsis; Neaxius.
c. Male Plp2 with appendix masculina, but without appendix interna.
Family Axiidae: Ralumcaris gen. nov. (Plps3-5 endopods without appendix interna).
3. Genus with male Plp1, but without male Plp2; Plps3-5 with appendix interna
Family Callianassidae: subfam. Callianassinae; Trypaea.
4. Families and genera with male Plps1-2; Plps3-5 with or without appendix interna
a. Male Plp1 without appendix interna; male Plp2 with appendices interna and masculina.
Family Axiidae: Axius; Boasaxius gen. nov.; Bouvieraxius∗ ); Bruceaxius gen. nov.;
Guyanacaris gen. nov.; Leonardsaxius gen. nov.; Levantocaris; Litoraxius; Oxyrhynchaxius; Spongiaxius.
Family Callianideidae: Callianidea; Paracallianidea.
Family Calocarididae: Ambiaxius; Briancaris; Calastacus; Calaxiopsis; Calocaris; Eucalastacus; Nipponcalaxiopsis gen. nov.; Paracalocaris.
Family Coralaxiidae: Coralaxius; Meinertaxius gen. nov.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
b.
c.
d.
e.
11
Family Eiconaxiopsididae fam. nov.: Eiconaxiopsis gen. nov.
Family Meticonaxiidae: Marcusiaxius; Meticonaxius; Tethisea.
Family Micheleidae: Michelea∗ ).
Family Callianassidae: subfam. Callichirinae sensu nov.; Glypturoides gen. nov.; Glypturus; Grynaminna; Lepidophthalmoides∗ ) gen. nov.; Lepidophthalmus; Neocallichirus;
Podocallichirus; Sergio∗ ) sensu nov.; Thailandcallichirus gen. nov.
Family Anacalliacidae: Anacalliaopsis gen. nov.; Anacalliax; Capecalliax gen. nov.
Family Callianopsidae: Callianopsis; Neocallianopsis gen. nov.
Family Eucalliacidae: Calliaxina∗ ) sensu nov.; Eucalliax; Eucalliaxiopsis gen. nov.;
Paraglypturus.
Family Ctenochelidae: Ctenocheles.
Family Gourretiidae: Dawsonius; Gourretia; Paragourretia.
Family Thomassiniidae: Crosniera; Heardaxius gen. nov.
Male Plp1 without appendix interna; male Plp2 with appendix interna, but without
appendix masculina.
Family Axiidae: Australocaris; Bouvieraxius∗ ); Parascytoleptus; Paraxius; Scytoleptus.
Family Calocarididae: Calocarisopsis gen. nov.
Family Micheleidae: Michelea∗ ).
Family Strahlaxiidae: Strahlaxius.
Family Bathycalliacidae: Vulcanocalliax.
Family Callianassidae: subfam. Callichirinae sensu nov.; Callichirus∗ ); Corallichirus
sensu nov.; Lepidophthalmoides∗ ) gen. nov.
Family Eucalliacidae: Calliapagurops.
Family Gourretiidae: Gourretia∗ ).
Male Plp1 without appendix interna; male Plp2 with appendix masculina, but without
appendix interna.
Family Axiidae: Eutrichocheles (Plps3-5 endopods without appendix interna); Marianaxius (male Plps3-5 endopods without appendix interna); Paraxiopsis (male Plps3-5 endopods without appendix interna).
Family Callianassidae: subfam. Callichirinae sensu nov.; Lepidophthalmoides gen. nov.;
Sergio∗ ) sensu nov.
Family Eucalliacidae: Andamancalliax gen. nov.; Calliax∗ ).
Male Plp1 without appendix interna; male Plp2 without appendices interna and masculina.
Family Axiidae: Bouvieraxius∗ ).
Family Callianassidae: subfam. Callianassinae; Callianassa; Cheramoides gen. nov.;
Cheramus; Notiax (= Rayllianassa).
Family Callianassidae: subfam. Callichirinae sensu nov.; Balsscallichirus gen. nov.;
Barnardcallichirus gen. nov.; Callichirus∗ ); Forestcallichirus gen. nov.; Michaelcallianassa; Tirmizicallichirus gen. nov.
Genera with male Plp1 with appendix interna; male Plp2 with appendices interna and
masculina; and Plps3-5 with appendix interna.
Family Axiidae: Planaxius.
Family Calocarididae: Lophaxius.
Family Eucalliacidae: Calliaxina∗ ) sensu nov.
∗ ) Genus with 2-3 forms of male Plp2.
12
CRM 013 – Katsushi Sakai
5. Genus with unknown male Plp2
Family Pseudogourretiidae (new status): Pseudogourretia.
6. Genera with unknown male Plps1-2
Family Axiidae: Amakusaxius gen. nov. (1 sp.); Balssaxius gen. nov. (1 sp.); Colemanaxius gen. nov. (1 sp.); Pillsburyaxius gen. nov. (1 sp.).
Family Calocarididae: Lophaxiopsis gen. nov. (1 sp.).
Family Callianassidae: Trypaea (12 spp.).
Family Callianopsidae: Pleurocalliax gen. nov. (1 sp.).
Family Gourretiidae: Laurentgourretia (1 sp.); Paracalliax (1 sp.).
Family Lipkecallianassidae (new status): Lipkecallianassa (1 sp.).
SUPERFAMILY AXIOIDEA HUXLEY, 1879
Not: Axioidea sensu De Saint Laurent, 1979: 1395; Poore, 1994: 96; Poore, 2004: 173.
Diagnosis. — Carapace without cardiac sulcus and linea thalassinica.
Maxilla 2 scaphognathite with or without posterior whip. Uropodal exopod
with or without distal flap. Setal row present or absent.
Remarks. — De Saint Laurent (1979) included three families (Axiidae Huxley, 1879; Callianideidae Kossmann, 1880; and Callianassidae Dana, 1852a)
in the superfamily Axioidea. Poore (1994, 2004) included four families (Axiidae Huxley, 1879; Calocarididae Ortmann, 1891 [as: Calocaridae]; Micheleidae Sakai, 1992c; and Strahlaxiidae Poore, 1994). In the present classification,
however, nine families are included in the Axioidea.
Families included. — Axiidae Huxley, 1879; Callianideidae Kossmann,
1880; Calocarididae Ortmann, 1891; Coralaxiidae Sakai & De Saint Laurent,
1989; Eiconaxiidae Sakai & Ohta, 2005; Eiconaxiopsididae fam. nov.; Meticonaxiidae Sakai, 1992c; Micheleidae Sakai, 1992c; and Strahlaxiidae Poore,
1994.
K EY
TO THE
FAMILIES
OF THE
S UPERFAMILY A XIOIDEA
1 – Uropodal exopod with distal flap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Uropodal exopod without distal flap and lateral notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 – Male Plp2 endopod bearing distally un- or bisegmented appendix masculina with
slender appendix interna with hooklets mesioproximally; hermaphroditic . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calocarididae
– Male Plp2 endopod bearing mesiomedially appendices masculina and interna, but
sometimes only appendix interna; gonochoristic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – P2 chela ovate; Mxp3 exopod with distal segment bending from distal margin of
penultimate segment with acute angle; no epipod on Ps1-4 . . . . . . . . . . . . . . Coralaxiidae
– P2 chela elongate; Mxp3 exopod with distal segment elongate, arising straight from
penultimate segment; epipod present on Ps1-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Axiidae
4 – Maxilla 2 scaphognathite with posterior whip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Maxilla 2 scaphognathite without posterior whip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5 – Plps2-5 without respiratory filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Meticonaxiidae
– Plps2-5 with respiratory filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – Rostrum pointed apically; dorsal surface flat . . . . . . . . . . . . . . . . . . . . . . . . . Callianideidae
14
CRM 013 – Katsushi Sakai
– Rostrum small, short, and obtusely triangular between orbits, located between branchiostegals, protruded to proximal half of eyestalks . . . . . . . . . . . . . . . . . . . . . Micheleidae
7 – Uropodal endopod trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strahlaxiidae
– Uropodal endopod oval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 – Male Plp1 absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eiconaxiidae
– Male Plp1 present, consisting of elongate, bisegmented proximal segment and circular
(regularly rounded) distal segment . . . . . . . . . . . . . . . . . . . . . . Eiconaxiopsididae fam. nov.
FAMILY AXIIDAE HUXLEY, 1879
Axiidae Huxley, 1879: 785; Bate, 1888: 36; Ortmann, 1891: 46; Stebbing, 1893: 187; Ortmann,
1898: 1141; Alcock, 1901: 186; Borradaile, 1903: 536; Borradaile, 1907: 476 (key);
Lagerberg, 1908: 47; Wollebæk, 1908: 3; Stebbing, 1910: 367; Balss, 1914: 85; Selbie, 1914:
88; Pesta, 1918: 190; Schmitt, 1921: 110; Balss, 1925: 209; Rathbun, 1929: 25; Gurney,
1938: 340; Makarov, 1938: 48; Bouvier, 1940: 93; Edmondson, 1946: 256; Barnard, 1950:
497 (key), 498; Vinogradov, 1950: 222; Balss, 1957: 1579; De Saint Laurent, 1979: 1395;
Sakai, 1987a: 303 (list); De Saint Laurent, 1988: 59; Kensley, 1989: 964; Sakai & De Saint
Laurent, 1989: 4; Sakai, 1992a: 158; Dworschak, 1992: 214; Poore, 1994: 96; Sakai, 1994:
175; Hendrickx, 1995: 386 (key), 392, figs.; Poore, 2004: 173; Sakai & Ohta, 2005: 88; Poore
& Collins, 2009: 222.
Axiinae — Sakai & De Saint Laurent, 1989: 11.
Diagnosis. — Carapace more or less compressed from each side, bearing
lateral lobes on posterior margin. Rostrum triangular, and distinct or short
with acute or obtuse apex. Linea thalassinica absent; cervical groove distinct
at full length. Gastric region flat, convex or concave, usually bearing lateral,
submedian, and median carinae. Postcervical carina present or absent. Eyestalks discernible; eyes rounded, and cornea usually pigmented. A2 scaphocerite distinct or reduced. Maxilla 2 scaphognathite with 1-2 long setae on
posterior margin. Mxp3 with exopod. Ps1-2 chelate. Ps3-4 propodi linear or
broadened; P4 coxa more or less cylindrical. Ps1-5 bases and ischia fused.
Abdominal somite 1 protruded posteroventrally; abdominal somite 2 less than
twice as long as somite 3. Male Plp1 uniramous and un- or bisegmented, or
absent; male Plp2 biramous, slender or foliaceous, endopod usually bearing
appendices interna and masculina, but sometimes only appendix interna; male
Plps3-5 usually with appendix interna; female Plp1 uniramous, un- or bisegmented, consisting of proximal segment and multiarticulate flagellum; female
Plps2-5 biramous, endopods with appendix interna. Telson without proximal
lobate process on each lateral margin. Uropodal endopod oval; uropodal exopod with or without transverse suture. Gonochoristic or hermaphroditic.
Remarks. — The Axiidae include 44 genera, 39 of which are characterized
by the uropodal exopod usually bearing a distal flap bordered by a transverse
suture. In Axius stirhynchus Leach, 1815, the type species of Axius Leach,
1815, type genus of the family, the uropodal exopod bears a distal flap, though
in a larger specimen examined the uropodal exopod seems to lack a distal
16
CRM 013 – Katsushi Sakai
flap, because it is indistinct and virtually indiscernible. In the following five
genera: Allaxius Sakai & De Saint Laurent, 1989, Anophthalmaxius De Man,
1905, Michelaxiopsis Poore & Collins, 2009, Parascytoleptus Sakai & De
Saint Laurent, 1989, and Pseudoaxiopsis gen. nov., the uropodal exopod lacks
a distal flap regardless of size.
Type genus. — Axius Leach, 1815.
Genera included. — Acanthaxius Sakai & De Saint Laurent, 1989; Albatrossaxius gen. nov.; Alienaxiopsis gen. nov.; Allaxiopsis gen. nov.; Allaxius
Sakai & De Saint Laurent, 1989; Amakusaxius gen. nov.; Anophthalmaxius
De Man, 1905; Australocaris Poore & Collins, 2009; Axiopsis Borradaile,
1903; Axiorygma Kensley & Simmons, 1988; Axius Leach, 1815; Balssaxius gen. nov.; Boasaxius gen. nov.; Bouvieraxius Sakai & De Saint Laurent,
1989; Bruceaxius gen. nov.; Calaxius Sakai & De Saint Laurent, 1989; Calocarides Wollebæk, 1908; Coelhocalaxius gen. nov.; Colemanaxius gen. nov.;
Dorphinaxius Sakai & De Saint Laurent, 1989; Eutrichocheles Wood-Mason,
1876b; Guyanacaris gen. nov.; Heterocalaxius gen. nov.; Leonardsaxius gen.
nov.; Levantocaris Galil & Clark, 1993; Litoraxius Komai & Tachikawa,
2007; Manaxius Kensley, 2003 (sensu nov.); Marianaxius Kensley, 1996e;
Michelaxiopsis Poore & Collins, 2009; Newzealandaxius gen. nov.; Oxyrhynchaxius Parisi, 1917; Parascytoleptus Sakai & De Saint Laurent, 1989; Paratrichocheles gen. nov.; Paraxiopsis De Man, 1905; Paraxius Bate, 1888; Pilbaraxius Poore & Collins, 2009; Pillsburyaxius gen. nov.; Planaxius Komai &
Tachikawa, 2008; Platyaxiopsis gen. nov.; Platyaxius Sakai, 1994; Pseudoaxiopsis gen. nov.; Ralumcaris gen. nov.; Scytoleptus Gerstaecker, 1856; Spongiaxius Sakai & De Saint Laurent, 1989.
K EY
TO THE
G ENERA
OF THE
FAMILY A XIIDAE
1 – Uropodal exopod without distal flap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Uropodal exopod with distal flap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2 – Male Plp1 absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Male Plp1 present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3 – Pleura 2-5 smooth and acutely angulated ventrally . . . . . . . Newzealandaxius gen. nov.
– Pleura 2-5 smooth and rounded or angulated ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – Eyestalks forming flat, quadrate plate; gastric region flat without submedian carinae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anophthalmaxius
– Eyestalks subglobose; gastric region convex, with five carinae . . . . . . . . . . . . . . . . . . . . 5
5 – Gastric lateral carinae with some obscure tubercles, median carina with two spinules,
running in anterior half; and two buttressed spinules between median and lateral
carinae; abdominal somites 3-5 without denticle ventrally . . . . Platyaxiopsis gen. nov.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
17
– Gastric lateral and submedian carinae smooth, median carina denticulate in anterior
half, bearing distinct hepatic tubercle, abdomial somites 3-5 with denticle ventrally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platyaxius
6 – Male Plp2 endopod with appendices interna and masculina . . . . . . . . . . . . . . Scytoleptus
– Male Plp2 endopod with appendix interna, but without appendix masculina . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parascytoleptus
7 – Male Plp1 absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– Male Plp1 present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
8 – Abdominal terga with carinae dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
– Abdominal terga without carinae dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
9 – Dorsal carinae of abdominal terga linearly striated . . . . . . . . . Pseudoaxiopsis gen. nov.
– Dorsal carinae of abdominal terga broadened and swollen . . Coelhocalaxius gen. nov.
10 – Telson convergent posteriorly to form subtriangle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
– Telson subquadrate or quadrate, and uropodal exopod with narrow distal flap . . . . . . 12
11 – Uropodal exopod bearing onion-shaped distal flap with strong distal spine . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alienaxiopsis gen. nov.
– Uropodal exopod bearing narrow distal flap . . . . . . . . . . . . . . . Heterocalaxius gen. nov.
12 – Rostrum short and triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
– Rostrum elongate and triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
13 – Gastric region without submedian carina dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
– Gastric region with submedian carina dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14 – Rostrum proximally with one pair of distinct spines . . . . . . . . . . . Ralumcaris gen. nov.
– Rostrum proximally with two pairs of distinct spines . . . . . . Paratrichocheles gen. nov.
15 – Rostrum trispinose distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
– Rostrum pointed distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
16 – Rostrum trispinose distally, median spine curved downward . . . . Allaxiopsis gen. nov.
– Rostrum trispinose distally, median spine not curved downward . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colemanaxius gen. nov.
17 – Rostrum without lateral spines, but with a pair of proximal spines . . . . . Dorphinaxius
– Rostrum with lateral spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 – Rostrum overreaching eyestalks . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amakusaxius gen. nov.
– Rostrum shorter than eyestalks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allaxius
19 – Gastric submedian carinae smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Axiorygma
– Gastric submedian carinae with denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 – Abdominal pleura sharply or moderately angulated ventrally . . . . . . . . . . . . . . . . . . . . 21
– Abdominal pleura truncate, rounded, or convex ventrally . . . . . . . . . . . . . . . . . . . . . . . . 23
21 – Gastric median carina linear . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaxius
– Gastric median carina tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 – Gastric median carina tuberculate anterior to median tubercle, and divided into two
subparallel carinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Albatrossaxius gen. nov.
– Gastric median carina tuberculate, and not divided into two carinae . . . . . . . . Manaxius
23 – Postcervical carina present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
– Postcervical carina absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
18
CRM 013 – Katsushi Sakai
24 – Fingers of cheliped slender and longer than palm . . . . . . . . . . . Pillsburyaxius gen. nov.
– Fingers of cheliped shorter than palm . . . . . . . . . . . . . . . . . . . . . . . . . Balssaxius gen. nov.
25 – Fingers of cheliped slender and longer than palm . . . . . . . . . . . . . . . . . . . . . . Acanthaxius
– Fingers of cheliped shorter than palm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 – Male Plp2 endopod with appendix interna, but without appendix masculina . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Michelaxiopsis
– Male Plp2 endopod with appendices interna and masculina . . . . . . . . . . . . . . . . . . . . . . 27
27 – Many tubercles among five gastric carinae; P1 palms smooth on lateral surface . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Axiopsis
– No tubercles among five gastric carinae; P1 palms tuberculate on lateral surface . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calocarides
28 – Dactylus of larger cheliped semicircular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spongiaxius
– Dactylus of larger cheliped slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 – Gastric region without submedian carinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
– Gastric region with five carinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
30 – Rostrum distinct and triangular [male Plp1 uniramous and bisegmented, distal segment
spatulate, bearing mesiodistally patch of hooklets; hermaphroditic] . . . . . Levantocaris
– Rostrum short and triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31 – Rostrum lacking lateral spines; telson with row of 4 mobile postlateral spines;
male Plp1 uniramous and unsegmented; male Plp2 endopod with distinct appendix
masculina medially, but without appendix interna; Plps3-5 endopods without appendix
interna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marianaxius
– Rostrum bearing lateral teeth; telson without postlateral spines . . . . . . . . . . . . . . . . . . 32
32 – Male Plp1 uniramous, bisegmented, and rod-shaped; distal segment shorter than
proximal one; male Plp2 endopod with appendix interna, but without appendix
masculina; Plps3-5 endopods with appendix interna . . . . . . . . . . . . . . . . . . . . . . . Paraxius
– Male Plp1 uniramous, unsegmented, spatulate, mesially with hooks, and distally bilobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Planaxius
33 – Gastric submedian carinae smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
– Gastric submedian carinae with spines or denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
34 – A2 scaphocerite short, and bifurcate distally . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraxiopsis
– A2 scaphocerite elongate, and simple or armed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35 – A2 scaphocerite with basal mesial spine, and spines on ventral margin . . Australocaris
– A2 scaphocerite without basal mesial spine and spines on ventral margin . . . . . . . . . 36
36 – Telson unarmed on lateral margins; maxilla 2 scaphognathite with two posterior whips;
median and submedian gastric carinae present; male Plp1 uniramous and bisegmented;
distal segment narrower than proximal one, tapering distally . . . . . . . . . . . . . . . . . . Axius
– Telson armed with spines on lateral margins; maxilla 2 scaphognathite with one
posterior whip; submedian and median gastric carinae distinctly present, bearing no
denticles; male Plp1 uniramous and bisegmented, distal segment elongate and longer
than proximal one . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leonardsaxius gen. nov.
37 – Postcervical carina absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
– Postcervical carina present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
38 – Gastric submedian carina with row of three spines; A2 scaphocerite distinctly protruded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bouvieraxius
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
19
– Gastric submedian carina with row of denticles; A2 scaphocerite horn-shaped . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boasaxius gen. nov.
39 – Eyestalks distinct, reaching nearly to distal end of rostrum . . . . . . . . . . . . . . . . . . . . . . 40
– Eyestalks subglobose, not reaching middle of rostrum . . . . . . . . . . . . . . . . . . . . . . . . . . 42
40 – Eyestalks elongate; carapace covered with scale-like tubercles . . . . . . Oxyrhynchaxius
– Eyestalks stout; carapace tuberculate, or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41 – Carapace tuberculate; P1 chelae without prominent tufts of plumose setae . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Guyanacaris gen. nov.
– Carapace not tuberculate; P1 chelae with prominent tufts of plumose setae . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoraxius
42 – Chelipeds setose; A2 scaphocerite short, simple, bi- or trifurcate; male Plp1 uniramous
and unsegmented, as stubby protrusion with truncate or slightly pointed apex; male
Plp2 biramous, endopod with appendix masculina bearing distal setae, but without
appendix interna; male Plps3-5 endopods lacking appendix interna . . . Eutrichocheles
– Body setose; A2 scaphocerite short, horn-shaped, and directed outside dorsodistal
spine of A2 segment 2; male Plp1 uniramous and bisegmented, distal segment ovate;
male Plp2 biramous, endopod and exopod elongate, endopod with appendices interna
and masculina at proximal third; male Plps3-5 endopods with appendix interna . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bruceaxius gen. nov.
Genus Acanthaxius Sakai & De Saint Laurent, 1989
Acanthaxius Sakai & De Saint Laurent, 1989: 4, 12, 14, 66; Poore, 1994: 98 (key); Sakai, 1994:
192; Kensley, 1996b: 70; Ngoc-Ho, 2006b: 58; Poore & Collins, 2009: 224.
Diagnosis. — Rostrum narrow, protruded anteriorly with pointed tip, lateral
margins extending posteriorly onto gastric region. Gastric region slightly
convex with 5-7 parallel rows of spines, equivalent to median, submedian, and
lateral carinae. Anterolateral margin of carapace usually unarmed. Cervical
groove present at full length. Postcervical carina absent. Cardiac prominence
present. Eyestalks subglobose; cornea pigmented. A2 scaphocerite protruded
straightly forward or laterally. P1 unequal; chelae obliquely positioned, lateral
surface covered with tubercles and setae, and armed with sharp teeth on
dorsal margin; fingers of both chelipeds obviously longer than palm. No
pleurobranchs. Abdominal somites without longitudinal carinae between terga
and pleura; pleuron 1 narrow, pleura 2-5 truncate or rounded ventrally. Male
Plp1 absent, male Plp2 biramous and elongate, endopod with appendices
interna and masculina at proximal third to half; male Plps3-5 endopods
with appendix interna; female Plp1 composed of proximal segment and
multiarticulate flagellum; female Plps2-5 endopods with appendix interna.
Telson subsquare; lateral margins bearing proximal lobate process with spine,
and posterior margin with median tooth. Uropodal exopod with transverse
suture. Gonochoristic.
20
CRM 013 – Katsushi Sakai
Remarks. — The type species of the genus Acanthaxius, Axiopsis (Axiopsis)
pilocheira (Sakai, 1987a) was described from a female, so that the features of
male pleopods 1-2 of this species are uncertain. However, the males of the
other species included in Acanthaxius suggest that the male Plp1 is absent, the
male Plp2 is biramous, and that the endopod of Plp2 bears appendices interna
and masculina at its midpoint. It can safely be stated that in A. pilocheirus the
cardiac prominence is present as in A. miyazakiensis, though in the original
description of A. pilocheirus it was undescribed because of lack of attention.
Recently, Ngoc-Ho (2006b) reviewed Acanthaxius, which included the
following 10 species: A. clevai Ngoc-Ho, 2006b from the Solomon Islands;
A. formosa Kensley & Chan, 1998 from Taiwan; A. gadaletae Ngoc-Ho,
2006b from the Solomon Is.; A. grandis Kensley & Chan, 1998 from Taiwan;
A. kirkmilleri Kensley, 1996b from off Anguilla in the Caribbean Sea; A.
miyazakiensis (Yokoya, 1933) from Miyazaki, Japan; A. pilocheirus (Sakai,
1987a) from Kumano-nada, Japan; A. polyacanthus (Miyake & Sakai, 1967)
from the East China Sea; A. polychaetes Sakai, 1994 from the Great Barrier
Reef, eastern Australia; and A. spinosissimus (Rathbun, 1906) from French
Polynesia.
However, Acanthaxius polychaetes Sakai, 1994 is different from the type
species of Acanthaxius, A. pilocheirus, because in A. polychaetes the male
Plp1 is present, the granulated postcervical carina is present, P1 chelae are
thickly setose on the lateral surface, the palm of the larger cheliped is stout
and longer than the fingers, the telson is longer than wide and oblong with
a rounded posterior margin, and the flap of the uropodal exopod is broad;
whereas in A. pilocheirus, the male Plp1 is absent, the postcervical carina
is also absent, P1 chelae bear spines on the dorsal and ventral margins, the
palms of the larger and smaller chelipeds are shorter than the fingers, the
telson is as long as wide, and the flap of the uropodal exopod is narrow. Those
differences between the two species suggest that A. polychaetes Sakai, 1994
is to be excluded from Acanthaxius and is herein reclassified under the new
genus Bruceaxius gen. nov. Another species, A. kirkmilleri Kensley, 1996b
is, though similar to the type species of Acanthaxius, A. pilocheirus, also
to be excluded from Acanthaxius and is herein reclassified under the new
genus Pillsburyaxius gen. nov., because of the following characteristics: the
carapace bears an obscure postcervical carina, the A2 scaphocerite is protruded
forward, and the abdominal somites 4-5 bear a spinule at the anteroventral
angle; although the male Plp1 is unknown because the holotype is a female and
no males have been reported, the female characteristics are deemed sufficient
to establish a new genus below.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
21
Type species: Axiopsis (Axiopsis) pilocheira Sakai, 1987a, by original
designation (Sakai & De Saint Laurent, 1989). The gender of the current
generic name, Acanthaxius, is masculine.
Species included. — Acanthaxius clevai Ngoc-Ho, 2006b; A. gadaletae
Ngoc-Ho, 2006b; A. garawa Poore & Collins, 2009; A. miyazakiensis (Yokoya,
1933) [= A. polyacanthus (Miyake & Sakai, 1967); A. formosa (Kensley &
Chan, 1998)]; A. ningaloo Poore & Collins, 2009; A. pilocheirus (Sakai,
1987a) [= A. grandis Kensley & Chan, 1998; A. gathaagudu Poore & Collins,
2009]; A. spinosissimus (Rathbun, 1906).
K EY
TO THE
S PECIES
OF THE
G ENUS ACANTHAXIUS
1 – Posterior margin of cervical groove unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Posterior margin of cervical groove with denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 – Gastric region bearing double submedian carina with spines . . . . . . . . . . . . . A. gadaletae
– Gastric region bearing single submedian row of spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – P1 chelae with strong spines on dorsal and ventral margins . . . . . . . . . . . . . A. pilocheirus
– P1 chelae with few short spines on dorsal and ventral margins . . . . . . . . . . . . . . . A. clevai
4 – Gastric region with single submedian row of spines . . . . . . . . . . . . . . . . . A. spinosissimus
– Gastric region with double submedian row of spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Rostral median carina unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. miyazakiensis
– Rostral median carina denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – Abdominal somite 2 smooth on ventral margin; P1 chelae spinous laterally . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ningaloo
– Abdominal somite 2 armed with anterolateral spine; P1 chelae rugose laterally . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. garawa
Acanthaxius clevai Ngoc-Ho, 2006
Acanthaxius clevai Ngoc-Ho, 2006b: 59, figs. 1, 2, 3A-C; Poore & Collins, 2009: 225, figs. 1-3,
37.
Material examined. — MNHN Th 1492, paratype, 1 male (TL/CL, 79.0/27.0 mm); 1 ovig.
female (TL/CL, 60.0/22.0 mm), Solomon Islands, Valla Gulf, R/V “Alis” Salomon 2Expedition, Sta. 2260, 8◦ 03.5′ S 156◦ 54.5′ E, 399-427 m, 27.ix.2001, leg. Ph. Bouchet,
A. Warren, and S. Samadi.
Diagnosis. — A2 scaphocerite protruded forward. Gastric lateral carinae
with spine. Male Plp1 absent; male Plp2 with appendices interna and masculina at proximal third, appendix masculina much longer than appendix interna; male Plps3-5 endopods with appendix interna. Female Plp1 consisting
of proximal segment and multiarticulate flagellum; Plps2-4 with appendix interna.
22
CRM 013 – Katsushi Sakai
Type locality. — Valla Gulf, Solomon Islands, 8◦ 03.5′ S 156◦ 54.5′ E, 399427 m.
Distribution. — Valla Gulf, Solomon Islands; off Mermaid Reef and off
Point Leveque, WA, Australia.
Acanthaxius gadaletae Ngoc-Ho, 2006
Acanthaxius gadaletae Ngoc-Ho, 2006b: 62, fig. 3D-I.
Material examined. — MNHN Th 1495, paratype, 1 male (TL/CL, 49.0/17.0 mm), Solomon
Islands, “Alis” Salomon 1-Expedition, Sta. 1761, [no coordinates], 191-290 m, 27.ix.2001,
leg. Ph. Bouchet, B. Dayrat, A. Warren, and B. Richer de Forges.
Diagnosis. — A2 scaphocerite protruded laterally. Gastric region with
double submedian carina with spines. Male Plp1 absent; male Plp2 endopod
with appendices interna and masculina; male Plps3-5 endopods with appendix
interna.
Type locality. — Solomon Islands, 135-325 m.
Distribution. — Only known from the type locality.
Acanthaxius garawa Poore & Collins, 2009
Acanthaxius garawa Poore & Collins, 2009: 227, figs. 4, 5.
Diagnosis. — Rostrum 0.2 times length of front to cervical groove, acutely
triangular, bearing 4 lateral spines anterior to supraocular spine, weakly
continuous with lateral gastric carinae. Supraocular spines prominent. Eyestalk
1.6 times length of rostrum; cornea weakly pigmented. A1 peduncle reaching
to middle of A2 segment 5. A2 segment 1 with a spine and three spinules
on distoventral margin; segment 2 with slender distal spine, directed slightly
inwards, reaching distally to middle of A2 segment 4; scaphocerite slightly
curved, reaching distally to just beyond midpoint of segment 4; segment 3
with 4 spines on mesioventral margin; segment 4 about two-thirds length of
segment 2 (excluding distal spine), bearing distoventral spine; segment 5 about
half length of segment 4. Carapace densely spinulose. Gastric lateral carina
with 10 spines; gastric submedian carina duplicated, external row with 10-12
spines and internal with 10-12 spines; gastric median carina with about 20
spines. Mxp3 basis with spine; ischium with two spines on ventral margin;
crista dentata with 24 teeth; merus with three spines; carpus with spine.
P1 asymmetrical, propodus of larger cheliped more swollen, 1.5 times as
long as that of smaller one. In larger (left) cheliped coxa with two spines
on ventral margin; basis without spine on ventral margin; ischium with three
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
23
spines on ventral margin; merus with 4 spines on convex dorsal margin, bearing 10 spines on ventral margin, lateral face spinose distally near dorsal margin, mesial face rugose distally; carpus with 6 spines on dorsal margin and 3
spines laterally on ventral margin and one spine mesially, lateral face tubercular and mesial face with 3 spines plus other smaller ones; propodus with
a row of 7 spines on dorsal margin, and lateral row of 12 spines and obsolete mesial row on ventral margin, lateral face covered with dome-like tubercles and mesial face covered with larger, distinct tubercles, bearing spine near
gape; fixed finger 1.2 times length of dorsal palm, cutting edge straight, bearing about 20 irregular, rounded teeth; dactylus with 8 spines on dorsal margin,
lateral face with row of blunt spines and mesial face with row of 6 spines and
denticles, cutting edge as in fixed finger; both fingers bearing setae. In smaller
cheliped, coxa and ischium as in larger cheliped; merus with 5 spines on dorsal
margin and 7 spines on ventral margin; carpus with 6 spines on dorsal margin,
4 spines laterally and one spine mesially on ventral margin, lateral face weakly
spinose and mesial face with three spines; propodus with 5 spines on dorsal
margin, and lateral row of 7 spines and mesial row of 7 spines on ventral margin, lateral face weakly tuberculate and mesial face grossly tuberculate, bearing two spines near gape; fixed finger 2.5 times length of dorsal palm, cutting
edge with about 40 irregular teeth; dactylus with 6 spines on dorsal margin,
lateral face with proximal row of 5 spines and mesial face with row of 4 denticles, cutting edge as in fixed finger, both fingers setose as in larger cheliped.
P2 ischium with distal spine on ventral margin; merus with 4 spines on ventral
margin; carpus as long as chela; propodus dorsal margin 0.8 length of dactylus.
P3 merus with 5 strong spines on ventral margin; carpus with spine on ventral
margin; propodus 1.9 times as long as dactylus, bearing 7 marginal robust setae (some duplicated) ventrally. P4 missing. P5 propodus 2.5 times as long as
dactylus and subchelate; fixed finger short; dactylus with broad blade on cutting edge. Abdominal pleuron 1, 1.4 times as deep as middorsal length, and
obtuse ventrally; pleuron 2 broad, lateral length as long as dorsal length, and
rounded anteroventrally; pleura 3-5 becoming more quadrate ventrally, each
with small anteroventral tooth; pleuron 6 rounded, bearing small anteroventral tooth; pleura 1-5 slightly flared laterally, bearing no lateral crease. Plp2-5
appendix interna one quarter length of endopod; appendix masculina just exceeding appendix interna. Telson about as long as broad and broadly convex on
posterior margin with median spine, bearing proximal process with spine on
lateral margin, and minute tooth and two robust setae at posterolateral angle;
dorsal surface with two symmetrical, oblique rows of two spines. Uropodal
24
CRM 013 – Katsushi Sakai
endopod twice as long as wide, bearing two lateral and one distolateral spines,
dorsal surface with longitudinal ridge with 5 spines (including marginal one).
Uropodal exopod 1.8 times as long as wide, bearing 4 spines on distal half of
lateral margin, and fixed spine and robust seta at posterolateral angle; dorsal
surface with two longitudinal ribs (outer rib with spine) and transverse suture
with 6 spines. [Adapted from Poore & Collins, 2009.]
Remarks. — The present species, Acanthaxius garawa is similar to A.
miyazakiensis, but differs from that species, because in A. garawa the rostral
median carina is armed with a row of teeth, and the abdominal somite 2 is
armed with an anteroventral spine, whereas in A. miyazakiensis the rostral
median carina is smooth, and abdominal somite 2 is unarmed on the ventral
margin.
Type locality. — Australia, Queensland, Gulf of Carpentaria, 12◦ 10.5′ S
139◦ 56.7′ E, 59 m.
Distribution. — Known from the type locality and from 11◦ 25.2′ S 139◦
25.2′ E.
Acanthaxius miyazakiensis (Yokoya, 1933)
(fig. 1)
Axius miyazakiensis Yokoya, 1933: 51, fig. 26.
Axiopsis (Axiopsis) polyacantha Miyake & Sakai, 1967: 303, fig. 1, pl. 4A; Sakai, 1987a: 303
(list). [Type locality. — East China Sea, 27◦ 16′ N 125◦ 34′ 5′′ E, 118 m.]
Axiopsis (Axiopsis) miyazakiensis — Sakai, 1987a: 303 (list).
Acanthaxius polyacanthus — Sakai & De Saint Laurent, 1989: 73; Ngoc-Ho, 2006b: 66, fig. 4.
Acanthaxius miyazakiensis — Sakai & De Saint Laurent, 1989: 67, figs. 17-19; Kensley, 1996b:
71 (list); Komai, 2000b: 344 (list).
Acanthaxius formosa Kensley & Chan, 1998: 256, figs. 1A, 2; Komai, 2000b: 344 (list). [Type
locality. — Ping Tong Country, southwestern coast of Taiwan, 350 m.]
Material examined. — NSMT, holotype, 1 female (TL/CL, 33.0/11.3 mm, damaged), R/V
“Soyo-maru”, Sta. 305, east of southern Miyazaki-Pref., Japan, 137 m, 12.vii.1928; SMF 31971,
1 female (TL/CL, 48.0/16.0 mm), Tosa-Saga, Kochi Prefecture, 17.vi.2007, leg. S. Shinomiya;
NTOU-H-1996-8-5, holotype of Acanthaxius formosa, male (TL/CL, 62.0/21.5 mm), Ping Tong
Country, Formosa, south-western coast, sandy mud bottom, ca. 350 m.
Diagnosis. — Rostrum elongate and triangular. Eyestalks thick, barely
overreaching or reaching tip of rostrum. A2 scaphocerite directed outside.
Gastric region with double submedian row of denticles; posterior margin of
cervical groove with denticles; postcervical carina absent. Fingers of both
chelipeds slender and longer than each palm. Abdominal pleura rounded
ventrally. Abdominal terga without carinae dorsally. Male Plp1 absent; male
Plp2 endopod with appendices interna and masculina. Telson subquadrate, and
uropodal exopod with narrow distal flap.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
25
Fig. 1. Acanthaxius miyazakiensis (Yokoya, 1933). Gastric region of carapace, dorsal view.
Lectotype, 1 female (TL/CL, 33.0/11.3 mm, damaged), R/V “Soyo-maru”, Sta. 305, east of
southern Miyazaki-Prefecture, Japan, 137 m, 12.vii.1928. Scale 1 mm.
Remarks. — The cardiac prominence is present on the carapace. The two
species, Acanthaxius formosa Kensley & Chan, 1998 from Taiwan and A. polyacanthus Miyake & Sakai, 1967 from the East China Sea, are synomymous
with A. miyazakiensis, because those two species are the same as A. miyazakiensis in the characteristics of Ps1-2, tailfan, and the gastric region, which is
provided with a double submedian row of spines and lateral rows of spines, the
anterior two spines of which are strong, followed posteriorly by small spines
(fig. 1).
Type locality. — East of southern Miyazaki-Prefecture, Pacific side of
Kyushu Island, 137 m.
Distribution. — Off Miyazaki, Kyushu, 137 m (Yokoya, 1933); East China
Sea, 118 m (Miyake & Sakai, 1967); Taiwan — Tong-Kong, Ping Tong
26
CRM 013 – Katsushi Sakai
County, Su-Aou, I-Lan County, and Ta-Chi, I-Lan County, 350-400 m (Kensley & Chan, 1998); Philippines (Sakai & De Saint Laurent, 1989); New Caledonia, 137-210 m (Sakai & De Saint Laurent, 1989); Macassar Strait, Indonesia, 120-200 m (Sakai & De Saint Laurent, 1989); Solomon Islands, 191-290 m
(Ngoc-Ho, 2006b).
Acanthaxius ningaloo Poore & Collins, 2009
Acanthaxius ningaloo Poore & Collins, 2009: 234, figs. 9, 10, 38.
Diagnosis. — Rostrum 0.4 times length of front to cervical groove, acute,
with 3-4 lateral spines anterior to supraocular spine, continuous with definite
gastric lateral carinae. Supraocular spines prominent. Eyestalk 0.8 length of
rostrum, and cornea weakly pigmented. A1 peduncle reaching to proximal part
of A2 segment 5. A2 segment 1 with spine and two spinules on ventrodistal
margin; segment 2 with slender distal spine, directed slightly inwards, reaching
distally to middle of A2 segment 4; scaphocerite slender and straight, reaching
distally almost to end of segment 4; segment 3 with 4 spines on mesioventral
margin; segment 4 about half length of segment 2 (excluding distal spine),
bearing distoventral spine; segment 5 about two-thirds length of segment
4. Carapace densely spinulose; gastric lateral carina with 9 spines; gastric
submedian carina duplicated, external row with 7-10 spines and internal with
12-13 spines; gastric median carina with 21 spines. Mxp3 basis with spine;
ischium with two spines on ventral margin; crista dentata with 16 teeth; merus
with three spines on ventral margin; carpus unarmed.
P1 scarcely differentiated, and subequal in length, propodus of larger
cheliped more swollen than that of smaller one. In larger (right) cheliped, coxa
with two spines on ventral margin; basis with spine on ventral margin; ischium
with 4 spines on ventral margin; merus with 4 spines on convex dorsal margin
and 9 spines on ventral margin, lateral face spinose distally and mesial face
spinose distally; carpus with 6 spines on dorsal margin, and 4 spines laterally
and one spine mesially on ventral margin, lateral face spinose, mesial face with
spine; propodus with two rows of 4 spines each on dorsal margin, and lateral
row of 21 spines and mesial row of 8 spines ventrally, lateral face spinose, and
mesial face tuberculate, bearing 8 spines concentrated near gape; fixed finger
twice length of dorsal palm, cutting edge straight, bearing about 20 irregular,
rounded teeth; dactylus with 9 spines on dorsal margin, lateral face with
proximal row of 5 submarginal spines and mesial face with proximal spine
followed by short row of denticles, cutting edge as in fixed finger; both fingers
with setae. In smaller cheliped, coxa, ischium, and merus as in larger cheliped;
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
27
carpus with 4 spines on dorsal margin and one spine on ventral margin, lateral
face spinose and mesial face with spine; propodus with two rows of 4 spines
each on dorsal margin, and lateral row of 23 spines and mesial row of 5 spines
ventrally, lateral face spinose, and mesial face tuberculate, bearing 6 spines
concentrated near gape; fixed finger 2.5 times length of dorsal palm, cutting
edge with about 30 oblique, sharp teeth; dactylus with 10 spines on dorsal
margin, lateral face with proximal row of 5 submarginal spines, and mesial
face with row of denticles, cutting edge as in fixed finger; both fingers setose as
in larger cheliped. P2 ischium with distal spine on ventral margin; merus with
three spines on ventral margin; carpus slightly shorter than chela; propodus
dorsal margin 0.8 length of dactylus. P3 merus with 3 spines on ventral margin;
propodus 2.8 times as long as dactylus, bearing 6 marginal robust setae (some
duplicated). P4 merus with 4 spines on ventral margin; propodus 2.6 times
as long as dactylus, bearing 7 marginal robust setae (some duplicated). P5
propodus 2.9 times as long as dactylus, and subchelate, fixed finger short;
dactylus with broad blade on cutting edge. Female Plp2-5 appendix interna
one-third length of endopod. Abdominal pleuron 1 twice as deep as middorsal
length, acute ventrally; pleuron 2 broad, lateral length 1.2 times dorsal length,
and rounded anteroventrally; pleura 3-5 becoming less broad and gradually
angled ventrally; pleuron 6 with small spine on ventral margin; pleura 1-5
with lateral crease. Telson about as long as broad, and convex on posterior
margin with spine, bearing proximal process with spine on lateral margin, and
one or two minute teeth and two robust setae at posterolateral angle; dorsal
surface with two symmetrical, oblique rows of two spines. Uropodal endopod
1.6 times as long as wide, bearing three spines (including posterolateral one)
on lateral margin and median longitudinal ridge with three spines (including
marginal one) on dorsal surface. Uropodal exopod 1.6 times as long as wide,
bearing 4 spines on lateral margin, and one fixed spine and one robust seta at
posterolatral angle, dorsal surface with two longitudinal ribs (outer rib with
two spines) and transverse suture with 7 spines.
Male P1 grossly differentiated, and subequal in length, propodus of larger
cheliped more swollen than that of smaller cheliped. In larger (right) cheliped,
coxa with two small spines on ventral margin; basis with spine on ventral
margin; ischium with 5 spines on ventral margin; merus with 4 spines on
convex dorsal margin, and 10 spines on ventral margin, lateral and mesial
faces spinose distally; carpus with 6 spines on dorsal margin, and three spines
laterally and one spine mesially on ventral margin, lateral face spinose and
densely setose in dorsal half, and mesial face with spine; propodus with two
28
CRM 013 – Katsushi Sakai
rows of 4 spines each (mesial row obsolete) on dorsal margin, and lateral
row of 19 spines and mesial row of 14 spines diminishing ventrally, lateral
face spinose and densely setose in dorsal half, and mesial face tuberculate,
bearing 8 spines concentrated near gape; fixed finger as long as dorsal palm,
cutting edge straight, bearing about 20 irregular, rounded teeth; dactylus with 6
spines on dorsal margin, lateral face densely setose, bearing proximal row of 5
submarginal spines, and mesial face with proximal spine and row of denticles,
cutting edge as in fixed finger, both fingers with setae. Male Plp2 with appendix
masculina about one-third length of endopod, and attached at proximal onethird of endopod; appendix interna slightly shorter than appendix masculina.
[Adapted from Poore & Collins, 2009.]
Type locality. — Off Ningaloo North, Western Australia; 165 m.
Distribution. — Only known from the type locality.
Acanthaxius pilocheirus (Sakai, 1987)
Axiopsis (Axiopsis) pilocheira Sakai, 1987a: 296, figs. 1, 2.
Acanthaxius pilocheirus — Sakai & De Saint Laurent, 1989: 73.
Acanthaxius pilocheira — Kensley, 1996b: 71 (list).
Acanthaxius grandis Kensley & Chan, 1998: 259, figs. 1B, 3-4; Komai, 2000b: 344 (list). [Type
locality. — Ping Tong County, Taiwan, 400 m.]
Acanthaxius gathaagudu Poore & Collins, 2009: 230, figs. 6-8. [Type locality. — Shark Bay,
Western Australia, 404-407 m.]
Material examined. — USNM 231418, holotype, 1 female (TL/CL, 10.0/35.0 mm),
Kumano-nada, 60 m, 5.iv.1937, coll. I. Kubo, Tokyo University of Fisheries, det. K. Sakai;
NTOUH-1996-8-5, holotype of Acanthaxius grandis, 1 male (TL/CL, 120.0/41.1 mm), Ping
Tong County, south-western coast of Taiwan, sandy mud bottom, about 400 m.
Diagnosis. — A2 scaphocerite protruded laterally. Male Plp1 absent; male
Plp2 biramous, endopod with appendices interna and masculina at proximal
third, appendix masculina with setae, and longer than appendix interna.
Remarks. — It seems that Acanthaxius grandis is a synonym of Acanthaxius
pilocheirus (Sakai, 1987a), because A. grandis has the following characteristics in common with A. pilocheira: the rostrum is narrowly triangular with a
pair of supraorbital teeth; the gastric lateral carinae are each armed with two
triangular teeth, the submedian carinae bear four smaller teeth, respectively,
and the median carina is provided with a hepatic tubercle and 6 denticles; the
Ps1-2 fingers are longer than the palm, and the P1 chelae are asymmetrical,
bearing anteriorly-directed denticles on the dorsal margin; the telson is subquadrate, bearing a proximal lobate process on each lateral margin, and a median spine on the posterior margin. It was found in the present examination of
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
29
the type specimen of Acanthaxius grandis (NTOUH-1996-8-5, holotype) that
the cardiac prominence is present.
Type locality. — Kumano-nada, off east coast of Kii Peninsula, Japan,
360 m.
Distribution. — Japan — Kumano-nada, 360 m (Sakai, 1987a); Ping Tong
County, south-western coast, Formosa (= Taiwan), sandy mud bottom, ca.
350-400 m (Kensley & Chan, 1998); Macassar, 120-200 m (Sakai & De Saint
Laurent, 1989); Shark Bay, Western Australia, 404-407 m (Poore & Collins,
2009).
Acanthaxius spinosissimus (Rathbun, 1906)
Axius spinosissimus Rathbun, 1906: 894, fig. 50a-b.
Axiopsis (Axiopsis) spinosissima — De Man, 1925d: 6 (list), 70 (key), 98, pl. 8 fig. 18-18g.
Acanthaxius spinosissimus — Sakai & De Saint Laurent, 1989: 67; Kensley, 1996b: 70; NgocHo, 2005: 49, fig. 1.
Material examined. — USNM 30534, type, male (TL/CL, 18.5/7.5 mm), south coast of
Molokai Island, Sta. Hawaii, 8.iv.1902, leg. R/V “Albatross”, Albatross Sta. 3847, det. M.
Rathbun.
Diagnosis. — A2 scaphocerite protruded forward. Male Plp1 absent; male
Plp2 endopod with appendices interna and masculina, male Plps3-5 endopods
with appendix interna.
Remarks. — Rathbun (1906) gave no description of the sex of the holotype
from Molokai Island, Hawaii. However, the holotype has turned out to be a
male in the present examination. Recently, Ngoc-Ho (2005) reported a female
from French Polynesia.
Type locality. — South coast of Molokai Island, Hawaiian Islands, 41-43 m.
Distribution. — Hawaiian Islands — Molokai Island, 41-43 m (Rathbun,
1906); Indonesia — between Wowoni and Buton Island [= Butung Is.] (De
Man, 1925d); Nuku Hiva, Marquesas Islands, French Polynesia, 90-155 m
(Ngoc-Ho, 2005); Shark Bay, Western Australia, 404-407 m (Poore & Collins,
2009).
Genus Albatrossaxius gen. nov.
Diagnosis. — Rostrum triangular and pointed at tip; lateral margins armed
with teeth, extending posteriorly to lateral carinae of gastric region. Gastric
region forming horseshoe-shaped prominence with five longitudinal carinae;
median carina tuberculate anterior to median tubercle, and divided into two
subparallel carinae reaching posteriorly as far as submedian and lateral carinae; submedian carinae tuberculate in anterior half and smooth in posterior
30
CRM 013 – Katsushi Sakai
half, lateral carinae each with a single tubercle. Anterolateral margin of carapace unarmed. Cervical groove present. Eyestalks subglobose; cornea pigmented. A2 scaphocerite strongly protruded forward. P1 chelate and subequal;
in larger cheliped fingers shorter than palm, and palm broad and slightly longer
than wide, whereas in smaller cheliped fingers about as long as palm, and
palm about as long as wide; dorsal margin of palm unarmed. P2 chelae shorter
than palm. No pleurobranchs. Abdominal pleura smooth on surface, pleuron 2
broadly truncate ventrally, pleura 3-5 with small denticle at anteroventral angle, and pleuron 6 triangular with ventral spine. Male Plp1 absent; male Plp2
biramous and slender, and endopod with appendices interna and masculina;
male Plps3-5 endopods with appendix interna. Female Plp1 uniramous and
bisegmented, consisting of proximal segment and multiarticulate flagellum;
female Plps2-5 leaf-like, endopods with appendix interna. Telson subsquare,
bearing median tooth on posterior margin. Uropodal exopod with transverse
suture.
Remarks. — Kensley (1996e: 471) described a species of Axiopsis,
A. bythos, showing in his figures (Kensley, 1996e, fig. 1H, I) that male Plp1 is
absent, and the Plp2 endopod and exopod are slender as in Axiopsis. However,
A. bythos does not belong in Axiopsis, because in A. bythos, the gastric median
carina is tuberculate in the anterior half and divided in the posterior half into
two subparallel carinae, P1 is subequal in size and similar in shape, and the
P1 meri are thick; whereas in the type species of Axiopsis, A. serratifrons A.
Milne-Edwards, 1873, the median gastric carina is straight and denticulate, P1
is unequal in size and dissimilar in shape, and the P1 meri are slender. His
species is rather reminiscent of a species belonging to the sponge-commensal
genus Eiconaxius, in which the gastric region bears a median carina divided
medially into two subparallel carinae, P1 is subequal in size and similar in
shape, and the P1 meri are thick. Yet, his species does not belong to Eiconaxius either, in which the maxilla 2 scaphognathite has no posterior whip, and
the Ps3-5 dactyli are short, subspatulate, and armed with spinules on the ventral margin. Those differences suggest that Axiopsis bythos belongs neither to
Axiopsis nor to Eiconaxius, so that a new genus Albatrossaxius is proposed for
Kensley’s species, Axiopsis bythos.
Type species. — Axiopsis bythos Kensley, 1996e, by present designation.
The gender of the new generic name, Albatrossaxius, is masculine.
Species included. — Albatrossaxius bythos (Kensley, 1996e); Albatrossaxius sp. Tirmizi, 1983.
Etymology. — The new generic name Albatrossaxius is derived from the
name of R/V “Albatross”, by which the type specimen was collected during
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31
the United States Philippines Expedition. The name of the vessel is used as a
prefix for the existing generic name Axius.
Albatrossaxius bythos (Kensley, 1996)
Axiopsis bythos Kensley, 1996e: 469, fig. 1, table 1.
Material examined. — USNM 243508, paratype, male (TL/CL, 40.0/10.0 mm), Dumaguett,
Negros Island, Negros Oriental, Philippines, Mindanao Sea, 09◦ 15.45′ N 123◦ 22.00′ E, 279 m,
19.viii.1909, leg. Philippines Expedition, R/V “Albatross”, Sta. 5536, det. B. Kensley; USNM
243509, 1 ovig. female (TL/CL, 39.0/12.0 mm), same data as paratype.
Diagnosis. — Rostrum triangular and pointed at tip; lateral margins armed
with teeth, extending posteriorly to lateral carinae of gastric region. Gastric
region forming horseshoe-shaped prominence with five longitudinal carinae;
median carina tuberculate anterior to median tubercle, and then divided
into two subparallel carinae reaching posteriorly as far as submedian and
lateral carinae; submedian carinae tuberculate in anterior half and smooth
in posterior half, lateral carinae each with a single tubercle. Anterolateral
margin of carapace unarmed. Cervical groove present. Eyestalks subglobose;
cornea pigmented. A2 scaphocerite strongly protruded forward. P1 chelate
and subequal; in larger cheliped fingers shorter than palm, and palm broad
and slightly longer than wide, whereas in smaller cheliped fingers about
as long as palm, and palm about as long as wide; dorsal margin of palm
unarmed. P2 chelae shorter than palm. No pleurobranchs. Abdominal pleura
smooth on surface, pleuron 2 broadly truncate ventrally, pleura 3-5 with
small denticle at anteroventral angle, and pleuron 6 triangular with ventral
spine. Male Plp1 absent; male Plp2 biramous and slender, and endopod with
appendices interna and masculina; male Plps3-5 endopods with appendix
interna. Female Plp1 uniramous and bisegmented, consisting of proximal
segment and multiarticulate flagellum; female Plps2-5 leaf-like, endopods with
appendix interna. Telson subsquare, bearing median tooth on posterior margin.
Uropodal exopod with transverse suture.
Type locality. — Off Apo Island, Philippines, 9◦ 15′ 45′′ N 123◦ 11′ 00′′ E,
“Albatross” Sta. 5536, 510 m.
Distribution. — Philippines — off Apo Island (Kensley, 1996e); 510 m.
Albatrossaxius sp.
Axius sp., Tirmizi, 1983: 94, fig. 5.
Remarks. — Tirmizi described the present species with figures as a species
in the genus Axius. However, it is different from the species included in Axius,
32
CRM 013 – Katsushi Sakai
but similar to the type species of Albatrossaxius, A. bythos (Kensley, 1996e),
because the rostrum is triangular, though unarmed on the lateral margins; the
eyestalks are stout; the A2 scaphocerite extends to the end of the penultimate
segment; the chelipeds are stout, the palm of the larger cheliped is thick and
granulate in its distal part, and the cutting edge of the fixed finger is denticulate.
She failed to make a precise description of the abdomen except for its length
(ca. 7 mm) of her fragmentary specimen, which is possibly a female.
Locality. — Close to Tg. Ani, Piru Bay, Ceram, Maluku Islands, Indonesia,
72-78 m, mud; R/V “Samudera”, Sta. 1/3, 8.i.1973, leg. Lon-Rumphis.
Genus Alienaxiopsis gen. nov.
Definition. — Rostrum acutely triangular with pointed tip; lateral margins
each armed with subdistal spine and proximal spine anterior to supraorbital
spine; median gastric carina distinct with hepatic tubercle and terminated with
pyramidal protuberance. Anterolateral margin of carapace triangularly convex
just below supraorbital spine. Cervical groove entire. Eyestalks subglobose;
cornea pigmented. A2 scaphocerite strongly protruded forward. P1 chelate
and subequal. P2 chelate. Ps3-4 propodi with rows of ventrolateral spinules
and strong ventrodistal spine; dactyli simple, tapering distally. Abdominal
pleura smooth laterally, without longitudinal suture between terga and pleura,
pleura 2-5 truncate ventrally, pleura 3-4 with spinule at anteroventral angle.
Male Plp1 absent; male Plp2 endopod with appendices interna and masculina;
male Plps3-5 endopods with appendix interna. Telson convergent posteriorly,
bearing 4-5 spines on lateral margin and median spine on posterior margin.
Uropodal exopod bearing onion-shaped distal flap with strong distal spine.
Remarks. — The type species of the new genus Alienaxiopsis, A. lizardensis
sp. nov. is defined as follows: the rostrum is acutely triangular with two
lateral spines, the median carina is terminated with a distinct pyramidal
gastric protuberance, the A2 scaphocerite is elongate, and the distal flap
of the uropodal exopod is protruded and onion-shaped. The other species,
Alienaxiopsis clypeata (De Man, 1888b), once included in the genus Allaxius,
is rather similar to A. lizardensis sp. nov. and considered to be a member of
Alienaxiopsis, because it has turned out to be very different from the type
species of Allaxius, A. aethiopicus (Nobili, 1904), in which the rostrum is
triangular with 3-5 sharp lateral spines, the gastric region bears no median
carina but a median row of spines, the A2 scaphocerite is short and hornshaped, and the distal flap of the uropodal exopod is short and wider than long.
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33
In consequence, Allaxius clypeatus (De Man, 1888b) is reclassified under the
new genus Alienaxiopsis.
Type species. — Alienaxiopsis lizardensis sp. nov., by present designation.
The gender of the new generic name is feminine.
Species included. — Alienaxiopsis lizardensis sp. nov.; A. clypeata (De
Man, 1888b).
Etymology. — The new generic name Alienaxiopsis is a compound of
the Latin word alienus, meaning “strange”, and the generic name Axiopsis,
because this new genus is distinctly different in morphological features,
enough to be separated from Axiopsis, even though it has the same pleopodal
form.
K EY
TO THE
S PECIES
OF THE
G ENUS ALIENAXIOPSIS
1 – Lateral carina in gastric region distinctly developed, and terminated with pyramidal
protuberance; another protuberance present as a submedian carina on each side of
median carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. lizardensis sp. nov.
– Lateral and submedian carinae in gastric region not developed, but three teeth present
on each side of median carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. clypeata
Alienaxiopsis clypeata (De Man, 1888)
Axius clypeatus De Man, 1888b: 470, pl. 20 fig. 2.
Axiopsis (Axiopsis) clypeata — De Man, 1925d: 5 (list), 70 (key).
Allaxius clypeatus — Sakai & De Saint Laurent, 1989: 75; Kensley, 2003: 360; Poore & Collins,
2009: 237.
Diagnosis. — Rostrum acutely triangular with pointed tip; lateral margins
each armed with subdistal spine and proximal spine anterior to supraorbital
spine; gastric median carina distinct, with hepatic tubercle, and terminated
with pyramidal protuberance. Three short teeth present on either side of
median carina. Telson longer than wide, bearing two pairs of spines on
dorsal surface. Uropodal endopod with three teeth on lateral margin, posterior
margin denticulate with mesial and lateral spines, and median carina with 4
spines; uropodal exopod bearing 4 spines on lateral margin, 4 small spines
on sublateral margin, two spines on distal margin, distinct movable spine at
distolateral angle, and triangular movable distal flap with fine marginal spines
and strong distal spine (De Man, 1888b, pl. 20 fig. 2; 1925d: 70).
Remarks. — The present species A. clypeata is similar to the type species
of Alienaxiopsis, A. lizardensis sp. nov., but different in the characteristics of
the gastric region.
Type locality. — Ambon, Indonesia.
Distribution. — Indonesia — Ambon (De Man, 1888b); Yonge Reef, Lizard
Island, Queensland, Australia (Poore & Collins, 2009).
34
CRM 013 – Katsushi Sakai
Alienaxiopsis lizardensis sp. nov.
(fig. 2)
Material examined. — AM P25014, holotype, male (TL/CL, 16.0/4.9 mm), Yong Reef, near
Lizard Island, Queensland, 14◦ 38′ S 145◦ 38′ E, reef, 8.xi.1975, leg. N. Coleman.
Fig. 2. Alienaxiopsis lizardensis sp. nov. A, whole body, lateral view; B, carapace, dorsal view;
C, telson; D, uropod on right side. A-D, AM P25014, holotype, male (TL/CL, 16.0/4.9 mm),
Yong Reef, near Lizard Island, Queensland, reef. Scales 1 mm.
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35
Diagnosis and description of male holotype. — Rostrum (fig. 2B) acutely
triangular with pointed tip, lateral margins each with subdistal spine and
proximal spine anterior to supraorbital spine, extending posteriorly to gastric
lateral carinae terminated with pyramidal protuberance; median carina thick
with hepatic tubercle and terminated with pyramidal protuberance, another
protuberance as submedian carina on each side of median carina. Cervical
groove distinct and unarmed. Posteromedian carina absent. Posterolateral lobe
of carapace present. Eyestalks subglobose, reaching almost tip of rostrum.
A2 (fig. 2A, B) scaphocerite distinct and horn-shaped; A2 segment 2 with
distinct dorsodistal spine. P1 unequal; in larger cheliped ischium denticulate
on ventral margin; merus broad, slightly longer than wide, and denticulate
on ventral margin; carpus short, thick, and 1.5 times as long as wide, palm
1.5 times as long as wide, lateral surface with distinct longitudinal carina
located from distoventral part of palm to fixed finger, dorsal margin with distal
tooth; dactylus incurved distally, and deeply concave mesially along dorsal
margin; cutting edge proximally convex, and distally concave; fixed finger
short, with irregular denticles on cutting edge. In smaller cheliped palm 1.2
times as long as wide, and slightly shorter than fingers; lateral surface with two
longitudinal carinae extending to fixed finger; dorsal margin terminated with
distal tooth; fixed finger armed with denticles on proximal third of cutting
edge, then widely concave. P2 chelate, merus with median tooth on ventral
margin. Ps3-5 missing. Abdominal pleura smooth; pleuron 1 triangular, with
obtuse distal tooth ventrally, pleura 2 and 5 truncate ventrally, pleura 3-4 with
small spinule at anteroventral angle. Male Plp1 absent; male Plp2 biramous,
endopod with appendices interna and masculina; male Plps3-5 endopods with
appendix interna. Telson (fig. 2C) almost as long as wide, lateral margins
convergent distally, bearing proximal lobate process with spine, followed by
three spines; posterior margin convex and denticulate, bearing median spine;
dorsomedian surface with two pairs of distinct spines. Uropodal endopod
(fig. 2D) bearing two distinct spines on lateral margin, another distinct tooth
at distolateral angle, and small spinules and setae on distal margin, and also
bearing median carina with a row of 4 teeth, distal one of which located on
distal margin. Uropodal exopod convex on lateral margin, with four distinct
spines, bearing row of 4 denticles sublaterally; distal margin bearing distinct,
movable tooth at posterolateral angle, small median tooth, and sharp distinct
tooth at posteromesial angle; distal flap onion-shaped, with marginal spinules
and distinct distal spine.
Remarks. — The present new species, Alienaxiopsis lizardensis sp. nov. is
similar to Alienaxiopsis clypeata (De Man, 1888b), in the characteristics of
36
CRM 013 – Katsushi Sakai
rostrum, median carina, and tailfan. However, they are different from each
other, because in A. lizardensis the lateral gastric carinae are distinctly developed, and each terminated with a pyramidal protuberance; another protuberance is present as a submedian gastric carina on each side of the median
gastric carina, whereas in A. clypeata the lateral and submedian gastric carinae
are not developed, but three teeth are present on each side of the median carina.
Type locality. — Yong Reef, near Lizard Island, Queensland, 14◦ 38′ S
145◦ 38′ E, reef.
Distribution. — Only known from the type locality.
Etymology. — The species name lizardensis is derived from the type locality of the present species, Yong Reef, near Lizard Island, Queensland, with the
Latin suffix “-ensis”, indicating the locality. The name is an adjective agreeing
in gender with the generic name.
Genus Allaxiopsis gen. nov.
Allaxius Sakai & De Saint Laurent, 1989: 73; Poore, 1994: 98 (key) [partim].
Definition. — Rostrum trispinose distally, median spine curved downward;
triangular supraocular teeth present. Anterolateral margin of carapace unarmed. Cervical groove distinct. Eyestalks subglobose; cornea pigmented. A2
scaphocerite short and horn-shaped. P1 chelate and asymmetrical. No pleurobranchs. Abdominal pleura smooth on surface. Male Plp1 absent; male Plp2
biramous, endopod with appendices interna and masculina at proximal third;
male Plps3-5 endopods with appendix interna. Female Plp1 composed of proximal segment and multiarticulate flagellum; female Plps2-5 endopods with
appendix interna. Telson subsquare, bearing posteromedian tooth. Uropodal
exopod armed with distinct teeth on lateral margin, bearing distal flap with
marginal spines. [Adapted from Sakai & De Saint Laurent, 1989.]
Remarks. — The three species belonging to the new genus Allaxiopsis gen.
nov. were included in the genus Allaxius Sakai & De Saint Laurent, 1989,
though those two genera are fundamentally different from each other in the
shapes of rostrum and telson. In Allaxiopsis the rostrum is trispinose distally
and its median spine is curved downward distally, the telson is wider than
long and subsquare in shape, whereas in Allaxius the rostrum is triangular and
pointed at the apex, the telson is longer than wide and oblong in shape.
Type species. — Allaxiopsis bougainvillensis sp. nov. by present designation. The gender of the new name is feminine.
Species included. — Allaxiopsis bougainvillensis sp. nov.; A. picteti (Zehntner, 1894); A. spinimana (De Man, 1905).
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37
Etymology. — A Greek suffix “-opsis”, meaning “appearance” is added to
the generic name Allaxius, from which the new genus is different, though it is
similar in appearance.
K EY
TO THE
S PECIES
OF THE
G ENUS ALLAXIOPSIS
1 – Eyestalks reaching middle of rostrum . . . . . . . . . . . . . . . . . . . . A. bougainvillensis sp. nov.
– Eyestalks overreaching rostrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Palm of larger cheliped with conical tubercles on lateral surface . . . . . . . . . A. spinimana
– Palm of larger cheliped with imbricated sculpture on lateral and mesial surfaces . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. picteti
Allaxiopsis bougainvillensis sp. nov.
(fig. 3)
Material examined. — ZMB 14440, holotype, 1 female (TL/CL, 12.0/4.9 mm); paratype,
1 female (TL/CL, 11.0/4.5 mm, missing larger cheliped), Triop Bougainville, German New
Guinea, leg. H. Schoede.
Diagnosis of female holotype. — Rostrum (fig. 3A, B) trispinose distally,
armed with stout spine medially on lateral margins, extending posteriorly to
lateral gastric carinae. Anterolateral margin of carapace unarmed. Eyestalks
subglobose; cornea pigmented, reaching middle of rostrum. Gastric region
bearing elongate median carina with three tubercles; lateral carina with supraorbital spine at its anterior extremity, extending to cervical groove, medially
bearing a shield-shaped protuberance; distinct tooth on either side of median
carina, and some tubercles in posterior half of gastric region. Cervical groove
distinct. Posteromedian carina absent. A2 scaphocerite short and horn-shaped,
located under short distodorsal spine of A2 segment 2. Ps3-4 (fig. 3C, D) carpi
with distodorsal spine; Ps3-5 propodi with four vertical rows of two spines on
ventrolateral margin, and Ps3-5 dactyli with spines laterally. Telson (fig. 3E)
wider than long, bearing proximal lobate process with spine, followed distally by 2-3 strong spines on lateral margin, and another lobate process dorsally between two proximal ones on lateral margin and posterior to these a
pair of spines; posterior margin convex, with median spine. Uropodal endopod
bearing 11 distinct spines on posterior margin, three distinct spines on lateral
margin, distinct movable spine mesial to posterolateral spine, and median carina with three spines, distal one of which distinct. Uropodal exopod bearing
5 distinct spines on lateral margin, a row of three spines sublaterally in proximal half, transverse distal suture with 5 teeth, and distal flap with 5 distinct
spines.
Remarks. — Allaxiopsis bougainvillensis sp. nov. is similar to A. picteti
(Zehntner, 1894), however, they differ from each other. In A. bougainvillensis
38
CRM 013 – Katsushi Sakai
Fig. 3. Allaxiopsis bougainvillensis sp. nov. A, carapace, dorsal view; B, anterior part of
carapace, lateral view; C, P3; D, P4; E, tail-fan. A-E, ZMB 14440, lectotype, 1 female (TL/CL,
12.0/4.9 mm); Triop Bougainville, German New Guinea. Scales A, E, 1 mm; B, C, D, 0.5 mm.
the eyestalks reach the middle of the rostrum, the gastric region bears a median
carina with three tubercles, and the lateral carinae extend to the cervical
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39
groove, with a shield-shaped protuberance medially, whereas in A. picteti the
eyestalks overreach the tip of the rostrum, and the gastric region bears neither
median nor lateral carinae, but spiniform teeth.
The present type specimens are registered as Axiopsis (Axiopsis) picteti
var. spinimana De Man, 1905 in the collection of the Zoological Museum of
Berlin, but they are not to be considered the type specimens of the species,
because those two specimens examined are females measuring 11 mm and
12 mm in body length from Bougainville, German New Guinea, leg. H.
Schoede, whereas De Man described an egg-bearing female measuring 25 mm
in body length and a younger male from off the south point of KabaënaIsland, Indonesia, “Siboga” Sta. 209, 22 m (De Man, 1905: 597; 1925d: 96).
In consequence, the specimens examined are determined as a new species, A.
bougainvillensis sp. nov.
Type locality. — Anchorage off south of Kabaena Island, Indonesia, “Siboga” Sta. 209, 22 m.
Distribution. — Indonesia — Anchorage off south of Kabaena Island, 22 m
(De Man, 1905, 1925d); Papua New Guinea — Bougainville; 22 m.
Etymology. — The species name bougainvillensis is derived from the type
locality of the specimens, Triop Bougainville, German New Guinea. It is an
adjective agreeing in gender with the generic name.
Allaxiopsis picteti (Zehntner, 1894)
Paraxius Picteti Zehntner, 1894: 196, pl. 9 fig. 25.
?Axiopsis Picteti — Borradaile, 1903: 539.
Axiopsis (Axiopsis) Picteti — De Man, 1925d: 6 (list), 70 (key), 92, pl. 7 fig. 16-16b.
Allaxius picteti — Sakai & De Saint Laurent, 1989: 75; Kensley, 2003: 361, pls. 5, 6.
Material examined. — MHNG, holotype, female (TL/CL, 29.0/10.0 mm), Amboina, Indonesia.
Diagnosis. — Rostrum narrowly triangular and trispinose distally, bearing subdistal teeth and proximal supraocular tooth on lateral margin, and a
swelling ventrally; the slender median spine of the distal three curved downward, eyestalks overreaching rostrum. Gastric region bears distinct tubercles,
lacking lateral and median carinae, anterior extremity of gastric region narrow
and triangular, reaching as far forward as supraocular spines. P1 unequal in
size and dissimilar in shape; palm in both larger and smaller chelipeds with
imbricated sculpture on lateral and mesial surfaces. Telson wider than long.
[Cf. Zehntner, 1894: 196, pl. 9 fig. 25a.]
Type locality. — Ambon, Indonesia.
40
CRM 013 – Katsushi Sakai
Distribution. — Indonesia — off Kabaena Island (De Man, 1925d); Ambon
(Zehntner, 1894).
Allaxiopsis spinimana (De Man, 1905)
Axiopsis Picteti var. spinimana De Man, 1905: 597.
Axiopsis (Axiopsis) Picteti var. spinimana — De Man, 1925d: 6 (list), 70 (key), 96, pl. 7 fig. 1717a.
Allaxius spinimanus — Sakai & De Saint Laurent, 1989: 75.
Diagnosis. — Rostrum narrowly triangular and trispinose at apex, eyestalks
overreaching rostrum; gastric region bearing lateral carina with three teeth,
and submedian carina with 5 teeth. Gastric median carina originating from
proximal part of rostrum between supraocular teeth, running posteriorly to
hepatic denticle, bearing two anterior denticles, and branched into two carinae
with two denticles each. P1 unequal in size and dissimilar in shape; palm in
both larger and smaller chelipeds with conical tubercles on lateral surface.
Telson wider than long. [Cf. De Man, 1925d.]
Remarks. — The present species, Allaxiopsis spinimana is similar to
A. picteti, however, the two species differ as follows: in A. spinimana the palm
of the larger cheliped bears conical tubercles on the lateral surface and 7 spines
on the dorsal margin (De Man, 1925b, pl. 7 fig. 17), whereas in A. picteti the
palm of the larger cheliped bears imbricated sculpture on the lateral and mesial
surfaces, and is smooth on the dorsal margin (Zehntner, 1894, pl. 9 fig. 25a).
Type locality. — “Siboga” Sta. 209, off the south point of Kabaëna-Island,
Indonesia, 22 m.
Distribution. — Only known from the type locality.
Genus Allaxius Sakai & De Saint Laurent, 1989
Allaxius Sakai & De Saint Laurent, 1989: 73; Poore, 1994: 98 (key); Poore & Collins, 2009:
225, figs. 1-3, 37.
Diagnosis. — Rostrum triangular, with 3-5 sharp lateral teeth. Gastric region
convex, bearing longitudinal rows of spinules, or irregularly arranged spinules.
Anterolateral margin of carapace unarmed. Cervical groove distinct except for
anterolateral part. Eyestalks subglobose; cornea pigmented. A2 scaphocerite
short and horn-shaped. P1 chelate and asymmetrical; chelae unarmed on dorsal
margin. No pleurobranchs. Abdominal pleura smooth on surface and unarmed
on ventral margin. Male Plp1 absent; male Plp2 endopod with appendices
interna and masculina; male Plps3-5 endopods with appendix interna. Female
Plp1 composed of proximal segment and multiarticulate flagellum; female
Plps2-5 endopods with appendix interna. Telson longer than wide, bearing 2-3
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
41
pairs of spines on dorsal surface, proximal lobe with spine on lateral margin,
and median tooth on convex posterior margin. Uropodal exopod armed with
teeth on lateral margin and with transverse suture. [Adapted from Sakai & De
Saint Laurent, 1989.]
Remarks. — The genus Allaxius Sakai & De Saint Laurent, 1989, was established on the basis of Axiopsis aethiopica Nobili, 1904, because A. aethiopica
is distinguishable from the type species of Axiopsis, A. serratifrons. They are
fundamentally different from each other in the features of the gastric region
and the A2 scaphocerite: in Axiopsis aethiopica the gastric region bears longitudinal rows of spinules, and the A2 scaphocerite is short and horn-shaped,
whereas in Axiopsis serratifrons the gastric region has a horseshoe-shaped
prominence bearing five longitudinal carinae with denticles, and numerous
denticles scattered among those carinae, and the A2 scaphocerite is elongate.
Type species. — Axiopsis aethiopica Nobili, 1904, by original designation
and monotypy. The gender of the current generic name, Allaxius, is masculine.
Species included. — Allaxius aethiopicus (Nobili, 1904).
Allaxius aethiopicus (Nobili, 1904)
(figs. 4, 5)
Axiopsis aethiopica Nobili, 1904: 235; Tattersall, 1921: 394; Dworschak, 1992: 215, fig. 15a-f.
Fig. 4. Allaxius aethiopicus (Nobili, 1904). Whole body, lateral view. ZMB 15555, ovig. female
(TL/CL, 39.0/12.0 mm), Republic of Djibouti. Scale 1 mm.
42
CRM 013 – Katsushi Sakai
Fig. 5. Allaxius aethiopicus (Nobili, 1904). A, gastric region of carapace, dorsal view; B, gastric
region of carapace, dorsal view; C, gastric region of carapace, lateral view; D, abdominal somite
6 and tail-fan, dorsal view. A, D, ZMB 15555, ovig. female (TL/CL, 39.0/12.0 mm), Republic
of Djibouti; B, C, NHMW 6894, 1 ovig. female (TL/CL, 34.0/11.0 mm), Red Sea, 18.ii.1895,
leg. Pola (?) Expedition. Scales 1 mm.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
43
Axiopsis (Paraxiopsis) aethiopica — Nobili, 1906b: 93, pl. 6 fig. 1; Balss, 1915: 1; De Man,
1925d: 6 (list), 72 (key).
Allaxius aethiopica — Sakai & De Saint Laurent, 1989: 75.
Material examined. — MNHN Th 142, syntypes, 6 females (TL/CL, 26.0/10.0-37.0/
13.0 mm), Djibouti, 1892, leg. Jousseumi; MNHN Th 143, 1 female (TL/CL, 32.0/11.0 mm),
Djibouti, 1892, leg. Jousseumi; MNHN Th 471, 1 male (TL/CL, 21.0/8.5 mm); 1 ovig.
female (TL/CL, 36.0/12.0 mm), Red Sea, “Calypso”; ZMB 15555, 1 ovig. female (TL/CL,
39.0/12.0 mm, lacking larger cheliped), Republic of Djibouti, biotope, ii-iii.1909, leg. Wache;
ZMH-K 32030, 1 ovig. female (TL/CL, 35.0/10.9 mm), Bay of Tadjura, Republic of Djibouti,
leg. E. Wache; NHMW 6894, 1 ovig. female (TL/CL, 34.0/11.0 mm), Red Sea, 18.ii.1895, leg.
Pola (?) Expedition.
Diagnosis. — Rostrum triangular, with 3-5 sharp lateral spines. Gastric
region convex (fig. 4), bearing either lateral carinae, two spines in place of
median carina, and submedian double carina (fig. 5B), or 5 rows of spines
in place of median, submedian, and lateral carinae (fig. 5A). A2 scaphocerite
horn-shaped and shorter than distodorsal spine of A2 segment 2. Mxp3 merus
with two sharp spines on mesial margin. P1 chelate and asymmetrical; in
young male larger cheliped with broad merus, 1.8 times as long as wide, dorsal
margin entire, but ventral margin denticulate proximally; carpus triangular,
ventral margin protruded as truncate tooth; palm broadened, 1.5 times as
long as wide and unarmed; dactylus about 3/4 length of palm; in smaller
cheliped merus more than twice as long as wide; carpus 1.5 times as long
as wide, and palm three times as long as wide, dactylus 2/3 times as long
as palm; cutting edges of dactylus and fixed finger unarmed. In adult female
cutting edge of dactylus roughly denticulate, and that of fixed finger with two
distinct teeth at proximal and distal third. P2 fingers longer than propodus.
P3 (fig. 4) propodus rectangular, P4 propodus elongate, Ps3-4 propodi bearing
transverse rows of spines on ventrolateral margin, and dactylus bearing row of
lateral spinules. Ps3-5 simple, bearing distoventral spine. Abdominal pleuron 1
narrow and slightly convergent posteroventrally; pleura 2-5 truncate ventrally,
pleuron 6 triangular, with ventral spine. Male Plp1 absent; male Plp2 endopod
with appendices interna and masculina at proximal third; appendix masculina
longer than appendix interna. Female Plp1 composed of proximal segment
and multiarticulate flagellum; female Plps2-5 endopods with appendix interna.
Telson longer than wide, laterally bearing proximal lobe with spine, followed
by 2-3 spines posterolaterally, two pairs of spines on dorsal surface, and
median spine on distal margin (fig. 5D). Uropodal endopod bearing spines
on distal margin, and another two spines on median carina, distal one of
which located on posterior margin; uropodal exopod spinulate on lateral
44
CRM 013 – Katsushi Sakai
margin, bearing transverse suture with spinules subdistally, and distal flap with
spinulate distal margin.
Remarks. — Nobili (1904: 235) stated in his description that “Le rostre est
pourvu de trois épines marginales sur chaque bord, et d’une épine terminale
aussi longue que les deux latéroterminales; une série de spinules sur sa surface
dorsale. Sur la carapace, il y a deux séries latérales (une pour chaque partie) de
spinules; entre ces deux séries, il y en a deux autres plus petites”. [The rostrum
bears three spines on each lateral margin, and an apical spine as large as the two
distolateral spines; a series of spines on the dorsal surface [= median carina].
The carapace is armed with a lateral row of spines on each side, and another
two rows of spinules between the median and lateral rows of spines.] However,
in the present specimens examined, the spinous rows in the gastric region are
indistinct and the number and distribution of spines vary; in the female (ZMB
15555) the gastric region bears one spine anteriorly between the median and
submedian rows of spines (fig. 5A), and in another female (ZMH K32030)
the gastric region is convex, with a triangular protuberance at short distance
from the base of the rostrum, bearing lateral carinae with a spine, no median
carina but two anterior spines mediodorsally, and submedian double carinae,
each followed anteriorly by 1-2 spines (NHMW 6894) (fig. 5B, C).
In the young male (MNHN Th 471, TL/CL, 21.0/8.5 mm) the dactylus
is 2/3 times as long as the palm; the cutting edges of the dactylus and the
fixed finger are unarmed, whereas in the adult female (NHMW 6894, TL/CL,
34.0/11.0 mm) the cutting edge of the dactylus is roughly denticulate, and that
of the fixed finger is armed with two distinct teeth at the proximal and distal
third.
Type locality. — Djibouti and Massawa.
Distribution. — Red Sea — Red Sea and Massawa (Nobili, 1904, 1906),
Haleib (Dworschak, 1992), Daedalus Riff, Mersa Halaib, Raveiyah, Khor
Dongonab. Gulf of Aden — Djibouti (Nobili, 1904, 1906).
Genus Amakusaxius gen. nov.
Definition. — Rostrum triangular, lateral margins with teeth. Anterolateral
margin of carapace unarmed. Gastric region situated on same level as rostrum, bearing five longitudinal carinae; lateral carina consisting of two long
carinate teeth, submedian carina of 5 short carinate teeth, and median carina with tubercles, reaching cervical groove; postcervical carina present. Eyestalks thick, overreaching rostrum. A2 scaphocerite short and horn-shaped.
P1 chelate; larger cheliped unknown; in smaller cheliped fingers longer than
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
45
palm. Abdominal pleura smooth laterally and unarmed ventrally. Male Plps1-2
unknown. Telson subsquare, bearing posteromedian tooth. Uropodal endopod
of narrow width; uropodal exopod also narrow, bearing distal flap. [Adapted
from Sakai & De Saint Laurent, 1989.]
Remarks. — The present new genus is established on the base of Calocaris (Calastacus) amakusana Miyake & Sakai, 1967, which was included
in the genus Acanthaxius Sakai & De Saint Laurent, 1989, because Calocaris
(Calastacus) amakusana is similar in the features of the gastric region, smaller
cheliped, and abdominal pleura to the type species of the genus Acanthaxius,
A. pilocheirus (Sakai, 1987a) from Kumano-nada, off east coast of Kii Peninsula, Japan, 360 m. However, it has turned out that Calocaris (Calastacus)
amakusana is distinctly different from A. pilocheirus in that the posterocervical carina is present, and is, therefore, not included in Acanthaxius, in which
the posterocervical carina is not present. Accordingly, Calocaris (Calastacus)
amakusana is reclassified under the new genus Amakusaxius, as A. amakusanus.
Type species. — Calocaris (Calastacus) amakusana Miyake & Sakai, 1967,
by present designation and monotypy. The gender of the new generic name is
masculine.
Species included. — Amakusaxius amakusanus (Miyake & Sakai, 1967).
Etymology. — The new generic name Amakusaxius is derived from the type
locality of the type species, Amakusa, northwestern Kyushu, Japan, with the
generic name Axius added.
Amakusaxius amakusanus (Miyake & Sakai, 1967)
Calocaris (Calastacus) amakusana Miyake & Sakai, 1967: 306, fig. 2, pl. 4B.
Axiopsis (Axiopsis) amakusana — Sakai, 1987a: 303 (list).
Acanthaxius amakusana — Sakai & De Saint Laurent, 1989: 73; Kensley, 1996b: 71 (list).
Remarks. — The holotype is a female, so it is at present impossible to
describe the forms of male Plps1-2.
Type locality. — Off Tomioka, Amakusa-shoto [= Amakusa Islands], Kumamoto Pref., Japan, 20-40 m.
Distribution. — Japan — Amakusa-shoto (Miyake & Sakai, 1967); 2040 m.
Genus Anophthalmaxius De Man, 1905
Anophthalmaxius De Man, 1905: 593; De Man, 1925d: 2 (key), 60; Sakai & De Saint Laurent,
1989: 14; Poore, 1994: 97 (key).
46
CRM 013 – Katsushi Sakai
Diagnosis. — Rostrum badly damaged at distal part; lateral margins extending onto gastric region. Anterolateral margins of carapace unarmed. Gastric region situated at same level as rostrum, bearing median carina. Cervical groove
discernible only on dorsal part, not extending to anterolateral part. Eyestalks
flat and quadrate in dorsal view; cornea unpigmented. A2 scaphocerite simple
and prominent. P1 chelate and equal; dorsal margin of chela unarmed. P2 fingers longer than palm. Ps2-4 with pleurobranchs. Abdominal pleura smooth
and unarmed ventrally. Male Plp1 absent; male Plp2 biramous and broad, endopod with appendices interna and masculina; male Plps3-5 biramous and
broad, endopods with appendix interna. Telson longer than wide, bearing posteromedian tooth. Uropodal exopod without distal flap. [Adapted from Sakai
& De Saint Laurent, 1989.]
Type species. — Anophthalmaxius eccoptodactylus De Man, 1905, by original designation and monotypy. The gender of the generic name is masculine.
Species included. — Anophthalmaxius eccoptodactylus De Man, 1905.
Anophthalmaxius eccoptodactylus De Man, 1905
(fig. 6)
Anophthalmaxius eccoptodactylus De Man, 1905: 594; De Man, 1925d: 5 (list), 61, pl. 5
fig. 11-11d, pl. 6 fig. 11e-n; Sakai & De Saint Laurent, 1989: 15.
Anophthalmaxius enoptodactylus — Balss, 1925: 210 (miss-spelling).
Material examined. — ZMA Crust De. 202840, holotype, male (TL/CL, 16.0/6.0 mm, P1
missing on right side), Banda Sea, Indonesia, 5◦ 46.7′ S 134◦ 0′ E, “Siboga” Sta. 271, 1788 m.
Diagnosis. — Rostrum badly damaged at distal part, but supposed to be
triangular and flattened dorsally as in fig. 6A-C, lateral margins extending
posteriorly to middle of gastric region. Gastric region flat, bearing longitudinal
median carina in anterior half, but no submedian carinae. Anterolateral margin
of carapace unarmed. Cervical groove discernible. Eyestalks quadrate in
dorsal view as in callianassid species. A2 segment 2 with dorsodistal spine,
scaphocerite distinct. P1 missing on right side, but present on left. Male Plp1
absent; male Plp2 endopod with appendices interna and masculina; appendix
interna with patch of hooklets distally and shorter than the appendix masculina,
which bears distal setae (fig. 6D), male Plps3-5 endopods with appendix
interna. Telson 1.5 times as long as wide, lateral margins slightly convex
proximally, bearing 2-3 spinules medially; posterior margin rounded with
median spine. Uropodal exopod without distal flap.
Remarks. — The male type specimen is fragile and partly damaged, lacking
the distal part of the rostrum, the mouthparts on the left side, P1 on the right
side, and Plp1 on the left side.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
47
Fig. 6. Anophthalmaxius eccoptodactylus De Man, 1905. A, anterior part of carapace, dorsal
view; B, anterior part of carapace, lateral view from right side; C, same from left side;
D, male Plp2. A-D, ZMA Crust De. 202840, holotype male (TL/CL, 16.0/6.0 mm), Banda
Sea, Indonesia, 5◦ 46.7′ S 134◦ 0′ E, “Siboga” Sta. 271, 1788 m. Scales 1 mm.
48
CRM 013 – Katsushi Sakai
Type locality. — Off west coast of Aru, Indonesia, 5◦ 46.7′ S 134◦ 0′ E,
“Siboga” Sta. 271, 1788 m.
Distribution. — Only known from the type locality.
Genus Australocaris Poore & Collins, 2009
Australocaris Poore & Collins, 2009: 238.
Diagnosis. — Rostrum styliform, elongate, denticulate laterally, three times
as long as eyestalks, on level with carapace, and continuous with definite lateral gastric carinae; supraocular spines (spine at anterior end of lateral gastric carina and base of rostrum) prominent. Eyestalks cylindrical and articulating; cornea unpigmented. A2 scaphocerite with mesial spine at base and
spines on ventral margin, extending beyond A2 peduncle. Carapace smooth;
lateral gastric carina unarmed except for anterior supraocular spine; submedian gastric carina present, but obsolete; median gastric carina as weak
ridge; postcervical carina absent; cervical groove short and weakly defined.
Mxp3 exopod not clearly bent at base of flagellum. P1 symmetrical; propodus broad and laterally flattened, and carinate on dorsal and ventral margins; carpus-dactylus with strong spines on dorsal and ventral margins. Pleurobranchs absent on Ps2-4; podobranchs and arthrobranchs well developed;
epipods present on Mxp2 to P4. Abdominal somite 1 pleuron produced; pleuron 2 broad, rounded anteriorly and posteriorly, and flat ventrally; pleura 35 rounded posteriorly. Plp3-5 appendix interna present. Male Plp1 minute.
Male Plp2 without appendix masculina. Uropodal exopod with transverse suture.
Type species. — Australocaris pinjarup Poore & Collins, 2009, by original
designation and monotypy. The gender of the generic name, Australocaris, is
feminine.
Species included. — Only the type species, Australocaris pinjarup Poore &
Collins, 2009, is known.
Australocaris pinjarup Poore & Collins, 2009
Australocaris pinjarup Poore & Collins, 2009: 238, figs. 11, 12, 39.
Diagnosis. — Rostrum narrow and 0.8 times length of front to cervical
groove, bearing 5-6 short, oblique lateral spines anterior to supraocular spine,
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
49
continuous with lateral gastric carinae. Supraocular spines prominent. Eyestalks 0.3 length of rostrum, and cornea unpigmented. A1 peduncle reaching
to midpoint of A2 segment 4; segment 1 swollen proximally, bearing small
lateral spinule. A2 segment 1 unarmed; segment 2 with slender distal spine, directed slightly upwards, reaching distally to middle of A2 segment 4; scaphocerite reaching distally beyond distal margin of segment 5, with three strong
spines on ventral margin, one spine on mesial margin; segment 3 with sharp
mesiodistal spine on ventral margin; segment 4 about as long as segment 2
(excluding distal spine), bearing mesiodistal spine (left side only); segment 5
about half length of segment 4. Carapace smooth except for slight rugosity at
base of rostrum, bearing few setae on gastric region and rostrum; lateral gastric carina unarmed; submedian gastric carina obsolete and curved mesially
between supraocular spines; median gastric carina sharp on rostrum, and unarmed. Mxp3 coxa and basis with distal spine on each ventral margin; ischium
unarmed; crista dentata with about 20 teeth; merus with two spines; carpus
with spine.
P1 symmetrical, flattened, and carinate; coxa with two spines on ventral
margin; basis with spine on ventral margin; ischium with two spines on ventral
margin; merus with three distal spines on barely convex dorsal margin, and
7 spines on ventral margin, lateral and mesial faces smooth; carpus with 6
spines on dorsal margin, and three spines laterally on ventral margin, mesial
face smooth; propodus with row of 4 spines on dorsal margin, and 9 spines
on convex ventral margin, lateral face tuberculate and mesial face smooth;
fixed finger 1.2 times as long as dorsal palm, and cutting edge convex, bearing
about 15 irregular, triangular teeth; dactylus with 4 spines on proximal half
of dorsal margin, lateral and mesial faces smooth; cutting edge denticulate;
both fingers bearing setae. P2 ischium and merus unarmed on ventral margin;
carpus 0.7 length of chela; propodus dorsal margin 0.5 length of dactylus. P3
or 4 merus with spine on ventral margin; propodus half as long as dactylus.
P5 missing. Abdominal somite 1 with pleuron truncate ventrally; pleuron
2 asymmetrical and rounded posteriorly; pleura 3-6 rounded. Plp1 minute
and cylindrical. Plp2 without appendix masculina; appendix interna one-third
length of endopod. Telson 1.2 times as long as wide, widest proximally,
then approximately parallel-sided, bearing no spines on lateral margin and at
rounded posterolateral angle, but median spine on truncate-convex posterior
margin; dorsal surface without oblique ridges or spines. Uropodal endopod
1.5 times as long as wide, bearing longitudinal median ridge with 4 spines
(none marginal) on dorsal surface, but no lateral spines. Uropodal exopod 1.6
50
CRM 013 – Katsushi Sakai
times as long as wide, bearing 6 obscure spines on distal half of lateral margin,
and one fixed spine and one robust seta at posterolateral angle; dorsal surface
with unarmed transverse suture, but no longitudinal ribs. [Adapted from Poore
& Collins, 2009.]
Type locality. — Off Bumbury, Western Australia; 423-397 m.
Distribution. — Only known from the type locality.
Genus Axiopsis Borradaile, 1903
Axiopsis Borradaile, 1903: 538; Nobili, 1906b: 91; Schmitt, 1921: 110; De Man, 1925d: 2 (key),
66; Vinogradov, 1950: 223; Balss, 1957: 1579; Poore & Griffin, 1979: 224; Sakai, 1986: 11;
Sakai, 1987a: 303 (list); Sakai & De Saint Laurent, 1989: 76; Sakai, 1992a: 173; Poore,
1994: 98 (key); Sakai, 1994: 198; Kensley, 1996e: 469; Poore & Collins, 2009: 241.
Axiopsis s. str., De Man, 1925d: 2 (key), 68; Balss, 1957: 1579.
Not: Axiopsis, Poore, 2004: 173 (= Michelaxiopsis Poore & Collins, 2009 and Leonardsaxius
gen. nov.).
Diagnosis. — Rostrum triangular and pointed at apex; lateral margins
armed with denticles, extending onto gastric region. Gastric region forming
horseshoe-shaped prominence with 5 longitudinal carinae or rows of denticles. Anterolateral margin of carapace unarmed. Cervical groove present at
full length. Eyestalks subglobose; cornea pigmented. A2 scaphocerite strongly
protruded forward. P1 unequal and vertically positioned; chelae unarmed
on dorsal margin. P2 chelate. No pleurobranchs. Abdominal pleura smooth
on surface, pleuron 2 broadly truncate ventrally, and pleuron 6 triangular.
Male Plp1 absent; male Plp2 biramous and narrow, endopod with appendices interna and masculina at proximal third, appendix masculina longer
than appendix interna; male Plps3-5 endopods with appendix interna. Telson subsquare, bearing median tooth on posterior margin. Uropodal exopod with transverse suture. [Adapted from Sakai & De Saint Laurent,
1989.]
Remarks. — Poore (2004: 173) included two Australian species, Axiopsis
australiensis De Man, 1925 and Axiopsis werribee (Poore & Griffin, 1979)
in Axiopsis. However, the former species is here classified as Michelaxiopsis
australiensis (De Man, 1925), and the latter as Leonardsaxius werribee (Poore
& Griffin, 1979). This is because Axiopsis differs from both Michelaxiopsis
and Leonardsaxius in major external morphological characteristics, as shown
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
51
in the scheme below:
Axiopsis
Michelaxiopsis
Leonardsaxius
gen. nov.
Gastric region
Gastric region forming horseshoeshaped
prominence with 5
longitudinal carinae
or rows of denticles
Gastric region forming
convex platform, bearing tuberculate lateral,
submedian, and median carinae, and additionally a short row
of tubercles between
median and submedian
carinae
Gastric region convex,
bearing median carina
with 1-3 teeth originating from base of
rostrum, and smooth
submedian and lateral
carinae
Male Plps1-2
Male Plp1 absent;
male Plp2 endopod
with appendices interna and masculina
at proximal third
Male Plp1 absent;
male Plp2 endopod
with appendix interna,
but without appendix
masculina
Male Plp1 uniramous
and bisegmented, distal segment triangular;
male Plp2, endopod
with appendices interna and masculina
Type species. — Axius affinis De Man, 1888b, by original designation.
[Synonym of Axia serratifrons A. Milne-Edwards, 1837]. The gender of the
current generic name, Axiopsis, is feminine.
Species included. — Axiopsis baronai Squires, 1977; A. consobrina (De
Man, 1905) [= A. irregularis Edmondson, 1930; A. tsushimaensis Sakai,
1992a]; A. serratifrons (A. Milne-Edwards, 1873); A. sp., Sakai, 1992a. [A.
eximia Kensley & Williams, 1990 is not considered here, as it is a fossil
species.]
K EY
TO THE
S PECIES
OF THE
G ENUS AXIOPSIS
1 – Eyestalks robust . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sp., Sakai, 1992
– Eyestalks subglobose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – In larger and smaller chelipeds carpi and chelae thickly setose and tuberculate on lateral
surface, and fingers longer than palm; P2 fingers longer than palm . . . . . . . . . A. baronai
– In larger and smaller chelipeds carpi and chelae not setose, but smooth, or roughly
scattered with flattened tubercles on lateral surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Merus of larger cheliped unarmed dorsally; palm of larger cheliped roughly scattered
with flattened tubercles on lateral surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. serratifrons
– Merus of larger cheliped armed with subdistal spine dorsally; palm of larger cheliped
smooth on lateral surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. consobrina
Axiopsis baronai Squires, 1977
Axiopsis (Axiopsis) baronai Squires, 1977: 1885, 4 text-figs., 1 table.
52
CRM 013 – Katsushi Sakai
Axiopsis baronai — Sakai & De Saint Laurent, 1989: 78; Lemaitre & León, 1992: 44;
Hendrickx, 1995: 390 (list), 393, 395, figs.; Kensley, 1996e: 469 (list); Hendrickx et al.,
2005: 171.
Material examined. — USNM 152653, holotype, male (TL/CL, 74.0/24.0 mm), Togoroma,
Colombia, Pacific Ocean, 04◦ 22′ N 77◦ 26.5′ W, 7 m, 8.vi.1971, leg. H.J. Squires; USNM
152655, paratype, female (TL/CL, 115.0/39.0 mm), Pizarro, Colombia, Pacific Ocean, 05◦ 11′ N
77◦ 26′ W, 9 m, 11.vi.1971, leg. H.J. Squires.
Remarks. — Squires (1977: 1888) described the male Plp1 as missing.
However, considering that the males of all species included in Axiopsis bear
no Plp1, the absence of the male Plp1 may safely be considered as one of the
characteristics of Axiopsis.
Type locality. — Togorama, Colombia.
Distribution. — Colombia — Togorama (Squires, 1977).
Axiopsis consobrina De Man,1905 (fig. 7)
Axiopsis consobrina De Man, 1905: 595 (partim, not specimen from Sulu Archipelago); Balss,
1925: 209; Sakai & De Saint Laurent, 1989: 77; Sakai, 1994: 198, 201 (key), fig. 14; Kensley,
1996e: 469 (list); Komai et al., 2002: 29; Poore & Collins, 2009: 241.
Axiopsis (Axiopsis) consobrina — De Man, 1925d: 5 (list), 69 (key), 80, pl. 6 fig. 13-13c;
Estampador, 1937: 500; Ngoc-Ho, 2005: 55, fig. 4.
Axiopsis (Axiopsis) irregularis Edmondson, 1930: 10, fig. 4, pl. 1A. [Type locality. — Pearl and
Hermes Reef, Hawaiian Islands, shallow waters.]
Axiopsis (Axiopsis) aff. serratifrons — Sakai, 1970a: 37, fig. 1a-b; Sakai, 1987a: 303 (list);
Poore & Collins, 2009: 242.
Axiopsis irregularis — Sakai & De Saint Laurent, 1989: 78.
Axiopsis tsushimaensis Sakai, 1992a: 173, figs. 14, 15; Kensley, 1996e: 469 (list); Komai et al.,
2002: 21, figs. 1-3, tables 1, 3. [Type locality. — Fukuoka, Korea Strait, 34◦ 11′ N 130◦ 2′ E,
102 m.]
Not: Axiopsis (Axiopsis) consobrina — Poore & Griffin, 1979: 230, fig. 4 (= Axiopsis serratifrons A. Milne-Edwards, 1873 and Pseudoaxiopsis carpentariaensis sp. nov.).
Not: Axiopsis (Axiopsis) consobrina — Tirmizi, 1983: 91, fig. 4 (= Axiopsis serratifrons
A. Milne-Edwards, 1873).
Material examined. — ZMA Crust. De. 102672, paralectotype female (TL/CL, 31.0/
10.2 mm), Indonesia, 4◦ 20′ S 122◦ 58′ E, between islands of Wowoni and Buton, northern
entrance of Button Strait, 75-94 m, sand with dead shells [“Buton” or “Button” = Butung];
ZMB 13853, 1 juv. (TL/CL, 12.0/4.1 mm), Banda Sea, R/V “Planet”, 27.viii.1909, leg. Graef;
SMF. 32778, 1 ovig. female (TL/CL, 64.0/22.2 mm), Suruga-Bay, fisher trawling boat “Seishinmaru”, 27.ix.1977, leg. S. Ohta.
Diagnosis. — Rostrum (fig. 7A, B) triangular and acuminate, projecting
downward distally, reaching anteriorly middle of A1 penultimate segment, lateral margins with 3-5 teeth, extending posteriorly onto gastric region as lateral
carinae with 8-9 denticles; submedian rows of 8-9 denticles; median carina
with 15 denticles including hepatic tubercle; no tubercles between submedian
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
53
Fig. 7. Axiopsis consobrina De Man, 1905. A, whole body, lateral view; B, carapace, dorsal
view. A, B, Axiopsis consobrina, ZMA Crust. De. 102672, paralectotype female (TL/CL,
31.0/10.2 mm), Indonesia, 4◦ 20′ S 122◦ 58′ E, between islands of Wowoni and Buton, northern
entrance of Button Strait, 75-94 m [“Buton” or “Button” = Butung]. Scales 1 mm.
rows and median carina. Cervical groove strong and clearly defined. Postcervical carina absent. Male Plp1 absent; male Plp2 endopod bearing appendices
interna and masculina with distal setae; male Plps3-5 endopods with appendix
interna.
Remarks. — The present species, Axiopsis consobrina is similar to Axiopsis
serratifrons A. Milne-Edwards, 1873, but they differ from each other as
54
CRM 013 – Katsushi Sakai
follows. In A. consobrina the merus of the larger cheliped is armed with a
subdistal spine on the dorsal margin, the gastric region bears lateral carinae
with 8-9 denticles, submedian rows of 8-9 denticles, and a median row of 15
denticles including a hepatic tubercle, but no tubercles between the submedian
and median rows, and the palm of the male larger cheliped is smooth on the
lateral surface. In A. serratifrons, the merus of the larger cheliped is unarmed
on the dorsal margin, the gastric region bears lateral carinae with 16 tubercles,
a median row of tubercles including a hepatic one, submedian rows of 12-13
tubercles, and numerous tubercles between the median and submedian rows of
tubercles, and the palm of the male larger cheliped is roughly scattered with
flattened tubercles on the lateral surface.
The juvenile specimen from the Banda Sea (ZMB 13853) is identified as
A. consobrina, because no denticles are observed between the median and
submedian rows of denticles in the gastric region.
Axiopsis (Axiopsis) irregularis Edmondson, 1930 from Hawaii, is synonymous with the present species, Axiopsis consobrina, because the gastric region
bears no denticles between the median and submedian rows of spines (Edmondson, 1930, fig. 4a, b), and the merus of the larger cheliped is armed with
a subdistal spine on the dorsal margin, as in A. consobrina (cf. Sakai, 1994,
fig. 14).
Poore & Griffin (1979: 230) described A. consobrina based on a specimen
from the Gulf of Carpentaria, Queensland, Australia, however, it is found that
the female specimen (Poore & Griffin, 1979, fig. 4e) is not A. consobrina but
A. serratifrons, because the merus of the larger cheliped bears no subdistal
spine on the dorsal margin, though this female specimen is now considered
lost (G.C.B. Poore, in litt.). The same was found when examining the male
specimen (AM P16308), which is not to be assigned to A. consobrina but to
Pseudoaxiopsis carpentariaensis sp. nov. (see p. 178), because in the former
the gastric region bears a simple median row of spines, and the dorsal surface
of the abdomen is smooth, whereas in the latter the gastric region bears
a tuberculate median carina with short double carina with tubercles in the
middle, running from the middle of the rostrum nearly to the cervical groove
(fig. 7A, B; Poore & Griffin, 1979, fig. 4a), and the abdominal somites 1-6 are
provided with a characteristic dorsal pattern of carinae arranged as longitudinal
stripes.
Tirmizi (1983) assigned a female from Bali, Indonesia to A. consobrina.
However, Sakai (1994: 199) rejected the identification, because the gastric
region is scattered with denticles between the median and submedian carinae,
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
55
unlike that of the type specimen of A. consobrina. In the present revision, it
is found by examination of the lectotype of A. serratifrons that her specimen
may safely be assigned to A. serratifrons.
Axiopsis tsushimaensis Sakai, 1992a was established from a young female
specimen from the Tsushima Islands, Fukuoka, Korean Strait, and distinguished from Axiopsis consobrina, because A. tsushimaensis is characterized
by its simple A2 scaphocerite, versus A. consobrina by its proximally bifurcate A2 scaphocerite. However, it would be better to regard A. tsushimaensis as
synonymous with A. consobrina, taking into consideration that the two species
are almost the same in all other characteristics, and A. tsushimaensis is young
and small (TL/CL, 18.0/5.8 mm) and its A2 scaphocerite is safely considered
to be undeveloped; even in the larger specimen of A. consobrina from Suruga
Bay, the scaphocerite is poorly developed and not as distinctly bifurcate as in
the type specimen of A. consobrina.
It is also noted that in A. consobrina another morphological transformation
is observed with growth. In the smaller specimens, pleuron 1 is armed with a
spinule ventrally, pleura 3-5 with an anteroventral spinule, and pleuron 6 with a
spinule ventrally, whereas in the larger specimen from Suruga Bay, pleura 1-3
are ventrally unarmed, pleura 4-5 bear no spinule but an anteroventral tubercle,
and pleuron 6 is unarmed.
Komai et al. (2002) made reference to the podobranchs on the thoracic
somites of A. consobrina under the name of A. tsushimaensis, and described
that they are present as rudimentary structures on thoracic somites 2-6.
However, in A. consobrina they are not present as rudimentary ones but as
small structures on thoracic somites 3-6 (table II).
Type locality. — Solor Strait, Indonesia, “Siboga” Sta. 305, 113 m.
Distribution. — Hawaiian Islands — Pearl and Hermes Reef (Edmondson,
1930); Indonesia — Buton [= Butung] Strait and Solor Strait (De Man, 1905,
1925); Australia — North West Shelf 75-275 m (Sakai, 1994; Poore & Collins,
TABLE II
Axiopsis consobrina (De Man, 1905), gill formula (SMF 32778, from Suruga Bay, Japan)
Maxillipeds
Epipods
Podobranchs
Arthrobranchs
Pleurobranchs
Pereiopods
1
2
3
1
2
3
4
5
1
–
–
–
1
–
1
–
1
1
2
–
1
1
2
–
1
1
2
–
1
1
2
–
1
–
2
–
–
–
–
–
56
CRM 013 – Katsushi Sakai
2009); Tahiti, Society Is., French Polynesia; Japan — Tsushima (Sakai,
1970a), Fukuoka, Korean Strait (Sakai, 1992a); N.E. of Iki Island, 33◦ 57.23′ N
129◦ 54.24′ E, 86 m, Yamaguchi Pref. (Komai et al., 2002); Suruga Bay,
Shizuoka Pref.
Axiopsis serratifrons A. Milne-Edwards, 1873
(figs. 8-9)
Axia serratifrons A. Milne-Edwards, 1873: 263, pl. 13 (not 2), figs. 6, 6a.
Axius serratifrons — Rathbun, 1906: 895; Edmondson, 1923: 27.
Axiopsis serratifrons — Sendler, 1923: 44, pl. 21 fig. 10 (not 9); Sakai & De Saint Laurent,
1989: 76; Werding & Müller, 1989: 251; Manning & Chace, 1990: 31, figs. 16, 17;
Dworschak, 1992: 215; Lemaitre & León, 1992: 44; Lemaitre & Ramos, 1992: 346, figs.
2, 3; Sakai, 1992a: 173; Dworschak & Ott, 1993: 280; Hendrickx, 1995: 390 (list); Kensley,
1996e: 469 (list); Rodrigues et al., 1998: 379; Debelius, 1999: 226, fig.; Melo, 1999: 312,
figs. 209-210; Vargas & Cortés, 1999b: 905; Abed-Navandi, 2000: 295; Komai, 2000b: 344
(list); Ngoc-Ho, 2002b: 224, fig.; Kensley, 2003: 366, pls. 2, 8, table 1; Ngoc-Ho, 2005: 53,
fig. 3 (not holotype); Dworschak, 2004: 15, fig. 5; Komai & Tachikawa, 2008: 20-22, fig. 1;
Poore & Collins, 2009: 241; Robles et al., 2009: 316.
?Axiopsis serratifrons — Borradaile, 1903: 538.
Fig. 8. Axiopsis serratifrons (A. Milne-Edwards, 1873). A, whole body, lateral view; B, C,
gastric region, dorsal view. A, B, ZMH K 8405 (Mus. Godeffroy 1723), lectotype, female
(TL/CL, 31.0/11.2 mm), Upolu, Samoa; C, MNHN Th 147, paralectotype, male (dried) (CL,
11.5; abd + telson 19.5 mm). Scales 1 mm.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
57
Fig. 9. Axiopsis serratifrons (A. Milne-Edwards, 1873). A, larger cheliped, lateral view;
B, smaller cheliped, lateral view; C, P2, lateral view; D, P3, lateral view; E, P4, lateral view.
A-E, ZMH K 8405 (Mus. Godeffroy 1723), lectotype, female, Upolu, Samoa. Scales 1 mm.
58
CRM 013 – Katsushi Sakai
Axiopsis (Axiopsis) serratifrons — De Man, 1925d: 6 (list), 68 (key), 72, pl. 6 fig. 12-12i;
Holthuis, 1953b: 51; Kensley, 1981a (not 1980): 1253, figs. 1-5; Rodrigues, 1983: 18, figs.
1-22.
Axius spinipes De Man, 1888b: 464, pl. 19 fig. 6; Zehntner, 1894: 195. [Type locality.
— Noordwachter (North Light House), nearest town or city, Tanjungpandan, Indonesia,
5◦ 12.24′ S 106◦ 27.36′ E.]
Axius affinis De Man, 1888b: 469, pl. 20 fig. 1 (fig. P1, cheliped). [Type locality. — Amboina,
Indonesia.]
Axiopsis spinipes — Borradaile, 1903: 538; Nobili, 1906b: 91; Borradaile, 1910: 262.
Axiopsis affinis — Borradaile, 1903: 538; Borradaile, 1904: 752; Nobili, 1906b: 92; Makarov,
1938: 47.
Axiopsis sculptimana Ward, 1942: 62. [Type locality. — Diego Garcia, Chagos Archipelago.]
Axiopsis sp. D, Coelho, 1971: 231.
Axiopsis (Axiopsis) sp. D, Coelho & Ramos, 1972: 161.
Allaxius sculptimanus — Sakai & De Saint Laurent, 1989: 75.
Axiopsis brasiliensis — Coelho & Ramos-Porto, 1985: 73, fig. 3; Melo, 1999: 310, figs. 207208; Coelho et al., 2007: 6, tab. 2.
Axiopsis pica Kensley, 2003: 361, figs. 1, 2, pl. 1, table 1; Ngoc Ho, 2005: 51, fig. 2. [Type
locality. — Anae Island, Guam.]
Axiopsis consobrina — Tirmizi, 1983: 91, fig. 4; Ngoc-Ho, 1995: 55, fig. 4.
Material examined. — Indo-West Pacific region. — ZMH K 8405 (Mus. Godeffroy 1723),
lectotype, female (TL/CL, 31.0/11.2 mm), Upolu, Samoa; ZMH K 8385 (Museum Godeffroy
16193), 1 male (TL/CL, 39.0/14.3 mm), 2 ovig. females (TL/CL, 49.0/17.1; 76.0/23.9 mm),
Tahiti; ZMH K 8389 (Museum Godeffroy 16367), 1 male (TL/CL, 65.0/19.8 mm), Caroline Is.;
ZMH K 24556, 1 male (TL/CL, 49.0/16.2 mm), Bismarck Archipelago, Südsee-Expedition, no.
90, 10.viii-25.ix.1908, leg. Dr. Dunker; ZMB 8904, 1 male (TL/CL, 53.0/16.0 mm), 1 ovig.
female (TL/CL, 65.0/22.0 mm), 1 female (TL/CL, 62.0/23.0 mm), Jaluit Island, Marshall
Islands; MNHN Th 147, paralectotype, 1 male, (TL/CL, 23.0/11.5 mm), Hawaii, Sandwich
Islands, det. Nobili, 1905; MNHN Th 691, male (TL/CL, 67.0/22.5 mm), Île des Pins, New
Caledonia, 19.xii.1961; MNHN Th 864, 1 male (TL/CL, 39.0/12.2 mm), Japan, leg. K. Sakai;
MNHN Th 690, 1 male (TL/CL, 26.0/8.5 mm), Calypso “Aldara, 1954”, det. De Saint. Laurent;
MNHN Th 4179 (= UF 2782), 1 ovig. female (TL/CL, 48.0/16.0 mm), Mariana Islands, Guam
Is., Apra Harbour; MNHN Th 1426, 1 male (TL/CL, 17.0/6.0 mm), Tahiti, Society Islands,
French Polynesia, 2.5 m, 17.i.1995 (Ngoc-Ho, 2005: 55, fig. 4, identified as A. consobrina);
NHM 1973: 629: 2, 1 male (TL/CL, 49.0/15.1 mm), 1 ovig. female (TL/CL, 52.0/17.4 mm),
Dar es Salaam, Tanzania, leg. R.G. Hartnoll; RMNH D 1168, 1 male (TL/CL, 21.0/6.5 mm),
“Siboga” Exp. vii.1929; RMNH D 50168, 1 male (TL/CL, 46.0/14.7 mm), Indonesia, S.W.
Sulawesi, Spermonde Archipelago, west side of Badi Island, 22 m, 26.v.2001, leg. et don. A.
Gittenberger.
Atlantic Ocean. — ZMB 26887, 1 male (TL/CL, 56.0/18.0 mm), Karibik coast, Punte de
Betin, Santa Marta, Colombia, 1-2 m, 26.iii.1977, leg. L. Werding, det. Müller; DOUFPe 314,
holotype, male, 5◦ 15′ S 34◦ 59′ W, 69 m, det. as Axiopsis braziliensis; DOUFPe 311, paratype,
female (broken), Recife, PE, Brazil, det. as Axiopsis braziliensis; DOUFPe 8584, 7◦ 52′ S
34◦ 32′ W, 45.5 m, det. as Axiopsis braziliensis; DOUFPe 1873, 1 male (TL/CL, 64.0/22.6 mm),
1 ovig. female (TL/CL, 54.0/19.6 mm), 1 female (TL/CL, 53.0/18.7 mm), Atol das Rocas, RN,
Brazil, 19.vii.1992, det. as Axiopsis braziliensis; DOUFPe 8583, Atol das Rocas, RN, Brazil,
20.viii.1992, det. as Axiopsis braziliensis.
Description of lectotype. — Rostrum (fig. 8B, C) stoutly triangular, with
5-6 teeth on lateral margins extending posteriorly onto gastric region as lateral
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59
carinae; gastric region bearing lateral carinae with 16 tubercles, median row of
tubercles including hepatic one, submedian rows of 12-13 tubercles, and numerous tubercles between median and submedian rows. Cervical groove strong
and distinct. Surface of carapace posterodorsally pitted, lacking postcervical
carina. P1 unequal; in left larger cheliped (fig. 9A), ischium with five teeth on
ventral margin; merus with four distinct teeth on mesioventral margin, series
of small teeth on ventrolateral margin, and unarmed on dorsal margin; carpus
unarmed; palm 1.7 times as long as wide, carinate and unarmed dorsolaterally, and flattened tubercles on lateral surface and along ventral margin; cutting
edge of fixed finger with proximal fine denticulation followed by median tooth,
and smooth distally; dactylus largely concave on cutting edge. In right smaller
cheliped (fig. 9B), ischium obtusely denticulate on dorsal margin, and armed
with 5 teeth on ventral margin; merus with 4 sharp teeth on ventral margin and
two subdistal teeth on dorsal margin; carpus unarmed; palm twice as long as
wide, and distinctly carinate on ventral and dorsal margins; fixed finger denticulate on cutting edge, and dactylus unarmed on cutting edge. P2 (fig. 9C)
merus with 4 teeth on ventral margin; fingers slightly shorter than palm. Ps3-4
(fig. 9D, E) propodi elongate, bearing short vertical rows of 1-3 spinules ventrolaterally. Abdominal pleuron 1 (fig. 8A) narrow ventrally; pleuron 2 broad,
and rounded at antero- and posteroventral angles; pleura 3-6 truncate ventrally,
bearing small tooth on anteroventral angle; pleuron 6 narrow and rounded, with
small tooth ventrally. Lateral surfaces of pleura 2-4 pitted dorsolaterally. Male
Plp1 absent; male Plp2 endopod with appendices interna and masculina; male
Plps3-5 endopods with appendix interna. Female Plp1 consisting of proximal
segment and multiarticulate flagellum. Plps2-5 leaf-like, endopods with appendix interna. Telson longer than wide, bearing proximal lobe with spine,
followed posteriorly by tooth on lateral margin, three spinules at posterolateral
angle, median tooth on posterior margin, and two pairs of spines medially on
dorsal surface. Uropodal endopod oval, bearing three spines including distolateral one, and longitudinal row of 5 spines medially on dorsal surface. Uropodal
exopod broadened, bearing 4-5 spines on distolateral margin, a submarginal
row of three spines distolaterally, and a transverse suture with spinules.
Remarks. — A. Milne-Edwards (1873a) published two type localities for
the two syntypes of the present species, Axiopsis serratifrons A. MilneEdwards, 1873, i.e., Upolu, Samoa, and Sandwich Islands, Hawaii, in the
Journal of the Museum Godeffroy. One of the syntypes from Upolu, Samoa
is preserved in good condition in the Zoological Museum of Hamburg, under
catalogue number ZMH K 8405 (Mus Godeffroy 1723) (fig. 8A), and the
60
CRM 013 – Katsushi Sakai
other from the Sandwich Islands, Hawaii is preserved as the “Holotype”,
designated without righteous reasons (Ngoc-Ho, 2005: 53), in the Muséum
national d’Histoire naturelle, Paris, under catalogue number MNHN Th 147.
According to the ICZN, 2000, calling a syntype “Holotype” after 1999 does
not imply its automatic designation as Lectotype (Segment 74.7) as in the cases
before 2000 (Art. 74.6). Therefore, the two specimens have still to be regarded
as syntypes, and the present author herewith selects the female specimen from
Upolu, Samoa in the Hamburg collection (ZMH K 8405) as the lectotype of the
species. This brings the name-bearing type back to the institution to which the
Godeffroy collection has always belonged. The remaining male syntype from
the Sandwich Islands, Hawaii, preserved in the Muséum national d’Histoire
naturelle, Paris, under catalogue no. MNHN Th 147, is, therefore, herewith
defined as the paralectotype.
The reasons why the female specimen is selected as the lectopype are as
follows. The figures of the type specimen published by A. Milne-Edwards
(1873a, pl. 13 fig. 6 and 6a) show that the type specimen bears a larger cheliped
on the left side, which suggests that the type specimen he figured is the female
preserved in the collection of the Zoological Museum in Hamburg (Panning,
1956), but not the male that was deposited by A. Milne-Edwards, and has been
preserved, in the Muséum national d’Histoire naturelle in Paris, and which was
misdescribed by Ngoc-Ho (2005) as the “holotype” of Axiopsis serratifrons
(MNHN Th 147) (fig. 8C). So, the female type specimen can safely be defined
as the lectotype of Axiopsis serratifrons, and the male type specimen as the
paralectotype.
The female lectotype from Upolu, Samoa is characterized, through careful
examination, as follows. The gastric region bears lateral carinae with numerous
tubercles, median and submedian rows of tubercles, and numerous tubercles
between these; the palm of the left, larger cheliped is scattered with flattened
tubercles on the lateral surface except for the median part, the dorsal margin is
carinate and unarmed (fig. 9A); the fixed finger bears one tooth medially on the
cutting edge, the palm of the smaller cheliped is not scattered with flattened
tubercles but smooth.
Kensley (2003: 361) described a new species, Axiopsis pica. However, the
male holotype of Axiopsis pica Kensley, 2003 from Guam, is almost the same
in characteristics as Axiopsis serratifrons, except that it has a different colour
pattern. This difference, however, is not sufficiently decisive to distinguish
the two species, and is to be regarded as a variation at species level, because
there are also other differences regarded as mere variations between the two
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61
species, such as the number of the median teeth on the cutting edge of the
larger cheliped, and the presence or absence of flattened tubercles on the
lateral surface of the larger and smaller chelipeds. Even though he (Kensley,
2003) described in his remarks, citing Knowlton’s notes, that “in several
groups, while shade and intensity of color may vary, the actual pattern of
pigmentation is consistent and can be used to separate morphologically very
similar species”. Therefore, A. pica is considered to be symonymous with A.
serratifrons.
Ngoc-Ho described Axiopsis pica Kensley, 2003 based on specimens from
the Tuamotu Archipelago, French Polynesia, and Mariana Island (Ngoc-Ho,
2005, fig. 2C), showing that the larger cheliped is scattered with flattened
tubercles on the mesial and lateral surfraces of the P1 palm. However, the
feature agrees, though the colour is different, with that of the female lectotype
of Axiopsis serratifrons (fig. 9A), and in the male and female specimens of
A. serratifrons (ZMH K 8385 (Museum Godeffroy 16193), 1 male (TL/CL,
39.0/14.3 mm), 2 ovig. females (TL/CL, 49.0/17.1; 76.0/23.9 mm), Tahiti;
NHM 1973: 629: 1 ovig. female (TL/CL, 52.0/17.4 mm), Dar es Salaam,
Tanzania), it is observed that the P1 palms of the larger and smaller chelipeds
are scattered with flattened tubercles on both lateral and mesial surfaces.
Lemaitre & Ramos (1992, fig. 2) described A. serratifrons from a juvenile
female specimen from Gorgona Island, Pacific coast of Colombia, in which
the palms of both larger and smaller chelipeds are scattered with flattened
tubercles on the lateral surface as in the specimen of A. serratifrons from
Kur Island, Indonesia (De Man, 1925d, pl. 6 fig. 12h, i) and the holotype
of Axiopsis pica from Guam (Kensley, 2003). The number of the tubercles
on the gastric region also varies conspicuously among the specimens of A.
serratifrons, as observed in the female lectotype (ZMH K 8405) (fig. 8B) and
the male paralectotype (MNHN Th 147) (Ngoc-Ho, 2005, fig. 3B), which
suggests that A. pica described by Ngoc Ho (2003) should be synonymous
with A. serratifrons.
The type specimen of Axiopsis sculptimana Ward, 1942 is beyond my
access, which makes it difficult to review Ward’s species. Ward (1942)
described that “this species is allied to Axiopsis serratifrons (A. M.-Edwards,
1873) but they differ; (1) The distal surfaces of both chelae are strongly
sculptured, having coarsely flattened tubercles appearance in A. sculptimana,
(2) The whole surface of A. serratifrons is covered with short stiff setae,
(3) The sculpture on the gastric region is different. In sculpture there are
two outer lines of spines, but the median portions are covered with spines
62
CRM 013 – Katsushi Sakai
irregularly arranged in A. sculptimana, whereas they are laid out in rows in
A. serratifrons”. However, he made errors and confused descriptions of the
differences between the two species, because in A. serratifrons the whole
surface of the body is not covered with short stiff setae, and tubercles (not
spines) are not always laid out in rows except for lateral rows of tubercles
on the gastric region, because the gastric region is usually covered with
tubercles irregularly arranged between the lateral rows (fig. 8A). This shows
that he failed to perform a careful examination of A. serratifrons, and gave
misdescriptions of its characteristics. Judging from the characteristics of A.
sculptimana described by Ward (1942), his species is sufficiently similar to be
synonymous with A. serratifrons.
It is here confirmed that Axiopsis brasiliensis Coelho & Ramos-Porto, 1985
is a junior synonym of the present species, Axiopsis serratifrons, as suggested
by Coelho (1997). In the female specimen (DOUFPe 1873) the palm of the
smaller cheliped is tuberculate on the lateral and mesial surfaces, whereas in
the male (DOUFPe 1873) it is not tuberculate but smooth on both surfaces.
Taking all this into consideration, the presence or absence of the tubercles on
the palm of the smaller cheliped depends on the individual, and hence cannot
be regarded as a species-specific character.
Type locality. — Upolu, Samoa.
Distribution. — Bermuda — North Rocks (Kensley, 1981a); Isla San José,
Islas Murciélago, Guanacaste; Punta Esmeralda, Bahia Culebra, Costa Rica
(Vargas & Cortés, 1999b: 905); Florida — Pigeon Key, Monroe County
(Kensley, 1981a); Belize — Carrie Bow Cay (Kensley, 1981a; Dworschak,
1992); Colombia — Gorgonia Island (Lemaitre & Ramos, 1992); Punta de
Betin, Santa Marta, (present data); Brazil — Rio Grande do Norte; Camocim;
Fortaleza; Ceará, Recife; Pernambuco; Porto de Pedras, Alagoas; Cape Verde
Islands (Abed-Navandi, 2000); Ascension Island (Manning & Chace, 1990).
South Africa — Sodwana Bay, Zululand (Kensley, 1981); Madagascar —
Tuléar (Sakai & De Saint Laurent, 1989); Kenya — Mombasa (Sakai &
De Saint Laurent, 1989); Aldabra Island (Sakai & De Saint Laurent, 1989);
Seychelles (Sakai & De Saint Laurent, 1989); Red Sea (Nobili, 1906b);
Gulf of Aden — Obock (Nobili, 1906b; Sakai & De Saint Laurent, 1989);
Maldives — Male Atoll (Borradiale, 1904); Chagos Archipelago — Salomon
Atoll (Borradaile, 1910); Sri Lanka — Colombo (Manning & Chace, 1990);
Indonesia — Noordwachter Island (De Man, 1888b), Ambon (De Man,
1888b, 1925; Zehntner, 1894), Obi Island, Damar Island, Salayar Island,
Kepulauan Lucipara, Kur Island, and Roti Island (De Man, 1925d), Kei Islands
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
63
(Sakai, 1992a); Palau Is. (Sendler, 1923); Australia — Gulf of Carpentaria,
Queensland (Poore & Collins, 2009); Marshall Islands — Narnu Island,
Bikini Atoll (Kensley, 1981a), Jaluit; Gilbert Islands — Onotoa (Kensley,
1981a); New Caledonia (Sakai & De Saint Laurent, 1989); Upolu, Samoa
(A. Milne-Edwards, 1873); north of South China (Liu & Zhong, 1994);
Hawaiian Islands (A. Milne-Edwards, 1873; Sakai & De Saint Laurent, 1989);
Tabuaeran (Edmondson, 1923); Bonin Islands — Chichi-jima, Miyano-hama;
Washington Beach (Komai & Tachikawa, 2008).
Axiopsis sp., Sakai, 1992
Axiopsis sp., Sakai, 1992a: 174, fig. 16.
Distribution. — Kenya — off Mombasa (Sakai, 1992a); 40 m.
Genus Axiorygma Kensley & Simmons, 1988
Axiorygma Kensley & Simmons, 1988: 657; Sakai & De Saint Laurent, 1989: 92; Poore, 1994:
98 (key).
Diagnosis. — Rostrum narrowly triangular and pointed at apex; lateral margins each armed with 5 teeth, extending onto gastric region as unarmed lateral carinae. Gastric region convex, bearing unarmed median carina originating from midlength of rostrum and reaching midway to cervical groove, and
unarmed submedian carinae starting posterior to rostrum and reaching midway to cervical groove. Cervical groove present at whole length; postcervical
carina absent. Anterolateral margins of carapace unarmed. Eyestalks elongate.
A2 scaphocerite elongate. P1 unequal in male, merus inflated dorsally, bearing
4 sharp spines ventrally; palms with cluster of setae at dorsoproximal angle,
in larger cheliped palm robust, and in smaller cheliped palm slender; female
having slender chelipeds subequal in length and shape. Ps2-5 slender; Ps34 propodi elongate, without transverse rows of spines on ventrolateral margin. No pleurobranchs. Abdominal pleura 3-5 with small bundle of setae, and
rounded on ventral margin. Male Plp1 absent; male Plp2 biramous, endopod
and exopod slender and leaf-like, endopod with appendix interna, but without
appendix masculina, male Plps3-5 endopods with appendix interna. Female
Plp1 uniramous and slender, female Plps2-5 endopods with appendix interna.
Telson subsquare with median tooth on posterior margin. Uropodal exopod
with transverse suture.
Type species. — Axiorygma nethertoni Kensley & Simmons, 1988, by
original designation and monotypy. Gender of the generic name, Axiorygma,
feminine.
Species included. — Axiorygma nethertoni Kensley & Simmons, 1988.
64
CRM 013 – Katsushi Sakai
Axiorygma nethertoni Kensley & Simmons, 1988
Axiorygma nethertoni Kensley & Simmons, 1988: 658, figs. 4-7; Sakai & De Saint Laurent,
1989: 92; Melo, 1999: 314, figs. 211-212.
Material examined. — USNM 211440, holotype, male (TL/CL, 21.0/7.0 mm); allotype,
ovig. female (TL/CL, 24.0/7.5 mm), Key Largo Marine Sanctuary, Florida, leg. S. Simmons,
det. B. Kensley.
Diagnosis of male holotype. — Rostrum narrowly triangular and pointed
at apex; lateral margins denticulate, extending onto gastric region. Anterolateral margins of carapace with two triangular protrusions. Gastric region convex; postrostral carina continued to gastric median carina, submedian and lateral carinae not serrated but smooth. Cervical groove present at full length;
postcervical carina absent. Eyestalks elongate, reaching level of rostral apex.
A2 scaphocerite elongate. P1 unequal, in both chelipeds ischium with slender
sharp spine on ventral margin; merus inflated, rounded on dorsal margin and
straight on ventral margin, bearing three slender sharp spines proximally on
ventral margin; in left larger cheliped palm thick, slightly less than twice as
long as wide, bearing two clusters of branched setae at dorsoproximal corner
(missing at present examination), fixed finger bearing strong triangular spine
midway on cutting edge, and dactylus largely gaped in proximal half of cutting
edge, while in right smaller cheliped palm slender, twice as long as wide, bearing two clusters of branched setae at dorsoproximal corner, fixed finger bearing tooth at proximal third, and dactylus regularly denticulate on each cutting
edge. P2 slender, fingers shorter than palm; Ps3-4 propodi elongated, bearing
no transverse rows of spines on ventrolateral margins. No pleurobranchs. Abdominal pleura 3-5 rounded on ventral margin, bearing small bundle of setae.
Male Plp1 absent. Male Plp2 biramous, endopod and exopod slender and leaflike, endopod with appendix interna, but without appendix masculina. Plps3-5
endopods with appendix interna. Telson subsquare, bearing proximal lobe followed by four spines on lateral margin, and median tooth on posterior margin.
Uropodal exopod with transverse suture.
Diagnosis of female. — Female Plp1 uniramous and slender. Plps2-5
endopods with appendix interna.
Type locality. — Key Largo National Marine Sanctuary, Florida, 30 m.
Distribution. — West coast of Florida (Kensley & Simmons, 1988; Sakai &
De Saint Laurent, 1989), Gulf of Mexico; 30-58.6 m.
Genus Axius Leach, 1815
Axius Leach, 1815: 343; Leach, 1816 (no number of page); Latreille, 1817: 35; Bell, 1846: 227;
Dana, 1852a: 13; Dana, 1852b: 509; Norman, 1868: 177; Meinert, 1877: 212; Boas, 1880:
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
65
98; Stebbing, 1893: 187; Borradaile, 1903: 536; Balss, 1914: 85; Selbie, 1914: 88 (key),
89; De Man, 1925d: 1 (key), 8; Rathbun, 1929: 25; Bouvier, 1940: 93; Zariquiey Alvarez,
1946: 95 (key), 103; Zariquiey Alvarez, 1968: 223; Poore & Griffin, 1979: 235; Sakai, 1987a:
303 (list); Sakai & De Saint Laurent, 1989: 26; Squires, 1990: 333; Sakai, 1992a: 165; Poore,
1994: 97 (key); Ngoc-Ho, 2003: 447; Ingle & Christiansen, 2004: 77; Poore & Collins, 2009:
242.
Axia H. Milne Edwards, 1837a: 310; H. Milne Edwards, 1837b: 150.
Axius s. str., Borradaile, 1903: 537; De Man, 1925d: 1 (key), 11 (key).
Diagnosis. — Rostrum triangular, bearing median carina, lateral margins
armed with denticles, extending onto gastric region. Gastric region convex.
Anterolateral margin of carapace unarmed. Cervical groove present at full
length. Eyestalks subglobose; cornea pigmented. A2 scaphocerite prominent.
P1 subequal in length and dissimilar in shape, and vertically positioned.
P3 propodus broadened, lacking transverse rows of spines ventrolaterally, but
setose on ventral margin; P4 propodus elongate, bearing median and ventral
rows of setae on lateral surface; dactylus bearing median row of spines on
lateral surface. P5 subchelate. Pleurobranchs on Ps2-4. Abdominal pleura 3-5
with tuft of soft setae laterally, and truncate or rounded on ventral margin. Male
Plp1 uniramous and bisegmented; distal segment narrower than proximal one,
tapering distally; male Plp2 biramous, endopod with appendices interna and
masculina; and male Plps3-5 biramous, broad and lamellar with marginal tufts
of soft setae, and endopods with appendix interna. Female Plp1 uniramous and
bisegmented, consisting of proximal segment and multiarticulate flagellum;
Plps2-5 biramous, broad and lamellar with marginal tufts of soft setae,
endopods with appendix interna. Telson longer than wide, bearing pair of
spinules on dorsal surface and median tooth on posterior margin. Uropodal
exopod with transverse suture.
Remarks. — In the type species of the genus Axius, A. stirhynchus,
the uropodal exopod bears a transverse suture near the posterior margin,
but sometimes that is inconspicuous, as mentioned by Sakai & De Saint
Laurent (1989). It was found through present examination of the specimens
of A. serratus that the transverse suture is distinct in the smaller specimens
from Georges Bank, North America (NHM 1898.5.7.833-4, 2 ovig. females,
TL 80.0; 87.0 mm), but is inconspicuous in the larger specimen (NHM
1898.5.7.832, 1 female, TL 92.0 mm) from the same locality, although Kensley
(2001b: 961, table I) described for A. serratus based on his specimens that the
transverse suture is absent.
Type species. — Axius stirhynchus Leach, 1815, by original designation and
monotypy. The gender of the generic name, Axius, is masculine.
66
CRM 013 – Katsushi Sakai
Species included. — Axius armatus Smith, 1881; A. mediterraneus (Caroli,
1921); A. serratus Stimpson, 1852; A. stirhynchus Leach, 1815.
K EY
TO THE
S PECIES
OF THE
G ENUS AXIUS
1 – Rostrum serrate on lateral margin, distributed in western Atlantic . . . . . . . . . . . . . . . . . . 2
– Rostum armed with teeth with rounded apex on lateral margin, distributed in eastern
Atlantic and Mediterranean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 – Gastric median carina with two spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. armatus
– Gastric median carina unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. serratus
3 – Gastric median carina distinct, running from rostral median carina to cervical groove
except for interrupted median part; gastric lateral carinae linear, extending to cervical
groove. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. stirhynchus
– Gastric median carina thick and distinct; gastric lateral carinae present anteriorly . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Axius mediterraneus
Axius armatus Smith, 1881
Axius armatus Smith, 1881: 433; Kensley, 2001a: 951, figs. 1, 2, 4; Heard et al., 2007: 18,
fig. 15.
Axius (Paraxius) armatus — Borradaile, 1903: 538.
Axius (Axius) armatus — De Man, 1925d: 3 (list), 9, 11 (key).
?Calocarides armata — Sakai & De Saint Laurent, 1989: 79.
Glypturus acanthochirus — Manning, 1987 [fig. 4 = G. armata, after A. Milne-Edwards, 1870;
fig. 5 after Kensley, 1975].
Material examined. — USNM 35391, syntype, female (CL, 16.0; abdomen and telson,
33.0 mm), off Martha’s Vineyard, off Newport, R/V “Fish Hawk”, Sta. 878, 142 fms (259.7 m),
13.ix.1880.
Remarks. — Smith (1881: 433) described as follows “The rostrum is
narrow, acuminate, spiniform at the tip and armed along each edge with
four or five slender, acute, and spiniform teeth, directed forward and slightly
upward”. Those characteristics of the rostrum seem to be similar to those of
the rostrum of Axius serratus Stimpson, 1852 (USNM 80514) from Sheepscot
River, Barters Is., Maine (fig. 11). However, Smith (1881: 435) also mentioned
that “This species is distinguished from A. stirhynchus and A. serratus by the
narrower and acuminate rostrum, the teeth on the dorsal carina, the form of the
chelipeds, and the more slender second, third, and fourth pairs of legs”.
Kensley (2001b, fig. 2) distinguished those three species by the form of
the sternal plate between the P4 coxae; in A. armatus the sternal plate is
rounded; in A. serratus it is rectangular, bearing a pair of lateral spines; and in
A. stirhynchus it is rectangular, lacking a pair of lateral spines.
Type locality. — South coast of New England, 180-259 m.
Distribution. — Massachusetts to South Carolina, 108-260 m (from Kensley, 2001b).
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
67
Axius mediterraneus (Caroli, 1921)
(fig. 10)
Axiopsis mediterranea Caroli, 1921b: 254, fig. 1, pl. 9 figs. 1-4, pl. 10 figs. 5-16.
Axiopsis (Axiopsis) mediterranea — De Man, 1925d: 6 (list), 70 (key).
Material examined. — RMNH D 31798, holotype male (TL/CL, 60.0/20.3 mm, missing
Plp1), Posillipo, Naples, Italy, 17.xii.1915, leg. E. Caroli.
Remarks. — The male holotype of the present species (RMNH D 31798) is
similar to A. stirhynchus in both chelipeds and other parts (fig. 10B, C, D, E),
but they differ in that in A. mediterraneus the gastric median carina is thick and
distinct at full length (fig. 10A), whereas in A. stirhynchus the gastric median
carina, extending to the cervical groove, is interrupted in the median part of
the gastric region (fig. 12C).
Type locality. — Posillipo, Naples, Italy.
Distribution. — Posillipo, Naples, Italy.
Axius serratus Stimpson, 1852
(fig. 11)
Axius serratus Stimpson, 1852: 222; Smith, 1879: 55, pl. 10 figs. 4, 4a; Smith, 1881: 435;
Fowler, 1911: 571; Rathbun, 1929: 25, fig. 32; Williams, 1974b: 17, fig. 46A, B; Pemberton
et al., 1976: 790; Williams & Wigley, 1977: 3, 4, 5, 8, 16; Williams, 1984: 185, 187, fig. 130;
Sakai & De Saint Laurent, 1989: 28; Squires, 1990: 333, figs. 175-177; Kensley, 2001b: 955,
figs. 3, 4.
Axius armatus Smith, 1881: 433.
Axius (Paraxius) serratus — Borradaile, 1903: 538.
Axius (Axius) serratus — De Man, 1925d: 3 (list), 11 (key).
Material examined. — NHM 1976: 320, 2 ovig. females (TL/CL, ca. 89.0/27.2, carapace
broken — 28.3/31.3 mm), Nantucket, Massachusetts, Atlantic Ocean, ii.1928, leg. C. Biroseye,
det. W.L. Schmitt; NHM 1898.5.7.832, 1 female (TL/CL, 92.0/32.3 mm), Georges Bank,
North America, leg. Norman; NHM 1898.5.7.833-4, 2 ovig. females (TL/CL, 80.0/24.3;
87.0/25.6 mm), 1 female (TL/CL, 51.0/17.4 mm), ?Georges Bank, leg. Norman; USNM 80514,
1 ovig. female (TL/CL, 10.7/3.3 mm), Sheepscot River, Barters Island, Lincoln County, Maine,
leg. L.W. Scattergood, 24.x.1941, id. by W.L. Schmitt; USNM 213546, 1 ovig. female (TL/CL,
80.0/30.0 mm, lacking telson), Baltimore Canyon, United States, 1984, leg. A.B. Williams;
USNM 31451, 1 male (42.0/15.0 mm), Long Island Sound, 15 1/3 fms (28 m), leg. U.S. Fish
Commission, “Albatross” Sta. 1804.
Remarks. — The present species is closely similar to Axius stirhynchus
Leach, 1815, but differs as follows. In Axius serratus, the gastric lateral
and submedian carinae are short (fig. 11A); the maxilla 2 scaphognathite
bears one long whip, whereas in A. stirhynchus the gastric lateral carinae
are linear, extending to the cervical groove; and the maxilla 2 scaphognathite
has two long whips (Ngoc-Ho, 2003, fig. 1G). It is to be noted that in the
large female specimen (TL/CL, 51.0/17.4 mm) (NHM 1898.5.7.833-4), the
68
CRM 013 – Katsushi Sakai
Fig. 10. Axius mediterraneus (Caroli, 1921). A, carapace, dorsal view; B, larger cheliped, lateral
view; C, smaller cheliped, on left side, lateral view; D, P2, lateral view; E, male Plp2. AE, RMNH D 31798, holotype, male (TL/CL, 60.0/20.3 mm), Posillipo, Naples, Italy. Scales A,
2 mm; B, 0.8 mm; C, D, E, 1 mm.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
69
Fig. 11. Axius serratus Stimpson, 1852. Carapace, dorsal view. USNM 80514, 1 ovig. female
(TL/CL, 10.7/3.3 mm), Sheepscot River, Barters Island, Lincoln County, Maine. Scale 1 mm.
uropodal exopod bears a transverse suture, whereas in the far larger female
specimen (TL/CL, 92.0/32.3 mm) (NHM 1898.5.7.832) it bears no transverse
suture.
Type locality. — Scituate, Massachusetts Bay.
Distribution. — North America — Canada — Bay of Fundy to Long Island
Sound (Rathbun, 1929); U.S.A. — Massachusetts Bay (Stimpson, 1852), Cape
Ann (Smith, 1879), Scituate and George’s Banks, Massachusetts, Narragansett
70
CRM 013 – Katsushi Sakai
Bay, Rhode Island, off Stratford Point and Norwalk, Connecticut, 18-36 m.
Nova Scotia to Maryland, 18-220 m (Kensley, 2001b).
Axius stirhynchus Leach, 1815
(fig. 12)
Axius stirynchus Leach, 1815: 343; Leach, 1816, pl. 33; Bell, 1846: 228, fig. 1; White, 1847: 70;
White, 1850: 32; Stebbing, 1893: 187; Sinel, 1907: 217; Poulsen, 1940: 236 (key); Holthuis,
1962: 252; Zariquiey Alvarez, 1968: 223, fig. 88a.
Axius stirhynchus — Desmarest, 1825: 207, pl. 36 fig. 1; H. Milne Edwards, 1837a: 311;
H. Milne Edwards, 1837b: 131, pl. 48 fig. 2; White, 1857: 25; Norman, 1868: 177; Carus,
1885: 490; Lovett, 1885: 16; Koehler, 1886: 59; Gourret, 1888: 40; Norman & Scott, 1906:
13; Norman, 1907: 357; Sinel, 1907: 217; Schlegel, 1912: 237; Selbie, 1914: 89, pl. 14 figs.
1-4; Caroli, 1921a: 241; Caroli, 1921b: 254; Webb, 1921: 406, pl. 3 figs. 3, 6; Williamson,
1927: 554, fig. 208a, b, c; Perrier, 1929: 197; Monod, 1931: 119, 123, fig. 6E; Delphy &
Magne, 1938: 84; Bouvier, 1940: 93, fig. 65; Zariquiey Alvarez, 1946: 103, fig. 131; Drensky,
1951: 213; Kurian, 1956: 72, fig. 141; Gordon, 1957: 249; Bourdillon-Casanova, 1960: 99,
fig. 35; Holthuis, 1962: 247; O’Céidigh, 1962: 163; ICZN, 1964: 341; Bourdon, 1965: 15;
Allen, 1967: 17, 57 (key), 89, fig.; Števčić, 1969: 128; Thiriot, 1976: 349, 367; Holthuis,
1977: 57, pl. 5 fig. a; Beaubrun, 1979: 69, fig. 44; Moncharmont, 1979: 69; d’Udekem
d’Acoz, 1986: 101, fig. 1; Sakai & De Saint Laurent, 1989: 27, fig. 7; d’Udekem d’Acoz,
1989: 175; Moyse & Smaldon, 1990: 517, fig. 10.12; Števčić 1990: 214; Dworschak, 1992:
214; Koukouras et al., 1992: 223; Froglia, 1995: 7; Hayward et al., 1995: 432, fig. 8.51
(partim); d’Udekem d’Acoz, 1995: 47, fig. 1; Barnich, 1996: 130, figs. 28, 66; Falciai &
Minervini, 1996: 143, 1 fig.; Astall et al., 1997: 668; Pin et al., 1999a: 103, fig. 1H, 2A; Pin
et al. 1999b: 1461, figs. 1B, 4, tabs. 1-6; d’Udekem d’Acoz, 1999: 153, cover fig.; Costello
et al., 2001: 289; González-Gordillo et al., 2001: 279; Livory, 2001: 30; Martin, 2001: 75, 1
fig.; Türkay, 2001: 289; Ngoc-Ho, 2003: 447, figs. 1, 2; Ingle & Christiansen, 2004: 78, figs.
59, 62; Sakai & Sawada, 2006: 1357, figs. 9, 10.
Callianassa bisulcata Risso, 1844: 94; Monod, 1931: 119, 123, fig. 6E. [Nomen nudum.]
Axius Stirhynchus — Hope, 1851: 14.
Axius (Axius) stirhynchus — Borradaile, 1903: 537; De Man, 1925d: 3 (list), 11 (key).
Axia stirhynchus — ?Gibert I Olive, 1920: 51, 1 fig.; Ngoc-Ho, 2003: 447.
Axius Bisculcata Risso, no date: MS, figure (f); Holthuis, 1977: 57; Ngoc-Ho, 2003: 447.
Material examined. — NHM, 1950: 9.29.9-10, 2 males (TL/CL, 64.0/18.8, 59.0/17.4 mm),
Dusaihe Bay, Jersey, leg. R. Pike; NHM 82.15, 1 male (TL/CL, 67.0/20.3 mm), southern British
coast; NHM 2005.397, 1 male (TL/CL, 60.0/18.7 mm), Plymouth Laboratory, Wembury Chued
reef, 23.iii.1940, leg. R. Ingle; NHM 1910.2.48 8, 1 ovig. female (TL/CL, 85.0/24.3 mm),
Jersey, 1883, leg. Norman; NHM 1968: 52, 1 juv. (TL/CL, 14.0/4.4 mm), Ponlock Bay, leg.
J. Crothees Nettlecomke; NHM 1911.11.8.1038, 1 male (TL/CL, 39.0/12.6 mm), Seaford,
Sussex, U.K., leg. Norman; NHM 1992.1344, 1 male (TL/CL, 49.0/15.5 mm), Iles Chansey,
Atlantic coast of France, W. of Grand Ile, under rocks, 28.viii.1992, leg. A. Ceoguiee; ZMB
17127, 1 female (TL/CL, 57.0/17.5 mm), Nice, leg. Peters; ZMB 4878, 1 female (TL/CL,
55.0/17.0 mm), Venice, Italy, leg. Peter; MNHN Th 202, 1 male (TL/CL, 66.0/20.0 mm),
locality unknown, det. De Saint Laurent.
Diagnosis. — Rostrum stoutly triangular and obtuse at apex (fig. 12A-C),
lateral margins each with 4-7 teeth with rounded apex, extending posteriorly to
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
71
Fig. 12. Axius stirhynchus Leach, 1815. A, C, carapace, dorsal view; B, rostrum, dorsal view;
D, male Plp1; E, male Plp2. A, B, ZMB 17127, female (TL/CL, 57.0/17.5 mm), Nice; C, D, E,
MNHN Th 202, male (TL/CL, 66.0/20.0 mm), locality unknown, det. De Saint Laurent. Scales
1 mm.
72
CRM 013 – Katsushi Sakai
cervical groove as gastric lateral carinae. Gastric region convex, bearing submedian carinae, and median carina continued from rostral median carina extending posteriorly to cervical groove, except for interrupted median part with
hepatic pit of gastric region. Anterolateral margin of carapace unarmed. Cervical groove present at full length. Eyestalks subglobose; cornea pigmented.
A1 peduncle reaching level of distal margin of A2 penultimate segment. A2
scaphocerite prominent. P1 subequal in length, but dissimilar in shape, and
vertically positioned. P2 chelate. P3 propodus broadened, bearing setae on
ventral margin. P4 propodus elongate, bearing irregularly arranged spines ventrally on lateral surface. P5 subchelate. Pleurobranchs on Ps2-4. Abdominal
pleuron 1 triangular ventrally; pleura 1-5 smooth laterally, pleura 3-5 rounded,
with marginal tufts of soft setae ventrally. Male Plp1 uniramous and trisegmented; distal segment short, tapering distally (fig. 12D); male Plp2 biramous,
endopod with appendices interna and masculina at proximal third (fig. 12E);
Plps3-5 biramous, and broad and lamellar, endopods bearing appendix interna,
with marginal setae. Female Plp1 uniramous and bisegmented, consisting of
proximal segment and multiarticulate flagellum; Plps2-5 biramous, and broad
and lamellar, bearing marginal setae, endopods with appendix interna. Telson
longer than wide, bearing median tooth on posterior margin. Uropodal exopod
with transverse suture.
Remarks. — Ngoc-Ho (2003: 447, fig. 2E) described for this species that
the male Plp1 is small and unsegmented. However, it was confirmed through
the present observation that it is uniramous and bisegmented, and it was also
observed that the younger specimens bear plumose, thick setae on the margins
of the pleura, pleopods, and tail-fan.
Axius stirhynchus from Europe is closely similar to Axius serratus from
North America, as Smith (1879: 55) mentioned that A. serratus Stimpson,
1852 is very likely to be identical with the European S. [misspelled by Smith
for A.] stirhynchus. However, they differ from each other as follows. In the
type specimen of A. serratus, the rostrum is armed with sharp teeth on the
lateral margins (fig. 11), whereas in A. stirhynchus the rostrum is armed with
distinct teeth with a rounded apex on the lateral margins (fig. 12A-C).
De Man (1925d: 11) separated those two species, Axius stirhynchus from
Europe and Axius serratus from the western Atlantic, in his key as “Abdomen
less broad than carapace. Upper border of the palm in both chelipeds of 1st
pair thick and rounded, . . . in Axius stirhynchus” and “Abdomen broad and
depressed, expanding laterally in the middle, much broader than the carapace.
Upper border of the palm in both chelipeds of 1st pair thin and strongly
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
73
carinated, in Axius serratus”. However, it has turned out through careful
observations that no difference is found in the morphology of the abdomen
and the palms of both chelipeds between these two species, though they are
distinguishable from each other by the difference in shape of the lateral teeth
of the rostrum.
The rostrum is variable in shape; in the female specimen (ZMB 17127), it
is acutely triangular.
Type locality. — Sidmouth, England.
Distribution. — England — Sidmouth (Leach, 1815; Sakai & De Saint
Laurent, 1989), Weymouth (Sakai & De Saint Laurent, 1989), Cornwall,
Falmouth, Polperro, Plymouth, Seaford; Ireland (Selbie, 1914); Channel Islands; France — coasts of France (Sakai & De Saint Laurent, 1989), Roscoff;
Mediterranean — Marseille (Bouvier, 1940), Nice, Gulf of Naples (Caroli, 1921a), Tarragona (Zariquiey Alvarez, 1968); Adriatic Sea — Piran
(Dworschak, 1992); shallow water, in sand.
Genus Balssaxius gen. nov.
Diagnosis. — Rostrum acutely triangular and pointed at apex; lateral margins armed with teeth, extending posteriorly onto gastric region. Anterolateral
margin of carapace usually unarmed. Gastric region convex, bearing five carinae; median carina smooth, and submedian and lateral carinae serrated. Cervical groove distinct at full length. Eyestalks subglobose; cornea pigmented or
not. A2 scaphocerite elongate. Pl unequal; carpi and chelae characteristically
tuberculate on lateral surface, and dorsal margin denticulate or tuberculate.
P2 fingers longer than palm. Abdominal pleura smooth on surface, and convex to rounded ventrally. No pleurobranchs. Male Plps1-2 unknown. Female
Plp1 uniramous, lacking distal flagellum. Plps 2-5 biramous, similar, and slender; endopods with appendix interna. Telson longer than wide, bearing median
tooth on posterior margin. Uropodal exopod with transverse suture. Gonochoristic.
Remarks. — Axius habereri Balss, 1913, was included in Axiopsis by De
Man (1925d), and later in Calocarides by Sakai & De Saint Laurent (1989).
However, it does not belong to Calocarides either, because a distinct difference
is found between the species A. habereri and the species of Calocarides.
In Axius habereri, the carapace bears a sharp postcervical carina, whereas
in Calocarides coronatus (Trybom, 1904), the type species of Calocarides,
the carapace bears no postcervical carina, though they are similar in that the
rostrum is acutely triangular with denticulate lateral margins, and abdominal
74
CRM 013 – Katsushi Sakai
pleura 2-5 are rounded ventrally. Therefore, Axius habereri Balss, 1913 is
reclassified under the new genus Balssaxius as B. habereri (Balss, 1913).
Type species. — Axius habereri Balss, 1913, by present designation and
monotypy. The gender of the generic name, Balssaxius, is masculine.
Species includes. — Balssaxius habereri (Balss, 1913).
Etymology. — Balss is added as a prefix to the generic name Axius, in
honour of Heinrich Balss, German carcinologist in Munich, who described
Axius habereri.
Balssaxius habereri (Balss, 1913)
(fig. 13A, B)
Axius habereri Balss, 1913: 238; Nakazawa, 1927: 1036, fig. 1995; Yokoya, 1933: 49.
Axius Habereri — Balss, 1914: 85, figs. 46-47.
Axiopsis (Axiopsis) Habereri — De Man, 1925d: 5 (list), 70 (key).
Axiopsis (Axiopsis) habereri — Miyake, 1982: 89, pl. 30 fig. 3; Sakai, 1987a: 303 (list).
?Calocarides habereri — Sakai & De Saint Laurent, 1989: 83.
Material examined. — ZSM A20061782, lectotype female (TL/CL, 44.0/15.7 mm, larger
cheliped on left side missing), Fukuura, Sagami Bay, Japan, iii.1903, leg. Haberer, det. H. Balss;
ZLKU 9619, 1 female (TL/CL, 43.0/15.0 mm), Yellow Sea, 35◦ 02′ N 121◦ 56′ E, 50 m, by trawlnet, 19.x.1962, leg. H. Yamashita.
Diagnosis of female lectotype. — Rostrum acutely triangular, reaching middle of A1 penultimate segment, and denticulate on lateral margins; gastric region with five carinae; median carina distinct, with spine anteriorly, submedian
carinae denticulate, and lateral carinae smooth (fig. 13A); postcervical carina
present, extending to posterior margin of carapace, on which a pair of posterior
protrusions is present. Eyestalks subglobose, overreaching middle of rostrum;
cornea pigmented. A2 scaphocerite slenderly elongate, reaching distal third
of penultimate segment. A1 peduncle reaching middle of A2 penultimate segment. P1 unequal; in larger left cheliped, merus robust in shape, bearing two
subdistal denticles dorsally and 4 sharp teeth on ventromesial margin. Carpus
short and rounded on dorsal margin, and sharply pointed distoventrally. Palm
sharp on dorsal and ventral margins, convex and smooth on lateral surface.
Fixed finger bearing a row of tubercles laterally, and dactylus denticulate on
dorsal margin, bearing a row of spines on lateral surface; both fingers incurved
distally (Balss, 1914, fig. 46). Smaller cheliped (fig. 13B) on right side different in shape from larger one on the left; in smaller right cheliped merus
two times as long as wide, bearing two subdistal spines on dorsal margin and
6 sharp teeth on ventral margin; carpus triangular; chela setose along dorsal
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
75
Fig. 13. Balssaxius habereri (Balss, 1913) and Boasaxius princeps (Boas, 1880). A, anterior
part of carapace, dorsal view; B, smaller cheliped; C, A2 peduncle. A, B, Balssaxius habereri,
ZSM A20061782, lectotype female (TL/CL, 44.0/15.7 mm, larger cheliped on left side
missing), Fuku-ura, Sagami Bay, Japan; C, Boasaxius princeps, ZMM N2732, male (TL/CL,
114.0/36.4 mm), Peter the Great Bay.
and ventral margins; palm slightly longer than wide, bearing two small subdistal spines on dorsal margin; fingers longer than palm and crossed with each
other distally. Abdominal pleuron 1 narrow and triangular, with curved anteroventral and straight posteroventral margins, pleuron 2 long, and straight
76
CRM 013 – Katsushi Sakai
ventrally, with rounded antero- and posteroventral angles, pleura 3-5 moderately long, and rounded ventrally. Telson longer than wide, bearing proximal
lobe with spine, followed by denticles on lateral margins, but no median tooth
on rounded posterior margin. Uropodal exopod with transverse suture.
Type locality. — Fukuura, Sagami Bay, Japan.
Distribution. — Japan — Sagami Bay (Balss, 1913, 1914; Yokoya, 1933;
Sakai & De Saint Laurent, 1989), off Sioya-zaki (Yokoya, 1933), Aomori
Prefecture, Maizuru; Korea — off Pusan (Yokoya, 1933); Yellow Sea.
Genus Boasaxius gen. nov.
Diagnosis. — Body covered with tufts of setae. Rostrum triangular and upturned anteriorly, bearing 3-4 teeth on each lateral margin. Gastric region with
deep inverted U-shape surrounded by smooth lateral carinae extending posteriorly to near cervical groove; median and submedian carinae denticulate;
postcervical carina absent. Eyestalks subglobular, cornea pigmented, overreaching middle of rostrum. A2 scaphocerite horn-shaped (fig. 13C). Mxp3
ischial crest denticulate and protruded distally. P1 unequal and large, covered
with tufts of soft setae, fingers longer than palm in both larger and smaller chelipeds. P2 chelate. Ps3-5 simple. P3 propodus rectangular, armed with transverse rows of spines on ventrolateral surface, dactylus with spines on lateral
surface. Ps4-5 propodi elongate, armed with transverse rows of spines on ventrolateral surface; dactyli with spines on lateral surface. Abdominal somites
setose laterally and smooth ventrally. Male Plp1 uniramous, unsegmented, and
crenulate, terminating in narrow distal protrusion (fig. 14A); male Plp2 biramous, endopod with appendices interna and masculina (fig. 14B); male Plps35 biramous, endopods with appendix interna. Telson longer than wide, bearing
proximal lobe with spine followed by 1-2 spines on lateral margin, and median
spine on rounded posterior margin. Uropodal endopod with distinct spines laterally, bearing median carina with teeth. Uropodal exopod armed with sharp
teeth laterally and distinct, movable spine at posterolateral angle, bearing transverse suture with spinules, and distal flap with distinct spine, and median carina with spines on dorsal surface.
Remarks. — Axius princeps Boas, 1880 was once included in Axiopsis by
Borradaile (1903), and later included in Allaxius by Sakai & De Saint Laurent
(1989). However, it does not belong in Allaxius either, because Axius princeps
is clearly different from the type species of Allaxius, A. aethiopicus (Nobili,
1904). In Axius princeps the male Plp1 is present, and conspicuously sinuate
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
77
in shape (fig. 14A), whereas in Allaxius aethiopicus it is absent. Hence, Axius
princeps is reclassified under the new genus Boasaxius as B. princeps.
Type species. — Axius princeps Boas, 1880, by present designation and
monotypy. The gender of the name, Boasaxius, is masculine.
Species included. — Boasaxius princeps (Boas, 1880).
Etymology. — The generic name Boasaxius is dedicated to Johan Erik Vesti
Boas, Danish carcinologist, who published various carcinological studies in
the latter half of the 19th century.
Boasaxius princeps (Boas, 1880)
(figs. 13C, 14A, B)
Axius princeps Boas, 1880: 98, pl. 7 figs. 214-217; Balss, 1914: 88.
Axiopsis princeps — Borradaile, 1903: 538; Makarov, 1938: 48, fig. 15; Vinogradov, 1950: 223;
Komai et al., 1992: 196.
Axiopsis (Axiopsis) princeps — De Man, 1925d: 6 (list), 69 (key); Miyake, 1982: 90, pl. 30
fig. 4; Sakai, 1987a: 303 (list).
Allaxius princeps — Sakai & De Saint Laurent, 1989: 74.
Material examined. — ZMM N2732, 1 male (TL/CL, 114.0/36.4 mm), Peter the Great Bay,
1977, det. Zarenkov; ZLKU, 12084, 1 male (TL/CL, 95.0/35.0 mm), off Yoichi, Hokkaido,
Fig. 14. Boasaxius princeps (Boas, 1880) and Bouvieraxius rudis (Rathbun, 1906). A, C, male
Plp1; B, D, male Plp2. A, B, Boasaxius princeps, ZMM N2732, male (TL/CL, 114.0/36.4 mm),
Peter the Great Bay; C, D, Bouvieraxius rudis, MNHN Th 1004, male (TL/CL, 32.0/11.5 mm),
New Caledonia, 25◦ 15.40′ S 159◦ 54.80′ E, 280-295 m. Scales 1 mm.
78
CRM 013 – Katsushi Sakai
7.iii.1959, leg. H. Kurata; ZLKU 12085, 1 female (TL/CL, 49.2/19.7 mm), off Manazuru,
Sagami Bay, leg. T. Sakai; ZLKU 12084, 1 male (TL/CL, 95.0/35.0 mm), off Yoichi, Hokkaido,
10-30 m, by gill-net for flat fish, 7.iii.1956, leg. H. Kurata; ZLKU 12085, 1 ovig. female
(TL/CL, 49.2/19.7 mm), off Manazuru, Sagami Bay, Kanagawa Prefecture, leg. T. Sakai; SMF
32728, 1 female (TL/CL, 130.0/40.6 mm), East fishery-harbour, Okirai, Sanriku-cho, Ohfunatocity, Iwate Prefecture, Pacific side, gill-net, 140 m, 23.iv.1981, leg. H. Mukai.
Diagnosis. — Body covered with tufts of setae. Rostrum triangular and upturned anteriorly, bearing smooth median carina dorsally and 3-4 teeth on each
of lateral margins extending onto gastric region. Gastric region bearing deep
inverted U-shape surrounded by smooth lateral carinae extending posteriorly
to near cervical groove, median carina with 4 denticles anteriorly, divided in
middle into two carinae with 2-4 denticles each, then meeting together posteriorly into a single carina with 7 denticles, and submedian rows of denticles.
A2 scaphocerite horn-shaped, overreaching dorsodistal spine of segment 2
(fig. 13C). P1 unequal; in male larger cheliped palm convex with denticles on
distal margin, bearing 6 spines dorsally, fixed finger roughly provided with obtuse teeth on cutting edge, dactylus also roughly provided with obtuse teeth on
cutting edge; in smaller cheliped fingers longer than palm; dactylus finely denticulate on cutting edge, fixed finger irregularly denticulate on cutting edge.
P2 chelate, fingers about as long as palm. P3 propodus rectangular, armed
with transverse rows of spines on ventrolateral surface, dactylus with row of
spines on lateral surface. Ps4-5 propodi elongate, armed with transverse rows
of spines on ventrolateral surface; dactylus with a row of spines on lateral surface. Abdominal pleuron 1 narrow, and rounded ventrally, pleuron 2 long, and
straight with rounded angles ventrally, pleuron 3 moderately long, and straight
ventrally with rounded angles; pleura 4-5 moderately broad and straight ventrally, with rounded anteroventral and angulate posteroventral angles; pleuron
6 triangular ventrally. Male Plp1 uniramous, unsegmented, and sinuate, terminating in narrow distal protrusion (fig. 14A); male Plp2 biramous, endopod
with appendices interna and masculina (fig. 14B); male Plps3-5 endopods with
appendix interna. Telson longer than wide, bearing proximal lobe with spine
followed by 1-2 spines on lateral margin, and median spine on rounded posterior margin. Uropodal endopod with distinct spines laterally, bearing median
carina with spines dorsally. Uropodal exopod armed with sharp teeth laterally and distinct, movable spine at posterolateral angle, bearing transverse suture with spinules, and distal flap with distinct spine; with median carina with
spines on dorsal surface.
Type locality. — Vladivostok.
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79
Distribution. — Vladivostok (Boas, 1880; Balss, 1914); Japan — off Yoichi,
Hokkaido, Sagami Bay (Sakai & De Saint Laurent, 1989), Oga Peninsula
(Miyake, 1982); 300 m.
Genus Bouvieraxius Sakai & De Saint Laurent, 1989
Bouvieraxius Sakai & De Saint Laurent, 1989: 45; Sakai, 1992a: 165; Poore, 1994: 97 (key);
Sakai, 1994: 176; Poore, 2007: 161; Poore & Collins, 2009: 244.
Posthonocaris Kensley, 1989: 964.
Diagnosis. — Rostrum elongate and acutely triangular with or without lateral spine, bearing a pair of supraocular teeth at base. Gastric region convex,
bearing smooth lateral carinae extending posteriorly to anterior half, submedian rows of 3-4 spines, and smooth median carina with hepatic tubercle only.
Anterolateral margin of carapace unarmed. Cervical groove distinct at full
length. Eyestalks subglobose; cornea pigmented. A2 scaphocerite distinctly
protruded. P1 subequal; palm tuberculate on dorsal margin, bearing small dorsodistal spine. Ps3-4 propodi with transverse rows of spines ventrolaterally;
dactyli with spines ventrally. P5 propodus with spinules at ventrodistal corner.
Abdominal pleura smooth on lateral surface, and truncate ventrally. Male Plp1
uniramous and bisegmented; distal segment rounded apically, proximally bearing mesial lobe with patch of hooklets; male Plp2 biramous; endopod bearing
appendix interna and elongate appendix masculina at proximal third, or endopod bearing appendix interna, but no appendix masculina, or endopod without
appendices interna and masculina; male Plps3-5 endopods without appendix
interna. Telson longer than wide, bearing proximal lobe on lateral margin, and
median spine on posterior margin. Uropodal endopod bearing spines on lateral margin and median carina with spines on dorsal surface. Uropodal exopod
bearing transverse suture with strong, movable spine at posterolateral angle.
Hermaphroditic.
Remarks. — The present genus Bouvieraxius is hermaphroditic, because
the Ps3 and 5 coxae bear genital pores. However, it was observed that there is
difference in the form of Plp2 between males and females (Sakai, 1992a, fig.
8E), namely, this genus has three types of male Plp2 endopod: (1) Plp2 endopod with appendices interna and masculina as in B. longipes (Bouvier, 1905);
(2) Plp2 endopod with appendix interna, but without appendix masculina as
in B. rudis (Rathbun, 1906); (3) Plp2 endopod without appendices interna and
masculina as in B. springeri Kensley, 1996. The present genus is characteristic
also in that Plps3-5 endopods bear no appendix interna as in Paratrichocheles
gen. nov. (see below).
80
CRM 013 – Katsushi Sakai
Type species. — Axius longipes Bouvier, 1905, by original designation. The
gender of the generic name, Bouvieraxius, is masculine.
Species included. — Bouvieraxius keiensis Sakai, 1992a; B. longipes (Bouvier, 1905); B. michelae Poore, 2008; B. rudis (Rathbun, 1906); B. springeri
Kensley, 1996e.
K EY
TO THE
S PECIES
OF THE
G ENUS BOUVIERAXIUS
1 – Rostrum bearing 4 spines on each lateral margin including supraorbital spine . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
– Rostrum with one spine or none on lateral margin, except for supraorbital spine . . . . . 3
2 – Gastric submedian carina with 3 spines; A2 article 2 armed with two ventral spines . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .B. longipes
– Gastric submedian carina with 4 spines; A2 article 2 unarmed ventrally . . . B. michelae
3 – Rostrum without lateral spine except for supraorbital spine . . . . . . . . . . . . . . B. springeri
– Rostrum with lateral spine except for supraorbital spine . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – Gastric submedian carina with row of 4 spines . . . . . . . . . . . . . . . . . . . . . . . . . . . B. keiensis
– Gastric submedian carina with row of 3 spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. rudis
Bouvieraxius keiensis Sakai, 1992
Bouvieraxius keiensis Sakai, 1992a: 166, figs. 8, 9; Kensley, 1996e: 472 (list); Poore & Collins,
2009: 245, figs. 15, 16, 41.
Type locality. — Kei Island, 5◦ 37′ S 132◦ 23′ E, 245 m, sand.
Distribution. — Mauritius (Sakai, 1992a); Indonesia — Kei Islands (Sakai,
1992a); 73-245 m; Australia — North West Shelf to Shark Bay, Western
Australia (Poore & Collins, 2009); 80-100 m.
Bouvieraxius longipes (Bouvier, 1905)
Axiopsis longipes Bouvier, 1905: 804; Balss, 1925: 209; Bouvier, 1925: 466, figs. 26, 27, pl. 9
fig. 5, pl. 10 fig. 5.
Axiopsis (Axiopsis) longipes — De Man, 1925d: 6 (list), 70.
Bouvieraxius longipes — Sakai & De Saint Laurent, 1989: 45; Kensley, 1996e: 472 (list).
Diagnosis. — Rostrum with 4 spines on each lateral margin. Gastric region
with submedian carina with 3 elongate spines. Male Plp1 uniramous and
bisegmented, distal segment rounded distally, bearing mesial lobe; male Plp2
biramous, endopod with appendices interna and masculina (Bouvier, 1925,
fig. 27).
Type locality. — Barbados, 225 m.
Distribution. — Porto Rico; Barbados (Bouvier, 1925).
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81
Bouvieraxius michelae Poore, 2008
Bouvieraxius michelae Poore, 2008: 162, fig. 1.
Diagnosis. — Rostrum with 4 spines on each lateral margin. Gastric
submedian carina with 4-5 spines. Male Plp1 a broad blade, dilated distally,
with oblique apex ending in two lobes with three minute coupling hooks; Plp2
endopod with appendices interna and masculina at proximal two-thirds. Telson
longer than wide, bearing three spines on each lateral margin, and a posterior
spine that is longer and movable on posterolateral angle; with median tooth
on posterior margin. Uropodal endopod broadened, bearing 3 teeth on lateral
margin and median carina with 4 teeth on dorsal surface. Uropodal exopod
bearing two teeth on lateral margin, a distinct, movable spine at posterolateral
angle, and a transverse suture with spinules dorsally. [Adapted from Poore,
2008.]
Type locality. — Timor Sea, 18 m.
Distribution. — Only known from the type locality.
Bouvieraxius rudis (Rathbun, 1906)
(fig. 14C, D)
Axius rudis Rathbun, 1906: 894, fig. 51; Balss, 1925: 209.
Axiopsis (Axiopsis) rudis — De Man, 1925d: 6 (list), 70 (key).
Bouvieraxius rudis — Sakai & De Saint Laurent, 1989: 46, figs. 12-14; Sakai, 1994: 177, 200
(key); Kensley, 1996e: 472 (list).
Lophaxius rudis — Kensley, 1989: 964.
Material examined. — USNM 30535, lectotype, female (TL/CL, 22.0/6.5 mm), S. coast of
Molokai Is., Hawaii, 4.iv.1902, leg. “Albatross” Sta. 3838, 92-212 fms (168.2-387.7 m), det.
M. Rathbun; MNHN Th 445, 1 female (TL/CL, 55.0/23.0 mm), Madagascar, 8.xi.1972, leg.
A. Crosnier; MNHN Th 1004, 1 male (TL/CL, 32.0/11.5 mm), New Caledonia, 25◦ 15.40′ S
159◦ 54.80′ E, MUSORSTOM V, Sta. 0255, 280-295 m.
Diagnosis. — Male Plp1 (fig. 14C) uniramous and bisegmented, proximal
segment foliaceous, distal segment ovate distally, mesially bearing triangular
protrusion without cluster of hooklets; male Plp2 (fig. 14D) biramous, endopod slender, distally bearing rod-shaped appendix interna with cluster of hooklets at proximal third, but no appendix masculina; male Plps3-5 biramous, endopods without appendix interna. Female Plp1 uniramous and bisegmented,
consisting of stem-like proximal segment and multiarticulate flagellum. Female Plps2-5 biramous, endopods without appendix interna.
Type locality. — Molokai Island, Hawaiian Islands, 168-388 m.
Distribution. — Madagascar (Sakai & De Saint Laurent, 1989); Western
Australia (Sakai, 1994); New Caledonia (Sakai & De Saint Laurent, 1989);
82
CRM 013 – Katsushi Sakai
Hawaiian Islands — Molokai Island and Kauai Island (Rathbun, 1906); 72419 m.
Bouvieraxius springeri Kensley, 1996
Bouvieraxius springeri Kensley, 1996e: 472, figs. 2, 3, table 2; Kensley, 2003: 366, pl. 4.
Material examined. — USNM 252832, 1 herm. (TL/CL, 49.0/17.0 mm), N. Alutom Island,
Agat Bay, Guam, 2.5-6 m, 29.xi.2000, leg. H. Conley; USNM 296339, 1 herm. (TL/CL,
45.0/15.1 mm), Piti Reef, Guam Is., 1.5-2.5 m, ix.1998, leg. H.T. Conley; USNM 296398,
1 herm. (TL/CL, 50.0/17.7 mm), breakwater, Apra Harbor, Guam, 5.9 m, 21.vi.1998, leg.
H.T. Conley.
Remarks. — This species is hermaphroditic, because the Ps3 and 5 coxae
have genital pores. In the specimen USNM 252832, the Plp2 endopod has
appendices interna and masculina, and the appendix masculina is much longer
than the appendix interna. However, in the other specimens (USNM 296339;
296398) the Plp2 endopod lacks appendices interna and masculina as in the
Plps3-5 endopods, which was also indicated by Kensley (2003: 367), who
described in his remarks that “B. springeri is a functionally hermaphroditic
species, but whether protandrous or protogynous, it is not yet possible to
establish”.
Type locality. — Sulu Sea, 10◦ 52′ 30′′ N 120◦ 56′ 00′′ E, 0-14 m.
Distribution. — Sulu Sea (Kensley, 1996e); 0-14 m.
Genus Bruceaxius gen. nov.
Diagnosis. — Body covered with setae. Rostrum narrow and elongate,
acutely triangular and deflected downward anteriorly, bearing spines on lateral margins. Gastric region slightly convex, bearing lateral, submedian, and
median carinae, each with denticles almost in parallel with one another.
Postcervical carina present. Anterolateral margin of carapace with spine. Cervical groove distinct at full length. Eyestalks subglobose; cornea pigmented.
A2 scaphocerite horn-shaped and directed outside dorsodistal spine of A2 segment 2. P1 unequal; chelae obliquely positioned, surface covered with thick
setae and unarmed on dorsal margin; fingers of larger cheliped shorter than
palm, but those of smaller cheliped distinctly longer than palm. No pleurobranchs. Abdominal somites 1-5 each with longitudinal carina between tergum
and pleuron. Pleuron 1 narrow, pleura 2-5 truncate or rounded at ventral margin. Male Plp1 uniramous and bisegmented, distal segment digitiform, male
Plp2 biramous and elongate, endopod with appendices interna and masculina
at proximal third; male Plps3-5 biramous, endopods with appendix interna.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
83
Female Plp1 uniramous and bisegmented, composed of proximal segment and
multiarticulate flagellum; female Plps2-5 endopods with appendix interna. Telson longer than wide, bearing proximal lobe with spine on each lateral margin,
and median tooth on posterior margin. Uropodal endopod broadened, bearing
two teeth on lateral margin and median carina with three teeth on dorsal surface. Uropodal exopod bearing 5-6 teeth on lateral margin and distinct, movable spine at posterolateral angle, and transverse suture with spinules. Gonochoristic.
Remarks. — Acanthaxius polychaetes Sakai, 1994 is closely similar to
Acanthaxius pilocheirus Sakai, 1987a, the type species of Acanthaxius. However, they differ in that in the A. polychaetes male, Plp1 is uniramous and
bisegmented, and the distal segment is digitiform; the anterolateral margin of
the carapace is armed with a spine; the postcervical carina is present; in P1
chelae, the lateral surfaces are covered with thick setae, and unarmed on the
dorsal margin; the fingers of the larger cheliped are shorter than the palm, but
those of the smaller cheliped are distinctly longer than the palm, and abdominal
somites 1-5 bear a longitudinal carina between the tergum and the pleuron. In
A. pilocheirus, the male Plp1 is absent; the anterolateral margin of the carapace
is unarmed, the postcervical carina of the carapace is absent; in the P1 chelae
the lateral surfaces are covered with tubercles and armed with sharp teeth on
the dorsal margin; the fingers of both chelipeds are longer than the palm, and
abdominal somites 1-5 bear no longitudinal carina between tergum and pleuron. Those differences show that they are distinctly different from each other
at the generic level, which suggests that A. polychaetes should be reclassified
under the new genus Bruceaxius as B. polychaetes.
Type species. — Acanthaxius polychaetes Sakai, 1994, by present designation and monotypy. The gender of the new name, Bruceaxius, is masculine.
Species included. — Bruceaxius polychaetes (Sakai, 1994).
Etymology. — The new genus is named Bruceaxius in honour of Dr.
A.J. Bruce of the Northern Territory Museum, Port Darwin, Australia, who
sent me this interesting species for study in 1994, by adding his name as a
prefix to the generic name Axius.
Bruceaxius polychaetes (Sakai, 1994)
Acanthaxius polychaetes Sakai, 1994: 193, 200 (key), figs. 11-13; Kensley, 1996b: 71 (list);
Poore & Collins, 2009: 237.
Diagnosis. — Male Plp1 uniramous and bisegmented, proximal segment
rod-like, distal segment digitiform; male Plp2 biramous; both endopod and
84
CRM 013 – Katsushi Sakai
exopod slenderly foliaceous, endopod with appendices interna and masculina
at proximal two-thirds, appendix masculina longer than appendix interna.
Type locality. — Great Barrier Reef, eastern Australia, 17◦ 57.5′ S
147◦ 03.5′ E, 259-260 m.
Distribution. — Queensland, eastern Australia (Sakai, 1994); continental
slope, 259-260 m.
Genus Calaxius Sakai & De Saint Laurent, 1989
Calaxius Sakai & De Saint Laurent, 1989: 84; Poore, 1994: 97 (key); Sakai, 1994: 192.
Manaxius Kensley, 2003: 367 [sensu nov.]; Clark et al., 2007: 64; Poore & Collins, 2009: 248.
Diagnosis. — Rostrum elongate and acutely triangular, with 4-5 sharp lateral spines including supraorbital spine, lateral margins extending posteriorly
onto gastric region as lateral carinae, with at least 1-2 distinct, buttressed teeth;
median carina originating from near tip of rostrum, running to cervical groove;
submedian rows of teeth; postcervical carina present. Anterolateral margin of
carapace with protrusion or spine. Cervical groove distinct at full length. Eyestalks stout or elongate; cornea pigmented. A2 scaphocerite protruded straight
forward. P1 subequal; palm armed with 3-4 interspaced, strong teeth on dorsal
margin; fingers longer than palm. P2 fingers shorter than palm. P2-4 with pleurobranchs. Abdominal somites with or without longitudinal carina between
terga and pleura, pleura sharply pointed ventrally. Male Plp1 absent; male Plp2
biramous, endopod with appendices interna and masculina at proximal third;
male Plps3-5 endopods with appendix interna. Female Plp1 uniramous and
bisegmented, consisting of proximal segment and flagellum; female Plps25 endopods with appendix interna. Telson longer than wide, bearing proximal lobe on lateral margin and no median tooth on rounded posterior margin.
Uropodal exopod with transverse suture.
Remarks. — The type species of the genus Calaxius, C. acutirostris Sakai &
De Saint Laurent, 1989, is similar to Eutrichocheles modestus (Herbst, 1796)
in that the abdominal pleura are triangular or sharply pointed on the ventral
margin. However, they are utterly different from each other in Plps1-5. In C.
acutirostris, the male Plp1 is absent, the male Plp2 endopod bears appendices
interna and masculina at its proximal third, and male Plps3-5 endopods bear
an appendix interna. In contrast, in E. modestus the male Plp1 is present as
a stubby protrusion, the male Plp2 endopod bears an appendix interna but no
appendix masculina, and male Plps3-5 endopods lack an appendix interna.
Type species. — Calaxius acutirostris Sakai & De Saint Laurent, 1989, by
original designation. The gender of the generic name, Calaxius, is masculine.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
85
Species included. — Calaxius acutirostris Sakai & De Saint Laurent, 1989
[= C. tungi Zhong, 2000; C. manningi Kensley, Lin & Yu, 2000]; Calaxius sp.
Robles et al., 2009.
Calaxius acutirostris Sakai & De Saint Laurent, 1989
Calaxius acutirostris Sakai & De Saint Laurent, 1989: 86, figs. 23-25; Sakai, 1994: 192, 200
(key), fig. 10; Komai, 2000b: 344 (list); Kensley & Hickman, 2001: 487; Poore & Collins,
2009: 250, figs. 18-20, 43.
Calaxius tungi Zhong, 2000: 33, figs. 1-11. [Type locality. — Northern continental slope of the
South China Sea, 367-402 m.]
Calaxius manningi Kensley, Lin & Lu, 2000: 207, figs. 2, 2, 7A; Tsang et al., 2008: 218, 219;
Chu et al., 2008. [Type locality. — Tai-Shi, N.E. Taiwan, 500 m.]
Material examined. — MNHN Th 447, holotype male (TL/CL, 46.0/19.0 mm), Madagascar,
leg. A. Crosnier; MNHN Th 449, 1 male (TL/CL, 60.0/24.0 mm), 15◦ 19.1′ S 46◦ 11.8′ E,
P4 chalutage 46, 400 m, 7.xi.1972, leg. A. Crosnier; MNHN Th 1176, 1 male (TL/CL,
23.0/8.5 mm), Guinea, Guinea Trawling Survey, Sta. 11, 25.viii.1966 (paratype of Calaxiopsis
serrata).
Diagnosis. — Male Plp1 absent; male Plp2 endopod with appendices
interna and masculina; male Plps3-5 endopods with appendix interna. Ps3 and
5 coxae with genital pores. Hermaphroditic.
Remarks. — Kensley et al. (2000: 209) mentioned that “Calaxius acutirostris (from the Philippines) appears to differ from C. manningi in having
4 spines on the posterior (= ventral) meral margin of pereiopod, the lateral
margin of the inner uropodal ramus bears three spines [though 1-2 spines in
Sakai, 1989, fig. 25], while the telson margins are serrate”. However, those differences found between the two species are not to be regarded as differences
that are decisive to distinguish one species from the other, but as acceptable
morphological variations at the species level. They may have been caused by
sexual difference, considering that the holotype specimen of C. acutirostris is
a male and that of C. manningi is a female, and they are almost of the same
size.
The male holotype of Calaxius tungi Zhong, 2000 (TL/CL, 59.5/23.0 mm)
from the northern continental slope of the South China Sea, 367-402 m, which
is larger than the male holotype of C. acutirostris (TL/CL, 46.0/19.0 mm) and
the female holotype of C. manningi (CL, 19 mm), is characterized as follows.
The ventral margin of the P1 merus is armed with 3-4 spinules, the lateral
margin of the uropodal endopod bears two spines, and the telson bears one
lateral spine. The difference in the number of spines or teeth among C. tungi,
C. manningi, and C. acutirostris (see table III) is considered acceptable as
a variation, which safely suggests that C. manningi and C. tungi should be
regarded synonymous with C. acutirostris.
86
CRM 013 – Katsushi Sakai
TABLE III
Comparison of relevant characters in three nominal species of the genus Calaxius
C. acutirostis Sakai
& De Saint Laurent,
from Madagascar
C. manningi Kensley,
Lin & Lu,
from N.E. Taiwan
C. tungi Zhong,
from the South
China Sea
Uropodal endopod
lateral margin
with 2-3 spines
including
distolateral one
with 1 distolateral
spine
with 2 spines
including
distolateral one
Telson lateral
margins
with 2-3
denticles
with 2 spines
with 2 spines
P1, palm of
larger chela on
anterior margin
unarmed (holotype
male, MNHN Th 447)
(left chela), or with
1 triangular tooth
(male, MNHN Th 944)
(left chela)
with tubercles
(left chela)
with 2 triangular
teeth
P1, palm of
larger chela
on dorsal margin
with 3 spines
with 3 spines
with 1 spine
P1, merus of
larger chela
on ventral margin
with 3-4 spines
with 3-4 spines
unarmed
Type locality. — Madagascar.
Distribution. — East Africa (Sakai & De Saint Laurent, 1989); Madagascar
(Sakai & De Saint Laurent, 1989); Australia — northern and eastern Australia
(Sakai, 1994); northern continental slope of the South China Sea, 367-402 m
(Zhong, 2000); Philippines — south of Mindoro and Tablas Strait (Sakai & De
Saint Laurent, 1989); Tai-Shi, N.E. Taiwan, 500 m (Kensley et al., 2000).
Calaxius sp., Robles et al., 2009
Calaxius sp. Robles et al., 2009: 316.
Genus Calocarides Wollebæk, 1908
Euconaxius Trybom, 1904: 383.
Calocaris (Calocarides) Wollebæk, 1908: 3, 23.
Axiopsis (Calocarides) — De Man, 1925d: 1, 2 (key), 6, 67, 71; Balss, 1957: 1579.
Calocarides — Bouvier, 1940: 97; Sakai & De Saint Laurent, 1989: 4, 78; Sakai, 1992a: 176;
Poore, 1994: 98 (key); Kensley, 1996a: 53; Ngoc-Ho, 2003: 451; Ingle & Christiansen,
2004: 81.
Diagnosis. — Rostrum triangular and pointed at apex; lateral margins armed
with teeth, extending posteriorly onto gastric region. Anterolateral margin of
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
87
carapace usually unarmed. Gastric region convex, bearing median and lateral carinae with teeth, and submedian carina with or without teeth. Cervical
groove distinct at full length. Eyestalks subglobose; cornea pigmented or not.
A2 scaphocerite elongate. Pl unequal; carpi and chelae characteristically tuberculate on lateral surfaces, and their dorsal margins denticulate or tuberculate.
P2 chelae longer than palm. Abdominal pleura smooth on lateral surface and
convex to rounded ventrally. No pleurobranchs. Male Plp1 absent; male Plp2
biramous, endopod with appendices interna and masculina at proximal third,
appendix interna slender and longer than appendix masculina; male Plps3-5 biramous and slender, endopods with appendix interna. Telson longer than wide
and oblong, bearing median tooth on posterior margin. Uropodal exopod with
transverse suture. Gonochoristic. [Adapted from Sakai & de Saint Laurent,
1989.]
Type species. — Euconaxius coronatus Trybom, 1904, by subsequent
designation by Sakai & De Saint Laurent (1989). The gender of the generic
name, Calocarides, is masculine.
Species included. — Calocarides capensis Kensley, 1996a [= C. longispinis
Stebbing, 1910]; C. chani Kensley, Lin & Yu, 2000; C. coronatus (Trybom,
1904) [= C. crassipes (Trybom, 1904)]; C. laevis (Bouvier, 1915a); C. lev
(Zarenkov, 1989); C. longispinis (McArdle, 1901); C. macphersoni Kensley,
1996a; C. okhotskensis sp. nov.; C. quinqueseriatus (Rathbun, 1902); C.
rostriserratus (Andrade & Báez, 1977); C. rudolfi (Zarenkov, 1989); C. soyoi
(Yokoya, 1933); C. tenuicornis (De Man, 1905); C. vigilus Sakai, 1992c.
K EY
TO THE
S PECIES
OF THE
G ENUS CALOCARIDES
1 – A2 scaphocerite short; gastric submedian carinae without teeth; telson 1.7 times as
long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laevis
– A2 scaphocerite elongate; gastric submedian carinae with teeth; telson less than 1.5
times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – A2 scaphocerite failing to reach end of A2 penultimate segment . . . . . . . . . . . . . . . . . . 3
– A2 scaphocerite reaching or overreaching A2 penultimate segment . . . . . . . . . . . . . . . 10
3 – P1 palm finely tuberculate on dorsal margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– P1 palm roughly denticulate on dorsal margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4 – P1 meri with one subdistal spine dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. chani
– P1 meri serrate on distal half of dorsal margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. soyoi
5 – Carapace shorter than length of abdominal somites 1-5 . . . . . . C. okhotskensis sp. nov.
– Carapace equal to length of abdominal somites 1-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – P1 fingers as long as palm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. quinqueseriatus
– P1 fingers shorter than palm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 – Gastric median carina with strong spine anterior to hepatic tubercle . . . . . . . . . . . . . . . 8
88
CRM 013 – Katsushi Sakai
– Gastric median carina without strong spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 – A2 scaphocerite reaching distal third of A2 penultimate segment . . . . . . C. longispinis
– A2 scaphocerite reaching distal end of A2 penultimate segment . . . . . . C. macphersoni
9 – Telson unarmed on lateral margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rudolfi
– Telson with proximal lobe followed by 1-2 spinules laterally . . . . . . . . C. rostriserratus
10 – Gastric submedian carinae with 1-2 spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
– Gastric submedian carinae with more than 4 spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 – Gastric median carina unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. capensis
– Gastric median carina armed with 1-2 spines anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . 13
12 – In larger cheliped carpus with two subdistal spines on dorsal margin; cutting edge of
dactylus concave in proximal half, bearing subproximal protrusion . . . . . C. coronatus
– In larger cheliped carpus denticulate on subdistal part of dorsal margin; cutting edge
of dactylus concave in proximal half, lacking subproximal protrusion . . . . . . . . . C. lev
13 – Gastric median carina denticulate; gastric submedian carinae with a row of 7 spines . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vigilus
– Gastric median carina with 4 interspaced spines; gastric submedian carinae with a row
of 3-4 spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tenuicornis
K EY
TO THE
S PECIES
OF
CALOCARIDES BY G EOGRAPHICAL D ISTRIBUTION
Indo-West Pacific species. — C. chani; C. longispinis; C. okhotskensis sp. nov.; Calocarides
soyoi.
West Atlantic species. — C. capensis; C. coronatus; C. laevis; C. macphersoni; C. rudolfi; C.
tenuicornis; C. vigilus.
East Pacific species. — C. lev; C. quinqueseriatus; C. rostriserratus.
Calocarides capensis Kensley, 1996
Calastacus longispinis Stebbing, 1910: 367 [not McArdle, 1901] [Type locality. — [Off] Table
Mountain, 457 m.]
Calocaris (Calastacus) longispinis — Barnard, 1950: 502 (key), 503, fig. 93d-f [not McArdle,
1901].
Calocaris longispinis — Kensley, 1981b: 30 [not McArdle, 1901]; Bianchi et al., 1993: 34.
Calocarides capensis Kensley, 1996a: 54 (list), 55, fig. 2 [not Trybom, 1904].
Calocarides coronatus — Sakai & De Saint Laurent, 1989: 80, fig. 20 (not 21).
Material examined. — USNM 243561, holotype, male (TL/CL, 74.0/24.0 mm), South
Atlantic Ocean, Namibia, N.W. of Swakopmund, 20◦ 55′ S 12◦ 23′ E, 475-480 m, 9.ix.1980, R/V
“Benguela II”, Sta. 66, det. as Calastacus longispinis Stebbing, 1910; MNHN Th 1044, 1 male
(TL/CL, 72.5/23.5 mm); MNHN Th 1045, 1 male (TL/CL, 68.0/23.0 mm), 26◦ 26′ S 14◦ 25′ W,
300 m, 1981, leg. R/V “Benguela IV” (Sakai & De Saint Laurent, 1989: 80, fig. 20, det. as
Calocarides coronatus (Trybom, 1904)).
Diagnosis. — Rostrum acutely triangular, bearing 5-6 lateral teeth. A2
scaphocerite slightly overreaching distal margin of penultimate segment.
Gastric median carina distinct, with hepatic tubercle, extending posteriorly
from middle of rostrum; gastric submedian rows of 1-2 buttressed spines;
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
89
lateral carinae with 3-4 spines. In larger cheliped merus with five distal spines
on dorsal margin and one distinct spine on ventral margin; palm with about 11
strong spines on dorsal margin; in smaller cheliped, merus with three spines on
dorsal margin and one distinct spine on ventral margin, and palm with about
11 strong spines on dorsal margin. Telson bearing 3-4 movable denticles at
posterolateral corner and a pair of spines on dorsal surface; posterior margin
gently convex with median spine. [Synopsis from Kensley, 1996a.]
Type locality. — Forty (40) miles N 79◦ E of Table Mountain, Cape of Good
Hope, South Africa, 1280 m.
Distribution. — Tropical west Africa (Sakai & De Saint Laurent, 1989);
Namibia — off Lüderitz (Sakai & De Saint Laurent, 1989); South Africa —
off Table Mountain (Stebbing, 1910; Kensley, 1996a).
Calocarides chani Kensley, Lin & Yu, 2000
Calocarides chani Kensley et al., 2000: 209, figs. 3, 4, 7B-D; Tsang et al., 2008: 218, 219; Chu
et al., 2008.
Type locality. — Tai-Shi fishing port, I-Lan County, N.E. Taiwan, 400 m.
Distribution. — I-Lan County, N.E. Taiwan; 300-600 m.
Calocarides coronatus (Trybom, 1904)
Euconaxius coronatus Trybom, 1904: 384, pl. 20 figs. 1-10, 13-14, pl. 21 figs. 1-8; Lagerberg,
1908: 48, pl. 2 fig. 7.
Euconaxius crassipes Trybom, 1904: 390, pl. 20 figs. 11, 12. [Type locality. — Norway,
Kosterfjord.]
Calocaris (Calocarides) coronatus — Wollebæk, 1908: 3, 5, 13, 23.
Calocaris (Calocarides) crassipes — Wollebæk, 1908: 3, 23, pls. 1-7.
Calocarides coronatus — Wollebæk, 1909a: 3; Björck, 1913: 7; Balss, 1925: 209; Balss, 1926:
26; Grieg, 1927: 43; Bouvier, 1940: 97; Poulsen, 1940: 208, 216, fig. 4; Christiansen, 1955:
1; Elofsson, 1959: 1, figs. 1-20; Brattegard, 1966: 45, 1 text-fig., 2 tables; Allen, 1967: 57
(key), 89 (fig.); Christiansen, 1972: 40, fig. 45; Noël, 1992: 79; Cates et al., 1994: 137;
García Raso, 1996: 738: 56; Kensley, 1996a: 54 (list), 56; Brattegard & Christiansen, 1997:
220; d’Udekem d’Acoz, 1999: 153; Christiansen, 2000: 231; Costello et al., 2001: 289;
González-Gordillo et al., 2001: 279; Türkay, 2001: 289; Ngoc-Ho, 2003: 451, figs. 2, 3;
Ingle & Christiansen, 2004: 81, figs. 60, 63.
Axiopsis (Calocarides) coronata — De Man, 1925d: 6 (list), 67, 71 (key).
Non: Calocarides coronatus — Sakai & De Saint Laurent, 1989: 80, fig. 20 (not 21)
[= Calocarides capensis Kensley, 1996a: 55].
Material examined. — SMF 29463, 1 male (TL/CL, 60.0/18.8 mm), 2 females (TL/CL,
54.0/18.7-57.0/18.6 mm), North Sea, Skagerrak, S. LarvikG-101 Ku (58◦ 26.35′ N 10◦ 40.6′ E),
164.3-179.3 m, 05.vi.1986, RV “Gauss”; SMF 19774, 1 ovig. female (TL/CL, 64.0/20.6 mm),
North Sea, N. Skagen, Skagerrak, Alk87-10VV (58◦ 03.11′ N 09◦ 40.06′ E), 320 m, 22.vii.1987,
R/V “Alkor”; SMF 11835, 2 males (TL/CL, 37.0/11.8-39.0/13.2 mm), 1 ovig. female (TL/CL,
90
CRM 013 – Katsushi Sakai
48.0/16.1 mm), 3 females (TL/CL, 47.0/12.1, rostrum broken – 40.0/12.6 mm), Norway,
Nordfjord (61◦ 55.4′ N 05◦ 25.3′ E), 556-578 m, 18.vi.1983, leg. T. Brattegard; SMF 25687,
1 male (TL/CL, 49.0/14.3 mm), North Sea, Skagerrak, S. Arendal, V53-115Ku (58◦ 08.940′ N
08◦ 53.370′ E – 58◦ 08.890′ N 08◦ 54.190′ E), 441-448 m, 25.ii.1987, R/V “Valdivia”; SMF
29464, 1 female (TL/CL, 48.0/15.9 mm), North Sea, Skagerrak, S. Kristiansand G-103 Ku
(57◦ 59.58′ N 7◦ 55.67′ E – 57◦ 58.9′ N 7◦ 53.18′ E), 189.3-226.8 m, 06.vi.1986, R/V “Gauss”;
SMF 29465, 1 male (TL/CL, 45.0/15.7 mm), North Sea, Skagerrak, S. Larvik G-101 Ku
(58◦ 26.35′ N 10◦ 40.04′ E – 58◦ 28.01′ N 10◦ 40.66′ E), 164.3-179.3 m, 05.vi.1986, R/V “Gauss”;
RMNH D 17319, 1 female (TL/CL, 19.0/4.4 mm), Langenuen, S. of Bergen, Norway (60◦ 4′ N
5◦ 20′ E), 570 m, Agassiz trawl, 7.ix.1961.
Diagnosis. — Male Plp1 absent; male Plp2 biramous, endopod and exopod
slender, endopod with appendices interna and masculina; male Plps3-5 biramous, endopods and exopods slender, endopods with appendix interna. Female
Plp1 uniramous, consisting of proximal segment and multiarticulate flagellum;
female Plps2-5 similar and narrow, endopods with appendix interna.
Type locality. — Skagerrak, North Sea, 500 m.
Distribution. — Norway — Kosterfjord (Trybom, 1904), Sorfjord,
Kvinnheradfjord, Osafjord, Alfjord, Fusafjord, and Bomlafjord (Kensley,
1996a), Hjoerund Fjord, Byfjord; Skagerrak (Trybom, 1904; Poulsen, 1940);
80-700 m.
Calocarides laevis (Bouvier, 1915)
Axius laevis Bouvier, 1915a: 182.
Axius (Neaxius) laevis — De Man, 1925d: 4 (list), 9, 11, 13 (key).
Axius laevis — Sakai & De Saint Laurent, 1989: 80.
Eiconaxius laevis — Sakai & De Saint Laurent, 1989, fig. 21.
Remarks. — The present species was once considered a synonym of
Calocarides coronatus (Trybom, 1904) by Sakai & De Saint Laurent (1989:
80), who studied the type specimen of Calocarides laevis. However, it has
turned out through the present reconsideration that those two species are
different. In C. laevis, the telson is distinctly longer than wide, and the gastric
median carina bears three distinct teeth, whereas in C. coronatus the telson is
slightly longer than wide, and the gastric median carina bears two interspaced
spinules.
Type locality. — West Africa, 698 m.
Distribution. — Only known from the type locality.
Calocarides lev (Zarenkov, 1989)
Calocaris lev Zarenkov, 1989: 24, fig. 1.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
91
Calocarides lev — Kensley, 1996a: 54 (list).
Diagnosis. — Rostrum acutely triangular, lateral margins with 4 teeth each,
extending posteriorly onto anterior half of gastric region as lateral carinae.
Gastric region bearing lateral carinae with two sharp spines, median carina
with two spines in anterior part, and another spine submedially on each
side of median carina. A2 scaphocerite strong, reaching distal margin of A2
penultimate segment. P1 chelate and unequal; in larger left cheliped fingers
shorter than palm; in smaller right cheliped carpus and chela granulate on
lateral surfaces, fingers slightly longer than palm. P2 fingers slightly longer
than palm. Female Plp1 uniramous and elongate; female Plp2 biramous,
endopod slender, bearing appendix interna at middle. [Adapted from Zarenkov,
1989: 24, fig. 1.]
Remarks. — Zarenkov’s (1989) fig. 1, m2, shows that the maxilla 2
scaphognathite bears no posterior whip, which was probably misfigured,
because the maxilla 2 scaphognathite with a posterior whip is a distinct feature
of Calocarides.
Type localities. — Gulf of California, Sta. Cr. 1541, 27◦ 02′ .7N 111◦ 22′ .8W,
2000 m; Cr. 1551, 27◦ 00.3N 111◦ 25′ .2W, 2023-2026 m; Sta. 1572, 27◦ 00′ .3N
111◦ 25′ .3W, 2025-2036 m.
Distribution. — Gulf of California; 2000-2036 m.
Calocarides longispinis (McArdle, 1901)
Calastacus longispinis McArdle, 1901: 522; Alcock & McArdle, 1902, pl. 57 fig. 2, 2a;
MacGilchrist, 1905: 239; De Man, 1925d: 8 (partim), 118; Balss, 1925: 209 (list).
Calocaris (Calastacus) longispinis — Borradaile, 1903: 539.
Calocarides longispinis — Sakai & De Saint Laurent, 1989: 79; Kensley, 1996a: 54 (list).
Not: Calocaris (Calastacus) longispinis — Barnard, 1950: 503, fig. 93d-f [= Calocarides
capensis Kensley, 1996a: 55].
Not: Calastacus longispinis — Stebbing, 1910: 367 [= Calocarides capensis Kensley, 1996a:
55].
Not: Calocaris longispinis — Kensley, 1981b: 30 [= Calocarides capensis Kensley, 1996a].
Not: Calocarides longispinis — Macpherson, 1983: 45, fig. 26; Macpherson, 1991: 405
[= Calocarides macphersoni Kensley, 1996a].
Diagnosis. — Rostrum with 5 lateral spines. Eyestalks reaching almost
middle of rostrum. A2 scaphocerite reaching distal third of A2 penultimate
segment. Gastric median carina with 2 spines followed posteriorly by low blunt
tubercle; gastric region with submedian rows of two spines on left and 4 spines
on right. [Adapted from De Man, 1925d: 118.]
Remarks. — Kensley (1996a: 60) compared Calastacus longispinis McArdle, 1901 from the Arabian Sea, with the Calastacus longispinis specimen of
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CRM 013 – Katsushi Sakai
Macpherson (1983) from Namibia, and he considered the latter a misidentification, so that Macpherson’s (1983) C. longispinis was renamed Calocarides
macphersoni Kensley, 1996a.
Type locality. — Arabian Sea, “Investigator” Sta. 279, 300 fath. (549 m).
Distribution. — Arabian Sea (McArdle, 1901); Gulf of Oman (MacGilchrist,
1905); 549-1280 m.
Calocarides macphersoni Kensley, 1996
(fig. 15)
Calocarides longispinis — Macpherson, 1983: 45, fig. 26; Macpherson, 1991: 405. [Not
McArdle, 1901.]
Calocarides macphersoni Kensley, 1996a: 54 (list), 58, fig. 3.
Material examined. — USNM 243561, holoytype, male (TL/CL, 74.0/24.0 mm), South
Atlantic Ocean, Namibia, NW of Swakopmund, 20◦ 55′ S 12◦ 23′ E, 475-480 m, 9.ix.1980, R/V
“Benguela II”, Sta. 66, det. Stebbing, 1910 as Calastacus longispinis; USNM 243562, paratype,
male (TL/CL, 54.0/21.0 mm), same data as holotype, det. Kensley, 1996a as Calocarides
macphersoni.
Diagnosis. — Rostrum (fig. 15) sharply triangular, with four lateral teeth;
gastric region bearing median carina with a single tooth and hepatic tubercle,
running from rostrum to cervical groove, submedian carinae poorly defined,
with 2-3 teeth, and lateral carinae with 5 teeth including supraocular spine. A2
scaphocerite reaching end of A2 penultimate segment. Mxp3 basis with obtuse
subdistal tooth ventrally; ischium unarmed; merus with three strong ventral
spines, carpus with small ventrodistal spine. In larger cheliped, merus roughly
denticulate on dorsal and ventral margins; carpus evenly tuberculate on lateral
and mesial surfaces, and dorsal and ventral margins. Palm about 1.5 times as
long as fingers, entire surface evenly and fairly densely tuberculate, fingers
widely gaping proximally and meeting distally. In smaller cheliped, merus
roughly denticulate on dorsal and ventral margins; carpus densely tuberculate;
palm about as long as fingers and tuberculate, bearing scanty tubercles on distal
margin; cutting edges of fixed finger and dactylus slightly concave proximally,
bearing small, regularly spaced teeth medially to distally.
Abdominal pleuron 1 narrow and rounded ventrally, pleura 2-6 broadly
rounded ventrally, pleura 3-6 with small spine anteroventrally. Male Plp1
absent; male Plp2 biramous, endopod and exopod slender, endopod with
appendices interna and masculina; male Plps3-5 biramous, endopods and
exopods slender, endopods with appendix interna. Telson bearing proximal
lobe with spine followed by 3-4 spinules on lateral margins, and median
spine on rounded posterior margin; dorsal surface with a pair of spinules
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93
Fig. 15. Calocarides macphersoni Kensley, 1996. Carapace, dorsal view. USNM 243561,
holoytype, male (TL/CL, 74.0/24.0 mm), South Atlantic Ocean, Namibia, N.W. of Swakopmund, 20◦ 55′ S 12◦ 23′ E, 475-480 m, 9.ix.1980, R/V “Benguela II”, Sta. 66. Scale 1 mm.
centrally, and a row of 3-4 spinules at each posterolateral angle. Uropodal
endopod bearing a single posterolateral spine, and dorsal carina with about
4 spinules. Uropodal exopod bearing 2-3 distal spinules on lateral margin, a
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CRM 013 – Katsushi Sakai
single movable spine at posterolateral angle, and a transverse suture with 1315 spinules.
Type locality. — Off Namibia, 20◦ 55′ S 12◦ 23′ E, 475-480 m.
Distribution. — Off Namibia (Macpherson, 1983; Kensley, 1996a); 300480 m.
Calocarides okhotskensis sp. nov.
Calastacus quinqueseriatus — Kobjakova, 1937: 142, pl. 2 fig. 8; Komai, 2000a: 229.
Material examined. — ZIASP, 1 specimen, holotype, Sea of Okhotsk, 1150 m, Sta. 229,
07.viii.1932, leg. P.V. Ushakov:
“At one deep water station in the Sea of Okhotsk, there was a chance to find
a peculiar representative of the Pacific fauna identified as Calastacus quinqueseriatus [= Calocarides okhotskensis]. Unfortunately, this single specimen
was somewhat damaged, which makes me unable to make a full comparison
with Rathbun’s description. It was possible to find some differences from the
typical form, although those differences are not essential and probably fall under intraspecific variability. They are shown as follows: (1) the carapace length
is shorter than the length of 5 abdominal somites (according to Rathbun it is
equal to the length of 5 abdominal somites); (2) the rostrum is slightly shorter
and reaches the mid-length of the 2nd segment of the antennular peduncle;
(3) intermediate ridge beween the lateral and median ones is armed with two
spines (2-3 spines according to Rathbun) and begins at the level of the 3rd
spine of the dorsal ridge (in typical specimen this ridge is more extended forward as Rathbun’s figures indicate); (4) fourth pereiopods are nearly equal but
the left is markedly stronger than the right one (according to Rathbun, they
are not equal); (5) chela and all segments of second pair of legs without setae
on margins (according to Rathbun, all margins are beset with setae); (6) the
lateral carinae on the telson extending toward posterior margin and terminated
with spinules are missing in our specimen. Further records of this species from
the Sea of Okhotsk will make it possible to clear up its taxonomic status. The
occurrence of the species in the Sea of Okhotsk is no doubt related to the penetration of the warm Pacific waters, and is, therefore, of considerable interest”.
[Translated from the Russian by V. Spiridonov.]
Remarks. — Kobjakova (1937) described Calastacus quinqueseriatus Rathbun, 1902, based on a specimen from the Sea of Okhotsk, 1150 m. However, it
was later identified as Calocarides quinqueseriatus by Sakai & De Saint Laurent (1989), because her species does not belong to Calastacus; the species is
different in characteristics from the type species of Calastacus, C. stilirostris
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95
Faxon, 1893, though she adopted the generic name from Rathbun’s original
description. The type specimen from the Sea of Okhotsk is inaccessible at
present, which prevents me from making its characteristics clear, but it seems
that Kobjakova’s Calocarides quinqueseriatus is geographically and morphologically different from Rathbun’s Calocarides quinqueseriatus from eastern
North America, 288-700 m. In Kobjakova’s species the palm of the larger cheliped is about 1.5 times as long as wide, and the dactylus is curved downward in the distal half of the cutting edge, and slightly shorter than the palm
(Kobjakova, 1937, pl. 2 fig. 8). In Rathbun’s species, however, the palm of
the larger cheliped is more than twice as long as wide, and the dactylus is
straight, and distinctly shorter than the palm. Therefore, Kobjakova’s Calocarides quinqueseriatus is here defined as a new species of Calocarides, Calocarides okhotskensis.
Type locality. — Sea of Okhotsk, 1150 m.
Distribution. — Only known from the type locality.
Etymology. — The new species is named okhotskensis after the type
locality, the Sea of Okhotsk, with the Latin suffix “ensis”, indicating a
geographic region. The name is an adjective agreeing in gender with the
generic name.
Calocarides quinqueseriatus (Rathbun, 1902)
Calastacus quinqueseriatus Rathbun, 1902: 887; Rathbun, 1904: 151, fig. 92; Schmitt, 1921:
113, fig. 76; Balss, 1925: 209; Hart, 1982: 50, fig. 11.
Calocaris (Calastacus) quinqueseriata — Borradaile, 1903: 539; De Man, 1925d: 8 (list), 118
(key).
Calocarides quinqueseriatus — Sakai & De Saint Laurent, 1989: 79; Hendrickx, 1995: 390
(list), 393, 396, figs.; Kensley, 1996a: 54 (list), 61, figs. 4, 5; Hendrickx et al., 2005: 171.
Material examined. — USNM 25240, holotype, male (TL/CL, 74.0/25.0 mm), San Luis
Obispo Bay, California, 5.iv.1890, “Albatross” Sta. 3196, 200 fms (365.8 m), det. M. Rathbun.
Diagnosis. — Male Plp1 absent; male Plp2 biramous, endopod with appendices interna and masculina; male Plps3-5 endopods with appendix interna.
Remarks. — Kensley (1996a: 54) remained uncertain of including Calastacus quinqueseriatus Rathbun, 1902 in the present genus Calocarides. However, it is safely included in Calocarides by the forms of male pleopods 1
and 2.
Type locality. — Off San Luis Obispo Bay, South California, “Albatross”
Sta. 3196, 360 m.
Distribution. — Southern California — off San Luis Obispo Bay, Point
Sur, San Simeon Bay, Point Conception, Santa Barbara Channel, and Anacapa
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CRM 013 – Katsushi Sakai
Island (Rathbun, 1902, 1904), off Point Sur, off San Simeon Bay, off San Luis
Obispo Bay, off Point Conception, Santa Barbara Channel, off Anacapa Island,
off San Nicolas Island (Schmitt, 1921), California coast and Oil Island (Sakai
& De Saint Laurent, 1989), off San Luis Obispo Bay, Santa Barbara Channel,
San Miguel Island, off Point Conception, off Point Sur, off San Simeon Bay,
off Anacapa Island, off San Nicolas Island, off San Diego, off Santa Rosa
Island, N.E. of Point Loma, San Cristobal Bay, and mouth of Columbia River
(Kensley, 1996a); 292-2013 m.
Calocarides rostriserratus (Andrade & Báez, 1977)
Calastacus rostriserratus Andrade & Báez, 1977: 65, fig. 1.
Calocarides rostriserratus — Kensley, 1996a: 54 (list), 64, fig. 6.
Type locality. — Pichidangui, Chile, 32◦ 08′ S 71◦ 50′ W, 320-400 m.
Distribution. — Gulf of Panama (Kensley, 1996a); Peru — off Tumbes
(Kensley, 1996a); Chile — Pichidangui (Andrade & Báez, 1977); 320-825 m.
Calocarides rudolfi (Zarenkov, 1989)
Calocaris rudolfi Zarenkov, 1989: 27, fig. 3.
Calocarides rudolfi — Kensley, 1996a: 54 (list).
Diagnosis. — Rostrum triangular with acute tip, lateral margins with 34 teeth, extending posteriorly onto middle of gastric region. Gastric region
bearing lateral carinae with supraorbital spine, followed posteriorly by 24 sharp spines; median carina unarmed; and submedian carinae with 34 spines. Postcervical carina absent. A2 scaphocerite strong, reaching near
distal margin of A2 penultimate segment. Mxp3 merus with 1-2 small and
3 sharp spines on ventral margin; merus with subterminal spine on ventral
margin. P1 chelate and unequal; in larger (left) cheliped merus with two
sharp spines on ventral margin, and two sharp spines followed distally by
another two spinules on dorsal margin; carpus and chela granulate on lateral
surface, fingers shorter than palm. P2 fingers longer than palm. Plp1 unknown.
Plp2 biramous, endopod slender, bearing appendices interna and masculina at
middle part. Telson almost square, lateral margins unarmed, posterior margin
slightly convex with median tooth, and dorsal surface with a pair of spinules
proximally and two pairs of spines in distal half. Uropodal endopod with 5
spines on median carina. Uropodal exopod with serrated transverse suture.
[Adapted from Zarenkov, 1989: 29, fig. 3.]
Remarks. — Zarenkov (1989) described a male specimen without referring
to male Plp1, but he showed in his fig. 3 that the Plp2 endopod bears
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appendices interna and masculina mesiomedially, which suggests that this
male specimen is not included in Calocaris, but in Calocarides. Accordingly,
the present species is reclassified under the genus Calocarides as C. rudolfi.
Type locality. — Off Angola, 490-510 m.
Distribution. — Only known from the type locality.
Calocarides soyoi (Yokoya, 1933)
Axius soyoi Yokoya, 1933: 49, fig. 25; Horikoshi et al., 1982: 30, 33, 38, 39, 120, 145, 168.
Axiopsis (Axiopsis) soyoi — Sakai, 1987a: 303 (list).
Calocarides soyoi — Sakai & De Saint Laurent, 1989: 4 (list), 83, tab. 3; Kensley & Komai,
1992: 81, fig. 1, tabs. 1-2; Kensley, 1996a: 54 (list), 66.
Axiopsis soyoi — Takeda, 1997: 250 (list).
Material examined. — SMF-32430, 1 male (TL/CL, 76,0/26.0 mm), Japan, off Heta, Suruga
Bay, 380-400 m, bottom trawl, 6.ii.1992, leg. H. Mukai & K. Sakai; SMF-32431, 1 male
(TL/CL, 62.0/20.5 mm), Japan, off Heta, Suruga Bay, 200-400 m, bottom trawl, 29.iii.1983, leg.
H. Mukai; SMF-32432, 1 ovig. female (TL/CL, 60.0/19.5 mm, missing both chelipeds), Japan,
off Heta, Suruga Bay, 200-400 m, bottom trawl, 29.iii.1983, leg. H. Mukai; SMF-32433, 1 male
(TL/CL, 61.0/21.7 mm), Japan, off Heta, Suruga Bay, 200-400 m, bottom trawl, 29.iii.1983,
leg. H. Mukai; SMF-32434, 1 male (TL/CL, 59.0/19.1 mm, missing larger cheliped), Japan, off
Heta, Suruga Bay, 200-400 m, bottom trawl, 29.iii.1983, leg. H. Mukai; SMF-32435, 1 male
(TL/CL, 60.0/20.1 mm, missing larger cheliped), Japan, off Heta, Suruga Bay, 200-400 m,
bottom trawl, 29.iii.1983, leg. H. Mukai; SMF-32436, 1 male (TL/CL, 82.0/25.2 mm, missing
larger cheliped) Japan, off Heta, Suruga Bay, 200-400 m, bottom trawl, 29.iii.1983, leg. H.
Mukai; SMF-32437, 1 male (TL/CL, 73.0/22.8 mm), Japan, N. Japan, Iwate Pref., off Otsuchi
(39◦ 22.99′ N 142◦ 03.19′ E – 39◦ 22.82′ N 142◦ 03.14′ E), 118-120 m, 1 m, dredge, R/V “Tanseimaru”, KT-93-15 Sta. 01 CB, 12-16.xi.1993, leg. K. Sakai & S. Ohta; SMF-32438, 1 male
(TL/CL, 79.0/25.5 mm), Japan, Iwate, off Otsuchi, 128 m (70 fathoms), gill-net, leg. H. Mukai;
SMF-32439, 1 male (TL/CL, 71.0/22.5 mm), Japan, off Heta, Suruga Bay, 380-400 m, bottom
trawl, 6.ii.1992, leg. H. Mukai & K. Sakai.
Remarks. — Calocarides soyoi from Japan is similar to Calocarides chani
from Taiwan in the carapace spination and the shape of the first pereiopods,
as Kensley et al. (2000: 210) mentioned, but still differs in several features. In
C. soyoi the carapace length is 12.0-18.0 mm; the chela of the larger cheliped
is three times as long as wide; the P1 merus is denticulate both on the distal
half of the dorsal margin and on the whole ventral margin, the P1 carpus has
numerous tubercles on the lateral surface; and the uropodal endopod bears
six spines on the longitudinal carina. In C. chani the carapace length is 10.212.2 mm; the chela of the larger cheliped is 2.2 times as long as wide; the P1
merus bears a single spine on the dorsal margin and about three spines on the
ventral margin; the P1 carpus has tubercles along the dorsal margin; and the
uropodal endopod bears 5 spines on the longitudinal carina.
Type locality. — South of Shioya-zaki, Fukushima Pref., Japan, 142 m.
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CRM 013 – Katsushi Sakai
Distribution. — Suruga Bay (Takeda, 1997); 120-270 m; Inubo-zaki, off
Shioya-zaki, Kii Strait, off Satsuma, and Tsushima (Yokoya, 1933), off
Fukushima Prefecture and off Hachinohe (Kensley & Komai, 1992; Kensley,
1996a).
Calocarides tenuicornis (De Man, 1905)
Axiopsis tenuicornis De Man, 1905: 596; Balss, 1925: 209.
Axiopsis (Axiopsis) tenuicornis — De Man, 1925d: 6 (list), 69 (key), 84, pl. 7 fig. 14-14l.
?Calocarides tenuicornis — Sakai & De Saint Laurent, 1989: 83.
Material examined. — ZMA Crust. De. 202874, lectotype, 1 female (dissected, CL, 7.4 mm,
abdomen somite 5-tail-fan missing).
Remarks. — It is confirmed by examining the female holotype of the
present species, Axiopsis tenuicornis De Man, 1905, and referring to De Man’s
figures (De Man, 1925d, pl. 7 fig. 14-14l), that this female type specimen is
what he really figured as holotype, though it is separated into several parts
(cephalothorax, thoracic somites and abdomen, and first chelipeds).
The present species C. tenuicornis is closely similar to C. vigilus Sakai,
1992c, because in both species the A2 scaphocerite is elongate and protruded
forward; P1 carpus and chela are not tuberculate on the lateral surfaces; the
P1 fingers are longer than the palm in the smaller cheliped; and the lateral
carinae of the gastric region bear distinct spines. However, the present species
is distinguished from C. vigilus, because in C. tenuicornis the gastric region
bears a median carina with 4 spinules, and submedian rows of 3-4 spinules,
whereas in C. vigilus the gastric region bears a spinulate median carina and
submedian rows of 7-8 spinules.
Type locality. — Java Sea, 7◦ 46′ S 114◦ 30.5′ E, “Siboga” Sta. 5, 330 m.
Distribution. — Indonesia — Java Sea; 330 m.
Calocarides vigilus Sakai, 1992
Calocarides vigila Sakai, 1992a: 176, figs. 17, 18; Kensley, 1996a: 54 (list).
Diagnosis. — Male Plp1 absent, male Plp2 biramous, endopod with appendices interna and masculina (Sakai, 1992a: 177, fig. 18G).
Type locality. — Hawke Bay, New Zealand, 39◦ 36′ S 177◦ 42′ E, 126 m.
Distribution. — New Zealand — Hawke Bay and Cook Strait; 60-146 m.
Genus Coelhocalaxius gen. nov.
Diagnosis. — Rostrum elongate and narrow, with 4-5 sharp spines on each
lateral margin, extending posteriorly onto gastric region as lateral carinae with
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two distinct buttressed teeth. Gastric region with median carina bearing tooth
and hepatic tubercle, originating from base or from near tip of rostrum to
cervical groove, and submedian rows of teeth; postcervical carina present,
posteriorly bearing intestinal ridge. Anterolateral margin of carapace with
triangular tooth. Cervical groove distinct at full length. Eyestalks elongate but
slightly shorter than rostrum; cornea dilated. A1 peduncle overreaching rostral
tip. A2 scaphocerite protruded straight forward. P1 subequal; palm armed
with 4 interspaced strong teeth on dorsal margin; fingers longer than palm.
P2 fingers shorter than palm. Abdominal somites 2-5 with broadened dorsal
carina and lateral carina between tergum and pleuron, pleura sharply pointed
ventrally. Male Plps1-2 unknown. Female Plp1 uniramous and bisegmented,
consisting of proximal segment and flagellum; female Plps2-5 endopods with
appendix interna. Telson longer than wide, bearing proximal lobe with spine
on lateral margin, but no median tooth on rounded posterior margin. Uropodal
exopod with transverse suture.
Remarks. — Calaxius spinosus Coelho, 1973 is similar to the type species
of Calaxius, C. acutirostris Sakai & De Saint Laurent, 1989, in the pleura,
which are sharply pointed ventrally. But it is distinguished from Calaxius
at the generic level, because in C. spinosus the rostrum is elongate and
narrow; the postcervical carina is present, posteriorly bearing an intestinal
ridge; the eyestalks are elongate but slightly shorter than the rostrum; the
cornea is dilated; abdominal somites 2-5 bear a broadened dorsal carina and
a longitudinal lateral carina between tergum and pleuron. In C. acutirostris
in contrast, the rostrum is elongate and triangular; the postcervical carina
is absent; the eyestalks are stout and short; the cornea is pigmented; and
abdominal somites 2-5 bear a longitudinal lateral carina between tergum and
pleuron, but are smooth dorsally. Therefore, a new genus, Coelhocalaxius gen.
nov., is erected for Calaxius spinosus Coelho, 1973.
Type species. — Calaxius spinosus Coelho, 1973, by present designation
and monotypy. The gender of the generic name, Coelhocalaxius, is masculine.
Species included. — Coelhocalaxius spinosus (Coelho, 1973).
Etymology. — The new generic name Coelhocalaxius is dedicated to Dr.
P.A. Coelho, Universidade Federal de Pernambuco, PE, Brazil, who described
the present species under the genus Calaxius.
Coelhocalaxius spinosus (Coelho, 1973)
(fig. 16)
Axiopsis sp. C, Coelho, 1971: 231.
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CRM 013 – Katsushi Sakai
Fig. 16. Coelhocalaxius spinosus (Coelho, 1973). A, carapace, dorsal view; B, carapace, lateral
view; C, cheliped on left side, lateral view; D, abdomen and tail-fan, dorsal view; E, abdomen
and tail-fan, lateral view; F, telson and uropod on right side. A-F, DOUFPe 1870, holotype,
1 female (TL/CL, 19.0/7.4 mm), W. [of] Amapá, Brazil. Scales 1 mm.
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Calocaris (Calastacus) sp., Coelho & Ramos, 1973: 161.
Calastacus spinosus Coelho, 1973: 345; Coelho & Ramos-Porto, 1983: 82, fig. 9; Rodrigues et
al., 1998: 382; Melo, 1999: 330, figs. 223-224.
Calocaris (Calastacus) oxypleura Williams, 1974a: 457, figs. 11-18. [Type locality: Straits of
Florida W. of Riding Rocks, 25◦ 15′ N 79◦ 13′ W, 365 m.]
Axiopsis oxypleura — Rabalais et al., 1981: 98.
Calaxius oxypleura — Sakai & De Saint Laurent, 1989: 86; Kensley & Hickman, 2001: 487;
Clark et al., 2007: 72.
Calaxius spinosus — Coelho et al., 2007: 4.
Material examined. — DOUFPe 1870, holotype, female (TL/CL, 19.0/7.4 mm), 3◦ 37′ S
50◦ 01′ W, Amapá, Brazil, 75, 82-94 m; RMNH D 41760, 1 male (TL/CL, 33.0/11.7 mm),
Pillsbury Sta. p-1367, 18◦ 21.4′ N 69◦ 08.7′ W, 165 m, ottertrawl, 9.vii.1971, USNM Acc. No.
381702; RMNH D 41761, 1 male (TL/CL, 33.0/11.7 mm), Pillsbury Sta. p-756, 11◦ 33.1′ N
69◦ 12.6′ W 11◦ 34.5′ N, 69◦ 11′ W, 9-21 fms (= 16.5-38.4 m), 10′ ottertrawl, 27.vii.1968, USNM
Acc. No. 381702, det. B. Kensley; USNM 151211, paratype, 1 male (TL/CL, 50.0/19.0 mm),
Quita Sueno Bank, Nicaragua, Caribbean Sea, 14◦ 12.00′ N 81◦ 57.30′ W, 146 m, trawl,
02.xii.1967, det. A.B. Williams.
Diagnosis. — Integument minutely (i.e., microscopically) granulated. Rostrum slenderly elongate, bearing 4-5 spines on each lateral margin, extending onto gastric region; anterolateral margin of carapace with triangular tooth.
Gastric region bearing lateral rows of two interspaced buttressed teeth, submedian rows of three interspaced teeth, and median carina with anterior spine
and hepatic tubercle. Postcervical carina present, posteriorly bearing intestinal
ridge (fig. 16A, B).
Eyestalks elongate and slightly shorter than rostrum; cornea dilated. A1 peduncle overreaching rostral tip. A2 penultimate segment extending beyond A1
peduncle; segment 1 with short distoventral spine; segment 2 with strong dorsolateral spine; scaphocerite slender and curved slightly downward, exceeding
distodorsal spine of segment 2; segment 3 with small mesioventral spine; distal
segment half length of penultimate segment. Mxp3 pediform, densely setigerous ventrally and mesially, a few long setae on carpus and merus laterally;
coxa with ventral spine; ischium with two spinules on ventral margin, and
along mesial margin a prominent crest with about 16 teeth, terminating in enlarged distal spine curved mesially; merus with three ventral spines increasing
in size and acuteness distally; carpus with small distoventral spine laterally;
dactylus unarmed.
P1 strong and asymmetrical, chelae elongate; in larger cheliped chela
subequal in length to palm, whereas in smaller cheliped chela longer than
palm. P2 chelate. Ps3-4 simple, propodi elongate, bearing transverse rows
of spinules ventrolaterally. P5 subchelate, propodus with small, rounded
distoventral protrusion.
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CRM 013 – Katsushi Sakai
Abdominal somites uniformly arched dorsally, abdominal somite 1 short
and drawn ventrally to acuminate tip, bearing low longitudinal carina laterally;
abdominal somites 2-5 with broadened dorsal carina (fig. 16D), and acutely
angulate posteroventrally (Coelho & Ramos-Porto, 1973, fig. 9), bearing
longitunal carina laterally; abdominal somites 3-5 with a single spine in middle
on anteroventral margin; abdominal somite 6 triangular ventrally, with tiny
blunt spine anterior to tip of pleuron.
Male Plp1 absent; male Plp2 biramous, endopod bearing appendices interna
and masculina mesiomedially; Plps3-5 biramous and slender, endopods with
appendix interna. Female Plp1 uniramous and bisegmented; Plps2-5 slender
and biramous, endopods bisegmented, bearing appendix interna at proximal
margin of distal segment.
Telson longer than wide, bearing proximal lobe with spine followed by
another two spines on lateral margin, two pairs of spines on dorsal surface
(fig. 16F), but no median spine on rounded posterior margin; dorsal surface
with two pairs of spines in proximal half around fairly broad longitudinal
median sulcus, and transverse row of three spines each at posterolateral
corner. Uropodal endopod as long as telson, bearing thick longitudinal median
ridge with 4 nearly equidistant spines; lateral margin with 4 spines including
strong one at posterolateral angle; uropodal exopod with transverse suture
with spinules; two thick longitudinal carinae centrally on dorsal surface;
lateral margin with 1-4 unevenly spaced spines, distalmost flanked mesially
by submarginal movable spine.
Remarks. — Calastacus spinosus Coelho, 1973 from Amapá, Brazil is
closely similar to Calocaris (Calastacus) oxypleura Williams, 1974 from
Florida, though the type specimen of C. spinosus is a female and small in size,
measuring 19.0 mm in total length. In both species the A2 scaphocerite is slender and curved slightly downward, exceeding the distodorsal spine of segment
2 and reaching the distal third of the A2 penultimate segment; the eyestalks are
elongate and reach nearly to the distal end of the rostrum; abdominal somites
2-5 bear broadened dorsal carinae (Wiliams, 1974, fig. 11). Therefore, Calocaris (Calastacus) oxypleura is safely synonymized with Calastacus spinosus,
which is here reclassified under the new genus Coelhocalaxius.
Type locality. — Amapá, Brazil, 75, 82-94 m.
Distribution. — U.S.A. — Florida (Williams, 1974a), Texas (Rabalais et al.,
1981); Nicaragua — Quita Sueño Bank (Williams, 1974a); 65-365 m; Amapá,
Brazil, 75, 82-94 m (Coelho, 1973).
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
103
Genus Colemanaxius gen. nov.
Diagnosis. — Rostrum trispinose and carinate on lateral margins, with
distinct supraorbital spines proximally, extending posteriorly onto gastric region as lateral carinae with tooth. Gastric region convex, bearing median carina with three tubercles, and submedian carinae with two tubercles. Cervical groove distinct. Eyestalks subglobose; cornea pigmented. A2 scaphocerite
horn-shaped. P1 unequal; in larger cheliped merus denticulate on ventral margin, carpus and chela smooth on lateral surface; palm with irregularly denticulate double carina on dorsal margin, and stout distodorsal tooth. P2 fingers
longer than palm. Ps3-4 propodi with transverse rows of spines on ventrolateral surface. Abdominal pleura smooth on surface, and convex to rounded
ventrally. Male Plps1-2 unknown. Female Plp1 unknown; female Plps2-5 biramous, endopods with appendix interna. Telson convergent posteriorly, lacking
posteromedian tooth. Uropodal exopod with transverse suture. Gonochoristic.
Remarks. — Axiopsis (Axiopsis) pitatucensis De Man, 1925b is not included
in Axiopsis, because A. (A.) pitatucensis is different from the type species of
Axiopsis. It differs from Axiopsis serratifrons (A. Milne-Edwards, 1873) in the
features of rostrum, gastric region, chela of larger cheliped, and telson, so that
A. (A.) pitatucensis is reclassified here under the new genus Colemanaxius as
C. pitatucensis (De Man, 1925b).
Kensley (2003: 368) described from specimens originating from Agana,
Guam, along with a corresponding figure, that “the strong setation on the upper
surface of the chelae of pereopod 1, the strong spine on the lateral palmar
surface at the base of the dactylus, and the articulated submarginal spine at
the first lateral spine of the telson are the features that distinguished this genus
from all the other axiid genera described”, and defined his specimens amiss
as Manaxius pitatucensis (De Man, 1925b), without examining De Man’s type
specimen of Axiopsis (Axiopsis) pitatucensis De Man, 1925b, and made it the
type species of his new genus Manaxius. However, Kensley’s pitatucensis from
Agana, Guam is clearly different from De Man’s pitatucensis. In Kensley’s
pitatucensis the rostrum is acutely triangular, the lateral margins are armed
with 2-4 teeth and a supraorbital tooth, the gastric region bears lateral carinae
with 1-5 buttressed teeth, and median and submedian carinae with denticles;
the anterolateral margin of the carapace is unarmed; and abdominal pleura 26 are triangular ventrally. By contrast, in De Man’s pitatucensis, the rostrum
is trispinose, bearing no teeth but a distinct supraorbital spine proximally on
the lateral margins, extending posteriorly onto the gastric region as lateral
carinae with a buttressed tooth, and reaching to near the cervical groove
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CRM 013 – Katsushi Sakai
(fig. 17C; De Man, 1925b, fig. 5); the gastric region bears a median carina
with three tubercles including the hepatic one, and medially interrupted
submedian carinae with two tubercles; the anterolateral margin of the carapace
is armed with a small spine; and abdominal pleura 2-6 are broadly or simply
rounded ventrally. Those differences between De Man’s Axiopsis (Axiopsis)
pitatucensis (= Colemanaxius pitatucensis (DeMan, 1925b)) and Kensley’s
Manaxius pitatucensis suggest, that they are utterly different, not only at the
species level but also at the generic level. Therefore, Kensley’s Manaxius
pitatucensis is a junior homonym of Colemanaxius pitatucensis (De Man,
1925b), and the generic name Manaxius is invalid. His species is to be renamed
herein (see below) as Manaxius aganaensis nom. nov., as the type species of
the genus Manaxius Kensley (sensu nov.).
Type species. — Axiopsis (Axiopsis) pitatucensis De Man, 1925b, by
present designation, and monotypy. The gender of the new generic name,
Colemanaxius, is masculine.
Species included. — Colemanaxius pitatucensis (De Man, 1925b).
Etymology. — The new genus is named Colemanaxius in honour of Dr.
Oliver Coleman, Curator of the Zoologisches Museum zu Berlin, who kindly
gave me a chance to study the precious type specimen for this revision, with
Axius, the generic name of the type genus of the Axiidae added.
Colemanaxius pitatucensis (De Man, 1925)
(fig. 17)
Axiopsis (Axiopsis) pitatucensis De Man, 1925b: 133, fig. 5-5f; De Man, 1925d: 6 (list), 69
(key).
Spongiaxius pitatucensis — Sakai & De Saint Laurent, 1989: 44.
Not: Manaxius pitatucensis — Kensley, 2003: 368, figs. 3, 4 [invalid: not Axiopsis (Axiopsis)
pitatucensis De Man, 1925b].
Material examined. — ZMB 14439, holoype, female (TL/CL, 14.0/4.9 mm), Pitatuki, Buka
Island, German Papua New Guinea, 23.x.1909, leg. H. Schoede.
Diagnosis of holotype. — Species of small size (fig. 17A). Rostrum
(fig. 17B; De Man, 1925b, fig. 5) trispinose, bearing no teeth but distinct supraorbital spine proximally on lateral margins, which extend posteriorly onto gastric region as lateral carinae with buttressed tooth, and reaching to near cervical
groove; gastric region bearing median carina with three tubercles including hepatic one, and medially interrupted submedian carinae also with two tubercles.
Anterolateral margin of carapace with a small spine. A2 scaphocerite short and
horn-shaped. Cervical groove distinct. In larger cheliped (fig. 17C); ischium
armed with two low, thick proximal teeth, a strong distal one on the ventral
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105
Fig. 17. Colemanaxius pitatucensis (De Man, 1925). A, whole body, lateral view; B, carapace,
dorsal view; C, cheliped on left side, lateral view; D, P3, lateral view; E, P4, lateral view;
F, abdominal somite 6, telson, and uropod on left side, dorsal view. A-F, ZMB 14439, holotype,
1 female (TL/CL, 14.0/4.9 mm), Pitatuki, Buka Island, German Papua New Guinea. Scales
1 mm.
106
CRM 013 – Katsushi Sakai
margin, and a sharp subdistal tooth on the dorsal margin; merus unarmed on
dorsal margin, but armed with 5 thick teeth including distal triangular tooth
on ventral margin, and protrusion at ventrodistal angle; carpus short, bearing
obtuse subdistal tooth on dorsal margin; palm 1.5 times as long as wide, bearing three buttressed teeth mesiodorsally, and some small, irregularly arranged
teeth and a distinct distal tooth dorsolaterally; distal margin largely convex in
ventral half; fixed finger bearing triangular median tooth, followed distally by
two low continuous teeth on cutting edge. P2 chelate; fingers longer than palm.
Ps3-4 propodi (fig. 17D, E) with a row of transverse setae. Abdominal pleuron 1 acutely triangular ventrally, abdominal pleura 2-6 (fig. 17A) convex or
rounded ventrally. Female Plp1 unknown; Plps2-5 biramous, endopods with
appendix interna. Telson (fig. 17F) convergent toward rounded posterior margin without median spine. Uropodal endopod bearing spines on lateral margin,
including distinct spine at posterolateral corner, and median row of 4 spines on
dorsal surface; uropodal exopod bearing 4 spines on lateral margin, sublateral
carina with spines, and serrated oblique suture with distal flap dorsally. Female
genital pores confirmed on P3 coxa.
Remarks. — De Man (1925b) stated that the Plp1 is absent in the female
type specimen, but the present re-examination of the female type specimen
(ZMB 14439, holoype) suggests that the female Plp1 is simply missing,
because it is usually present in females and consists of a proximal segment
and a multiarticulate flagellum. The female Plp2 endopod is provided with an
appendix interna as De Man described (1925b: 137, fig. 5d), but Plps3-5 are
very much shrunk in the present condition, which makes it difficult to examine
them precisely.
Type locality. — Pitatuki, Buka Island, Papua New Guinea.
Distribution. — Papua New Guinea — Buka Island (De Man, 1925b; Sakai
& De Saint Laurent, 1989).
Genus Dorphinaxius Sakai & De Saint Laurent, 1989
Dorphinaxius Sakai & De Saint Laurent, 1989: 33; Poore, 1994: 97 (key); Davie, 2002: 452;
Poore, 2004: 175; Poore & Collins, 2009: 253.
Diagnosis. — Rostrum distinct and triangular, with supraorbital spines at
base; lateral margins unarmed, posteriorly extending shortly onto gastric region as lateral carinae; gastric region steeply elevated from base of rostrum,
bearing median carina with tooth halfway along, and submedian carinae with a
blunt tooth at anterior end. Anterolateral margin of carapace unarmed. Cervical
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107
groove present at full length. Eyestalks small; cornea pigmented. A2 scaphocerite comma-shaped and directed anteriorly. P1 unequal; palms of both chelipeds much longer than dactyli. Pleurobranchs present on Ps2-4. Abdominal
pleura smooth on surface, and truncate or rounded on ventral margins. Male
Plp1 absent; male Plp2 biramous and slender, endopod with appendices interna and masculina; male Plps3-5 endopods with appendix interna. Female
Plp1 uniramous, consisting of proximal segment and multiarticulate flagellum; female Plps2-5 biramous and slender, endopods with appendix interna.
Telson longer than wide, bearing no median spine on rounded posterior margin. Uropodal exopod with transverse suture.
Remarks. — This genus Dorphinaxius Sakai & De Saint Laurent, 1989 is
similar to the genus Ralumcaris gen. nov. in that the male Plp1 is absent, and
the gastric region is steeply elevated from the base of the rostrum. However,
they differ from each other, because in Dorphinaxius the gastric region is
provided with median, submedian, and lateral carinae, and the male Plp2
endopod bears appendices interna and masculina, whereas in Ralumcaris the
gastric region bears only a median carina, and the male Plp2 endopod bears an
appendix masculina, but no appendix interna.
Type species. — Axiopsis (Paraxiopsis) appendiculis Poore & Griffin, 1979,
by original designation and monotypy. Gender of the new generic name,
Dorphinaxius, masculine.
Species included. — Dorphinaxius kermadecensis (Chilton, 1911).
Dorphinaxius kermadecensis (Chilton, 1911)
(fig. 18)
Iconaxiopsis kermadecensis Chilton, 1911: 550, figs. 1, 2; Balss, 1925: 211.
Axius (Eiconaxius) kermadecensis — De Man, 1925d: 4 (list), 15 (key).
Eiconaxius kermadecensis — Sakai & De Saint Laurent, 1989: 23.
Axiopsis (Paraxiopsis) appendiculis Poore & Griffin, 1979: 224, fig. 1; Poore & Collins, 2009:
254, figs. 21-23. [Type locality. — Shellharbour, New South Wales, Australia.]
Dorphinaxius appendiculis — Sakai & De Saint Laurent, 1989: 34; Sakai, 1994: 200 (key).
Dorphinaxius kermadecensis — Davie, 2002: 452; Poore, 2004: 175, figs. 45e, f, 46c.
Material examined. — AM P 9359, holotype, ovig. female (TL/CL, 39.0/13.7 mm), Shellharbour, New South Wales, tide marks; AM P 9063, 1 ovig. female (TL/CL, 36.0/11.9 mm);
AM P 6365, 1 male (TL/CL, 26.0/9.6 mm); AM P 18557, 1 male (TL/CL, 51.0/18.5 mm),
Long Reef, Collaroy, N.S.W., 30.xii.1971, leg. D. Hoese, det. G. Poore; AM P 24674, 1 male
(TL/CL, 25.0/8.2 mm), Mince Waters near Graftor, N.S.W., intertidal rock platform, i.1967, leg.
G. Biddle; AM P 8693, 1 female (TL/CL, 36.0/12.9 mm), Bottle & Glass Rocks, Port Jackson,
Australia.
108
CRM 013 – Katsushi Sakai
Fig. 18. Dorphinaxius kermadecensis (Chilton, 1911). Carapace, dorsal view. AM P 9359,
holotype, ovig. female (TL/CL, 39.0/13.7 mm), Shellharbour, New South Wales, tide marks.
Scale 1 mm.
Diagnosis. — Rostrum (fig. 18) distinct and triangular in dorsal view,
bearing distinct supraorbital tooth proximally. Male Plp1 absent; male Plp2
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
109
biramous, endopod and exopod slender, endopod with appendices interna and
masculina; Plps3-5 biramous, endopods and exopods slender, endopods with
appendix interna. Female Plp1 uniramous, consisting of proximal segment
and multiarticulate flagellum; Plps2-5 biramous and slender, endopods with
appendix interna.
Remarks. — Axiopsis (Paraxiopsis) appendiculis Poore & Griffin, 1979 was
synonymized with Iconaxiopsis kermadecensis Chilton, 1911 by Davie (2002:
452) and Poore (2004: 175). However, the former species had been designated
as the type species of the genus Dorphinaxius Sakai & De Saint Laurent, 1989,
so that as a result the valid name for the present species now is Dorphinaxius
kermadecensis (Chilton, 1911).
Type locality. — Meyer Is. and Kermadec Islands, littoral.
Distribution. — Meyer Is. and Kermadec Islands (Chilton, 1911); Australia — Grafton, Collaroy, Port Jackson, Bass Point, and Shellharbour, N.S.W.
(Poore & Griffin, 1979); intertidal to 5 m; Emily Bay, Norfolk Is. (Poore &
Collins, 2009).
Genus Eutrichocheles Wood-Mason, 1876
Eutrichocheles Wood-Mason, 1876b: 264; Chopra, 1933: 277; Balss, 1957: 1580; Sakai & De
Saint Laurent, 1989: 51; Sakai, 1992b: 215; Poore, 1994: 97 (key); Sakai, 1994: 185, 200
(key); Ngoc-Ho, 1998: 364; Ngoc-Ho et al., 2005: 199, figs. 1-3.
Diagnosis — Rostrum triangular with pointed apex, bearing denticles on lateral margins extending posteriorly onto gastric region as lateral carinae; gastric
region convex, bearing smooth median carina with hepatic tubercle, submedian rows of denticles, and lateral carinae in anterior half with supraorbital
tooth followed posteriorly by another tooth; anterolateral margin of carapace
usually with spine. Cervical groove distinct. Postcervical carina occasionally
present posteriorly. Posterior margin of carapace with a pair of posterolateral lobes. Eyestalks subglobose; cornea pigmented. A1 segment 1 slightly
inflated, segments 2-3 short. A2 scaphocerite short, and bi- or trifurcate. Mxp3
pediform, exopod slender; ischium with strong, dentate crest on inner surface;
merus dentate on mesial margin. P1 chelate and asymmetrical. P2 chelate. Ps34 simple, and propodi with transverse rows of spines laterally. P5 simple or
subchelate, propodus with setae on distoventral margin. Epipod present. Pleurobranchs absent. Abdominal somites 1-5 with or without longitudinal carina
between tergum and pleuron, pleuron 1 narrow and protruded ventrally, pleuron 2 broad and truncate ventrally, pleura 3-6 triangular or rounded ventrally,
pleura 2-5 usually without anteroventral spine. Male Plp1 uniramous and unsegmented, as stubby protrusion or slightly curved truncate protrusion; male
110
CRM 013 – Katsushi Sakai
Plp2 biramous, endopod bearing appendix masculina with distal setae, but no
appendix interna; male Plps3-5 biramous, endopods without appendix interna.
Female Plp1 consisting of a proximal segment and a multiarticulate flagellum;
female Plps2-5 biramous, and endopods without appendix interna. Telson subsquare, bearing proximal lobe with spine on lateral margin, and median spine
on posterior margin; dorsal surface with paired, divergent carinae. Uropodal
endopod bearing lateral spines and median carina with spines on dorsal surface. Uropodal exopod with transverse suture with spinules. Gonochoristic,
but some species are hermaphroditic.
Remarks. — Eutrichocheles is closely similar to Paraxiopsis, as evident
from those features of the two genera that the male Plp2 is biramous, and
the endopod bears an appendix masculina with distal setae but no appendix
interna; male Plps3-5 are biramous, and the endopods lack an appendix
interna; female Plp1 consists of a proximal segment and a multiarticulate
flagellum; female Plps2-5 are biramous, and the endopods bear no appendix
interna. However, Eutrichocheles is different from Paraxiopsis, because in the
type species of Eutrichocheles, E. modestus (Herbst, 1796), the male Plp1 is
uniramous and unsegmented, as a stubby protrusion (fig. 20E), or as a slightly
curved truncate protrusion (fig. 20D; Ngoc-Ho et al., 2005: 206), and the
gastric region bears submedian rows of teeth. In contrast, in the type species
of Paraxiopsis, P. brocki De Man, 1888, the male Plp1 is a small, thornlike protrusion (WAM 34-75 from Garden Island, Western Australia, Poore &
Griffin, 1979, fig. 3i), or a slender bisegmented one (fig. 25C, AM P 6824
from Rail Pier, Port Darwin, Northern Territory, Australia), though it is absent
in a young specimen (SMF 20362, from Bora Bora, French Polynesia; Sakai,
1992b: 216), as Kensley (1996c) mentioned in the definition of the genus
Paraxiopsis, and the gastric region bears smooth submedian carinae.
It is here considered that the differences in the features of the postcervical
carina, P1 chela, and the median notch of the telson, which are found in the
species included in Eutrichocheles, result from the difference in size of the
specimens; in E. modestus, the postcervical carina is distinct posteriorly in
larger specimens, but indistinct in a smaller mature specimen (NHM 1879.32,
1 male, CL, 31.3 mm, from Pinang, Malaysia).
Type species. — Cancer modestus Herbst, 1796, designated by WoodMason, 1876b, and by monotypy. The gender of the generic name, Eutrichocheles, is masculine.
Species included. — Eutrichocheles austrinus Sakai, 1994; E. defensus
(Rathbun, 1901); E. foveolatus (Kensley, 1996c); E. granulimanus (Kensley,
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
111
1996c); E. modestus (Herbst, 1796) [= E. crosnieri Ngoc-Ho, 1998]; E.
spinipleurus (Kensley, 1996c); E. tuamotu Ngoc-Ho, 1998.
K EY
TO THE
S PECIES
OF THE
G ENUS EUTRICHOCHELES
1 – Pleuron 2 triangular ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Pleuron 2 truncate or rounded ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 – P1-2 fingers longer than palm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. modestus
– P1-2 fingers shorter than palm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. spinipleurus
3 – P1 smaller cheliped with fingers longer than palm; P1 chela densely setose and tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. granulimanus
– P1 smaller cheliped with fingers shorter than palm; P1 chela poorly setose . . . . . . . . . . 4
4 – Carapace obviously pitted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. foveolatus
– Carapace smooth or setose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Carapace, pereiopods, and abdomen covered with setae . . . . . . . . . . . . . . . . . E. austrinus
– Carapace smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – Pleura 3-5 triangular with obtuse tip ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . E. tuamotu
– Pleura 3-5 rounded ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. defensus
Eutrichocheles austrinus (Sakai, 1994)
Eutrichocheles austrinus Sakai, 1994: 185, 200 (key), figs. 6, 7.
Paraxiopsis austrinus — Kensley, 2003, table 2; Animal Diversity Web, Univ. Michigan; Poore
& Collins, 2009: 266, fig. 29.
Remarks. — In Eutrichocheles austrinus Sakai, 1994, the submedian carinae of the gastric region are denticulate as in E. modestus.
Type locality. — Darwin, Australia.
Distribution. — Australia — Darwin (Sakai, 1994).
Eutrichocheles defensus (Rathbun, 1901)
Axius defensus Rathbun, 1901: 95, text-fig. 17.
?Axiopsis defensus — Borradaile, 1903: 539.
Axiopsis (Paraxiopsis) defensus — De Man, 1925d: 7 (list), 71 (key); Schmitt, 1935a: 191,
fig. 51; Coelho & Ramos-Porto, 1985: 76, fig. 5.
Eutrichocheles defensus — Sakai & De Saint Laurent, 1989: 3, 52, 101.
Paraxiopsis defensus — Kensley, 1996c: 715, figs. 3, 4; Melo, 1999: 320, figs. 215-216.
Material examined. — USNM 23780, holotype, female (TL/CL, 30.0/10.8 mm), off Boca
Prieta, Puerto Rico, 8.1/2 fms, leg. R/V “Fish Hawk”, Sta. 6075, US EC, det. M. Rathbun.
Remarks. — Paraxiopsis defensus is to be included in Eutrichocheles,
because the gastric region bears no smooth submedian carinae, but submedian
rows of teeth.
112
CRM 013 – Katsushi Sakai
Type locality. — Off Boca Prieta, Puerto Rico, 15.5 m (= 8.5 fms), coral
and sand.
Distribution. — Puerto Rico — off Boca Prieta (Rathbun, 1901; Kensley,
1996c); Dominican Republic — Barahona Harbour (Kensley, 1996c); 15 m.
Eutrichocheles foveolatus (Kensley, 1996)
Paraxiopsis foveolata Kensley, 1996c: 718, fig. 5.
Type locality. — Florida shelf, Gulf of Mexico, 25◦ 45′ 42′′ N 83◦ 11′ 04′′ W,
53.7 m.
Distribution. — Gulf of Mexico — Florida shelf (Kensley, 1996c); 53.7 m,
soft sand and mud bottom.
Eutrichocheles granulimanus (Kensley, 1996)
Paraxiopsis granulimana Kensley, 1996c: 722, fig. 8.
Remarks. — In the present species, the submedian carina of the gastric
region is denticulate; abdominal somites 2-6 are smooth, and the pleura are
triangular ventrally.
Type locality. — S.W. Florida shelf, Gulf of Mexico, 53.7 m.
Distribution. — Gulf of Mexico — S.W. Florida shelf (Kensley, 1996c); off
Trinidad (Kensley, 1996c); 53-95 m.
Eutrichocheles modestus (Herbst, 1796)
(figs. 19, 20, 21A-G)
Homarus scaber Weber, 1795: 94 [nomen nudum].
Cancer modestus Herbst, 1796: 173, pl. 43 fig. 2; Latreille, 1816: 35; Stebbing, 1893: 206;
Balss, 1933: 87.
Astacus scaber Fabricius, 1798: 407; Bosc, 1801-1802: 62; Latreille, 1802-1803: 241; Latreille,
1806: 52; Desmarest, 1830: 41; H. Milne Edwards, 1837a: 335; H. Milne Edwards, 1837b:
383; Broderip, 1838: 274; Zimmsen, 1964: 652. [Type locality: Indian Ocean.]
Astacus Scaber — Lamarck, 1818: 217.
Axius biserratus Von Martens, 1869: 612; Nobili, 1903: 12; De Saint Laurent, 1979 (in De Saint
Laurent & Le Loeuff, 1979): 32. [Type locality: Indian Ocean.]
Astacus modestus — Wood-Mason, 1876a: 231; Wood-Mason,1876b: 264; Bouvier, 1917: 14;
De Saint Laurent, 1979: 32.
Eutrichocheles modestus — Wood-Mason, 1876a: 231; Wood-Mason, 1876b: 264; Miers, 1880:
380; Stebbing, 1893: 206; De Man, 1916: 96; Bouvier, 1917: 14; Balss, 1933: 87; Chopra,
1933: 277, text-fig. 1, pl. 6; Balasubrahmanyan & Jacob, 1961: 830, fig. 1; Kensley, 1961c:
711; De Saint Laurent & Le Loeuff, 1979: 32; Holthuis, 1986: 249; Sakai & De Saint
Laurent, 1989: 51, 102; Ngoc-Ho, 1998: 374, fig. 6C-F; Komai, 2000b: 345; Xuân & NgocHo, 2006: 1153-1166, figs. 1-4.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
113
Fig. 19. Cancer modestus Herbst, 1796 (= Eutrichocheles modestus (Herbst, 1796)), copy of
original figure of Herbst (1796, pl. 43 fig. 2).
?Axiopsis biserratus — Borradaile, 1903: 538.
Axiopsis (Paraxiopsis) biserrata — De Man, 1925b: 138, fig. 6; De Man, 1925d: 7 (list), 71
(key).
Eutrichocheles crosnieri Ngoc-Ho, 1998: 371, figs. 4-6; Komai, 2000b: 345; Ngoc-Ho et al.,
2005: 206. [Type locality: Ha-tien, Vietnam, 30 m.]
Eutrichocheles Holthuis, 1946: 71.
Paraxiopsis crosnieri — Kensley, 2003, table 2.
Material examined. — RMNH D 38354, 1 female, Trat Province Koa Chang, Gulf of
Thailand, Thailand, 1980, don. S. Chaitiamvong; ZMB 3226, 1 male (TL/CL, 64.0/21.4 mm,
lacking P1 on left side) [Axius biserrata Von Martens, 1869, lectotype (fig. 21A)]; ZMB
x3226, 1 male (TL/CL, 55.0/20.3 mm, lacking both P1) [Axius biserrata Von Martens, 1869,
paralectotype], Malacca, leg. S. Baumgarten; NHM 1879.32 (fig. 20A, B), 1 male (CL 31.3 mm,
damaged), Pinang, Malaysia, Indopacific side, from Indian Museum, det. De Saint Laurent,
vi.1977; MNHN Th 1267, 1 male (TL/CL, 74.0/29.5 mm), Singapore fishmarket, i.1914,
leg. Falshwn; MNHN Th 1499, 1 female (TL/CL, 82.0/33.0 mm, scaphocerite with 1 spine)
Cangio District, about 10 km north of Vung Tan (Cap Sta. Jacques), 10 m, bottom sand and
mud, iv.2000, leg. N. Van Xuan; MNHN Th 1500, 3 males (TL/CL, 76.0/32.0-78.0/32.0 mm,
scaphocerite with 1-3 spines), Vietnam, Cangio District, about 21 km north of Vung Tan (Cap
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Fig. 20. Eutrichocheles modestus (Herbst, 1796). A, whole body, lateral view; B, C, abdomen,
lateral view; D, E, male Plp1. A, ZMB3226, 1 male (TL/CL, 64.0/21.4 mm), Malacca
[= Eutrichocheles biserrata (Von Martens, 1869), lectotype]; B, MNHN Th 1267, male
(TL/CL, 74.0/29.5 mm), Singapore fishmarket; C, MNHN Th 1314, female (TL/CL, 83.5/
33 mm [= Eutrichocheles crosnieri Ngoc-Ho, 1998, holotype]; D, E, NHM 1879.32, 1 male
(CL, 31.3 mm, damaged), Pinang, Malaysia, Indo-Pacific side. Scales A, 0.8 mm; B, C, 1 mm;
D, E, 0.5 mm.
Sta. Jacques), 6-8 m, iv-viii.2000, leg. N. Van Xuan; MNHN Th 1314, 1 female (TL/CL,
83.5/33 mm) [= Eutrichocheles crosnieri Ngoc-Ho, 1998, holotype], Ha-tien, Vietnam, 30 m.
Diagnosis. — Rostrum (figs. 19, 21A-B) acutely triangular and denticulate
on lateral margins, with supraorbital spine at rostral base, extending posteriorly
onto anterior half of gastric region as lateral carinae with tooth. Gastric
region bearing smooth median carina with hepatic tubercle, and submedian
carinae with 6 spines. Postcervical carina present in posterior fourth in larger
specimen. A2 scaphocerite (fig. 21A) short and bi- or trifurcate distally. P1
chelate and unequal (figs. 19, 20A); in smaller (fig. 21C) and larger chelipeds
of both sexes, cutting edge of fixed finger denticulate, proximally bearing
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115
Fig. 21. Eutrichocheles modestus (Herbst, 1796) and Levantocaris hornungae Galil & Clark,
1993. A, H, gastric region, dorsal view; B, same, lateral view; C, smaller cheliped, on
right side, lateral view; D, P2, lateral view; E, distal parts of P3, lateral view; F, distal
parts of P4, lateral view; G, tail-fan, dorsal view. A-G, Eutrichocheles modestus, ZMB3226,
1 male (TL/CL, 64.0/21.4 mm), Malacca [= Eutrichocheles biserrata (Von Martens, 1869),
lectotype]; H, Levantocaris hornungae, NHM 1192: 608, holotype, hermaphroditic male
(TL/CL, 28.0/10.9 mm), north Haifa, Israel, 33◦ 00′ N 34◦ 35′ E, 1400 m. Scales 1 mm.
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CRM 013 – Katsushi Sakai
distinct conical tooth; P2 (fig. 21D) chelate, fingers longer than palm. P3
(fig. 21E) simple. P4 simple; propodus with spinules laterally and ventrally.
P5 (fig. 21F) subchelate, propodus unarmed laterally, bearing protrusion with
distal spinules distoventrally. Abdominal somites 1-5 (figs. 19, 20A, B) with
longitudinal carina between tergum and pleuron. Male Plp1 uniramous and
unsegmented, as a stubby protrusion or a slightly curved truncate protrusion
(fig. 20D, E); male Ppl 2 biramous, endopod bearing appendix masculina with
distal setae, but no appendix interna; Plps3-5 biramous, endopods lacking
appendix interna. Telson (fig. 21G) longer than wide, bearing proximal lobe
with spine followed by 2-4 spines on lateral margin, median notch with or
without spine on posterior margin, and paired divergent carinae with spines on
dorsal surface. Uropodal endopod broad and armed with teeth or unarmed on
lateral margin, and bearing median carina with 4-5 teeth dorsally. Uropodal
exopod broad and armed with teeth or unarmed on lateral margin, and bearing
smooth median carina and transverse suture with spinules on dorsal surface.
Remarks. — The type specimen of Eutrichocheles modestus (Herbst, 1796)
is missing (Balss, 1933: 87), but the figure (fig. 19; Herbst, 1796, pl. 43
fig. 2) was made clear enough to distinguish the species from the other species
included in Eutrichocheles, though the figure shows no proximal conical tooth
on the cutting edge of the fixed finter of the larger cheliped on the right side.
Herbst defined the specimen as Cancer modestus, but it was later described as
Astacus scaber (Flusskrebs) by Fabricius (1798).
Wood-Mason (1876b) designated this species as Eutrichocheles modestus,
though he thought that it might be a species of Thaumastochelidae, Thaumastocheles zaleucus (Thompson, 1873), or a species of the Laomediidae, Jaxea
nocturna Nardo, 1847 (= Calliaxis adriatica Heller, 1863). Chopra (1933) reported, by following Balss’ (1933) suggestion, on the type specimen of Eutrichocheles modestus that it should be a species of the Axiidae, though the
specimen she defined as a female by mentioning that “The pleopods of the first
pair in the single female example that I have seen are very much reduced, and
are represented by short, uniramous, stumpy structures (Chopra, 1933: 280)”,
has turned out to be a male, because the feature mentioned above by her is a
decisive feature of the males of Eutrichocheles modestus.
De Man (1925b: 138) described two types of Axiopsis (Paraxiopsis) biserrata (= Eutrichocheles biserrata), the lectotype (fig. 20A) and a paralectotype, and mentioned that those two types are probably males. It is confirmed through the present examination that those types of E. biserrata are
males, because the genital pores are located on the P5 coxae; Plp1 bears a
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117
movable thorn-shaped ramus; and the Plp2 endopod bears an appendix masculina, but no appendix interna, as shown in De Man’s (1925b) description of
Plp2, that “der Endopod der 2. Pleoden keine mit Cincinnuli versehene Stylamblys, sondern eine Appendix masculina und die folgenden Pleopoden keine
Stylamblys” [= Plp2 without appendix interna, but with appendix masculina,
and Plps3-5 without appendix interna]. The lectotype and paralectotype have
the following characteristics in common with each other: the scaphocerite is
bifurcate, the carapace bears a longitudinal postcervical carina; pleuron 2 is
unarmed on the anteroventral margin, but pleura 3, 4, and 5 bear an anteroventral spinule. However, they differ from each other in that in the lectotype the
anterolateral margin of the carapace bears a distinct antennal spine and below
it an indistinct spinule as in E. crosnieri Ngoc-Ho, 1998, while in the paralectotype the anterolateral margin of the carapace bears an antennal spine, but
below it a second, obscure protrusion. Those characteristics are observed also
in E. modestus in the Paris collections under MNHN Th 1499 and Th 1500.
Ngoc-Ho (1998: 376) compared a large-sized male specimen of E. modestus
and the female holotype of E. crosnieri Ngoc-Ho, 1998 from Ha-tien, Vietnam,
and later Ngoc-Ho et al. (2005: 206) compared again two female specimens
of E. modestus and the female holotype of E. crosnieri, and summed up
the differences between the females of the two species, E. modestus and E.
crosnieri, in the same way they had done between the male of E. modestus and
the female of E. crosnieri in 1998, describing as follows. (1) The anterolateral
border of the carapace often bears a spinule and a tubercle in E. modestus (with
two spinules in E. crosnieri); (2) the postcervical region of the carapace has a
short median dorsal carina in the posterior quarter in E. modestus (absent in E.
crosnieri); (3) the abdominal pleura 3-5 carry an anterior spine in E. modestus
(unarmed in E. crosnieri); (4) the antennal acicle (= scaphocerite) is bifid in
E. modestus (trifid in E. crosnieri); (5) the 4th thoracic sternite is armed with a
lateral spine in E. modestus (unarmed in E. crosnieri). They (Ngoc-Ho et al.,
2005) reached the same conclusion as Ngoc-Ho in 1998, mentioning that “the
above distinguishing characters permit an unambiguous separation of females
of E. modestus from E. crosnieri and confirm the validity of the latter species”,
without making any reference to the differences in the features between the
males and females of the two species.
However, those differences are to be considered allowable as variations
in the species and are not to be regarded as species differences between E.
modestus and E. crosnieri, because of the small number of specimens she
compared: only one female holotype of E. crosnieri (MNHN-Th 1314, TL/CL
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CRM 013 – Katsushi Sakai
83.5/33 mm), and a male (MNHN Th 1267, TL/CL 74.0/29.5 mm) and two
females (ZRC 1999.0088, TL/CL 84.0/33.0 mm; RMNH D 38354 TL/CL
70.5/30.0 mm) of E. modestus. Accordingly, it is too hasty a conclusion that
those two species are different from each other and that E. crosnieri would
thus be valid.
Besides, it is also found through the present comparison of the female
holotype of E. crosnieri (MNHN-Th 1314) with the male specimen of E.
modestus (MNHN Th 1267), that the postcervical carina of the carapace is
present in both E. crosnieri and E. modestus; the armature of the ventral
margins of the abdominal pleura is variable, and it has also turned out by
examining the specimens (MNHN Th 1499, 1 female, and MNHN Th 1500,
3 males) that in the female (MNHN Th 1499) pleura 3 and 4 bear a spine on
the anteroventral margin, though in the female (MNHN Th 1314, E. crosnieri,
holotype) pleura 2 and 6 are indeed unarmed (fig. 20C). In two of the three
males (MNHN Th 1500) pleura 4 and 5 bear a spine on the anteroventral
margin, but pleura 2, 3, and 6 are unarmed, and in the other male (MNHN
Th 1500) pleura 3, 4, and 5 bear a spine on the anteroventral margin, though
pleura 2 and 6 are unarmed. Those differences in the armature of the pleural
ventral margin are, however, not to be regarded as specific characteristics, but
as intraspecific variation of the species.
The present observation shows that the A2 scaphocerite is simple, bi-, or
trifurcate in E. modestus, though Ngoc-Ho reported that it is bifurcate in E.
modestus (trifurcate in E. crosnieri) (Ngoc-Ho, 2005).
Those variations found in the males of E. modestus are also observed in
the males of E. biserrata (figs. 20A, 21A-G). Furthermore, her comparison of
E. crosnieri with E. modestus is of little value, because she discussed those
two species regardless of the sexual differences of the specimens, namely the
specimen of E. crosnieri she used for comparison with the male specimen
of E. modestus is a female, and such a comparison is not essential for the
separation of one species from the other, considering that spinulation often
varies by size and by sex. So, it is safely concluded without further discussion
that E. crosnieri is synonymous with E. modestus.
Type locality. — Indian Ocean.
Distribution. — India — Porto Novo; Malacca (Von Martens, 1869; De
Man, 1925b); Malaysia — Pinang (Ngoc-Ho, 1998, as E. crosnieri); Singapore
(Nobili, 1903; Ngoc-Ho, 1998, as E. crosnieri); Vietnam — Ha-tien (Ngoc-Ho,
1998 as E. crosnieri).
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Eutrichocheles spinipleurus (Kensley, 1996)
Paraxiopsis spinipleura Kensley, 1996c: 726, figs. 10-12.
Remarks. — In Eutrichocheles spinipleurus (Kensley, 1996c), the submedian carinae of the gastric region are denticulate; abdominal somites 2-5 are
provided with a longitudinal carina between tergum and pleuron; and pleura
2-6 are triangular ventrally as in E. modestus.
Type locality. — Sta. Thomas, U.S. Virgin Islands.
Distribution. — Florida — Looe Key reef (Kensley, 1996); Belize — Carrie
Bow Cay (Kensley, 1996c); U.S. Virgin Islands — Sta. Thomas (Kensley,
1996c).
Eutrichocheles tuamotu Ngoc-Ho, 1998
Eutrichocheles tuamotu Ngoc-Ho, 1998: 368, figs. 2, 3.
Paraxiopsis tuamotu — Kensley, 2003, table 2.
Material examined. — MNHN, 1 male (TL/CL, 19.0/7.8 mm), ?Bali, Indonesia, leg. A.
Anker.
Remarks. — In Eutrichocheles tuamotu Ngoc-Ho, 1998, the submedian
gastric carinae are denticulate as in E. modestus; Ps1-2 fingers are shorter than
the palm; the abdominal somites are smooth; and pleuron 2 is truncate distally
as in Paraxiopsis brocki De Man, 1888b; however, pleura 3-5 are triangular
ventrally as in E. modestus.
Type locality. — Mururoa, Tuamotu Archipelago, 21◦ 53.12′ S 139◦ 2.62′ W,
140 m.
Distribution. — Only known from the type locality, and only by the material
mentioned here.
Genus Guyanacaris gen. nov.
Diagnosis. — Carapace tuberculate. Rostrum narrowly protruded, bearing
3-4 spinules on the lateral margins. Gastric region bearing lateral, submedian,
and median carinae each with spinules; postcervical carina present. Eyestalks
subglobose and large; cornea pigmented. A2 scaphocerite elongate. P1 subequal in size and dissimilar in shape. Ps3-4 propodi with a ventrolateral row
of spinules. Abdominal pleuron 2 broadly truncate ventrally and abdominal
pleura 3-5 rounded ventrally; terga and pleura separated by weak longitudinal
carinae with tubercles. Male Plp1 uniramous and bisegmented, distal segment
shorter than proximal; male Plp2 biramous and foliaceous, endopod with appendices interna and masculina, appendix interna slightly longer than appendix
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CRM 013 – Katsushi Sakai
masculina; male Plps3-5 biramous, endopods with appendix interna. Telson
longer than wide, bearing proximal lobe on lateral margin, and median spine
on convex posterior margin. Uropodal endopod broad, bearing median carina
with teeth on dorsal surface. Uropodal exopod broadened, bearing transverse
suture with denticles.
Remarks. — The two species, Axiopsis (Axiopsis) caespitosa Squires, 1979
from San Juan del Sur, Colombia, Pacific Ocean, and Calocaris (Calastacus)
hirsutimana Boesch & Smalley, 1972 from off British Guiana, which were
included in the genus Acanthaxius by Sakai & De Saint Laurent (1989),
are reclassified under the new genus Guyanacaris gen. nov., because the
two species are distinctly different from the type species of Acanthaxius,
A. pilocheirus (Sakai, 1987a). In the latter, the A2 scaphocerite is directed
outward; and male Plp1 is absent; whereas in the former two species the
A2 scaphocerite is strongly elongate and directed forward, and male Plp1 is
uniramous and bisegmented.
Type species. — Calocaris (Calastacus) hirsutimana Boesch & Smalley,
1972, by present designation. The gender of the new name, Guyanacaris, is
feminine.
Species included. — Guyanacaris caespitosa (Squires, 1979); G. hirsutimana (Boesch & Smalley, 1972).
Etymology. — The generic name Guyanacaris is derived from the type
locality, British Guyana.
Guyanacaris caespitosa (Squires, 1979)
Axiopsis (Axiopsis) caespitosa Squires, 1979: 1584, figs. 1-3, tabs. 1-2.
Acanthaxius caespitosa — Sakai & De Saint Laurent, 1989: 73; Lemaitre & León, 1992: 44;
Hendrickx, 1995: 390 (list), 393, 394, figs.; Hendrickx et al., 2005: 171.
Material examined. — USNM 172265, holotype female (TL/CL, 90.0/29.0 mm), San Juan
del Sur, Colombia, Pacific Ocean, 90 m, 16.ii.1971, leg. Choco, det. H.J. Squires.
Diagnosis. — Rostrum narrowly protruded forward, bearing 4 spines laterally. Eyestalks stout and subglobose, and longer than rostrum; eyes wider
than stalk, cornea pigmented. A2 scaphocerite present as a long, spine-like
process. Gastric region with 5 longitudinal carinae each with a few or several spines directed forward; postcervical carina distinct. Abdominal pleuron
2 truncate ventrally, and abdominal pleura 3-5 rounded ventrally, pleura 1-5
with thin border and a double row of denticles along ventral margin. Female
Plp1 uniramous; female Plps2-5 endopods with appendix interna. [Adapted
from Squires, 1979: 1584, figs. 1-3.]
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121
Remarks. — This species was described only from the female holotype, so
male Plps1-2 are unknown. However, it is safely included in the present new
genus Guyanacaris, because the rostrum is narrowly protruded, the eyestalks
are thick, overreaching the rostrum, the carapace bears a postcervical carina,
and the abdominal pleura, and the tail-fan are similar to those of the type
species of the genus Guyanacaris, G. hirsutimana.
Type locality. — Off Rio San Juan del Sur, Pacific coast of Colombia.
Distribution. — Colombia — off Rio San Juan del Sur (Squires, 1979).
Guyanacaris hirsutimana (Boesch & Smalley, 1972)
(fig. 22A-B)
Calocaris (Calastacus) hirsutimana Boesch & Smalley, 1972: 45, figs. 1-9.
Acanthaxius hirsutimanus — Sakai & De Saint Laurent, 1989: 73; Blanco-Rambla, 1995: 62,
figs. 11, 12; Heard et al., 2007: 18, fig. 10.
Axiopsis hirsutimana — Vargas & Cortés, 1999a: 882.
Material examined. — Carribean Sea: USNM 137428, holotype, male (TL/CL, 75.0/
27.0 mm), off British Guiana, Surinam, 6◦ 50′ N 54◦ 47′ W, leg. U.S. Fish and Wildlife Service,
28 fathoms (51.2 m), leg. R/V “Coquette”, Sta. 422. — Pacific side: RMNH D 31563, 1 male
(TL/CL, 116.0/40.8 mm), Caleta la Cruz, Peru, 180 m, v.1952, leg. M. del Solar; SMF 31827,
1 female (TL/CL, 135.0/48.0 mm), Golfo Dulce, SW Rio Coto, Costa Rica, Pacific side, 8◦ 28′ N
83◦ 10.3′ W, 12.3-23 m, xii.1993-ii.1994, R/V “Victor Hensen”.
Diagnosis. — Carapace (fig. 22A, B) granulate, rostrum slender and acutely
triangular, lateral margins with 3-4 spinules; gastric region bearing distinct median carina with 4 anterior spinules, submedian rows of 3-4 irregularly sized
spinules, and lateral carinae with 5 anteriorly directed spinules; postcervical
carina present. Eyestalks stout and subglobose, exceeding rostrum; cornea
pigmented. A2 scaphocerite elongate, extending to distal fourth of A2 segment 3. P1 dissimilar in shape; carpi and chelae densely covered with thick
setae. Abdominal pleura 1-3 denticulate on ventral margin; pleura 2-3 bearing
longitudinal carina with tubercles between tergum and pleuron. Male Plp1 is
small, uniramous, and bisegmented, distal segment leaf-like and shorter than
proximal one. Male Plp2 biramous and foliaceous, endopod with appendices
interna and masculina, appendix interna slightly longer than appendix masculina; male Plps3-5 endopods with appendix interna. Female Plp1 uniramous
and leaf-like; female Plps2-5 biramous, endopods with appendix interna. Telson subsquare, bearing median spine on convex posterior margin. Uropodal
exopod with transverse suture.
Type locality. — Off Guyana, 6◦ 50′ N 54◦ 47′ W, 50 m.
Distribution. — Atlantic side: Florida; Guyana (Abele & Kim, 1986;
Boesch & Smalley, 1972); Costa Rica (Vargas & Corté, 1999a); Mississippi
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CRM 013 – Katsushi Sakai
Fig. 22. Guyanacaris hirsutimana (Boesch & Smalley, 1972) and Manaxius euophthalmus
(De Man, 1905). A, whole body with left P1-4, lateral view; B, carapace, dorsal view;
C, larger cheliped, lateral side; D, smaller cheliped, lateral view; E, abdomen, lateral view.
A, B, Guyanacaris hirsutimana, USNM 137428, holotype, male (TL/CL, 75.0/27.0 mm), off
British Guiana, 6◦ 50′ N 54◦ 47′ W, 28 fathoms (51.2 m); C, D, E, Manaxius euophthalmus, ZMA
Crust. De. 240080, holotype, male (TL/CL, 220./8.6 mm), north of the island of Batjan, 0◦ 11′ S
127◦ 25′ E, “Siboga” Sta. 139, 4.viii.1899-1900, mud, stones, and coral. Scale A, 5 mm; B,
4 mm; C, D, E, 1 mm.
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(Boesch & Smalley, 1972); 11-50 m. Pacific side: Caleta la Cruz, Peru, 180 m;
Herradura, Costa Rica, Pacific side.
Genus Heterocalaxius gen. nov.
Diagnosis. — Rostrum stoutly triangular with obtuse and upturned apex,
bearing two similar lateral spines and supraorbital one at base; lateral margins extending posteriorly onto gastric region as lateral carinae with obtuse
median spine; gastric region bearing median carina in anterior half as weak
crest with worn tubercle near posterior end, and submedian carinae originating
from anteriormost of two slightly offset submedian teeth, becoming ill-defined
posteriorly; postcervical carina absent. Anterolateral margin of carapace unarmed. Cervical groove remarkable with broadened V-shape posteromedially.
Eyestalks subglobose; cornea pigmented. A2 scaphocerite short and protruded
forward. P1 similar in shape; carpus and palm tuberculate on dorsal margin,
beset with tufts of setae on lateral surface; fingers of both chelipeds shorter
than palms. P2 fingers slightly longer than palm. Abdominal pleuron 1 narrow
and protruded ventrally; pleuron 2 broadened ventrally, bearing small, acute
tooth at posteroventral angle; pleura 3-5 truncate ventrally, bearing small acute
tooth on anteroventral margin and another one on posteroventral margin; pleuron 6 triangular ventrally, bearing small acute tooth on anteroventral tip and
broad triangular flange at posteroventral angle. Male Plp1 absent; male Plp2
biramous, bearing thick setae; endopod with appendix interna, but without appendix masculina; male Plps3-5 biramous, bearing thick setae, endopods with
appendix interna. Tail-fan with thick plumose setae; telson distinctly longer
than basal width, bearing at proximal fourth a fixed spine followed posteriorly
by serration on lateral margin, and tapering towards rounded posterior margin.
Uropodal endopod broad, bearing posterolateral spine; uropodal exopod with
transverse suture with few denticles.
Remarks. — Calaxius carneyi was described by Felder & Kensley (2004)
as one of the species belonging to the genus Calaxius. However, it has turned
out through the present examination of the holotype that their species is not
to be included in Calaxius, because C. carneyi is different in morphological
features from the type species of Calaxius, C. acutirostris Sakai & De Saint
Laurent, 1989. In C. carneyi the male Plp2 endopod bears an appendix interna,
but no appendix masculina; the carapace lacks a postcervical carina; the
scaphocerite is short; in the larger cheliped the carpus and palm are weakly
tuberculate on the dorsal margin, in the smaller cheliped the carpus and palm
are more distinctly tuberculate than in the larger cheliped; the fingers of both
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CRM 013 – Katsushi Sakai
chelipeds are shorter than the palms; and the telson is triangular, tapering
posteriorly. In C. acutirostris, on the other hand, the male Plp2 endopod bears
appendices interna and masculina; the carapace bears a postcervical carina; the
scaphocerite is strongly protruded forward; in the larger and smaller chelipeds
the palms are armed with distinct interspaced teeth on the dorsal margins, and
the fingers are longer than the palms; and the telson is rectangular. For those
reasons, C. carneyi is not to be included in Calaxius, and is reclassified here
under the new genus Heterocalaxius, which is established for the species C.
carneyi, as Heterocalaxius carneyi.
Type species. — Calaxius carneyi Felder & Kensley, 2004, by present
designation and monotypy. The gender of the new name, Heterocalaxius, is
masculine.
Species included. — Heterocalaxius carneyi (Felder & Kensley, 2004).
Etymology. — The new genus is named Heterocalaxius, derived from its
heterogenic characteristics, which differentiate the new genus from Calaxius,
by adding a Greek prefix “hetero”, meaning “different”, to the generic name
Calaxius.
Heterocalaxius carneyi (Felder & Kensley, 2004)
Calaxius carneyi Felder & Kensley, 2004: 69, figs. 1, 2; Clark et. al., 2007: 72.
Material examined. — USNM 1009165, holotype, male (TL/CL, 30.0/11.0 mm),
27◦ 46.904′ N 91◦ 30.286′ W, chemosynthetic communities of the Louisiana continental slope,
Gulf of Mexico, 544 m.
Diagnosis. — Male Plp1 absent; Plps2-5 endopods with appendix interna.
Type locality. — 27◦ 46.904′ N 91◦ 30.286′ W, chemosynthetic communities
of the Louisiana continental slope, Gulf of Mexico, 544 m.
Distribution. — Only known from the type locality.
Genus Leonardsaxius gen. nov.
Diagnosis. — Rostrum triangular with upturned apex; lateral margins
armed with teeth, extending posteriorly onto gastric region as lateral carinae.
Gastric region convex, bearing median carina with 1-3 teeth originating from
base of rostrum, and smooth submedian and lateral carinae. Cervical groove
distinct at full length; postcervical carina absent. Anterolateral margin of
carapace unarmed. Eyestalks subglobose, not reaching middle of rostrum. A2
scaphocerite elongate. P1 chelate and unequal; palm of cheliped unarmed on
dorsal margin. Ps3-4 propodi elongate without transverse rows of spines on
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125
ventrolateral margin. Abdominal pleura rounded on ventral margin. Male Plp1
uniramous and bisegmented, distal segment triangular; male Plp2 biramous,
endopod with appendices interna and masculina; male Plps3-5 endopods with
appendix interna. Female Plp1 uniramous and bisegmented, proximal segment
simple and distal one multiarticulate; female Plps2-5 endopods with appendix
interna. Telson longer than wide, bearing median tooth on posterior margin.
Uropodal endopod shorter than telson, bearing strong distal spine on lateral
margin, and median carina with spines on dorsal surface. Uropodal exopod
with transverse suture.
Remarks. — The type species Axiopsis werribee was described as one of
the species included in Axiopsis by Poore & Griffin (1979). However, it is not
included in Axiopsis, because in Axiopsis werribee the male Plp1 is uniramous
and bisegmented, and the gastric region bears a median carina with 1-3 teeth
originating from the base of the rostrum and smooth submedian and lateral
carinae. In the type species of Axiopsis, in contrast, Axiopsis serratifrons
A. Milne-Edwards, 1873, the male Plp1 is absent and the gastric region forms
a horseshoe-shaped prominence, bearing 5 longitudinal rows of denticles. So
Axiopsis werribee is reclassified here under the new genus Leonardsaxius,
which is established for Axiopsis werribee, as L. werribee (Poore & Griffin,
1979).
Type species. — Axiopsis (Axiopsis) werribee Poore & Griffin, 1979, by
present designation. The gender of new name, Leonardsaxius, is masculine.
Species included. — Leonardsaxius amurensis (Kobjakova, 1937); L. spinulicauda (Rathbun, 1902); L. werribee (Poore & Griffin, 1979); and L. sp.
aff. L. werribee Poore & Collins, 2009.
Etymology. — The new genus is named Leonardsaxius, derived from the
type locality of the type species, Sta. Leonards, Port Phillip Bay, Victoria, by
adding Leonards as a prefix to the generic name Axius.
Leonardsaxius amurensis (Kobjakova, 1937)
Axius spinulicauda amurensis Kobjakova, 1937: 142, pl. 2 fig. 9; Kensley, 1996a: 68; Komai,
2000a: 229.
Axiopsis spinulicauda amurensis — Vinogradov, 1950: 223, pl. 24 fig. 102.
Diagnosis. — Carapace and rostrum shorter than or equal in length to 4
abdominal somites; rostrum straight, not curved; gastric submedian carinae
considerably longer than lateral ones, and not straight but curved anteriorly,
merging to median carina; A1 peduncle longer than rostrum; long setae on
margin of rostrum and on gastric area of carapace; abdominal somite 6 and
telson markedly setose. [Adapted from translation by V. Spiridonov.]
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CRM 013 – Katsushi Sakai
Remarks. — Kobjakova (1937) included her (sub)species in Axius, as Axius
spinulicauda amurensis. The type specimen of her species is inaccessible,
which makes its exact observation impossible. However, it is characterized
by Kobjakova’s (1937) figure and description, translated by V. Spiridonov
as follows: “the rostrum is straight and slightly elongate, spinulate, and the
submedian carinae of the gastric region are considerably longer than the lateral
ones, and are not straight but curved anteriorly and merging to the median
one”. Those morphological features are different from those of the type species
of Axius, Axius stirhynchus Leach, 1815, in which the rostrum is straight but
stoutly triangular with a median carina and denticles on the lateral margins, the
gastric region bears short submedian carinae, and a median carina is continued
to the rostral median carina. Therefore, Kobjakova’s (1937) species does not
belong in Axius, but is here assigned to a new genus, Leonardsaxius, as L.
amurensis (Kobjakova, 1937).
On the other hand, in Leonardsaxius spinulicauda (Rathbun, 1902), which
was once included in Axius and later in Calocarides, the rostrum is curved
downward, the submedian carinae are short and curved anteriorly, and not
merging with the median one (Kensley, 1996a, fig. 7c; cf. also Rathbun, 1904,
fig. 90a). It is thus evident that L. spinulicauda amurensis and L. spinulicauda
are characteristically and geographically different, so that those two forms are
considered as valid species, i.e., as L. amurensis (Kobjakova, 1937) and L.
spinulicauda (Rathbun, 1902).
Type locality. — Amur Bay, Sea of Japan, 31 m.
Distribution. — Amurskiy Bay, part of Peter the Great Bay, Sea of Japan,
31 m.
Leonardsaxius spinulicauda (Rathbun, 1902)
Axius spinulicauda Rathbun, 1902: 886, 10; Rathbun, 1904: 149, fig. 90a-b.
Axius (Paraxius) spinulicauda — Borradaile, 1903: 538.
Axiopsis spinulicauda — Schmitt, 1921: 111, fig. 74; Hart, 1982: 44, fig. 8, 1 colour fig.
Axiopsis (Axiopsis) spinulicauda — De Man, 1925d: 6 (list), 69 (key).
Acanthaxius spinulicaudus — Sakai & De Saint Laurent, 1989: 66.
Calocarides spinulicauda — Kensley, 1996a: 54 (list), 66, fig. 7.
Material examined. — USNM 25239, holotype, female (TL/CL, 57.0/19.0 mm), Bodega
Head, California, “Albatross” Sta. 3172, 113 m.
Diagnosis of holotype. — Female Plp1 uniramous and bisegmented, distal
segment slender and longer than proximal one; female Plp2 biramous, endopod
on left side missing, endopod on right side lacking appendix interna; detached
Plps3-5 endopods with appendix interna.
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127
Remarks. — The species Axius spinulicauda Rathbun, 1902 was included
in the genus Calocarides by Kensley (1996a: 66). However, it is not to be
included in Calocarides, in which the gastric median carina bears a few
denticles anteriorly; the submedian and lateral carinae are serrate; and the
P1 carpi and chelae are characteristically tuberculate on the lateral surfaces.
It is probably better included in the genus Leonardsaxius, because it is very
similar to the type species of Leonardsaxius, L. werribee (Poore & Griffin,
1979) in that the rostrum is elongate and triangular with an upturned apex;
the lateral margins are armed with teeth, extending onto the gastric region as
lateral carinae; the gastric region is convex, bearing a median carina with 3
teeth anterior to the hepatic one, originating from the base of rostrum, and
smooth submedian and lateral carinae; and the P1 carpi and chelae are not
tuberculate on the lateral surfaces. Although it is impossible to compare this
species with those of Calocarides in regard of the male Plps1-2, because this
species was described only on the female type specimen, it is safely concluded
that Axius spinulicauda Rathbun, 1902 can be reclassified under the new genus
Leonardsaxius as L. spinulicauda (Rathbun, 1902).
Type locality. — Off Bodega Head, California, “Albatross” Sta. 3172,
113 m.
Distribution. — California — off Bodega Head (Rathbun, 1902; Kensley,
1996a); 113 m.
Leonardsaxius werribee (Poore & Griffin, 1979)
Axiopsis (Axiopsis) werribee Poore & Griffin, 1979: 232, figs. 5, 6; Poore & Collins, 2009: 244,
fig. 14b.
Calocarides werribee — Sakai & De Saint Laurent, 1989: 84; Sakai, 1994: 201 (key).
Axiopsis werribee — Kensley, 1996e: 469 (list); Poore, 2004: 174, figs. 45c, d, 46b.
Material examined. — AM P 25276, paratype, 1 female (TL/CL, 23.0/7.5 mm), Port Phillip
Bay, det. G. Poore; AM P 25285, paratype, 1 female (TL/CL, 22.0/8.2 mm), Fisheries &
Wildlife Dept., Sp. No. 050, Sample 969/3; AM P 25277, paratype, 2 juvs. (TL/CL, 9.0/3.0;
10.0/3.5 mm), Fisheries & Wildlife Dept., Sp. No. 340, Sample 917/2; AM P 25286, paratype,
1 female (TL/CL, 13.0/4.8 mm), Fisheries & Wildlife Dept., Sp. No. 7409, Sample 983/3; AM P
25278, paratype, 1 juv. (TL/CL, 8.0/3.2 mm), Fisheries & Wildlife Dept., Sp. No. 303, Sample
917/4; AM P 72885, 1 juv. (TL/CL, 6.0/2.8 mm), Malabar Deep Ocean Outfall Study, Sydney,
benthos samples, site Sept. 95, no. 22, 1995-1998: USNM 1082580, paratype, 1 male (TL/CL,
35.0/11.0 mm); USNM 1082581, paratype, 1 female (TL/CL, 29.0/9.5 mm), Port Phillip Bay,
38◦ 16.3′ S 144◦ 51.5′ E, 24 m, 8.xii.1971, leg. Port Phillip Bay Environmental Study Benthic
Survey, 1969-1973, Sta. 977, det. G. Poore.
Diagnosis. — Small-sized species. P1 subequal and dissimilar in shape, in
larger cheliped fingers shorter than palm, but in smaller cheliped fingers longer
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CRM 013 – Katsushi Sakai
than palm. Male Plp1 uniramous and bisegmented, distal segment slenderly
leaf-shaped with distal setae, slightly convex on mesial margin, and slightly
longer than proximal segment; male Plp2 biramous, endopod with appendices interna and masculina; male Plps3-5 endopods with appendix interna.
Female Plp1 uniramous and bisegmented, distal segment multiarticulate; female Plps2-5 endopods with appendix interna.
Remarks. — The present species, Leonardsaxius werribee was described on
the basis of the female holotype as a species included in Axiopsis, as follows:
“female Plp1 is uniramous and bisegmented, the distal segment is smaller
and medially lobed; Plp1 is absent in male”. However, it is found through
the examination of the type specimens (NMV J. 280, holotype, and USNM
1082580, paratype), that the figure drawn as female Plp1 (Poore & Griffin,
1979, fig. 6h) is considered to be that of a male Plp1, which was subsequently
admitted by G.C.B. Poore (in litt.), who also admitted that he had by mistake
regarded his holotype as a female (Poore & Griffin, 1979: 235), but that it is
a male, and they had made a misdescription. Thus, the characteristics of the
female Plp1 shown in their figure (Poore & Griffin, 1979, fig. 6h) have turned
out to be those of a male Plp1.
It is here confirmed by examining the male specimen (USNM 1082580)
that male Plps1-2 are characterized as follows; male Plp1 is uniramous and
bisegmented, and the distal segment bears two rami mesially, as shown by
Poore (cf. Poore & Griffin, 1979, fig. 6h), but as features of a female Plp1; the
male Plp2 endopod bears appendices interna and masculina.
Type locality. — 15◦ E of Sta. Leonards, Port Phillip Bay, Victoria, Australia, 24 m.
Distribution. — Australia — Port Phillip Bay, Victoria (Poore & Griffin,
1979); Tasmania — Isthmus Bay (Poore & Griffin, 1979; Poore & Collins,
2009); 2-25 m.
Leonardsaxius sp. aff. werribee (Poore & Collins, 2009)
Axius sp. aff. werribee, Poore & Collins, 2009: 244.
Remarks. — Poore & Collins (2009) described Axius sp. aff. werribee based
on a juvenile male from east of Nowra, N.S.W., Australia. However, it would
be better to treat their species as an undefined species belonging to the genus
Leonardsaxius in the present review, because it is indicated as “aff. werribee”.
Distribution. — East of Nowra, N.S.W., Australia.
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Genus Levantocaris Galil & Clark, 1993
Levantocaris Galil & Clark, 1993: 48; Poore, 1994: 97 (key).
Diagnosis. — Rostrum narrowly protruded, lateral margins with two spines,
extending posteriorly onto gastric region as unarmed lateral carinae; gastric
region slightly convex, bearing median carina developed anteriorly and posteriorly except on interrupted median part with median tubercle, but no submedian carinae; postcervical carina absent; posterior margin of carapace sinuous,
medially convex. Eyestalks flattened, without cornea. A2 scaphocerite prominent. P1 chelate and subequal; ischium and merus with a row of interspaced
spines ventrally; carpus triangular; palm with triangular tooth on distal margin but without dorsodistal tooth; in right cheliped fingers slightly shorter than
palm, whereas in left cheliped fingers slightly longer than palm; in both chelipeds cutting edge of fixed finger roughly denticulate, proximally serrated, and
armed medially with several teeth including first strong one followed distally
by teeth gradually reducing in size; cutting edge of dactylus widely concave
at proximal third, and distal to it finely denticulate. P2 chelate; Ps3-5 simple;
Ps3-4 propodi elongate, lacking rows of spines ventrally. Male Plp1 uniramous and bisegmented, proximal segment slender, and distal segment spatulate and mesially protruded triangularly with minute setae; male Plp2 biramous
and foliaceous, endopod bearing mesiomedially appendices masculina and interna; male Plps3-5 biramous, endopods with appendix interna. Telson longer
than wide, bearing median spine on rounded posterior margin. Pleurobranchs
present on Ps2-4. Hermaphroditic, genital pores on Ps3 and 5 coxae.
Remarks. — The genus Levantocaris Galil & Clark, 1993 is closely similar
to Calocaris in the family Calocarididae, in that the gastric region bears no
submedian carinae; the eyestalks are flattened; the male Plp1 bears a spatulate
distal segment with a mesial patch of hooklets; and pleurobranchs are present.
However, they differ from each other, because in L. hornungae, type species
of Levantocaris in the family Axiidae, the A2 scaphocerite is distinct; Plp2
is biramous, and the endopod bears mesiomedially slender appendices interna
and masculina with distal setae. Whereas in Calocaris macandreae Bell, 1846,
type species of the genus Calocaris in the family Calocarididae, the A2
scaphocerite is a small and reduced spine; Plp2 is biramous, and the endopod
bears distally an elongate, stout appendix masculina with marginal setae and a
slender and shorter appendix interna with a distal patch of hooklets.
Type species. — Levantocaris hornungae Galil & Clark, 1993, by original
designation and monotypy. The gender of the generic name, Levantocaris, is
feminine.
Species included. — Levantocaris hornungae Galil & Clark, 1993.
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CRM 013 – Katsushi Sakai
Levantocaris hornungae Galil & Clark, 1993
(fig. 21)
Levantocaris hornungae Galil & Clark, 1993: 49, figs. 1-4; d’Udekem d’Acoz, 1999: 154;
Costello et al., 2001: 289; Türkay, 2001: 289; Ngoc-Ho, 2003: 455, fig. 4.
Material examined. — Levantocaris hornungae Galil & Clark, 1993 NHM 1192: 608,
holotype, hermaphroditic male (TL/CL, 28.0/10.9 mm), north of Haifa, Israel, 33◦ 00′ N
34◦ 35′ E, 1400 m, xii.1991, leg. B.S. Galil.
Diagnosis. — Rostrum (fig. 21H) narrowly triangular, lateral margins
with two spines, extending posteriorly onto anterior third of gastric region.
A2 scaphocerite strong, reaching middle of A2 penultimate segment.
Remarks. — Levantocaris bears pleurobranchs on Ps2-4 and genital pores
on the Ps3 and 5 coxae, and is hermaphroditic as Calocaris, in which the
Plp2 endopod bears a distinct appendix masculina distally provided proximally
with an appendix interna. However, Levantocaris is not included in the
Calocarididae but in the family Axiidae, because the Plp2 endopod bears
mesiomedially slender appendices interna and masculina.
Type locality. — Fifty km N.W. of Haifa, Israel, 33◦ 00′ N 34◦ 35′ E, 1400 m.
Distribution. — Only known from the type locality.
Genus Litoraxius Komai & Tachikawa, 2007
Litoraxius Komai & Tachikawa, 2007: 114.
Diagnosis. — Carapace not tuberculate, bearing median, submedian, and
lateral carinae in gastric region; postcervical median carina present. Eyestalks
stout, reaching nearly to distal end of rostrum. P1 asymmetrical, chelae
bearing several prominent tufts of plumose setae. Male Plp1 consisting of
two partially fused segments, distal segment spatulate with proximomesial
clump of adhesive hooks; Plp2 biramous, endopod with appendices interna
and masculina; Plps3-5 with appendix interna.
Type species. — Litoraxius boshu Komai & Tachikawa, 2007, by original
designation and monotypy. The gender of the generic name, Litoraxius, is
masculine.
Species included. — Only the type species is known so far.
Litoraxius boshu Komai & Tachikawa, 2007
Litoraxius boshu Komai & Tachikawa, 2007: 115, figs. 1-6.
Material examined. — CBM-ZC 9226, paratype, male (TL/CL, 43.0/15.7 mm), Ubara,
Katuura, Boso Peninsula, Chiba Pref., Japan, SCUBA diving, vi.1994, leg. M. Aizawa.
Diagnosis. — As mentioned in the diagnosis of the genus.
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Type locality. — Pacific side of Boso Peninsula, Ubara, Katsuura, Chiba
Prefecture, at 10 m depth.
Distribution. — Japan — Pacific side of Boso Peninsula, Katsuura; Ubara
Inlet, Katsuura; Kominato; Tateyama at the tip of Boso Peninsula, Chiba Pref.;
Misaki at the tip of Miura Peninsula, Kanagawa Pref.
Genus Manaxius Kensley, 2003 (sensu nov.)
Manaxius Kensley, 2003: 367 (sensu nov.)
Diagnosis. — Rostrum acutely triangular, lateral margins with 2-4 teeth and
supraorbital tooth. Gastric region bearing lateral carinae with 1-5 buttressed
teeth, median and submedian carinae with denticles; postcervical carina absent. Anterolateral margin of carapace unarmed. Cervical groove distinct almost at full length. Eyestalks subglobose; cornea pigmented. A2 scaphocerite
horn-shaped. P1 unequal; in both chelipeds dorsal margin of palm armed with
interspaced teeth; P2 chelate; fingers shorter than palm. Ps2-4 with pleurobranchs. Abdominal somites smooth laterally. Abdominal pleuron 1 narrow
and triangular ventrally; pleuron 2 broad and triangular ventrally, pleura 3-5
triangular ventrally. Male Plp1 absent; male Plp2 biramous, endopod with appendices interna and masculina at proximal third; male Plps3-5 endopods with
appendix interna. Telson longer than wide; posterior margin rounded, without
median tooth. Uropodal exopod with transverse suture.
Remarks. — Kensley (2003: 367, figs. 3, 4) described Manaxius pitatucensis as the type species of the genus Manaxius Kensley, 2003. However, it
was found through the present examination of his blue-stained male specimen
(USNM 243379) that Manaxius pitatucensis Kensley, 2003 is different from
Axiopsis (Axiopsis) pitatucensis De Man, 1925b. This is the classical case of a
misidentified type-species, and art. 70.3 of the ICZN allows the first revising
author to decide upon the further usage of the name. I consider it preferable to
fix the name in the sense of Kensley (2003) by leaving of what he thought to be
pitatucensis De Man as the type species of Manaxius; the different genus attributed to pitatucensis proper already received the name Colemanaxius in this
revision (see p. 103). Besides the decision taken here, pitatucensis Kensley
[not De Man] needs a new name, M. aganaensis, as characterized below.
Manaxius is similar to Calocarides Wollebæk, 1908, in that the rostrum
is protruded forward and triangular, bearing two lateral and supraorbital
spines; the gastric region is slightly convex, bearing 5 longitudinal carinae;
the A2 scaphocerite is distinct and horn-shaped; male Plp1 absent; male Plp2
biramous, and the endopod bears appendices interna and masculina at its
132
CRM 013 – Katsushi Sakai
proximal third; the appendix interna is slender and longer than the appendix
masculina with apical setae; male Plps3-5 endopods bear an appendix interna.
However, Manaxius and Calocarides differ from each other, because in
Manaxius the P1 carpus and chela are granulate on the lateral surface, bearing
some sharp, interspaced spines on the dorsal margin; the abdominal pleura are
triangular ventrally; and the telson is rounded on the posterior margin without a
median spine. In Calocarides, the P1 carpus and chela are smooth on the lateral
surface, bearing granules or spinules on the dorsal margin; the abdominal
pleura are rounded ventrally; and the telson is convex on the posterior margin,
with a median spine.
Type species. — Manaxius aganaensis sp. nov. by present designation. The
gender of the generic name, Manaxius, is masculine.
Species included. — Manaxius aganaensis sp. nov.; M. angulatus (Coelho,
1973); M. euophthalmus (De Man, 1905); M. galapagensis Kensley & Hickman, 2001; M. inaequalis (Rathbun, 1901); M. jenneri (Williams, 1974b);
M. kensleyi (Clark, Galil & Poore, 2007); M. mimasensis (Sakai, 1967); M.
ohsumiensis sp. nov.; M. pailoloensis (Rathbun, 1906); M. sibogae (De Man,
1925d); M. thailandensis sp. nov.
K EY
TO THE
S PECIES
OF THE
G ENUS MANAXIUS
1 – Rostrum trispinose distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. galapagensis
– Rostrum simply pointed distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Eyestalks reaching or overreaching tip of rostrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Eyestalks reaching middle of rostrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3 – Eyestalks distinct and dumbbell-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. inaequalis
– Eyestalks elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – Gastric submedian carina with 4 slender teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . M. jenneri
– Gastric submedian carina with buttressed denticles. . . . . . . . . . . . . . . . . . . .M. angulatus
5 – Gastric submedian carina terminating anteriorly with denticle . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. thailandensis sp. nov.
– Gastric submedian carina with denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – Gastric lateral carina consisting of anterior buttressed tooth and following teeth . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. mimasensis
– Gastric lateral carina consisting of 1-2 buttressed teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 – Abdominal pleuron 2 rounded ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.
– Abdominal pleuron 2 angulated ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 – Ischium of larger cheliped armed with minute denticle on ventral margin . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. euophthalmus
– Ischium of larger cheliped armed with 3 sharp spines on ventral margin . . . M. sibogae
9 – A2 segment 2 armed with ventral spine . . . . . . . . . . . . . . . . . . . . . M. aganaensis sp. nov.
– A2 segment 2 unarmed ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
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133
10 – Dorsal surface of telson with one pair of spines . . . . . . . . . . . . . . . . . . . . . . . . M. kensleyi
– Dorsal surface of telson with two pairs of spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 – Telson with rounded posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pailoloensis
– Telson subquadrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. ohsumiensis sp. nov.
Manaxius aganaensis sp. nov.
Manaxius pitatucensis — Kensley, 2003: 368, figs. 3, 4 [misidentification; partim also Manaxius paulayi Kensley, 2003].
Diagnosis. — Rostrum simply pointed distally. Eyestalks reaching middle
of rostrum. Gastric submedian carina with denticles. A2 segment 2 armed
with ventral spine. Gastric lateral carina consisting of 1-2 buttressed teeth.
Abdominal pleuron 2 angulated ventrally.
Remarks. — Kensley (2003: 268, figs. 5, 6) described Manaxius pitatucensis (De Man, 1925b) with figures. However, his M. pitatucensis has turned out
to be invalid (as shown in the remarks of the genus Manaxius sensu Sakai).
Type locality. — Agana, Guam Island.
Etymology. — The new species is named M. aganaensis, derived from the
type locality, Agana, Guam Island, with a Latin suffix “-ensis” added, referring
to a geographic locality.
Manaxius angulatus (Coelho, 1973)
Calastacus angulatus Coelho, 1973: 344; Coelho & Ramos-Porto, 1985: 82, figs. 7, 8;
Rodrigues et al., 1998: 382; Melo, 1999: 328, figs. 221-212.
Acanthaxius angulatus — Coelho et al., 2007: 4.
Diagnosis. — Rostrum simply pointed distally. Eyestalks elongate, overreaching tip of rostrum. Gastric submedian carina with buttressed denticles.
Abdominal pleuron 2 triangular ventrally. [Adapted from Coelho & RamosPorto, 1985, figs. 7, 8.]
Remarks. — This species was classified under the genus Acanthaxius by
Coelho et al. (2007). However, it does not belong in Acanthaxius, because
in Acanthaxius the fingers of the chelipeds are longer than the palm, and
pleura 2-5 are truncate or rounded ventrally, whereas in the present species, the
fingers of the chelipeds are shorter than the palm, and pleura 2-5 are triangular
ventrally (Coelho & Ramos-Porto, 1985, fig. 8b), so that it is reclassified here
under the genus Manaxius.
Type locality. — Cabo Caciporé, Amapá, Brazil, 108-118 m.
Distribution. — Only known from the type locality.
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CRM 013 – Katsushi Sakai
Manaxius euophthalmus (De Man, 1905)
(fig. 22C-E)
Calastacus euophthalmus De Man, 1905: 598; Balss, 1925: 209.
Calocaris (Calastacus) euophthalma — De Man, 1925d: 8 (list), 118 (key), 122, pl. 10 fig. 2222g.
Calaxius euophthalmus — Sakai & De Saint Laurent, 1989: 86; Kensley & Hickman, 2001:
487.
Material examined. — ZMA Crust. De. 240080, lectotype, male (TL/CL, 22.0/8.6 mm),
north of the island of Batjan, 0◦ 11′ S 127◦ 25′ E, “Siboga” Sta. 139, 4.viii, 1899-1900, mud,
stones and coral [det. as C. euophthalmus]; MNHN Th 1421, 1 female (TL/CL, 19.0/6.3 mm),
French Polynesia, Marquesas Islands, Nuju Hiva, MUSORSTOM 9, R/V “Alis”, Sta. 1298,
305 m, 9.ix.1997, leg. P. Bouchet, B. Dayrat, B. Richer de Forges [det. as M. sibogae];
USNM 243379, 1 male (CL, 6.0 mm; abdomen + telson, 15.9 mm), Okinawa, Japan,
70.1 m, 11.vi.1981, leg. R. Bolland; USNM 243378, 1 male (TL/CL, 10.0/3.2 mm), 1 female
(TL/CL,10.0/3.2 mm), Agana Bay, Guam, 30 ft, 23.x.1984, leg. J.H. Dominguez, det. B.
Kensley; USNM 243380, 1 female (TL/CL, 6.3/2.1 mm, rostrum broken), on reef front 300 ms
north of boat basin channel, Agana Bay, Guam, 42 ft [ca. 12.6 m], 26.iv.1984, leg. R.K. Kropp,
det. B. Kensley; USNM 243381, 1 male (TL/CL, 10.5/3.2 mm), Piti Channel, Cabras Island,
Guam, 40 ft [ca. 12 m], 31.v.1984, leg. H. Dominguez & R.K. Kropp, det. B. Kensley [det. as
Manaxius pitatucensis Kensley, 2003].
Diagnosis. — Rostrum narrowly triangular, lateral margins with spines and
distinct supraorbital spine. Gastric region with 5 carinae. P1 (fig. 22C, D)
subequal; in larger cheliped merus unarmed on dorsal margin but denticulate
on ventral margin; carpus with irregularly arranged spines on dorsal margin;
palm with irregularly arranged teeth on dorsal margin, and 1-2 distinct subdistal spines on lateral surface. Abdominal somites (fig. 22E) smooth laterally;
pleuron 1 rounded ventrally; pleura 2-5 truncate ventrally, without marginal
spines; pleuron 6 rounded ventrally in anterior half. Male Plp1 absent; Plp2
biramous and slender, endopod with appendices interna and masculina, appendix masculina extremely longer than appendix interna, bearing setae; Plps3-5
endopods with appendix interna. Telson longer than wide, bearing proximal
lobe with spine followed distally by denticles on lateral margin; posterior margin rounded without median tooth. Uropodal endopod oval, bearing spines on
lateral margin, movable spine distolaterally, and median carina with spines on
dorsal surface. Uropodal exopod oval, bearing 5 denticles on lateral margin
including posterolateral one, and distal flap with distinct, movable spine laterally, as well as some denticles sublaterally on dorsal surface.
Remarks. — Axius pailoloensis Rathbun, 1906, Calocaris (Calastacus) sibogae De Man, 1925, and Manaxius aganaensis sp. nov. are closely similar to
M. euophthalmus (De Man, 1905), because of the denticles on the five gastric
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135
carinae, the spines on the dorsal margin of the P1 palm, the denticulation of
the cutting edges of the P1 chela, and the form of the A2 scaphocerite. However, those three species are different, as shown in the key, in the features of
the ventral margin of A2 segment 2, the denticulation on the ventral margin of
the ischium of the larger cheliped, and the form of the telson.
Type locality. — North of Batjan Island, Indonesia, 0◦ 11′ S 127◦ 25′ E,
“Siboga” Sta. 139, 397 m.
Distribution. — Indonesia — north of Batjan Island (De Man, 1905, 1925),
397 m; between the islands Wowoni and Buton [= Butung] (De Man, 1925d).
Hawaiian Islands — Pailolo Channel (Rathbun, 1906; Sakai & De Saint
Laurent, 1989); 74-252 m.
Manaxius galapagensis (Kensley & Hickman, 2001)
Calaxius galapagensis Kensley & Hickman, 2001: 484, fig. 1, table 1.
Material examined. — USNM 308977, holotype, 1 ovig. female (41.0/14.0 mm), west end
of North Plaza, Santa Cruz Island, Galapagos Islands, 6 m, 28.viii.1997, leg. C.P. Hickman, Jr.
Type locality. — West end of North Plaza, off Isla Santa Cruz, Galapagos
Islands, 6 m.
Distribution. — Only known from the type locality.
Manaxius inaequalis (Rathbun, 1901)
Axius inaequalis Rathbun, 1901: 96, fig. 18a-b.
?Axiopsis inaequalis — Borradaile, 1903: 539.
Axiopsis (Axiopsis) inaequalis — De Man, 1925d: 6 (list), 69 (key); Schmitt, 1935a: 190, fig. 50.
Calaxius inaequalis — Sakai & De Saint Laurent, 1989: 85; Kensley & Hickman, 2001: 487.
Type locality. — Mayagüez Harbor, Puerto Rico, 289-309 m.
Distribution. — Puerto Rico — Mayagüez Harbor (Rathbun, 1901); 289309 m.
Manaxius jenneri (Williams, 1974)
Calocaris (Calastacus) jenneri Williams, 1974b: 451, figs. 1-10.
Calaxius jenneri — Sakai & De Saint Laurent, 1989: 86; Kensley & Hickman, 2001: 487; Clark
et al., 2007: 72; Heard et al., 2007: 19, fig. 12.
Material examined. — USNM 150472, holotype, 1 male (TL/CL, 38.0/15.0 mm), east
of Cape Lookout, North Carolina, 34◦ 18′ N 74◦ 01.2′ W – 34◦ 17.1′ N 76◦ 01.3′ W, 100-85 m,
3.x.1969, leg. R/V “Eastwing”, Sta. 12885, dredge, C.E. Jenner & class; USNM 150473,
paratype, 1 male (TL/CL, 26.0/9.5 mm), SSW of Cape Lookout, North Carolina, 33◦ 43.3′ N
136
CRM 013 – Katsushi Sakai
76◦ 42′ W – 34◦ 44′ N 76◦ 41′ W, 91-98 m, 4.xi.1968, leg. R/V “Eastwing”, Sta. 10789, dredge, J.
& W. Vernberg.
Diagnosis. — Male Plp1 absent; male Plp2 biramous and slender; endopod
with appendices interna and masculina; male Plps3-5 biramous and slender,
endopods with appendix interna.
Type locality. — S.W. of Cape Lookout, North Carolina, 33◦ 43.1′ N
76◦ 40.7′ W – 33◦ 43.4′ N 76◦ 40.3′ W, 100 m.
Distribution. — North Carolina — Cape Lookout (Williams, 1974b); 85110 m.
Manaxius kensleyi Clark, Galil & Poore, 2007
Calaxius kensleyi Clark et al., 2007: 63-73, figs. 1-6.
Remarks. — The present species M. kensleyi was described by Clark et
al. (2007) as the first member of Calaxius from the western Atlantic Ocean.
Most of the species in Calaxius are reclassified under Manaxius sensu Sakai,
except C. acutifrons Sakai & De Saint Laurent, 1989, in which the abdominal
pleura are acutely angulate ventrally. M. kensleyi is very similar to the IndoWest Pacific species, M. euophthalmus (De Man, 1905), but differs from it,
because in M. kensleyi the Mxp3 ischium is armed with 4 spines on the
mesial margin, and abdominal pleuron 2 is triangular ventrally, whereas in
M. euophthalmus the Mxp3 ischium is armed with two spines on the mesial
margin, and abdominal pleuron 2 is truncate ventrally (fig. 22E).
Type locality. — Off Gabon, 0◦ 5′ S 9◦ 3′ E, 49.2-49.3 m.
Distribution. — Off Gabon, 0◦ 4-5′ S 9◦ 3′ E; 49.2 m.
Manaxius mimasensis (Sakai, 1967)
Calocaris (Calastacus) mimasenis Sakai, 1967: 41, pl. 3 fig. 1.
Calocaris mimasensis — Sakai, 1987a: 303 (list).
Calaxius mimasensis — Sakai & De Saint Laurent, 1989: 86; Kensley & Hickman, 2001: 487;
Komai et al., 2002: 30, figs. 4-6, tables 1, 4.
Diagnosis. — Male Plp1 absent; Plp2 biramous, endopod with appendices
interna and masculina at proximal third (cf. Komai et al., 2002: 37, fig. 6K).
Type locality. — Tosa Bay, Japan.
Distribution. — Japan — Tosa Bay (Sakai, 1967), littoral; off S. Izuhara,
Tsushima Island, 33◦ 56.21′ N 129◦ 25.36′ E, 1333 m.
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137
Manaxius ohsumiensis sp. nov.
(figs. 23, 24)
Material examined. — SMF 31814, holotype, female (TL/CL, 21.0/8.2 mm), Ohsumi shoto,
between Yakushima and Tanegashima, Kagoshima Pref., Japan, 80-82 m, KT93-09, Sta. YT
12(1), 17.vi.1993, by 1 m biol. dredge, leg. S. Ohta.
Description of female holotype. — Rostrum (fig. 23A, B) acutely triangular, bearing two teeth on each lateral margin, which extend short posteriorly
as gastric lateral carinae with paired teeth anteriorly and another tooth posterior to lateral carina on left side. Carapace studded with tubercles and setae.
Gastric region convex, bearing distinct median carina with some short or elongate tubercles extending to cervical groove, and submedian rows of irregularly
arranged denticles submedially between median and lateral carinae. Cervical
groove distinct at full length. Cardiac prominence present. Eyestalks subglobose, reaching distal third of rostrum. A1 peduncle overreaching rostrum at
distal end of penultimate segment; proximal segment unarmed and longer than
penultimate and distal segments combined; penultimate segment unarmed and
slightly longer than distal one. A2 scaphocerite distinct and horn-shaped, and
about twice as long as dorsodistal tooth of segment 2; penultimate segment
about twice as long as distal one, overreaching rostrum at its middle.
Mxp3 (fig. 23C) exopod absent. Coxa with sharp tooth on ventral margin;
basis with sharp ventral tooth; ischium 2.5 times as long as wide, bearing tooth
on mesial margin, and ischial crest with more than 16 teeth; merus about as
long as ischium, bearing two sharp teeth on mesial margin; carpus about twothirds as long as merus and subequal to propodus; dactylus slightly shorter
than propodus.
P1 (fig. 24A, B) subequal. In right larger cheliped coxa carinate proximally
on mesial margin, bearing two teeth; basis unarmed; ischium with three ventral
teeth. Merus about 1.8 times as long as wide; dorsal margin largely convex in
distal part, bearing two subterminal teeth, and ventral margin almost straight,
bearing series of 5 distinct teeth; carpus wider than long, bearing two subdistal
teeth on dorsal margin and median tooth on ventral margin. Chela twice
as long as wide; palm 1.2 times as long as wide, bearing three interspaced
teeth on dorsal margin, one subdistal tooth on dorsolateral and dorsomesial
margins, and another three small teeth at mesiodistal part of dorsal margin,
ventral surface unarmed. Fixed finger with two w-shaped teeth proximally and
convex subdistally on cutting edge. Dactylus shorter than palm, and concave
proximally. In smaller left cheliped, ischium bearing 4 teeth on ventral margin
and distinct subdistal spine on dorsal margin; merus broad, bearing 5 teeth
138
CRM 013 – Katsushi Sakai
Fig. 23. Manaxius ohsumiensis sp. nov. A, whole body, lateral view; B, carapace, dorsal view;
C, Mxp3, mesial face; D, P3-5 coxae and thoracic sternite between coxae 4; E, female Plp2.
A-E, SMF 31814, holotype, female (TL/CL, 21.0/8.2 mm), Ohsumi shoto, between Yakushima
and Tanegashima, Kagoshima Pref., Japan, 80-82 m. Scales 1 mm.
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139
Fig. 24. Manaxius ohsumiensis sp. nov. A, large cheliped on right side; B, small cheliped on
left side; C, P3; D, P5; E, telson and uropod on left side. A-E, SMF 31814, holotype, female
(TL/CL, 21.0/8.2 mm), Ohsumi shoto, between Yakushima and Tanegashima, Kagoshima Pref.,
Japan, 80-82 m. Scales 1 mm.
140
CRM 013 – Katsushi Sakai
on ventral margin and subterminal spine on dorsal margin; carpus slightly
wider than long, bearing two spines on dorsal margin and two tubercles
on ventral margin; chela narrower than that of larger cheliped, and more
than twice as long as wide; palm slightly longer than wide, bearing three
interspaced teeth on dorsal margin; fixed finger with triangular subproximal
tooth, and concave distally. P2 merus elongate, bearing three teeth on ventral
margin; fingers about half length of palm. P3 (fig. 24C) merus with 4 teeth
on ventral margin; propodus twice as long as carpus, bearing 5 transverse
rows of transparent articulate spinules on ventrolateral surface; dactylus with
7 transparent spinules on dorsal margin. P4 similar in shape to, but shorter
than, P3; merus unarmed on ventral margin; propodus about 1.5 times as long
as carpus, bearing transverse rows of transparent articulate spinules laterally;
dactylus sharp and armed with transparent articulate spinules ventrolaterally.
P5 simple; coxa (fig. 24D), basis, and ischium unarmed; propodus 1.8 times
as long as carpus, and ventral margin protruded distally to form subchela with
dactylus.
Abdominal somites 1-6 (fig. 23A) with longitudinal carina between tergum
and pleuron. Abdominal pleuron 1 narrow, protruded ventrally into obtuse
angle, and curved posteriorly. Pleura 2-6 angulated ventrally. Female Plp1
uniramous, consisting of proximal segment and multiarticulate flagellum;
female Plps2-5 (fig. 23E) biramous, with slender foliaceous endopod and
exopod; endopods without appendix interna.
Telson (fig. 24E) longer than wide, slightly converging posteriorly, and
slightly longer than somite 6; lateral margins bearing proximal lobe with spine
followed distally by three interspaced teeth including one at posterolateral corner; posterior margin rounded without median spine; dorsal surface longitudinally furrowed medially, bearing pair of sharp teeth at each posterolateral angle
and paired oblique rows of three spines on posterior half. Uropodal endopod
bearing 6-7 spines on lateral margin, including one at posterolateral angle, and
a row of three spines medially on dorsal surface. Uropodal exopod denticulate
on distal half of lateral margin, on dorsal surface bearing a submarginal row of
teeth and distal flap with distinct, movable posterolateral spine.
Remarks. — The present new species is counted as the 11th species of the
genus Manaxius, because, as in the other species of Manaxius, the P1 chelae
are setose, the dorsal margin of the palm bears teeth, the abdominal pleura
are triangular ventrally, and the telson bears no median tooth on the posterior
margin. The present species is characteristic in bearing paired spines on each
side of the rostral base.
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141
Type locality. — Ohsumi shoto, between Yakushima and Tanegashima,
Kagoshima Pref., Japan, 80-82 m.
Distribrution. — Only known from the type locality.
Etymology. — The present new species is named after the type locality,
the Ohsumi Islands between Tanegashima und Yakushima in Kagoshima
Prefecture, where the specimen was collected by dredge [S. Ohta & H. Watabe,
in litt.]
Manaxius pailoloensis (Rathbun, 1906)
Axius pailoloensis Rathbun, 1906: 893, fig. 49.
Axiopsis pailoloensis — Balss, 1925: 209.
Axiopsis (Axiopsis) pailoloensis — De Man, 1925d: 6 (list), 69 (key), 89, pl. 7 fig. 15-15b.
Calaxius pailoloensis — Sakai & De Saint Laurent, 1989: 85, fig. 22; Poore, 1994: 114, fig. 3a;
Kensley & Hickman, 2001: 487.
Material examined. — USNM 30533, lectotype, male (TL/CL, 31.0/11.0 mm), Pailolo
Channel, Hawaii Islands, 9.iv.1902, leg. R/V “Albatross” Sta. 3859, 138-140 m, det. Rathbun.
Diagnosis. — Rostrum simply pointed distally. Eyestalks reaching middle
of rostrum. A2 segment 2 unarmed ventrally. Gastric submedian carina with
denticles. Gastric lateral carina consisting of 1-2 buttressed teeth. Abdominal
pleuron 2 angulated ventrally. Telson rectangular with rounded posterior
margin.
Type locality. — Pailolo Channel, Hawaiian Islands, 248-252 m.
Distribution. — Only known from the type locality.
Manaxius sibogae (De Man, 1925)
Calocaris (Calastacus) Sibogae De Man, 1925d: 8 (list), 118, pl. 9 fig. 21-21e.
Calaxius sibogae — Sakai & De Saint Laurent, 1989: 86; Kensley & Hickman, 2001: 487;
Ngoc-Ho, 2005: 57, fig. 5.
Diagnosis. — Rostrum simply pointed distally. Eyestalks reaching middle
of rostrum. Gastric submedian carina with denticles. Gastric lateral carina
consisting of 1-2 buttressed teeth. Ischium of larger cheliped armed with 3
sharp spines on ventral margin. Abdominal pleuron 2 rounded ventrally.
Type locality. — North of Batjan Island, Indonesia, 0◦ 11′ S 127◦ 25′ E,
“Siboga” Sta. 139, 397 m.
Distribution. — Only known from the type locality.
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CRM 013 – Katsushi Sakai
Manaxius thailandensis sp. nov.
Eutrichocheles sp., Sakai, 1992a: 169, fig. 10.
Material examined. — ZMUC, holotype, male (TL/CL, 9.5/3.8 mm), paratype, 1 female
(TL/CL, 8.0/2.2 mm), Gulf of Thailand, 7◦ 00′ N 103◦ 18′ E, 55 m, mud, a little sand, and shell,
GE Sta. 381-I, 8.vi.1951 (Sakai, 1992a).
Diagnosis. — Rostrum simply pointed distally, and armed with lateral
denticles. Eyestalks subglobose, reaching middle of rostrum. Anterolateral
margin of carapace with two denticles. Gastric median carina with anterior
spine and hepatic tubercle. Gastric submedian carina with anterior buttressed
denticle. Abdominal pleuron 2 angulated ventrally. Telson longer than broad,
bearing proximal lobe with spine followed posteriorly by 3-4 spines on lateral
margin. [Adapted from Sakai, 1992.]
Remarks. — The present species was treated in 1992 (Sakai, 1992a) as
Eutrichocheles sp. However, the present revision suggests that it is to be
considered a species included in Manaxius and not in Eutrichocheles, because
in the present species the A2 scaphocerite is short and simple, and the telson is
unarmed on the posterior margin as in Manaxius, while in Eutrichocheles the
A2 scaphocerite is short and usually bifurcate, and the telson is armed with a
median spine on the posterior margin.
Type locality. — Gulf of Thailand (Sakai, 1992a), 55 m.
Distribution. — Only known from the type locality.
Genus Marianaxius Kensley, 1996
Marianaxius Kensley, 1996e: 483.
Diagnosis. — Carapace slightly convex. Rostrum armed with teeth on
lateral margins; gastric region anteriorly bearing short, entire median and
lateral carinae, but no submedian carinae. Anterolateral margin of carapace
without supraorbital spine; postcervical carina and spines absent. Eyestalks
subglobose, cornea well pigmented, facets hexagonal. A2 scaphocerite fairly
well developed as a spike, less than half length of A2 segment 4. Mxp3 ischial
crest with about 20 denticles; merus with 3 teeth on ventral margin, distalmost
largest. P1 asymmetrical, not sexually dimorphic; P2 chelate, fingers longer
than palm; Ps3-5 simple. Male Plp1 uniramous and unsegmented; male Plp2
biramous, endopod with strong appendix masculina in middle, but without
appendix interna; male Plps3-5 biramous, endopods without appendix interna.
Telson longer than wide, tapering toward posterior margin without median
spine, and bearing a row of 4 articulate spines at each posterolateral angle;
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143
dorsal surface with non-articulate submedian spines. Uropodal endopod broad,
with three lateral spines including one at posterolateral angle, and a row of
4 spines medially on dorsal surface. Uropodal exopod broad, bearing three
lateral spines including one at posterolateral angle, and distal flap with strong,
movable spine at posterolateral angle. [Adapted from Kensley, 1996e.]
Remarks. — Kensley (1996e) mentioned that “Marianaxius superficially
appears most similar to Paraxiopsis (Kensley, 1995 — [not 1995, but 1996c]),
especially in the lack of appendices internae on the pleopods. They differ,
however, in that Marianaxius does not have a bifid, shortened antennal scale,
possesses pleopod 1 in the male (absent in Paraxiopsis), lacks a submedian
carapace carina and spines” (Kensley, 1996c: 484).
However, it must be pointed out that he made a mistake in comparing Marianaxius with Paraxiopsis, because the male Plp1 is present in Paraxiopsis,
though he mentioned in his remarks, as shown above, that male Plp1 is absent
in Paraxiopsis.
Type species. — Marianaxius kroppi Kensley, 1996, by original designation. Gender of generic name, Marianaxius, masculine.
Species included. — Marianaxius kroppi Kensley, 1996.
Marianaxius kroppi Kensley, 1996
Marianaxius kroppi Kensley, 1996e: 484, figs. 9, 10, table 4; Kensley, 2003: 368.
Material examined. — USNM 243530, holotype, male (TL/CL, 17.0/6.2 mm), Piti Bay,
Guam, 15 m, 11.vi.1986, leg. L. Kropp; USNM 43531, allotype, female (TL/CL, 14.0/5.5 mm),
Piti Bay, Guam, 15 m, 18.v.1986, leg. L. Kropp.
Type locality. — Piti Bay, Guam, outer reef, flat rubble, to 1.5 m.
Distribution. — Guam — Piti Bay (Kensley, 1996e); to 1.5 m.
Genus Michelaxiopsis Poore & Collins, 2009
Michelaxiopsis Poore & Collins, 2009: 000.
Diagnosis. — Rostrum triangular, bearing median carina; lateral margins
armed with teeth, extending posteriorly onto gastric region. Gastric region
forming convex platform, bearing tuberculate lateral, submedian, and median
carinae, and additionally a short row of tubercles between median and submedian carinae. Anterolateral margin of carapace unarmed. Cervical groove
present at full length. Eyestalks subglobose; cornea pigmented. A2 scaphocerite prominent. P1 subequal in length and shape. Ps3-4 propodi narrow.
Abdominal pleura 3-5 truncate ventrally, bearing soft marginal setae. Male
144
CRM 013 – Katsushi Sakai
Plp1 absent; male Plp2 endopod with appendix interna, but without appendix
masculina; male Plps3-5 biramous and broad, bearing soft marginal setae; endopods with appendix interna. Telson longer than wide, lateral margins armed
with a few spinules or unarmed, slightly convergent toward posterior margin
with median tooth. Uropodal exopod with distal flap.
Remarks. — Axiopsis (Axiopsis) australiensis De Man, 1905b was included
in the genus Axius by Sakai & De Saint Laurent (1989), but it does not belong
in Axius, in which the male Plp1 is present and the male Plp2 endopod bears
appendices interna and masculina, because in A. (Axiopsis) australiensis the
male Plp1 is absent and the male Plp2 endopod bears an appendix interna
but no appendix masculina. The species seems to belong rather to Axiorygma
Kensley & Simmons, 1988 by the features of Plps1-2. However, A. (Axiopsis)
australiensis is also different from the type species of Axiorygma, A. nethertoni
Kensley & Simmons, 1988. In the former, the gastric region is provided with
tuberculate lateral, median, and submedian carinae, and additionally a row
of tubercles between the median and submedian carinae; the eyestalks are
subglobose and shorter than the rostrum; the male P1 chelae bear no cluster
of slender feathered setae; and the telson is longer than wide, with the lateral
margins armed with a few spinules or unarmed, slightly convergent toward
the posterior margin with a median tooth. Whereas in the latter species, the
gastric region is provided with five smooth carinae; the eyestalks are elongate,
reaching the level of the rostral apex; the male P1 chelae proximally bear a
dorsal cluster of slender feathered setae; and the telson is subsquare, bearing
distinct spines on the lateral margins. For those reasons mentioned above,
A. (Axiopsis) australiensis De Man, 1925b is reclassified under the genus
Michelaxiopsis Poore & Collins, 2009.
Type species. — Axiopsis (Axiopsis) australiensis De Man, 1925b, by
original designation. The gender of the generic name, Michelaxiopsis, is
feminine.
Species included. — Michelaxiopsis australiensis (De Man, 1925b).
Michelaxiopsis australiensis (De Man, 1925)
(figs. 25, 26A, B)
Axiopsis (Axiopsis) australiensis De Man, 1925b: 127, fig. 4-4j; De Man, 1925d: 5 (list), 69
(key); Poore & Griffin, 1979: 226, fig. 2.
Axius australiensis — Sakai & De Saint Laurent, 1989: 29; Sakai, 1992a: 165, fig. 7; Sakai,
1994: 200 (key).
Axiopsis australiensis — Kensley, 1996e: 469 (list); Poore, 2004: 174, figs. 45a, b, 46a, 176.
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145
Fig. 25. Michelaxiopsis australiensis (De Man, 1925b). A, carapace, dorsal view; B, cheliped
on right side, lateral view; C, distal part of cheliped on right side, mesial view; D, cheliped on
left side, lateral view. A-D, ZMB 14260, holotype, female (TL/CL, 22.0/9.1 mm), Port Jackson,
Australia. Scales 1 mm.
146
CRM 013 – Katsushi Sakai
Fig. 26. Michelaxiopsis australiensis (De Man, 1925b). A, telson and uropodal endopod
and exopod on left side; B, telson and uropod with endopod and exopod on right side.
A, Michelaxiopsis australiensis, ZMB 14260, holotype, female (TL/CL, 22.0/9.1 mm), Port
Jackson, Australia; B, Michelaxiopsis australiensis, AM P 29545, 1 male (TL/CL, 62.0/
20.8 mm), Long Reef, Collaroy, N.S.W., Australia, 33◦ 45′ S 151◦ 19′ E, intertidal. Scales 1 mm.
Michelaxiopsis nauo Poore & Collins, 2009: 263, figs. 27, 28. [Type locality. — Roxby I., Sir
Joseph Banks Group, South Australia].
Material examined. — ZMB 14260, holotype, female (TL/CL, 22.0/9.1 mm), Port Jackson,
Australia, leg. Oschat; RMNH D10605, 1 male (TL/CL, 43.0/13.6 mm), Long Reef, Collaroy,
N. of Sydney, New South Wales, Australia, 22.iv.1955, leg. L.B. Holthuis; AM P 29545, 1
male (TL/CL, 62.0/20.8 mm), 1 ovig. female (TL/CL, 66.0/21.6 mm), Long Reef, Collaroy,
N.S.W., Australia, 33◦ 45′ S 151◦ 19′ E, intertidal, under boulders, intertidally, 16.x.1978, leg.
I. Bennet; AM P23493, 1 male (TL/CL, 38.0/12.6 mm), Long Reef, Collaroy, coast N. of
Sydney, 33◦ 44′ S 151◦ 19′ E, under stones, low tide, 25.i.1956, leg. E. Pope; AM P24700, 1
male (TL/CL, 44.0/14.5 mm), Angourie, Clarence River mouth, N.S.W., 29◦ 29′ S 153◦ 22′ E,
1.ix.1961, leg. A.A. Cameron; AM P 72882, 1 male (TL/CL, 46.0/14.8 mm), N. side of Long
Reef, Collaroy, N.S.W., intertidal, 28.viii.1976, leg. I. Ioch & J. Farguhar; AM P 11360, 1 male
(TL/CL, 37.0/12.0 mm), Angourie, near Yamba, N.S.W., leg. Allan & Cameron; AM P 8693,
1 male (TL/CL, 52.0/16.8 mm), 1 ovig. female (TL/CL, 59.0/17.5 mm), Long Reef, Collaroy,
near Sydney, N.S.W., intertidal, 27.i.1964, leg. J.C. Yaldwyn; AM P 64235, 1 female (TL/CL,
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25.0/8.7 mm), northwest side of Bare Island, Botany Bay, N.S.W., 33◦ 59.6′ S 151◦ 13.8′ E,
13.8 m, 18.v.1998, leg. S. Ahyong; AM P 7472, 1 male (TL/CL, 35.0/11.2 mm), 1 ovig.
female (TL/CL, 47.0/14.4 mm); 1 female (TL/CL, 40.0/13.4 mm), Long Reef, Port Jackson;
AM P 36104, 1 ovig female (TL/CL, 50.0/17.0 mm), Muccumbulga Point, Two Gold Bay,
N.S.W., intertidal rocks, 11.xii.1984, leg. S. Eab; AM P 72883, 1 female (TL/CL, 29.0/9.4 mm),
Angourie, Clarence River, N.S.W., 4.x.1940, leg. A.A. Cameron.
Diagnosis. — Rostrum (fig. 25A) stoutly triangular, blunt at apex, and
slightly longer than basal width, bearing 5-6 blunt teeth on each lateral margin. Gastric region with longitudinal, tuberculate submedian carinae curving
together anteriorly about midway along tuberculate median carina to form
deep inverted U-shape; median carina continuing to base of rostrum; posterior
part of gastric region smooth; anterolateral margin of carapace smooth. A1 peduncle reaching middle of A2 penultimate segment, scaphocerite present as a
strong spine. A2 segment 2 with distolateral spine; scaphocerite present as a
single long spine, reaching about midpoint of segment 4; segment 3 smooth.
Eyestalks half length of rostrum. Mxp3 ischium with strong teeth on ischial
crest, merus with 3-4 large mesial spines distally. P1 (fig. 25B, D) subequal
and not sexually dimorphic. P1 ischium dentate on ventral margin; merus also
dentate on ventral margin, including strong spines, and smooth dorsally; carpus and chela smooth; distal margin of palm convex, with denticles in ventral
half; cutting edge of fixed finger with strong tooth medially and another one
near distal end; dactylus as long as fixed finger, bearing strongly curved tip
(fig. 25C). P2 chelate; fingers shorter than palm. Ps3-5 propodi with rows of
spines ventrolaterally; dactyli with longitudinal row of spines laterally. Male
Plp1 absent; male Plp2 endopod with appendix interna, but without appendix masculina; male Plps3-5 endopods with appendix interna. Female Plp1
uniramous, consisting of proximal segment and multiarticulate flagellum, this
flagellum about twice as long as proximal segment; female Plps2-5 endopods
with appendix interna. Telson (fig. 26A, B) longer than wide, armed with a few
denticles or unarmed on lateral margins, which are slightly convergent toward
rounded posterior margin with a median spine, bearing two pairs of spines
on dorsal surface. Uropodal endopod bearing small lateral spine at distal end,
sometimes additionally 1-2 lateral spines, and dorsal longitudinal ridge with
3-5 spines; uropodal exopod bearing spines on distal half of lateral margin,
movable spine distolaterally, and sublateral row of spinules and transverse suture with spinules dorsally. [Adapted from Poore & Griffin, 1979.]
Remarks. — Poore & Collins (2009: 263) established a new species,
Michelaxiopsis nauo, and distinguished their species from M. australiensis,
because in M. nauo the telson always bears three spines on the lateral margin,
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while in M. australiensis it bears no spine on the lateral margin. However,
the difference in the number of teeth on the lateral margins of the telson is
observed in the series of the specimens of M. australiensis examined in the
present study and, moreover, De Man (1925: 129, fig. 4c) described from
the holotype of his species, M. australiensis as follows: “. . . der vorderste
Zahn des Seitenrandes [of the telson], whelcher bei Ax. consobrina ungefähr
am vorderen Drittel gelegen ist, liegt bei der neuen Art [= M. australiensis]
mehr nach hinten, nahezu in der Mitte des Telsons, den medianen Endstachel
nicht mitgerechnet. Der vordere Teil der Seitenränder zwischen der Basis
und dem vorderen Zahn erscheint schwach gebogen, beinahe recht, bei Ax.
consobrina ein bißchen konkav; bei Ax. consobrina tragen die Seitenränder
zwei Zahne zwischen dem vordersten und der hinteren Seitenecken, bei Ax.
australiensis nur einen, gerade in der Mitte zwischen dieser Ecke und dem
fordersten Zahne. . . ” [= . . . the most anterior spine is located rather posteriorly
on the middle of the lateral margin of the telson in M. australiensis, and there
is another spine between the anterior spine and the posterolateral angle. . . ]
(fig. 26A).
As mentioned above, the difference in the number of spines on the lateral
margin of the telson is not so decisive a character as to distinguish those two
species from each other, and it is true of the difference in number of the spines
on the gastric median carina in M. australiensis, which suggests that those two
species, M. nauo and M. australiensis are not to be distinguished from each
other, and therefore M. nauo is synonymized with M. australiensis.
Type locality. — Port Jackson, Australia.
Distribution. — Australia — Port Jackson, N.S.W. (De Man, 1925b);
Clarence River, Collaroy; North Head, Shellharbour, N.S.W., Broken Bay,
about 50 km north of Sydney on the coast of N.S.W. (Sakai, 1992a), and
Shoreham, Victoria (Poore & Griffin, 1979); intertidal zone; Roxby Is., Sir
Joseph Banks Group, Reevesby Is., Marum Is., South Australia; 3-6 m (Poore,
2009).
Genus Newzealandaxius gen. nov.
Diagnosis. — Rostrum prominent and furrowed on surface; lateral margins
armed with erect spines, extending posteriorly onto gastric region. Gastric region convex, bearing 5 carinae; median carina originating from middle keel of
rostrum and running to anterior two-thirds of gastric region, bearing in middle an elongate-triangular patch of granules gradually narrowing anteriorly
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and becoming a single row of spines; submedian carinae with strip of granules becoming spines just behind base of rostrum; lateral carinae with spines,
posterior smaller ones and anterior larger ones including anteriormost stout,
thorn-like one at some distance from base of rostrum. Anterolateral margins
of carapace unarmed. Cervical groove distinct. Eyestalks subglobose; cornea
pigmented. A2 scaphocerite prominent and horn-shaped. P1 unequal in size
and dissimilar in shape. P2 chelate, fingers longer than palm. Ps3-4 propodi
elongate, bearing transverse rows of spines ventrolaterally. Abdominal pleura
1-5 convex between terga and pleura, and acutely triangular ventrally; abdominal pleura 3-5 in male with spinule on anteroventral margin, while abdominal pleura 3-5 in females unarmed ventrally. Male Plp1 absent; male Plp2
biramous, endopod and exopod slender, endopod with appendices interna and
masculina; male Plps3-5 biramous, endopods with appendix interna. Female
Plp1 uniramous, consisting of proximal segment and multiarticulate flagellum;
female Plps2-5 endopods with appendix interna. Telson longer than wide, bearing proximal lobe with spine on lateral margin; posterior margin with median
spine. Uropodal exopod without distal flap.
Remarks. — It has turned out through the examination of the male type
specimen of Axius (Axius) novae-zealandiae Borradaile, 1916 (NHM 1917.1.
29.106-110, TL 50 mm), which was included in Spongiaxius by Sakai & De
Saint-Laurent (1989), that Spongiaxius novaezealandiae does not belong in
the genus Spongiaxius, because it has distinct morphological characteristics
different from those of the type species of the genus Spongiaxius, S. brucei
(Sakai & De Saint-Laurent, 1989). In Spongiaxius novaezealandiae, the male
Plp1 is absent, and the median gastric carina originates from the middle keel
of the rostrum up to the anterior two-thirds of the gastric region, bearing in its
middle an elongate-triangular patch of granules gradually narrowing anteriorly
and becoming a single row of spines, and the uropodal exopod bears no transverse suture. In Spongiaxius brucei, however, the male Plp1 is present, bearing
a spatulate distal segment, the gastric median carina is serrated, and the uropodal exopod bears a transverse suture. Therefore, Spongiaxius novaezealandiae
is here reclassified under the new genus Newzealandaxius as N. novaezealandiae. The male type specimen of N. novaezealandiae examined, seems to be
an immature one.
Type species. — Axius (Axius) novae-zealandiae Borradaile, 1916, by
present designation and monotypy. The gender of the new generic name,
Newzealandaxius, is masculine.
Species included. — Newzealandaxius novaezealandiae (Borradaile, 1916).
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Etymology. — The new genus is named Newzealandaxius after the type
locality, New Zealand, where the type specimen was collected, with the
existing generic name Axius added.
Newzealandaxius novaezealandiae (Borradaile, 1916)
Axius (Axius) novae-zealandiae Borradaile, 1916: 91, fig. 5a-b; De Man, 1925d: 3 (list), 12
(key).
Axius (Axius) novaezealandiae — Balss, 1933: 87.
Spongiaxius novaezealandiae — Sakai & De Saint Laurent, 1989: 42, fig. 11.
Material examined. — NHM 1917.1.29.106-110, types, 1 male (TL/CL, 50.0/19.6 mm),
2 females (TL/CL, 27.0/9.7-37.0/14.0 mm), 1 damaged female, New Zealand, “Terra Nova”
Expedition.
Type locality. — Seven miles [± 13 km] east of North Cape, New Zealand,
126 m.
Distribution. — New Zealand — North Cape (Borradaile, 1916; Sakai &
De Saint Laurent, 1989); 70-720 m.
Genus Oxyrhynchaxius Parisi, 1917
Oxyrhynchaxius Parisi, 1917: 17; De Man, 1925d: 2 (key); Sakai, 1987a: 303 (list); Sakai & De
Saint Laurent, 1989: 65; Poore, 1994: 98 (key); Poore & Collins, 2009: 203.
Diagnosis. — Rostrum styliform, lateral margins denticulate and not extending onto gastric region. Carapace covered with scale-like granules. Gastric region not convex, and situated at same level as rostrum, bearing 5 almost parallel carinae. Anterolateral margins of carapace armed with a denticle
each. Cervical groove present at full length. Eyestalks elongate, overreaching
rostrum; cornea pigmented. A2 scaphocerite prominent. P1 subequal; chelae
obliquely positioned; lateral surface granulate, with setae. Pleurobranchs absent. Abdominal pleura unarmed ventrally. Male Plp1 uniramous and unsegmented; male Plp2 biramous, endopod with appendices interna and masculina;
male Plps3-5 endopods with appendix interna. Female Plp1 uniramous, consisting of proximal segment and multiarticulate flagellum. Telson longer than
wide, lateral margins with proximal lobe with spine, and posterior margin convex with median spine. Uropodal exopod with transverse suture. [Adapted
from Sakai & De Saint Laurent, 1989.]
Type species. — Oxyrhynchaxius japonicus Parisi, 1917, by original designation and monotypy. The gender of the generic name, Oxyrhynchaxius, is
masculine.
Species included. — Oxyrhynchaxius japonicus Parisi, 1917; O. manningi
Lin, Kensley & Chan, 2000; O. tricarinatus Lin, 2006.
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G ENUS OXYRHYNCHAXIUS
1 – Postcervical carina absent; abdominal somites 2-5 with distinct dorsal carina . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. manningi
– Postcervical carina present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Abdominal somites 3-5 with 3 distinct longitudinal dorsal carinae . . . . . O. tricarinatus
– Abdominal somites lacking distinct longitudinal dorsal carina . . . . . . . . . . . O. japonicus
Oxyrhynchaxius japonicus Parisi, 1917
Oxyrhynchaxius japonicus Parisi, 1917: 18, figs. 5, 6; De Man, 1925d: 7 (list); Froglia & Grippa,
1986: 260, fig. 4; Sakai, 1987a: 303 (list); Sakai & De Saint Laurent, 1989: 65; Komai,
2000b: 345; Lin et al., 2000: 199, figs. 1, 2, 3a-c.
Material examined. — MNHN Th 428, 1 female (TL/CL, 69.0/25.5 mm), Hong Kong.
Diagnosis. — Female Plp1 uniramous, consisting of proximal segment and
multiarticulate flagellum; female Plps2-5 endopods with appendix interna.
Type locality. — Enoshima, Sagami Bay, Japan.
Distribution. — Japan — Sagami Bay (Parisi, 1917); Hong Kong (Sakai &
De Saint Laurent, 1989).
Oxyrhynchaxius manningi Lin, Kensley & Chan, 2000
Oxyrhynchaxius manningi Lin et al., 2000: 199-206, figs. 3c, d, f, 4; Davie, 2002: 203; Poore
& Collins, 2009: 266.
Remarks. — Only the female type specimens are known.
Type locality. — North West Shelf between Port Hedland and Dampier,
Western Australia, 134 m.
Distribution. — Only known from the type locality.
Oxyrhynchaxius tricarinatus Lin, 2006
Oxyrhynchaxius tricarinatus Lin, 2006: 238, figs. 3, 4, table 2.
Material examined. — MNHN Th 1467, holotype, male (TL/CL, 53.0/19.0 mm, dissected),
New Caledonia, 4, 20◦ 16.68′ S 163◦ 50.26′ E, BATHUS (French expedition), 500-600 m,
3.viii.1994.
Diagnosis. — Male Plp1 missing; male Plp2 endopod with appendices
interna and masculina; male Plps3-5 endopods with appendix interna.
Remarks. — Male Plp1 of the male holotype (MNHN TH 1467) is missing,
but it is described as rudimentary, as shown in the figure (Lin, 2000: 238,
fig. 4G).
Type locality. — New Caledonia, [Sta.] 4, 20◦ 16.68′ S 163◦ 50.26′ E, 500600 m.
Distribution. — Only known from the type locality.
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Genus Parascytoleptus Sakai & De Saint Laurent, 1989
Parascytoleptus Sakai & De Saint Laurent, 1989: 39; Sakai, 1992b: 212.
Diagnosis. — Rostrum triangular and short, barely reaching distal end of
eyes, tuberculiform tooth on either side. Lateral margins extending posteriorly
onto gastric region as lateral carinae. Gastric region steeply elevated from rostrum, bearing median carina reaching cervical groove, but no submedian carinae. Anterolateral margin of carapace unarmed. Cervical groove discernible
only in dorsal part. Eyestalks short and subglobose; cornea pigmented. A1
peduncles reaching proximal third of A2 penultimate segment. A2 scaphocerite reduced. P1 chelate and distinctly unequal; palm bearing no dorsodistal spine. P2 chelate, fingers shorter than palm. Ps3-4 propodi with transverse
rows of spines. Pleurobranchs present on Plps2-4. Abdominal somites smooth;
terga 2-5 with 1-2 tufts of long setae, and pleura truncate ventrally. Male Plp1
present as small unsegmented protrusion; male Plp2 biramous, endopod bearing appendix interna, but no appendix masculina; male Plps3-5 endopods with
appendix interna. Female Plp1 uniramous, slender, and composed of proximal segment and multiarticulate flagellum; female Plps2-5 slender and biramous, endopods with appendix interna. Telson triangular and convergent towards rounded posterior margin, without median spine, bearing proximal lobe
with spine followed distally by another two spines on lateral margin. Uropodal endopod elongate and oval, bearing distolateral spine, and median carina
with spine on posterior margin. Uropodal exopod with two lateral spines, but
without distal flap dorsally. [Adapted from Sakai & de Saint Laurent, 1989.]
Remarks. — The genus Parascytoleptus is characterized by the male
pleopods; male Plp1 is present as a small, unsegmented protrusion, male Plp2
is biramous, and the endopod bears an appendix interna but no appendix
masculina; male Plps3-5 endopods bear an appendix interna. This genus is
similar to the genus Scytoleptus in that the gastric region is elevated.
Type species. — Paraxius tridens Rathbun, 1906, by monotypy. The gender
of the generic name, Parascytoleptus, is masculine.
Species included. — Parascytoleptus tridens (Rathbun, 1906).
Parascytoleptus tridens (Rathbun, 1906)
(fig. 27A-B)
Paraxius tridens Rathbun, 1906: 895, fig. 53; Sendler, 1923: 44.
Axius (Paraxius) tridens — De Man, 1925d: 5 (list), 18 (key).
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Fig. 27. Parascytoleptus tridens (Rathbun, 1906) and Paraxiopsis brocki (De Man, 1888b).
A, carapace, dorsal view; B, abdomen, pleopods 2-5, and tail-fan, lateral view; C, male Plp1 on
left side. A-B, Parascytoleptus tridens, SMF 20361, male (TL/CL, 9.0/3.8 mm), Bora Bora,
French Polynesia, riff-edge by Baitape, 0.5-1 m; C, Paraxiopsis brocki, AM P 6824, male
(TL/CL, 45.0/13.5 mm), rail pier, Port Darwin, Northern Territory, Australia. Scales 1 mm.
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Parascytoleptus tridens — Sakai & De Saint Laurent, 1989: 40, fig. 10; Sakai, 1992b: 212,
fig. 2; Kensley, 2003: 370.
Material examined. — SMF 4936, 1 female (TL/CL, 24.0/6.9 mm, lacking small cheliped),
Makatea, Paumotu-Island, 10-20.vii.1909, Hanseatische Süde-Expedition, leg. E. Wolf (Lit.
Sendler, 1923: 44, as Paraxius tridens); SMF 20358, 2 males (TL/CL, 8.0/3.1-12.0/4.3 mm),
ca. 2.6 km W. of airport, Moorea, French Polynesia, Bareer-reef at Maharepa, 0.5 m, dead
coral, iii.1988, leg. H.G. Müller; SMF 20359, 1 male (TL/CL, 10.0/4.2 mm), 1 female (TL/CL,
12.0/4.8 mm), ca. 2.6 km W. of airport, Moorea, French Polynesia, Bareer-reef at Maharepa,
0.5 m, dead coral, iii.1988, leg. H.G. Müller; SMF 20360, 1 ovig. female (TL/CL, 22.0/6.9 mm),
1 juv., Moorea, French Polynesia, riff-edge by Afareaitu, 1-2 m, dead coral, 26.iii.1988, leg.
H.G. Müller; SMF 20361, 2 males (TL/CL, 13.0/5.3; 9.0/3.8 mm), Bora Bora, French Polynesia,
riff-edge by Baitape, 0.5-1 m, dead coral, 27.ii.-06.iii.1988, leg. H.G. Müller.
Diagnosis. — Rostrum small, short, triangular, and concave on dorsal surface, lateral margins each with distinct proximal tooth, extending posteriorly
onto gastric region (fig. 27A). Gastric region steeply elevated from rostrum;
median carina entire, originating from proximal part of rostrum and running
to cervical groove; submedian carinae absent. Anterolateral margins of carapace unarmed. Cervical groove discernible only in dorsal part. Eyestalks short
and subglobose; cornea pigmented. A1 peduncles reaching proximal third of
A2 penultimate segment. A2 scaphocerite reduced. P1 chelate and unequal.
In male larger cheliped ischium unarmed; merus unarmed on dorsal margin,
and unarmed or armed with some obscure denticles on ventral margin; carpus broad and two-thirds as wide as long; palm broad and more than twice as
long as carpus, bearing obtuse dorsodistal spine, and obtuse subdorsal tooth
on distal margin. Fixed finger short, cutting edge slightly convex in proximal
half, bearing 2-3 obtuse denticles. Dactylus thick and curved downward, and
unarmed on cutting edge. In smaller cheliped carpus as long as wide, and palm
twice as long as carpus. Dactylus and fixed finger slender, about 2/3 length of
palm, cutting edges of dactylus and fixed finger with spinules. P2 chelate and
unarmed. Ps3-4 propodi with transverse rows of spines. Pleurobranch present
on Ps2-4.
Abdominal pleuron 1 triangular ventrally, pleura 2-5 truncate ventrally with
some marginal setae (fig. 27B). Male Plp1 present as small, unsegmented
protrusion (SMF 20359, male; SMF 20361, larger male), Plp2 slender and
biramous, endopod bearing appendix interna but no appendix masculina;
Plps3-5 biramous, endopods with appendix interna. Female Plp1 uniramous,
composed of proximal segment and multiarticulate flagellum; female Plps2-5
slender and biramous, endopods with appendix interna.
Type locality. — French Frigate Shoal, Hawaiian Islands, 36-59 m.
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Distribution. — Madagascar (Sakai & De Saint Laurent, 1989); Hawaiian
Islands — French Frigate Shoal and off Modu Manu (Rathbun, 1906); Society
Islands — Moorea and Bora Bora (Sakai, 1992b); Tuamotu Archipelago —
Makatea; 36-59 m.
Genus Paratrichocheles gen. nov.
Diagnosis. — Rostrum acutely triangular with pointed apex, located at level
lower than that of anterior carapace, lateral margins each armed with 1-2 teeth
proximally, and spinule or none at proximal third, extending posteriorly onto
anterior half of gastric region as lateral carinae; gastric region convex, bearing
short smooth median carina. Hepatic tubercle present. Anterolateral margin of
carapace unarmed. Cervical groove distinct. Postcervical carina absent. Posterior margin of carapace with lateral lobes. Eyestalks subglobose; cornea pigmented. A1 peduncle with slightly inflated basal segment; segments 2-3 short.
A2 scaphocerite short and bifurcate. Mxp3 pediform, ischium with strongly
dentate crest on inner surface; merus dentate on mesial margin; exopod slender. P1 chelate and symmetrical. P2 chelate, fingers shorter than palm; Ps3-5
unknown. Abdominal somites 1-5 smooth, pleura rounded, bearing no tuft of
setae ventrally. Male Plp1 absent; male Plp2 biramous, endopod bearing appendix interna, but no appendix masculina; male Plps3-5 biramous, endopods
lacking appendix interna. Female Plp1 uniramous and bisegmented; Plps2-5
biramous, endopods without appendix interna. Telson longer than wide, bearing proximal lobe with spine followed posteriorly by 1-2 teeth on lateral margin, and two posteriorly-divergent carinae with single and double spines originating from two paired spines centrally on dorsal surface; posterior margin
without median spine. Uropodal endopod broad, bearing sharp spine at posterolateral angle and median carina with spines dorsally. Uropodal exopod
bearing movable spine at distolateral angle, and denticulate transverse suture
dorsally. [Adapted from Sakai, 1994.]
Remarks. — Eutrichocheles pumilus Sakai, 1994 was treated as one of
the species included in the genus Eutrichocheles in an earlier identification,
but this species has turned out to be clearly different from the type species
of Eutrichocheles, E. modestus (Herbst, 1796), as follows. In Eutrichocheles
pumilus Sakai, 1994 from Port Essington, Cobourg Peninsula, NT, the gastric
region bears no submedian rows of denticles; male Plp1 is absent; and the male
Plp2 endopod bears an appendix interna, but no appendix masculina. In the
type species of Eutrichocheles, by contrast, the gastric region bears submedian
rows of denticles; the male Plp1 is present as a simple protrusion; and the male
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Plp2 endopod bears an appendix masculina, but no appendix interna. For those
reasons, E. pumilus is to be excluded from Eutrichocheles, and reclassified
under the new genus Paratrichocheles gen. nov., based on E. pumilus as the
type species.
Type species. — Eutrichocheles pumilus Sakai, 1994, by present designation and monotypy. The gender of the name, Paratrichocheles, is masculine.
Species included. — Paratrichocheles pumilus (Sakai, 1994).
Etymology. — The new genus is named Paratrichocheles by replacing
the Greek prefix “eu” meaning “good or true” with the Greek prefix “para”
meaning “near”.
Paratrichocheles pumilus (Sakai, 1994)
Eutrichocheles pumilus Sakai, 1994: 188, 200 (key), figs. 8, 9.
Paraxiopsis pumilus — Kensley, 2003, table 2; Animal Diversity Web, Univ. Michigan; Poore
& Collins, 2009: 266, figs. 30, 31, 44.
Paraxiopsis dianae Poore, 2008: 165, fig 2. [Type locality. — Dampier Archipelago, Western
Australia, 6.0-14.0 m.]
Remarks. — The diagnosis is given under the genus Paratrichocheles.
Type locality. — Port Essington, Cobourg Peninsula, Australia, 11◦ 21.5′ S
132◦ 13′ E.
Distribution. — Australia — Port Essington and North West Shelf (Sakai,
1994); off Barrow Is.; Port George IV, Bonaparte Archipelago, Western
Australia; Port Essington, west side of Barrow Bay; Northern Territory;
Arafura Sea (Poore & Collins, 2009).
Genus Paraxiopsis De Man, 1905
Axiopsis (Paraxiopsis) [s. str.] De Man, 1905: 597; Poore & Griffin, 1979: 221 (key); Sakai &
De Saint Laurent, 1989: 3, 51; Poore, 2008: 164.
Axiopsis (Paraxiopsis) — De Man, 1925d: 2 (key), 71, 101; Gurney, 1942: 240; Balss, 1957:
1579; Sakai, 1987a: 303 (list).
Paraxiopsis — Kensley, 1996c: 709; Kensley, 2003: 372; Poore, 2004: 175. [Type species:
Axius Brocki De Man, 1888b.]
Diagnosis. — Rostrum acutely triangular and furrowed on dorsal surface;
lateral margins with denticles followed by surpaorbital spine, extending posteriorly onto gastric region as lateral carinae. Gastric region convex; median
carina smooth with hepatic tubercle; submedian carinae smooth and lateral
carinae with 1-2 anterior teeth; anterolateral margin of carapace usually armed
with a spine. Cervical groove distinct at full length. Postcervical carina absent.
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157
Posterior margin of carapace with a pair of lateral lobes. Eyestalks subglobose; cornea pigmented. A1 peduncle with slightly inflated proximal segment;
segments 2-3 short. A2 scaphocerite short and simple, and bi- or trifurcate.
Mxp3 pediform, exopod slender; ischium with denticulate crest on inner surface; merus with teeth on mesial margin. P1 chelate and asymmetrical, fingers
with wide gape. P2 chelate. Ps3-4 simple, propodi with transverse rows of
spines ventrolaterally. P5 simple or subchelate, propodus with setae on distoventral margin. Epipod present. Pleurobranchs absent.
Abdominal somites 1-5 smooth laterally, pleura 2-5 broadly rounded or
triangular ventrally. Male Plp1 uniramous, unsegmented or bisegmented; male
Plp2 biramous, endopod bearing appendix masculina with distal setae, but
no appendix interna; male Plps3-5 biramous, endopods without appendix
interna. Telson longer than wide, lateral margins bearing proximal lobe with
spine; dorsal surface with divergent carinae with spines in posterior two-thirds,
posterior margin with median spine. Uropodal endopod broad, bearing spine
posterolaterally and median carina with spinules on dorsal surface. Uropodal
exopod bearing distinct, movable spine at distolateral angle, and denticulate
transverse suture dorsally.
Type species. — Axiopsis (Paraxiopsis) brocki (De Man, 1888b), by original
designation. The gender of the generic name, Paraxiopsis, is feminine.
Species included. — Paraxiopsis brocki (De Man, 1888b); P. gracilimana
Kensley, 1996c; P. hispida Kensley, 1996c; P. johnstoni Edmondson, 1925; P.
majuro Kensley, 2003; P. paulayi (Kensley, 2003); P. pindatyba (Rodrigues &
Kensley, 1991); P. plumosimanus (Kensley, 2003); P. vicina Coelho & RamosPorto, 1985; P. sp., Robles et al., 2009.
K EY
TO THE
S PECIES
OF THE
G ENUS PARAXIOPSIS
1 – P2 fingers longer than palm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. johnstoni
– P2 fingers shorter than palm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Pleura 2-5 triangular ventrally, and pleura 3-5 armed with anterolateral spinule . . . . . . 3
– Pleura 2-5 rounded ventrally and unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 – Carapace smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. paulayi
– Carapace finely pitted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. majuro
4 – Uropodal endopod smooth or 2-3 spinules on lateral margin . . . . . . . . . . . . . . . . . . . . . . . 5
– Uropodal endopod serrate on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 – P1 palm with denticulate ventral plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. vicina
– P1 palm without denticulate ventral plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. brocki
6 – Carapace scattered with short setae; P1 chela with granules . . . . . . . . . . . . . . . . . . . . . . . . 7
– Carapace smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
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CRM 013 – Katsushi Sakai
7 – Gastric submedian carinae entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. plumosimanus
– Gastric submedian carinae with 2-3 small tubercles . . . . . . . . . . . . . . . . . . . . . . P. hispidus
8 – P1 propodus with numerous low tubercles, and finely serrate on ventral margin . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. gracilimana
– P1 propodus roughly tuberculate and distinctly serrate on ventral margin . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pindatyba
Paraxiopsis brocki (De Man, 1888)
(fig. 27C)
Axius Brocki De Man, 1888b: 475, pl. 20 fig. 3.
?Axiopsis Brocki — Borradaile, 1903: 539.
Axiopsis (Paraxiopsis) Brocki — De Man, 1905: 597; De Man, 1925d: 7 (list), 71 (key) 101,
pl. 8 fig. 19-19f.
Axiopsis (Paraxiopsis) brocki — Poore & Griffin, 1979: 228, fig 3; Tirmizi, 1983: 88, fig. 3;
Sakai, 1987a: 304 (list); Morgan, 1990: 6, 63.
Eutrichocheles brocki — Sakai & De Saint Laurent, 1989: 3, 8, 52, fig. 4B; Sakai, 1992b: 215;
Sakai, 1994: 185, 200 (key); Ngoc-Ho, 1998: 365, fig. 1.
Eutrichocheles aff. brocki — Sakai, 1992a: 168.
Paraxiopsis brocki — Kensley, 1996c: 712, figs. 1, 2; Kensley, 2003, table 2; Poore, 2004: 176;
Poore & Collins, 2009: 266.
Material examined. — ZMB 20454, 5 males (TL/CL, 11.0/3.3-21.0/6.3 mm); 1 ovig.
female (TL/CL, 20.0/6.7 mm); 1 female (TL/CL, 16.0/5.1 mm), Piatuki, Buka Island, German
New Guinea, biotope, 23.x.1909, leg. Schoede, det. De Man; SMF 16569, 1 male (TL/CL,
17.0/5.3 mm), Amboina, Moluccas, 1891, leg. Strubell; SMF 20362, 1 ovig. female (TL/CL,
20.0/6.6 mm), 2 males (TL/CL, 14.0/4.0; 12.0/3.9 mm), Bora Bora, French Polynesia, 0.51 m, in dead stone coral, 27.ii.-06.iii.1988, leg. H.G. Müller; SMF 20363, 1 juvenile (TL/CL,
6.0/2.3 mm), Bora Bora, French Polynesia, 1-1.5 m, in dead stone coral, 07.iii.1988, leg.
H.G. Müller; RMNH D1167, 1 male (TL/CL, 36.0/8.5 mm), 0◦ 7.2′ N 130◦′ 25.5′ E, Singapore
Expedition vii.1929; NHM 189311.3.74-75, 2 males (TL/CL, 23.0/8.4-21.0/7.0 mm), 1 female
(TL/CL, 33.0/10.6 mm), Macclesfield Bank, South China Sea, 25-40 fms [ca. 40-72 m], H.M.S.
“Egeria”, leg. Bassett-Smith; AM P20357, 1 female (TL/CL, 94.0/31.6 mm), Waigait Reef,
Darwin, Northern Territory, Australia; AM P 6824, 1 male (TL/CL, 45.0/13.5 mm), Rail Pier,
Port Darwin, Northern Territory, Australia.
Diagnosis. — Carapace smooth. Rostrum acutely triangular and concave on
dorsal surface, lateral margins with 4-6 spines followed by supraorbital spine,
extending posteriorly onto gastric region as lateral carinae with 1-2 spines.
Gastric region convex, with 5 distinct carinae; median carina originating from
base of rostrum, extending to posterior third, bearing thick hepatic tubercle;
submedian carinae with tooth at anterior end. Anterolateral margins of carapace with distinct spine just above A2 basis. A1 peduncle barely reaching distal margin of A2 penultimate segment; A2 segment 2 with dorsodistal spine;
scaphocerite bifurcate; segment 3 with distoventral spine. Eyestalks subglobose, less than half length of rostrum. Mxp3 ischium with toothed crest on
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159
mesial margin, merus with 2-3 large distinct spines and several smaller proximal spines on mesial margin, propodus with distoventral spine.
Pl subequal, not sexually dimorphic. P1 ischium with 3-4 spines on ventral
margin; merus with spine on dorsal margin and 3-4 sharp spines on ventral
margin; carpus short, bearing 2-3 tubercles on ventral margin; palm about 1.5
times as long as dactylus, bearing subdistal spine on dorsal margin; cutting
edge of fixed finger denticulate, bearing triangular convexity at proximal
third; dactylus as long as fixed finger; cutting edge concave at proximal
third, denticulate in distal two-thirds, and curved downward distally. P2
chelate; ischium and merus with distinct teeth on ventral margin; palm slightly
longer than dactylus. Ps3-5 propodi with numerous small, articulate spines
ventrolaterally.
Abdominal somite 1 narrow and protruded ventrally, with rounded tip,
somite 2 smooth with broadened pleuron, somites 3-4 smooth with rounded
pleura ventrally, somite 5 smooth with slightly truncate pleuron, and somite 6
smooth and concave on posterior half of ventral margin. Male Plp1 present
as slender, unsegmented or bisegmented protrusion (fig. 27C); male Plp2
bearing appendix masculina with distal setae, but no appendix interna; Plps35 biramous, endopods lacking appendix interna. Female Plp1 uniramous,
consisting of proximal segment and multiarticulate flagellum about three times
as long as proximal segment; Plps2-5 biramous, endopods lacking appendix
interna.
Telson longer than wide and parallel-sided, bearing proximal lobe with
spine followed posteriorly by 1-3 spines on lateral margin, strong, movable
spine posterolaterally, and posteriorly-divergent carinae each with 3-4 spines
in posterior two-thirds of dorsal surface; posterior margin rounded with median
spine. Uropodal endopod smooth laterally, bearing spine at posterolateral
angle, and median longitudinal carina with 5-8 spines extending to posterior
margin on dorsal surface. Uropodal exopod bearing 2-5 distal spines on lateral
margin, distinct, movable spine at distolateral angle, and denticulate transverse
suture dorsally. Hermaphroditic, genital pores present on Ps3 and 5 coxae.
Remarks. — Paraxiopsis brocki is characterized as shown above, but it is
found by examining and comparing the specimens with one another that this
species has some variation in the characteristics of its abdominal pleura and
male Plp1: (1) abdominal pleura 3 and 4 bear a denticle at the anteroventral
angle, and abdominal pleuron 5 also bears a denticle at the posteroventral
angle (NHM 1893.11.3.74-75, 2 males and 1 female from Macclesfield Bank,
South China Sea; USNM 95562, 1 female from Bikini Atoll; from Kensley,
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CRM 013 – Katsushi Sakai
1996c, fig. 1), while the abdominal pleura lack such denticles (SMF 20362,
1 ovig. female, 2 males from Bora Bora, French Polynesia; MNHN Th.
1313, 1 female; from Ngoc-Ho, 1998, fig. 1); (2) male Plp1 is present as
a short conical projection (ZMB 20454, 1 male, TL/CL, 21.0/6.3 mm, from
Buka Island, former German New Guinea; RMNH 1167, 1 male, TL/CL,
36.0/8.5 mm, from Singapore), but it is absent in 3 males from Buka Island; or
it is a slender simple protrusion tapering distally (SMF 16569, 1 male, TL/CL,
17.0/5.3 mm, from Amboina); or it is a slender, bisegmented protrusion (AM
P 6824, 1 male, TL/CL, 45.0/13.5 mm, Rail Pier, Port Darwin, Northern
Territory, Australia) (fig. 27C); or it is absent (SMF 20362, 1 male, TL/CL,
14.0/4.0 mm; 1 male, 12.0/3.9 mm, from Bora Bora, French Polynesia). The
result of the present examination and comparison of the specimens shows,
that whether the pleura are armed with denticles or unarmed, is not to be
considered a valid character to distinguish one species from the other, but
instead considered as intraspecific variation, and male Plp1 is absent in the
smaller specimens, whereas in the larger ones, it is present and varies in shape
from a simple conical protrusion to a slender bisegmented protrusion.
Type locality. — Ambon, Indonesia.
Distribution. — Kenya — off Mombasa (Sakai, 1992a); Indonesia —
Ambon and Edam Island, Jakarta (De Man, 1888b), Madura Bay, Borneo
Bank, anchorage off Beo, and north of Waigeo Island (De Man, 1925d);
Papua New Guinea — Buka Island; Timor Sea — Ashmore Reef (Sakai,
1994); Australia — Darwin, Broome, Point Peron, Geladton, Mullaloo Beach,
Rottnest Island, Cockburn Sound near Fremantle, Garden Island, Safety Bay,
and Augusta (Poore & Griffin, 1979), Darwin (Kensley, 1996c); Marshall
Islands — Bikini Atoll (Kensley, 1996c); Society Islands — Bora Bora (Sakai,
1992b); Tuamotu Archipelago — Fangataufa (Ngoc-Ho, 1998); reef to 91 m;
Japan — Okinawa (Sakai & De Saint Laurent, 1989).
Paraxiopsis gracilimana Kensley, 1996
Paraxiopsis gracilimana Kensley, 1996c: 719, figs. 6, 7; Heard et al., 2007: 19, fig. 14.
Material examined. — RMNH D 41747, 1 female (TL/CL, 16.0/5.5 mm), Pillsbury Sta. P1208 West Atlantic, Jamaica, 17◦ 45.5′ N 76◦ 59.1′ W, 15 m, 10′ ottertrawl, 5.vii.1970, USNM
Acc. No. 381702.
Type locality. — Bonaire Harbor, Netherlands Antilles.
Distribution. — Off South Carolina (Kensley, 1996c); Florida — Looe
Key reef and off Panama City (Kensley, 1996c); Gulf of Mexico (Kensley,
1996c); Netherlands Antilles — Bonaire Harbor (Kensley, 1996c); Tobago –
Charlottesville (Kensley, 1996c); Belize — Carrie Bow Cay (Kensley, 1996c).
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161
Paraxiopsis hispida Kensley, 1996
Paraxiopsis hispida Kensley, 1996c: 724, fig. 9.
Remarks. — Hermaphroditic species (Kensley, 1996c).
Type locality. — Off Yucatan Peninsula, 15◦ 59.2′ N 86◦ 02′ W, 35-37 m.
Distribution. — Off Yucatan Peninsula (Kensley, 1996c); Belize — Carrie
Bow Cay (Kensley, 1996c); 24-37 m.
Paraxiopsis johnstoni Edmondson, 1925
Axiopsis (Paraxiopsis) johnstoni Edmondson, 1925: 20, fig. 4, pl. 1A.
Eutrichocheles johnstoni — Sakai & De Saint Laurent, 1989: 54.
Paraxiopsis johnstoni — Kensley, 2003, table 2.
Type locality. — John Stone Island, Hawaiian Islands.
Distribution. — Hawaiian Islands — John Stone Island (Edmondson, 1925).
Paraxiopsis majuro Kensley, 2003
Paraxiopsis majuro Kensley, 2003: 380, figs. 9, 10, pl. 7, table 2.
Type locality. — Majuro Island, Marshall Islands, patch reef in north-northeast lagoon, under rock, 10 m.
Distribution. — Only known from the type locality.
Paraxiopsis paulayi Kensley, 2003
Paraxiopsis paulayi Kensley, 2003: 372, figs. 5, 6, pl. 3, table 2.
Material examined. — USNM 296395, holotype, hermaphrodite (TL/CL, 36.0/10.5 mm),
on reef flat, Luminao Reef, Guam, 10 m, 21.v.1999, leg. L. Kirkendale, det. B. Kensley.
Diagnosis of holotype specimen (USNM 296395). — Rostrum sharply
triangular, with proximal spine on lateral margin. Gastric region with 5
carinae; smooth median carina with hepatic tubercle starting at rostral base,
smooth submedian carinae with anterior spine, and lateral carinae with two
spines including supraorbital one. A2 scaphocerite short and simple. P1 merus
bearing two teeth on dorsal margin, and 4 interspaced, sharp teeth on ventral
margin, carpus and palm tuberculate on lateral surface, palm tuberculate on
dorsal and ventral margins, bearing distodorsal tooth and another tooth on
distal margin. Cutting edge of fixed finger denticulate, and cutting edge of
dactylus concave proximally and denticulate toward distal end. Genital pores
present on Ps3 and 5 coxae. Plp1 present as a short, simple protrusion tapering
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CRM 013 – Katsushi Sakai
distally. Plp2 biramous, endopod on left side without appendix interna. Plps3-5
biramous, endopods without appendix interna. Hermaphroditic.
Remarks. — The present re-examination of the holotype specimen shows
that Plp2 on the right side is missing, and Plp2 on the left side is biramous,
and the endopod bears neither an appedix interna nor an appendix masculina.
Considering that in Paraxiopsis the male Plp2 endopod bears an appendix
masculina but no appendix interna, it may safely be said that the holotype
specimen of P. paulayi was damaged and lost Plp2 on the right side and the
appendix masculina of the Plp2 endopod on the left side.
Type locality. — Luminao Reef flat, Guam, 10 m.
Distribution. — Guam, Luminao Reef flat, 10 m; Double Reef, 3-4 m;
south-east of Orote Peninsula, under rock.
Paraxiopsis pindatyba (Rodrigues & Kensley, 1991)
Eutrichocheles pindatyba Rodrigues & Kensley, 1991: 556, figs. 1-16, table 1; Rodrigues et al.,
1998: 379; Melo, 1999: 322, figs. 217-218.
Remarks. — In P. pindatyba (Rodrigues & Kensley, 1991), the submedian
carinae of the gastric region are smooth, the Ps1-2 fingers are shorter than the
palm, and abdominal somites 1-6 are smooth laterally as in Paraxiopsis brocki
(De Man, 1888), though pleura 2-3 are truncate ventrally, and pleura 4-5 are
triangular ventrally as in the species of Eutrichocheles.
Type locality. — Santa Cruz, Espirito Santo, Brazil, 19◦ 57′ S 40◦ 08′ W,
intertidal, under stones.
Distribution. — Only known from the type locality.
Paraxiopsis plumosimanus Kensley, 2003
Paraxiopsis plumosimanus Kensley, 2003: 376, figs. 7, 8, table 2.
Remarks. — Kensley mentioned in the remarks (Kensley, 2003: 377) that
“The present specimen has both male and female genital orifices, but pleopod
1 is present, while pleopod 2 lacks an appendix masculina. The specimen is
thus regarded as a female”. However, the presence or absence of an appendix
interna and an appendix masculina are generic features and have nothing to
do with the definition of the sex of specimens. So, the present species is a
hermaphroditic species.
Type locality. — Apra Harbour, Sasa Bay, Guam, sand bottom, 3-5 m.
Distribution. — Only known from the type locality.
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163
Paraxiopsis vicina Coelho & Ramos-Porto, 1985
(fig. 28)
Axiopsis (Paraxiopsis) vicina Coelho & Ramos-Porto, 1985: 79, fig. 6; Melo, 1999: 324, figs.
219-220.
Material examined. — Holotype, male TL/CL, 23.0/7.9 mm, 2◦ 13′ S 40◦ 43′ W, Brazil.
Diagnosis. — Rostrum sharply triangular with 5 spines on each of the lateral
margins, which extend posteriorly onto anterior half of gastric region as lateral
Fig. 28. Paraxiopsis vicina Coelho & Ramos-Porto, 1985. A, carapace, dorsal view; B, carapace, lateral view; C, cheliped on left side, lateral view; D, cheliped on right side, lateral view;
E, abdomen and tail-fan, lateral view. Scale 1 mm.
164
CRM 013 – Katsushi Sakai
carinae. Gastric region with smooth median carina starting at rostral base and
extending to anterior half of gastric region, and smooth submedian carinae
extending also to anterior half of gastric region (fig. 28A). A2 scaphocerite
short and bilobed distally (fig. 28B). P1 subequal (fig. 28C, D); merus bearing
subdistal tooth on dorsal margin, and 4 interspaced, sharp teeth on ventral
margin, carpus denticulate on ventral margin, palm tuberculate on dorsal
margin, bearing some tubercles on lateral surface and tuberculate plate on
ventral margin. P1 fixed finger denticulate on whole cutting edge, concave
proximally, bearing triangular convexity at proximal fourth followed distally
by irregular denticulation towards distal end; dactylus convex and smooth
proximally on cutting edge, bearing triangular convexity at proximal third
followed distally by denticulation towards distal end. Abdominal somites
smooth on lateral surface, pleura 3-5 bearing spine at anteroventral angle,
and pleuron 6 with spine at posteroventral angle (fig. 28E). Genital pores
present on P3. Male Plp1 present as a small, unsegmented protrusion; male
Plp2 biramous, endopod proximally bearing appendix masculina without distal
setae, but no appendix interna; Plps3-5 biramous, endopods lacking appendix
interna. Uropodal exopod with distal flap (fig. 28E).
Remarks. — The present species, P. vicina from Brazil is closely similar to
P. brocki from the Indo-West Pacific region, but differs in that in P. vicina
the male P1 palm is distinctly tuberculate on the dorsal margin, bearing a
tuberculate plate on the ventral margin, whereas in P. brocki it is slightly
tuberculate on the dorsal margin (Kensley, 1996c), bearing no tuberculate plate
on the ventral margin.
Type locality. — Brazil, 2◦ 13′ S 40◦ 43′ W.
Distribution. — Only known from the type locality.
Paraxiopsis sp. Robles et al., 2009
Paraxiopsis sp., Robles et al., 2009: 316.
Genus Paraxius Bate, 1888
Paraxius Bate, 1888: 37; Stebbing, 1893: 188; Sakai & De Saint Laurent, 1989: 34; Poore,
1994: 98 (key).
Paraxius s. str. — Borradaile, 1903: 538; De Man, 1925d: 1 (key), 18 (key); Balss, 1957: 1579.
Diagnosis. — Rostrum short, triangular, and concave dorsally, lateral margins armed with teeth, posteriorly extending shortly onto gastric region.
Anterolateral margins of carapace unarmed. Gastric region convex, bearing
median carina with hepatic tubercle in anterior two-thirds, and short, obscure
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165
lateral carinae, but no submedian carinae. Cervical groove distinct only on dorsal surface. Eyes fixed on conical peduncles, cornea faded. A2 scaphocerite
short and directed outward. P1 unequal; palm of larger cheliped with dorsodistal spine. No pleurobranchs. Abdominal pleura smooth on lateral surface, and
rounded ventrally. Male Plp1 uniramous, bisegmented, and rod-shaped, distal segment shorter than proximal one; male Plp2 biramous, endopod bearing
appendix interna with patch of hooklets distally, but no appendix masculina.
Plps3-5 endopods with appendix interna. Telson longer than wide, serrate on
lateral margins, and slightly convergent toward rounded posterior margin without median spine; dorsal surface with two pairs of spines medially. Uropodal
endopod with two spines laterally. Uropodal exopod with transverse suture.
Deep-water habitat. [Adapted from Borradaile, 1903 and Sakai & De Saint
Laurent, 1989.]
Type species. — Paraxius altus Bate, 1888, by original designation and
monotypy. The gender of the generic name, Paraxius, is masculine.
Species included. — Paraxius altus Bate, 1888.
Paraxius altus Bate, 1888
(fig. 29)
Paraxius altus Bate, 1888: 37, pl. 5 fig. 1d-z; Balss, 1925: 210; Sakai & De Saint Laurent, 1989:
35, fig. 8.
Axius (Paraxius) altus — Borradaile, 1903: 538; De Man, 1925d: 5 (list), 18 (key).
Material examined. — NHM 88.22, holotype, male (CL 8.1; Abd with telson, 19.0 mm,
damaged, P1 left missing), north of Papua, 23◦ 3′ S 144◦ 4′ E, 1070 fathoms (1956.8 m), leg.
“Challenger”; MNHN Th 996, 1 male (TL/CL, 17.0/5.5 mm), Philippines.
Diagnosis. — Rostrum (fig. 29A) short, triangular, and concave dorsally,
lateral margins armed with 4 distinct teeth, posteriorly extending shortly onto
gastric region as obscure lateral carinae. Gastric region distinctly convex, bearing short, smooth median carina in middle of dorsal surface, but no submedian
carinae. Cervical groove distinct only in dorsal part. Eyes fixed on conical peduncles, cornea faded. A1 reaching distal margin of A2 penultimate segment,
proximal segment with dorsal denticle in middle; scaphocerite minute and directed outward. P1 unequal; in cheliped on right side (fig. 29B) ischium with
subdistal spine on ventral margin; merus with subdistal spine on dorsal margin,
and denticulate with sharp subdistal spine on ventral margin; carpus short and
unarmed; palm longer than fingers, dorsal margin carinate with distal tooth;
dactylus with proximal and median teeth on cutting edge; and fixed finger with
a series of tubercles in proximal third of cutting edge, which gradulally become
166
CRM 013 – Katsushi Sakai
Fig. 29. Paraxius altus Bate, 1888. A, carapace, dorsal view; B, cheliped on right side; C, P2,
lateral view; D, P3, lateral view; E, P4, lateral view. A-E, NHM 88.22, holotype, male (CL, 8.1;
abd. with telson, 19.0 mm; damaged), Papua. Scales 1 mm.
smaller in size towards distal part. Cheliped on left side missing. P2 (fig. 29C)
fingers of chela about as long as palm. P3 propodus (fig. 29D) elongate, bearing a row of spinules on ventrolateral surface and strong ventrodistal spine.
P4 propodus (fig. 29E) unarmed on lateral surface, but bearing oblique row of
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167
setae near distal end and distoventral spine. P5 (fig. 29E) simple. Male Plp1
uniramous and bisegmented: distal segment simple, rod-shaped, and shorter
than proximal one; male Plp2 biramous, endopod bearing appendix interna
with patch of hooklets distally, but no appendix masculina; male Plps3-5 endopods with appendix interna. Telson longer than wide and denticulate on lateral margins, slightly tapering towards rounded posterior margin without median spine. Uropodal exopod with irregularly denticulate suture and movable
spine at posterolateral angle. [Adapted from Sakai & De Saint Laurent, 1989,
fig. 8.]
Remarks. — Bate (1888: 40) described Paraxius altus on the basis of
a female, but the “holotype” examined (NHM 88.22) is a male, which is
dissected into some parts. The gastric region is obscurely provided with
median and lateral carinae. Male Plp1 on the left side and Plp2 on the right
side are missing, and the exopod of Plp2 on the left side is also missing.
Type locality. — North of Papua New Guinea, 1958 m, blue mud.
Distribution. — Philippines — Sibuyan Sea (Sakai & De Saint Laurent,
1989); north of Papua New Guinea (Bate, 1888; Sakai & De Saint Laurent,
1989); 1926-1958 m.
Genus Pilbaraxius Poore & Collins, 2009
Pilbaraxius Poore & Collins, 2009: 267.
Diagnosis. — Rostrum acutely triangular, with a pair of lateral spines,
longer than eyestalk, not depressed below level of carapace, and continuous
with definite lateral carinae; supraocular spines prominent. Eyestalk cylindrical and articulating; cornea weakly pigmented. A2 scaphocerite long. Carapace smooth or tuberculate; gastric lateral carinae each with a spine; gastric
submedian carinae present, each bearing spine; gastric median carina present
as unarmed, weak ridge; cervical groove visible laterally over third distance
to anterolateral margin; postcervical carina absent. Mxp3 exopod not clearly
bent at base of flagellum. P1 slightly asymmetrical; propodus flattened; carpuspropodus unarmed and sparsely setose on dorsal margin. Ps3-5 propodi with
transverse rows of robust setae; dactyli tapering, and bearing longitudinal row
of robust setae. Pleurobranchs present above Ps2-4; podobranchs and arthrobranchs well developed; epipods present on Mxp2 to P4. Abdominal somites
with pleura 1-2 acute, pleura 3-5 acute, bearing anteroventral tooth. Male Plp1
absent; male Plp2 endopod with appendix interna, but without appendix masculina; Plps3-5 endopods with appendix interna. Telson rounded on posterior
margin, bearing fixed spines on lateral margin and robust setae at posterolat-
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CRM 013 – Katsushi Sakai
eral angle. Uropodal endopod without lateral and distolateral spines. Uropodal
exopod with transverse suture. [Adapted from Poore & Collins, 2009.]
Type species. — Pilbaraxius kariyarra Poore & Collins, 2009, by original
designation and monotypy. Gender of generic name, Pilbaraxius, masculine.
Species included. — Only known by the type species.
Pilbaraxius kariyarra Poore & Collins, 2009
Pilbaraxius kariyarra Poore & Collins, 2009: 269, figs. 32, 33, 45.
Calaxiopsis sp., Robles et al., 2009: 314.
Diagnosis. — Rostrum 0.4 times length of front-to-cervical groove, acute,
elongate, with long lateral spine anterior to supraocular spine, continuous with
definite gastric lateral carinae. Supraocular spines prominent. Eyestalk 0.3
length of rostrum; cornea weakly pigmented. A1 peduncle reaching to end of
A2 segment 4. A2 segment 1 with two small spines on distal margin; segment 2
with straight distal spine, approximately half length of segment 2; scaphocerite
simple and straight, reaching distally almost to end of segment 4; segment 3
with spine on ventral margin; segment 4 about as long as segment 2 (excluding
distal spine); segment 5 about half as long as segment 4. Carapace covered with
small tubercles; gastric lateral carinae each with a spine; gastric submedian
carinae each with a spine; gastric median carina present as ridge extending
anteriorly onto rostrum, bearing tubercle located midway between rostrum
and cervical groove. Postcervical carina absent. Mxp3 basis with spine; crista
dentata with 15 teeth; merus with two spines (one large and one small) on
ventral margin; carpus unarmed.
P1 differentiated, and propodus of larger cheliped longer and more swollen
than that of smaller cheliped. In larger (right) cheliped, coxa with spine on
ventral margin; basis unarmed on ventral margin; ischium with spine on ventral
margin; merus with hooked spine on convex dorsal margin, and spine and
obsolete tubercle laterally on ventral margin, lateral face with broad tubercle
distally and mesial face smooth; carpus tuberculate on dorsal margin, and
unarmed on ventral margin, lateral face tuberculate and mesial face smooth;
propodus with distal spine on dorsal margin, and low lateral carina on ventral
margin, lateral face tuberculate and mesial face smooth; fixed finger 1.2
times length of dorsal palm, and cutting edge with 4 large, irregular, rounded
teeth; dactylus smooth on dorsal margin, lateral and mesial faces smooth,
and cutting edge with narrow proximal notch. In smaller cheliped, coxa,
ischium, merus, and carpus as in larger cheliped; propodus similar except
mesial face with spine near gape; fixed finger about as long as dorsal palm,
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
169
cutting edge with two large triangular teeth and smaller intermediate denticles;
dactylus excavated proximaly on cutting edge. P2 unarmed; carpus slightly
shorter than chela; propodus as long as dactylus. P3 unarmed; propodus 2.2
times as long as dactylus, bearing 6 marginal robust setae (some duplicated).
P4 unarmed; propodus 2.2 times as long as dactylus, bearing 7 marginal
robust setae (some duplicated). P5 propodus subchelate and 3.9 times as
long as dactylus; fixed finger short, bearing 5 distinct robust setae; dactylus
slightly flattened. Pleurobranchs present above Ps2-4; arthrobranchs on Mxp2
(rudimentary) to P4; epipods with well developed podobranchs (with up to
10 gill filaments) on Mxp2 to P3. Abdominal pleuron 1, 2.6 times as deep
as middorsal length, bearing small ventral spine; pleuron 2 asymmetrical,
lateral length 1.3 times dorsal length, concave ventrally, bearing two distinct
ventral spines; pleura 3 and 4 tapering to ventral spine, bearing another spine
located anteroventrally; pleuron 5 tapering to ventral spine, bearing another
two spines located anteroventrally; pleuron 6 with ventral spine. Plp1 absent.
Plp2 endopod without appendix masculina, but with slender appendix interna,
and about quarter length of endopod. Telson 1.3 times as long as broad,
bearing 4 spines on lateral margin, one robust seta at posterolateral angle,
but no median spine on rounded posterior margin. Uropodal endopod 1.9
times as long as wide, bearing no lateral spines; dorsal surface with unarmed
longitudinal ridge. Uropodal exopod 1.9 times as long as wide, bearing robust
seta at posterolateral angle, but no lateral spines; dorsal surface with unarmed
longitudinal ribs and unarmed transverse suture. [Adapted from Poore &
Collins, 2009.]
Type locality. — Off Port Hedland, Western Australia, 401-405 m.
Distribution. — Only known from the type locality.
Genus Pillsburyaxius gen. nov.
Diagnosis. — Rostrum slender and rod-shaped; lateral margins armed with
2-3 spines, posteriorly extending shortly onto gastric region. Gastric region
convex, bearing 5 carinae with teeth; median carina with two teeth anterior, and
one posterior to hepatic tubercle, and running from rostral base to gastric region; submedian carinae with 4 teeth; lateral carinae with one buttressed tooth
posterior to tooth at rostral base. Anterolateral margin of carapace usually unarmed. Cervical groove distinct. Postcervical carina poorly present. Eyestalks
subglobose; cornea pigmented. A2 scaphocerite protruded forward, overreaching A2 penultimate segment. P1 asymmetrical; chelae slender in form, fingers distinctly longer than palm. Abdominal somites smooth on lateral surface, and convex to rounded ventrally; abdominal somites 4-5 bearing spinule
170
CRM 013 – Katsushi Sakai
at anteroventral angle. Male Plps1-2 unknown. Female Plp1 uniramous and
bisegmented, distal segment multiarticulate; female Plp2 biramous, endopod
bisegmented, simple proximal segment and multiarticulate distal segment with
appendix interna at base. Female Plps3-5 slender, endopods with appendix interna. Telson longer than wide, bearing proximal process with spine on lateral
margin, and two movable spines at each posterolateral angle and median spine
on posterior margin; posteriorly-divergent carinae with 2-3 spines medially on
dorsal surface. Uropodal endopod bearing two lateral spines including distinct
one at posterolateral angle, and median carina with 4 spines including distalmost one on posterior margin on dorsal surface. Uropodal exopod bearing two
spines on lateral margin and movable spine at posterolateral angle, and transverse suture with spinules. [Adapted from Kensley, 1996b.]
Remarks. — It has turned out that Acanthaxius kirkmilleri Kensley, 1996b
does not belong in Acanthaxius, because in A. kirkmilleri the rostrum is slender
and rod-shaped, the carapace bears a poor postcervical carina, the A2 scaphocerite is protruded forward, overreaching the A2 penultimate segment, and abdominal somites 4-5 bear a spinule at the anteroventral angle. Whereas in the
type species of Acanthaxius, Axiopsis (Axiopsis) pilocheira (Sakai, 1987)
the rostrum is acutely triangular, the carapace bears no postcervical carina, the
A2 scaphocerite is protruded either forward or laterally, not overreaching the
A2 penultimate segment, and abdominal somites 4-5 are unarmed ventrally.
Hence, Acanthaxius kirkmilleri Kensley, 1996b is reclassified here under the
new genus Pillsburyaxius as P. kirkmilleri (Kensley, 1996b).
Type species. — Acanthaxius kirkmilleri Kensley, 1996b by present designation and monotypy. The gender of the generic name, Pillsburyaxius, is
masculine.
Species included. — Pillsburyaxius kirkmilleri (Kensley, 1996b).
Etymology. — The new genus Pillsburyaxius is derived from the name of
R/V “John Elliott Pillsbury”, by which the type specimen was collected from
the Caribbean Sea.
Pillsburyaxius kirkmilleri (Kensley, 1996)
Acanthaxius kirkmilleri Kensley, 1996b: 71, figs. 1, 2.
Type locality. — Off Anguilla, Caribbean Sea, 18◦ 26.4′ N 63◦ 12.6′ W, 421439 m.
Distribution. — Only known from the type locality.
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171
Genus Planaxius Komai & Tachikawa, 2008
Planaxius Komai & Tachikawa, 2008: 22; Poore & Collins, 2009: 272.
Diagnosis. — Rostrum acutely triangular, with three teeth on each lateral
margin, extending posteriorly to short lateral carina. Eyestalks rounded distally, reaching middle of rostrum; cornea distinct and located in distal half.
Gastric region elevated dorsally, and lacking median and submedian carinae.
P1 subequal. Abdominal somites 1-6 smooth laterally; abdominal somite 2 almost straight ventrally and protruded on ventral half of posterior margin. Male
Plp1 uniramous, unsegmented, spatulate, and bilobed distally, bearing hooklets mesiomedially; male Plp2 with slender appendices interna and masculina;
Plps3-5 with appendix interna. Telson with posterolateral spines; Plps3-5 with
appendices internae. Uropodal exopod with distal flap. [Adapted from Komai
& Tachikawa, 2008.]
Remarks. — The present genus, Planaxius, is similar to Paraxius Bate, 1888
in the shapes of the rostrum and the gastric region, but they differ in that in
Planaxius the male Plp1 is unsegmented and spatulate, and the Plp2 endopod
is provided with appendices interna and masculina, whereas in Paraxius the
male Plp1 is two-segmented and rod-shaped, and the Plp2 endopod bears an
appendix interna, but no appendix masculina. Planaxius is also similar to
Levantocaris Galil & Clark, 1993 the form of the male Plps1-2, but differs
from that genus, because in Levantocaris the gastric region, with a shorter
median carina, is not as elevated as it is in Planaxius.
Type species. — Planaxius brevifrons Komai & Tachikawa, 2008, by
original designation and monotypy. Gender of the generic name, Planaxius,
masculine.
Species included. — Only the type species is known.
Planaxius brevifrons Komai & Tachikawa, 2008
Planaxius brevifrons Komai & Tachikawa, 2008: 24-29, figs. 2-6, tab. 1; Poore & Collins, 2009:
272, figs. 34, 46.
Material examined. — CBM-ZC 9226, paratype, male (TL/CL, 13.0/4.4 mm), off Kirime,
Kii-Minabe, Wakayam Pref., Japan, gill-net, 80-100 m, leg T. Komai.
Diagnosis. — Rostrum acutely triangular, with three teeth on each lateral
margin. Gastric region elevated with short lateral carinae, but without median
and submedian carinae. Mxp3 with exopod. P1 subequal. Abdominal somites
1-6 smooth laterally. Male Plp1 uniramous, unsegmented, spatulate, and
bilobed distally, bearing hooklets mesiomedially. Plp2 endopod with slender
appendices interna and masculina; Plps3-5 with appendices internae. Telson
172
CRM 013 – Katsushi Sakai
longer than wide, slightly convergent posteriorly, bearing distinct midspine
followed by 3 or 4 spinules on lateral margin and spine at posterolateral angle,
but no median tooth on slightly convex posterior margin. Uropodal exopod
with distal flap. [Adapted from Komai & Tachikawa, 2008.]
Type locality. — Off Kirime, Minabe, Kii Peninsula, Wakayama Pref., 80100 m.
Distribution. — Japan — Minabe, Kii Peninsula and Ogasawara Is., 47100 m; off Jurien Bay, Western Australia.
Genus Platyaxiopsis gen. nov.
Diagnosis. — Rostrum triangular, lateral margins spinulate, extending onto
gastric region. Gastric region slightly elevated from base of rostrum, bearing
lateral carinae with some obscure tubercles, median carina with two spinules,
running in anterior half, and two buttressed spinules between median and
lateral carinae. Eyestalks subglobose, cornea pigmented. A2 scaphocerite
prominent, peduncular segment 2 with strong dorsodistal spine. Anterolateral
margin of carapace unarmed. Abdominal somite 1 deflected posteroventrally;
somite 2 broad and unarmed ventrally; somites 3-5 subequal and unarmed
ventrally, bearing no denticle at anteroventral angle. Male Plp1 absent; male
Plp2 biramous, endopod and exopod elongate and leaf-like, endopod with
appendices interna and masculina at proximal third; male Plps3-5 endopods
with appendix interna. Telson longer than wide, bearing proximal lobe with
spine on lateral margin, and median tooth on rounded posterior margin.
Uropodal exopod without distal flap.
Remarks. — Axius (Neaxius?) odontorhynchus De Man, 1905 was included
in Spongiaxius Sakai & De Saint Laurent, 1989, but it does not belong in
Spongiaxius, because in Axius (Neaxius?) odontorhynchus the gastric region
bears lateral carinae armed with some obscure tubercles, a median carina
with two spinules running in the anterior half, and two buttressed spinules
between median and lateral carinae; abdominal somite 2 is broad and unarmed
ventrally; somites 3-5 are subequal and unarmed ventrally, bearing no denticle
at anteroventral angle; male Plp1 is absent, and the uropodal exopod lacks a
distal flap. In contrast, in the type species of Spongiaxius, S. brucei (Sakai,
1986), the gastric region is convex, bearing serrated median, submedian, and
lateral carinae; the abdominal somites are almost smooth on lateral surface
and acutely triangular ventrally; the male Plp1 is uniramous and bisegmented;
distal segment spatulate, and the uropodal exopod bears a distal flap. For those
reasons, Axius (Neaxius?) odontorhynchus is reclassified here under the new
genus Platyaxiopsis gen. nov. as P. odontorhyncha (De Man, 1905).
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
173
Type species. — Axius (Neaxius?) odontorhynchus De Man, 1905, by
present designation and monotypy. The gender of the new name, Platyaxiopsis,
is feminine.
Species included. — Platyaxiopsis odontorhyncha (De Man, 1905).
Etymology. — The new genus is named Platyaxiopsis, derived from its
similarity in appearance to the type species of Platyaxius, P. brevirostris Sakai,
1994.
Platyaxiopsis odontorhyncha (De Man, 1905)
(fig. 30)
Axius (Neaxius?) odontorhynchus De Man, 1905: 591.
Axius (Axius) odontorhynchus — De Man, 1925d: 3 (list), 12 (key), 18, pl. 1 fig. 1-1m.
Spongiaxius odontorhynchus — Sakai & De Saint Laurent, 1989: 42.
Material examined. — ZMA, lectotype, 1 male (TL/CL, 9.0/3.0 mm), “Siboga” Sta. 260,
north point of Nuhu Jaan Kei Island, 5◦ 36.5′ S 132◦ 55.2′ E, 90 m, 16 & 18.xii.1899.
Diagnosis. — Eyestalks stout, reaching tip of rostrum. Abdominal somite
1 deflected posteroventrally, somite 2 broad and twice as long as somite
1; somites 3-5 subequal and shorter than somite 2, bearing no denticle at
anteroventral angle (fig. 30). Uropodal exopod without distal flap.
Remarks. — The present species is similar to Platyaxius brevirostris Sakai,
1994 in lacking a transverse suture on the uropodal exopod.
Fig. 30. Platyaxiopsis odontorhynchus (De Man, 1905). Lectotype, 1 male (TL/CL, 9.0/
3.0 mm), “Siboga” Sta. 260, north point of Nuhu Jaan Kei Island.
174
CRM 013 – Katsushi Sakai
Type locality. — Kepulauan Kai, Indonesia, 5◦ 36.5′ S 132◦ 55.2′ E, “Siboga”
Sta. 260, 90 m.
Distribution. — Indonesia — Kepulauan Kai (De Man, 1905), Kepulauan
Kai and south coast of Timor (De Man, 1925d); 73-90 m.
Genus Platyaxius Sakai, 1994
Platyaxius Sakai, 1994: 180; Poore & Collins, 2009: 274.
Diagnosis. — Rostrum stoutly triangular, lateral margins spinulate, extending onto gastric region. Gastric region slightly elevated from base of rostrum,
bearing 5 longitudinal carinae; lateral and submedian carinae smooth, and median carina present in anterior half, bearing denticles and distinct hepatic tubercle. Anterolateral margin of carapace unarmed. Cervical groove distinct.
Eyestalk subglobose, cornea pigmented. A2 scaphocerite prominent, peduncular segment 2 with strong dorsodistal spine. P1 subequal. Ps3-4 propodi
with transverse rows of spines ventrolaterally; dactyli with longitudinal rows
of spines on lateral surface. Abdominal somite 1 deflected posteroventrally,
somite 2 broadly rounded ventrally; somites 3-5 rounded ventrally, bearing
denticle at anteroventral angle. Male Plp1 absent; male Plp2 biramous, endopod and exopod elongate and leaf-like, endopod with appendices interna and
masculina at proximal third; male Plps3-5 endopods with appendix interna. Female Plp1 uniramous and bisegmented, proximal segment and multiarticulate
flagellum; female Plps2-5 biramous, endopods with appendix interna. Telson
longer than wide, lateral margins each with proximal lobe with spine; posterior
margin rounded, without median tooth. Uropodal endopod bearing spinules on
distal half of lateral margin, serrated protrusion at posterolateral angle, and a
longitudinal row of 4 interspaced teeth medially on dorsal surface. Uropodal
exopod without distal flap.
Type species. — Platyaxius brevirostris Sakai, 1994, by original designation. Gender of generic name, Platyaxius, masculine.
Species included. — Platyaxius brevirostris Sakai, 1994; P. bardi Poore &
Collins, 2009.
K EY
TO THE
S PECIES
OF THE
G ENUS PLATYAXIUS
1 – Gastric submedian carina smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. brevirostris
– Gastric submedian carina denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bardi
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
175
Platyaxius bardi Poore & Collins, 2009
Platyaxius bardi Poore & Collins, 2009: 274, figs. 35, 36, 47.
Diagnosis. — Rostrum 0.3 times length of front-to-cervical groove, narrowly triangular, with 4 weak lateral spines anterior to supraocular spine, and
continuous with definite gastric lateral carinae. Supraocular spines similar in
size to other rostral spines. Eyestalk 0.8 length of rostrum: cornea pigmented.
A1 peduncle reaching to distal part of A2 segment 4. A2 segment 1 with two
spinules on ventrodistal margin; segment 2 with well developed distal spine,
reaching to proximal part of A2 segment 5; scaphocerite strongly curved downwards, reaching distally almost to end of segment 4; segment 3 with spine on
ventral margin; segment 4 approximately as long as segment 2 (excluding distal spine); segment 5 about two-thirds length of segment 4. Carapace smooth;
gastric lateral carina with two prominent spines in addition to supraocular
spine; gastric submedian carina with 4 spines; gastric median carina with 6
spines. Mxp3 basis with spine; ischium with spine on ventral margin; crista
dentata with numerous small, even teeth; merus with three spines on ventral
margin; carpus unarmed.
P1 differentiated, and propodus of larger cheliped longer and more swollen
than that of smaller cheliped. In larger (left) cheliped, coxa with spine on
ventral margin; basis unarmed on ventral margin; ischium with 4 small spines
and one larger on ventral margin; merus unarmed and strongly convex on
dorsal margin, bearing 10 teeth on ventral margin, lateral and mesial faces
smooth; carpus carinate on dorsal margin, and unarmed on ventral margin,
lateral and mesial faces smooth; propodus with blunt tooth on carinate dorsal
margin, and weak lateral carina along ventral margin, better defined on fixed
finger, lateral face tuberculate over dorsal and ventral distal third, and mesial
face tuberculate near base of fixed finger; fixed finger 0.6 length of dorsal
palm, cutting edge straight; dactylus smooth on margins and faces, cutting
edge with few blunt teeth. Smaller cheliped narrower than larger cheliped.
In smaller cheliped, coxa, ischium, merus and carpus, as in larger cheliped;
propodus with blunt distal tooth on carinate dorsal margin, and weak lateral
carina along ventral margin, better defined on fixed finger, lateral face with
few tubercles and mesial face with few tubercles near base of fixed finger;
fixed finger as long as dorsal palm, cutting edge with regularly spaced sharp
teeth; dactylus smooth on margins and faces, and cutting edge denticulate. P2
ischium unarmed on ventral margin; merus unarmed on ventral margin; carpus
slightly longer than chela; propodus dorsal margin 0.4 length of dactylus. P3
merus with three weak spines on ventral margin; propodus 2.9 times as long
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CRM 013 – Katsushi Sakai
as dactylus, bearing 8 transverse rows each of 4-7 robust setae; dactylus with
13 robust setae on mesial face plus unguis. P4 propodus 4.1 times as long
as dactylus, bearing 8 transverse rows each of 3-8 robust setae; dactylus with
12 robust setae on mesial face plus unguis. P5 propodus 3.5 times as long as
dactylus, and weakly subchelate, bearing distally two transverse rows each of
2 or 3 robust setae; dactylus with 6 robust setae on distal margin, and three
robust setae on mesial face.
Abdominal pleuron 1, 2.6 times as deep as middorsal length, and acute
ventrally; pleuron 2 asymmetrical, lateral length 1.2 times dorsal length, and
quadrate posteroventrally; pleura 3-5 angled posteroventrally, each bearing
small anteroventral spine; pleuron 6 with small spine on ventral margin. Plp2
appendix masculina as long as appendix interna. Plps2-5 with appendix interna
one-third length of endopod. Telson ovate, 1.1 times as long as wide, and
slightly tapering posteriorly, bearing three spines on lateral margin, robust seta
at posterolateral angle, and median spine on convex posterior margin; dorsal
surface with two oblique rows of two spines. Uropodal endopod 1.7 times as
long as wide, bearing 7 lateral spines; dorsal surface with longitudinal ridge
with 4 spines (including marginal one). Uropodal exopod 1.6 times as long
as wide, bearing 11 spines on lateral margin, and fixed spine and robust seta
at posterolateral angle; dorsal surface with two longitudinal ribs (inner rib
ending in marginal spine, outer rib with 7 spines) but without transverse suture.
[Adapted from Poore & Collins, 2009.]
Type locality. — Near Mermaid Reef, Western Australia, 178-184 m.
Distribution. — Continental slope of North-West Shelf; 178-184 m.
Platyaxius brevirostris Sakai, 1994
Platyaxius brevirostris Sakai, 1994: 181, 200 (key), figs. 4-5; Poore & Collins, 2009: 277.
Diagnosis. — Abdominal somite 1 deflected posteroventrally, somite 2
broadly rounded ventrally; somites 3-5 rounded ventrally, bearing denticle at
anteroventral angle. Male Plp1 absent; male Plp2 endopod with appendices
interna and masculina (Sakai, 1994, fig. 4f); male Plps3-5 endopods with
appendix interna. Female Plp1 uniramous and bisegmented, proximal segment
and multiarticulate flagellum; female Plps2-5 endopods with appendix interna.
Type locality. — North West Shelf, Western Australia, 18◦ 33.4′ S
118◦ 35.4′ E, 141 m.
Distribution. — Australia — North West Shelf (Sakai, 1994); 141 m.
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177
Genus Pseudoaxiopsis gen. nov.
Diagnosis. — Rostrum triangular and pointed at apex; lateral margins spinulate, extending posteriorly onto gastric region as lateral carinae. Gastric region convex, bearing 5 longitudinal carinae; median carina tuberculate, running from middle of rostrum nearly to cervical groove, and bearing short double carina with tubercles in middle; submedian carinae tuberculate; and lateral
carinae denticulate anteriorly but tuberculate posteriorly. Anterolateral margin
of carapace armed with a small spine just above A2. Cervical groove distinct.
Eyestalks subglobose; cornea pigmented. A2 scaphocerite strongly protruded
forward. P1 subequal; fingers longer than palm. P2 fingers longer than palm.
Abdominal terga 1-6 with carina in longitudinal stripes. Abdominal somite 1
narrow and pointed ventrally; somites 2-4 provided laterally with longitudinal
carina between tergum and pleuron; somites 5-6 without longitudinal carina
laterally; pleura 2-4 triangular and pleura 5-6 rounded ventrally; pleura 4-5
with spinule on anteroventral margin, and pleuron 6 with spinule ventrally.
Male Plp1 absent; male Plp2 biramous, endopod with appendices interna and
masculina at proximal third; appendix interna slightly longer than appendix
masculina with distal setae; male Plps3-5 endopods with appendix interna.
Telson longer than wide, bearing posteromedian tooth. Uropodal exopod with
transverse suture.
Remarks. — Poore & Griffin (1979: 230, fig. 4) described Axiopsis (Axiopsis) consobrina on the basis of two specimens (male and female), but their
male specimen is different from Axiopsis consobrina De Man, 1905 as follows.
In De Man’s (1905) A. consobrina the abdominal somites are smooth dorsally
and laterally, and their pleura are truncate or triangularly protruded on the ventral margin; the P1 are unequal; the fingers are shorter than the palm; the P2
fingers are shorter than the palm; and the gastric median carina is denticulate
throughout its length. In Poore & Griffin’s (1979) A. consobrina the abdominal
somites 2-4 are each provided with a longitudinal carina on the dorsal surface,
pleura 2-4 are triangular ventrally, pleura 4-5 bear a spinule on the anteroventral margin, and pleuron 6 also bears a spinule ventrally; P1 are subequal; the
fingers are longer than the palm; the P2 fingers are longer than the palm; and
the gastric median carina, running from the middle of the rostrum to near the
cervical groove, is tuberculate with a short double carina with tubercles in
the middle. For those reasons, the specimen identified as A. (A.) consobrina
by Poore & Griffin (1979) is considered to be a new species, and, therefore,
reclassified here under the new genus Pseudoaxiopsis as the type species, P.
carpentariensis sp. nov.
178
CRM 013 – Katsushi Sakai
Type species. — Pseudaxiopsis carpentariensis gen. et sp. nov., by present
designation and monotypy. The gender of the new generic name, Pseudaxiopsis, is feminine.
Species included. — Pseudaxiopsis carpentariensis gen. et sp. nov.
Etymology. — The Greek prefix “pseud-”, meaning “false”, is added to the
generic name Axiopsis, because the type species, P. carpentariensis is closely
similar to the species of the genus Axiopsis.
Pseudoaxiopsis carpentariensis sp. nov.
(figs. 31, 32)
Axiopsis (Axiopsis) consobrina — Poore & Griffin, 1979: 230, fig. 4a-d [not fig. 4e = Axiopsis
consobrina De Man, 1905].
Material examined. — AM P 16308, holotype, 1 male (TL/CL, 52.0/18.2 mm), Gulf of
Carpentaria, Queensland, Sta. 363, less than 14 fms [less than 25 m], 1963-64, leg. CSIRO
Prawn Survey, det. G. Poore as Axiopsis consobrina.
Diagnosis of male holotype (fig. 31A). — Body scattered with short
stiff setae and granules. Rostrum (fig. 31B) acutely triangular and deflexed
downward distally and almost twice as long as wide at base, lateral margins
each with about 7 spines, extending posteriorly onto gastric region as lateral
carinae. Gastric region convex, bearing 5 longitudinal carinae; median carina
tuberculate, originating from middle of rostrum to near cervical groove, and
bearing short double carina with tubercles in middle; submedian carinae
tuberculate, lateral carinae denticulate anteriorly but tuberculate posteriorly,
and both lateral and submedian carinae also extending to near cervical groove.
Anterolateral margin of carapace with a small spine just above rounded
convexity at level of A2 peduncle, and below it two swellings; branchial region
with anterior spine. A1 peduncle reaching proximal half of A2 penultimate
segment. A2 peduncular segment 1 denticulate, with distinct distolateral
tooth on dorsodistal margin; segment 2 elongate to form dorsodistal spine,
bearing spine dorsoproximally; scaphocerite strong, reaching distal margin of
penultimate segment, segment 3 terminated with ventrodistal spine. Eyestalks
subglobose, reaching half length of rostrum. Mxp3 (fig. 31C) coxa with
stout mesiodistal tooth; ischium with three mesial teeth; merus with 5 mesial
spines increasing in length distally; propodus unarmed; and dactylus setose
on distal half of ventral margin. P1 subequal. In left cheliped (fig. 32A), coxa
with mesiodistal spine; merus with scale-like granules with setae on lateral
surface, scale-like denticles on ventrolateral margin, 4 distinct ventral spines
on ventromesial margin, and subdistal spine on dorsal margin; carpus smooth
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Fig. 31. Pseudoaxiopsis carpentariaensis sp. nov. A, whole body, lateral view; B, carapace,
dorsal view; C, Mxp3. A-C, AM P 16308, holotype, 1 male (TL/CL, 52.0/18.2 mm), Gulf of
Carpentaria, Queensland. Scales A, 2 mm; B, C, 1 mm.
on ventral margin; palm with distodorsal spinule and unarmed ventrally,
cutting edges of fingers dentate; dactylus as long as fixed finger; ratio of dorsal
lengths of merus : carpus : propodus = 1.0 : 0.6 : 1.0. Right cheliped similar to
left cheliped, but more slender: distal parts of fingers lost. P2 chelate; coxa
with mesiodistal spine; ischium with distinct proximal and distal teeth, and
three denticles on ventral margin; merus with 4 spines on ventral margin;
fingers longer than palm. P3 merus with row of denticles and subdistal spines
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Fig. 32. Pseudoaxiopsis carpentariaensis sp. nov. A, P1 on left side; B, abdominal somites 15, dorsal view; C, abdominal somite 6, telson, and uropod on right side. A-C, AM P 16308,
holotype, 1 male (TL/CL, 52.0/18.2 mm), Gulf of Carpentaria, Queensland. Scales 1 mm.
on ventral margin; propodus elongate, with spinules on ventrolateral margin.
P4 merus similar to P3 merus, but more slender. P5 missing. Abdominal
somites 1-6 bearing characteristic tergum composed of carinae arranged in
longitudinal stripes (fig. 32B, C). Male Plp1 absent; male Plp2 endopod with
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appendices interna and masculina at proximal third, appendix interna slightly
longer than appendix masculina, with distal setae; Plps3-5 endopods with
appendix interna. Telson and uropods setose on posterior margin. Telson (fig.
32C) longer than wide; lateral margins almost parallel, each bearing proximal
lobate process with spine, followed distally by 1-2 spines; posterior margin
rounded with median spine; two pairs of spines medially on dorsal surface.
Uropodal endopod denticulate, with spine at distolateral angle, bearing median
carina with three distal spines dorsally. Uropodal exopod bearing 5 spines on
lateral margin, and longitudinal ridge with spinules and transverse suture with
spinules on dorsal surface.
Remarks. — Poore & Griffin (1979: 230, fig. 4) described one male (AMP
16308) and one female (AMP 24677), both from the Gulf of Carpentaria,
Queensland, and regarded them as Axiopsis consobrina De Man, 1905, though
the female specimen is inaccessible, because it was lost (G.C.B. Poore, in litt.).
However, it is found in the present revision, by carefully examining the male
specimen, that the gastric region is convex, bearing five longitudinal carinae;
the median carina is tuberculate, originating from the middle of the rostrum
to near the cervical groove, and bearing a short double carina with tubercles
in the middle; the submedian carinae are tuberculate, and the lateral carinae
are denticulate anteriorly but tuberculate posteriorly, both the lateral and
submedian carinae are also extending to near the cervical groove (fig. 31B);
the chelipeds on the left and right side are slender; the cheliped on the right
side is cut off at the distal half of the chela (fig. 31A), and the larger cheliped
propodus on the left side (fig. 32A) is slender with smooth dorsal and ventral
margins, the surface is scattered with short vertical rows of setae, the cutting
edges of the fingers are minutely and regularly dentate; the ratio of dorsal
lengths of merus : carpus : propodus = 1 : 0.6 : 0.4; and male abdominal
somite 1 is narrow and pointed ventrally; somites 2-4 are provided laterally
with a longitudinal carina between tergum and pleuron; somites 5-6 are not
provided laterally with a longitudinal carina; pleura 2-4 are triangular, and
pleura 5-6 are rounded ventrally, pleura 4-5 bear a spinule on the anteroventral
margin, and pleuron 6 is provided ventrally with a spinule (fig. 31A).
It is noted that there are some differences between Poore & Griffin’s (1979)
description and the present description. Nonetheless, the male specimen
identified as Axiopsis consobrina by Poore & Griffin (1979) is clearly different
from Axiopsis consobrina De Man, 1905, in which the gastric region bears
lateral carinae with 8-9 denticles; submedian rows of 8-9 denticles, and a
median row of 15 denticles including a hepatic tubercle, lacking a short double
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carina in the middle. Also, “the lower angle of the pleura of the 1st somite
is obtuse, presenting no small tooth; the regularly rounded, posterior margin
of the pleura of the 2nd somite curves regularly into lower border, which is
also the case in the pleura of the 3rd ; those of the 4th an angle is still barely
distinguishable, but only in the 5th an obtuse angle is observed” (De Man,
1925d: 81) and in the paralectotype of A. consobrina examined, abdominal
somites 1-6 are smooth and bear no longitudinal carina between tergum and
pleuron, and pleura 2-5 are convex or rounded ventrally (see fig. 7A). For
those reasons mentioned, the male specimen must clearly be assigned to a new
species and included into a new genus, Pseudoaxiopsis, as P. carpentariensis
sp. nov. Regarding the missing female specimen from the same locality, it is
probable that the female specimen was not Axiopsis consobrina De Man either,
because its larger cheliped (Poore & Griffin, 1979, fig. 4e) is quite different
from that of Axiopsis consobrina De Man.
Type locality. — Gulf of Carpentaria, Queensland.
Distribution. — Only known from the type locality.
Etymology. — The new species is named Pseudaxiopsis carpentariensis sp.
nov. after its type locality, Carpentaria, Queensland, where the holotype was
collected.
Genus Ralumcaris gen. nov.
Diagnosis. — Rostrum short and triangular, unarmed on lateral margins,
and proximally with a pair of distinct supraorbital spines. Gastric region
distinctly elevated, bearing smooth median carina running to near cervical
groove in male, though only in anterior half in female, but no submedian
and lateral carinae. P1 chelate; merus with low triangular tooth on ventral
margin. Abdominal somites smooth laterally. Male Plp1 absent; male Plp2
endopod without appendix interna, but with appendix masculina at proximal
third; male Plps3-5 endopods without appendix interna. Uropodal exopod with
distal flap.
Remarks. — Axiopsis (Paraxiopsis) bisquamosa De Man, 1905 was included in the genus Eutrichocheles by Sakai & De Saint Laurent (1989) as
E. bisquamosa (De Man, 1905). However, it has turned out that E. bisquamosa
is very different from the type species of Eutrichocheles, E. modestus (Herbst,
1796). E. modestus is a large-sized species, the scaphocerite is bifurcate, the
gastric region bears median, submedian, and lateral carinae, and the postcervical carina is often present; and the male Plp1 is a simple stubby protrusion.
Whereas E. bisquamosa is a small-sized species; the scaphocerite is small and
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simple; the gastric region bears only a smooth median carina; the male Plp1 is
absent. For those reasons, E. bisquamosa is excluded from Eutricholeles and
reclassified here under the new genus Ralumcaris as R. bisquamosa (De Man,
1905).
Type species. — Axiopsis (Paraxiopsis) bisquamosa De Man, 1905, by
present designation and monotypy. The gender of the new generic name,
Ralumcaris, is feminine.
Species included. — Ralumcaris bisquamosa (De Man, 1905).
Etymology. — The new generic name Ralumcaris is derived from the type
locality, “Ralum, New Pommerania”, now a camp at Rabaul on New Britain
(Bismarck Archipelago, Papua New Guinea); “New Pommeriana” comes from
the German colonial name of the island that was called “Neu Pommern”.
Ralumcaris bisquamosa (De Man, 1905)
(figs. 33, 34)
Axiopsis (Paraxiopsis) bisquamosa De Man, 1905: 597; De Man, 1925d: 7 (list), 72 (key), 109,
pl. 8 fig. 20-20c, pl. 9 fig. 20d-m; Holthuis, 1953b: 51.
Eutrichocheles bisquamosa — Sakai & De Saint Laurent, 1989: 53, fig. 15; Kensley, 1994: 822.
Fig. 33. Ralumcaris bisquamosa (De Man, 1905). A, B, whole body, lateral view. A, ZMA
Crust. De. 102.674, lectotype, 1 male (TL/CL, 10.5/3.5 mm), anchorage off Lirung, SalibabuIsland, “Siboga” Sta. 133, up to 36 m; B, ZMB 20455, 1 ovig. female (TL/CL, 20.0/6.1 mm),
Ralum, New Pomerania. Scales 1 mm.
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Fig. 34. Ralumcaris bisquamosa (De Man, 1905). A, gastric region of carapace, dorsal view;
B, same, lateral view; C, larger cheliped, lateral view. A-C, ZMA Crust. De. 102.674, lectotype,
1 male (TL/CL, 10.5/3.5 mm), anchorage off Lirung, Salibabu-Island, “Siboga” Sta. 133, up to
36 m. Scales 1 mm.
Paraxiopsis bisquamosa — Kensley, 1996c: 711, 712; Kensley, 2003: 372, table 2.
Material examined. — ZMA Crust. De. 102.674, lectotype, 1 male (TL/CL, 10.5/3.5 mm)
(fig. 33A), anchorage off Lirung, Salibabu-Island, “Siboga” Sta. 133, up to 36 m; ZMB 20455,
1 male (TL/CL, 11.0/4.2 mm), 2 ovig. females (TL/CL, 18.0/5.7; 20.0/6.1 mm) (fig. 33B),
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Ralum, New Pommerania, 24.x.1896, leg. S. Dahl, det. De Man (De Man, 1925d: 190, shown
as No. 18713 of the Berlin Museum).
Diagnosis. — Rostrum (figs. 33A, B, 34A, B) small and triangular, and
slightly upturned, lateral margins unarmed, bearing proximally a pair of
distinct supraorbital spines. Gastric region steeply elevated from base of
rostrum, bearing median carina running to near cervical groove in male (ZMB
20455), though only in anterior half in female, and lateral carinae originating
from behind supraorbital spines to near cervical groove, but no submedian
carinae. Cervical groove distinct; no linea thalassinica. Posterior lobes present.
A2 peduncular segment 2 not forming distodorsal spine; A2 scaphocerite
small and simple. Mxp3 with denticulate ischial crest. P1 chelate, merus with
low triangular tooth on ventral margin (fig. 34C). Ps3-4 propodi with series
of ventral setae, bearing slender but distinct distoventral spines; P5 simple.
Abdominal somites smooth laterally (fig. 33A, B). Male Plp1 absent; male
Plp2 endopod with appendix masculina at proximal third, but without appendix
interna; male Plps3-5 biramous, endopods without appendix interna. Female
Plp1 uniramous, consisting of proximal segment and multiarticulate flagellum,
and female Plps2-5 biramous, endopods without appendix interna. Telson
longer than wide, bearing proximal lobe with spine on lateral margins, and
median spine on rounded posterior margin. Uropodal endopod bearing three
spines including distal one on distal half of lateral margin, and longitudinal
median carina with 4 spines on dorsal surface. Uropodal exopod bearing 4
spines on lateral margin and movable spine at distolateral angle, and transverse
suture with spinules dorsally.
Type locality. — Anchorage off Lirung, Salibabu Island, “Siboga” Sta. 133,
up to 36 m.
Distribution. — Indonesia — Salibabu Island (De Man, 1905, 1925; Sakai
& De Saint Laurent, 1989); Mariana Islands — Saipan; Papua New Guinea –
Ralum, Gazelle Peninsula (De Man, 1925d); up to 36 m.
Genus Scytoleptus Gerstaecker, 1856
Scytoleptus Gerstaecker, 1856: 155; Borradaile, 1903: 539; De Man, 1925d: 1 (key), 49;
Barnard, 1950: 498 (key), 499; Poore & Griffin, 1979: 243; Sakai & De Saint Laurent, 1989:
36; Poore, 1994: 97 (key); Sakai, 1994: 180; Poore & Collins, 2009: 277.
Evaxius Kingsley, 1882: 126.
Diagnosis. — Rostrum small, short, and triangular, with pointed apex, bearing supraorbital spines at base. Anterolateral margin of carapace unarmed.
Gastric region extremely elevated from base of rostrum, bearing median carina
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and lateral carinae, but no submedian carinae. Eyestalks subglobose; cornea
pigmented. A2 scaphocerite present as small plate. P1 unequal; palm much
longer than fingers, and unarmed on dorsal margin. Ps3-4 propodi with transverse rows of spines ventrolaterally, and dactyli with lateral row of spines.
Ps3-5 subchelate. Pleurobranchs on Ps2-4. Male Plp1 uniramous, and unsegmented or bisegmented; male Plp2 biramous, endopod with appendices interna and masculina; male Plps3-5 biramous, endopods with appendix interna.
Female Plp1 uniramous, consisting of proximal segment and multiarticulate
flagellum; female Plps2-5 endopods with appendix interna. Telson strongly
convex dorsally and ovate, tapering towards rounded posterior margin with or
without median spine. Uropods convex dorsally; exopod without distal flap.
Type species. — Scytoleptus serripes Gerstaecker, 1856, by original designation and monotypy. Gender of the generic name, Scytoleptus, masculine.
Species included. — Scytoleptus barbatus sp. nov. and S. serripes Gerstaecker, 1856.
K EY
TO THE
S PECIES
OF THE
G ENUS SCYTOLEPTUS
1 – Gastric median carina running from base of rostrum onto vertically elevated anterior
gastric region, bearing two spines on its vertical slope . . . . . . . . . . . . S. barbatus sp. nov.
– Gastric median carina running from base of rostrum onto slantingly elevated anterior
gastric region with three spines on its slanting slope . . . . . . . . . . . . . . . . . . . . . . S. serripes
Scytoleptus barbatus sp. nov.
(figs. 35, 36)
Scytoleptus serripes — Strahl, 1862a: 1055 [type locality: Luzon]; Strahl, 1862b: 383; Estampador, 1937: 500; Poore & Griffin, 1979: 243, fig. 11; Sakai, 1994: 180, 200 (key), fig. 3;
Poore & Collins, 2009: 277.
Material examined. — AM P 72878, holotype, male (TL/CL, 69.0/22.6 mm), between Cape
Bossutt and Broome, W. Australia, dredged, 11.ix.1929; AM P 72879, allotype, 1 female
(TL/CL, 61.0/23.5 mm), between Cape Bossutt and Broome, Australia, dredged, 11.ix.1929;
AM P 72875, paratypes, 2 females (TL/CL, 44.0/15.5; 68.0/23.0 mm), East Poiny, Port Darwin,
Australia, 27.vi.1929; AM P 72876, paratype, 1 male (TL/CL, 40.0/14.3 mm), N.W. of South
Shell Is., Port Darwin, dredged, 5 fms [ca. 9 m], 24.vii.1929; AM P 72877, paratype, 1 female
(TL/CL, 74.0/23.2 mm), between Cape Bossutt and Broome, Australia, dredged, 10.ix.1929;
AM P 72880, paratypes, 1 male (TL/CL, 68.0/20.9 mm), 2 females (TL/CL, 66.0/22.4;
78.0/25.8 mm), Dudley Rey, Darwin, Northern Territory, 13.x.1965, leg. B. Pope; AM P 72881,
paratype, 1 male (TL/CL, 75.0/23.3 mm), Dudley Rey, Darwin, Northern Territory, 13.x.1965,
leg. B. Pope; RMNH D 37756, 1 male (TL/CL, 38.0/13.3 mm), Australia, North Kimberley, east
of Montaliver Island, Sta. 50, intertidal, 15.vii.1988, WAM 1023-88; NHM 1843.6, 1 female
(TL/CL, 35.0/13.8 mm, smaller cheliped missing), S. Philippines, leg. Cuming.
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Fig. 35. Scytoleptus barbatus sp. nov. Whole body, lateral view. AM P 72878, holotype, 1 male
(TL/CL, 69.0/22.6 mm), between Cape Bossutt and Broome, Western Australia. Scale 1 mm.
Description of male holotype. — Rostrum (fig. 35) small, short, and
triangular, with ventral convexity just above eyestalks, bearing a pair of
supraorbital teeth at base. Carapace setose on lateral margins. Gastric region
bearing median carina with hepatic tubercle, originating from base of rostrum
onto vertically elevated anterior gastric region with two spines on vertical
slope, and extending posteriorly to anterior two-thirds of gastric region;
lateral carinae short and terminating anteriorly in rounded corner; but without
submedian carinae. Cervical groove indistinct. Posteromedian carina obscure.
A1 peduncle slightly overreaching rostrum. A2 peduncle segment 1 with 4
small ventrodistal spines and median spinule on ventral margin; segment 2
unarmed, scaphocerite present as small triangular plate; penultimate segment
elongate, distal segment one-third of penultimate segment. Eyestalks small,
cornea pigmented. Mxp3 ischial crest curved and dentate; merus with strong
distal spine mesially. P1 (fig. 36A, B) unequal; larger cheliped unarmed
on margins, merus smoothly carinate on ventral margin; palm two times as
long as dactylus, and granulate in distal parts of lateral and mesial surfaces;
fixed finger with obtuse median tooth on cutting edge; dactylus little longer
than fixed finger, stout, curved downward distally, and unarmed on cutting
edge. Smaller cheliped missing. P2 chelate. Ps3-4 propodi with transverse
rows of spines ventrolaterally; dactyli with a row of spines laterally. Ps3-5
subchelate. Abdominal pleura 2-5 beset with thick setae on ventral margin.
Male Plp1 uniramous and unsegmented, but slightly concave at distal third
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Fig. 36. Scytoleptus barbatus sp. nov. A, larger cheliped; B, smaller cheliped; C, male Plp1;
D, male Plp2. A-D, AM P 72878, holotype, male (TL/CL, 69.0/22.6 mm), between Cape
Bossutt and Broome, Western Australia. Scales 1 mm.
and setose marginally; male Plp2 biramous and slender, with thick marginal
setae, endopod with appendices interna and masculina; Plps3-5 biramous,
with thick marginal setae, endopods with appendix interna. Telson longer
than wide, bearing proximal lobe with spine on lateral margins, small spine
at posterolateral angle, and small median spine on rounded posterior margin;
dorsal surface strongly convex, spinous, and setose. Uropodal endopod slightly
shorter than telson, bearing spinous longitudinal carina medially on dorsal
surface. Uropodal exopod convex dorsally, scattered with spinules on lateral
half, and rounded at posterolateral angle.
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In smaller cheliped of paratype (AM P 72881) merus smooth on ventral
margin; palm more than twice as long as dactylus; fixed finger minutely
denticulate.
Remarks. — The present new species, Scytoleptus barbatus from Australia
is closely similar to the commonly known species Scytoleptus serripes Gerstaecker, 1856 from South Africa, but they are also clearly different from each
other. In Scytoleptus barbatus sp. nov. the median gastric carina with a hepatic tubercle is running from the base of the rostrum onto the vertically elevated anterior gastric region, bearing two spines on its vertical slope, and
extending to the posterior third of the gastric region; and the lateral carinae
are short and terminating anteriorly with a rounded corner; in the larger cheliped the fixed finger is armed with an obtuse median tooth on the cutting
edge; the abdominal pleura 2-5 are beset ventrally with thick setae. The male
Plp1 (fig. 36C) is uniramous and unsegmented, slender and setose marginally,
bearing a shallow concavity at its distal third; the proximal part is broader and
longer than the distal section; the male Plp2 (fig. 36D) with slender protopod is biramous, the endopod and exopod are slender and setose marginally,
and the endopod bears appendices interna and masculina mesiomedially; male
Plps3-5 with slender protopod are biramous, the endopods and exopods are
setose marginally, and the endopods bear an appendix interna; the telson bears
a proximal lobe without a spine on the lateral margin. In Scytoleptus serripes
by contrast, the gastric median carina is running from the base of the rostrum
onto the slantingly elevated anterior gastric region, with three spines on its
slanting slope, and extending to the cervical groove; the gastric lateral carinae are terminating anteriorly with a tooth, extending to the posterior third of
the gastric region; in the larger cheliped the fixed finger is armed with a distinct median tooth on the cutting edge; the abdominal pleura 3-5 do not bear
marginal setae ventrally. Male Plp1 (fig. 37C) is uniramous and leaf-like, and
bisegmented, and the proximal segment is broader and longer than the distal one; and male Plp2 (fig. 37D) is biramous, the protopod is broadened and
oval, and the endopod bears mesiomedially appendices interna and masculina;
Plps3-5 protopods are broadened and oval, and the endopods bear an appendix
interna; the telson is provided with a proximal lobe with a strong spine on the
lateral margin.
The small specimen from the Philippines (NHM 1843.6, female, TL/CL,
35.0/13.8 mm) is identified as S. barbatus sp. nov. from Australia, because
the gastric median carina is running from the base of the rostrum onto the
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CRM 013 – Katsushi Sakai
vertically elevated anterior gastric region, and the abdominal pleura are setose
on the ventral margin, though the telson bears a proximal lobe with a reduced
tooth on the lateral margins.
Most of the specimens recorded from the Indian Ocean belong to Scytoleptus serripes, because the gastric median carina is running from the base of the
rostrum onto the slantingly elevated anterior gastric region, and the gastric lateral carinae are distinctly discernible, bearing a short, acute spine anteriorly
(Bouvier, 1915b, fig. 8, from Mauritius Is.; Sakai & De Saint Laurent, 1989,
fig. 9A, B, from Tuléar, Madagascar).
It has turned out by the present, careful examination of the material from
Australia that the male Plp2 endopod bears slender appendices interna and
masculina, though Sakai (1994: 180, fig. 3A, D) described that the male
Plp2 endopod bears an appendix masculina, but no appendix interna. Such
misdescription may have been caused by the thick setae covering the pleopods,
which obstructed an exact observation.
Type locality. — Between Cape Bossutt and Broome, W. Australia.
Distribution. — Australia — Cape Wessel, Darwin, Cockatoo Island, Broome, Port Hedland, Dampier Archipelago, and Exmouth Gulf (Poore & Griffin,
1979); Darwin and Port Essington (Sakai, 1994), up to 36 m; Philippines —
Luzon (Strahl, 1862b; Estampador, 1937).
Etymology. — The present new species is named Scytoleptus barbatus,
by applying the Latin word “barbatus”, meaning “bearded”, because it is
characterized by its marginal, thick setae on the carapace and the abdominal
somites. The new species name is an adjective, agreeing in gender with the
(masculine) generic name.
Scytoleptus serripes Gerstaecker, 1856
(fig. 37)
Scytoleptus serripes Gerstaecker, 1856: 158, pl. 6 figs. 1-4; Hilgendorf, 1878: 827; Borradaile,
1903: 540; Lenz, 1905: 379; Bouvier, 1914: 702; Bouvier, 1915b: 198, figs. 8, 9; De Man,
1925d: 5 (list), 49 (in part), pl. 4 fig. 9-9h; Barnard, 1950: 499; Fourmanoir, 1955: 30;
Kensley, 1981b: 30; Sakai & De Saint Laurent, 1989: 37, fig. 9.
Evaxius tricarinatus Kingsley, 1882: 130, pl. 1 fig. 1 [type locality: Zanzibar]; Borradaile, 1903:
540.
Scytoleptus tricarinatus — De Man, 1925d: 5 (list).
Material examined. — RMNH D 1487, holotype, ovig. female, damaged (CL 14.8), Seba,
Savu, “Siboga”-Exp., 25.iv.1899; ZMB 1135, 2 males (TL/CL, 45.0/16.7-47.0/17.9 mm),
1 ovig. female (TL/CL, 67.0/22.3 mm), 1 female (TL/CL, 59.0/21.4 mm), Mozambique, leg.
Peters.
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191
Fig. 37. Scytoleptus serripes Gerstaecker, 1856. A, whole body, lateral view; B, larger cheliped,
lateral view; C, male Plp1; D, male Plp2. A-D, ZMB 1135, male (TL/CL, 47.0/17.9 mm),
Moçambique. Scales 1 mm.
Diagnosis. — Rostrum (fig. 37A) thorn-shaped, lateral margins posteriorly
extending shortly. Gastric region bearing median carina with hepatic tubercle
running from base of rostrum onto slantingly elevated anterior gastric region
with three spines on its slanting slope, and extending posteriorly to cervical
groove; lateral carinae terminating anteriorly with tooth, extending to posterior third; submedian carinae absent. Cervical groove distinct. Postcervical ca-
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CRM 013 – Katsushi Sakai
rina obscurely developed in posterior part. Pair of postlateral lobes present.
A1 segment 2 slightly overreaching rostrum. A2 segment 1 with ventral spine;
segment 2 unarmed, scaphocerite present as a small triangular plate; penultimate segment elongate, 4 times as long as terminal one. Eyestalks about
half length of rostrum. Mxp3 ischial crest curved and dentate on mesial surface; merus with strong mesiodistal spine. P1 unequal, not sexually dimorphic.
Larger cheliped (fig. 37B) usually unarmed; merus smoothly carinate on ventral margin; palm two-thirds length of dactylus; fixed finger tuberculate on
ventral margin, bearing obtuse median tooth on cutting edge; dactylus slightly
longer than fixed finger, stout, curved downward distally, and unarmed on cutting edge. In smaller cheliped merus carinate on ventral margin; palm more
than twice as long as dactylus; fixed finger denticulate on cutting edge. P2
chelate. Ps3-4 propodi with transverse rows of spines ventrolaterally; dactyli
with rows of spines laterally. Ps3-5 subchelate. Abdominal pleura 1-6 without
thick setae on ventral margin. Male Plp1 (fig. 37C) uniramous, leaf-like, and
bisegmented, proximal segment broader and longer than distal one, distal segment slenderly protruded; male Plp2 (fig. 37D) with ovate protopod, biramous
and slender, endopod with appendices interna and masculina; Plps3-5 biramous, endopods with appendix interna. Female Plp1 slender and uniramous,
consisting of proximal segment and multiarticulate flagellum; female Plps2-5
biramous, endopods with appendix interna. Telson longer than broad, tapering
towards rounded posterior margin with small median spine; lateral margins
bearing proximal lobe with strong spine; dorsal surface spinous and setose,
and strongly convex. Uropodal endopod slightly shorter than telson, bearing
longitudinal median carina with spinules. Uropodal exopod convex dorsally
and rounded at distolateral angle, bearing spinous row sublaterally on dorsal
surface.
Remarks. — The figure of Evaxius tricarinatus Kingsley, 1882 from
Zanzibar clearly shows that his species is safely regarded as Scytoleptus
serripes, because the gastric median carina is running from the base of
the rostrum onto the slantingly elevated anterior gastric region, and the
gastric lateral carinae are clearly developed, bearing a distinct anterior spine.
Therefore, E. tricarinatus is here synonymized with S. serripes.
Poore & Collins (2009: 277) included Scytoleptus serripes in the Indo-West
Pacific fauna as an Australian species. However, most probably the Australian
specimens are to be referred to the new species here described, i.e., S. barbatus,
because these cannot be identified with S. serripes in view of the present
descriptions.
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Type locality. — Port Natal, South Africa.
Distribution. — South Africa — Port Natal (Gerstaecker, 1856); Mozambique (Strahl, 1862a; Hilgendorf, 1878); Port Natal, South Africa (Fourmanoir,
1955); Mayotte (Sakai & De Saint Laurent, 1989); Kenya — Iwatin Island and
Jadini (Sakai & De Saint Laurent, 1989); Zanzibar (Kingsley, 1882); Comoros
(Sakai & De Saint Laurent, 1989); Madagascar — Sta. Marie (Bouvier, 1915b;
Sakai & De Saint Laurent, 1989), Nosy Bé and Tuléar (Sakai & De Saint Laurent, 1989); Aldabra (Lenz, 1905); Mauritius (Bouvier, 1915b; Sakai & De
Saint Laurent, 1989); Indonesia — Savu Islands and Salayar Island (De Man,
1925d).
Genus Spongiaxius Sakai & De Saint Laurent, 1989
Spongiaxius Sakai & De Saint Laurent, 1989: 41; Poore, 1994: 97 (key); Sakai & Ohta, 2005:
88; Poore, 2008: 168; Poore & Collins, 2009: 277.
Sakaiocaris Kensley, 1989: 964.
Diagnosis. — Rostrum prominent and furrowed on dorsal surface; lateral
margins armed with erect spines, extending onto gastric region. Gastric region convex, bearing serrated median, submedian, and lateral carinae. Cervical groove distinct except for anterolateral part. Eyestalks subglobose; cornea
pigmented. A2 scaphocerite prominent and horn-shaped. P1 subequal; chelae
thick and semicircular. P2 chelae densely pubescent on lateral surface. Ps34 simple. P5 subchelate. Ps3-4 coxae with knobs ventrodistally. Both P3 and
P5 coxae with genital pores. Pleurobranchs on Ps2-4. Abdominal somites 2-5
furrowed between tergum and pleuron. Abdominal pleura almost smooth on
lateral surface and acutely pointed ventrally. Male Plp1 uniramous and bisegmented; distal segment spatulate; male Plp2 biramous and slender, endopod
with appendices interna and masculina; male Plps3-5 endopods with appendix
interna. Female Plp1 uniramous, consisting of proximal segment and multiarticulate flagellum; female Plps2-5 endopods with appendix interna. Telson
twice as long as wide. Uropodal exopod bearing short transverse suture with
spinules. Usually living in sponges. Hermaphroditic. [Adapted from Sakai &
Ohta, 2005.]
Type species. — Axiopsis brucei (Sakai, 1986), by original designation and
monotypy. The gender of the generic name, Spongiaxius, is masculine.
Species included. — Spongiaxius brucei (Sakai, 1986).
Spongiaxius brucei (Sakai, 1986)
Axiopsis brucei Sakai, 1986: 12, figs. 1-6.
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CRM 013 – Katsushi Sakai
Spongiaxius brucei — Sakai & De Saint Laurent, 1989: 44; Sakai, 1994: 200 (key); Sakai &
Ohta, 2005: 89, fig. 11 (in colour); Poore, 2008: 168; Poore & Collins, 2009: 277; Robles et
al., 2009: 316.
Material examined. — RMNH D 36516, paratype, 1 ovig. female (TL/CL, 68.0/22.6 mm),
North West Shelf, Western Australia, 18◦ 52.2′ S 116◦ 11.1′ E, 30.i.1984, leg. A.J. Bruce; NHM
1986: 300, paratype, 1 female (TL/CL, 77.0/25.5 mm), same data as above; NTM Cr. 00605,
1 female (TL/C 71.0/24.9 mm), R/V “Soela” T/3, 17◦ 55.5′ S 118◦ 19.5′ E, 450-454 m, in
hexactinellid sponge, 27.i.1984, leg. A.J. Bruce; NHM 1992.1087, paratype, 1 female (TL/CL,
48.0/15.8 mm), Philippines, 12◦ 20′ N 121◦ 42′ E, MUSORSTOM 3, Sta. CP 122, 1985, N.O.
“Caviolis”, 673-675 m.
Diagnosis. — Rostrum prominent and furrowed on dorsal surface; lateral
margins armed with 9 erect spines, extending onto gastric region. Anterolateral margin of carapace unarmed. Gastric region convex, bearing serrated
median, submedian, and lateral carinae. Cervical groove distinct except for
anterolateral part. Posterior margin of carapace protruded as lateral lobes.
Eyestalks subglobose; cornea pigmented. A2 scaphocerite prominent, hornshaped, and of same length or slightly longer than dorsal spine of segment
2. Mxp3 ischium-merus subpediform; propodus rectangular; dactylus slender;
exopod present. P1 subequal, palm broadened and tuberculate on lateral surface; chelae thick and semicircular. P2 chela densely pubescent on lateral surface. Ps3-4 simple, propodi bearing 15 transverse rows of spinules ventrolaterally, a row of spinules along distoventral margin, a strong spine at setose
distoventral angle; dactyli scattered with 4-7 distinct, translucent spines on
lateral surface, bearing translucent spinules on ventral margin, and distinct
translucent spine at distal tip. P5 subchelate, propodus protruded ventrodistally and rounded on distal margin, with translucent spinules, bearing a row
of thick setae distolaterally; dactylus truncate on distal margin, with slender
translucent spines including strong one at distodorsal angle. Ps3-4 coxae with
knobs ventrodistally. Both P3 and P5 coxae with genital pores. Pleurobranchs
on Ps2-4. Abdominal somites 1-4 smoothly and shallowly furrowed on lateral
side of tergum and convex on dorsal side of pleuron; abdominal somites 5-6
broadly convex on dorsal side of pleuron; pleura 1-6 sharply pointed ventrally;
pleura 3-5 bearing small spine on anteroventral margin. Telson twice as long
as wide, bearing proximal lobe with spine followed posteriorly by three spines
on lateral margin, and two pairs of spines on anterior half of dorsal surface
and a pair of spines at each posterolateral corner; posterior margin with median spine. Uropodal endopod as long as telson, bearing two spines including
posterolateral one on lateral margin, and proximal spine medially on dorsal
surface and median spine on posterior margin. Uropodal exopod with 5 spines
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195
including distolateral one on lateral margin, bearing a row of 4 spines sublaterally and a short transverse suture with spinules on dorsal surface, and median
spine on distal margin. Usually living in sponges. Hermaphroditic. [Adapted
from Sakai & Ohta, 2005.]
Type locality. — North West Shelf, Western Australia, 18◦ 43.7′ S
117◦ 02.2′ E, R/V “Soela”, Sta. NWS-43, trawl, 454 m.
Distribution. — Australia — North West Shelf, Western Australia, 296456 m (Sakai, 1986; Sakai & De Saint Laurent, 1989); W. and WNW Lacépède
Archipelago, 434-444 m (Poore, 2008); Philippines — Sulu Sea, 08◦ 06.36′ N
118◦ 26.49′ E – 08◦ 07.04′ N 118◦ 26.06′ E, 688-693 m (Sakai & Ohta, 2005).
FAMILY CALLIANIDEIDAE KOSSMANN, 1880
Callianideinae Kossmann, 1880: 80.
Callianideidae Kossmann, 1880: 80; Gurney, 1938: 343; De Saint Laurent, 1979: 1395; Poore
& Griffin, 1979: 221; Sakai, 1987a: 303 (list); Kensley & Heard, 1991: 497; Sakai, 1992c: 9;
Poore, 1994: 103; Poore, 1997: 348.
Callianideinae — De Man, 1928b: 20, 30; Melin, 1939: 4; Balss, 1957: 1582; De Saint Laurent,
1973: 515; Sakai, 1992c: 10.
Diagnosis. — Rostrum distinct or short between deep-set orbits. Carapace
with setal rows, but without cardiac sulcus and cardiac prominence; linea thalassinica present but short posterior to eyestalk, or absent. Cervical groove
present. Eyestalks flattened and contiguous. Maxilla 2 scaphognathite with
posterior whip. P1 chelate, unequal, and dissimilar. P2 chelate. P3 propodus
square. P4 propodus rectangular, coxa flattened. Abdominal somite 1 without
anterolateral lobes. Male Plp1 uniramous and unsegmented, or bi- or trisegmented; male Plp2 biramous, endopod and exopod each with respiratory filaments; endopod bearing appendices interna and masculina; male Plps3-5 biramous, endopods and exopods each with respiratory filaments, endopods with
appendix interna. Female Plp1 uniramous and bisegmented, distal segment
slender; female Plps2-5 biramous, endopods and exopods each with respiratory filaments, endopods with appendix interna. Pleurobranchs absent. Uropodal exopod without suture and dorsal plate.
Remarks. — Kensley & Heard (1991) included 7 genera (Callianidea
H. Milne Edwards, 1837a, Michelea Kensley & Heard, 1991, Mictaxius
Kensley & Heard, 1991, Thomassinia De Saint Laurent, 1979, Crosniera
Kensley & Heard, 1991, Marcusiaxius Rodrigues & De Carvalho, 1972, and
Meticonaxius De Man, 1905) in the family Callianideidae Kossmann, 1880.
Poore (1994) revised the Callianideidae and reclassified it into three families
(Callianideidae and Thomassiniidae in the superfamily Callianassoidea Dana,
and Micheleidae in the superfamily Axioidea Huxley, 1879). He included only
one genus (Callianidea H. Milne Edwards, 1837a) in the Callianideidae, and
three genera (Crosniera Kensley & Heard, 1991, Mictaxius Kensley & Heard,
1991, and Thomassinia De Saint Laurent, 1979) in the Thomassiniidae, and
four genera (Marcusiaxius Rodrigues & De Carvalho, 1972, Meticonaxius De
Man, 1905, Michelea Kensley & Heard, 1991, and Tethisea Poore, 1994) in the
198
CRM 013 – Katsushi Sakai
Micheleidae. Though he included four genera (Marcusiaxius, Meticonaxius,
Tethisea, and Michelea) in the Micheleidae, the first three genera are different
from Michelea, the type genus of the family Micheleidae, because in those
three genera the rostrum is distinct, whereas in Michelea it is located between
the deep-set orbits and small, so that Michelea is safely included in the family
Micheleidae as the type genus, but the other three genera (Marcusiaxius,
Meticonaxius, and Tethisea) are not to be included in the family Micheleidae
because of their distinct rostrums. Therefore, they are excluded from the
Micheleidae, and reclassified under the family Meticonaxiidae Sakai, 1992c.
Type genus. — Callianidea H. Milne Edwards, 1837a, by designation of
ICZN, 1989: 61; and as the nominate genus.
Genera included. — Callianidea H. Milne Edwards, 1837a; Paracallianidea
Sakai, 1992c.
Genus Callianidea H. Milne Edwards, 1837
Isea Guérin-Méneville, 1832: 295 [suppressed by ICZN, 1989: 61].
Callianidea H. Milne Edwards, 1837a: 319; H. Milne Edwards, 1838: 388; Dana, 1852a: 14;
Dana, 1852b: 509; Guérin-Méneville, 1857: 45; Boas, 1880: 108, 110; Kossmann, 1880: 80;
Bate, 1888: 10; Stebbing, 1893: 184; Ortmann, 1899: 1143; Rathbun, 1901: 94; Borradaile,
1903: 548; Balss, 1914: 90; Gurney, 1938: 301, 342; Melin, 1939: 4; Balss, 1957: 1582; De
Saint Laurent, 1973: 515; Le Loeuff & Intès, 1974: 23; Poore & Griffin, 1979: 281; Sakai &
Holthuis, 1987: 93; Sakai, 1987a: 303 (list); ICZN, 1989: 61; Kensley & Heard, 1991: 498;
Sakai, 1992c: 11; Poore, 1994: 103; Poore, 1997: 349.
Callianisea H. Milne Edwards, 1837a: 321. [Replacement name for Isea Guérin-Méneville.]
Callisea Dana, 1852a: 11; Dana, 1852b: 510. [Replacement name for Isea Guérin-Méneville.]
Callianassa (Calliactites) — Borradaile, 1903: 545; De Man, 1928b: 25, 91, 95. [Type species,
Callianassa secura Lanchester, 1902.]
Calliactites — Stebbing, 1910: 369; Gurney, 1944: 83.
Diagnosis. — Cephalothorax laterally compressed. Rostrum short and
broad. Linea thalassinica short. Mxp3 merus unarmed on ventral margin.
Ps2-4 propodi with setal rows centrally or proximally. Male Plp1 uniramous
and bisegmented; male Plp2 biramous, endopod and exopod with respiratory
filaments, endopod with appendices interna and masculina; male Plps3-5
biramous, endopods and exopods with respiratory filaments, endopods bearing
leaf-like appendix interna. Female Plp1 uniramous and bisegmented; distal
segment elongate and twisted in distal half, with marginal setae (Sakai, 1992a,
fig. 5G). Female Plps2-5 biramous, endopods and exopods with respiratory
filaments, endopods bearing leaf-like appendix interna. [Adapted from Sakai,
1992c.]
Type species. — Callianidea typa H. Milne Edwards, 1837a, through
designation by ICZN, 1989: 61. The gender of the genric name, Callianidea,
is feminine.
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199
Species included. — Callianidea madagascariensis sp. nov.; C. ryukyu sp.
nov.; C. typa H. Milne Edwards, 1837a (= Callianassa secura Lanchester,
1902).
K EY
TO THE
S PECIES
OF THE
G ENUS CALLIANIDEA
1 – Mxp3 merus with subterminal tooth on mesial margin; merus of larger cheliped 1.5
times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. madagascariensis sp. nov.
– Mxp3 merus unarmed on mesial margin; merus of larger cheliped 1.8 times as long as
wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Male Plp1 with elongate distal segment bearing digitiform projection . . . . . . . . . C. typa
– Male Plp1 with distal segment shorter than proximal one, bearing no projection . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ryukyu sp. nov.
Callianidea madagascariensis sp. nov.
Callianidea sp., Poore, 1997: 354, figs. 4D-I.
Remarks. — The present new species was described as Callianidea sp. from
Madagascar by Poore (1997). His species is clearly different from C. typa;
in his Callianidea sp. the Mxp3 merus bears a subdistal tooth on the mesial
margin, and the merus of the larger cheliped is about 1.5 times as long as
broad, while in C. typa the Mxp3 merus is unarmed on the mesial margin, and
the merus of the larger cheliped is about 1.8 times as long as broad. Therefore,
Poore’s (1997) species is here defined as a new species under the name of
Callianidea madagascariensis.
Type specimen. — MNHN Th. 139, female.
Type locality. — Madagascar — Maromandia, 14◦ 10′ S 48◦ 06′ E.
Distribution. — Only known from the type locality.
Etymology. — The species name is derived from the type locality, Madagascar. It is an adjective agreeing in gender with the (feminine) generic name.
Callianidea ryukyu sp. nov.
(figs. 38, 39)
Callianidea typa — Balss, 1914: 90; Miyake, 1956: 304, figs. 1-3; Sakai, 1992c: 12, figs. 3-5.
Material examined. — SMF 24918, holotype, male (TL/CL, 55.0/12.0 mm), Taketomi-jima
between the islands Iriomote and Ishigaki, Ryukyu Islands, Japan, 1-8.v.1973, leg. Uchida &
Uda; BLT 18238 (SMF 32730), paratypes, 2 males (TL/CL, 20.0/5.2; 24.0/4.3 mm), 2 females
(TL/CL, 26.0/6.5; 39.0/8.8 mm), Mitara, Iriomote-Jima, Ryukyu Islands, 15.vi.2003, leg. H.O.
Gusey; SMF 32731, paratype, 1 female (TL/CL, 40.0/7.0 mm), Maenohama, Aka-jima Island,
Kerama Islands, Ryukyu Archipelago, Japan, 12.v.1991, leg. N. Iwasaki.
Diagnosis of male holotype (fig. 38). — Rostrum flat, triangular, with obtuse tip in dorsal view, and less than one-third as long as eyestalks (fig. 39A);
carapace protruded backward medially on posterior margin, bearing submar-
200
CRM 013 – Katsushi Sakai
Fig. 38. Callianidea ryukyu sp. nov. Whole body, lateral view. SMF 24918, holotype, 1 male
(TL/CL, 55.0/12.0 mm), Taketomi-jima between Iriomote and Ishigaki Islands, Ryukyu Islands,
Japan. Scale 1 mm.
ginal row of setae posterior to rostrum, vertical row of setae posterior to base of
each antenna, and cervical groove dorsally. Eyestalks flattened and contiguous,
with obtuse apex; cornea pigmented distolaterally. A1 segment 1 overreaching eyestalks; segment 2 shorter than segment 3. A2 peduncle overreaching
A1 peduncle, reaching distal fourth of penultimate segment; terminal segment
two-thirds as long as penultimate segment. Mxp3 ischium with denticulate ischial crest along mesial margin; merus without subterminal tooth mesially;
ischium-merus with mesial row of long, dense setae; carpus-dactylus longer
than ischium-merus; exopod with flagellum reaching middle of ischium. P1
unequal. In larger cheliped (fig. 39B) ischium denticulate on ventral margin;
merus convex on ventral and dorsal margins, 1.8 times as long as wide; carpus with 4 teeth mesiodistally; palm with obsolete teeth on ventral margin;
fixed finger two-thirds length of palm, and inconspicuously dentate on proximal half of cutting edge with obtuse median tooth, bearing a row of denticles mesially along cutting edge; dactylus bearing concavity with three distinct teeth on proximal half of cutting edge, followed by curved, denticulate
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
201
Fig. 39. Callianidea ryukyu sp. nov. A, carapace and abdominal somite 1, dorsal view; B, larger
cheliped, lateral view; C, P2; D, abdominal somites 1-2, lateral view; E, male Plp1; F, Plp2;
G, pleopodal gills. SMF 24918, holotype, 1 male (TL/CL, 55.0/12.0 mm), Taketomi-jima,
between Iriomote and Ishigaki Islands, Ryukyu Islands, Japan. Scales 1 mm.
202
CRM 013 – Katsushi Sakai
distal half. Smaller cheliped with ischium and carpus unarmed and narrow, carpus much longer than that of larger cheliped; propodus elongate; fixed finger
one-third length of palm, bearing two distal teeth; dactylus longer than fixed
finger. P2 (fig. 39C) merus to propodus with marginal setae ventrally; carpus
longer than half length of merus; chela as long as carpus; dactylus slightly
longer than fixed finger. P3 (fig. 38) subchelate; propodus about 1.3 times as
long as wide, bearing spiniform setae on ventrodistal corner. P4 coxa disc-like;
propodus three times as long as wide, bearing spine at distal corner on ventral
margin; P4 on left side missing. P5 subchelate; dactylus twice as long as fixed
finger; P5 on right side missing.
Abdominal somite 1 (fig. 38) broadening posteriorly, bearing dorsal carina
followed posteriorly by boss, and lateral row of setae on each side of dorsal
carina; pleuron weakly rounded, bearing subventral row of setae. Abdominal
somite 2 slightly longer than somite 1, pleuron without transverse setal row,
not overlapping somite 1. Abdominal somite 6 with transverse row of about
6 setae on each side near posterior margin. Male Plp1 (fig. 39E) uniramous
and bisegmented, proximal segment longer than distal one, and distal segment
finger-like. Male Plp2 (fig. 39F) biramous, endopod with small appendix
interna and elongate appendix masculina. Male Plps3-5 essentially similar to
Plp2, endopods with leaf-like appendix interna, bearing elongate, tri-branched
filaments (fig. 39G). Female Plp1 uniramous and bisegmented; distal segment
leaf-like. Female Plps2-5 biramous, endopods with appendix interna. Telson
longer than wide, parallel-sided in proximal half, and semicircular in distal
half. Uropodal endopod oval and twice as long as wide; uropodal exopod twice
as long as wide and convex laterally but straight on distal margin with short,
spiniform setae.
Remarks. — The present new species Callianidea ryukyu is different
from the Pan-Indopacific species C. typa, because in C. ryukyu the Plp1
is uniramous, the distal segment is shorter than the proximal segment and
fingerlike (fig. 39E), whereas in C. typa the male Plp1 is uniramous and the
distal segment is elongate, bearing a small digitiform projection distolaterally
(fig. 40B).
Regarding the specimens collected in the area around Japan, Balss (1914:
90) described C. typa based on two female specimens from the Yaeyama
Islands (= Iriomote-Ishigaki Islands) and Kagoshima. However, their localities
suggest that those specimens are safely determined as the present new species,
C. ryukyu.
Type locality. — Taketomi-jima between Iriomote and Ishigaki Islands,
Ryukyu Archipelago, Japan.
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203
Distribution. — Ryukyu Islands and Takara-jima, Tokara-retto (Miyake,
1956), Akajima (Akashima), Kerama Island group, Ryukyu Archipelago; and
Kagoshima (Balss, 1914).
Etymology. — The new species is named Callianidea ryukyu after its type
locality, the Ryukyu Archipelago, Japan. The name is to be considered a noun
in apposition to the generic name.
Callianidea typa H. Milne Edwards, 1837
(fig. 40A-E)
Isea elongata Guérin-Méneville, 1832: 300, pl. 10A figs. 1-7. [Type locality: Mariana.]
[Suppressed under the plenary powers of the ICZN (Sakai & Holthuis, 1987: 92).]
Callianidea typa H. Milne Edwards, 1837a: 320, pl. 25bis figs. 8-14; H. Milne Edwards,
1838: 389; Lockington, 1878: 302; Neumann, 1878: 34; Borradaile, 1898: 1000, 1001,
1015; Borradaile, 1900: 420; De Man, 1902: 751; Borradaile, 1903: 548; Borradaile, 1904:
752; Nobili, 1906b: 113; Pesta, 1913: 678; De Man, 1928b: 21 (list), 31, pl. 1 fig. 3-3f;
Edmondson, 1944: 38, fig. 2; Holthuis, 1959: 129; Sakai, 1987a: 303 (list); ICZN, 1989: 61;
Dworschak, 1992: 218, fig. 17a-d; Liu & Zhong, 1994: 562; Poupin, 1994: 7, fig. 3, colour
pl. 1b; Poore, 1994: 113, fig. 2a; Poore, 1997: 349, figs. 1-3, 4A-B; Komai, 2000a: 236;
Sakai & Sawada, 2006: 1357, figs. 11, 12; Komai, 2008: 29; Komai & Tachikawa, 2008: 29;
Robles et al., 2009: 317.
Callianisea elongata — H. Milne Edwards, 1837a: 325.
Callianidea typus White, 1847: 71. [Type locality. — Philippine Islands.]
Callianidea elongata — Guérin-Méneville, 1857: 45; De Man, 1928b: 21, 30; Holthuis,
1953b: 51.
Callianidea mucronata Kossmann, 1880: 80; Borradaile, 1903: 548. [Type locality. — Red Sea.]
Callianassa secura Lanchester, 1902: 555, pl. 34 fig. 2. [Type locality. — Kota Bharu, Kelantan,
Malaysia.]
Callianassa (Calliactites) secura — Borradaile, 1903: 545; De Man, 1928a: 5; De Man, 1928b:
26 (list), 93, 96 (key).
Callianidea planocula Melin, 1939: 5, figs. 1-4. [Type locality. — Port Lloyd, west side of
Chichi-jima, Bonin Islands.]
Callianidea sp., Edmondson, 1944: 40, fig. 3.
Material examined. — SMF 23636, 1 female (TL/CL, 59.0/15.1 mm), Saudi-Arabia, Jana
Island (27◦ 22.240′ N 049◦ 54.000′ E), eastern coast, under stone in shallow water, 25.iv.1993,
leg. M. Apel; SMF 26511, 1 female bearing eggs, inside larvae with nauplius-eyes (TL/CL,
63.0/12.2 mm), Sokotra, Abd al-Kuri, NE coast, 12◦ 11.750′ N 052◦ 20.170′ E, 0.5 m intertidal,
6.iv.2000, leg. T. Wehe; SMF 7936, 1 male (TL/CL, 35.0/6.5 mm), Ternate, Indonesia, 1894,
leg. Kükenthal; USNM 243606, 1 male (TL/CL, 62.0/22.0 mm), Piti Bay, Guam, 20.xii.1996,
leg. L.A. Ward; USNM 1096793, 1 male (TL/CL, 36.0/8.0; 37.0/9.0 mm) and 1 female (TL/CL,
36.0/8.5 mm), Picard Island, Seychelles, Indian Ocean; ZMUC, 1 male (TL/CL, 29.0/6.4 mm),
Sunda Strait, Kei Island, 5◦ 57′ N 105◦ 32′ E, sandy mud with pumice, Danish Expedition,
31.vii.1922.
Diagnosis. — Male Plp1 uniramous and bisegmented, proximal segment
short and distal segment long, distal segment bearing small digitiform projection mesiodistally (fig. 40B, C); male Plp2 biramous, endopod with appendices
interna and masculina (fig. 40D); male Plps3-5 biramous, endopods with leaf-
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CRM 013 – Katsushi Sakai
Fig. 40. Callianidea typa H. Milne Edwards, 1837 and Paracallianidea laevicauda Gill, 1859.
A, carapace and A1-2 peduncles; B, C, F, male Plp1; D, G, male Plp2; E, male Plp3; H, male
Plp3. A, Callianidea typa SMF 26511, 1 female bearing eggs, inside larvae with naupliuseyes (TL/CL, 63.0/12.2 mm), Socotra, Abd al-Kuri, N.E. coast, 12◦ 11.750′ N 052◦ 20.170′ E,
0.5 m, intertidal; B, D, Callianidea typa, SMF 7936, male (TL/CL, 35.0/6.5 mm), Ternate,
Indonesia; C, E, Callianidea typa, ZMUC, 1 male (TL/CL, 29.0/6.4 mm), Sunda Strait, Kei
Island, Danish Expedition; F, G, H, Paracallianidea laevicauda, SMF 23607, 1 male (TL/CL,
53.0/12.1 mm), Colombia, Bahia Nenguangue, Magdalena, ca. 25 km N.E. of Santa Marta,
littoral. Scales 1 mm.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
205
like appendix interna (fig. 40E). Female Plp1 uniramous, bisegmented; distal
segment leaf-like. Female Plps2-5 biramous, endopods with leaf-like appendix
interna.
Remarks. — The female type specimen of Callianidea planocula is inaccessible, however, Melin’s (1939) description and figures of C. planocula show
that the abdominal pleuron 1 bears anterior and median setal rows subventrally
(Melin, 1939: 6, fig. 2), as in C. typa (cf. Sakai, 1992c, fig. 3A). Though some
difference is found between the two species, i.e., in C. planocula the distal
segment of female Plp1 is slender (Melin, 1939, fig. 2), while in C. typa it is
swollen proximally (Sakai, 1992c, fig. 5G), this difference probably relates to
the stage of development, which makes it difficult to distinguish one species
from the other, thus suggesting that the difference observed is not sufficiently
distinct to distinguish two separate species. Therefore, C. planocula is safely
considered synonymous with C. typa.
Type locality. — New Ireland, Bismarck Islands.
Distribution. — Red Sea — Massawa (Kossmann, 1880), Isles Muscha and
Eritrea (Nobili, 1906b), Isles Muscha (Poore, 1997); Gulf of Aden — Djibouti and Obock (Nobili, 1906b; Poore, 1997); Tanzania — Tanga (Balss,
1914); Zanzibar (Poore, 1997); Comoro Island (Poore, 1997); Madagascar
— Nosy Bé and Île Juan de Nova (Poore, 1997); Aldabra Islands (Poore,
1997); Maldives — Goidu, Goifurfejendu Atoll (Borradaile, 1904); Malay
Peninsula — Kota Bharu, Kelantan (Lanchester, 1902; De Man, 1928a); Port
Lloyd, west side of Chichi-jima (= Peel Island), Bonin Islands (Melin, 1939;
Sakai, 1992c; Komai & Tachikawa, 2008); Taiwan; Philippines — Gulf of
Davao (Poore, 1997); Indonesia — Ternate (De Man, 1902), Lombok, Paternoster Islands, Sawu, Adonara, Tanah Djampeah, and Atjatuning (De Man,
1928b), Sunda Strait, Karapang, E. Java, and Ternate (Sakai, 1992c); Papua
New Guinea — New Ireland (H. Milne Edwards, 1837a; Poore, 1997), Sariba
(Borradaile, 1900); Mariana Islands — (Guérin-Méneville, 1832; H. Milne Edwards, 1837a), Guam (Edmondson, 1944); Wake Island (Edmondson, 1944);
Rotuma (Borradaile, 1898); Funafuti (Borradaile, 1898); Western Samoa —
Savaii (Pesta, 1913; Dworschak, 1992); Tahiti (Edmondson, 1944); Tuamotu
Archipelago — Taiaro (Poore, 1997); La Paz, Baja California, Mexico (Lockington, 1878).
Genus Paracallianidea Sakai, 1992
Paracallianidea Sakai, 1992c: 17; Poore, 1997: 354.
Diagnosis. — Linea thalassinica short, extending posteriorly from orbit.
Mxp3 merus bearing subdistal spine on ventral margin. Ps2-3 propodi with
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CRM 013 – Katsushi Sakai
setal row proximally. Male Plp1 uniramous and bisegmented, distal segment
broadly triangular, bearing patch of hooklets mesially; male Plp2 biramous,
endopod with short appendix interna and elongate, rod-shaped appendix masculina, bearing unsegmented and 2- to 4-segmented respiratory filaments;
male Plps3-5 biramous, endopods with appendix interna, bearing both unsegmented and 2- to 4-segmented respiratory filaments. Female Plp1 uniramous
and bisegmented, distal segment leaf-like; female Plps2-5 biramous, endopods
with appendix interna, bearing unsegmented and 2- to 4-segmented respiratory
filaments. Uropodal exopod without transverse suture. [Adapted from Sakai,
1992c.]
Remarks. — Poore (1997: 354) did not accept the new genus Paracallianidea Sakai, 1992c, established for Callianidea laevicauda, mentioning that
“Sakai (1992c) erected a separate genus Paracallianidea, for C. laevicauda.
The general habitus of this species is very similar to that of the other species
and the differences of the pleopods 1 and 2, the male appendix interna on
pleopod 2, and the pleopodal filaments are not adequate to justify a second
genus for a single species”. However, he failed to pay attention to the features found on Plps1-2, which are important in separating one genus from the
other. Paracallianidea laevicauda, though similar to C. typa, is very different
at the generic level as follows. In P. laevicauda the male Plp1 is uniramous and
bisegmented, and the distal segment is triangular, bearing a patch of hooklets
mesially (fig. 40F); whereas in C. typa the male Plp1 is uniramous and bisegmented, and the elongate distal segment mesiodistally bears a small digitiform
projection (fig. 40B, C). As a consequence, Paracallianidea, which is distinguishable from Callianidea, should definitely be admitted as a valid genus.
Type species. — Callianidea laevicauda Gill, 1859, by original designation
and monotypy. The gender of the generic name, Paracallianidea, is feminine.
Species included. — Only the type species, Paracallianidea laevicauda
(Gill, 1859).
Paracallianidea laevicauda Gill, 1859
(fig. 40F-H)
Callianidea laevicauda Gill, 1859: 167; Rathbun, 1901: 94; Borradaile, 1903: 548; Schmitt,
1924: 79; De Man, 1928b: 21, 30; Schmitt, 1935a: 193, fig. 54; Schmitt, 1936: 375;
Rodriguez, G., 1980: 207; Rodrigues, 1983: 93; Kensley & Heard, 1991: 499, figs. 3, 4,
tab. 1A; Dworschak, 1992: 218, fig. 16a-f; Lemaitre & León, 1992: 45; Lemaitre & Ramos,
1992: 352; Hendrickx, 1995: 390 (list); Poore, 1997: 354, fig. 4C; Vargas & Cortés, 1999b:
906; Hendrickx et al., 2005: 170.
Callianidea Steenstrupii Boas, 1880: 108.
Callianidea Steenstrupi — Borradaile, 1903: 548.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
207
Callianidea laevicauda occidentalis Schmitt, 1939: 10, 26.
Paracallianidea laevicauda — Sakai, 1992c: 17.
Material examined. — SMF 23606, 2 females (TL/CL, 35.0/7.4; 55.0/11.2 mm), Colombia,
Ensenada Taganga, Magdalena, ca. 5 km N.E. of Santa Marta, intertidal, ii.1977; SMF 23607,
1 male (TL/CL, 53.0/12.1 mm), Colombia, Bahia Nenguangue, Magdalena, ca. 25 km N.E.
of Santa Marta, littoral, 26.iii.1975; SMF 23625, 1 female (TL/CL, 20.0/4.3 mm), Colombia,
Bahia Nenguangue, Magdalena, ca. 25 km N.E. of Santa Marta, root of Thalassia, 28.v.1975,
leg. M. Mercedes-Criales; SMF 9755 (ex Mus. Heidelberg), 1 male (TL/CL, 45.0/11.2 mm),
1 female (TL/CL, 48.0/10.8 mm), locality unknown.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment triangular, bearing patch of hooklets on mesial margin (fig. 40F); male Plp2 biramous, endopod and exopod with simple and bisegmented respiratory filaments, endopod subproximally bearing elongate appendix masculina with setae, and small appendix interna with distal cluster of hooklets (fig. 40). Male
Plps3-5 biramous, exopods with simple and bisegmented respiratory filaments,
endopods with appendix interna (fig. 40H). Female Plp1 uniramous and bisegmented, distal segment leaf-like. Female Plps2-5 biramous, endopods with
slender appendix interna.
Type locality. — Barbados.
Distribution. — Bahamas — San Salvador Is. (Kensley & Heard, 1991);
Jamaica (Kensley & Heard, 1991); Puerto Rico — Ensenada Honda, Culebra
(Rathbun, 1901; Kensley & Heard, 1991), near Guanica (Schmitt, 1935a);
Playa Blanca, Parque Nacional Santa Rosa (Vargas & Cortés, 1999b); U.S.
Virgin Islands — St. Johns (Kensley & Heard, 1991); British Virgin Islands —
Norman Island (Poore, 1997); St. Maarten — Grande Bay (Kensley & Heard,
1991); Antigua — English Harbour (Kensley & Heard, 1991); Barbados —
Needham’s Point and Pelican Island (Schmitt, 1924; Kensley & Heard, 1991);
Grenadines — Baradal, Tobago Cays (Kensley & Heard, 1991); Belize —
Carrie Bow Cay (Kensley & Heard, 1991); Panama — Caledonia Bay (Kensley
& Heard, 1991); Curaçao — Caracas Bay (Schmitt, 1924; Kensley & Heard,
1991); Bonaire, Kralendijk (Schmitt, 1936; Kensley & Heard, 1991); Bonaire
(Dworschak, 1992); Venezuela (Dworschak, 1992); Mexico — Socorro Island
(Schmitt, 1939); Colombia — Santa Marta (Sakai, 1992c); Gorgona Island
and Ensenada de Utr’a (Lemaitre & Ramos, 1992); Galapagos Islands —
Albemarle Island (Schmitt, 1939); Barrington Island (Poore, 1997); in shallow
water under stones.
FAMILY CALOCARIDIDAE ORTMANN, 1891
Calocaridae Ortmann, 1891: 47, 50; Lagerberg, 1908: 47 (key), 50; Stephenson, 1910: 75, 77,
189; Runnstrøm, 1925: 14.
Calocarididae — Kensley, 1989: 960; Poore, 1994: 98; Kensley, 1996d: 158; Ngoc-Ho, 2003:
443; Kensley & Chan, 1998: 262; Felder, 2001: 440; Sakai & Ohta, 2005: 82.
Diagnosis. — Rostrum acutely or narrowly triangular, lateral margins
extending posteriorly onto gastric region. Gastric region with median carina;
submedian and lateral carinae present or absent; postcervical carina present
or not; linea thalassinica absent; cervical groove distinct. Eyestalks reduced
and cornea not pigmented. A1 segment 1 twice as long as segment 2; A2
scaphocerite distinct or indistinct. Maxilla 2 scaphognathite with posterior
whip. Mxp3 with exopod. Abdominal pleuron 1 narrow; pleuron 2 broadened.
Ps3-4 propodi more or less elongate; P4 coxa more or less cylindrical. Plp1
uniramous and bisegmented; distal segment diversified in shape, bearing
cluster of hooklets; Plp2 biramous, endopod bearing distally enlarged un- or
bisegmented appendix masculina with slender appendix interna with hooklets
mesioproximally, or bearing no appendix masculina; exopod consisting of
proximal segment and multiarticulate flagellum; Plps3-5 biramous, endopods
with rod-like appendix interna. Uropodal endopod oval and uropodal exopod
with transverse suture. Hermaphroditic.
Remarks. — The family Calocarididae includes 11 genera that are characterized as follows. The male Plp2 is biramous, the endopod bears distally
an un- or bisegmented appendix masculina provided proximally with a small
appendix interna with a cluster of hooklets, but exceptionally in Calocarisopsis gen. nov., it bears an appendix interna, lacking an appendix masculina.
Kensley (1989: 965) mentioned that three synapomorphies separate the Calocarididae from the Axiidae (s. l.) “1. Invariable hermaphroditism. 2. Enlargement of the appendix masculina of pleopod 2, along with loss of the setose
distal element of the endopod. 3. Eye reduction and loss of eye pigment”.
However, it was suggested by Xuân & Ngoc-Ho (2006: 1161) that Eutrichocheles modestus (Herbst, 1796) in the family Axiidae is hermaphroditic, so
that hermaphroditism is not always so important a characteristic as to separate
the Calocarididae from the Axiidae. The following genera, such as Bouvieraxius, Calaxius (C. acutirostris Sakai & De Saint Laurent, 1989), Paraxiopsis
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CRM 013 – Katsushi Sakai
(P. brocki (De Man, 1888b), P. hispidus Kensley, 1996c, P. paulayi (Kensley,
2003)), and Levantocaris are not included in the Calocarididae but in the Axiidae, though they are hermaphroditic. Instead, the male Plp2 is important in
separating one family from the other, because the male Plp2 shows very distinct differences in morphology. In the Calocarididae the male Plp2 endopod
bears distally an enlarged appendix masculina provided proximally with an appendix interna, whereas in the Axiidae the male Plp2 endopod bears medially
or at its proximal third an appendix masculina provided proximally with an
appendix interna.
Type genus. — Calocaris Bell, 1846.
Genera included. — Ambiaxius Sakai & De Saint Laurent, 1989; Briancaris
Sakai & Ohta, 2005; Calastacus Faxon, 1893; Calaxiopsis Sakai & De Saint
Laurent, 1989; Calocaris Bell, 1846; Calocarisopsis gen. nov.; Eucalastacus
Sakai, 1992a; Lophaxiopsis gen. nov.; Lophaxius Kensley, 1989; Nipponcalaxiopsis gen. nov.; Paracalocaris Sakai, 1991.
K EY
TO THE
G ENERA
OF THE
FAMILY C ALOCARIDIDAE
1 – Plp2 endopod without appendix masculina . . . . . . . . . . . . . . . . Calocarisopsis gen. nov.
– Ppl2 endopod with appendix masculina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Plp2 appendix masculina boot-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Plp2 appendix masculina simple, un- or bisegmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 – Plp2 endopod bearing distally a boot-shaped appendix masculina, in such a shape, that
between the “foot part” of the “boot” and the “leg” there is an angle of 90◦ . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ambiaxius
– Plp2 endopod bearing distally a boot-shaped appendix masculina, in such a shape, that
between the “foot part” of the “boot” and the “leg” there is an angle of 30◦ . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Briancaris
4 – Appendix masculina bisegmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Appendix masculina unsegmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 – Pleura 3-5 with sharp anteroventral spine, and triangular on posteroventral angle . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaxiopsis
– Pleura 3-5 unarmed ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calastacus
6 – Carapace granulate or setose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Carapace smooth; Plp1 distal segment elongate-triangular or pear-shaped . . . . . . . . . . 8
7 – Carapace granulate; Plp1 distal segment trispinose distally . . . . . . . . . . . . Paracalocaris
– Carapace setose; Plp1 unknown . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophaxiopsis gen. nov.
8 – Plp1 distal segment shaped as elongate triangle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
– Plp1 distal segment pear-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9 – Uropodal exopod without distal flap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucalastacus
– Uropodal exopod with distal flap . . . . . . . . . . . . . . . . . . . . . . Nipponcalaxiopsis gen. nov.
10 – Eyes flattened, mesially contiguous, cornea unpigmented . . . . . . . . . . . . . . . . . Calocaris
– Eyes rounded, not mesially contiguous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophaxius
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
211
Genus Ambiaxius Sakai & De Saint Laurent, 1989
Ambiaxius Sakai & De Saint Laurent, 1989: 54; Poore, 1994: 99 (key); Sakai, 1994: 177; Sakai,
1995: 79; Kensley, 1996e: 486; Sakai & Ohta, 2005: 82; Poore & Collins, 2009: 237.
Callistocaris Kensley, 1989: 961 [partim = Briancaris Sakai & Ohta, 2005].
Not: Ambiaxius Kensley, 1996e: 486 [= Briancaris Sakai & Ohta, 2005].
Diagnosis. — Rostrum styliform and curved upward, lateral margins unarmed, extending posteriorly onto gastric region. Gastric region convex, bearing median and lateral carinae, but no submedian carinae. Anterolateral margins of carapace unarmed or armed with a pair of small teeth. Cervical groove
distinct. Eyes and stalks not differentiated, flattened; cornea unpigmented. A2
scaphocerite prominent; penultimate segment long. Pl asymmetrical. Ps3 and 5
coxae with genital pores. All gills poorly developed or devoid of gill-branches.
No pleurobranchs. Abdominal pleura smooth on lateral surface and truncate
ventrally. Plp1 uniramous and bisegmented; proximal segment elongate and
flattened, and distal segment divergent distally, bearing terminal plate with
patch of hooklets; Plp2 biramous and slender; endopod bearing distally a bootshaped appendix masculina with marginal setae, with an angle of 90◦ between
the “foot” of the “boot” and the “leg”, provided proximally with fused appendix interna with cluster of hooklets; exopod consisting of proximal segment and multiarticulate flagellum; Plps3-5 biramous and slender, endopods
with appendix interna. Telson longer than broad, bearing obscurely marked
carinae on dorsal surface, but no posteromedian spine. Uropodal exopod with
non-dentate transverse suture. Hermaphroditic. [Adapted from Sakai & Ohta,
2005.]
Remarks. — The type species, Ambiaxius alcocki, which was described
on the basis of an adult female as Calocaris alcocki by McArdle (1900),
is different from the type species of Calocaris, C. macandreae Bell, 1846,
because he described it as “Rostrum long, narrow, and curved upwards,
deeply grooved on its upper surface” (McArdle, 1900: 476), whereas in
C. macandreae the rostrum is stoutly triangular and not curved upwards,
but horizontal or curved downward. Distinctive differences are also observed
between their Plps1-2.
Type species. — Calocaris Alcocki McArdle, 1900, by original designation.
The gender of the generic name, Ambiaxius, is masculine.
Species included. — Ambiaxius alcocki (McArdle, 1900); A. cf. alcocki
(McArdle, 1901); A. franklinae Sakai, 1994; A. surugaensis Sakai & Ohta,
2005; Ambiaxius sp. Poore & Collins, 2009.
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CRM 013 – Katsushi Sakai
Ambiaxius alcocki (McArdle, 1900)
Calocaris Alcocki McArdle, 1900: 476; Balss, 1925: 209.
Calocaris alcocki — Alcock, 1901: 190; Alcock & McArdle, 1901, pl. 50 fig. 4; Stebbing,
1915: 59; Stebbing, 1917: 27, pl. 91.
Calocaris (Calocaris) Alcocki — Borradaile, 1903: 539; De Man, 1925d: 7 (list), 116 (key).
Calocaris (Calocaris) alcocki — Barnard, 1950: 502, fig. 93g-h; Kensley, 1981b: 30.
Ambiaxius alcocki — Sakai & De Saint Laurent, 1989: 54; Sakai, 1995: 80, figs. 1-5; Kensley,
1996e: 487 (list); Takeda, 1997: 250; Sakai & Ohta, 2005, fig. 8.
Type locality. — Off northeast Sri Lanka, 8◦ 36.15′ N 81◦ 20.30′ E, 992 m.
Distribution. — South Africa — Cape Natal (Stebbing, 1915; Sakai, 1995);
S.W. Indian Ocean (Sakai, 1995); Sri Lanka (McArdle, 1900); Japan — Suruga
Bay (Sakai, 1995; Takeda, 1997); New Caledonia (Sakai & De Saint Laurent,
1989); 792-1630 m.
Ambiaxius cf. alcocki (McArdle, 1901)
Calocaris cf. alcocki — McArdle, 1901.
Ambiaxius cf. alcocki — Kensley, 1996e: 487 (list).
Distribution. — Southwest Indian Ocean, 1000 m.
Ambiaxius franklinae Sakai, 1994
Ambiaxius franklinae Sakai, 1994: 177, 200 (key), figs. 1, 2; Poore & Collins, 2009: 238.
Type locality. — North West Shelf, Western Australia, 17◦ 19.76′ S
147◦ 28.05′ E, 1310-1357 m.
Distribution. — Western Australia — North West Shelf (Sakai, 1994); 13101357 m.
Ambiaxius surugaensis Sakai & Ohta, 2005
Ambiaxius alcocki — Sakai, 1995: 80, figs. 1-5; Takeda, 1997: 250.
Ambiaxius surugaensis Sakai & Ohta, 2005: 84, figs. 9, 10.
Type locality. — W. off Matsuzaki, Suruga Bay, 490-547 m.
Distribution. — Suruga Bay; 490-547 m.
Ambiaxius sp. Poore & Collins, 2009
Ambiaxius sp., Poore & Collins, 2009: 238.
Distribution. — Off Cape Leveque, Western Australia; 698-700 m.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
213
Genus Briancaris Sakai & Ohta, 2005
Callistocaris Kensley, 1989: 961 (partim).
Ambiaxius — Kensley, 1996e: 486.
Briancaris Sakai & Ohta, 2005: 87.
Diagnosis. — Rostrum short and triangular, denticulate on lateral margins.
Gastric region convex, bearing median carina and short lateral carinae, but no
submedian carinae. Anterolateral margins of carapace unarmed or armed with
a pair of small teeth. Cervical groove distinct. Eyes rounded in lateral view and
contiguous to carapace; cornea unpigmented. A2 scaphocerite prominent; A2
penultimate segment long. P1 asymmetrical. Plp1 uniramous and bisegmented,
proximal segment flattened, and distal segment divergent distally, bearing
ovate terminal plate with patch of hooklets; male Plp2 biramous, endopod
stem-like, bearing distally a boot-shaped appendix masculina with setae on
distal margin and distal protrusion with apical setae, with an angle of 30◦
between the “foot” of the “boot” and the “leg”, and provided proximally with
appendix interna bearing a patch of hooklets; exopod consisting of proximal
segment and multiarticulate flagellum. Telson longer than wide, and unarmed
on lateral and posterior margins. Uropodal exopod with transverse suture.
Dimorphic.
Remarks. — Briancaris is separated from Ambiaxius, because in Briancaris
the rostrum is triangular and denticulate on the lateral margins; Plp2 is
biramous, and the endopod bears distally a boot-shaped appendix masculina,
with an angle of 30◦ between the “foot” of the “boot” and the “leg”, that is
provided proximally with an appendix interna with a patch of hooklets. In
Ambiaxius, by contrast, the rostrum is styliform and almost unarmed laterally;
Plp2 is biramous, and the endopod distally bears a boot-shaped appendix
masculina, with an angle of 90◦ between the “foot” of the “boot” and the “leg”,
that is slightly concave on the distal margin, with marginal setae and an apical
protrusion with setae, and is provided mesioproximally with a small appendix
interna.
Type species. — Ambiaxius japonicus (Kensley, 1996e), by original designation. The gender of the generic name, Briancaris, is feminine.
Species included. — Briancaris aberrans (Bouvier, 1905); B. japonica
(Kensley, 1996e); B. foveolata (Kensley, Lin & Yu, 2000).
Briancaris aberrans (Bouvier, 1905)
Calocaris aberrans Bouvier, 1905: 805; Bouvier, 1917: 120; Bouvier, 1925: 453, figs. 17-21,
pl. 10 figs. 6, 7; Balss, 1925: 209.
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CRM 013 – Katsushi Sakai
Calocaris (Calocaris) aberrans — De Man, 1925d: 7 (list), 116 (key).
Ambiaxius aberrans — Sakai & De Saint Laurent, 1989: 55; Kensley, 1996e: 487 (list).
Type locality. — Santa Lucia, Antilles, 772 m.
Distribution. — Santa Lucia (Bouvier, 1905); 772 m.
Briancaris japonica (Kensley, 1996)
Ambiaxius japonicus Kensley, 1996e: 487, fig. 11, table 5.
Type locality. — Off Ose-saki, Japan, 34◦ 58′ 40′′ N 138◦ 43′ 35′′ E, “Albatross” Sta. 5056, 472 m.
Distribution. — Japan — Off Ose-saki (Kensley, 1996e); 472 m.
Briancaris foveolata (Kensley, Lin & Yu, 2000)
Ambiaxius foveolatus Kensley, Lin & Yu, 2000: 215, figs. 6, 7E.
Type locality. — Tai-Shi fishing port, I-Lan County, N.E. Taiwan, 400500 m.
Distribution. — Only known from the type locality.
Genus Calastacus Faxon, 1893
Calastacus Faxon, 1893: 194; Faxon, 1895: 105; Borradaile, 1903: 539; Stebbing, 1910: 367;
Schmitt, 1921: 112; Vinogradov, 1950: 223; Balss, 1957: 1580; De Saint Laurent, 1972: 353,
354; Kensley, 1989: 961; Sakai & De Saint Laurent, 1989: 59; Poore, 1994: 98 (key); Poore
& Collins, 2009: 247.
Calastacus s. str. — Alcock, 1901: 191; Borradaile, 1903: 53; De Man, 1925d: 2 (key), 116;
Barnard, 1950: 502.
Diagnosis. — Rostrum acutely elongate or triangular, lateral margins unarmed, shortly extending posteriorly onto gastric region as lateral carinae with
supraorbital spine. Gastric region convex, bearing median carina, no submedian carinae, but submedian rows of 2-3 teeth may be present or not. Anterolateral margin of carapace unarmed. Cervical groove distinct only in dorsal region; no postcervical carina. Eyestalks obsolete and flattened on surface; cornea also obsolete. A2 scaphocerite slender and elongate. P1 chelate
and subequal. Ps3-5 simple. Ps3 and 5 coxae with genital pores. No pleurobranchs. Abdominal pleura smooth on lateral surface and truncate ventrally,
with or without teeth. Plp1 uniramous and bisegmented; proximal segment
stem-shaped, and distal segment broad or elongate, and double-lobed by longitudinal median suture, mesial lobe proximally with a patch of hooklets. Plp2
biramous, endopod bearing slender, bisegmented appendix masculina fused
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
215
proximally with appendix interna bearing a patch of hooklets; exopod consisting of proximal segment and multiarticulate flagellum; Plps3-5 endopods
with appendix interna. Telson longer than wide and rounded on posterior
margin without median spine. Uropodal exopod with transverse suture. Hermaphroditic. [Adapted from Sakai & De Saint Laurent, 1989.]
Remarks. — The genus Calastacus Faxon, 1893 is closely similar to
the genus Calaxiopsis Sakai & De Saint Laurtent, 1989 in that the Plp1
is uniramous and bisegmented, consisting of an elongate proximal segment
and a broad distal segment, and Plp2 is biramous, and the endopod bears a
slender bisegmented appendix masculina with setae on the mesial margin; the
proximal segment is fused at its base with a small appendix interna with a
cluster of hooklets. However, those two genera are also clearly different from
each other: in Calastacus the rostrum is unarmed on the lateral margins, and
the gastric region bears no submedian carinae, whereas in Calaxiopsis the
rostrum is armed with lateral spines, and the gastric region bears submedian
carinae.
Calastacus crosnieri is very characteristic and different from the other
species in the genus Calastacus, in that the Plp1 distal segment is broad and
sinuate on the mesial margin, bearing mesioproximally an appendage with
hooklets; however, Plp2 is biramous, and the endopod bears a bisegmented
appendix masculina with a small appendix interna proximally, as in C. colpos,
C. stilirostris, C. mexicanus, and C. laevis, so that C. crosnieri is regarded as a
member of the species included in Calastacus.
Type species. — Calastacus stilirostris Faxon, 1893, by original designation
and monotypy. The gender of the generic name, Calastacus, is masculine.
Species included. — Calastacus colpos Kensley, 1996d; C. crosnieri Kensley & Chan, 1998; C. inflatus Komai et al., 2009; C. laevis (De Saint Laurent,
1972); C. mclaughlinae (Lin & Komai, 2006); C. mexicanus Kensley, 1996d;
C. myalup Poore & Collins, 2009; C. stilirostris Faxon, 1893.
K EY
TO THE
S PECIES
OF THE
G ENUS CALASTACUS
1 – Uropodal exopod with unarmed suture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Uropodal exopod with armed suture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 – Mxp3 merus with two distinct spines on ventral margin . . . . . . . . . . . . . . . . . . . C. myalup
– Mxp3 merus with only one distinct spine on ventral margin . . . . . . . . . . . . . . . C. inflatus
3 – Gastric region with submedian rows of 2-3 spines . . . . . . . . . . . . . . . . . . . C. mclaughlinae
– Gastric region without submedian rows or carinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – P1 palm with spines on dorsal margin and lateral surface . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– P1 palm with one subdistal spine on dorsal margin, but unarmed on lateral surface . . . 6
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CRM 013 – Katsushi Sakai
5 – Telson armed with posteromedian spine and posterolateral spines . . . . . . . C. mexicanus
– Telson unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stilirostris
6 – Plp1 bisegmented, distal segment broad and double-lobed distally, mesial lob sinuate,
bearing mesioproximally appendage with hooklets . . . . . . . . . . . . . . . . . . . . . . C. crosnieri
– Plp1 bisegmented, distal segment elongate and double-lobed in plough-shape . . . . . . . 7
7 – Eyes flattened; A2 scaphocerite longer than distodorsal spine of A2 segment 2; in Plp1
mesial lobe of distal segment triangular, distally with seta . . . . . . . . . . . . . . . . . . C. colpos
– Eyes subspherical; A2 scaphocerite about same length of distodorsal spine of A2
segment 2; in Plp1 mesial lobe of distal segment rounded, distally without seta . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laevis
Calastacus colpos Kensley, 1996
Calastacus colpos Kensley, 1996d: 159, figs. 1, 2.
Material examined. — USNM 243393, holotype, male (TL/CL, 30.0/10.0 mm), Gulf of
Mexico, 27◦ 17.5′ N 95◦ 08.5′ W, 109 m, 21.xi.1968, leg. W.E. Pequegnat.
Diagnosis. — Carapace faintly rugose. Rostrum spiniform and unarmed,
reaching anteriorly to level of A2 penultimate segment; supra-orbital spines
directed anterodorsally. Median carina weakly defined, bearing tubercle at anterior third of gastric region; cervical groove distinct dorsally, becoming obsolete anteroventrally; suprabranchial groove faint. Eyes with flattened cornea,
contiguous along midline, and unpigmented. A1 segment 1 subequal in length
to segments 2 and 3 combined. A2 segment 2 with slender distolateral spine;
scaphocerite slender, almost reaching distal part of penultimate segment. Maxilla 2 scaphognathite with elongate posterior whip. Mxp3 ischium smooth on
lateral margin; ischial crest bearing about 20 denticles; merus with two spines
mesiodistally. P1 subequal and similar in shape; ischium with subdistal spine
on ventral margin; merus with about 6 spines on ventral margin, and subdistal spine on dorsal margin; carpus unarmed; palm slightly longer than fingers,
bearing subdistal spine on dorsal margin; dactylus strongly curved downward;
cutting edges of fingers finely serrated. P2 fingers shorter than palm.
Abdominal pleura 2-5 truncate and pleuron 6 rounded ventrally, bearing
no longitudinal carinae between terga and pleura; pleuron 1 shortest and narrowed posteroventrally; pleuron 2 broadest. Plp1 uniramous and bisegmented,
distal segment double-lobed, mesial lobe longer than lateral one and triangular,
with distal seta and proximal patch of hooklets. Plp2 biramous, endopod with
bisegmented appendix masculina distally, proximal segment bearing mesioproximally appendix interna with patch of hooklets, and distal segment shorter
than proximal one and tapering; exopod consisting of proximal segment and
multiarticulate flagellum.
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217
Telson 1.6 times as long as its greatest width, and roughly rectangular
in shape, bearing tiny, movable posterolateral spine on each side; posterior
margin slightly convex. Uropodal endopod with one or two spines on lateral
margin. Uropodal exopod bearing distinct, movable spine on lateral margin and
transverse suture with 3-4 spinules on dorsal surface. [Adapted from Kensley,
1996d.]
Type locality. — North-western Gulf of Mexico, 27◦ 17.5′ N 95◦ 08.5′ W,
1098-1171 m.
Distribution. — Gulf of Mexico (Kensley, 1996d); 339-1171 m.
Calastacus crosnieri Kensley & Chan, 1998
Calastacus crosnieri Kensley & Chan, 1998: 262, figs. 1C, 5; Tsang et al., 2008: 218, 219; Chu
et al., 2008.
Material examined. — USNM 253355, paratype, male (TL/CL, 30.0/12.0 mm), Ta-Chi, ILan County, N.E. Taiwan, 350 m, 10.vi.1993, leg. commercial trawler.
Diagnosis. — Carapace somewhat inflated and glabrous. Rostrum spiniform
and dorsally grooved, bearing strong supraocular spines at base; median carina
weakly indicated, rounded and stronger at base of rostrum. Cervical groove
faintly discernible dorsally. Eyes rounded, unpigmented, not contiguous. A2
scaphocerite slender and acute, reaching middle of penultimate segment.
P1 chelipeds subequal, ischium bearing 5 ventral spines; merus with 9-10
spines on ventral margin and single distal spine on dorsal margin; carpus
unarmed; palm with a single distal tooth on carinate dorsal margin; fingers
0.7 times length of palm. Ps2-5 unarmed, P4 longest. Pleuron 1 triangular and
subacute ventrally. Pleura 2-6 broadly rounded ventrally. Plp1 uniramous and
bisegmented, distal segment broad, sinuate mesially, and double-lobed distally,
bearing mesioproximally appendage with hooklets. Plp2 biramous, endopod
bearing distally bisegmented appendix masculina with setae on mesial margin,
provided mesioproximally with small, proximally fused appendix interna;
exopod consisting of proximal segment and multiarticulate flagellum. Telson
1.7 times as long as wide, bearing proximal lobe on lateral margins and a
single movable spinule on each posterolateral angle; posterior margin rounded.
Uropodal endopod with distolateral spine. Uropodal exopod bearing transverse
suture with 9 spinules and movable spine at distolateral angle. [Adapted from
Kensley & Chan, 1998.]
Type locality. — Ta-Chi, I-Lan County, north-eastern coast of Taiwan, sandy
mud bottom, 350 m.
Distribution. — Only known from the type locality.
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CRM 013 – Katsushi Sakai
Calastacus inflatus Komai, Lin & Chan, 2009
Calastacus inflatus Komai et al., 2009: 24-30.
Description. — Carapace stout; integument fragile and naked. Rostrum
spiniform, unarmed, failing to reach distal margin of A1 second segment.
Eyestalks slightly flattened dorsoventrally, hardly mobile, lacking pigment.
A1 peduncle moderately stout; segment 1 subequal in length to distal two
segments combined. A2 peduncle moderately stout, longer than A1 peduncle;
second segment with weakly produced, rounded dorsolateral distal angle; third
segment with small spine at ventromesial distal angle; fourth segment longest;
scaphocerite slender and spiniform, reaching midlength of fourth segment;
flagellum missing. Epistome produced in rounded lobe and naked.
Carapace somewhat compressed laterally and faintly rugose on branchiostegites, otherwise almost smooth; dorsal surface strongly convex, and gastric region much higher than base of rostrum; supraorbital spines small and directed
anterodorsally; gastric median carina obsolete; cervical groove very shallow.
Mxp3 moderately stout; ischium unarmed on ventral margin, crista dentata
(not illustrated) with row of about 20 small acute teeth; merus with small subdistal spine on ventral margin; carpus with blunt dorsodistal projection; exopod
overreaching distal margin of merus.
P1 slightly unequal (right larger than left), but similar in shape. In larger
cheliped coxa with minute denticle on ventrodistal margin; ischium with some
minute denticles mesially on ventral margin; merus slightly widened distally,
dorsal and ventral margins each with moderately small subterminal spine,
ventral margin also with two tiny denticles mesially; carpus about half length
of palm; palm slightly widened distally, about twice as long as wide, with small
subdistal spine dorsally and with sparse tufts of short setae on lateral surface;
dorsal and ventral surfaces rounded, each bearing a row of long, stiff setae, and
a row of minute denticles is also present on ventral surface; dactylus subequal
in length to palm, terminating in slender, curved tip, lateral surface with weak
longitudinal carina adjacent to dorsal margin, cutting edge with row of blunt
or subacute teeth; fixed finger also terminating in slender, curved tip crossing
tip of dactylus, and cutting edge with a row of small triangular teeth; scattered
tufts of stiff setae on surfaces of both fingers.
Smaller (left) cheliped broken. P2 chelate and moderately slender, reaching
anterolateral margin of carapace by tip of merus; ischium very short and
unarmed; merus with few setae subdistally on dorsal margin and a row of
long setae on ventral margin, bearing no spination; carpus slighty shorter than
palm, bearing row of long setae on dorsal surface, and few setae on ventral
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
219
surface; palm about 2.7 times as long as wide, bearing a row of long setae
on sharply carinate dorsal and ventral margins; dactylus about half length of
palm, bearing a row of stiff setae on sharply carinate dorsal surface, and a row
of minute corneous spinules on cutting edge; fixed finger also with a row of
minute spinules on cutting edge.
P3 long, slightly overreaching anterolateral margin of carapace by tip of
merus; merus and ischium unarmed; carpus about half length of propodus;
dactylus 0.3 times as long as propodus, and slightly depressed dorsoventrally;
propodus with tufts of stiff setae on dorsal and ventral margins, bearing slender
terminal spine at ventrodistal margin. Ps4-5 damaged.
Plp1 uniramous and bisegmented; proximal segment stem-shaped, and distal segment broad or elongate, and double-lobed by longitudinal median suture, mesial lobe proximally with patch of hooklets. Plp2 biramous, endopod
bearing slender bisegmented appendix masculina, fused proximally with appendix interna bearing patch of hooklets; exopod consisting of proximal segment and multiarticulate flagellum.
Abdomen dorsoventrally flattened. Pleuron 1 shortest and narrowed posteroventrally; pleura 2-6 rounded ventrally (pleuron 4 damaged on left side).
Telson 1.8 times as long as wide and roughly rectangular, bearing faint proximal convexity but no spines on lateral margins and on gently convex posterior margin. Uropodal endopod broadened distally and tear-shaped, bearing
no marginal spines; dorsal carinae faint. Uropodal exopod without marginal
spines, bearing unarmed suture on dorsal surface; lateral margin slightly convex. [Adapted from Komai et al., 2009.]
Type locality. — Platas Island, South China Sea, 1400 m.
Distribution. — Only known from the type locality.
Calastacus laevis De Saint Laurent, 1972
(fig. 41C-D)
Calastacus laevis De Saint Laurent, 1972: 348, figs. 1-10; Sakai & De Saint Laurent, 1989: 60;
Kensley, 1996d: 159 (list); Costello et al., 2001: 289.
Material examined. — MNHN TH 152, holotype, 1 male (TL/CL, 45.0/15.0), Bay of Biscay,
Spain, “Thalassa” Sta. W 377.
Diagnosis. — Plp1 (fig. 41C) bisegmented; proximal segment stem-like,
distal segment double-lobed, mesial lobe rectangular, with patch of hooklets
but without distal seta, and lateral lobe elongate. Male Plp2 (fig. 41D) biramous, endopod bearing bisegmented appendix masculina provided mesioproximally with small, proximally fused appendix interna with a patch of hooklets;
220
CRM 013 – Katsushi Sakai
Fig. 41. Calastacus stilirostris Faxon, 1893 and Calastacus laevis De Saint Laurent, 1972.
A, carapace, lateral view; B, D, male Plp2; C, male Plp1. A, B, Calastacus stilirostris, USNM
21064, syntype, male (TL/CL, 51.0/16.0 mm, distal part of rostrum missing), off Acapulco; C,
D, Calastacus laevis MNHN Th 152, holotype, male (TL/CL, 45.0/15.0), Bay of Biscay, Spain.
Scales 1 mm.
exopod consisting of proximal segment and multiarticulate flagellum. Plps3-5
endopods with appendix interna.
Remarks. — The male Plps1-2 are similar to those of C. colpos Kensley,
1996d, C. mexicanus Kensley, 1996d, and C. mclaughlinae (Lin & Komai,
2006).
Type locality. — Bay of Biscay, Spain, 43◦ 43.5′ N 04◦ 27′ W, 950-1000 m.
Distribution. — Only known from the type locality.
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221
Calastacus mclaughlinae (Lin & Komai, 2006)
Calaxiopsis mclaughlinae Lin & Komai, 2006: 399, figs. 1-4.
Remarks. — Lin & Komai (2006) described the present species as Calaxiopsis mclaughlinae, while their species does not belong in the genus Calaxiopsis, but instead in the genus Calastacus, because they differ from each
other as follows. In Calaxiopsis the rostrum is elongate and armed with lateral spines; pleura 3-5 bear a sharp spine anteroventrally; Plp1 uniramous and
bisegmented, with the proximal segment short and the distal segment slender
and elongate, distally forming a spear-head in shape; Plp2 biramous, with the
endopod rectangular, bearing bisegmented appendix masculina, proximal segment elongate and provided basally with a small appendix interna, and distal
segment short. In Calastacus, in contrast, the rostrum is spiniform and unarmed on the lateral margins; and pleura 3-5 bear no sharp spine anteroventrally; Plp1 uniramous and bisegmented with proximal segment stem-shaped,
and distal segment broad or elongate, and double-lobed by a longitudinal median suture, mesial lobe proximally with patch of hooklets; Plp2 biramous, endopod bearing slender bisegmented appendix masculina fused proximally with
appendix interna bearing patch of hooklets. Therefore, Calaxiopsis mclaughlinae is reclassified under the genus Calastacus, as C. mclaughlinae, though its
gastric region bears submedian rows of interspaced spines.
Type locality. — Vicinity of Taiwan 24◦ 52.02′ N 122◦ 03.11′ E; 572-605 m.
Distribution. — Off northern Taiwan, 24◦ 51.17′ N 122◦ 04.97′ E – 24◦ 52.02′ N
122◦ 03.11′ E; 572-800 m.
Calastacus mexicanus Kensley, 1996
Calastacus mexicanus Kensley, 1996d: 162, figs. 3, 4.
Material examined. — USNM 211496, holotype, male (TL/CL, 45.0/15.5 mm), Gulf of
Mexico, 28◦ 01.04′ N 85◦ 39.38′ W, 629 m, 16.v.1985, det. Kensley.
Type locality. — Gulf of Mexico, 28◦ 01.04N 85◦ 39.38W, 629 m.
Distribution. — Gulf of Mexico (Kensley, 1996d); 624-640 m.
Calastacus myalup Poore & Collins, 2009
Calastacus myalup Poore & Collins, 2009: 248, figs. 17, 42.
Description. — Rostrum 0.5 times length of front-to-cervical groove, slender and acutely triangular, and unarmed anterior to supraocular spine on lateral
222
CRM 013 – Katsushi Sakai
margin, not continuous with gastric lateral carina. Supraocular spines prominent. Eyestalk 0.4 length of rostrum; cornea unpigmented. A1 peduncle reaching two-thirds along A2 segment 4. A2 segment 1 with sharp spine and spinule
on ventrodistal margin; segment 2 with slender distal spine, directed slightly
inwards, reaching distally to middle of segment 4; scaphocerite slender and
directed slightly upwards, reaching distally to middle of A2 segment 4; segment 3 with sharp distal spine on ventral margin; segment 4 little longer than
segment 2 (excluding distal spine); segment 5 about half length of segment 4.
Carapace smooth except for slight rugosity between supraocular spines; gastric
median carina unarmed; gastric lateral and submedian carinae absent.
P1 coxa with spine; basis unarmed; remainder missing. P2 ischium unarmed; merus unarmed; carpus 0.7 length of chela. P3 merus unarmed; propodus three times as long as dactylus, bearing oblique rows of simple setae but
no robust setae. P4 unarmed; propodus three times as long as dactylus, bearing
oblique rows of simple setae distally but no robust setae. P5 unarmed; propodus with oblique rows of simple setae distally but without robust setae; fixed
finger short and rounded; dactylus broken. Maxilliped 3 basis unarmed; ischium unarmed on ventral margin, bearing crista dentata with 20 teeth; merus
with two long spines on distal half of ventral margin; carpus unarmed.
Abdominal pleuron 1 twice as deep as middorsal length, short, and rounded
ventrally; pleuron 2 broad, lateral length 1.3 times dorsal length, and rounded
anteroventrally; pleura 3-6 rounded ventrally. Plp1 bisegmented, distal segment 0.8 length of proximal segment and leaf-like, bearing hooklets on proximal broad triangular lobe; Plp 2 endopod with bisegmented appendix masculina bearing proximally thumb-like appendix interna; exopod half length
of endopod. Telson 1.6 times as long as wide, widest proximally, approximately parallel-sided posterior to proximal process, and semicircular in posterior third, bearing no marginal spines; dorsal surface with unarmed obsolete
oblique rib. Uropodal endopod two times as long as wide and unarmed marginally, bearing longitudinal ridge without spines on dorsal surface. Uropodal
exopod twice as long as wide, bearing no spines on lateral margin but one
fixed spine and one robust seta at distolateral angle; dorsal surface with two
unarmed obsolete longitudinal ribs, and transverse suture with 11 robust setae
of varying lengths. [Adapted from Poore & Collins, 2009.]
Type locality. — Off Bunbury, Western Australia, 397-423 m.
Distribution. — Only known from the type locality.
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223
Calastacus stilirostris Faxon, 1893
(fig. 41A-B)
Calastacus stilirostris Faxon, 1893: 194; Faxon, 1895: 106, pl. 27 fig. 1-1f; Sakai & De Saint
Laurent, 1989: 59; Hendrickx, 1995: 390 (list); Kensley, 1996d: 159 (list).
Calocaris (Calastacus) stilirostris — Borradaile, 1903: 539; De Man, 1925d: 8 (list), 117 (key).
Calastacus stylirostris — Balss, 1925: 209.
Material examined. — USNM 21064, syntype, 3 males (TL/CL, 51.0/16.0 mm, distal part of
rostrum missing; TL/CL, 14.0/4.9-16.0/5.3 mm), off Acapulco, Mexico, 11.iv.1891, leg. U.S.
Fish Commission Steamer “Albatross”, Sta. 3418.
Remarks. — Faxon (1893: 107) examined 9 male specimens, three of
which were examined in the present revision as follows. The carapace bears
a median carina, as shown by Faxon (1895: 106), who described that “from
the root of rostrum the median dorsal line of the carapace is convex; a light
median carina runs along the back from the base of the rostrum, fading out
before reaching the hind border of the carapace”. The A2 scaphocerite is
shorter than the distodorsal spine of segment 2 (fig. 41A). The male Plp2
(fig. 41B) is biramous, and the endopod distally bears a bisegmented appendix
masculina, the proximal segment of which is mesioproximally provided with
a tiny appendix interna.
Type locality. — Off Acapulco, 1098 m.
Distribution. — Acapulco (Faxon, 1893); Bay of Panama; 1098-1200 m.
Genus Calaxiopsis Sakai & De Saint Laurent, 1989
Calaxiopsis Sakai & De Saint Laurent, 1989: 60; Poore, 1994: 99 (key).
Diagnosis. — Rostrum elongate and acutely triangular, lateral margins with
4 distinct spines, extending posteriorly onto anterior part of gastric region.
Anterolateral margins of carapace unarmed. Gastric region convex with 5 carinae. Cervical groove only on dorsal region; no postcervical carina. Eyes subspherical and contiguous to carapace; cornea unpigmented. P1 subequal. Both
Ps3 and 5 coxae with genital pore. Pleura 2-5 triangular or subsquare, bearing small anteroventral spine or not, and sharp distinct posteroventral spine or
not. Pleurobranchs absent. Plp1 uniramous and bisegmented; proximal segment short and distal segment slender and elongate to form distally spearhead shape. Plp2 biramous; endopod rectangular, bearing bisegmented appendix masculina, proximal segment elongate and provided basally with small
appendix interna, and distal segment short; exopod consisting of proximal
segment and multiarticulate flagellum. Plps3-5 endopods with appendix interna. Telson longer than broad, and rounded on posterior margin without me-
224
CRM 013 – Katsushi Sakai
dian spine. Uropodal exopod with transverse suture. Hermaphroditic. [Adapted
from Sakai & de Saint Laurent, 1989.]
Remarks. — The genus Calaxiopsis was established on the basis of the type
species C. serrata Sakai & De Saint Laurent, 1989, and is different from the
genus Calastacus not only in the shapes of male Plps1-2, but also as follows.
In Calaxiopsis the gastric region bears submedian carinae, and pleura 2-5 are
triangular or subsquare, bearing a small anteroventral spine in some species
(not in all), and a sharp, distinct posteroventral spine in some species (not in
all). In Calastacus, in contrast, the gastric region bears no submedian carinae,
and pleura 3-6 are rounded ventrally and pleura 2-6 bear no spine ventrally.
Type species. — Calaxiopsis serrata Sakai & De Saint Laurent, 1989, by
original designation. The gender of the generic name, Calaxiopsis, is feminine.
Species included. — Calaxiopsis serrata Sakai & De Saint Laurent, 1989;
Calaxiopsis sp. Robles et al., 2009.
Calaxiopsis serrata Sakai & De Saint Laurent, 1989
(figs. 42, 43A-E)
Calaxiopsis serrata Sakai & De Saint Laurent, 1989: 61, fig. 16 [partim; mis-described; the
second D is for E, female MP 1173].
Material examined. — MNHN Th 1172, holotype, male (TL/CL, 21.0/9.0 mm), Guinea,
Guinea Trawling Survey, Sta. 11, 11.viii.1966; MNHN Th 1173, paratype, female, (TL/CL,
18.0/7.8 mm), Guinea, Trawling Survey, 240-250 m, 19.viii.1966; MNHN Th 1174, paratype,
juv. male (TL/CL, 10.0/3.5 mm), Guinea, Trawling Survey, Radial B, Sta. 7, 200 m, 25.xi.1966;
MNHN Th 1175, paratype, male (TL/CL, 19.0/8.0 mm), Guinea, Guinea Trawling Survey,
Radial D, Sta. 8, dredged, 100 m, 24.ii.1966; MNHN Th 1176, paratype, male (TL/CL,
23.0/8.5 mm), Guinea, Guinea Trawling Survey, Sta. 11, 25.viii.1966.
Diagnosis. — Rostrum elongate, acutely triangular, furrowed on dorsal surface, overreaching distal margin of penultimate segment of A1 peduncle, lateral margins with three teeth posteriorly extending shortly onto gastric region
as smooth lateral carinae with supraorbital spines. Carapace smooth. Anterolateral margin of carapace unarmed. Gastric region convex, bearing median
and submedian carinae; median carina running short from base of rostrum;
submedian carinae incurved, with two spines anteriorly. Cervical groove only
on dorsal part. Eyes subspherical and contiguous to carapace; cornea unpigmented. A1 peduncle reaching distal third of A2 penultimate segment. A2
segment 1 with strong ventral tooth; segment 2 elongated to form dorsodistal tooth reaching distal third of penultimate segment; scaphocerite (fig. 42D)
directed outward, and about as long as dorsodistal tooth of segment 2; segment 3 with short ventrodistal tooth; penultimate segment about three times as
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
225
Fig. 42. Calaxiopsis serrata Sakai & De Saint Laurent, 1989. A, C, male abdominal somites 1-6;
B, female abdominal somites 1-6; D, A2 peduncle, lateral view; E, male abdominal somites 1-6
and tail-fan. A, MNHN Th 1174, paratype, juv. male (TL/CL, 10.0/3.5 mm), Guinea; B, MNHN
Th 1173, paratype, female (TL/CL, 18.0/7.8 mm), Guinea; C, MNHN Th 1175, paratype, male
(TL/CL, 19.0/8.0 mm), Guinea; D, E, MNHN Th 1176, male (TL/CL, 23.0/8.5 mm), Guinea.
Scales 1 mm.
long as distal segment; antennal flagellum about twice as long as antennular
flagella. Mxp3 pediform; coxa and basis each with ventrodistal tooth; ischium
more than twice as long as wide, bearing two teeth on mesial margin, and ischial crest with 11 interspaced marginal denticles, terminated with triangular
tooth. Merus about as long as ischium, armed with two sharp teeth distally on
mesial margin. Carpus broadened distally, about three-fourths length of merus
and about as long as propodus. Dactylus about three-fourths length of propodus, and brimmed with dense bristles on ventral margin.
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CRM 013 – Katsushi Sakai
P1 subequal; in larger cheliped on left side, coxa and basis unarmed;
ischium with subterminal tooth on ventral margin; merus armed with two sharp
teeth on ventral margin and subterminal tooth on dorsal margin; carpus with
ventrolateral tooth, and subterminal tooth on dorsal margin; chela 2.5 times as
long as wide; fingers setose and longer than palm; palm slightly longer than
wide, lateral surface smooth, furnished with sharp subterminal tooth; dorsal
margin carinate with distal tooth; cutting edge of fixed finger armed with two
sharp teeth. Dactylus clearly longer than palm, cutting edge armed proximally
with obtuse triangular tooth, followed distally by large concavity. In smaller
cheliped on right side, ischium with two teeth on ventral margin; merus less
than twice as long as wide, armed with three teeth on ventral margin and
subterminal one on dorsal margin; carpus about half length of merus, armed
with subterminal tooth on dorsal margin; chela about 2.5 times as long as
wide; palm with subterminal tooth on carinate dorsal margin, lateral surface
bearing subterminal tooth; fingers longer than palm; fixed finger with two teeth
on cutting edge; dactylus 1.6 times as long as palm, and slightly sigmoid on
cutting edge. Both Ps3 and 5 coxae with genital pore. Pleurobranchs absent.
Abdominal somites smooth laterally. Somite 1 triangular. In young male
specimen (MNHN Th 1174, paratype, juv. male, TL/CL, 10.0/3.5 mm) pleuron 2 broad, bearing strong posteroventral spine (fig. 42A); pleuron 3 rounded
posteroventrally, bearing small anteroventral spine; pleuron 4 unarmed; pleuron 5 with small posteroventral spine. In adult male specimen (MNHN Th
1176) pleuron 2 almost straight ventrally, bearing strong posteroventral spine
(fig. 42E); pleuron 3 with reduced anteroventral and distinct posteroventral
spines; pleuron 5 with small anteroventral spine; and pleuron 6 rounded and
unarmed ventrally. In female specimen (MNHN Th 1173) pleuron 2 broad
and triangular ventrally; pleuron 3 with small anteroventral spine; pleura 3-4
with antero- and posteroventral spines; and abdominal somite 6 triangular and
unarmed ventrally (fig. 42B). In male specimen (MNHN Th 1172, holotype)
abdominal pleuron 2 with triangular posteroventral tooth; pleura 3-5 bearing
sharp anteroventral teeth and triangular posteroventral teeth. In male specimen
(MNHN Th 1175) pleuron 2 declining ventrally towards posteroventral angle
and unarmed; pleuron 3 rounded posteroventrally, bearing a single small anteroventral spine; pleura 3-4 with antero- and posteroventral teeth; pleuron 5
with a single anteroventral spine; and pleuron 6 triangular with a small ventral
spine (fig. 42C).
In male specimens (MNHN Th 1174, 1173) Plp1 uniramous and bisegmented; proximal segment short and distal segment slender and elongate distally to form spear-head shape (fig. 43A, B). In male specimens (MNHN Th
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
227
Fig. 43. Calaxiopsis serrata Sakai & De Saint Laurent, 1989 and Nipponcalaxiopsis manningi
(Komai, 2000). A, B, male Plp1; C, D, E, F, male Plp2. A, E, Calaxiopsis serrata MNHN Th
1175, paratype, male (TL/CL, 19.0/8.0 mm), Guinea; B, D, Calaxiopsis serrata, MNHN Th
1176, male (TL/CL, 23.0/8.5 mm), Guinea; C, Calaxiopsis serrata, MNHN Th 1172, holotype,
male (TL/CL, 21.0/9 mm), Guinea; F, Nipponcalaxiopsis manningi (Komai, 2000), CBM-ZC
4934, holotype (TL/CL, 19.2/7.1), off Takeoka, Uchibo coast of Boso peninsula. Scales 1 mm.
1172, 1176) Plp2 biramous and slender, endopod bearing distally bisegmented
appendix masculina, proximal segment provided mesioproximally with small,
proximally fused appendix interna, and short distal segment with marginal setae (fig. 43C, D); in the male specimen (MNHN Th 1175), an aberrant specimen, Plp2 is biramous, endopod with multiarticulate distal half, bearing mesiomedially appendices interna and masculina (fig. 43E). Plps3-5 biramous, endopods with appendix interna. Telson longer than wide, slightly convergent
posteriorly; lateral margins with proximal lobe with distinct spine followed
posteriorly by two teeth; dorsal surface with a pair of teeth; posterior margin
rounded and unarmed. Uropodal endopod with distinct spine at distolateral angle, bearing smooth median carina. Uropodal exopod unarmed on lateral margin, bearing movable spine at distolateral angle and distinct transverse suture
with 2-3 spinules.
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CRM 013 – Katsushi Sakai
Remarks. — The male holotype (MNHN 1172 from Guinea, Guinea
Trawling Survey, Sta. 11) is slightly different from the male paratype (MNHN
1176 from Guinea, Guinea Trawling Survey, Sta. 11) in the shape of the
distal segment of the bisegmented appendix masculina, because in the male
holotype (MNHN 1172) that distal segment is short with an obtuse tip
(fig. 43C), whereas in the male paratype (MNHN 1176) the distal segment
of the bisegmented appendix masculina is slender and pointed distally (fig.
43D). Another male paratype (MNHN 1175 from Guinea, Guinea Trawling
Survey, Radial D, Sta. 8, dredged, 100 m) is different from those two males
(MNHN 1172, 1176), because in the male specimen (MNHN 1175) the Plp2
endopod is slender and multiarticulate in its distal half, bearing mesiomedially
appendices interna and masculina (fig. 43E). It seems that the male specimen
(MNHN 1175) is aberrant in the form of the male Plp2 from the other type
specimens.
Type locality. — Guinea, Guinea Trawling Survey, Sta. 11, 240-250 m.
Distribution. — Only known from the type locality.
Calaxiopsis sp. Robles et al., 2009
Calaxiopsis sp., Robles et al., 2009: 316.
Genus Calocaris Bell, 1846
Calocaris Bell, 1846: 231; Dana, 1852a: 13; Dana, 1852b: 509; Ortmann, 1891: 50, pl. 1 fig. 5dI; Stebbing, 1893: 190; Alcock, 1901: 187; Borradaile, 1903: 539; Lagerberg, 1908: 50;
Selbie, 1914: 88 (key), 92; Pesta, 1918: 191; De Man, 1925d: 2 (key), 114; Rathbun, 1929:
25; Bouvier, 1940: 95; Barnard, 1950: 498 (key), 502; Balss, 1957: 1580; Zariquiey Alvarez,
1968: 225; De Saint Laurent, 1972: 353; Sakai, 1987a: 303 (list); Kensley, 1989: 962; Sakai
& De Saint Laurent, 1989: 55; Squires, 1990: 338; Poore, 1994: 98 (key); Felder, 2001: 440;
Ngoc-Ho, 2003: 459.
Callocaris — Zariquiey Alvarez, 1946: 95 (key), 104.
Calocaris s. str. — Borradaile, 1903: 539; De Man, 1925d: 2 (key), 115; Balss, 1957: 1580.
Diagnosis. — Rostrum triangular and grooved on dorsal surface, lateral
margins with 0-8 teeth, extending posteriorly onto anterior half of gastric
region as lateral carinae. Anterolateral margins armed or unarmed. Gastric
region convex; median carina smooth, submedian carinae absent. Cervical
groove present at full length. Postcervical carina present. Eyestalks and cornea
not differentiated, cornea unpigmented, anteriorly flattened and contiguous
along midline. A2 scaphocerite reduced. P1 subequal in length and similar
in shape; palm of larger cheliped with dorsodistal spine; fingers longer than
palm. P2 fingers longer than palm. Ps3-4 propodi without spinous rows. Ps35 dactyli simple. Ps3 and 5 coxae with genital pore. Pleurobranchs on Ps2-4.
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Abdominal pleura convex or rounded ventrally, and distinguished from terga
by obscure longitudinal carina. Plp1 uniramous and bisegmented; proximal
segment elongate and distal segment pear-shaped, bearing apically a small lobe
with a patch of hooklets. Plp2 biramous, exopod elongate and endopod bearing
distally elongate appendix masculina and small, slender appendix interna.
Plps3-5 biramous with slender endopod and exopod, endopod with appendix
interna. Telson longer than wide, bearing proximal lobe with spine followed
posteriorly by denticles on lateral margin; posterior margin with or without
median spine. Uropodal exopod bearing movable spine at posterolateral angle,
and transverse suture dorsally. Hermaphroditic.
Type species. — Calocaris macandreae Bell, 1846, by original designation
and monotypy. The gender of the generic name, Calocaris, is feminine.
Species included. — Calocaris caribbaea Kensley, 1996d; C. granulosa
Grebenjuk, 1975; C. isochela Zarenkov, 1989; C. macandreae Bell, 1846 (syn.
Calocaris barnardi Stebbing, 1914); C. sp. Kensley, 1996d.
K EY
TO THE
S PECIES
OF THE
G ENUS CALOCARIS
1 – Plp1 with subtriangular distal segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. isochela
– Plp1 with pear-shaped distal segment with distal patch of hooklets . . . . . . . . . . . . . . . . . 2
2 – Carapace covered with tubercles with setae; abdominal somite 6 with pair of longitudinal setal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. granulosa
– Carapace smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Lateral margins of rostrum and lateral gastric carinae continuous, each with 2-3 spines;
dorsal surface of telson bearing medially paired divergent carinae with 4-5 spines . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. macandreae
– Lateral margins of rostrum and lateral gastric carinae continuous, each with 4-8 spines;
dorsal surface of telson bearing medially paired divergent carinae with 1-2 spines . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. caribbaea
Calocaris caribbaea Kensley, 1996
(fig. 44)
Calocaris caribbaeus Kensley, 1996d: 164, figs. 5, 6; Robles et al., 2009: 316.
Material examined. — RMNH 41755 (= USNM 243421), 1 female (TL/CL, 31.2/11.2 mm),
western Atlantic, N. of Tobago, 11◦ 37.8′ N 60◦ 37.4′ W – 11◦ 38.8′ N 60◦ 37.5′ W, 659-1126 m,
10′ ottertrawl, 2.vii.19769, Pillsbury Sta. P-846; RMNH D 41746 (USNM Acc. No. 381702),
2 males (TL/CL, 36.0/12.2 mm; CL 12.1 mm, Abd4-telson missing), off coast of Surinam,
07◦ 56.5′ N 54◦ 39′ W – 08◦ 08′ N 54◦ 36′ W, 1042-1070 m, 41′ ottertrawl, 11.vii.1968.
Diagnosis. — Rostrum (fig. 44A, B) triangular and pointed distally, lateral
margins extending posteriorly onto gastric region as lateral carinae, and
230
CRM 013 – Katsushi Sakai
Fig. 44. Calocaris caribbaea Kensley, 1996. A, B, anterior part of carapace; C, A2 peduncle,
lateral view; D, Mxp3; E, telson. A-E, RMNH D 41746, male (TL/CL, 36.0/12.2 mm), off coast
of Surinam, 07◦ 56.5′ N 54◦ 39′ W – 08◦ 08′ N 54◦ 36′ W, 1042-1070 m. Scales 1 mm.
bearing series of 4-8 (5-6) spines. Carapace smooth. A1 segment 1 subequal in
length to segments 2 and 3 together, bearing small tooth on lateral margin. Eyes
flattened and contiguous along midline, cornea unpigmented. A2 segment 3
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231
with short distolateral tooth; scaphocerite (fig. 44C) reduced to tiny scale;
segment 4 three times as long as segment 5. Maxilla 2 scaphognathite with
posterior whip. Mxp3 (fig. 44D) ischium unarmed on mesial margin, ischial
crest bearing 9 denticles; merus with a single small distomesial tooth. P1
ischium with small distal spine on ventral margin; merus bearing 4 ventral
spines increasing in size distally, and a single subdistal spine on dorsal
margin; carpus unarmed; palm two-thirds length of fingers, bearing strong
distal spine on dorsal margin, and strong carina on lateral surface reaching
fixed finger; fixed finger bearing on cutting edge distinct proximal tubercle
followed distally by 5 small tubercles and finely denticulate straight distal end;
dactylus bearing on cutting edge distinct proximal tubercle followed distally
by well-interspaced low tubercle about in middle and finely denticulate straight
distal end. Plp1 uniramous and bisegmented, proximal segment fusiform;
distal segment broadened distally to form pear-shape with rounded distal lobe
bearing subapical patch of hooklets. Plp2 biramous, endopod bearing distally
appendix masculina with dense setae in proximal two-thirds, and appendix
interna. Plps3-5 endopods with appendix interna at proximal third. Abdominal
pleura rounded ventrally, bearing no spine. Telson (fig. 44E) longer than
wide, bearing 0-6 spinules on lateral margin; posterior margin rounded; dorsal
surface medially concave between divergent carinae, each with 0-4 spinules.
Uropodal endopod without spines on lateral margin. Uropodal exopod bearing
a single spine at posterolateral angle, and transverse suture. [Adapted from
Kensley, 1996d.]
Remarks. — Plps1-2 are the same as those of Calocaris macandreae Bell,
1846, which were figured by me (Sakai, 1991, fig. 5C, D).
Type locality. — Lesser Antilles, off Antigua, 17◦ 06′ N 62◦ 17′ W, 589 m.
Distribution. — Gulf of Mexico (Kensley, 1996d); Lesser Antilles —
off Antigua (Kensley, 1996d); off Surinam (Kensley, 1996d); off Trinidad
(Kensley, 1996d); 589-1272 m.
Calocaris granulosa Grebenjuk, 1975
Calocaris (Calocaris) granulosus Grebenjuk, 1975: 299, figs. 1, 2.
Calocaris granulosus — Sakai & De Saint Laurent, 1989: 59; Kensley, 1996d: 163 (list).
Paracalocaris granulosa — Sakai, 1991: 31.
Not: Calocaris (Calocaris) granulosus — Sakai, 1987a: 300, 303 (list) [= Paracalocaris
sagamiensis Sakai, 1991].
Diagnosis. — Carapace scattered with many clumps of setae. Rostrum
acutely triangular, lateral margins extending posteriorly onto gastric region as
lateral carinae, bearing three spines. A2 scaphocerite shorter than dorsal spine
232
CRM 013 – Katsushi Sakai
of A2 segment 2. Abdominal somite 6 bearing dorsomedially paired, divergent
rows of 6 clumps of setae. Plp1 uniramous and bisegmented, proximal segment
fusiform, distal segment pear-shaped. Plp2 biramous, endopod bearing distally
elongate appendix masculina and small appendix interna. Telson longer than
wide, bearing proximal lobe with spine, followed posteriorly by spines on
lateral margin, and divergent carinae with 6 spines medially on dorsal surface;
posterior margin slightly concave. [After Grebenjuk, 1975.]
Remarks. — Grebenjuk shows in his figure that the maxilla 2 scaphognathite
bears no posterior whip (Grebenjuk, 1975, fig. 3), however, that was presumably lost in the course of dissection, because all the other species in Calocaris
bear a posterior whip as a characteristic of the family Calocarididae.
Type locality. — Bay of Alaska.
Distribution. — Alaska (Grebenjuk, 1975); 756-1000 m.
Calocaris isochela Zarenkov, 1989
Calocaris isochela Zarenkov, 1989: 25, fig. 2; Kensley, 1996d: 163 (list).
Diagnosis. — Rostrum triangular, lateral margins extending posteriorly
onto gastric region as lateral carinae, bearing 6 spines. Gastric submedian
carinae absent, median carina unarmed, originating from base of rostrum and
running to cervical groove. Anterolateral margin of carapace armed with a
spine. A2 scaphocerite reduced. P1 chelate and unequal, merus with subdistal
tooth on dorsal margin; carpus unarmed; palm with subdistal tooth on dorsal
margin; fingers distinctly longer than palm. P2 fingers longer than palm.
Plp1 uniramous and bisegmented, distal segment subtriangular. Plp2 biramous,
endopod unsegmented, bearing mesiomedially distinct appendix interna and
elongate appendix masculina. Telson longer than wide, bearing unarmed
proximal lobe followed distally by denticles on lateral margin, and divergent
carinae with spinules medially on dorsal surface, posterior margin slightly
concave. Uropodal endopod with median carina. Uropodal exopod bearing a
spine at posterolateral angle, and a transverse suture. [After Zarenkov, 1989,
fig. 2.]
Remarks. — The type specimen is inaccessible, however, it is possible to
identify the species by examining in detail the figures given by Zarenkov
(1989, fig. 2). Zarenkov’s species, Calocaris isochela is then characterized
as follows: The A2 scaphocerite is reduced, Plp1 is uniramous and bisegmented, the distal segment is subtriangular; Plp2 is biramous, and the endopod bears mesiomedially a distinct appendix interna and an elongate appendix
masculina.
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233
Type locality. — New Zealand.
Distribution. — New Zealand — Plateau; 570 m.
Calocaris macandreae Bell, 1846
(figs. 45-47)
Calocaris Macandreae Bell, 1846: 233, figs.; White, 1850: 33; Smith, 1879: 55; Sars, 1883: 6;
Sars, 1884: 180, pls. 2, 6, 7; Carus, 1885: 490; Stebbing, 1893: 190; Adensamer, 1898: 621;
Lo Bianco, 1903: 187; Norman & Scott, 1906: 12; Hansen, 1908: 41; Lo Bianco, 1909: 603;
Stephensen, 1910a: 277; Selbie, 1914: 92, pl. 14 figs. 5-7; Bouvier, 1917: 119, pl. 11 figs. 5,
6; Balss, 1925: 209; De Man, 1925b: 140, fig. 7; Zariquiey Cenarro, 1935: 7.
Calocaris Macandrei — White, 1857: 99, pl. 8 fig. 4.
Calocaris macandreae — Kirk, 1879: 401; Sars, 1884: 166, pl. 2; Lovett, 1885: 16; Ortmann,
1891: 50, pl. 1 fig. 5; Meinert, 1893: 220; Anderson, 1896: 97; Adensamer, 1898: 621;
Appelhöf, 1906: 132; Lagerberg, 1908: 51, pl. 1 fig. 17; Wollebæk, 1909b: 251, figs. 1-6,
pls. 15-17; Björck, 1913: 1, figs. 1-6, pl. 1; Selbie, 1914: 92, pl. 14 figs. 5-7; Williamson,
1915: 447, figs. 211-214; Pesta, 1918: 191, fig. 59; Caroli, 1921a: 241; Caroli, 1921b: 264,
fig. 2; Runnstrøm, 1925: 14, pl. 1 figs. 1-2, 7, 10-15, pl. 2; Balss, 1926: 26; Grieg, 1927:
34; Williamson, 1927: 447, figs. 211-214; Rathbun, 1929, fig. 33 (after Bouvier, 1917, pl.
11 fig. 5); Miranda y Rivera, 1933: 21; Gustafson, 1934: 16; Poulsen, 1940: 208, 214, 236
(key), fig. 3; Gurney, 1942: 242, fig. 97; Zariquiey Alvarez, 1946: 104, fig. 132; Holthuis,
1950: 108, fig. 38; Zariquiey Alvarez, 1952: 19; Christiansen, 1955: 5; Soot-Ryen, 1955:
1; Holthuis, 1958: 8, fig. 5; Gordon, 1959: 191; Bourdillon-Casanova, 1960: 101, figs. 36,
37; Holthuis, 1962: 246; Buchanan, 1963: 729, figs. 1-15, tabl. 1; O’Riordan, 1963: 72;
ICZN, 1964: 340; Forest, 1965: 347; Picard, 1965: 104; Squires, 1965: 10, figs. 2Cm, 3Cm,
5Cm; Allen, 1967: 17, 57, 89 (fig.); Zariquiey Alvarez, 1968: 225, fig. 88b; Stevčić, 1969:
128; Christiansen, 1972: 40, fig. 46; Kattoulas & Koukouras, 1974: 344; Froglia, 1976: 76;
Holthuis & Heerebout, 1976: 62, fig. 82; Thiriot, 1976: 349; Beaubrun, 1979: 70, figs. 45,
46; Moncharmont, 1979: 70; Manning & Froglia, 1982: 323; Abel et al., 1983: 482, pl.
177; García Raso, 1983: 318; Macpherson, 1983: 44, fig. 25A; Riedl, 1983: 482, pl. 177, 2
figs; Nash et al., 1984: 425, figs. 1-3, pls. 1-3; Atkinson, 1986: 356, figs. 1K, 2A; Moore,
1986: 31; Thessalou-Legaki, 1986: 182; Atkinson & Taylor, 1988: 212, fig. 1h; Sakai & De
Saint Laurent, 1989: 56; Moyse & Smaldon, 1990: 117, figs. 10, 12; Števčić, 1990: 214;
Sakai, 1991: 31, fig. 5C-D; Dworschak, 1992: 215; Falciai & Minervini, 1992: 143, 1 fig.;
Koukouras et al., 1992: 223; Noël, 1992: 79; Kocatas & Katagan, 1993: 35; Pipitone &
Tumbiolo, 1993: 361; Anderson et al., 1994: 515, figs. 1-5; Cartes et al., 1994: 136, tables
2, 3; Diez et al., 1994: 47; Galil & Goren, 1994: 47; Froglia, 1995: 7; Hayward et al., 1995:
432, fig. 8.51 (partim); Falciai & Minervini, 1996: 143, 1 fig.; Barnich, 1996: 131, fig. 65;
García Raso, 1996: 738; Kensley, 1996d: 163 (list); Astall et al., 1997: 675, pl. 1a, tabls. 1-5;
Brattegard & Christiansen, 1997: 220; Hughes & Atkinson, 1997: 640; Johns et al., 1997:
127, figs. 1-4, tabl. 2; Pinn et al., 1998a; 243, fig. 1A; Pinn et al., 1998b: 211, figs. 1B, 2B;
Števčić, 1998: 652; Pinn et al., 1999a: 103, figs. 2B-D, 2H, 4A, 4B, 5F; Pinn et al., 1999b:
1461, figs. 1E, 2, 3, tabls. 1-6; Taylor et al., 1999: 163, figs. 1, 2, tabls. 1-4; d’Udekem
d’Acoz, 1999: 153; Christiansen, 2000: 230, fig. 1, tabl. 1; Taylor et al., 2000: 265, figs. 13, tabls. 1-6; Widdicombe et al., 2000: 371, figs. 1-8, tabls. 1-3; Costello et al., 2001: 289;
González-Gordillo et al., 2001: 279; Türkay, 2001: 289; Ngoc-Ho, N., 2003: 458, figs. 6, 7;
Atkinson & Taylor, 2004: 45, fig. 1, tabls. 1-2; Ingle & Christiansen, 2004: 84, fig. 64.
Calocaris Macandrae — Adensamer, 1898: 621.
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CRM 013 – Katsushi Sakai
Calocaris (Calocaris) Macandreae — Borradaile, 1903: 539; De Man, 1925d: 7 (list), 116
(key).
Calocaris barnardi Stebbing, 1914: 9, pl. 46; Kensley, 1981b: 30; Macpherson, 1983: 42,
fig. 25B-D; Sakai & De Saint Laurent, 1989: 58; Bianchi et al., 1993: 34; Kensley, 1996d:
163 (list). [Type locality. — Cape Castle, South Africa, 163 m.]
Calocaris (Calocaris) Barnardi — De Man, 1925d: 7 (list), 116 (key).
Calocaris Macandrae — Bull, 1933: 48, figs. 1-18.
Calocaris Mc Andreae — Bouvier, 1940: 96, pl. 4 figs. 1, 2 (cf. Bouvier, 1917).
Callocaris macandreae — Zariquiey Alvarez, 1946: 104, fig. 132.
Calocaris (Calocaris) barnardi — Barnard, 1950: 502 (key), 503, fig. 93i-k.
Not: Calocaris macandreae — Alcock, 1901: 189 (= Calocaris sp. Kensley, 1996d: 163);
Alcock & Anderson, 1894: 163; Whiteaves, 1901: 257 (= Calocarisopsis templemani
Squires, 1965).
Not: Calocaris macandreae — Rathbun, 1929: 25 (Gulf of St. Lawrence, Iceland).
Material examined. — RMNH D 38575, 4 samples, Turkey, Sea of Marmara, 52 m, beamtrawl, 23.xi.1990, leg. et don. B. Öztürk; ZMB 4878, 1 female (TL/CL, 41.0/14.4 mm), Korsfjord, Finnmark, Norway; ZMB 19882, 2 hermaphrodites (TL/CL, 50.5/17.3; 47.0/16.6 mm),
Biological Station Hardla at Bergen, 5.viii.1923, leg. Andt; SMF 20650, 4 hermaphrodites.
(TL/CL, 24.0/7.8-45.0/15.6 mm), North Sea, E. of Witch-Ground, 58◦ 00.00′ N 0◦ 55.90′ E –
57◦ 59.12′ N 0◦ 55.59′ E, 142.5 m, 22.vii.1989, R/V “Senckenberg”; SMF 29467, 1 hermaphrodite (TL/CL, 43.0/15.8 mm), North Sea, Fladengrund Val 103-46 Ku (58◦ 12.7′ N 0◦ 12′ W –
58◦ 12.7′ N 0◦ 12′ W), 133-133 m, 05.x.1990, R/V “Valdivia”; SMF 25683, 1 hermaphrodite
(TL/CL, 43.0/14.6 mm), North Sea, central North Sea, (55◦ 20.540′ N 0◦ 04.690′ W –
55◦ 20.570′ N 0◦ 030.370′ W, 101 m, 04.ii.1987, R/V “Valdivia”; SMF 20649, 1 hermaphrodite
(TL/CL, ca. 39.0/11.0 mm, carapace damaged), Fladengrund, North Sea, 58◦ 21.00′ N
0◦ 31.01′ W, 130.6 m, 22.vii.1989, R/V “Senckenberg”; SMF 20645, 1 ovig. hermaphrodite
(TL/CL, 48.0/17.4 mm), Fladengrund, North Sea, 58◦ 47.37′ N 0◦ 08.19′ W – 58◦ 46.56′ N
0◦ 06.77′ E, 137.3-146.6 m, 20.vii.1989, R/V “Senckenberg”. SMF 20647, 1 hermaphrodite
(TL/CL, 43.0/16.7 mm), S.W. Fladengrund-Rinne, North Sea, 58◦ 48.62′ N 0◦ 43.82′ W, 130.6 m,
21.vii.1989, R/V “Senckenberg”; SMF 29466, 1 hermaphrodite (TL/CL, 40.0/14.8 mm), Morocco between Tanger and Rabat M 60-12ST (34◦ 56′ N 07◦ 4′ W – 34◦ 56′ N 07◦ 7′ W, 784-829 m,
closing trawl, 23.i.1882, R/V “Meteor”; MNHN Th 1046 (TL, ca. 28.0, CR damaged), 26◦ 22′ S
13◦ 59′ E, R/V “Benguela II”, 380 m; ZMB 4878, 1 hermaphrodite (TL/CL, 42.0/14.0 mm),
Korsfjord entrance, Norway, biotope, 135-217 Faden (246.9-396.8 m), leg. Pommerania; ZMB
8170, 1 hermaphrodite (TL/CL, 32.0/12.0 mm), Häkonsund, Norway; ZMB 1015 (TL/CL,
30.0/11.0 mm), 1 hermaphrodite, Christian Fjord, Norway; ZMB 19882, 2 hermaphrodites
(TL/CL, 50.0/17.0; 51.0/18.0 mm), Biological Station Hardla near Bergen, 5.viii.1923, leg.
Biological Station Hardla; ZMB 18611, 1 hermaphrodite (TL/CL, 22.0/0.8 mm), Skagerrak,
Bohüslan, Sweden, 2.vii.1878, leg. A.W. Malm; SMF Ex-28339, 4 hermaphrodites (TL/CL,
47.0/16.9, 46.0/15.9, 45.0/14.8, 42.0/14.2 mm), Fiskebäckskil, Göteborg and Bohüslan, Sweden, 120 km N. of Göteborg, Kristineberg Marine Station, ix.1976, leg. U. Pettke; SMF 20651,
4 hermaphrodites (TL/CL, 42.0/15.7; 42.0/12.5; 41.2/15.2; 35.0/12.0 mm), Fladengrund, North
Sea, NWN 18 Ku (58◦ 21.00′ N 0◦ 30.88′ W, 58◦ 20.20′ N 0◦ 29.27′ W), 22.vii.1989, leg. R/V
“Senckenberg”, 4 m; SMF 28341, 1 hermaphrodite (TL/CL, 45.0/14.5 mm), Fladengrund, Vall
03-46 Ku (58◦ 12.7′ N 0◦ 12.0′ W), 133 m, 05.x.1990, leg. R/V “Valdivia”.
Material identified as Calocaris barnardi. — SAMC A 10697, 3 syntypes, (TL/CL,
28.03/11.2; ca. 32.0/11.1; 34.0/11.6 mm), alcohol specimens after having been dried, hermaphrodites, 29◦ S 16◦ 45′ E, 84 m, Fishery Survey; ZMB 26195, 1 ovig. female, hermaphrodite
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
235
(TL/CL, 50.0/15.6 mm), S. Atlantic, 26◦ 38′ S 13◦ 54′ E, 410-375 m, 23.v.1968, leg. E. Haeckel,
det. H.-E. Gruner; ZMB 26194, 1 hermaphrodite (TL/CL, 30.0/9.4 mm), S. Atlantic, 20◦ 16′ S
12◦ 16′ E, 30.iv.1968, leg. E. Haeckel, det. H.-E. Gruner.
Diagnosis. — Rostrum triangular with pointed apex, straight, horizontal or
slightly curved downward anteriorly (figs. 45A, 46A, 47A), lateral margins
extending posteriorly onto gastric region as lateral carinae, bearing 2-6 spines.
Carapace (fig. 45B, C) convex dorsally and carinate dorsomedially in gastric
and cardiac regions; gastric region convex, bearing smooth median carina.
Anterolateral margin of carapace with antennal spine. Posterolateral lobes
present. Eyes and eyestalks not differentiated and hardly mobile, cornea
flattened anteriorly, unpigmented, and contiguous to carapace. A1 peduncle
reaching distal two-thirds of A2 penultimate segment. A2 scaphocerite (figs.
46A, 47A, B) reduced, with pointed tip. P1 (figs. 46B, 47C) subequal; ischium
with two ventral spines; merus with 4-6, or 7 (ZMB 19882) interspaced
spines on ventral margin; carpus triangular; palm bearing triangular subdistal
tooth on dorsal margin and 1-2 teeth distally on ventrolateral carina; fingers
much compressed, and approximately of same length, 1.5-3 times as long
as palm; in larger cheliped cutting edge of dactylus bearing broad median
concavity, followed distally by triangular tooth and denticulate distal third;
cutting edge of fixed finger bearing proximally conical tooth, followed distally
by denticulation; in smaller cheliped dactylus smooth on cutting edge; and
fixed finger denticulate on cutting edge. P2 chelate. Ps3-5 simple. Ps3-4
propodus elongate, bearing no spinous rows. Plp1 uniramous and bisegmented,
distal segment pear-shaped with apical lobe with patch of hooklets (Sakai,
1991, fig. 5C); Plp2 biramous, and exopod and endopod elongate, endopod
distally bearing appendices masculina and interna (Sakai, 1991, fig. 5D).
Telson (figs. 46C, D, 47D) 1.5 times as long as wide, bearing proximal lobe
with spine followed distally by spinules on lateral margin, and also bearing
median concavity between divergent carinae with 3-7 spinules on dorsal
surface (table IV); posterior margin with median spine (SMF 25683, female).
Hermaphroditic, genital pore on Ps3 and 5 coxae.
Remarks. — In the specimens of Calocaris macandreae the rostral lateral
margins extend posteriorly onto the gastric region as lateral carinae, bearing
2-6 spines; in the larger cheliped the cutting edge of the dactylus bears a broad
median concavity, followed distally by a triangular tooth and a denticulate
distal third, the cutting edge of the fixed finger proximally bears a conical
tooth, followed distally by denticulation; in the smaller cheliped the cutting
edge of the dactylus is denticulate; and the dorsal surface of the telson is
concave medially between the divergent carinae, each with 1-6 sharp spinules.
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CRM 013 – Katsushi Sakai
Fig. 45. Calocaris macandreae Bell, 1846 (synonym: Calocaris barnardi Stebbing, 1914).
A, whole body with A1-2 peduncles, lateral view; B, C, anterior part of carapace and A1-2
peduncles, dorsal view. A, B, SAMC A 10697, syntype, from the South Atlantic Ocean (29◦ S
16◦ 45′ E, 84 m); C, ZMB 26195, 1 ovig. female, hermaphrodite (TL/CL, 50.0/15.6 mm), South
Atlantic, 26◦ 38′ S 13◦ 54′ E, 410-375 m. Scales 1 mm.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
237
Fig. 46. Calocaris macandreae Bell, 1846 (synonym: Calocaris barnardi Stebbing, 1914).
A, anterior part of carapace and A1-2 peduncles; B, smaller cheliped, lateral view; C, telson;
D, telson and uropod on right side. A, B, D, ZMB 26195, 1 ovig. female, hermaphrodite (TL/CL,
50.0/15.6 mm), South Atlantic, 26◦ 38′ S 13◦ 54′ E, 410-375 m; C, ZMB 26194, 1 ovig. female,
hermaphrodite (TL/CL, 50.0/15.6 mm), South Atlantic, 26◦ 38′ S 13◦ 54′ E, 410-375 m. Scales
1 mm.
In the present revision, 5 specimens of Calocaris barnardi including three
syntypes from the South Atlantic Ocean have been compared with many
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CRM 013 – Katsushi Sakai
Fig. 47. Calocaris macandreae Bell, 1846. A, anterior part of carapace; B, A2 peduncle, C, cheliped on right side; D, telson. A, B, SMF 29466, 1 hermaphroditic spm. (TL/CL, 40.0/14.8 mm),
Morocco, between Tanger and Rabat; C, ZMB 26195, 1 ovig. female, hermaphrodite (TL/CL,
50.0/15.6 mm), South Atlantic, 26◦ 38′ S 13◦ 54′ E, 410-375 m; D, SMF, 25683, 1 hermaphroditic
spm. (TL/CL, 43.0/14.6 mm), North Sea, central North Sea, (55◦ 20.540′ N 0◦ 04.690′ W –
55◦ 20.570′ N 0◦ 030.370′ W, 101 m, R/V “Valdivia”. Scales 1 mm.
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239
TABLE IV
Spines on rostral lateral margins, gastric lateral carinae, and divergent median carinae on dorsal
surface of telson, as defined for Calocaris macandreae Bell
Cat. No.
SMF Ex-28339
SMF Ex-28339
SMF Ex-28339
SMF Ex-28339
SMF 20651
SMF 20651
SMF 20651
SMF 20651
SMF 28341
ZMB 19882
ZMB 19882
ZMB 8170
ZMB 18611
ZMB 4878
ZMB 1015
TL/CL
(mm)
Localities
Spines on rostral
lateral margins
and gastric
lateral carinae
(left + right
sides)
Specimens det. as Calocaris macandreae
47.0/16.9 120 km N. Göteborg
46.0/15.9 120 km N. Göteborg
45.0/14.8 120 km N. Göteborg
42.0/14.2 120 km N. Göteborg
42.0/15.7 North Sea,
58◦ 21.00′ N 0◦ 30.88′ W –
58◦ 20.20′ N 0◦ 29.27′ W
42.0/12.5 North Sea,
58◦ 21.00′ N 0◦ 30.88′ W –
58◦ 20.20′ N 0◦ 29.27′ W
41.2/15.2 North Sea,
58◦ 21.00′ N 0◦ 30.88′ W –
58◦ 20.20′ N 0◦ 29.27′ W
35.0/12.0 North Sea,
58◦ 21.00′ N 0◦ 30.88′ W –
58◦ 20.20′ N 0◦ 29.27′ W
45.0/14.5 58◦ 12.7′ N 0◦ 12.0′ W
50.0/17.0 Bergen
51.0/18.0 Bergen
32.0/12.0 Häkonsund, Norway
22.0/0.8 Skagerrak, Bohüslan, Sweden
42.0/14.0 Korsfjord, Norway
30.0/11.0 Christiansfjord, Denmark
Spines on
median
carinae of
telson
(left + right
sides)
4+4
4+5
5+5
4+4
6+5
4+5
5+4
6+6
4+4
1+2
3+3
5+5
4+3
4+4
4+3
4+4
4+4
4+6
5+5
4+5
4+3
5+5
6+5
2+3
5+4
4+4
3+3
3+3
5+4
4+5
specimens of C. macandreae from the North and South Atlantic Oceans and
the Mediaterranea Sea.
In the three syntypes of Calocaris barnardi (SAMC A 10697, 3 syntypes
(TL/CL, 28.03/11.2; ca. 32.0/11.1; 34.0/11.6 mm)) from the South Atlantic
Ocean (29◦ S 16◦ 45′ E, 84 m), the rostral lateral margins extend backward
onto the gastric region as lateral carinae, bearing 2-3 spines; the A1 peduncle
reaches the distal two-thirds of the A2 penultimate segment; in the larger
cheliped the cutting edge of the dactylus bears a broad median concavity,
followed distally by a triangular tooth and a denticulate distal third, and the
cutting edge of the fixed finger proximally bears a conical tooth, followed
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CRM 013 – Katsushi Sakai
TABLE V
Spines on rostral lateral margins, gastric lateral carinae, and divergent median carinae on dorsal
surface of telson as defined for Calocaris macandreae Bell
Cat. No.
SAMC A 10697
SAMC A 10697
SAMC A 10697
ZMB 26194
ZMB 26195
TL/CL
(mm)
Localities
Spines on rostral
lateral margins
and gastric
lateral carinae
(left + right
sides)
Specimens det. as Calocaris barnardi Stebbing
28.03/11.2 29◦ S 16◦ 45′ E, 84 m
2+2
ca. 32.0/11.1
29◦ S 16◦ 45′ E, 84 m
3+3
2+2
34.0/11.6
29◦ S 16◦ 45′ E, 84 m
30.0/9.4
S. Atlantic,
2+2
20◦ 16′ S 12◦ 16′ E
50.0/15.6
S. Atlantic,
2+2
26◦ 38′ S 13◦ 54′ E,
410-375 m
Spines on
median
carinae of
telson
(left + right
sides)
5+5
4+4
4+6
7+6
5+4
distally by denticulation; in the smaller cheliped the dactylus is smooth on
the cutting edge, the fixed finger is denticulate on the cutting edge; and the
dorsal surface of the telson is concave medially between the divergent carinae,
each with 4-6 sharp spinules (table V).
In the specimen of C. barnardi (ZMB 26194, TL/CL, 30.0/9.4 mm) from
the South Atlantic (20◦ 16′ S 12◦ 16′ E, off Namibia), the rostral lateral margins
extend backward onto the gastric region as gastric lateral carinae with one
sharp spine followed by spinules and another sharp spine. The telson is
provided with a proximal lobe with a spine on the lateral margin, followed
posteriorly by three spinules only on the left, but no spinules on the right, and
the dorsal surface is concave medially between the divergent carinae, each with
6-7 interspaced, sharp spinules.
In the larger specimen of C. barnardi (ZMB 26195, 1 ovig., TL/CL,
50.0/15.6 mm) from the South Atlantic (26◦ 38′ S 13◦ 54′ E, 410-375 m, off
Namibia), the rostral lateral margins extend backward onto the gastric region
as lateral carinae with two spines. The telson is provided with a proximal
lobe with a spine on the lateral margin, followed posteriorly by three spinules
only on the left, but no spinules on the right (fig. 46D); the dorsal surface is
concave medially between the diviergent carinae, each with 4-5 interspaced
sharp spinules.
As shown above, C. barnardi is almost the same in morphological features
as C. macandreae, except for the difference in the number of spines both on the
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
241
rostral lateral margin and the gastric lateral carinae, and on the dorsal median
carinae of the telson, and there seems to be no significant difference between
the two species. Therefore, C. barnardi Stebbing, 1914 is safely synonymized
with C. macandreae Bell, 1846.
Type locality. — Loch Fyne and Mull of Galloway, Scotland.
Distribution. — North Atlantic Ocean — Canada — Gulf of St. Lawrence;
Iceland (Hansen, 1908); Norway — Trondheim Fjord, Christiania Fjord; Kattegat (Stephensen, 1910a; Poulsen, 1940); North Sea — (Poulsen, 1940;
Sakai & De Saint Laurent, 1989), Gullmar fjord, Sweden (Gustafson, 1934);
Norway — Mangerfjord, Bergen; Sweden — Kristineberg, and Skagerrak
(Dworschak, 1992); Scotland — Loch Fyne and Mull of Galloway (Bell,
1846); Ireland (Selbie, 1914; De Man, 1925d); England — Plymouth;
Sweden — Kristineberg, Bahusia [= Bohüslan]; Spain — Bay of Biscay
(Sakai & De Saint Laurent, 1989), Gulf of Cadiz (Bouvier, 1917).
Mediterranean — Sicilia and south of Monaco (Bouvier, 1917), Gulf of
Napoli (Caroli, 1921a), Marseille (Bouvier, 1940), Rosas, Arenys de Mar,
Badalona, Barcelona, and Melilla (Zariquiey Alvarez, 1968), Sardinia Island
(Manning & Froglia, 1982), south of Marseille and Canal of Mallorca (Sakai
& De Saint Laurent, 1989); Adriatic Sea (Adensamer, 1898; Dworschak, 1992;
Števčić, 1998); Aegean Sea (Koukouras et al., 1992); Turkey, Sea of Marmara.
South Atlantic Ocean — Mauritania (Sakai & De Saint Laurent, 1989);
Senegal — Dakar (Sakai & De Saint Laurent, 1989); Congo — PointeNoire (Sakai & De Saint Laurent, 1989); Namibia (Macpherson, 1983, det. as
C. barnardi); South Africa — Cape Castle (Stebbing, 1914, det. as
C. barnardi), near Port Nolloth (Barnard, 1950, det. as C. barnardi) 84-180 m.
Calocaris sp. Kensley, 1996
Calocaris macandreae — Alcock, 1901: 189.
Calocaris sp., Kensley, 1996d: 163 (list).
Remarks. — Calocaris macandreae Bell, 1846 was described by Alcock
(1901: 189) based on specimens from the Arabian Sea, 636 fathoms (1163 m);
Bay of Bengal, off Ceylon, 800-637 fathoms (1463-1165 m) as follows:
“the rostrum reaches the end of the basal joint of the antennal peduncle,
and is sharply serrated on the lateral border”. However, those characteristics
described by Alcock (1901) are different from those of C. macandreae Bell,
1846, in which the rostrum overreaches the A1 basal segment, and the lateral
margins of the rostrum are not serrated. Kensley (1996d: 163) distinguished
Calocaris macandreae from the Arabian Sea (Alcock, 1901: 189) from
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CRM 013 – Katsushi Sakai
C. macandreae from the Atlantic Ocean, and defined the former as Calocaris
sp. Indian ‘form’.
Distribution. — Arabian Sea; Bay of Bengal; 1163-1464 m.
Genus Calocarisopsis gen. nov.
Diagnosis. — Rostrum acutely triangular, lateral margins with 3-4 spines
extending posteriorly onto gastric region as lateral carinae with 4-5 spines, and
reaching cervical groove. Gastric region convex; median carina smooth, originating from anterior part of rostrum and running to cervical groove; submedian carinae absent. Cervical groove present at full length. Postcervical carina
present. Eyes and eyestalks not differentiated, cornea unpigmented, flattened
anteriorly, and contiguous along midline. A2 scaphocerite reduced. P1 subequal in length and similar in shape; palm of larger cheliped with dorsodistal
spine; fingers longer than palm. P2 fingers longer than palm. Ps3-4 propodi
without spinous rows. Ps3-5 dactyli simple. Ps3 and 5 coxae with genital pore.
Abdominal pleura convex or rounded ventrally, and distinguished from terga
by obscure longitudinal convexity. Plp1 uniramous and unsegmented, fusiform
in proximal half and spatulate in distal half. Plp2 biramous and slender, endopod bearing mesiomedially slender appendix interna, but no appendix masculina. Plp3 endopod with appendix interna. Telson longer than wide, bearing
proximal lobe with spine followed posteriorly by denticles on lateral margins,
and no median spine on posterior margin. Uropodal exopod with transverse
suture. Hermaphroditic.
Remarks. — The present new genus Calocarisopsis gen. nov. is different
from Calocaris, because in Calocarisopsis the carapace bears gastric lateral
carinae extending posteriorly to cervical groove, the median carina is developed anteriorly to the dorsal surface of the rostrum, Plp1 is unsegmented and
spatulate in its distal half, and Plp2 is biramous, the endopod is elongate, bearing mesiomedially an appendix interna, but no appendix masculina. Whereas
in Calocaris the carapace bears gastric lateral carinae in the anterior half of
the gastric region, the median carina is not developed anteriorly to the rostrum,
Plp1 is bisegmented, the distal segment is pear-shaped, and Plp2 is biramous,
the endopod bearing distally a distinct appendix masculina and a small appendix interna.
Type species. — Calocaris templemani Squires, 1965, by present designation and monotypy. The gender of the new name, Calocarisopsis, is feminine.
Species included. — Calocarisopsis templemani (Squires, 1965).
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
243
Etymology. — The new genus is named Calocarisopsis, by adding a Greek
suffix “-opsis”, meaning “appearance”, to the generic name Calocaris, from
which the new genus is different, though it is similar in appearance.
Calocarisopsis templemani (Squires, 1965)
Calocaris templemani Squires, 1965: 2, figs. 1, 2A-B, Ct, 3Ct, 4, 5Ct, 6, table 1; Sakai & De
Saint Laurent, 1989: 58; Kensley, 1996d: 163 (list); Ngoc-Ho, 2003: 465; Heard et al., 2007:
20, fig. 8.
Calocaris macandreae — Whiteaves, 1901: 257 [not C. macandreae Bell, 1846].
Diagnosis. — Rostrum acutely triangular, lateral margins with 3-4 spines
extending posteriorly onto gastric region as lateral carinae with 4-5 spines,
and reaching cervical groove. Gastric region convex; median carina smooth,
originating from anterior part of rostrum and running to cervical groove.
Cornea unpigmented and flattened obliquely laterally. A1 about half length of
A2, and about equal to total length of carapace and abdomen. A2 proximal
segment with 4 ventral spines. Mouthparts and chelae of pereiopods with
profuse, long setae. P1 chelate and unequal, in both chelipeds dactylus dorsally
carinate, bearing profuse long setae on each side, and fixed finger fringed
with profuse long setae ventrally, bearing somewhat oblique lower carina also
with profuse long setae running from distal part of fixed finger to proximal
part of palm. Abdomen rather longer than carapace and somewhat flattened
laterally; abdominal somites 2-6 fringed with long fine setae ventrally, dorsally
bearing two clumps of fine setae. Plp1 uniramous and unsegmented, fusiform
in proximal half and spatulate in distal half; Plp2 biramous and slender,
endopod bearing mesiomedially a slender appendix interna, but no appendix
masculina; Plp3 endopod with appendix interna. Telson longer than wide and
slightly convergent posteriorly, bearing proximal lobe with spine followed
posteriorly by two spinules on lateral margin, and divergent carinae with 2-3
spines medially on dorsal surface; posterior margin unarmed. Uropodal exopod
with transverse suture. Hermaphroditic. [Adapted from Squires, 1965.]
Remarks. — Squires (1965: 9) described in his holotype specimen, determined as Calocaris templemani, that the Plp1 is undivided and somewhat spatulate with a light fringe of setae and a curved foliaceous apex; Plps2-5 are
biramous, and the endopod has a long narrow appendix interna, and both exopod and endopod are fringed by long plumous setae (Squires, 1965, fig. 6
PL2). However, those features of Plps1-2 are different from those of the other
species of Calocaris, in which Plp1 is uniramous and bisegmented, and the
distal segment is pear-shaped apically, with a small lobe with a patch of hooklets (Sakai, 1991, fig. 5C); Plp2 is biramous with the exopod elongate, and the
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CRM 013 – Katsushi Sakai
endopod distally bears an elongate appendix masculina and a small, slender appendix interna. Therefore, Calocaris templemani Squires, 1965 is reclassified
under the new genus Calocarisopsis gen. nov. as Calocarisopsis templemani
(Squires, 1965).
Type locality. — Hermitage Bay, Newfoundland, 260 m.
Distribution. — Canada — Newfoundland, Hermitage Bay (Squires, 1965);
260 m, soft mud; northwest Atlantic, from Newfoundland, Gulf of Maine to
North Carolina, 200-785 m (Kensley, 1996d).
Genus Eucalastacus Sakai, 1992
Eucalastacus Sakai, 1992a: 170; Poore, 1994: 98 (key).
Diagnosis. — Rostrum styliform, lateral margins with 4 spines including
supraorbital spine, extending posteriorly onto gastric region as lateral carinae;
median and submedian carinae present; postcervical carina absent. Anterolateral margin of carapace unarmed. Eyes quadrate and contiguous mesially;
cornea unpigmented. A2 scaphocerite slenderly protruded. P1 symmetrical,
fingers gaping proximally. Plp1 uniramous and bisegmented; proximal segment elongate, and distal segment subtriangular with two apical setae and
patch of hooklets medially on convex mesial margin; Plp2 biramous and slender; endopod distally bearing slender appendix masculina with setae, provided
proximally with small appendix interna with patch of hooklets; Plps3-5 biramous with slender endopod and exopod; endopods bearing mesiomedially
appendix interna. Genital pores on Ps3 and 5 coxae. Telson longer than wide,
and convergent posteriorly, bearing proximal lobe with spine followed by two
spinules on right lateral margin but unarmed on left lateral margin, and divergent carinae with two spines medially on dorsal surface; posterior margin
rounded without median spine. Uropodal endopod with median carina. Uropodal exopod without transverse suture. Hermaphroditic. [Adapted from Sakai,
1992a.]
Remarks. — The present genus Eucalastacus is closely similar to Calocaris
in Plps1-2, but differs in that the uropodal exopod bears no transverse suture.
Type species. — Eucalastacus torbeni Sakai, 1992a, by original designation
and monotypy. The gender of the generic name, Eucalastacus, is masculine.
Species included. — Eucalastacus torbeni Sakai, 1992a.
Eucalastacus torbeni Sakai, 1992
Eucalastacus torbeni Sakai, 1992a: 170, figs. 11-13.
Type locality. — Milford Sound, New Zealand, 44◦ 37′ S 167◦ 52′ E, 290 m.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
245
Distribution. — Tasman Sea (Sakai, 1992a); New Zealand — Milford
Sound (Sakai, 1992a); 290-341 m.
Genus Lophaxiopsis gen. nov.
Diagnosis. — Rostrum triangular, lateral margins extending short posteriorly onto gastric region as lateral carinae each with 1-2 spines including
supraocular spine. Carapace covered with short, stiff, yellowish, forwardly directed setae, and carinate dorsomedially in gastric and cardiac regions; gastric
region with series of antrorse [= directed forward and upward] spines arranged
in form of a horseshoe with the convexity forward. Postcervical carina with irregular tubercles present. Cornea small and irregularly pigmented. A2 scaphocerite short. Abdominal somites with dorsal carina; pleura 3-5 somewhat angular ventrally. Male Plps1-2 unknown. Telson longer than wide. Uropodal
exopod with non-dentate transverse suture. Hermaphroditic. [Adapted from
Alcock & Anderson, 1899; Alcock, 1900, pl. 42 fig. 3.]
Remarks. — It has turned out that Calaxiopsis felix (Alcock & Anderson,
1899) (cf. Sakai & De Saint Laurent, 1989) is not to be included in Calaxiopsis,
because it is different from the type species of the genus, C. serrata Sakai &
De Saint Laurent, 1989. In Calaxiopsis felix the gastric region is covered with
short stiff, yellowish, forwardly-directed setae, bearing a series of antrorse
[this term was used by Alcock (1901)] spines arranged in the form of an
elongate horseshoe, a median carina originating from the base of the rostrum
and running to the posterior margin of the carapace, and short lateral carinae
with a couple of spines, submedian rows of spines, and the abdominal somites
bear a dorsal carina. In Calaxiopsis serrata, however, the gastric region is
smooth, bearing a median carina running short from the base of the rostrum,
and smooth submedian carinae, and the abdominal somites lack a dorsal carina.
Calaxiopsis felix is rather similar to the type species of Lophaxius, L. rathbunae, but it is also different from L. rathbunae, in which the gastric region
is tuberculate, and the abdominal somites lack a dorsal carina. Unfortunately,
inaccessibility to the type specimen of Calaxiopsis felix prevents further examination of its Plps1-2, of which no description was made, but those differences
mentioned above are sufficient to differentiate C. felix from L. rathbunae at the
generic level, and a new genus Lophaxiopsis gen. nov. is thus propopsed for
Calaxiopsis felix.
Type species. — Calastacus felix Alcock & Anderson, 1899, by present
designation and monotypy. The gender of the generic name, Lophaxiopsis, is
feminine.
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CRM 013 – Katsushi Sakai
Species included. — Lophaxiopsis felix (Alcock & Anderson, 1899).
Etymology. — The new generic name Lophaxiopsis is composed of a
generic name, Lophaxius, and a Latin suffix, -opsis, meaning “similar”.
Lophaxiopsis felix (Alcock & Anderson, 1899)
Calastacus felix Alcock & Anderson, 1899: 287; Alcock, 1900, pl. 42 fig. 3; Balss, 1925: 209.
Calocaris (Calastacus) felix — Alcock, 1901: 192; Borradaile, 1903: 539; De Man, 1925d: 8
(list), 117 (key).
Calaxiopsis felix — Sakai & De Saint Laurent, 1989: 64.
Diagnosis. — Rostrum triangular, lateral margins with two subdistal spines,
extending short posteriorly onto gastric region as outstanding lateral carinae
with two spines. Carapace covered with scanty growth of short, stiff, yellowish, forwardly-directed setae, springing either singly or in groups of two or
three setae from the bottom of small pits. Gastric region with series of antrorse
spines arranged in the form of an elongate horseshoe, median carina originating from base of rostrum and running to posterior margin of carapace, and
submedian carinae absent. Cornea small and irregularly pigmented. Abdominal somites carinate dorsally, most prominent on anterior three somites, gradually widening and becoming less and less marked on posterior three somites.
Abdominal pleura 3-5 rounded ventrally. Plps1-2 unknown. Telson longer than
wide. [Adapted from Alcock & Anderson, 1899: 287; Alcock, 1900, pl. 42 fig.
3.]
Type locality. — Off Cape Comorin, Arabian Sea, 430 m.
Distribution. — Arabian Sea — off Cape Comorin (Alcock & Anderson,
1899); 430-785 m.
Genus Lophaxius Kensley, 1989
Lophaxius Kensley, 1989: 962; Poore, 1994: 98 (key).
Diagnosis. — Rostrum triangular, lateral margins extending short posteriorly onto gastric region, bearing 1-2 spines including supraocular spine; submedian carinae absent. Carapace carinate dorsomedially in gastric region, and
postcervical carina present with irregular tubercles. Eyes rounded and not
mesially contiguous; cornea unpigmented. A2 scaphocerite short. Mxps1-3
with exopod and epipod; large podobranch on Mxps2 and 3; 2 arthrobranchs on
Mxp3. P1 symmetrical, fingers of chela gaping proximally. P2 subchelate. Ps35 dactyli simple. Abdominal somites with pleura rounded ventrally. Plp1 uniramous and bisegmented, distal segment spatulate, distally with patch of hooklets; Plp2 biramous, endopod bearing distally elongate appendix masculina and
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
247
slender appendix interna; Plps3-5 biramous, endopods with appendix interna.
Telson longer than wide, dorsally bearing non-articulating spines. Uropodal
exopod with non-dentate transverse suture. Hermaphroditic. [Adapted from
Kensley, 1989.]
Remarks. — Kensley (1989: 962-963) established a new genus, Lophaxius
Kensley, 1989 for Calastacus rathbunae Kensley, 1989 [= Calocaris investigatoris (Rathbun, 1904) from Alaska and Oregon, not Anderson, 1896 from
Arabian Sea, 1733 m], and characterized his new genus Lophaxius as “the
eyes are rounded and not mesially contiguous” (Kensley, 1989: 963), which
distinguishes Lophaxius from Calocaris, in which the eyes are flattened, and
mesially contiguous, though in those two genera Lophaxius and Calocaris the
male Plps1-2 are almost the same in shape as follows: the male Plp1 is uniramous and bisegmented; the proximal segment is elongate, and the distal segment is pear-shaped, apically bearing a small lobe with a patch of hooklets.
Plp2 is biramous, the exopod is elongate, and the endopod distally bears an
elongate appendix masculina and a small, slender appendix interna.
Type species. — Calastacus rathbunae Kensley, 1989, by original designation. The gender of the generic name, Lophaxius, is masculine.
Species included. — Lophaxius investigatoris (Anderson, 1896); L. rathbunae Kensley, 1989.
K EY
TO THE
S PECIES
OF THE
G ENUS LOPHAXIUS
1 – Carapace covered with small granules; lateral margins of rostrum extending short
posteriorly as gastric lateral carinae, bearing acute tooth posteriorly followed by
another considerably larger tooth; eyestalks short and conical, and cornea unpigmented;
distributed in the Arabian Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. investigatoris
– Carapace tuberculate; distributed from Alaska to California . . . . . . . . . . . . . L. rathbunae
Lophaxius investigatoris (Anderson, 1896)
Calastacus investigatoris Anderson, 1896: 97; Balss, 1925: 209; Rathbun, 1904: 151.
Axius (Eiconaxius) Investigatoris — Alcock & Anderson, 1896, pl. 25 fig. 1.
Calocaris (Calastacus) investigatoris — Alcock, 1901: 191; Borradaile, 1903: 539.
Calastacus investigatoris — Schmitt, 1921: 112 (partim), fig. 75a-b.
Calocaris (Calastacus) Investigatoris — De Man, 1925d: 8 (list), 117 (key).
Calocaris investigatoris — Hart, 1982: 48, fig. 10; Sakai & De Saint Laurent, 1989: 58.
Lophaxius investigatoris — Kensley, 1989: 963.
Not: Calastacus investigatoris — Rathbun, 1904: 151; Schmitt, 1921: 112 (not fig. 75a-b),
[= Lophaxius rathbunae Kensley, 1989].
Diagnosis. — Carapace covered with granules. Rostrum triangular and dorsally flattened, and lateral margins extending short posteriorly onto gastric region as lateral carinae with acute tooth followed posteriorly by another considerably larger tooth. Eyestalks short and conical, and cornea unpigmented.
248
CRM 013 – Katsushi Sakai
Dorsal margin of P1 merus armed with one large distal and 3 or 4 small spines.
[Adapted from Alcock & Anderson, 1896: 97-98.]
Type locality. — Arabian Sea, off the coast of Sind, 1733 m.
Distribution. — Arabian Sea — off the coast of Sind (Anderson, 1896);
1733 m.
Lophaxius rathbunae Kensley, 1989
Calastacus investigatoris — Rathbun, 1904: 151 (partim); Schmitt, 1921: 112 (partim, not
fig. 75a-b) [not Anderson, 1896].
Lophaxius rathbunae Kensley, 1989: 963.
Diagnosis. — Gastric region tuberculate, bearing smooth median carina and
short lateral carinae with 1-2 spines; submedian carina absent; postcervical
carina present. Plp1 uniramous and bisegmented, distal segment pear-shaped,
apically bearing a small patch of hooklets. Plp2 biramous, endopod bearing
distally setose appendix masculina and appendix interna. [Adapted from
Kensley, 1989.]
Remarks. — Calastacus investigatoris Rathbun, 1904, distributed from
Alaska to California, and Calastacus investigatoris Anderson, 1896 distributed
in the Arabian Sea, were reclassified by Kensley (1989: 963) under the
new genus Lophaxius, as Lophaxius investigatoris and Lophaxius rathbunae,
because he considered that L. investigaroris (cf. Rathbun, 1904), is not
conspecific with L. investigatoris (Anderson, 1896).
Type locality. — South of Sannak Islands, Alaska, 483 fathoms (883 m).
Distribution. — Alaska — south of Sannak Islands, 883 m (Rathbun, 1904);
Oregon — off Cascade Head, 631 m (Rathbun, 1904); California — off San
Diego, 763 m (Rathbun, 1904).
Genus Nipponcalaxiopsis gen. nov.
Diagnosis. — Rostrum narrowly elongate, lateral margins armed with
two spines, extending posteriorly onto gastric region as lateral carinae with
supraorbital spine. Anterolateral margins of carapace unarmed. Gastric region
convex with median and submedian carinae. Cervical groove present only on
dorsal surface; no postcervical carina. Eyes rounded, cornea unpigmented and
contiguous to carapace. P1 subequal. Ps3 and 5 coxae with genital pores.
Abdominal pleura 2-5 truncate and unarmed ventrally. Pleurobranchs absent.
Plp1 uniramous and bisegmented; proximal segment stem-shaped, and distal
segment bilobed distally, bearing proximal lobe with patch of hooklets. Plp2
biramous, endopod bearing distally slender, curved, unsegmented appendix
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
249
masculina with setae on mesial margin and apical tuft of setae, and appendix
interna with patch of hooklets; exopod consisting of proximal segment and
multiarticulate flagellum. Plps3-5 endopods with appendix interna. Telson
longer than wide, bearing 6-8 spines on lateral margin, and no median spine
on rounded posterior margin. Uropodal endopod with distolateral spine and
unarmed median carina dorsally. Uropodal exopod bearing fixed and movable
spines at distolateral angle, and median carina with a few tufts of setae and
distal spine, and transverse suture on dorsal surface.
Remarks. — It is found that Calaxiopsis manningi Komai, 2000a is not to
be included in Calaxiopsis Sakai & De Saint Laurent, 1989, to which, though,
it seems closely similar, because C. manningi is fundamentally different from
the type species of Calaxiopsis, C. serrata Sakai & De Saint Laurent, 1989.
In Calaxiopsis manningi the male Plp2 appendix masculina is unsegmented,
and pleura 2-5 are unarmed ventrally, whereas in C. serrata the male Plp2
appendix masculina is bisegmented, and pleura 2-5 are armed with 1-2
spines ventrally. Therefore, a new genus, Nipponcalaxiopsis is erected for
Calaxiopsis manningi Komai, 2000a.
Type species. — Calaxiopsis manningi Komai, 2000 by present designation
and monotypy. The gender of the new name, Nipponcalaxiopsis, is feminine.
Species included. — Nipponcalaxiopsis manningi (Komai, 2000).
Etymology. — The new generic name Nipponcalaxiopsis is a combination
of “Nippon” meaning “Japan”, where the type specimen was collected, and the
generic name Calaxiopsis.
Nipponcalaxiopsis manningi (Komai, 2000)
(fig. 43F)
Calaxiopsis manningi Komai, 2000a: 218, figs. 1-5.
Material examined. — CBM-ZC 4934, holotype (TL/CL, 19.2/7.1 mm), off Takeoka,
Uchibo coast of Boso Peninsula, 80-100 m, burrowing on clay rock, leg. J. Takayama, det.
T. Komai.
Remarks. — Komai (2000: 226) requested the late M. De Saint Laurent to
examine the type specimens of Calaxiopsis serrata Sakai & De Saint Laurent,
1989 (cf. Komai, 2000a: 226), and defined his specimen as a new species,
Calaxiopsis manningi.
Komai mentioned by comparing his new species Calaxiopsis manningi
with C. serrata that “In addition to the development of the podobranchs,
Calaxiopsis manningi is readily distinguished from C. serrata in the unarmed
gastric submedian carinae, unarmed pleura of the second to fourth abdominal
250
CRM 013 – Katsushi Sakai
somites, and much shorter antennal acicles”. However, he failed to pay
attention to the distinct difference in the form of the male Plp2 between
the two species C. manningi and C. serrata. Namely, those two species are
fundamentally different from each other, because in C. serrata the male
Plp2 endopod bears a bisegmented appendix masculina (fig. 43C), whereas
in C. manningi it bears an unsegmented appendix masculina (Komai, 2000a,
fig. 3L) (fig. 43F). The characteristics of the abdominal pleura described by
Komai and in the Plp2 appendix masculina pointed out herein are safely
considered to be sufficient to distinguish the specimens on the generic level,
which suggests that C. manningi does not belong to Calaxiopsis, but to the
new genus Nipponcalaxiopsis established for C. manningi.
Type locality. — Off Takeoka, Uchibo coast of Boso Peninsula, Tokyo Bay.
Distribution. — Only known from the type locality.
Genus Paracalocaris Sakai, 1991
Paracalocaris Sakai, 1991: 30.
Diagnosis. — Carapace studded with tubercles. Rostrum elongate and
styliform, lateral margins with three spines, extending posteriorly onto gastric
region as lateral carinae, each with a distinct spine. Gastric region convex,
bearing three longitudinal carinae; median carina smooth; submedian carinae
absent. Anterolateral margin unarmed. Postcervical carina smooth. Eyestalks
not contiguous mesially; cornea unpigmented and upraised. A2 scaphocerite
present as minute spine. Mxps1-3 with exopod and epipod; Mxp2 with
small podobranch; Mxp3 with large podobranch. Pereiopods lacking exopod;
epipod present on Ps1-4; large podobranch on Ps1-3; pleurobranchs absent.
P1 chelate and subequal; palm armed with dorsal spines. P2 chelate. Ps3-4
propodi with rows of spinules on ventrolateral surface. Plp1 uniramous and
bisegmented, distal segment leaf-like and trilobed distally, distomesial lobe
with apical cluster of hooklets. Plp2 biramous and elongate, endopod with
slender setose appendix masculina provided proximally with appendix interna;
exopod consisting of proximal segment and multiarticulate flagellum. Plps35 endopods with appendix interna. Telson longer than wide, bearing median
concavity between divergent carinae with 8-9 spines. Uropodal exopod bearing
fixed and movable spines at posterolateral angle, and transverse suture on
dorsal surface. Hermaphroditic. [Adapted from Sakai, 1991.]
Type species. — Paracalocaris sagamiensis Sakai, 1991, by original designation and monotypy. The gender of the generic name, Paracalocaris, is
feminine.
Species included. — Paracalocaris sagamiensis Sakai, 1991.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
251
Paracalocaris sagamiensis Sakai, 1991
Calocaris (Calocaris) granulosus — Sakai, 1987a: 300, 303 (list) [not Grebenjuk, 1975].
Paracalocaris sagamiensis Sakai, 1991: 30, figs. 1-5, table 1; Tsang et al., 2008: 218, 219; Chu
et al., 2008.
Remarks. — The holotype was described as a female, however, the presence
of the genital pore on Ps3 and 5 coxae shows that it is a hermaphroditic
specimen (Sakai, 1999c: 30, fig. 5A, B).
Type locality. — Off Ohiso, Sagami Bay, Japan, 250-280 m (Sakai, 1987a:
300).
Distribution. — Japan — Sagami Bay (Sakai, 1987a, 1991); 250-280 m.
FAMILY CORALAXIIDAE SAKAI & DE SAINT LAURENT, 1989
Coralaxiinae Sakai & De Saint Laurent, 1989: 10.
Diagnosis. — Carapace with supraocular spine. Rostrum short, acutely triangular, and deflexed, bearig 0-1 spine on lateral margin. Gastric region bearing median carina with or without spines extending short or to posterior third
of gastric region, and anteriorly teeth arranged in circle or not. No linea thalassinica. Cervical groove distinct. Postcervical carina absent. Posterior margin
of carapace with lateral lobes. Eyestalks stout; cornea pigmented. A2 scaphocerite present or absent. Maxilla 2 scaphognathite with posterior whip. Mxp3
exopod present; distal segment with flagellum deflexed from proximal segment
with acute angle. P1 chelate and unequal. P2 chela ovate. Ps3-4 propodi slender, bearing spines on ventral margin. Ps3-5 dactyli biunguiculate. Epipods
vestigial on Ps1-2, and absent on Ps3-5. Abdominal pleura 2-4 convex or truncate ventrally. Male Plp1 uniramous and bisegmented; proximal segment long,
and distal one ovate and bilobed distally; male Plp2 biramous, endopod bearing appendices masculina and interna mesiomedially; male Plps3-5 biramous,
endopods with appendix interna. Telson subtriangular, bearing proximal lobe
with or without spine, and stiff setae posterolaterally, and median spine on
posterior margin. Uropodal exopod with serrated transverse suture.
Remarks. — The family Coralaxiidae Sakai & De Saint Laurent, 1989 was
established on the basis of the genus Coralaxius Kensley & Gore, 1981, which
had been included in the family Axiidae Huxley, 1879 just because those
two families have some characteristics in common as follows. The carapace
bears no linea thalassinica, the maxilla 2 scaphognathite bears a posterior
whip, and they are gonochoristic. However, the family Coralaxiidae is clearly
different from the family Axiidae as follows. In the Coralaxiidae the Mxp3
exopod bears a distal segment deflected from the un- or bisegmented proximal
segment as in pagurid species; the P2 chela is ovate in shape; Ps3-5 dactyli
are biunguiculate; the male Plp1 is uniramous and bisegmented, the proximal
segment is long, and the distal one is ovate; the male Plp2 is biramous, and
the endopod bears a stout appendix masculina and a small appendix interna;
the telson bears stiff setae on the posterolateral margin. However, in the
Axiidae the Mxp3 exopod is straight, consisting of a proximal segment and
254
CRM 013 – Katsushi Sakai
a multiarticulate flagellum; the P2 chela is slender in shape, and Ps3-5 dactyli
are simple; the male Plp1 is absent or present, and when present, uniramous
and unsegmented or bisegmented; the male Plp2 is biramous, and the endopod
bears both appendices masculina and interna, or only an appendix interna, or
rarely neither of them; the telson bears no stiff setae posterolaterally.
Type genus. — Coralaxius Kensley & Gore, 1981, by original designation.
Genera included. — Coralaxius Kensley & Gore, 1981; Meinertaxius gen.
nov.
K EY
TO THE
G ENERA
OF THE
FAMILY C ORALAXIIDAE
1 – Gastric region with teeth arranged in circle anteriorly; male Plp2 endopod with broad
bilobed appendix masculina with numerous distal setae and small appendix interna . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coralaxius
– Gastric region without teeth arranged in circle anteriorly; male Plp2 endopod with broad
and ovate appendix masculina and small appendix interna . . . . . Meinertaxius gen. nov.
Genus Coralaxius Kensley & Gore, 1981
Coralaxius Kensley & Gore, 1981: 1278; Sakai & De Saint Laurent, 1989: 2 (list), 10; Kensley,
1994: 813; Poore, 1994: 97 (key).
Diagnosis. — Rostrum short, triangular, and deflexed on dorsal surface.
Gastric region bearing teeth arranged in circle anteriorly behind rostrum, and
median carina developed short, with or without spines, bearing medially pyramidal hepatic convexity. Cervical groove distinct. Eyestalks stout; cornea pigmented. A2 scaphocerite small. Mxp3 exopod consisting of proximal segment and deflected distal segment with flagellum. P1 chelate and unequal.
P2 chelate; chela ovate in shape. Abdominal pleuron 2 rounded ventrally and
broadened, and protruded antero- and posteroventrally. Male Plp1 uniramous
and bisegmented, proximal segment elongate, and distal one ovate and slightly
bilobed distally, mesiodistally bearing hooklets; male Plp2 biramous, endopod
bearing mesiomedially appendices masculina and interna; appendix masculina
longitudinally bilobed in distal half, and provided proximally with appendix
interna with distal patch of hooklets; male Plps3-5 biramous, endopods with
appendix interna. Female Plp1 consisting of proximal segment and multiarticulate flagellum. Female Plps3-5 endopods with appendix interna. Telson
subtriangular and converging posteriorly, bearing stiff setae posterolaterally
and two pairs of teeth medially on dorsal surface; posterior margin fringed
with elongate setae, bearing median spine. Uropodal exopod bearing strong,
movable spine at posterolateral angle, and spinous transverse suture on dorsal
surface. [Adapted from Sakai & De Saint Laurent, 1989.]
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
255
Remarks. — Kensley (1994) examined the type specimen of Axius nodulosus Meinert, 1877 from the North Sea, and synonymized the species Coralaxius abelei Kensley & Gore, 1981 from the Caribbean Sea, with Meinert’s
species. However, the present re-examination of Meinert’s type specimen has
revealed that Meinert’s species is clearly different from Kensley & Gore’s
species, as mentioned in the remarks of Meinertaxius gen. nov.
Type species. — Coralaxius abelei Kensley & Gore, 1981 by original
designation and monotypy. The gender of the generic name, Coralaxius, is
masculine.
Species included. — Coralaxius abelei Kensley & Gore, 1981; C. galapagensis Kensley, 1994; C. indopacificus Kensley, 1994.
K EY
TO THE
S PECIES
OF THE
G ENUS CORALAXIUS
1 – Rostrum elongate and triangular, reaching distal margin of eyestalks; gastric region with
median carina with 2-3 spines, and lateral carinae with 2-3 teeth; pleurobranchs present
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. indopacificus
– Rostrum short and triangular, reaching proximal third to half of eyestalks; gastric region
with short median carina, and teeth arranged in circle; pleurobranchs absent . . . . . . . . . 2
2 – P2 palm and dactylus unarmed on dorsal margin; rostrum with one lateral tooth . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. galapagensis
– P2 palm and dactylus armed with denticles on dorsal margin; rostrum unarmed . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. abelei
Coralaxius abelei Kensley & Gore, 1981
(fig. 48)
Axiopsis sp. B, Coelho, 1971: 231.
Axiopsis (Paraxiopsis) sp. B, Coelho & Ramos, 1972: 160.
Coralaxius abelei Kensley & Gore, 1981: 1278, figs. 1-6, 2 tables; Abele & Kim, 1986: 26, 289,
291 (fig.); Sakai & De Saint Laurent, 1989: 2 (list), 11; Coelho & Ramos-Porto, 1985: 68;
Werding & Müller, 1989: 252; Melo, 1999: 316, figs. 213-214. [Type locality. — Atlantic
Ocean, French Reef, off Key Largo, Monroe Co., Florida, 25◦ 02′ N 80◦ 19′ W, 76 m.]
Material examined. — SMF 16575, 1 spm., coral protection area, Punta de Betin, Santa
Marta, Dept. Magdalena, Colombia, 6 m, 4.viii.1985, leg. H.G. Müller; SMF 16576, 1 spm.,
Punta de Betin, Santa Marta, Dept. Magdalena, Colombia, 13 m, 3.vi.1985, leg. H.G. Müller;
SMF 16577, 5 spms., Punta de Betin, Dept. Magdalena, Colombia, 13 m, 15.ix.1985, leg.
H.G. Müller; SMF 16578, 1 spm., Punta de Betin, Santa Marta, Dept. Magdalena, Colombia,
30 m, 10.ii.1986, leg. H.G. Müller; SMF 16579, 10 spms., Isla del Morro, Santa Marta,
Dept. Magdalena, Colombia, 25 m, 18.ix.1985, leg. H.G. Müller; SMF 16580, 3 spms.,
Salva Clengue, ca. 15 km N.E. of Santa Marta, Bahia Chengue, Dept. Magdalena, Colombia,
27.ix.1985, leg. H.G. Müller; SMF 16581, 3 spms., coral protection area, ca. 25 km N.E. of
Santa Marta, Bahia Nenguangue, Dept. Magdalena, Colombia, 11-16 m, 23.ix.1985, leg. H.G.
Müller; SMF 16582, 1 spm., coral protection area, ca. 25 km N.E. of Santa Marta, Bahia
Nenguangue, Dept. Magdalena, Colombia, 16 m, 10.x.1985, leg. H.G. Müller; SMF 16583,
3 spms., coral protection area, ca. 5 km E. of Santa Marta, Bahia Nenguangue, Punta Ancon near
256
CRM 013 – Katsushi Sakai
Taganga, Dept. Magdalena, Colombia, 15 m, 2.viii.1985, leg. H.G. Müller; SMF 16584, 1 spm.,
coral protection area, ca. 4 km E. of Santa Marta, Punta de Aguja, Dept. Magdalena, Colombia,
10 m, 24.ix.1985, leg. H.G. Müller; SMF 16585, 1 male (TL/CL, 8.0/2.1 mm), 1 ovig. female
(TL/CL, 9.0/2.9 mm), 3 females (TL/CL, 8.0/2.3-9.0/2.9 mm), coral protection area, ca. 4 km
E. of Santa Marta, Punta de Aguja, Dept. Magdalena, Colombia, 16-18 m, 16.ix.1985, leg. H.G.
Müller; SMF 16586, 2 spms., ca. 4 km E. of Santa Marta, Punta de Aguja, Dept. Magdalena,
Colombia, 18 m, 2.x.1985, leg. H.G. Müller; ZMB 26888, 1 male (TL/CL, 7.0/2.0 mm), 1 ovig.
female (TL/CL, 9.0/2.9 mm), eggs with nauplius eyes, 3 females (TL/CL, 9.0/2.9-12.0/3.9 mm),
Morro Grande, Santa Marta, Colombia, 25 m, coral protection area, 18.ix.1985, leg. H.G.
Müller; RMNH D 32920, 2 males, 3 females, paratypes, Caribbean Sea, Carrie Bow Cay,
Belize, 65 feet, 22.iii.1979 (B. Kensley: male gonopods illustrated in publication as C. abelei);
RMNH D 41748, 3 spms., Pillsbury Sta. P-1196, west Atlantic, N.E. of Jamaica, 17◦ 27.5′ N
75◦ 57′ W, 26 m, 10′ ottertrawl, 3.vii.1970, USNM Acc. No. 381702; RMNH D 41749, 1 spm.,
Pillsbury Sta. P-1317, west Atlantic, off Panama, 09◦ 59′ N 82◦ 55′ W, 55-69 m, 10′ ottertrawl,
27.i.1971, USNM Acc. No. 381702; RMNH D 41750, 1 spm., Pillsbury Sta. P-1252, west
Atlantic, S.W. of Jamaica, 17◦ 09′ N 78◦ 57′ W, 26 m, 10′ ottertrawl, 14.vii.1970, USNM Acc.
No. 381702; RMNH D 41751, 1 spm., Pillsbury Sta. P-1202, west Atlantic, N. of Jamaica,
17◦ 51.1′ N 76◦ 23.8′ W, 18 m, 10′ ottertrawl, 4.vii.1970, USNM Acc. No. 381702; RMNH D
41752, 1 spm., Pillsbury Sta. P-1204, west Atlantic, N. of Jamaica, 17◦ 46.1′ N 76◦ 46.0′ W, 3792 m, 10′ ottertrawl, 4.vii.1970, USNM Acc. No. 381702; RMNH D 41753, 1 spm., Pillsbury
Sta. P-1196, west Atlantic, N. of Jamaica, 17◦ 23.6′ N 76◦ 02.2′ W, 9-18 m, 10′ ottertrawl,
3.vii.1970, USNM Acc. No. 381702.
Diagnosis. — Rostrum (fig. 48A, B) short, acutely triangular, and deflexed
on dorsal surface. Gastric region bearing short unarmed median carina with
pyramidal hepatic convexity, and teeth arranged in circle posteriorly behind
rostrum; submedian carinae absent. Cervical groove distinct. A1 peduncle
with proximal segment equal in length to two distal segments, bearing three
denticles on ventral margin, and small distal denticle on somewhat inflated
stylocerite; segment 3 slightly shorter than segment 2; flagella about three
times as long as carapace. A2 peduncle with basal segment bearing small
tooth ventral to aperture of antennal gland; segment 2 with broad, distallybifurcate or obscurely-trifurcate scaphocerite; segment 3 set obliquely on
segment 2; segments 4 and 5 subequal in length; flagellum about 3 times
as long as carapace. Maxilla 2 scaphognathite bearing modified setal whip.
Mxp3 ischium and merus each bearing about 6 spines on mesial margin,
increasing in size distally; carpus with a single distal spine on mesial margin;
exopod consisting of proximal segment and deflected distal segment with
multiarticulate flagellum. P1 larger cheliped bearing ischium with 5 low
spines on ventral margin; merus with about 9 strong spines ventrally; carpus
denticulate on dorsal margin, lateral surface tuberculate; palm robust, lateral
surface tuberculate, dorsal margin dentate; fixed finger irregularly denticulate
on cutting edge; dactylus strongly denticulate on dorsal margin, cutting edge
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Fig. 48. Coralaxius abelei Kensley & Gore, 1981. A, B, gastric region, dorsal view. A, B, ZMB
26888, female (TL/CL, 12.0/3.9 mm), Murro Grande, Santa Marta, Colombia, 25 m. Scales
1 mm.
denticulate. P1 smaller cheliped with ischium, merus, and carpus shaped as in
larger cheliped; chela markedly less broad than in larger cheliped; palm about
two-thirds length of fingers, mesial and lateral surfaces tuberculate, dorsal
margin spinose; cutting edge of fixed finger denticulate, medially with two
large, acutely triangular teeth; dactylus denticulate on dorsal margin, cutting
edge denticulate. P2 chela broad, fingers longer than palm. Ps3-5 dactyli
biunguiculate distally. Abdominal pleuron 2 broadened, protruding anteroand posteroventrally. Male Plp1 uniramous and bisegmented, distal segment
ovate and bilobed distally, bearing patch of hooklets mesiodistally. Male Plp2
biramous, exopod bearing marginal plumose setae; endopod pestle-shaped,
bearing mesiomedially short appendix interna and broad, strongly bilobed
appendix masculina with stout marginal setae. Female Plp1 uniramous and
bisegmented, distal segment elongate, consisting of two articles with plumose
distal setae. In female Plp2, and Plps3-5 of both sexes, exopod slightly shorter
than endopod; endopod with short appendix interna mesiomedially.
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CRM 013 – Katsushi Sakai
Telson subtriangular and convergent posteriorly, bearing proximal lobe with
spine followed posteriorly by movable spine in middle on lateral margin, and
two spines on posterolateral margin with stiff setae, of which distal one long
and strong; dorsal surface with two pairs of teeth medially; posterior margin
fringed with elongate setae, bearing median spine. Uropodal endopod with
three spines on lateral margin, bearing dorsal carina with five spines including
distalmost marginal spine. Uropodal exopod bearing 3-4 spines on lateral
margin and large movable spine at distolateral angle; dorsal surface with two
spines on dorsal carina, and transverse suture with 6-8 spines. [Adapted from
Kensley, 1994.]
Remarks. — In Coralaxius abelei, the female Plp1 is uniramous and
bisegmented; the distal segment is elongate and bisegmented, but in the other
species of Coralaxius, C. galapagensis Kensley, 1994 and C. indopacificus
Kensley, 1994, it is an unsegmented ramus with distal setae (Kensley, 1994,
figs. 1L, 3L).
Type locality. — Florida — Key Largo, 25◦ 02′ N 80◦ 19′ W, 76 m.
Distribution. — Florida — Key Largo (Kensley & Gore, 1981), Looe Key,
and southwest of Alligator Light (Sakai & De Saint Laurent, 1989); Strait of
Florida (Kensley, 1994); off Jamaica (Kensley, 1994); off Costa Rica (Kensley,
1994); Belize (Sakai & De Saint Laurent, 1989); Carrie Bow Cay, Belize
(Kensley & Gore, 1981), Glovers Reef, Curlew Cay, Belize (Kensley, 1994);
Colombia; Bahia, Espírito Santo & Rocas, Brazil (Coelho & Ramos-Porto,
1985); 6-230 m.
Coralaxius galapagensis Kensley, 1994
Coralaxius galapagensis Kensley, 1994: 815, figs. 1, 2.
Diagnosis. — Rostrum short, triangular, and deflexed on dorsal surface.
Gastric region bearing short unarmed median carina with pyramidal hepatic
convexity medially, and teeth arranged in circle posteriorly behind rostrum;
submedian carinae absent. Cervical groove distinct. A2 basal segment with
two short spines ventral to aperture of antennal gland; segment 2 with distally
bifid scaphocerite. Mxp3 ischium with a few small teeth and a single large
distal tooth on mesial margin; merus with three large mesiodistal teeth; carpus
with a single large mesiodistal tooth; exopod consisting of proximal segment
reaching distal margin of merus, and deflected distal segment with flagellum.
In P1 larger cheliped, ischium serrate on ventral margin; merus roughly serrate
and armed with strong distal tooth; carpus scattered with acute tubercles on
lateral surface, and serrate and armed with strong distal tooth on dorsal margin;
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palm 1.5 times length of fingers, strongly serrate on dorsal and ventral margins,
lateral surface fairly densely tuberculate, several tubercles acute; cutting edge
of fixed finger with a single moderately strong tubercle in middle; dactylus
strongly serrate on dorsal margin, bearing a row of tubercles and several ones
scattered on lateral surface, cutting edge with broad, rounded tubercles at
base. In P1 smaller cheliped, ischium denticulate on ventral margin; merus
with 10 teeth on ventral margin, increasing in size distally; carpus scattered
with acute tubercles on lateral surface and strong denticles on dorsal margin;
palm about three-fifths length of fingers, and strongly serrate dorsally, bearing
numerous acute tubercles scattered on lateral surface, and strongly serrate on
ventral margin; fixed finger denticulate on cutting edge, bearing two distinct
teeth in distal half; dactylus strongly serrate on dorsal margin, and cutting edge
denticulate. Abdominal pleura rounded ventrally. Telson subtriangular, about
1.4 times as long as wide, bearing two teeth laterally, a single strong, movable
spine at posterolateral angle, and two pairs of blunt teeth on dorsal surface.
Uropodal endopod bearing three obtuse teeth including distal one on lateral
margin, and 4 teeth on dorsomedian carina. Uropodal exopod bearing 4 teeth
on lateral margin and strong spine at distolateral angle; dorsal surface with
two teeth on sublateral carina and transverse suture with 6 teeth. [Adapted
from Kensley, 1994.]
Type locality. — Sullivan Bay, James Is., San Salvador, Galapagos.
Distribution. — Galapagos Islands (Kensley, 1994).
Coralaxius indopacificus Kensley, 1994
Coralaxius indopacificus Kensley, 1994: 820, figs. 3, 4.
Diagnosis. — Rostrum short, triangular, and deflexed on dorsal surface.
Gastric region bearing short median carina with 2-3 spines and pyramidal hepatic convexity medially, and lateral carinae with 2-5 teeth, submedian carinae
absent. Cervical groove distinct. A1 peduncle reaching far beyond eyestalks,
proximal segment equal in length to two distal segments, bearing three small
spines on dorsal margin; stylocerite somewhat inflated and terminating with a
single small distal spine on ventral margin; segment 2 slightly longer than segment 3. A2 proximal segment with a single short spine ventral to aperture of
antennal gland; segment 2 bearing distally bifid scaphocerite. Mxp3 ischium
with 4 small teeth on mesial margin; merus with two small proximal and two
large distal teeth on mesial margin; carpus with small distal spine; exopod
consisting of proximal segment and deflected distal segment with flagellum.
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CRM 013 – Katsushi Sakai
In larger cheliped ischium denticulate on ventral margin; merus roughly serrate on dorsal margin, and 6 small spines and a strong distal one on ventral
margin; carpus with ventral spine proximally, dorsal and lateral surfaces scattered with acute tubercles; palm 1.3 times as long as fingers, bearing acute
tubercles on lateral surface and dorsal and ventral margins; fixed finger irregularly tuberculate on cutting edge; dactylus roughly serrate on dorsal margin,
and scattered with acute denticles on lateral surface, cutting edge irregularly
tuberculate. In smaller cheliped, merus serrate and armed with two or three
distal teeth on dorsal margin, bearing 9 small teeth and a strong distal one on
ventral margin; carpus serrate on dorsal margin and scattered with acute denticles on lateral surface; palm 0.6 times as long as fingers and strongly serrate
on ventral margin, bearing acute denticles on lateral surface; fixed finger with
two strong teeth and some small ones on cutting edge; dactylus strongly serrate on dorsal margin, and barely serrate on cutting edge. Abdominal pleura
rounded ventrally. Telson hexagonal, bearing two pairs of teeth in distal half
of dorsal surface; lateral margins with two teeth, two small and one strong
movable spines, and stiff setae posterolaterally; posterior margin with median
spine. Uropodal endopod with three teeth including distal one on lateral margin. Uropodal exopod with three teeth including distal tooth on lateral margin,
movable spine at distolateral angle, and transverse suture with 6 teeth dorsally.
[Adapted from Kensley, 1994.]
Type locality. — Astrolabe Reef, Kandavu, Fiji, 5-8 m.
Distribution. — Comoro Islands (Kensley, 1994); Fiji (Kensley, 1994);
5-20 m.
Genus Meinertaxius gen. nov.
Diagnosis. — Rostrum short and acutely triangular. Gastric region with
median carina originating from base of rostrum and running to anterior half
of gastric region; posterior half of median carina forming elongate triangular
swelling in dorsal view, submedian and lateral carinae absent. Anterior margin
of carapace with three spines including supraocular spine on each side of small
rostrum. Cervical groove distinct. Eyestalks thick; cornean pigments faded in
yellow colour. Mxp3 exopod consisting of proximal segment and deflexed
distal segment with flagellum. P1 unequal. P2 chelate; chela ovate in shape.
Ps3-4 simple, propodi elongate, bearing spines on ventral margin; dactyli
biunguiculate. No pleurobranchs. Abdominal pleuron 1 rounded ventrally;
pleuron 2 protruded antero- and posteroventrally, and truncate ventrally. Male
Plp1 uniramous and bisegmented, distal segment bilobed distally; male Plp2
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261
biramous, endopod with stout appendix masculina and small bisegmented
appendix interna, distal segment with patch of hooklets distally; male Plps35 biramous, endopods bearing appendix interna. Telson subtriangular and
convergent posteriorly, and denticulate on posterolateral margin with stiff
setae, bearing median spine on convex posterior margin. Uropodal exopod
bearing spines distally on lateral margin, and denticulate transverse suture.
Remarks. — It has turned out that Coralaxius nodulosus (Meinert, 1877)
(cf. Sakai & De Saint Laurent, 1989) is not included in Coralaxius, though
it is similar to the type species of Coralaxius, Coralaxius abelei Kensley
& Gore, 1981 in that the gastric region bears a median carina originating
from the base of the rostrum and running to the anterior half of the gastric
region, and also in the shapes of the eyestalks, A1-2 peduncles, Mxp3, Ps1-2,
dactyli of Ps3-4 (probably also of P5), abdominal pleura 1-6, and male Plp1.
However, they are different from each other at the generic level as follows:
in Coralaxius nodulosus the gastric region bears no teeth arranged in a circle
or semicircle anteriorly behind the rostrum, the Mxp3 exopod consists of a
bisegmented proximal segment and a deflexed distal segment with a flagellum,
the male Plp2 appendix masculina is simple on its distal margin, and the Plp2
exopod is larger than the endopod. In contrast, in Coralaxius abelei the gastric
region bears teeth arranged in a circle or semicircle anteriorly behind the
rostrum, the Mxp3 exopod consists of an unsegmented proximal segment and
a deflexed distal segment with a flagellum, the male Plp2 appendix masculina
is bilobed distally, and the Plp2 exopod is shorter than the endopod. Therefore,
Coralaxius nodulosus is reclassified under the new genus Meinertaxius, and
distinguished from the other species included in Coralaxius.
Type species. — Axius nodulosus Meinert, 1877, by present designation and
monotypy. The gender of the new name, Meinertaxius, is masculine.
Species included. — Meinertaxius nodulosus (Meinert, 1877).
Etymology. — The new genus is named Meinertaxius in honour of F. Meinert, a Danish carcinologist who described this species in 1877, by adding his
name as a prefix to -axius.
Meinertaxius nodulosus (Meinert, 1877)
(figs. 49, 50)
Axius nodulosus Meinert, 1877: 212; Stephensen, 1910a: 276, figs. 1-5; Stephensen, 1910b: 75,
fig. 49; De Man, 1925d: 5, 18; Balss, 1926: 26; Poulsen, 1940: 208, 236 (key), fig. 2; Ingle
& Christiansen, 2004: 77.
?Axiopsis nodulosus — Borradaile, 1903: 539.
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CRM 013 – Katsushi Sakai
Fig. 49. Meinertaxius nodulosus (Meinert, 1877). A, whole body, lateral view; B, carapace,
dorsal view; C, gastric region, dorsolateral view. A-C, ZMC CRU-587, lectotype, male (TL/CL,
6.5/2.2 mm), off Nymindegab, North Sea. Scales 0.5 mm.
Coralaxius nodulosus — Sakai & De Saint Laurent, 1989: 2 (list), 11 [typing error, p. 101,
abelei], table 2; Kensley, 1994: 822, figs. 5, 6; Robles et al., 2009: 316.
Material examined. — ZMC CRU-587, lectotype male (TL/CL, 6.5/2.2 mm), off Nymindegab, North Sea, ca. 1840, leg. H. Krøyer.
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Diagnosis. — Rostrum (fig. 49B, C) short, triangular in dorsal view, unarmed laterally, protruded to nearly proximal fourth of eyestalks. Anterior margin of carapace with three spines including distinct supraocular spine on each
side of rostrum. Gastric region bearing median carina running from base of
rostrum to anterior half of gastric region, posterior half of median carina forming elongate triangular swelling with fine tubercles on surface in dorsal view
(considered a deformation of the hepatic tubercle), submedian and lateral carinae absent. Cervical groove distinct. No postcervical carina. Eyestalks thick;
cornean pigments faded in yellow colour. A1 peduncle not reaching distal margin of A2 penultimate segment. Mxp3 merus with strong subdistal spine on
ventral margin; propodus finely serrate on ventral margin; dactylus with long
setae ventrally; exopod consisting of bisegmented proximal segment not reaching distal margin of merus, and deflected distal segment with multiarticulate
flagellum. P1 chelate and unequal. In larger (right) cheliped (figs. 49A, 50A),
merus with 2-3 low teeth on dorsal margin and 4 strong teeth distally on ventral margin; carpus with 4 distally-directed spines on dorsal margin; palm distinctly crenulate on dorsal margin but indistinctly on ventral margin; dactylus
crenulate dorsally and distinctly denticulate on cutting edge; cutting edge of
fixed finger armed with teeth. In smaller (left) cheliped (fig. 50B), merus with
two lower teeth on dorsal margin and two distinct teeth distally on ventral
margin; carpus bearing distal spine on dorsal margin but smooth on ventral
margin; palm smooth on dorsal margin, but 4 lower teeth medially on ventral
margin; cutting edge of fixed finger bearing 6 sharp, distally-directed teeth;
dactylus twice length of palm, bearing 5 lower, distally-directed teeth. P2
(figs. 49A, 50C) ischium with subdistal tooth on ventral margin; merus denticulate with distinct subdistal tooth on ventral margin, but smooth on dorsal
margin; carpus with triangular subdistal tooth on ventral margin; chela rounded
in shape, and fixed finger proximally with three distinct spines on cutting edge;
dactylus 1.8 times as long as palm and unarmed on cutting edge. Ps3-4 simple,
propodi elongate, bearing ventral spines; dactyli blade-shaped and biunguiculate distally, carrying about 4-5 teeth and a couple of setae on ventral margin.
No pleurobranchs. Abdominal pleuron 1 (fig. 50D) rounded ventrally; pleuron
2 protruded antero- and posteroventrally, and rounded ventrally. Male Plp1
uniramous and bisegmented, distal segment emarginate distally. Male Plp2
(fig. 50F) biramous, endopod shorter than exopod, bearing mesiomedially distinct appendix masculina and small bisegmented appendix interna with distal
hooklets. Plps3-5 biramous, endopods with appendix interna. Telson (fig. 50E)
subtriangular and convergent, bearing laterally a low proximal lobe followed
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CRM 013 – Katsushi Sakai
Fig. 50. Meinertaxius nodulosus (Meinert, 1877). A, larger cheliped; B, smaller cheliped; C, P2;
D, abdomen and tail-fan, lateral view; E, abdominal somite 6, telson, and uropod on left side;
F, male Plp2. A-F, ZMC CRU-587, lectotype, male (TL/CL, 6.5/2.2 mm), off Nymindegab,
North Sea. Scales A, B, C, D, E, 1 mm; F, 0.1 mm.
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by followed by denticles with stiff setae at posterolateral angle, and median
spine on convex posterior margin; dorsal surface bearing transverse row of
irregularly-arranged spinules at proximal third. Uropodal exopod with transverse suture.
Remarks. — Poulsen (1940: 208) described the present species, Meinertaxius nodulosus as a young post-larval specimen of Axius nodulosus because
of its small size in comparison with other axiid specimens, which fact was
referred to also by Ingle & Christiansen (2004: 77).
Kensley (1994: 822) considered his species Coralaxius abelei Kensley &
Gore, 1981 from the Florida Keys and the western Caribbean Sea, to be
synonymous with Axius nodulosus Meinert, 1877 from Nymindegab, North
Sea, by developing his theory as follows; “a more likely scenario, however,
is that the evolution reflects a tethyan distribution from tropical Indo-West
Pacific to Western Atlantic and Eastern Pacific before the raising of the
Isthmus of Panama” (Kensley, 1994: 827). This theory of his prevented him,
unfortunately, from paying attention to the morphological differences between
the two species, Axius nodulosus Meinert, 1877 and Coralaxius abelei Kensley
& Gore, 1981. The mophological differences found between them make it
possible to distinguish the one from the other. It is also safely said even,
that those two species are clearly different in morphology from each other
at the generic level, because in A. nodulosus the gastric region bears no teeth
arranged in a circle posterior to the rostrum, and the male Plp2 endopod is
provided with a simple appendix masculina, whereas in C. abelei the gastric
region bears teeth arranged in a circle posteriorly behind the rostrum, and
the male Plp2 endopod bears a setose appendix masculina which is distally
bilobed. As a result, Axius nodulosus Meinert, 1877 from Nymindegab, North
Sea is reclassified under the new genus Meinertaxius as a valid species
Meinertaxius nodulosus (Meinert, 1877), and is distinguished from Coralaxius
abelei Kensley & Gore, 1981.
Type locality. — Nymindegab, North Sea.
Distribution. — North Sea — west coast of Jutland (Meinert, 1877; Poulsen,
1940; Kensley, 1994).
FAMILY EICONAXIIDAE SAKAI & OHTA, 2005
Eiconaxiidae Sakai & Ohta, 2005: 69.
Diagnosis. — Rostrum stout, conspicuous, and pointed or rounded apically,
and carinate medially with or without denticles; rostral median carina extending posteriorly as gastric median carina with or without bifurcate posterior end;
and rostral lateral margins extending posteriorly onto gastric region as gastric
lateral carinae; gastric submedian carinae absent. Carapace convex on dorsal
surface, linea thalassinica absent, cervical groove usually indistinct, posterior
margin of carapace with lateral lobes. Eyestalks indiscernible in lateral view;
eyes rounded or ovate, cornea pigmented or unpigmented. A1 segment 1 as
long as segment 2. Scaphocerite strong. Maxilla 2 scaphognathite without posterior whip. Mxp3 ischial crest crenulate, exopod present. Ps1-2 chelate. Ps3-5
dactyli short, subspatulate in form, and spinulate on ventral margin. P4 coxa
slightly flattened. Abdominal somite 1 with pleuron slightly protruded posteroventrally; abdominal somites without carina between tergum and pleuron.
Male Plp1 absent; male Plp2 biramous and narrow, endopod with appendices
interna and masculina or only with appendix interna; male Plps3-5 biramous,
endopods with appendix interna. Female Plp1 uniramous and bisegmented;
distal segment and flagellum with setae; Plps2-5 slender and biramous, endopods with appendix interna. Telson longer than wide, bearing no proximal
lobe on lateral margin. Uropodal exopod without transverse suture.
Remarks. — The family Eiconaxiidae Sakai & Ohta, 2005 is separated
from the family Axiidae Huxley, 1878, because it is very characteristic in that
Ps3-5 are short, subspatulate in shape, and spinulate on the ventral margin;
maxilla 2 bears no posterior whip; and the Mxp3 ischial crest is crenulate,
which features are not observed in the Axiidae. The Eiconaxiidae are divided
into two genera, Eiconaxius and Eiconaxiopsis gen. nov., by the distinct
morphological differences between them, especially in Plp1. They are different
from each other as follows. In Eiconaxius the eyes are rounded, and the cornea
is pigmented; male Plp1 is absent; and the telson bears a median spine on the
posterior margin. In Eiconaxiopis gen. nov. the eyes are ovate laterally, and the
cornea is unpigmented; male Plp1 is present as a uniramous and bisegmented
appendage, and the distal segment is elongate and slightly convex on its mesial
margin with setae; the telson bears no median spine on the posterior margin.
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CRM 013 – Katsushi Sakai
Type genus. — Eiconaxius Bate, 1888, as the nominate genus.
Genera included. — Eiconaxius Bate, 1888.
Genus Eiconaxius Bate, 1888
Eiconaxius Bate, 1888: 40; Ortmann, 1891: 49; Stebbing, 1893: 188; Lagerberg, 1908: 48
(partim); Sakai & De Saint Laurent, 1989: 15; Sakai, 1992a: 158; Poore, 1994: 97 (key);
Kensley, 1996e: 474; Sakai & Ohta, 2005: 69; Poore & Collins, 2009: 255.
Iconaxiopsis Alcock, 1901: 193 [type species: Eiconaxius kermadeci var. laccadivensis Alcock
& Anderson, 1894, through subsequent designation by Borradaile, 1903]; Borradaile, 1903:
537; Balss, 1925: 210.
Iconaxius Alcock, 1901: 193 (miss-spelling).
Iconaxiopsis s. str. — Borradaile, 1903: 537.
Axius (Eiconaxius) — Borradaile, 1903: 537; De Man, 1925d: 1 (key), 14 (in key); Sakai,
1987a: 303 (list).
Diagnosis. — Rostrum distinct, triangular, and concave dorsally, bearing
median carina extending posteriorly onto gastric region as gastric median
carina, and bifurcate posteriorly; rostral apex acute or rounded, and lateral
margins armed or unarmed, extending posteriorly onto gastric region. Cervical groove inconspicuous. Eyestalks indiscernible, eyes ovate, cornea pigmented or not. A2 peduncular segment 2 with distinct distodorsal spine; A2
scaphocerite conspicuously elongate. P1 subequal or unequal. P2 chelate. Ps35 dactyli short, subspatulate, and spinulose on ventral margin. Pleurobranchs
present on Ps2-4. Pleura and terga entire. Male Plp1 absent; male Plp2 biramous and foliaceous, endopod with appendices interna and masculina mesiomedially, or with appendix interna but without appendix masculina. Male
Plps3-5 endopods with appendix interna. Female Plp1 uniramous and bisegmented; proximal segment and distal flagellum; female Plps2-5 biramous, endopods with appendix interna. Telson longer than wide and denticulate on
lateral margin, bearing median spine on posterior margin. Uropodal exopod
without transverse suture. Deep-water habitat.
Remarks. — In the present genus Eiconaxius, the male Plp2 has two
types: (1) the appendices interna and masculina are present as in E. agassizi
Bouvier, 1905, E. albatrossae Kensley, 1996, E. antillensis Bouvier, 1905, and
E. consobrinus (De Man, 1907); (2) the appendix interna is present, but the
appendix masculina is absent as in E. acutifrons Bate, 1888.
Type species. — Eiconaxius acutifrons Bate, 1888, subsequently designated
by Borradaile, 1903. The gender of the generic name, Eiconaxius, is masculine.
Species included. — Eiconaxius acutifrons Bate, 1888; E. agassizi Bouvier, 1905; E. albatrossae Kensley, 1996e; E. andamanensis (Alcock, 1901);
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E. antillensis Bouvier, 1905; E. asper Rathbun, 1906; E. baja Kensley,
1996e; E. bandaensis sp. nov.; E. borradailei Bouvier, 1905; E. caribbaeus
Faxon, 1896; E. carinatus Bouvier, 1925; E. consobrinus (De Man, 1907);
E. cristagalli Faxon, 1893; E. demani Sakai, 1992a; E. farreae Ortmann, 1891;
E. faxoni sp. nov.; E. hakuhou Sakai & Ohta, 2005; E. indicus (De Man, 1907);
E. indonesicus sp. nov.; E. kermadeci Bate, 1888; E. kimbla Kensley, 1996e;
E. laccadivensis (Alcock & Anderson, 1894); E. mallacoota Poore & Collins,
2009; E. mortenseni Sakai, 1992a; E. parvus Bate, 1888; E. rotundifrons Bouvier, 1905; E. singularis (Zarenkov, 1983); E. spiniger (MacGilchrist, 1905);
E. weberi (De Man, 1907).
K EY
TO THE
S PECIES
OF THE
G ENUS EICONAXIUS
1 – Rostrum with smooth and rounded apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Rostrum with arcuate or triangular apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2 – Merus of larger cheliped with lateral carina . . . . . . . . . . . . . . . . . . . . . . . . . . . E. carinatus
– Merus of larger cheliped without lateral carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Cutting edge of fixed finger of larger cheliped smooth . . . . . . . . . . . . . E. andamanensis
– Cutting edge of fixed finger of larger cheliped denticulate . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – Rostral median carina armed with denticles . . . . . . . . . . . . . . . . . . E. bandaensis sp. nov.
– Rostral median carina unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Rostrum denticulate on lateral margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
– Rostrum unarmed on lateral margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6 – Cutting edge of fixed finger of larger cheliped with convexity with distinct tooth . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. rotundifrons
– Cutting edge of fixed finger of larger cheliped with two proximal covexities . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. borradailei
7 – Palm of larger cheliped armed with denticles ventrally on distal margin . . . . . . . . . . . . 8
– Palm of larger cheliped unarmed ventrally on distal margin . . . . . . . . . . . . . . . . . . . . . . . 9
8 – Palm of larger cheliped with two denticles ventrally on distal margin . . . . . . E. demani
– Palm of larger cheliped denticulate ventrally on distal margin . . . . . . . . . . . . . E. parvus
9 – Dactylus of larger cheliped with median tooth on cutting edge . . . . . . . . . E. mortenseni
– Dactylus of larger cheliped proximally convex on cutting edge . . . . . . . . E. caribbaeus
10 – Palm of larger cheliped denticulate on dorsal margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
– Palm of larger cheliped smooth on dorsal margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
11 – Chela of larger cheliped setose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. spinigerus
– Chela of larger cheliped not setose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 – Palm of larger cheliped unarmed on ventrodistal margin . . . . . . . . . . . . . . . . . . . E. faxoni
– Palm of larger cheliped convex on ventrodistal margin . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 – Rostral median carina denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. indicus
– Rostral median carina smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 – Dorsal margin of palm of smaller cheliped unarmed . . . . . . . . . . E. indonesicus sp. nov.
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CRM 013 – Katsushi Sakai
– Dorsal margin of palm of smaller cheliped denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 – Rostrum turned-up distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. baja
– Rostrum not turned-up distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. albatrossae
16 – Rostrum unarmed on lateral margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
– Rostrum serrate on lateral margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17 – Rostral median carina smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. kimbla
– Rostral median carina denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. hakuhou
18 – Pleura 2-4 less sharply pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
– Pleura 2-4 almost sharply pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
19 – Cutting edge of dactylus of larger cheliped unarmed . . . . . . . . . . . . . . . E. laccadivensis
– Cutting edge of dactylus of larger cheliped proximally with cusp . . . . . . . . . . . E. asper
20 – Rostrum short and as long as wide at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. weberi
– Rostrum more than half length of width at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 – Rostral median carina denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
– Rostral median carina smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22 – Cutting edge of fixed finger of larger cheliped proximally notched . . . . . . E. cristagalli
– Cutting edge of fixed finger of larger cheliped widely concave proximally . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. antillensis
23 – Cutting edge of fixed finger with row of denticles . . . . . . . . . . . . . . . . . . . . . E. singularis
– Cutting edge of fixed finger distintly serrated or denticulate, or armed with a cusp. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
24 – Cutting edge of fixed finger of larger cheliped strongly or moderately serrated, or
denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
– Cutting edge of fixed finger of larger cheliped bearing a cusp . . . . . . . . . . . . . . . . . . . . 27
25 – Cutting edge of fixed finger of larger cheliped irregularly denticulate in proximal half
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. mallacoota
– Cutting edge of fixed finger of larger cheliped distinctly serrated . . . . . . . . . . . . . . . . . 26
26 – Cutting edge of fixed finger of larger cheliped strongly serrated . . . . . . . . . E. acutifrons
– Cutting edge of fixed finger of larger cheliped compactly serrated . . . . . . E. kermadeci
27 – Merus of larger cheliped with distinct median tooth on ventral margin . . . . E. agassizi
– Merus of larger cheliped distally with denticles or unarmed . . . . . . . . . . . . . . . . . . . . . 28
28 – Cutting edge of fixed finger of larger cheliped notched proximally, and followed by
small cusp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. consobrinus
– Cutting edge of fixed finger of larger cheliped proximally with convexity bearing distal
cusp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. farreae
Eiconaxius acutifrons Bate, 1888
Eiconaxius acutifrons Bate, 1888: 40, fig. 4, pl. 5 fig. 2d, g, h, I, k, l, q; Sakai & De Saint
Laurent, 1989: 16 (partim); Hendrickx, 1995: 390 (list); Kensley, 1996e: 475 (list).
Axius (Eiconaxius) acutifrons — Borradaile, 1903: 538.
Iconaxius acutifrons — Balss, 1925: 209 (partim).
Not: Axius acutifrons — Faxon, 1895: 103, pl. 28 fig. 2 [= Eiconaxius faxoni sp. nov.].
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271
Not: Axius (Eiconaxius) acutifrons — De Man, 1925d: 4 (list), 15 (key), 37, pl. 3 fig. 5-5e
[= Eiconaxius indonesicus sp. nov.].
Diagnosis. — Male Plp2 biramous, endopod with appendix interna, but
without appendix masculina. [From Bate, 1888, pl. 5 fig. 2q.]
Remarks. — Eiconaxius acutifrons was reported by Bate in 1888 from the
Banda Sea for the first time, and was next reported by Faxon in 1895 from
southern Panama (Sta. 3358, 3359, 4◦ 31′ S 129◦ 57.20′ E, 360 fathoms (658 m)),
and thirdly reported by De Man in 1925 from the southeast coast of Great Kei
Island. However, it seems that those three acutifrons are different from one
another.
Bate’s specimens and De Man’s specimen are different from each other,
even though they were reported from almost the same locality, Indonesia, in the
forms of the rostrum and chelipeds. In Bate’s E. acutifrons (male) the rostrum
is narrowly triangular in males, or broadly triangular in females (Bate, 1888:
40), and in the male larger cheliped on the left side (Bate, 1888, pl. 5 fig. 2kl),
the cutting edge of the fixed finger is strongly serrated, the dactylus is broad,
and the cutting edge bears proximally a convexity and is then curved in S-shape
distally. In De Man’s E. acutifrons (male), by contrast, the rostrum is triangular
and acuminate at the tip, and a little longer than wide at its base; in the male
larger cheliped on the left side (De Man, 1925d: 39, fig. 5b), the cutting edge of
the fixed finger bears a median cusp followed distally by denticulation toward
the tip, and the cutting edge of the dactylus carries a distinct, truncate proximal
convexity in its proximal third, and then concave and smooth. Accordingly, it
is proposed that De Man’s E. acutifrons is renamed E. indonesicus sp. nov.
Faxon’s E. acutifrons from Panama is geographically and morphologically
different from Bate’s and De Man’s E. acutifrons, as shown in the figure of
the right cheliped given by Faxon (1895, pl. 28 fig. 2), in that the palm is
armed with 4 teeth including a distodorsal tooth on the distal half of the dorsal
margin, and the cutting edge of the fixed finger bears a distinct triangular tooth
at its proximal third, and then denticulate toward the tip, and the cutting edge
of the dactylus bears a broad, truncate convexity in its proximal third, and then
is further concave with a small median tooth in its distal third, which suggests
that Faxon’s E. acutifrons is also to be differentiated from Bate’s and De Man’s
E. acutifrons, and is here renamed E. faxoni sp. nov.
Type locality. — Off Banda, Challenger Sta. 194A, 4◦ 31.0′ S 129◦ 57.20′ E,
360 fathoms (658.4 m), volcanic mud.
Distribution. — Off Banda, Challenger Sta. 194A, 360 fathoms [ca. 658 m]
(Bate, 1888); Banda, 660 m (Balss, 1925); southeast coast of Great Kei Island,
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CRM 013 – Katsushi Sakai
“Siboga” Sta. 266, 5◦ 56.5′ S 132◦ 47.7′ E, 595 m, grey mud with coral and
stones (De Man, 1925d).
Eiconaxius agassizi Bouvier, 1905
Eiconaxius Agassizi Bouvier, 1905: 803.
Iconaxius Agassizi — Balss, 1925: 209.
Axius (Eiconaxius) agassizi — Bouvier, 1925: 458, fig. 22, pl. 7 fig. 5, pl. 9 fig. 2.
Axius (Eiconaxius) Agassizi — De Man, 1925d: 4 (list), 17 (key).
Eiconaxius agassizi — Sakai & De Saint Laurent, 1989: 20; Kensley, 1996e: 475 (list).
Material examined. — RMNH D 41754, 3 males (TL/CL, 17.0/5.6-18.0/6.6 mm), Pillsbury
Sta. P-1187, west Atlantic, N. of Jamaica, 18◦ 17′ N 75◦ 07′ W, 1033 m, 10′ ottertrawl, 2.vii.1970,
USNM Acc. No. 381702.
Diagnosis. — Male Plp1 absent, male Plp2 biramous, and endopod bearing
appendices interna and masculina; male Plps3-5 biramous, and endopods with
appendix interna.
Type locality. — West Indies to Gulf of Mexico, 523-1548 m.
Distribution. — Only known from the type locality.
Eiconaxius albatrossae Kensley, 1996
Eiconaxius albatrossae Kensley, 1996e: 475, figs. 4, 5.
Diagnosis. — Male Plp1 absent, male Plp2 biramous, endopod with appendices interna and masculina at proximal third.
Type locality. — Pacific Panama, 6◦ 22′ 20′′ N 81◦ 52′ 11′′ W, “Albatross” Sta.
3359, 851 m.
Distribution. — Pacific Panama, 851-1016 m.
Eiconaxius andamanensis (Alcock, 1901)
Iconaxiopsis andamanensis Alcock, 1901: 196, pl. 2 fig. 1; Alcock & McArdle, 1902, pl. 58
fig. 2; Balss, 1925: 209.
Axius (Iconaxiopsis) andamanensis — Borradaile, 1903: 537.
Axius (Eiconaxius) andamanensis — De Man, 1925d: 4 (list), 16 (key).
Eiconaxius andamanensis — Sakai & De Saint Laurent, 1989: 20; Kensley, 1996e: 475 (list).
Remarks. — Alcock (1901: 196) mentioned that this species is closely
similar to E. laccadivensis (Alcock & Anderson, 1894), but is distinguished
from the latter as follows. In E. andamanensis the rostrum is smooth on the
lateral margins, and the cutting edges of the dactylus and fixed finger are
smooth (Alcock, 1901: 196, pl. 2 fig. 1), whereas in E. laccadivensis the
rostrum is microscopicallly serrulate on the lateral margins, and the cutting
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273
edge of the dactylus is concave to correspond with a slightly enlarged proximal
tooth on the cutting edge of the fixed finger (Alcock & MacGilchrist, 1905, pl.
71 fig. 3).
Type locality. — Off the west coast of Andaman Islands, 428-522 m.
Distribution. — Only known from the type locality.
Eiconaxius antillensis Bouvier, 1905
Eiconaxius crista-galli var. antillensis Bouvier, 1905: 803.
Iconaxius cristagalli var. antillensis — Balss, 1925: 210.
Axius (Eiconaxius) crista-galli antillensis — Bouvier, 1925: 456, pl. 8 fig. 3, pl. 9 fig. 1.
Axius (Eiconaxius) crista-galli var. antillensis — De Man, 1925d: 4 (list), 33.
Econaxius antillensis — Sakai & De Saint Laurent, 1989: 21; Kensley, 1996e: 475 (list).
Material examined. — RMNH D 41756, 1 ovig. female (TL/CL, 17.0/6.3 mm), Pillsbury
Sta. P-1262, west Atlantic, S. of Jamaica, 17◦ 21.1′ N 77◦ 34.8′ W, 805-1089 m, 10′ ottertrawl,
10.vii.1970, USNM Acc. No. 381702; RMNH D 41757, 1 male (TL/CL, 17.0/6.3 mm),
Pillsbury Sta. P-944, west Atlantic, N. of Guadeloupe, 16◦ 33.2′ N 61◦ 36.8′ W – 16◦ 34.4′ N
61◦ 37.2′ W, 360-421 m, 5′ Black trawl, 17.vii.1969, USNM Acc. No. 381702.
Diagnosis. — Male Plp1 absent; male Plp2 biramous, endopod with appendix masculina and short appendix interna; Plps3-5 endopods with appendix
interna. Female Plp1 uniramous, consisting of proximal segment and flagellum; female Plps2-5 endopods with appendix interna.
Type locality. — Antilles and Gulf of Mexico, 518-536 m.
Distribution. — Gulf of Mexico (Bouvier, 1905); Antilles (Bouvier, 1905);
518-536 m.
Eiconaxius asper Rathbun, 1906
Eiconaxius asper Rathbun, 1906: 895, fig. 52; Sakai & De Saint Laurent, 1989: 22; Kensley,
1996e: 475 (list).
Iconaxius asper — Balss, 1925: 209.
Axius (Eiconaxius) asper — De Man, 1925d: 4 (list), 14 (key).
Type locality. — Off Kauai Island, Hawaiian Islands, 752-950 m.
Distribution. — Hawaiian Islands — Kauai Island (Rathbun, 1906); 752950 m.
Eiconaxius baja Kensley, 1996
Eiconaxius baja Kensley, 1996e: 478, fig. 6, table 3.
Remarks. — The present species, Eiconaxius baja is closely similar to E. albatrossae, because, as shown by Kensley (1996e), the external characteristics
274
CRM 013 – Katsushi Sakai
of the rostrum, median and lateral carinae on the gastric region, the shape and
denticulation of P1, pleura, and tail-fan, are fundamentally the same between
the two species. However, they differ as follows. In E. baja the rostrum is vertically turned up at the apex, and the lateral surface of the P1 palm is armed
with strong tubercles with setae, whereas in E. albatrossae the rostrum is not
turned up at the apex, and the lateral surface of the P1 palm bears no setose
tubercles.
Type locality. — Baja California, 28◦ 57′ 00′′ N 118◦ 14′ 30′′ W, “Albatross”
Sta. 2986, 1252 m.
Distribution. — So far only known from the type locality.
Eiconaxius bandaensis sp. nov.
Eiconaxius sibogae — Sakai, 1992a: 163, fig. 6 [not: De Man, 1925a: 218; De Man, 1925b: 34,
fig. 4-4L].
Material examined. — ZMC, holotype, female (TL/CL, 19.0/6.2 mm, with chelipeds);
paratype, female (TL/CL, 19.0/6.2 mm, without chelipeds), Banda Sea, 5◦ 47′ S 132◦ 51′ E,
348 m, trawl, KIE Sta. 51, 7.v.1922, leg. Danish Expedition to Kei Islands, 1922.
Diagnosis of female holotype. — Rostrum triangular with rounded apex and
unarmed on lateral margins, rostral median carina with 1-4 denticles extending
onto gastric region to form bifurcate gastric median carina with denticles
anterior to junction. In larger cheliped, ischium with subdistal spine on ventral
margin; merus 1.3 times as long as wide, lateral surface convex, ventral margin
straight with minute serration and triangular distal spine, dorsal margin largely
rounded; carpus less than half length of merus, half as long as wide, bearing
obtuse spine on ventral angle; chela about six times as long as carpus; palm 1.3
times as long as wide, lateral surface carinate along ventral margin, distal
margin convex medially and concave at ventral corner, then continued to
cutting edge of fixed finger; fixed finger half length of palm, cutting edge
with proximal tubercle in smooth proximal half, and armed in distal half with
denticles reducing in size distally; dactylus incurved at tip, cutting edge with
two obtuse teeth in proximal two-fifths, proximal tooth much larger than distal
one. Telson longer than wide, armed with 4-6 denticles on lateral margin, and
rounded on posterior margin with median spine. Uropodal endopod protruded
on serrated distolateral margin. [Adapted from Sakai, 1992a.]
Remarks. — The two female specimens from the Banda Sea, which were
identified as E. sibogae (De Man, 1925) (cf. Sakai, 1992a: 164, fig. 6),
have turned out to be different from E. sibogae, because the morphological
differences found between the female specimens and the holotype of E.
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275
sibogae are as follows. In the female specimens, the rostrum is smooth and
rounded at the apex, bearing 1-4 denticles on the median carina. In E. sibogae
the rostrum is subacute apically and smooth on the median carina. Therefore,
the females from the Banda Sea are distinguishable from the type specimens
of E. sibogae from the Sulu Sea, and are here renamed E. bandaensis sp. nov.
as a new species. In addition, Econaxius sibogae is transferred to the genus
Eiconaxiopsis by the presence of male Plp1 (see p. 295).
Type locality. — Banda Sea, 5◦ 47′ S 132◦ 51′ E, 348 m.
Distribution. — Only known from the type locality, Banda Sea.
Eiconaxius borradailei Bouvier, 1905
Eiconaxius Borradailei Bouvier, 1905: 803.
Iconaxius Borradailei — Balss, 1925: 209.
Axius (Eiconaxius) borradailei — Bouvier, 1925: 465, pl. 7 figs. 7, 8, pl. 9 fig. 4.
Axius (Eiconaxius) Borradailei — De Man, 1925d: 4 (list), 17 (key).
Eiconaxius borradailei — Sakai & De Saint Laurent, 1989: 21; Kensley, 1996e: 475 (list).
Type locality. — Caribbean Sea, 172-243 m.
Distribution. — 22◦ 09′ N 82◦ 21′ E (off the south of Cuba), Martinique,
Dominica, Grenada, and Barbados (Bouvier, 1905, 1925); 172-243 m.
Eiconaxius caribbaeus Faxon, 1896
Iconaxius caribbaeus Faxon, 1896: 155, pl. 1 figs. 1-4; Balss, 1925: 209.
Eiconaxius communis Bouvier, 1905: 803. [Type locality. — Antilles Islands and Gulf of
Mexico.]
Iconaxius communis — Balss, 1925: 210.
Axius (Eiconaxius) caribbaeus — Bouvier, 1925: 461, pl. 7 fig. 6; De Man, 1925d: 4 (list), 16
(key).
Axius (Eiconaxius) communis — De Man, 1925d: 4 (list), 17 (key).
Eiconaxius caribbaeus — Sakai & De Saint Laurent, 1989: 19; Kensley, 1996e: 475 (list).
Material examined. — RMNH D 41758, 1 female (TL/CL, 11.0/4.0 mm), Pillsbury Sta.
P-991, west Atlantic, N.E. of Anguilla, 18◦ 47′ N 64◦ 46.8′ W, 205-380 m, 5′ Black trawl,
23.vii.1970, USNM Acc. No. 381702.
Diagnosis. — Female Plp1 uniramous and slenderly elongate, consisting
of proximal segment and flagellum with setae; female Plps2-5 endopods with
appendix interna.
Type locality. — Caribbean Sea, 434 m.
Distribution. — Gulf of Mexico — Milligan Key (Bouvier, 1925), Milligan
Key and Havana (Sakai & De Saint Laurent, 1989); Guadeloupe (Bouvier,
1925; Sakai & De Saint Laurent, 1989); St. Lucia (Bouvier, 1925); St. Vincent
(Bouvier, 1925); Grenadines, Barbados (Bouvier, 1925; Sakai & De Saint
Laurent, 1989); Venezuela — Cariaco (Bouvier, 1925); 158-426 m.
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CRM 013 – Katsushi Sakai
Eiconaxius carinatus Bouvier, 1925
Axius (Eiconaxius) caribbaeus carinatus Bouvier, 1925: 465, pl. 9 fig. 3; Schmitt, 1935a: 191,
fig. 52.
Eiconaxius carinatus — Sakai & De Saint Laurent, 1989: 23.
Type locality. — Off St. Croix, Caribbean Sea, 158-536 m.
Distribution. — Milligan Key (Bouvier, 1925); Montana (Bouvier, 1925);
St. Croix (Bouvier, 1925); St. Vincent (Bouvier, 1925), 158-536 m.
Eiconaxius consobrinus (De Man, 1907)
(fig. 51)
Iconaxius (Iconaxiopsis?) consobrinus De Man, 1907: 129.
Axius (Eiconaxius) consobrinus — De Man, 1925d: 4 (list), 16 (key), 40, pl. 3 fig. 6-6d.
Eiconaxius consobrinus — Sakai & De Saint Laurent, 1989: 23 (partim); Kensley, 1996e: 475
(list).
Not: Iconaxiopsis consobrina — Balss, 1925: 209, 211, fig. 15 [= Eiconaxiopsis heinrichi sp.
nov.].
Material examined. — ZMA Crust. De. 102874, lectotype, 1 male (TL/CL, 24.0/8.5 mm,
carapace and abdomen separated), paralectotype, 1 ovig. female (TL/CL, 25.5/8.4 mm),
Indonesia between east point of Timor and Leti Island, 8◦ 17.4′ S 127◦ 30.7′ E, “Siboga” Sta.
280, 1224 m.
Diagnosis. — Rostrum elongate and three times as long as wide, lateral
margins armed with three obtuse denticles in male, or obscurely denticulate
in female; rostral median carina smooth, extending to anterior part of gastric
region. In larger cheliped fixed finger proximally notched (fig. 51A, B).
Abdominal somites smooth on surfaces (fig. 51C, D). Male Plp1 absent. Male
Plp2 biramous, endopod bearing appendices interna and masculina mesially at
proximal fourth.
Remarks. — The type specimens preserved in the Zoological Museum,
Amsterdam are a male and a female; the male is here designated as the
lectotype, and the ovigerous female as the paralectotype in order to avoid
confusion with the material of Iconaxiopsis consobrina in Balss (1925)
(= Eiconaxiopsis heinrichi sp. nov.).
Type locality. — Indonesia, between east point of Timor and Leti Island,
◦
8 17.4′ S 127◦ 30.7′ E, “Siboga” Sta. 280, 1224 m.
Distribution. — Indonesia — between east point of Timor and Leti Island
(De Man, 1925d); 1224 m.
Eiconaxius cristagalli (Faxon, 1893)
Axius crista-galli Faxon, 1893: 193; Faxon, 1895: 104, pl. 28 fig. 1-1h.
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Fig. 51. Eiconaxius consobrinus (De Man, 1907). A, larger cheliped in male, lateral view;
B, same in female; C, abdomen and tail-fan in male, lateral view; D, same in female. A, C,
ZMA Crust. De. 102874, lectotype, male (TL/CL, 24.0/8.5 mm), Indonesia, between east point
of Timor and Leti Island, 8◦ 17.4′ S 127◦ 30.7′ E, “Siboga” Sta. 280, 1224 m; B, D, ZMA Crust.
De. 102874, paralectotype, ovig. female (TL/CL, 25.5/8.4 mm), same data. Scale 1 mm.
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Axius (Eiconaxius) crista-galli — Borradaile, 1903: 538; De Man, 1925d: 4 (list), 14 (key).
Eiconaxius crista-galli — Rathbun, 1906: 895.
Iconaxius cristagalli — Balss, 1925: 210.
Eiconaxius cristagalli — Sakai & De Saint Laurent, 1989: 18; Hendrickx, 1995: 390 (list);
Kensley, 1996e: 475 (list), 481, fig. 7.
Remarks. — The present species, Eiconaxius cristagalli from south of
Panama, west coast of America, 837 m, is almost the same as E. asper from the
vicinity of Kauai Island, Hawaii, 418-528 m, except that in E. cristagalli the
dactylus of the larger cheliped is simply convex proximally on the cutting edge,
as shown by Kensley (1966e, fig. 7), whereas in E. asper it bears a prominent
tooth proximally on the cutting edge (Rathbun, 1906, fig. 52).
Type locality. — South off Panama, west coast of America, 837 m.
Distribution. — Panama (Faxon, 1893); 837 m.
Eiconaxius demani Sakai, 1992
Axius (Eiconaxius) parvus — De Man, 1925d: 42, pl. 3 fig. 7-7f [not: E. parvus Bate, 1888].
Eiconaxius demani Sakai, 1992a: 158, figs. 1, 2; Kensley, 1996e: 475 (list).
Type locality. — Kei Is., 5◦ 46′ S 132◦ 50′ E, 352 m.
Distribution. — Indonesia — Kepulauan Kai (De Man, 1925d; Sakai,
1992a); Arafura Sea (Sakai, 1992a); 352-560 m.
Eiconaxius farreae Ortmann, 1891
Eiconaxius farreae Ortmann, 1891: 49, pl. 1 fig. 4; Sakai & De Saint Laurent, 1989: 18; Sakai,
1992a: 162, fig. 5; Kensley, 1996e: 475 (list); Komai, 1999: 63; Sakai & Ohta, 2005: 70, figs.
1, 2.
“?Axius (Iconaxiopsis) farreae” — Borradaile, 1903: 537.
Axius (Eiconaxius) farreae — Balss, 1914: 88; De Man, 1925b: 125, fig. 3-3d; De Man, 1925d:
4 (list), 16 (key); Yokoya, 1933: 52; Miyake, 1982: 192 (list); Sakai, 1987a: 303 (list).
Iconaxiopsis farreae — Balss, 1925: 209, 211.
Axius farreae — Sakai, 1987a: 304, table 1.
Material examined. — Paralectotypes, MZS 313, 7 males (TL/CL: 19.0/5.7-12.0/3.8),
1 ovig. female (20.0/6.5), 2 females (21.0/6.5-17.0/5.9), 1 juv. (8.0/2.5). Lectotype, MZS 314
(1 specimen inside a sponge∗ ). Paralectotype, MZS 315 (1 specimen inside a sponge∗ ), Sagami
Bay, Japan; NSMT-Cr R547, fragments, Kannonzuka-dashi, Amadaiba, Sagami-Bay, Japan,
R/V “Hayama-Maru”, 400 m, 8.vi.1950, leg. His Majesty the Emperor of Japan, Hirohito et
al.; BLT 18245, 3 males (TL/CL, 19.0/6.4-2.4/7.9 mm), 1 ovig. female (TL/CL, 2.6/8.0 mm),
Sagami Bay, S. flank of Merase, 34◦ 50.06′ N 139◦ 43.10′ E – 34◦ 50.10′ N 139◦ 43.24′ E, 517528 m depth, R/V “Tansei-Maru”, Sta. MR-1-2, 13.v.2004, 08:03-08:12 h, 1 m, biological
dredge.
(∗ Farrea occa Bowerbank, 1862, Hexasterophora, det. C. Eckert.)
Type locality. — Sagami Bay, Japan, 182-364 m.
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Distribution. — Japan — Sagami Bay (Ortmann, 1891; De Man, 1925b;
Sakai, 1992a), Tokyo Bay and Sagami Bay (Balss, 1914), Omae-zaki and
Goto-retto (Yokoya, 1933); 77-732 m.
Eiconaxius faxoni sp. nov.
Axius acutifrons, Faxon, 1895: 103, pl. 28 fig. 2; Wicksten, 1982: 246 [not: Bate 1888].
Material considered. — Types, MCZ, 2 males and 4 ovig. females from R/V “Albatross”
Sta. 3358; 3 males, 3 females (2 ovig.) from Sta. 3359, southern Panama. [From Faxon, 1895.]
Remarks. — The type locality of E. acutifrons Bate, 1888 is the Banda Sea,
Indonesia, so that the specimens from southern Panama reported by Faxon
(1895) are geographically and morphologically different from those described
as E. acutifrons Bate, as shown in the Remarks on that species (p. 271).
Type locality. — South off Panama, 6◦ 30′ N 81◦ 44′ W to 6◦ 22.20′ N 81◦ 52′ W.
Distribution. — South of Panama; 850-1000 m (Faxon, 1895).
Etymology. — The new species is named Eiconaxius faxoni in honour
of Dr. W. Faxon, who described “Eiconaxius acutifrons” on the basis of
specimens from off southern Panama. The name thus is a noun in the genitive
singular.
Eiconaxius hakuhou Sakai & Ohta, 2005
Eiconaxius hakuhou Sakai & Ohta, 2005: 78, figs. 6, 7.
Type locality. — Sulu Sea; 1981-2019 m.
Distribution. — Only known from the type locality.
Eiconaxius indicus (De Man, 1907)
(fig. 52)
Iconaxius crista-galli var. indica De Man, 1907: 128.
Iconaxius cristagalli var. indica — Balss, 1925: 210.
Axius (Eiconaxius) crista-galli var. indica — De Man, 1925d: 4 (list), 15 (key), 31, pl. 2 fig. 33b.
Eiconaxius indica — Sakai & De Saint Laurent, 1989: 22.
Eiconaxius indicus — Kensley, 1996e: 475 (list); Sakai & Ohta, 2005: 73, figs. 3-5; Chu et al.,
2008.
Material examined. — ZMA Crust. De. 102465, holotype, 1 ovig. female (TL/CL, 32.0/10.6
mm) (fig. 52), off southeast coast of Great Kei Island, Indonesia, “Siboga” Sta. 267, 5◦ 54′ S
132◦ 56.7′ S, 984 m.
Type locality. — Off southeast coast of Great Kei, Indonesia, 5◦ 54′ S
132◦ 56.7′ E, “Siboga” Sta. 267, 984 m.
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CRM 013 – Katsushi Sakai
Fig. 52. Eiconaxius indicus (De Man, 1907). Whole body, lateral view. ZMA Crust. De. 102465,
holotype, 1 ovig. female (TL/CL, 32.0/10.6 mm), off south-east coast of Great-Kei island,
Indonesia, “Siboga” Sta. 267, 5◦ 54′ S 132◦ 56.7′ E, 984 m. Scale 1 mm.
Distribution. — Indonesia — Kepulauan Kai (De Man, 1907, 1925); 984 m;
Sulu Sea, Philippines, R/V “Hakuhou-Maru” Sta. 08◦ 06.36′ N 118◦ 26.49′ E –
08◦ 07.04′ N 118◦ 26.06′ E, 688-693 m; 08◦ 07.72′ N 118◦ 34.76′ E – 08◦ 08.51′ N
118◦ 34.44′ E, 1012-1015 m; 08◦ 06.41′ N 118◦ 29.18′ E – 08◦ 07.32′ N
118◦ 28.44′ E, 796-804 m; 08◦ 22.77′ N 118◦ 56.11′ E – 08◦ 24.31′ N 118◦ 55.61′ E,
1981-2019 m (Sakai & Ohta, 2005).
Eiconaxius indonesicus sp. nov.
Axius (Eiconaxius) acutifrons — De Man, 1925d: 4 (list), 15 (key), 37, pl. 3 fig. 5-5e.
Material. — Holotype, 1 male, south-east coast of Great Kei Island, “Siboga” Sta. 266,
5◦ 56.5′ S 132◦ 47.7′ E, 595 m, from a hexactinellid sponge; grey mud with coral and stones.
Diagnosis. — Rostrum stout and triangular, with apical tubercle, bearing
denticles on lateral margins. Eyestalks rounded. Gastric median carina unarmed. Mxp3 ischial crest crenulate. In male larger cheliped palm with distal
tooth on dorsal margin. Cutting edge of fixed finger with median cusp followed distally by denticulation toward tip. Ps3-5 short, subspatulate in shape,
and spinulate on ventral margin. Male Plp1 absent. Male Plp2 biramous and
foliaceous, endopod with appendices interna and masculina mesiomedially.
Telson rectangular, bearing median spine on posterior margin.
Remarks. — De Man (1925d: 37) described E. acutifrons on the basis of
specimens from the south of Great Kei Island, Indonesia, but his species is
clearly different from Bate’s E. acutifrons from the Banda Sea and Faxon’s
E. acutifrons from Panama, as descibed in the remarks of Bate’s acutifrons
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281
(p. 271). Therefore, De Man’s E. acutifrons is distinguished from Bate’s and
Faxon’s E. acutifrons, and here renamed Eiconaxius indonesicus sp. nov.
Type locality. — South-east coast of Great Kei Island, “Siboga” Sta. 266,
5◦ 56.5′ S 132◦ 47.7′ E, 595 m, grey mud with coral and stones.
Distribution. — Only known from the type locality.
Etymology. — The new species is named E. indonesicus, derived from the
type locality, Great Kei Island, Indonesia.
Eiconaxius kermadeci Bate, 1888
Eiconaxius kermadeci Bate, 1888: 43, pl. 5 fig. 3-3l; Balss, 1925: 210; Sakai & De Saint
Laurent, 1989: 16, fig. 5; Kensley, 1996e: 475 (list).
Axius (Eiconaxius) kermadeci — Borradaile, 1903: 538; De Man, 1925d: 4 (list), 16.
Iconaxius kermadeci — Balss, 1925: 210.
Type locality. — North of Kermadec Islands, 1080 m.
Distribution. — Kermadec Islands (Bate, 1888; Sakai & De Saint Laurent,
1989); 1080-1100 m.
Eiconaxius kimbla Kensley, 1996
(fig. 53)
Eiconaxius kimbla Kensley, 1996e: 481, fig. 8; Poore & Collins, 2009: 258.
Material examined. — AM P.25015, holotype, female (TL/CL, 16.0/4.8 mm); paratypes,
2 males (TL/CL, 13.0/4.9; 13.0/3.0 mm), 1 female (TL/CL, 16.0/5.5 mm), 14.5 km off Moreton
Bay, Queensland, Australia, 27◦ 06′ S 153◦ 37′ E, Sta. 6/6.9-1, 148 m, vi.1969, leg. HMAS
“Kimbla”, det. B. Kensley.
Diagnosis. — P1 chelate and unequal (fig. 53A, B). Male Plp1 absent; male
Plp2 with appendices interna and masculina; male Plps3-5 endopods with
appendix interna. Female Plp1 uniramous, elongate, and bisegmented; female
Plps2-5 endopods with appendix interna.
Remarks. — The present species is closely similar to Eiconaxiopsis sibogae
(De Man, 1925d) from the Sulu Sea, but they are different as follows. In
E. kimbla, the rostrum bears no apical spinule and the telson is subsquare,
whereas in Eiconaxiopsis sibogae the rostrum bears an apical spinule, and the
telson is longer than wide.
Type locality. — Fourteen-and-a-half (14.5) km off Moreton Bay, Australia,
27◦ 06′ S 153◦ 37′ E, 140-148 m.
Distribution. — Australia — off Moreton Bay (Kensley, 1996e); 140-148 m.
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CRM 013 – Katsushi Sakai
Fig. 53. Eiconaxius kimbla Kensley, 1996. A, larger cheliped on right side; B, smaller cheliped
on left side. A, B, AM P.25015, holotype, female (TL/CL, 16.0/4.8 mm), 14.5 km off Moreton
Bay, Queensland, Australia, 27◦ 06′ S 153◦ 37′ E, 148 m. Scale 1 mm.
Eiconaxius laccadivensis (Alcock & Anderson, 1894)
Eiconaxius kermadeci var. laccadivensis Alcock & Anderson, 1894: 162.
Iconaxiopsis laccadivensis — Alcock, 1901: 195; Alcock & MacGilchrist, 1905, pl. 71 fig. 3;
Balss, 1925: 209, 211.
Axius (Iconaxiopsis) laccadivensis — Borradaile, 1903: 537.
Axius (Eiconaxius) laccadivensis — De Man, 1925d: 4 (list), 16 (key).
Eiconaxius laccadivensis — Sakai & De Saint Laurent, 1989: 20; Kensley, 1996e: 475 (list).
Remarks. — The rostrum is microscopically serrated on the lateral margins;
the fixed finger of the larger cheliped bears on the cutting edge two enlarged
teeth separated by a characteristic notch (Alcock, 1901: 195).
Type locality. — Arabian Sea near Laccadive and Travancore, 1285 m
(705 fms).
Distribution. — Laccadive Islands (Alcock & Anderson, 1894); India —
Travancore coast (Alcock & Anderson, 1894); 1285 m.
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Eiconaxius mallacoota Poore & Collins, 2009
Eiconaxius mallacoota Poore & Collins, 2009: 258, fig. 24.
Diagnosis. — Rostrum 0.3 times length of front-to-posterior margin of carapace, concave dorsally, parallel-sided over eyes, then tapering to acute tip, with
10 marginal lateral tubercles on oblique margins, depressed below level of
median carina, anteriorly directed, continuous with lateral carinae. Supraocular spines absent. Eyestalk 0.5 length of rostrum; cornea unpigmented. A1
peduncle reaching to end of A2 segment 4; segment 1 unarmed. A2 segment
1 unarmed; segment 2 stylocerite as vertical blade, reaching to midpoint of
segment 5; scaphocerite as vertical blade, overreaching end of segment 5; segment 3 with mesial tooth on ventral margin; segment 4 as long as segment 2;
segment 5 about half length of segment 4. Carapace smooth; gastric lateral
carina unarmed; gastric submedian carina smooth, together semicircular and
converging on median carina; gastric median carina obsolete, only at base of
rostrum. Mxp3 coxa-ischium unarmed; crista dentata with about 15 similar
teeth; merus and carpus unarmed.
P1 asymmetrical and robust. In larger cheliped coxa with spinule; basis
unarmed; ischium with few irregular teeth on ventral margin; merus with two
small teeth on strongly convex dorsal margin, and 6 small teeth on ventral
margin; carpus with distal tooth on ventral margin; propodus with distal tooth
on dorsal margin and 5 small teeth on lateral submarginal ridge ventrally;
greatest depth equal to length of dorsal margin; fixed finger 0.7 times as long as
dorsal palm, cutting edge with irregular teeth in shallow proximal concavity,
and irregular small teeth distally; dactylus curved distally, and cutting edge
smooth. Smaller cheliped less robust; coxa with spinule; basis unarmed;
ischium with few irregular teeth on ventral margin; merus with two small
teeth on strongly convex dorsal margin, and 5 small teeth on ventral margin;
carpus with distal tooth on ventral margin; propodus with distal tooth on dorsal
margin, and 4 small teeth on lateral submarginal ridge ventrally; greatest depth
1.1 times length of dorsal margin; fixed finger 1.2 times as long as dorsal
palm, bearing lateral ridge along proximal half of cutting edge, cutting edge
armed proximally with irregular teeth; dactylus tapering, cutting edge smooth.
P2 ischium and merus unarmed on ventral margin; carpus slightly shorter
than chela; propodus dorsal margin three times as long as dactylus. P3 merus
unarmed; propodus 2.5 times as long as dactylus, bearing 6 rows of 1-2 robust
setae; dactylus spatulate, bearing 13 robust setae along oblique margin, plus
unguis. P4 identical with P3; propodus 2.5 times as long as dactylus, bearing
6 rows of 1-3 robust setae; dactylus spatulate, bearing 11 robust setae along
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CRM 013 – Katsushi Sakai
oblique margin, plus unguis. P5 propodus 2.8 times as long as dactylus, simple,
densely setose distally, bearing no robust setae; dactylus spatulate, bearing 8
robust setae along oblique margin, plus unguis.
Abdominal somite 1 pleuron rounded-truncate ventrally; pleuron 2 obliquely
angled, acutely produced posteroventrally; pleura 3-4 acutely produced posteroventrally; pleuron 5 less so; all with anteroventral tooth; pleuron 6 subacute; abdominal somite 6 with pairs of lateral teeth dorsally on posterior margin at base of telson, and 3 (uneven) denticles dorsally along dorsal posterior
margin. Plp2-5 each with appendix interna, and 0.3 length of endopod. Male
Plp2 appendix masculina 0.7 times as long as appendix interna. Telson 1.5
times as long as wide, widest proximally, then tapering distally, bearing 810 small lateral teeth and weak posterolateral teeth, but no median spine on
posterior margin; dorsal surface with obsolete longitudinal ridges. Uropodal
endopod 2.4 times as long as wide and oval, bearing 12 small irregular lateral
teeth, last tooth distal, but no longitudinal ridge. Uropodal exopod 1.6 times
as long as wide, oval, bearing 14-20 small irregular lateral teeth over distal
two-thirds, last tooth distal, but no longitudinal rib. [Adapted from Poore &
Collins, 2009.]
Type locality. — South of Point Hicks, Victoria, Australia, 930-951 m.
Distribution. — Off eastern Victoria, Australia; 930-1000 m.
Eiconaxius mortenseni Sakai, 1992
Eiconaxius mortenseni Sakai, 1992a: 160, figs. 3, 4; Kensley, 1996e: 475 (list).
Type locality. — Misaki, Sagami Bay, Japan, 366 m.
Distribution. — Japan — Sagami Bay (Sakai, 1992a); 366-732 m.
Eiconaxius parvus Bate, 1888
Eiconaxius parvus Bate, 1888: 44, pl. 5 figs. 4, 5; Sakai & De Saint Laurent, 1989: 18; Kensley,
1996e: 475 (list).
Axius (Eiconaxius) parvus — Borradaile, 1903: 538; De Man, 1925d: 4 (list), 16.
Iconaxius parvus — Balss, 1925: 210.
Not: Axius (Eiconaxius) parvus — De Man, 1925d: 42, pl. 3 fig. 7-7f [= Eiconaxius demani
Sakai, 1992a].
Remarks. — Bate (1888: 44, fig. 4) described from his specimen that the
rostrum is wanting, however, his figure shows that the rostrum is unarmed.
The female with 7 ova (not 7 ovig. females as in Sakai & De Saint Laurent,
1989: 18) was designated as the lectotype of the present species.
Type locality. — Off Kermadec Islands, 952 m, hard ground.
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285
Distribution. — Kermadec Islands (Bate, 1888; Sakai & De Saint Laurent,
1989); 520-950 m.
Eiconaxius rotundifrons Bouvier, 1905
Eiconaxius rotundifrons Bouvier, 1905: 803; Sakai & De Saint Laurent, 1989: 21, fig. 6.
Iconaxius rotundifrons — Balss, 1925: 210.
Axius (Eiconaxius) caribbaeus rotundifrons — Bouvier, 1925: 463, figs. 23-25, pl. 10 figs. 3, 4.
Axius (Eiconaxius) rotundifrons — De Man, 1925d: 4 (list), 17 (key).
Material examined. — RMNH D 41759, 1 male (TL/CL, 15.0/4.9 mm), 2 females (TL/CL,
18.0/6.5-21.0/6.4 mm), Pillsbury Sta. P-944, west Atlantic, N. of Guadeloupe, 16◦ 33.2′ N
61◦ 36.8′ W – 16◦ 34.4′ N 61◦ 37.2′ W, 360-421 m, 5′ Black trawl, 17.vii.1969, USNM Acc.
No. 381702.
Diagnosis. — Male Plp1 absent; male Plp2 endopod with appendices interna and masculina; male Plps3-5 endopods with appendix interna. Female
Plp1 uniramous and bisegmented, consisting of proximal segment and flagellum with setae; female Plps2-5 endopods with appendix interna.
Type locality. — West Indies to Gulf of Mexico.
Distribution. — Gulf of Mexico (Bouvier, 1905); Martinique (Bouvier,
1925); St. Lucia (Bouvier, 1925; Sakai & De Saint Laurent, 1989); Barbados
(Bouvier, 1925; Sakai & De Saint Laurent, 1989); Venezuela — Cariaco
(Bouvier, 1925; Sakai & De Saint Laurent, 1989); 277-2506 m.
Eiconaxius singularis Zarenkov, 1983
Axius (Eiconaxius) singularis Zarenkov, 1983: 83, fig. 2.
Eiconaxius singularis — Sakai & De Saint Laurent, 1989: 23; Kensley, 1996e: 475 (list).
Type locality. — Around submarine mountains Marcus-Necker near Hawaii,
100-1350 m.
Distribution. — Submarine mountains Marcus-Necker near Hawaii (Zarenkov, 1983); 100-1350 m.
Eiconaxius spiniger (MacGilchrist, 1905)
Iconaxiopsis spiniger MacGilchrist, 1905: 240; Alcock et al., 1907, pl. 78 fig. 1; Balss, 1925:
209.
Axius (Eiconaxius) spiniger — De Man, 1925d: 4 (list), 15 (key).
Eiconaxius spiniger — Sakai & De Saint Laurent, 1989: 22; Kensley, 1996e: 475 (list).
Type locality. — Bay of Bengal, “Investigator” Sta. 310, 1755 m.
Distribution. — Bay of Bengal (MacGilchrist, 1905); 1755 m.
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CRM 013 – Katsushi Sakai
Eiconaxius weberi (De Man, 1907)
(fig. 54)
Iconaxius Weberi De Man, 1907: 127; Balss, 1925: 210.
Axius (Eiconaxius) Weberi — De Man, 1925d: 5 (list), 17 (key), 44, pl. 3 fig. 8-8a, pl. 4 fig. 8b-o.
Eiconaxius weberi — Sakai & De Saint Laurent, 1989: 22; Sakai, 1992a: 163; Kensley, 1996e:
475 (list).
Material examined. — ZMA Crust. De. 102464, lectotype, 1 male (TL/CL, 16.5/5.49), leg.
“Siboga” Sta. 267, 5◦ 54′ S 132◦ 56.7′ E, 984 m; paralectotypes, 1 male (TL/CL, 17.0/5.6 mm),
2 ovig. females (TL/CL, 21.5/6.6; 21.0/7.1 mm), 3 females (TL/CL, 10.5/3.1-14.0/4.6 mm),
leg. “Siboga” Sta. 266, 5◦ 56′ S 132◦ 47.7′ E, 595 m; RMNH D 1172, 1 male (TL/CL, 18.0/6.0
mm), 1 female (TL/CL, 12.0/4.2 mm), 5◦ 56′ S 132◦ 47.7′ E, “Siboga” Sta. 266, vii.1929.
Diagnosis of male lectotype. — Rostrum stoutly triangular with obtuse
apex, lateral margins armed with 8-9 subacute denticles, extending posteriorly
onto anterior third of gastric region as lateral carinae. P1 asymmetrical in
shape, but almost same in size; in larger cheliped (fig. 54A) ischium with
two spines and denticles distally on ventral margin, and unarmed on dorsal
margin; merus unarmed on dorsal margin and serrated on ventral margin;
carpus divergent distally and twice as wide as long, bearing subdistal tooth on
ventral margin; chela length twice its width, palm 1.2 times as long as wide,
bearing one distal and 0-2 subdistal denticles on dorsal margin, and carinate
on ventral margin; fixed finger armed with 5 irregular teeth on proximal
half on cutting edge, and medially a distinct, conical tooth followed distally
by denticles (fig. 54A); dactylus furrowed along dorsal margin on lateral
surface, cutting edge armed with distinct tooth at proximal third, but smooth
in distal two-thirds. Abdominal pleuron 1 small, and rounded on distal margin
(fig. 54C); pleuron 2 broadened and protruded posteroventrally with acute
angle; pleuron 3 protruded posteroventrally with almost right angle, bearing
denticle anteriorly on ventral margin; pleuron 4 protruded posteroventrally
with right angle, bearing denticle anteriorly on ventral margin; pleuron 5
rounded, bearing denticle anteriorly on ventral margin. Male Plp1 absent;
male Plp2 biramous, endopod bearing appendices interna and masculina
mesiomedially; male Plps3-5 endopods with appendix interna. Female Plp1
slender and bisegmented, bearing setae distally; female Plps2-5 biramous,
endopods with appendix interna.
Female P1. — In smaller female, P1 (fig. 54B) ischium with two distal
spines on ventral margin and two closely-located distal denticles on dorsal
margin; merus with two distal spines on dorsal margin, and 4 distal denticles
on ventral margin; carpus twice as wide as long, bearing tooth on ventral
margin; chela 2.5 times as long as wide, palm 1.3 times as long as wide,
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Fig. 54. Eiconaxius weberi De Man, 1907. A, larger cheliped in male; B, smaller cheliped in
female; C, abdomen and tail-fan, lateral view. A, ZMA Crust. De. 102464, paralectotype, ovig.
female (TL/CL, 21.0/7.1 mm); B, ZMA Crust. De. 102464, female (TL/CL, 10.5/3.1 mm), leg.
“Siboga” Sta. 266, 5◦ 56′ S 132◦ 47.7′ E, 595 m; C, ZMA Crust. De. 102464, lectotype, male
(TL/CL, 16.5/5.49), leg. “Siboga” Sta. 267, 5◦ 54′ S 132◦ 56.7′ E, 984 m. Scales 1 mm.
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CRM 013 – Katsushi Sakai
bearing three distal teeth on dorsal margin, and carinate on ventral margin;
fixed finger denticulate on cutting edge, bearing distinct, obtuse conical tooth
at proximal two-fifths; dactylus bearing two teeth proximally on cutting edge,
proximal one small and triangular, distal one distinct and broad.
Remarks. — De Man (1925a: 44) described E. Weberi on the basis of 5 specimens (two males and three females) of the type series. In the present revision, all the specimens composing the type series were examined, including
the 5 specimens that De Man described: two males, two ovigerous females,
and 4 non-ovigerous females deposited in the Zoological Museum Amsterdam (ZMA Crust. De. 102464), and another male separately deposited in the
National Museum of Natural History, Naturalis, Leiden (RMNH D 1172), of
which the male from “Siboga” Sta. 267 is designated the lectotype, and the
others as the paralectotypes.
Type locality. — Off south-west coasts of Great Kei Island, Indonesia,
5◦ 54′ S 132◦ 56.7′ E, 984 m.
Distribution. — Indonesia — Kepulauan Kai (De Man, 1907, 1925a), Java
Sea (Sakai, 1992a); 200-984 m.
FAMILY EICONAXIOPSIDIDAE FAM. NOV.
Diagnosis. — Rostrum stout, conspicuous, and pointed apically; dorsal
surface concave, lateral margins extending posteriorly onto gastric region as
lateral carinae up to cervical groove. Gastric region bearing median carina
gradually broadening posteriorly, but no submedian carinae; cervical groove
usually indistinct; no linea thalassinica; posterior margin of carapace with
lateral lobes. Eyestalks indiscernible in lateral view; eyes rounded or ovate,
cornea pigmented or unpigmented. A1 segment 1 as long as segments 2-3 combined. Scaphocerite strong. Maxilla 2 scaphognathite without posterior whip.
Mxp3 ischial crest crenulate, exopod present. Ps1-2 chelate. Ps3-5 dactyli
short, tapering, and pointed distally. Abdominal somite 1 with pleuron slightly
protruded posteroventrally; abdominal somites without carinae between terga
and pleura. Male Plp1 uniramous and bisegmented; male Plp2 biramous and
slender, endopod with appendices interna and masculina mesially at proximal
third; Plps3-5 biramous, endopods with appendix interna. Female Plp1 uniramous and bisegmented; distal segment and flagellum with setae; Plps2-5 slender and biramous, endopods with appendix interna. Telson longer than wide,
bearing no proximal lobe laterally. Uropodal endopod oval. Uropodal exopod
without transverse suture.
Remarks. — The new genus Eiconaxiopsis gen. nov. is closely similar to
the type genus of the family Eiconaxiidae Sakai & Ohta, 2005, Eiconaxius
in the shapes of the rostrum, A2 scaphocerite, abdominal somites, and tailfan, but they differ from each other as follows. In Eiconaxiopsis gen. nov.
the gastric region of the carapace bears a median carina gradually broadening
posteriorly, the Ps3-5 dactyli are simple, tapering distally, and pointed at the
tip, the male Plp1 is present as a bisegmented appendage, and the telson lacks
a median spine on the posterior margin. In Eiconaxius, however, the gastric
region of the carapace bears a linear median carina with or without denticles,
which is simple or bifurcate posteriorly, the Ps3-5 dactyli are subspatulate and
are spinulose on the ventral margin, the male Plp1 is absent, and the telson
bears a median spine on the posterior margin. For those reasons, a new family
Eiconaxiopsididae fam. nov. is established for Eiconaxiopsis gen. nov.
Type genus. — Eiconaxiopsis gen. nov., by being the nominate genus.
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CRM 013 – Katsushi Sakai
Genus included. — Eiconaxiopsis gen. nov.
Etymology. — The new family is named Eiconaxiopsididae by adding the
Latin suffix “idae”, to the stem of the generic name Eiconaxiopsis, as indicated
in the ICZN (2000).
Genus Eiconaxiopsis gen. nov.
Diagnosis. — Rostrum triangular and furrowed dorsally, lateral margins
extending posteriorly onto gastric region as lateral carinae. Gastric region
convex, bearing median carina gradually broadening posteriorly, but no submedian carinae. Cervical groove indistinct. Eyestalks indiscernible, eyes ovate,
cornea unpigmented. A2 peduncular segment 2 with distinct distodorsal spine;
scaphocerite conspicuously developed. P1 subequal. P2 chelate. Ps3-5 dactyli
short and simple. No longitudinal carinae between abdominal pleura and terga.
Pleurobranchs present on Ps2-4. Male Plp1 uniramous and bisegmented; male
Plp2 biramous, endopod bearing appendices interna and masculina mesially at
proximal third; male Plps3-5 endopods with appendix interna. Telson longer
than wide and denticulate on lateral margins, bearing no median spine on posterior margin. Uropodal exopod without transverse suture. Deep-water habitat.
Remarks. — De Man (1907) listed a male (now lectotype) from Timor
as Iconaxius (Iconaxiopsis?) consobrinus De Man, 1907, and later listed that
same male (now lectotype) and a female (now paralectotype) from Timor as
Axius (Eiconaxius) consobrinus (cf. De Man, 1925d). Balss (1925) recorded a
male from Nias as Iconaxiopsis consobrina De Man, 1925d. It has turned out,
however, through the present careful examination of De Man’s male specimen
from Timor (ZMA Crust. De. 102874, lectotype) and Balss’s male specimen
from Nias (ZMB 19345, now holotype of E. heinrichi) that the former is
distinctly different from the latter. In De Man’s specimen, the Ps3-4 dactyli are
short, subspatulate in form, and spinulose on the ventral margin, the male Plp1
is absent (De Man, 1925d: 42) as in the other species in the genus Eiconaxius,
whereas in Balss’s specimen the Ps3-4 dactyli are simple, tapering, and pointed
distally, the male Plp1 is present as a uniramous and bisegmented appendage,
and the distal segment is elongate and slightly convex on its mesial margin
with setae (fig. 57C), which suggests that Balss’s Iconaxiopsis consobrina
is a misidentified species and different from De Man’s Axius (Eiconaxius)
consobrinus, so that Balss’s male from Nias is to be distinguished from
De Man’s male from Timor at the generic level. Accordingly, a new genus
Eiconaxiopsis gen. nov. is established based on Eiconaxius consobrinus (cf.
Balss, 1925: 209), and Balss’s species is renamed Eiconaxiopsis heinrichi sp.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
291
nov., distinguished from Iconaxius (Iconaxiopsis?) consobrinus De Man, 1907
from Timor, which is named Eiconaxius consobrinus (De Man, 1907).
Type species. — Eiconaxiopsis heinrichi sp. nov., by present designation.
The gender of the new name, Eiconaxiopsis, is feminine.
Species included. — Eiconaxiopsis heinrichi sp. nov.; E. sibogae (De Man,
1925c).
Etymology. — The new generic name Eiconaxiopsis is composed of a
generic name, Eiconaxius and a Latin suffix “-opsis”, meaning “similar”.
Eiconaxiopsis heinrichi sp. nov.
(figs. 55-57)
Iconaxiopsis consobrina — Balss, 1925: 209, 211, fig. 15. [Not: Iconaxius (Iconaxiopsis?)
consobrinus De Man, 1907.]
Eiconaxius consobrinus — Sakai & De Saint Laurent, 1989: 23 (partim).
Material examined. — ZMB 19345, holotype, male (TL/CL, 24.0/9.0 mm, eyestalks of right
side missing), South Canal, Nias, west coast of Sumatra, Deutsche Tiefsee Exped., 2.ii.1899,
St. 198, 677 m, trawl, det. Balss as Iconaxiopsis consobrinus De Man, 1907.
Diagnosis. — Carapace (figs. 55A, 56A) smooth. Rostrum lanceolate in
dorsal view and convex medially, with distinct smooth median carina originating from anterior third of rostrum, increasing in breadth, and extending
Fig. 55. Eiconaxiopsis heinrichi sp. nov. A, whole body, lateral view; B, telson and uropod on
right side. A-B, ZMB 19345, holotype, 1 male (TL/CL, 24.0/9.0 mm, eyestalks of right side
missing), Nias, west coast of Sumatra, Deutsche Tiefsee Expedition. Scales 1 mm.
292
CRM 013 – Katsushi Sakai
Fig. 56. Eiconaxiopsis heinrichi sp. nov. A, carapace dorsal view; B, anterior part of carapace;
C, anterior part of carapace, lateral view; D, P3, lateral view. A-D, ZMB 19345, holotype,
1 male (TL/CL, 24.0/9.0 mm, eyestalk of right side missing), Nias, west coast of Sumatra,
Deutsche Tiefsee Expedition. Scales 1 mm.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
293
Fig. 57. Eiconaxiopsis heinrichi sp. nov. A, P1 lateral view; B, same, mesial view; C, male Plp1;
D, male Plp2. A-D, ZMB 19345, holotype, male (TL/CL, 24.0/9.0 mm, eyestalk of right side
missing), Nias, west coast of Sumatra, Deutsche Tiefsee Expedition. Scales 1 mm.
posteriorly along dorsal line in gastric region; rostral tip turned up and subacute apically, not reaching distal end of A1 proximal segment; lateral margins
indistinctly dentate, extending posteriorly onto anterolateral part of gastric region as lateral carinae; submedian carinae absent (fig. 56A, B, C). Cervical
groove indistinct. Eyestalks indiscernible, eyes ovate, cornea unpigmented. A1
proximal segment subequal in length to segments 2-3 combined. A2 segment
1 short and unarmed, and segment 2 elongate. Scaphocerite extending forward
beyond dorsodistal spine of A2 segment 2; segment 5 twice length of segment 4. A2 flagellum about 1.3 times as long as A1 flagella. Mxp3 coxa, basis,
and ischium each with sharp mesiodistal tooth; ischial crest denticulate; merus
to dactylus unarmed; exopod consisting of proximal segment and flagellum,
reaching distal margin of merus.
P1 subequal and asymmetrical in shape. In larger cheliped (fig. 57A),
ischium with denticles on dorsal margin; merus about 1.2 times as long as
wide, and armed with distal tooth on ventral margin; carpus wider than long,
294
CRM 013 – Katsushi Sakai
two-thirds length of merus on dorsal margin, and armed with obtuse, low
tooth subterminally on ventral margin; chela wide and slightly less than twice
as long as wide; palm 1.3 times as long as wide, distal margin convex and
denticulate; cutting edge of fixed finger smooth in proximal half, bearing
medially a convexity followed posteriorly by denticles diminishing in size in
distal half; dactylus carinate along dorsal margin on lateral surface, cutting
edge armed with low obtuse double tooth in proximal half, and then distinctly
concave, smooth, and incurved distally. In smaller cheliped (fig. 57B) coxa and
basis unarmed, ischium armed with distal tooth on ventral margin; merus 1.8
times as long as wide, bearing distal tooth on ventral margin; carpus two-thirds
length of merus on dorsal margin; palm 2.8 times as long as carpus and 1.5
times as long as wide, convex and roughly denticulate on distal margin; fixed
finger double-edged on cutting edge, lateral edge armed with low, irregularlyshaped denticles, and mesial edge with sharp teeth; dactylus two-thirds length
of palm, and smooth and incurved distally on cutting edge. P2 chelate and
unarmed. P3 (fig. 56D) simple, propodus 1.2 times length of carpus, bearing
transverse rows of spinules ventrally; dactylus tapering and pointed distally.
P4 simple, propodus 1.5 times length of carpus, bearing rows of transverse
spinules ventrally. P5 simple, propodus twice length of carpus, bearing a row
of setae at ventrodistal angle.
Abdominal somites (fig. 55A) smooth, abdominal somite 1 short, somites 26 subequal dorsally; pleuron 1 narrow; pleuron 2 large and protruded posteroventrally to form subacute angle; pleura 3-5 subacute at posteroventral angle; pleuron 6 triangular ventrally and smooth on posterior margin.
Male Plp1 uniramous and bisegmented, distal segment with setae distally
(fig. 57C); male Plp2 biramous, endopod with appendices interna and masculina (fig. 57D); male Plps3-5 biramous, endopods with appendix interna.
Telson (fig. 55B) longer than wide, and much longer than somite 6; lateral
margins nearly parallel and denticulate; posterior margin rounded without median spine. Uropodal endopod serrate on lateral margin, and armed with two
distinct teeth at distolateral angle. Uropodal exopod largely convex and serrated on lateral margin to lateral half of distal margin.
Remarks. — The present new species, Eiconaxiopsis heinrichi sp. nov.
is established on the male specimen from Nias, an island off the western
coast of Sumatra, Indonesia, 677 m, which was identified by Balss (1925) as
Iconaxiopsis consobrina De Man, 1907. The present review shows that Balss’s
male specimen of “I. consobrina” from Sumatra has turned out to be clearly
different from De Man’s male specimen of Iconaxius consobrinus from Timor,
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
295
“Siboga Sta. 280”, 1224 m, in that the rostrum is lanceolate, and male Plp1
is present. Those differences between the two specimens mean that Balss’s
specimen is a new species and it is here renamed as Eiconaxiopsis heinrichi
sp. nov.
Type locality. — Nias, west coast of Sumatra, 677 m.
Distribution. — Only known from the type locality.
Etymology. — The new species is named Eiconaxiopsis heinrichi sp. nov.
in honour of Dr. Heinrich Balss, who described the holotype specimen with
figures of the larger cheliped in 1925. The name is a noun in the genitive
singular.
Eiconaxiopsis sibogae (De Man, 1925)
(fig. 58)
Axius (Eiconaxius) Sibogae De Man, 1925c: 218; De Man, 1925d: 4 (list), 15 (key), 34, pl. 2
fig. 4-4l.
Axius (Eiconaxius) sibogue — Estampador, 1937: 500 (misspelling).
Eiconaxius sibogae — Sakai & De Saint Laurent, 1989: 23; Sakai, 1992a: 163, fig. 6; Kensley,
1996e: 475 (list).
Eiconaxius sibogae — Sakai, 1992a: 163, fig. 6.
Material examined. — ZMA Crust. De. 102525, lectotype, male (TL/CL, 13.0/4.6 mm), Sulu
Sea, “Siboga” Sta. 95, 5◦ 43.5′ N 119◦ 40′ E, 522 m; paralectotype, female (TL/CL, 14.0/4.8 mm,
with left small cheliped, larger cheliped missing); Sulu Sea, “Siboga” Sta. 95, 5◦ 43.5′ N
119◦ 40′ E, 522 m.
Diagnosis. — Rostrum triangular and obtuse at tip with (male) or without
(female) apical spinule (fig. 58B), bearing smooth median carina extending
posteriorly onto gastric region as gastric median carina with bifurcate posterior end. A2 scaphocerite slightly overreaching A2 penultimate segment. P1
chelate and unequal. Abdominal somites smooth laterally, somites 3-5 each
with anteroventral spine (fig. 58A). Male Plp1 uniramous, slender, and trisegmented; male Plp2 endopod with appendices interna and masculina. Telson (fig. 58C) longer than wide, bearing 3-4 denticles on lateral margins, and
movable spine at posterolateral angle; convex posterior margin with (male) or
without (female) median spine.
Type locality. — Sulu Sea, 5◦ 43.5′ N 119◦ 40′ E, “Siboga” Sta. 95, 522 m.
Distribution. — Sulu Sea (De Man, 1925d; Estampador, 1937); 348-522 m.
296
CRM 013 – Katsushi Sakai
Fig. 58. Eiconaxius sibogae De Man, 1925. A, whole body, lateral view; B, anterior part of
carapace, dorsal view; C, telson and uropod on right side. A-C, ZMA Crust. De. 102525, female
(TL/CL, 14.0/4.8 mm, with left small cheliped, larger cheliped missing), Sulu Sea, “Siboga”
Sta. 95, 5◦ 43.5′ N 119◦ 40′ E, 522 m. Scales 1 mm.
FAMILY METICONAXIIDAE SAKAI, 1992
Meticonaxiinae Sakai, 1992c: 19.
Diagnosis. — Rostrum triangular; dorsal surface flat with median carina,
and lateral margins extending posteriorly onto gastric region as lateral carinae. Carapace compressed, bearing posterolateral lobes; linea thalassinica absent; cervical groove distinct. Eyestalks deformed. Maxilla 2 scaphognathite
with 1-2 posterior whips. P3 propodus rounded. P4 propodus oblong, P4 coxa
flattened. P5 subchelate. Abdominal somite 1 with anterolateral lobe. Male
Plp1 uniramous and bisegmented, distal segment cleaver-shaped; male Plp2
biramous, endopod with appendices interna and masculina; male Plps3-5 biramous, endopods with appendix interna. Pleopodal filament absent. Uropodal
exopod bearing neither distal flap nor lateral notch. Setal rows present on carapace, abdomen, and pereiopods.
Remarks. — Kensley & Heard (1991: 534, fig. 25) showed by the cladograms resulting from PAUP analysis of 22 characters and 11 taxa, that Michelea is distinguished from Marcusiaxius and Meticonaxius. Sakai (1992c) established a subfamily Meticonaxiinae for the genera Marcusiaxius and Meticonaxius, and another subfamily Micheleinae for the genus Michelea. Poore
(1997: 354), however, included the subfamilies Meticonaxiinae Sakai, 1992c
and Micheleinae Sakai, 1992c (not Poore, 1994) in the family Micheleidae Sakai, 1992c, because the phylogenetic analysis suggests that the Meticonaxiinae are polyphyletic, and must therefore be synonymized with the
Micheleinae. Poore (1997: 356) also mentioned that “there are so many
synapomorphies linking the genera that this division cannot be sustained. Further, the cladistic analysis (Poore, 1994) hypothesized that Michelea is a terminal taxon and not a sister group to other genera in the family”. However, such
characteristics as “length of Mxp2 exopod; setal rows on abdominal somite
6; Mxp3 ischial crest dentate; and Mxp3 meral spine” used for his phylogenetic analysis (Poore, 1994: 109) are of no significance in comparing the
genera Marcusiaxius, Meticonaxius, and Michelea. In the present revision,
the characteristics of rostrum, pleopods, Ps3-4 propodi, pleurobranchs, telson,
and uropodal endopod are, instead, applied in defining the family Meticonaxiidae for the genera Marcusiaxius, Meticonaxius, and Tethisea, and the family
Micheleidae for the genus Michelea.
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CRM 013 – Katsushi Sakai
Michelea, the type genus of the family Micheleidae, is clearly different
from Meticonaxius, the type genus of the family Meticonaxiidae, i.e., in
Michelea vandoverae (Gore, 1987), type species of Michelea, the rostrum
is short and bluntly triangular; the Ps3-4 propodi are rectangular, bearing
transverse rows of spinules, P5 is not chelate but simple, pleopodal filaments
are present, and pleurobranchs are absent. Whereas in Meticonaxius monodon
De Man, 1905, type species of Meticonaxius, the rostrum is conspicuous and
acutely triangular, with lateral margins extending posteriorly, the P3 propodus
is rounded, bearing no transverse rows of spinules, and the P4 propodus
is broadened without transverse rows of spinules, P5 is chelate, pleopodal
filaments are absent, and pleurobranchs are present above Ps2-4. As mentioned
above, the two families Meticonaxiidae and Micheleidae are clearly different
from each other.
Poore (1994: 109, table 1) showed character transformations used in the
phylogenetic analysis of 22 taxa of the Thalassinidea, and this genus-character
matrix (27 genera by 93 characters) was used in a Hennig 86 cladistic analysis
of the genera of the Thalassinidea (his table 2; see table VI herein), showing
(0) for Tethisea, Marcusiaxius, Meticonaxius, and Michelea. However, the
number of the matrix does not always show correctly the characteristics of the
genera, because in the case of No. 43, Maxilliped 2 exopod: almost reaching
end of merus (0); reduced or absent (1), in the type species of Michelea,
M. vandoverae (Gore, 1987) (cf. Kensley & Heard, 1991, fig. 21F) it is distinct
(0), but in M. pillsburyi Kensley & Heard, 1991 (cf. Kensley & Heard, 1991,
fig. 19J), it is reduced (1), which suggests that the character is not valid as a
generic one, because those two species, belonging to the same genus, do not
have the character in common, which means that those matrix numbers are
unreliable. In the case of No. 47, Mxp3 meral spine: present (0); absent (1),
the matrix of Michelea is shown as (1) in the table, meaning that the Mxp3
meral spine is absent, but in M. vandoverae the meral spine is really present
(Kensley & Heard, 1991, fig. 21G), so that it should be shown as (0). As a
result, the genus-character matrix has not been composed completely correctly.
Type genus. — Meticonaxius De Man, 1905, as the nominate genus.
Genera included. — Marcusiaxius Rodrigues & De Carvalho, 1972; Meticonaxius De Man, 1905; Tethisea Poore, 1994.
K EY
TO THE
G ENERA
OF THE
FAMILY M ETICONAXIIDAE
1 – Rostrum acutely triangular and flat, with median carina . . . . . . . . . . . . . . . . Meticonaxius
– Rostrum triangular with obtuse apex, bearing middorsally a raised area with flat surface
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Tethisea
(type sp.:
T. indica)
Marcusiaxius
(type sp.:
M. lemoscastroi)
Length of Mxp2
exopod: almost
reaching to end
of merus (0);
reduced or
absent (1)
(0), extending to
end of merus
(Poore, 1997,
fig. 26B)
Mxp3 ischial
crista dentata:
prominent
toothed ridge
(0); obsolete or
absent (1)
(0), present
(Poore, 1997,
fig. 26C)
(1), in M.
(1), absent
lemoscastroi
(Kensley et al.,
(cf. Kensley et
1991: 507)
al., 1991, fig. 9G);
(0) distinct in
M. colpos, overreaching carpus
(Kensley et al.,
1991, fig. 7I)
Mxp3 meral
spine:
present (0);
absent (1)
Male Plp1
Male Plp2
(1), absent
(Poore, 1997,
fig. 26C)
composed of stemlike proximal segment and subovate distal segment bearing
mesial lobe with
patch of hooklets
exopod ovate and endopod with appendix inelongate, leaf-like, with ap- terna
pendices interna and masculina
(1), absent
composed of stemlike proximal segment and cleavershaped distal segment
exopod broad and earshaped; endopod broadly
triangular, with appendices interna and masculina
Male Plps3-5
with appendix interna
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
TABLE VI
Genus-character matrix as used by Poore (1994), and generic differences on male Plps1-5
299
300
TABLE VI
(Continued; *) in present classification transferred to family Micheleidae)
Michelea*)
(type sp.:
Callianidea
vandoverae)
Mxp3 ischial
crista dentata:
prominent
toothed ridge
(0); obsolete or
absent (1)
(0), present in
M. monodon (cf.
Kensley et al.,
1991, fig. 15G)
(0), in M. vandoverae (cf.
Gore, 1983,
fig. 3B)
Mxp3 meral
spine:
present (0);
absent (1)
Male Plp1
Male Plp2
Male Plps3-5
(0), in M. monodon (cf. Kensley et al., 1991,
fig. 15G); M.
longispina (cf.
Kensley et al.,
1991, fig. 13L)
composed of stemlike proximal segment and cleavershaped distal segment (M. bouvieri,
cf. Kensley &
Heard, 1991,
fig. 11J)
composed of stemlike proximal segment and cleavershaped distal
segment
earshaped exopod and
oval endopod with
appendices interna and
masculina (M. bouvieri,
cf. Kensley & Heard,
1991, fig. 11K)
with appendix interna (M. bouvieri, cf. Kensley &
Heard, 1991,
fig. 11L)
broadened triangular exopod, and ovate, elongate endopod bearing appendices
interna and masculina, both
exopod and endopod bearing frond-like bisegmented
filaments marginally
endopods with
appendix interna,
both exopods
and endopods
bearing frondlike bisegmented
filaments marginally
(1), in M. lamellosa (cf. Kensley et al., 1991,
fig. 17G); in
M. microphylla
(cf. Poore, 1997,
fig. 20F)
CRM 013 – Katsushi Sakai
Meticonaxius
(type sp.:
M. monodon)
Length of Mxp2
exopod: almost
reaching to end
of merus (0);
reduced or
absent (1)
(1), in M. monodon (Kensley
& Heard, 1991,
fig. 15M); (0),
in M. bouvieri
(Kensley &
Heard, 1991,
fig. 11I)
(0), in M. vandoverae; (1), in
M. pillsburyi,
reduced (Kensley & Heard,
1991, fig. 17f)
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
301
2 – Uropodal endopod trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marcusiaxius
– Uropodal endopod oval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tethisea
Genus Marcusiaxius De Carvalho & Rodrigues, 1973
Marcusiaxius De Carvalho & Rodrigues, 1973: 553; Kensley & Heard, 1991: 506, fig. 2C;
Sakai, 1992c: 25; Poore, 1994: 99 (key); Poore, 1997: 357.
Diagnosis. — Rostrum prominent and triangular with obtuse apex, bearing
middorsally raised triangular area with flat surface. Carapace bearing anterolaterally short submarginal setal row on each side. Linea thalassinica absent.
Eyestalks deformed and subacute apically. Anterolateral cephalothorax with
two vertical rows of setae, anterior row shorter than posterior one. P1 chelate
and symmetrical. Abdominal somite 1 with a longitudinal row of setae, abdominal somite 2 with longitudinal row of setae and transverse one, abdominal somites 3-5 with transverse row of setae laterally, and abdominal somite
6 with three rows of setae, converging posteroventrally; abdominal somites
3-5 with dorsal patch of dense plumose setae. A1 peduncular segment 1 elongate. Scaphocerite less than half length of A2 peduncular segment 4. Mxp3
without crista dentata; merus mesially with weak row of setae; exopod absent
or rudimentary. P1 fixed finger with distinct, curved tooth on proximal third of
cutting edge. P2 chelate, fixed finger with denticles on cutting edge. Ps3-4 simple, propodi ovate. Pleurobranchs on Ps2-4. Male Plp1 uniramous and bisegmented, distal segment cleaver-shaped; male Plp2 biramous, endopod broadly
triangular, bearing mesiomedially appendices interna and masculina, exopod
broad; male Plps3-5 endopods with appendix interna. Male Plps2-5 without
marginal lamellae. Telson broader than long and rounded on posterior margin
without median spine. Uropodal endopod longer than wide and tear-shaped
or subtriangular with acute distolateral angle. Uropodal exopod broader than
endopod, bearing neither distal flap nor lateral notch. [Adapted from Sakai,
1992c; Poore, 1997.]
Remarks. — Meticonaxius longispina (Stebbing, 1920) and M. microps
(Bouvier, 1905), which were included in the genus Meticonaxius by Kensley &
Heard (1991), have turned out to be more properly included in Marcusiaxius,
because in those two species the rostrum is not so acutely triangular as in
Meticonaxius, but obtusely triangular as in Marcusiaxius.
Type species. — Marcusiaxius lemoscastroi De Carvalho & Rodrigues,
1973, by original designation and monotypy. The gender of the generic name,
Marcusiaxius, is masculine.
Species included. — Marcusiaxius colpos Kensley & Heard, 1991; M.
lemoscastroi De Carvalho & Rodrigues, 1973; M. longispina (Stebbing, 1920);
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CRM 013 – Katsushi Sakai
M. microps (Bouvier, 1905); M. minutus (Coelho, 1973); M. torbeni Sakai,
1992; M. wamsoi Poore, 1997; M. sp. Poore, 1997.
K EY
TO THE
S PECIES
OF THE
G ENUS MARCUSIAXIUS∗ )
1 – Rostrum with crenulate margin, median carina broad with rounded anterior end . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. longispina
– Rostrum with smooth margin, median carina convergent toward anterior end . . . . . . . . 2
2 – Telson slightly longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. microps
– Telson wider than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Telson rounded on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Telson pentagonal in shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4 – Mxp3 exopod overreaching distal margin of ischium . . . . . . . . . . . . . . . . . . . . . M. wamsoi
– Mxp3 exopod rudimentary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. torbeni
5 – Mxp3 exopod lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. lemoscastroi
– Mxp3 exopod reaching to distal margin of ischium . . . . . . . . . . . . . . . . . . . . . . . . M. colpos
∗ ) The present key does not include M. minutus, because it has been described only briefly.
Marcusiaxius colpos Kensley & Heard, 1991
Marcusiaxius colpos Kensley & Heard, 1991: 506, figs. 7, 8, tab. 1C; Poore, 1997: 358.
Diagnosis. — Male Plp1 uniramous and bisegmented, proximal segment
stem-like, distal segment cleaver-shaped; male Plp2 biramous, endopod
broadly triangular, bearing mesiomedially appendices interna and masculina,
appendix masculina twice as long as appendix interna; male Plps3-5 biramous,
endopods with appendix interna. [Cf. Kensley & Heard, 1991, fig. 8G, H, I.]
Type locality. — Gulf of Mexico, 29◦ 40′ N 86◦ 17′ W, 73 m.
Distribution. — Gulf of Mexico (Kensley & Heard, 1991); 43-175 m.
Marcusiaxius lemoscastroi De Carvalho & Rodrigues, 1973
Marcusiaxius lemoscastroi Rodrigues & De Carvalho, 1972: 337 [nomen nudum]; De Carvalho
& Rodrigues, 1973: 553, figs. 1-21; Kensley & Heard, 1991: 507, figs. 9, 10, tab. 1D; BlancoRambla, 1995: 60, figs. 5, 6; Poore, 1997: 358, fig. 5; Rodrigues et al., 1998: 381; Melo, 1999:
336, figs. 225-226.
Meticonaxius lemoscastroi — Coelho et al., 1980: 39; Coelho, 1987: 63, 68; Coelho & RamosPorto, 1987: 32 (key), 33.
Diagnosis. — Male Plp1 uniramous and bisegmented, proximal segment
stem-like and distal segment cleaver-shaped; male Plp2 biramous, endopod
and exopod broad, endopod with appendices interna and masculina, exopod
broad and ear-shaped; male Plps3-5 endopods with appendix interna. [Cf.
Kensley & Heard, 1991, fig. 9I, J, K.]
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Type locality. — Amapá, north coast of Brazil, 03◦ 40′ N 49◦ 55.5′ W, littoral.
Distribution. — Honduras, off Limóns (Poore, 1997); Panama, Culebra
Island (Poore, 1997); Colombia — Santa Marta (Poore, 1997); Venezuela —
Gulf of Venezuela (Kensley & Heard, 1991), N. of Margarita Island (Poore,
1997); Brazil — (Poore, 1997), Amapá, north coast of Brazil (De Carvalho &
Rodrigues, 1973), Amapá and Pará (Coelho & Ramos-Porto, 1987), Fortaleza
(Kensley & Heard, 1991); littoral to 73 m.
Marcusiaxius longispina (Stebbing, 1920)
(fig. 59A)
Axius longispina Stebbing, 1920: 265, pls. 106B, 107.
Meticonaxius ? longispina — De Man, 1925d: 5 (list).
Meticonaxius longispina — Barnard, 1950: 500, fig. 93a-c; Kensley, 1981b: 30; Coelho, 1987:
63; Kensley & Heard, 1991: 514, fig. 13, tab. 1F; Poore, 1997: 365 (key), 369.
Material examined. — SMA-A2754, 1 male (TL/CL, 14.0/4.4 mm), Pieter Faure Sta. 13039,
off East London, South Africa, 91 m.
Remarks. — Male Plp1 is missing and inaccessible, however, Kensley &
Heard (1991, fig. 13N) show in their figure that the male Plp1 consists of a
Fig. 59. Marcusiaxius longispina (Stebbing, 1920), Meticonaxius dentatus Lin, 2006, and
Michelea abranchiata Poore, 1997. A, male Plp2; B, anterior part of carapace; C, male
Plp1. A, Marcusiaxius longispina, SMA-A2754, 1 male (TL/CL, 14.0/4.4 mm), Pieter Faure
Sta. 13039, off East London, South Africa, 91 m; B, Meticonaxius dentatus, MNHN Th
1464, holotype, hermaphroditic female (TL/CL, 66.0/17.0 mm), New Caledonia, 20◦ 57.5′ S
165◦ 35.9′ E, BATHUS 1 (French Expedition), 302-335 m; C, Michelea abranchiata, USNM
291184, paratype, 1 male (TL/CL, 22.0/4.3 mm), Puerto Rico, Caribbean Sea. Scales 1 mm.
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CRM 013 – Katsushi Sakai
stem-like proximal segment and a cleaver-shaped distal segment. Male Plp2
(fig. 59A) endopod bears appendices interna and masculina, and male Plps3-5
endopods bear an appendix interna.
Type locality. — Off Cape Morgan, South Africa, 95 m.
Distribution. — South Africa, Cape Morgan (Stebbing, 1920; Poore, 1997),
off East London (Barnard, 1950; Kensley & Heard, 1991); 91-126 m.
Marcusiaxius microps (Bouvier, 1905)
Metaxius microps Bouvier, 1905: 804; Bouvier, 1925: 470, fig. 28; Balss, 1925: 210; De Man,
1925d: 8 (list); De Man, 1928b: 18, 20, 21; Schmitt, 1935a: 192, fig. 53; Balss, 1957: 1582.
Meticonaxius microps — Kensley & Heard, 1991: 516, fig. 14, tab. 1G; Poore, 1997: 365 (key),
369.
Diagnosis. — Male Plp1 consisting of stem-like proximal segment and
cleaver-shaped distal segment; male Plp2 biramous (probably endopod bearing
appendices interna and masculina), and male Plps3-5 endopods with appendix
interna. [Cf. Kensley & Heard, 1991, fig. 14I, J.]
Type locality. — Off St. Croix, Antilles, 210 m.
Distribution. — St. Croix (Bouvier, 1905, 1925; Kensley & Heard, 1991;
Poore, 1997); 210 m.
Marcusiaxius minutus (Coelho, 1973)
Meticonaxius minutus Coelho, 1973: 345; Coelho et al., 1980: 58; Coelho, 1987: 63, 68; Coelho
& Ramos-Porto, 1987: 32; Kensley & Heard, 1991: 516.
Marcusiaxius minutus — Poore, 1997: 358 (key), 360; Rodrigues et al., 1998: 381; Melo, 1999:
338, figs. 227-228.
Type locality. — Off Amapá, Brazil, 04◦ 18′ S 50◦ 17′ W, “Almirante Saldanba” Sta. 2413, 89-90 m.
Distribution. — Only known from the type locality.
Marcusiaxius torbeni Sakai, 1992
Marcusiaxius torbeni Sakai, 1992c: 26, figs. 9, 11; Poore, 1997: 358 (key), 360, fig. 6.
Diagnosis. — Eyestalks triangular and flattened on lateral surface, cornea
small and discrete subapically. [Cf. Sakai, 1992c: 27.]
Type locality. — West coast of Malay Peninsula, 9◦ 13′ 22′′ N 97◦ 50′ 11′′ E,
70 m.
Distribution. — Only known from the type locality.
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Marcusiaxius wamsoi Poore, 1997
Marcusiaxius wamsoi Poore, 1997: 358 (key), 361, figs. 7, 8.
Material examined. — RMNH D 45521, holotype, female (TL/CL, 48.0/13.4 mm, damaged,
Mxp1-3, Ps1-5, Plps2-5 on left side, Plps4-5 on right side picked off), Irian Jaya, 1 mile east of
Dauwi, Wamsoi Lagoon, Padaid Island, Indonesia, Adaido Group, 54-90 m, National Science
Foundation, Ostheimer, 4.ii.1956, leg. Orr. Powell.
Diagnosis. — Male Plps1-2 unknown. Female Plp1 uniramous and bisegmented; female Plps2-3 (4-5 missing) endopods with appendix interna.
Remarks. — It is observed, as Poore (1997: 364) mentioned, that the female
Plp1 is bisegmented: a proximal segment and a distal segment with distal setae.
Type locality. — One mile E. of Dauwi, Wamsoi lagoon, Padaido Islands,
Irian Jaya, Indonesia, 01◦ 17′ S 136◦ 46′ E, 54-90 m.
Distribution. — Indonesia — Padaido Islands, Irian Jaya (Poore, 1997);
54-90 m.
Marcusiaxius sp. Poore, 1997
Marcusiaxius sp., Poore, 1997: 364.
Distribution. — England, Gault, Folkestone (Poore, 1997).
Genus Meticonaxius De Man, 1905
Meticonaxius De Man, 1905: 592; De Man, 1925d: 2 (key), 53; De Man, 1928b: 18, 21, 30;
Barnard, 1950: 498 (key), 499; Balss, 1957: 1579; De Saint Laurent, 1973: 515; Le Loeuff &
Intès, 1974: 23; De Saint Laurent, 1979: 1397; Sakai & De Saint Laurent, 1989: 9; Kensley
& Heard, 1991: 510; Sakai, 1992c: 20; Poore, 1994: 99 (key); Poore, 1997: 357 (key), 364.
Metaxius Bouvier, 1905: 804; Bouvier, 1925: 469; De Man, 1925d: 2, 8 (list); De Man, 1928b:
18, 20, 21, 30; Balss, 1957: 1582.
Diagnosis. — Rostrum acutely triangular, usually carinate medially. Carapace with cardiac prominence, but without cardiac sulcus and linea thalassinica. Eyestalks visible in dorsal view, shorter than rostrum, rounded or
truncate-flattened distally; cornea weakly or moderately pigmented. Anterolateral cephalothorax with at least one vertical row of setae close to each lateral carina. Cervical groove distinct. Postcervical carina present posteriorly.
A1 peduncular segment 1 elongate. Scaphocerite less than half length of A2
penultimate segment. Mxp3 crista dentata reduced; merus with strong mesial
row of setae; exopod very short or absent. P1 chelate and symmetrical; fixed
finger with at least one sharp, curved tooth on distal third of cutting edge.
P2 fixed finger with spinules on cutting edge; dactylus as long as fixed finger. Ps3-4 propodi and dactyli without lateral spiniform setae (rarely one or
two on P4 dactylus); P4 propodus with one or two transverse rows of setae
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CRM 013 – Katsushi Sakai
laterally. Pleurobranchs present above Ps2-4. Abdominal somite 1 with longitudinal row of setae laterally, abdominal somites 2-5 with vertical rows of
setae laterally, and abdominal somite 6 with 2-3 rows of setae converging
posteroventrally; abdominal somites 3-5 with dense plumose setae laterally.
Male Plp1 uniramous and bisegmented, proximal segment stem-like and distal
segment cleaver-shaped; male Plp2 biramous, endopod bearing mesiomedially appendices interna and masculina, exopod ear-shaped; male Plps3-5 biramous, endopods with appendix interna, male Plps2-5 endopods without marginal lamellae. Telson longer than wide, lateral margins each with small proximal lobe, posterior margin rounded, without median spine. Uropodal endopod
shorter than wide and subtriangular with acute distolateral angle. Uropodal exopod bearing neither distal flap nor lateral notch. [Adapted from Sakai, 1992c;
Poore, 1997.]
Type species. — Meticonaxius monodon De Man, 1905, by original designation and monotypy. Gender of generic name, Meticonaxius, masculine.
Species included. — Meticonaxius bouvieri Kensley & Heard, 1991; M.
capricorni Coelho, 1987; M. dentatus Lin, 2006; M. monodon De Man, 1905;
M. noumea Poore, 1997; M. soelae Sakai, 1992c; M. spicatus Poore, 1997; M.
sp. Poore, 1997.
K EY
TO THE
S PECIES
OF THE
G ENUS METICONAXIUS
1 – Dorsal surface of rostrum with divergent dorsomedian carina . . . . . . . . . . . . . . . . . . . . . . 2
– Dorsal surface of rostrum with lineal dorsomedian carina . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 – P1 fixed finger with one tooth on cutting edge . . . . . . . . . . . . . . . . . . . . . . . . M. capricorni
– P1 fixed finger with 4-5 sharp spines on cutting edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Eyestalks overreaching rostrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. spicatus
– Eyestalks reaching half of rostrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. dentatus
4 – P1 fixed finger with one distinct median tooth on cutting edge . . . . . . . . . . . M. monodon
– P1 fixed finger with one distinct median tooth and proximal denticles on cutting edge
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
5 – Mxp3 merus urnarmed on mesial margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. bouvieri
– Mxp3 merus with one spine on mesial margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – Telson longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. soelae
– Telson wider than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. noumea
Meticonaxius bouvieri Kensley & Heard, 1991
Meticonaxius bouvieri Kensley & Heard, 1991: 512, figs. 11, 12, tab. 1E; Poore, 1997: 365
(key), 366, figs. 9, 10.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment cleaver-shaped; male Plp2 biramous, endopod triangular, bearing appendices interna and masculina, appendix masculina elongate and more than twice as long
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307
as appendix interna; male Plps3-5 biramous, endopods with appendix interna.
[Cf. Kensley & Heard, 1991, fig. 11J, K, L.]
Type locality. — Off Jamaica, 17◦ 13′ N 77◦ 45′ W, 722-768 m.
Distribution. — Caribbean Sea (Poore, 1997); Bahamas, north of Long
Island (Kensley & Heard, 1991); off Jamaica (Kensley & Heard, 1991); 180768 m.
Meticonaxius capricorni Coelho, 1987
Meticonaxius sp., Coelho & Ramos-Porto, 1987: 32 (key), 33.
Meticonaxius capricorni Coelho, 1987: 64, figs. 1-3; Kensley & Heard, 1991: 513; Poore, 1997:
365 (key), 369, fig. 11A; Rodrigues et al., 1998: 381; Melo, 1999: 340, figs. 229-230.
Remarks. — The present species, M. capricorni is similar to M. monodon
in that the fixed finger of the larger cheliped bears a distinct tooth on the
cutting edge, but differs in that in M. capricorni the P3 propodus is as long as
wide, convex on the dorsal margin, rounded on the ventral margin, and bilobed
distally, and the uropodal endopod is wider than long and trapezoid, while in
M. monodon the P3 propodus is subsquare, and the uropodal endopod is as
long as wide.
Type locality. — Brazil, 23◦ 52′ S 43◦ 11′ W, 156 m.
Distribution. — Brazil, Baia da Guanabara, Rio de Janeiro (Coelho &
Ramos-Porto, 1987), off Rio de Janeiro and Cabo de São Tomé (Coelho, 1987;
Poore, 1997); 139-214 m.
Meticonaxius dentatus Lin, 2006
(fig. 59B)
Meticonaxius dentatus Lin, 2006: 234, figs. 1, 2.
Material examined. — MNHN Th 1464, holotype, hermaphroditic female (TL/CL, 66.0/17.0
mm), New Caledonia, 20◦ 57.5′ S 165◦ 35.9′ E, BATHUS 1 (French expedition), 302-335 m,
18.iii.1993, leg. B. Richer de Forges.
Diagnosis. — Rostrum convex ventrally (fig. 59B). Plp1 uniramous, consisting of proximal segment and flagellum; Plps2-5 endopods with appendix
interna. Genital pore on Ps3 and 5 coxae (MNHN Th 1464). Hermaphroditic.
Type locality. — New Caledonia, 20◦ 57.5′ S 165◦ 35.9′ E, 302-335 m.
Distribution. — Only known from the type locality.
Meticonaxius monodon De Man, 1905
Meticonaxius monodon De Man, 1905: 593; De Man, 1925d: 5 (list), 54, pl. 4 fig. 10, pl. 5
fig. 10a-t; Balss, 1925: 210; De Man, 1928b: 19, 21, 30; Kensley & Heard, 1991: 516, fig. 15,
tab. 1H; Poore, 1997: 365 (key), 369.
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CRM 013 – Katsushi Sakai
Callianassa (Calliactites) coeca Balss, 1921: 175. [Type locality. — Dar es Salaam, Tanzania.]
Callianassa (?Scallasis) coeca — Balss, 1925: 212, fig. 16; De Man, 1928b: 30.
Material examined. — ZMA Crust. De 240583 (not RMNH), 1 female (damaged), off
northeast of Java, 7◦ 46′ S 114◦ 30.5′ E, “Siboga” Sta. 5, 330 m.
Diagnosis. — Rostrum triangular and pointed at apex, median carina
extending posteriorly onto slightly concave gastric region; lateral margins
extending posteriorly a bit onto gastric region. A2 scaphocerite overreaching
midlength of penultimate segment. P1 equal and chelate. Telson subsquare,
bearing proximal lobe on each of parallel lateral margins, posterior margin
straight, without median spine. Uropodal exopod bearing neither distal flap
nor lateral notch.
Type locality. — Off northeast Java, 7◦ 46′ S 114◦ 30.5′ E, “Siboga” Sta. 5,
330 M.
Distribution. — Tanzania — Dar es Salaam (Balss, 1921, 1925; Poore,
1997); Indonesia — northeast of Java (De Man, 1905, 1925; Kensley & Heard,
1991; Poore, 1997); 330-404 m.
Meticonaxius noumea Poore, 1997
Meticonaxius noumea Poore, 1997: 365 (key), 370, fig. 11B-K.
Type locality. — SSE of Yaté, New Caledonia, 22◦ 13′ S 167◦ 08′ E, 275320 m.
Distribution. — New Caledonia (Poore, 1997); 275-320 m.
Meticonaxius soelae Sakai, 1992
Meticonaxius soelae Sakai, 1992c: 21, figs. 6-8; Poore, 1997: 365 (key), 370; Tsang et al., 2008:
218, 219.
Type locality. — Coral Sea, E. Australia, 17◦ 59.4′ S 147◦ 64′ E, 298-300 m.
Distribution. — Only known from the type locality.
Meticonaxius spicatus Poore, 1997
Meticonaxius spicatus Poore, 1997: 365 (key), 370, fig. 12.
Type locality. — Caribbean Sea, “Atlantis” Sta. 3418, exact location unknown, 351 m.
Distribution. — Caribbean Sea (Poore, 1997); 351 m.
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Meticonaxius sp. Poore, 1997
Meticonaxius sp. Poore, 1997: 327.
Distribution. — Philippines (“Albatross” Sta.).
Genus Tethisea Poore, 1994
Tethisea Poore, 1994: 99; Poore, 1997: 357 (key), 393.
Diagnosis. — Rostrum prominent and flat. Linea thalassinica absent. Eyestalks invisible in dorsal view. P1 chelate and symmetrical; fixed finger with
thickened setae in proximal gape. Pleurobranchs present above Ps2-4. Ps3-4
without lateral spiniform setae. Male Plp1 uniramous and bisegmented; male
Plp2 biramous, endopod elongate and leaf-like, bearing appendices interna and
masculina, exopod small and ovate; male Plps3-5 biramous, endopods with
appendix interna. Telson longer than wide and trapezoid. Uropodal endopod
oval, uropodal exopod longer than wide, slightly concave on lateral margin
and broadly rounded on distal margin. Rows of setae present laterally on carapace and abdominal somites 1-2 and 6. [Adapted from Poore, 1994.]
Type species. — Tethisea indica Poore, 1994, by original designation and
monotypy. The gender of the generic name, Tethisea, is feminine.
Species included. — Tethisea indica Poore, 1994; T. mindoro Poore, 1997.
Tethisea indica Poore, 1994
Tethisea indica Poore, 1994: 100; Poore, 1997: 394, figs. 25, 26.
Material examined. — MNHN Th 1216, paratypes, 1 male (TL/CL, 18.0/6.0 mm); 1 female
(TL/CL, 20.0/6.5 mm), Indonesia, Macassar Strait, 0◦ 54.2′ S 119◦ 28.7′ E, det. G. Poore.
Diagnosis. — Rostrum triangular and 1.5 times as long as wide at base of
eyes. Carapace anterolaterally with a row of three setae. Mxp3 merus without
mesial tooth; exopod minute. P1 merus with two spines on ventral margin.
Male Plp1 uniramous and bisegmented, consisting of stem-like proximal
segment and subovate distal segment bearing small mesioproximal lobe with
hooklets; male Plp2 biramous, exopod ovate, and endopod elongate and leaflike, bearing mesiomedially appendices interna and masculina; male Plp3
with appendix interna. Female Plp1 uniramous, slender, and three-segmented;
female Plps2-5 endopods with appendix interna. Telson longer than wide and
trapezoid. Uropodal endopod oval; uropodal exopod subtriangular, slightly
concave on lateral margin, and broadly rounded distally.
Type locality. — Mozambique, 24◦ 64′ S 35◦ 20′ E, 165 m.
Distribution. — Mozambique (Poore, 1994, 1997); La Réunion; Indonesia;
New Caledonia; 165-460 m.
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CRM 013 – Katsushi Sakai
Tethisea mindoro Poore, 1997
Tethisea mindoro Poore, 1997: 398, fig. 27; Komai, 2000b: 346 (list); Robles et al., 2009: 316.
Material examined. — MNHN Th 1222, holotype, female (CL 51.0 mm), Philippines, W. of
Mindoro, 12◦ 31.2′ N 120◦ 39.3′ E, MUSORSTOM Sta. DR 117, 92-97 m, 3.vi.1985.
Diagnosis. — Male Plps1-2 uncertain. Female Plp1 uniramous and bisegmented, consisting of proximal segment and flagellum; female Plps2-5 endopods with appendix interna.
Type locality. — W. of Mindoro, Philippines, 12◦ 31.2′ N 120◦ 39.3′ E, 9297 m.
Distribution. — Philippines — W. of Mindoro (Poore, 1997); Australia —
North-West Shelf (Poore, 1997); 92-134 m.
FAMILY MICHELEIDAE SAKAI, 1992
Micheleinae Sakai, 1992c: 18; Poore, 1994: 99; Poore, 1997: 454; Felder, 2001: 440; Poore,
2004: 176.
Diagnosis. — Rostrum small, short, and triangular between orbits, located
between branchiostegals protruded to proximal half of eyestalks. Linea thalassinica absent; carapace with posterolateral lobes. A1 segment 1 extremely
elongate. Scaphocerite distinct. Maxilla 2 scaphognathite with 1-2 posterior
whips. P1 chelate, subequal, and similar in male and female. P2 chelate.
Ps3-4 propodi rectangular. P4 coxa flattened. P5 simple. Abdominal somite 1
with anteriorly-directed lobe laterally. Male Plp1 uniramous and un- or bisegmented, distal segment cleaver-shaped; male Plp2 biramous, endopod with appendices interna and masculina, or with appendix interna but without appendix
masculina; male Plps3-5 endopods with appendix interna. Male pleopods 2-5
with respiratory lamellae. Uropodal exopod bearing neither distal flap nor lateral notch. Rows of setae present on carapace, abdominal somites 1-6, and
Ps2-3 propodi.
Remarks. — Poore (1994: 99, 1997: 354) included four genera, Marcusiaxius Rodrigues & De Carvalho, 1973, Meticonaxius De Man, 1905, Tethisea
Poore, 1994, and Michelea Kensley & Heard, 1991 in the family Micheleidae
Sakai, 1992c. However, the genus Michelea is different from the other three
genera, Marcusiaxius, Meticonaxius, and Tethisea, because in Michelea the
rostrum is small, short, and triangular between the orbits, located between the
protruded branchiostegals, and the pleopods bear frond-like respiratory lamellae, whereas in the other three genera, the rostrum is prominent and protruded
anteriorly to form a flat triangular plate; and the pleopods lack frond-like respiratory lamellae. Therefore, Michelea is to be distinguished from the other
three genera at the family level.
Type genus. — Michelea Kensley & Heard, 1991, as the nominate genus.
Genus included. — Only the genus Michelea Kensley & Heard, 1991.
Genus Michelea Kensley & Heard, 1991
Michelea Kensley & Heard, 1991: 519; Poore, 1994: 99 (key); Poore, 1997: 357 (key), 373;
Poore, 2004: 176.
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CRM 013 – Katsushi Sakai
Diagnosis. — Rostrum small, short, obtusely triangular between orbits,
and shorter than eyestalks. Eyestalks visible in dorsal view. A1 peduncle
segment 1 extremely elongate. A2 scaphocerite distinct. Mxp3 with crista
dentata; merus lacking mesial spine; but with strong mesial row of setae;
exopod exceeding end of ischium. P1 chelate, subequal, and similar in male
and female. P2 chelate. Ps3-4 propodi rectangular. Ps2-3 propodi with row of
setae. P4 propodus with two transverse rows of setae. Pleurobranchs usually
absent. Male Plp1 uniramous and un- or bisegmented, distal segment cleavershaped; male Plp2 biramous, endopod elongate and ovate, bearing appendices
interna and masculina at proximal third, or bearing appendix interna but
no appendix masculina, and exopod broadened and triangular; male Plps3-5
biramous, endopods with appendix interna and frond-like respiratory lamellae
marginally. Uropodal endopod and exopod ovate. Telson convergent to form
rounded triangle. [Adapted from Poore, 1997.]
Type species. — Callianidea vandoverae Gore, 1987, by original designation. Gender of generic name, Michelea, feminine.
Species included. — Michelea abranchiata (Poore, 1997); M. dampieri
Poore, 2008; M. devaneyi Poore, 1997; M. hortus Poore, 1997; M. lamellosa
Kensley & Heard, 1991; M. lepta (Sakai, 1987a); M. leura (Poore & Griffin, 1979); M. microphylla Poore, 1997; M. novaecaledoniae Poore, 1997;
M. paraleura Poore, 1997; M. pillsburyi Kensley & Heard, 1991; M. vandoverae (Gore, 1987); M. sp., Robles et al., 2009: 316.
K EY
TO THE
S PECIES
OF THE
G ENUS MICHELEA
1 – Plps2-5 endopods with respiratory lamellae, but exopods without respiratory lamellae
............................................................................ 2
– Plps2-5 endopods and exopods with respiratory lamellae . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 – Telson parallel-sided in proximal half and tapering toward convex posterior margin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. lepta
– Telson tapering toward broadly rounded posterior margin . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Rostrum short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. novaecaledoniae
– Rostrum distinct, reaching middle of eyestalks . . . . . . . . . . . . . . . . . . . . . . . . . . M. hortus
4 – Plp2 endopod and exopod with respiratory lamellae on whole margin . . . . . . . . . . . . . . 5
– Plp2 endopod and exopod with respiratory lamellae in some parts only . . . . . . . . . . . . 6
5 – Respiratory lamellae bisegmented; maxilla 2 scaphognathite with two posterior whips
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. vandoverae
– Respiratory lamellae unsegmented; maxilla 2 scaphognathite with a single posterior
whip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. lamellosa
6 – Mxp3 merus without mesial tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
313
– Mxp3 merus with mesial tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7 – A1 peduncles reaching distal margin of A2 peduncles . . . . . . . . . . . . . . . M. abranchiata
– A1 peduncles overreaching distal margin of A2 penultimate segment . . . . . . . . . . . . . . 8
8 – A1 elongate proximal segment reaching distal fifth of A2 penultimate segment . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. dampieri
– A1 elongate proximal segment reaching proximal third of A2 penultimate segment . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. microphylla
9 – Cervical groove present at full length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
– Cervical groove short in median region of carapace . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 – Uropodal exopod with short marginal spiniform setae . . . . . . . . . . . . . . . . . . . . . M. leura
– Uropodal exopod with distinct dense marginal spiniform setae . . . . . . . . . M. paraleura
11 – A1 peduncle reaching end of A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. devaneyi
– A1 peduncle not reaching end of A2 penultimate segment . . . . . . . . . . . . . M. pillsburyi
Michelea abranchiata Poore, 1997
(fig. 59C)
Michelea abranchiata Poore, 1997: 374 (key), 375, figs. 13, 14.
Material examined. — USNM 291184, paratype, 1 male (TL/CL, 22.0/4.3 mm), Puerto Rico,
Caribbean Sea, 23.vi.1916, det. G. Poore.
Remarks. — The paratype (USNM 291184) examined seems to be different
from the holotype in the shape of Plp1, because it is bisegmented, and the
distal segment is cleaver-shaped with a patch of hooklets mesioproximally
(fig. 59C), though Poore described for M. abranchiata “Pleopod 1 of male with
a triangular distal blade” and showed by his figure that Plp1 is unsegmented
and the distal half is ovate, with a patch of hooklets medially on the convex
mesial margin (Poore, 1997, fig. 14L).
Type locality. — British West Indies, Barbuda, Spanish Point, Caribbean
Sea, 17◦ 41′ N 61◦ 48′ W, shore.
Distribution. — Caribbean Sea — Barbuda and Puerto Rico (Poore, 1997),
intertidal.
Michelea dampieri Poore, 2008
Michelea dampieri Poore, 2008: 175, figs. 6, 7.
Diagnosis. — Rostrum small, short, obtusely triangular between orbits, flat,
as long as broad at base, and about one-third as long as eyestalks. Cervical
groove weakly defined, reaching 0.6 length of cephalothorax, longitudinal
row of 4 setae level with lateral margin of eyestalk; marginal row of three
setae at base of eyestalks; lateral row of three setae. Eyestalks visible in
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CRM 013 – Katsushi Sakai
dorsal view. A1 peduncle segment 1 extremely elongate and waisted, and
half as long as cephalothorax. A2 scaphocerite distinct, peduncle reaching
to middle of A1 penultimate segment. Mxp3 ischium without crista dentata;
merus with strong mesial row of setae; exopod exceeding end of ischium.
P1 chelate and equal, ischium with weak ventral tooth; merus with weak
mid-tooth on slightly convex ventral margin, dorsal margin strongly convex,
especially proximally; carpus unarmed; propodus almost cylindrical; fixed
finger 0.4 length of propodus, cutting edge with obsolete tooth on proximal
half; dactylus with cutting edge irregular, curved distally, and equal to fixed
finger. P2 chelate, merus-propodus with rows of long setae on ventral margin;
carpus 0.6 length of merus; propodus little longer than carpus, with row of 7
short setae; fixed finger with 4 short spiniform setae on cutting edge; dactylus
longer than fixed finger, with 5 short spiniform setae on distal half of cutting
edge; each finger with corneous tip. P3 propodus twice as long as wide, bearing
4 spiniform setae and long setae on ventral margin, plus two on distoventral
and two on distodorsal mesial face, and two transverse rows of 4 setae on
lateral face; dactylus with spiniform seta on dorsomesial margin. P4 propodus
2.8 times as long as wide, 4-6 weakly aligned transverse rows of spiniform
setae on mesial face, concentrated near margins and strongest on ventral
margin and distally, and two transverse rows of 5 and 3 setae; propodus with
setae on ventral margin transforming to strongly pectinate distally; dactylus
with about 11 spiniform setae in weak rows on dorsomesial margin.
Male Plp1 curved mesially and expanded distally, bearing 8 minute hooklets
along mesiodistal margin; Plp2 endopod with probably 7 marginal lamellae
laterally; appendix interna saclike, and 3 times as long as wide, appendix
masculina 6 times as long as wide, and 1.7 times as long as appendix interna,
bearing 8 setae on distal third; exopod with 3 lamellae distolaterally. Plps3-4
endopods with 17 lamellae and exopods with 5 lamellae. Plp5 endopod with
5 lamellae and exopod with 2 lamellae. Telson 0.9 times as long as wide,
almost semicircular. Uropodal endopod ovate, and 1.3 times as long as wide,
bearing minute distal tooth. Uropodal exopod oval with oblique mesiodistal
margin, and 1.7 times as long as wide, bearing short spiniform setae on lateral
to rounded distolateral corner. [Adapted from Poore, 2008.]
Type locality. — Dampier Archipelago, Western Australia.
Distribution. — Only known from the type locality.
Michelea devaneyi Poore, 1997
Michelea devaneyi Poore, 1997: 375 (key), 376, fig. 15.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
315
Type locality. — North of Sand Island, Eniwetok Atoll, Marshall Islands,
11◦ 30′ S 162◦ 15′ E, 2 m.
Distribution. — Marshall Islands — Eniwetok Atoll (Poore, 1997), subtidal.
Michelea hortus Poore, 1997
Michelea hortus Poore, 1997: 374 (key), 378, fig. 16; Poore, 2004: 177.
Type locality. — One-and-a-half (1.5) miles west of south end of Garden
Island, Western Australia, 32◦ 12′ S 115◦ 40′ E.
Distribution. — Western Australia — Garden Island (Poore, 1997); shelf.
Michelea lamellosa Kensley & Heard, 1991
Michelea lamellosa Kensley & Heard, 1991: 519, figs. 16, 17, tab. 1I; Poore, 1997: 374 (key),
381.
Type locality. — Off Jamaica, 17◦ 36.3′ N 77◦ 02.6′ W, 24 m.
Distribution. — Only known from the type locality.
Michelea lepta (Sakai, 1987)
Callianidea lepta Sakai, 1987a: 300, 303 (list), fig. 3.
Michelea lepta — Kensley & Heard, 1991: 519; Poore, 1997: 374 (key), 381.
Material examined. — USNM 231419, holotype, male (TL/CL, 18.0/3.0 mm), Ryukyu
Islands, Japan, leg. S. Shokita, det. K. Sakai.
Diagnosis. — Plp2 biramous, exopod ovate and shorter than endopod,
endopod with marginal setae except on proximal margin and several simple
frond-like respiratory lamellae, bearing proximally slender appendix interna
with distal hooklets, and slender appendix masculina. Plp4 endopod with
marginal setae, but without frond-like respiratory lamellae.
Type locality. — Chatan, Okinawa Island, low tidal zone.
Distribution. — Japan — Okinawa Island (Sakai, 1987a); low tidal zone.
Michelea leura (Poore & Griffin, 1979)
Callianidea leura Poore & Griffin, 1979: 281, figs. 40, 41; Sakai, 1984: 104.
Michelea leura — Kensley & Heard, 1991: 519; Poore, 1997: 374 (key), 381, figs. 17-19.
Type locality. — Masthead Is., Capricorn Group, Queensland, Australia.
Distribution. — Australia — Masthead Island and Holbourne Island (Poore
& Griffin, 1979), Heron Island (Sakai, 1984), Masthead Island and Heron
Island (Poore, 1997).
316
CRM 013 – Katsushi Sakai
Michelea microphylla Poore, 1997
Michelea microphylla Poore, 1997: 374 (key), 386, figs. 20, 21; Poore, 2004: 177, fig. 47 (from
Poore, 1997).
Type locality. — Crib Point, Western Port, Victoria, Australia, 38◦ 19.92′ S
145◦ 13.95′ E, 19 m.
Distribution. — Only known from the type locality.
Michelea novaecaledoniae Poore, 1997
Michelea novaecaledoniae Poore, 1997: 374 (key), 388, fig. 22.
Type locality. — Baie de Prony, Île Ouen, New Caledonia, 22◦ 24′ S 166◦ 50′ E,
37 m.
Distribution. — New Caledonia — Île Ouen (Poore, 1997); 37-52 m.
Michelea paraleura Poore, 1997
Michelea paraleura Poore, 1997: 375 (key), 390, figs. 23, 24.
Type locality. — Holbourne Island, Queensland, Australia, 19◦ 42′ S
148◦ 21′ E.
Distribution. — Australia — Holbourne Island, Queensland, and Oxley
Island, Northern Territory (Poore, 1997); intertidal to 2 m.
Michelea pillsburyi Kensley & Heard, 1991
Michelea pillsburyi Kensley & Heard, 1991: 522, figs. 18, 19, tab. 1J; Poore, 1997: 374 (key),
393.
Type locality. — Caribbean Sea, off Panama, 9◦ 22.6′ N 78◦ 08.4′ W, 59 m.
Distribution. — Caribbean coast of Panama (Kensley & Heard, 1991);
28-59 m.
Michelea vandoverae (Gore, 1987)
Callianidea vandoverae Gore, 1987: 186, figs. 1-4.
Michelea vandoverae — Kensley & Heard, 1991: 523, figs. 20, 21, tab. 1K; Poore, 1997: 374
(key), 393.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment cleaver-shaped, bearing mesiomedially a patch of hooklets. Male Plp2 biramous,
with broadened triangular exopod, and elongate ovate endopod, bearing appendices interna and masculina at proximal third on mesial margin, both exopod and endopod with numerous frond-like bisegmented filaments marginally;
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
317
male Plps3-5 biramous, endopods with appendix interna, both exopods and
endopods bearing numerous frond-like bisegmented filaments marginally. [Cf.
Gore, 1987: 191; Kensley & Heard, 1991: 523, fig. 21I, J.]
Type locality. — Florida, east of St. Lucie Inlet, 27◦ 10.5′ N 80◦ 00.2′ W, 58 m.
Distribution. — Gulf of Mexico (Kensley & Heard, 1991) and S.W. Caribbean Sea (Poore, 1997); 37-162 m.
Michelea sp. Robles et al., 2009
Michelea sp., Robles et al., 2009: 316.
FAMILY STRAHLAXIIDAE POORE, 1994
Strahlaxiidae Poore, 1994: 100; Felder, 2001: 440; Poore, 2004: 178.
Diagnosis. — Rostrum narrowly or broadly rounded, with bifurcate apex;
gastric region flattened, bearing neither median nor submedian carinae; cardiac
sulcus and cardiac prominence absent. Cervical groove distinct; no linea
thalassinica; postcervical carina absent. Eyestalks distinct, cornea usually
pigmented. A1 peduncular segment 1 as long as segments 2-3 combined.
Scaphocerite distinct. Maxilla 2 scaphognathite with or without posterior
whip. Mxp3 with exopod. P1 chelate and unequal. P2 chelate. Ps3-4 simple;
P3 propodus broadly rectangular, and setose ventrally, lacking vertical rows
of spinules; P4 propodus elongate, lacking transverse rows of spinules, coxa
more or less flattened. P5 chelate. Pleurobranchs present or not. Abdominal
pleuron 1 protruded ventrally; abdominal pleuron 2 twice as long as pleuron 3;
abdominal pleura 3-5 with tufts of setae ventrally. Male Plp1 present or absent;
male Plp2 biramous, endopod bearing appendix interna, but no appendix
masculina; Plps3-5 biramous, endopods with appendix interna. Female Plp1
uniramous, consisting of proximal segment and multiarticulate flagellum;
female Plps2-5 biramous, endopods with appendix interna. Uropodal exopod
bearing neither distal flap nor lateral notch.
Remarks. — The family Strahlaxiidae Poore, 1994 was established based on
the genera Neaxius Borradaile, 1903, Neaxiopsis Sakai & De Saint Laurent,
1989, and Strahlaxius Sakai & De Saint Laurent, 1989 (type genus of the
Strahlaxiidae). Poore (1994: 100) mentioned that “the three strahlaxiid genera
are superficially similar to some members of the Axiidae where they have
traditionally been placed. The most significant diagnostic characters are the
acute outer angle of the uropodal endopod, the bifid rostrum, rugose gastric
region of the carapace, broad pleopodal rami, and possession of some setalrows on some pereopods and abdominal somites. Absence of the appendix
masculina is a synapomorphy but this is shared with some axiid genera”.
Type genus. — Strahlaxius Sakai & De Saint Laurent, 1989, as the nominate
genus.
Genera included. — Neaxiopsis Sakai & De Saint Laurent, 1989; Neaxius
Borradaile, 1903; and Strahlaxius Sakai & De Saint Laurent, 1989.
320
CRM 013 – Katsushi Sakai
K EY
TO THE
G ENERA
OF THE
FAMILY S TRAHLAXIIDAE
1 – Male Plp1 present; maxilla 2 scahphognathite with posterior whip . . . . . . . . Strahlaxius
– Male Plp1 absent; maxilla 2 scahphognathite without posterior whip . . . . . . . . . . . . . . . 2
2 – Rostrum rounded and dentate anteriorly, bearing apical notch. Telson without transverse
carinae; uropodal endopod simple on distolateral margin; uropodal exopod unarmed on
lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neaxiopsis
– Rostrum narrow and bifurcate anteriorly. Telson with transverse carinae; uropodal
exopod denticulate on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neaxius
Genus Neaxiopsis Sakai & De Saint Laurent, 1989
Neaxiopsis Sakai & De Saint Laurent, 1989: 32; Poore, 1994: 100 (key).
Diagnosis. — Rostrum broad and distinct, dentate on semicircular anterior
margin, with median notch, lateral margins extending posteriorly onto gastric
region. Gastric region flattened, bearing median carina anteriorly, but no submedian carinae. Anterolateral margin of carapace unarmed. Cervical groove
distinct, bearing no teeth. Eyestalks subglobose; cornea pigmented. Scaphocerite simple and distinct. Maxilla 2 scaphognathite without posterior whip.
Mxp3 with exopod. Pl unequal. P2 chelate. P3 propodus without transverse
rows of setae and spines. No pleurobranchs. Male Plp1 absent; male Plp2 biramous, endopod bearing appendix interna, but no appendix masculina; male
Plps3-5 biramous, endopods with appendix interna. Female Plp1 uniramous,
consisting of proximal segment and elongate multiarticulate flagellum; female
Plps2-5 endopods with appendix interna. Telson wider than long, or about as
long as wide, bearing median tooth on posterior margin. Uropodal endopod
rounded distolaterally; uropodal exopod bearing neither series of strong teeth
laterally, nor transverse suture.
Type species. — Callianidea gundlachi Von Martens, 1872, designated by
Sakai & De Saint Laurent, 1989. The gender of the generic name, Neaxiopsis,
is feminine.
Species included. — Neaxiopsis euryrhyncha (De Man, 1905); N. gundlachi
(Von Martens, 1872).
K EY
TO THE
S PECIES
OF THE
G ENUS NEAXIOPSIS
1 – Telson longer than wide, and slightly longer than uropods . . . . . . . . . . . . N. euryrhyncha
– Telson wider than long, and slightly shorter than uropods . . . . . . . . . . . . . . . N. gundlachi
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
321
Neaxiopsis euryrhyncha (De Man, 1905)
(fig. 60A-C)
Axius (Neaxius?) euryrhynchus De Man, 1905: 590.
Axius (Neaxius) euryrhynchus — De Man, 1925d: 3 (list), 12 (key), 25, pl. 1 fig. 2-2e, pl. 2
fig. 2f-k; Sakai, 1987a: 303 (list).
Neaxiopsis euryrhynchus — Sakai & De Saint Laurent, 1989: 33.
Not: Axius (Neaxius) euryrhynchus — Miyake, 1982: 90, 192 (list), pl. 30 fig. 5 [= Neaxiopsis
gundlachi (Von Martens, 1872)].
Material examined. — ZMA Crust. De 202879 (syntypes of Neaxiopsis euryrhynchus (De
Man, 1905)), two males (TL/CL, 11.0/4.2; 11.0/4.8 mm).
Diagnosis. — Male Plp1 absent; male Plp2 endopod with appendix interna,
but without appendix masculina. Male Plps3-5 biramous, endopods with
appendix interna.
Remarks. — De Man (1925d: 25-31) described this species as Axius (Neaxius?) euryrhynchus, based on two young female specimens measuring TL/CL,
11.06/4.56 and 10.5/4.34 mm. However, the present examination of these has
revealed that those two specimens regarded as females, are males. De Man
(1925d) mentioned that “A. euryrhynchus (= Axius (Neaxius) euryrhynchus)
should be considered as the young stage of the former (N. gundlachi var. N.
orientalis) or as a proper species (= N. gundlachi)”. However, N. euryrhyncha is clearly different in morphology from N. gundlachi as follows. In N.
euryrhyncha the rostrum is truncate anteriorly with a median notch, and dentate on the anterior margin (fig. 60A), the A1 peduncle overreaches the distal
margin of the P2 penultimate segment (fig. 60A, B), the telson is longer than
wide, slightly convergent toward the posterior margin, with a median tooth,
and distinctly longer than the uropodal endopod (fig. 60C), and the gastric region is elevated with a smooth and flattened surface bearing a short median
carina posterior to the flat rostrum (fig. 60A). In N. gundlachi in contrast, the
rostrum is broadly rounded posteriorly, with a median notch, and dentate on
the anterior margin (fig. 60D), the A1 peduncle fails to reach the distal margin
of the A2 penultimate segment (fig. 60D); the telson is wider than long and
slightly convergent towards the posterior margin, with a median tooth, and not
longer than the uropoal endopod (fig. 60E), and the gastric region is elevated
with a flattened surface with tubercles along the edge, the rostral median carina extends short posteriorly onto the elevated gastric region as a short gastric
median carina, and the rostral lateral margins extend onto the gastric region as
7-8 emarginated gastric lateral carinae (fig. 60D). Therefore, N. euryrhyncha
is not to be considered as the young stage of N. gundlachi.
322
CRM 013 – Katsushi Sakai
Fig. 60. Neaxiopsis euryrhyncha (De Man, 1905) and Neaxiopsis gundlachi (Von Martens,
1872). A, D, carapace, dorsal view; B, anterior part of carapace, lateral view; C, E, telson and
uropod. A, B, C, Neaxiopsis euryrhyncha, ZMB 8915, male (TL/CL, 58.0/21.4), Maupi, New
Pomerania; D, E, Neaxiopsis gundlachi, ZMB 4572, holotype, female (TL/CL, 92.0/32.0 mm),
Cuba. Scales 1 mm.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
323
Type locality. — Anchorage off Dongala, Palos-bay, Sulawesi, Indonesia,
“Siboga” Sta. 86, 36 m.
Distribution. — Indonesia — off Dongala, Sulawesi (De Man, 1905, 1925);
New Britain — Matupi near Rabaul (De Man, 1925b); 36 m.
Neaxiopsis gundlachi (Von Martens, 1872)
(fig. 60D-E)
Callianidea Gundlachi Von Martens, 1872: 132, pl. 5 figs. 15, 15b-c.
?Axius (Neaxius) Gundlachi — Borradaile, 1903: 537.
Axius gundlachi — Boas, 1880: 187; Rathbun, 1920: 327.
Axius (Neaxius) Gundlachi — De Man, 1925b: 120, fig. 1; De Man, 1925d: 4 (list), 9, 12 (key),
31.
Axius (Neaxius) Gundlachi var. orientalis De Man, 1925b: 122, fig. 2-2b; De Man, 1925d:
4 (list), 12 (key), 31. [Type locality. — Matupi near Rabaul, New Britain, Bismarck
Archipelago.]
Axius (Neaxius) euryrhynchus — Miyake, 1982: 90, 192 (list), pl. 30 fig. 5.
Neaxiopsis gundlachi — Sakai & De Saint Laurent, 1989: 32.
Neaxiopsis orientalis — Sakai & De Saint Laurent, 1989: 33.
Material examined. — ZMB 4572, holotype, female (TL/CL, 92.0/32.0 mm), Cuba, leg.
J. Gundlach; RMNH D 2300, 1 male (TL/CL, 83.0/28.5 mm), harbour of Willemstad, Curaçao,
31.iii.1905, leg. Boecke; ZMB 8915 (syntypes of Neaxiopsis orientalis (De Man, 1925b)),
2 males (TL/CL, 58.0/21.4; CL 22.1 mm, posterior part of abdomen missing), 2 females
(TL/CL, 48.0/17.1-74.0/25.9 mm), Matupi, New Pommerania (= Matupit Island near Rabaul,
New Britain, Bismarck Archipelago, Papua New Guinea), leg. Finsch.
Diagnosis. — Rostrum broadly rounded anteriorly, with median notch,
and dentate on anterior margin (fig. 60D). Male Plp1 absent; male Plp2
biramous, endopod with appendix interna, but without appendix masculina.
Female Plp1 uniramous, consisting of proximal segment and multiarticulate
flagellum; Plps2-5 endopods broad and leaf-like in shape, bearing appendix
interna. No pleurobranchs. Telson wider than long and slightly convergent
toward posterior margin, with median tooth, and not longer than uropodal
endopod (fig. 60E).
Remarks. — The syntypes of N. orientalis from Matupi (= Matupit Island
near Rabaul, New Britain, Bismarck Archipelago, Papua New Guinea) were
examined, and it was found that in N. orientalis the ventral margin of the
P1 palm is carinate ventrally, with a row of tubercles along the carina as
in N. gundlachi (De Man, 1925b, fig. 1), but not as in N. euryrhynchus, in
which it is carinate without any row of tubercles along the carina. De Man
(1925b) described the difference between N. orientalis and N. gundlachi as
“die Palmarportion der grösseren Schere des Männchens, deren Aussen-und
Innenseite ein wenig minder fein gekörnt sind als bei der westindischen Art
324
CRM 013 – Katsushi Sakai
(= in N. gundlachi)” [“the lateral and mesial surfaces of the male Pl palm
are a little less granulate than those in the western Indian species (= in N.
gundlachi)”]. However, such a difference is not to be regarded as characteristic
enough to distinguish one species from the other. So, it seems that N. orientalis
is synonymous with N. gundlachi.
Miyake (1982: 90) described Axius (Neaxius) euryrhynchus on the basis of
a large female specimen measuring TL/CL, 69.0/28.0 mm from Tanabe Bay,
west coast of Kii Peninsula, Japan, sandy-mud, 20 m. The Japanese female,
which is much larger than the male syntypes of Neaxiopsis euryrhynchus from
Celebes, 36 m, measuring TL/CL, 11.0/4.2 and 11.0/4.8 mm, was described
in Japanese as follows: “the gastric region bears tubercles along the 7-8
emarginated lateral carinae extended from the rostral lateral margins, and the
telson is slightly longer than the uropods”, though his colour photo (Miyake,
1982, pl. 30 fig. 5) shows that the uropods on the left side slightly overreach
the telson (= the telson is slightly shorter than the uropods), and additionally
the A1 peduncle is shorter than the distal end of the A2 penultimate segment
as in N. gundlachi. Those characteristics suggest that his large Japanese
specimen regarded as N. euryrhyncha should be considered to be N. gundlachi,
but not N. euryrhyncha, in which the gastric region bears neither tubercles
nor emarginated lateral carinae, and the telson is distinctly longer than the
uropods. In consequence, the Japanese female specimen is safely identified as
N. gundlachi.
Type locality. — Cuba.
Distribution. — Cuba (Von Martens, 1872); Curaçao (De Man, 1925b),
Schottegat and harbour of Willemstad (Rathbun, 1920), Schottegat (Sakai &
De Saint Laurent, 1989), Tanabe Bay, west coast of Kii Peninsula (on the
Pacific side), Japan, sandy-mud (Miyake, 1982); 20 m.
Genus Neaxius Borradaile, 1903
Neaxius s. str. Borradaile, 1903: 537; De Man, 1925d: 1 (key), 3 (list), 12 (key); Sakai, 1987a:
303 (list).
Neaxius — Sakai & De Saint Laurent, 1989: 29; Poore, 1994: 100 (key); Sakai, 1994: 176, 200
(key).
Diagnosis. — Rostrum narrow and bifurcate distally, lateral margins armed
with teeth, and median carina extending onto gastric region. Gastric region flattened. Anterolateral margin of carapace armed with teeth. Cervical groove distinct almost at full length, and usually armed with lateral teeth. Eyestalks subglobose; cornea pigmented. A2 scaphocerite elongate and either unarmed, or
armed with marginal teeth. Maxilla 2 scaphognathite without posterior whip.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
325
Pl unequal. P2 chelate. Ps3-4 simple. P3 propodus broadened and setose on
ventral margin. P4 propodus elongate. P5 chelate. Ps2-5 with pleurobranch.
Abdominal pleura 3-5 with stout tuft of setae ventrally. Male Plp1 absent;
male Plp2 biramous, endopod with appendix interna but without appendix
masculina; male Plps3-5 biramous, endopods with appendix interna. Female
Plp1 uniramous, consisting of proximal segment and flagellum; Plps2-5 biramous, broad and setose, endopods with appendix interna. Telson usually wider
than long, bearing 1-3 transverse carinae. Uropodal endopod protruded distolaterally; uropodal exopod denticulate on lateral margin, bearing no transverse
suture.
Type species. — Axius glyptocercus Von Martens, 1869 [= synonym,
Axia acanthus A. Milne-Edwards, 1878, designated by Borradaile, 1903], by
original designation and monotypy. Gender of the generic name, Neaxius,
masculine.
Species included. — Neaxius frankeae Lemaitre & Ramos, 1992; N. glyptocercus (Von Martens, 1869) [= Neaxius acanthus (A. Milne-Edwards, 1878);
N. trondlei Ngoc-Ho, 2005]; N. mclaughlinae Ngoc-Ho, 2006; N. vivesi (Bouvier, 1895).
K EY
TO THE
S PECIES
OF THE
G ENUS NEAXIUS
1 – Both carpus and palm of male larger cheliped armed with spines on dorsal margins
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
– Both carpus and palm of male larger cheliped unarmed on dorsal margins . . . . . . . . . . . 3
2 – Telson with three transverse carinae, second and third carinae curved posteriorly at both
ends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. frankeae
– Telson with three transverse carinae, second carina straight, and third one interrupted
medially and curved posteriorly at both ends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. vivesi
3 – Telson with three transverse carinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. mclaughlinae
– Telson with two transverse carinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . N. glyptocercus (= N. acanthus; N. mauritianus; N. taliliensis; N. trondlei)
Neaxius frankeae Lemaitre & Ramos, 1992
Neaxius frankeae Lemaitre & Ramos, 1992: 344, fig. 1; Lemaitre & León, 1992: 44; Hendrickx,
1995: 390 (list).
Type locality. — La Barra, Gorgona Island, Pacific coast of Colombia.
Distribution. — Only known from the type locality.
326
CRM 013 – Katsushi Sakai
Neaxius glyptocercus (Von Martens, 1869)
(figs. 61, 62)
Axius glyptocercus Von Martens, 1869: 613; Haswell, 1882: 165. [Type locality. — Cape York,
Australia.]
Axia acantha A. Milne-Edwards, 1878: 110. [Type locality. — New Caledonia.]
Eiconaxius acanthus — De Man, 1896: 491; De Man, 1898: 700, pl. 34 fig. 57-57a.
Fig. 61. Neaxius glyptocercus (Von Martens, 1869). A, anterior part of carapace and A2
peduncle; B, P2, lateral view; C, P4, lateral view; D, P5, lateral view; E, male Plp2. A-E, ZMB
2973, holotype, male (TL/CL, 82.0/ 26.4 mm), Cape York, Australia. Scales 1 mm.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
327
Fig. 62. Neaxius glyptocercus (Von Martens, 1869) (= Neaxius acanthus (A. Milne-Edwards,
1878)). A-D, anterior parts of carapace and A2 peduncles; E-F, P2, lateral view. A, ZMH
K 8392 (Godeffroy No. 7430), male (TL/CL, 44.0/14.3 mm), Viti-Levu Island; B, ZMH
K 41226 (Museum Godeffroy 16193), male (TL/CL, 88.0/27.3 mm), Tahiti; C, RMNH
42448, male (TL/CL, 94.0/30.6 mm), island of Barang Lompo, off Ujung Pandang, Sulawesi,
Indonesia, reefflat, in seagrass bed of Thalassia hemprichii Aschers; D, F, ZMH K 8411, female
(TL/CL, 55/1.07.1 mm), Palau Island; E, ZMH K 8392 (Godeffroy No. 7430), male (TL/CL,
44.0/14.3 mm), Viti-Levu Island. Scales 1 mm.
328
CRM 013 – Katsushi Sakai
Axius (Neaxius) acanthus — Borradaile, 1903: 537; Bouvier, 1915b: 196, fig. 7; De Man,
1925d: 3 (list), 10-12, 14 (key), 74; Holthuis, 1953b: 51; Poore & Griffin, 1979: 235, fig. 7;
Miyake, 1982: 192 (list); Tirmizi, 1983: 85, figs. 1, 2; Sakai, 1987a: 303 (list).
Eiconaxius taliliensis Borradaile, 1900: 420, fig. 15a-c. [Type locality. — Talili Bay, New
Britain.]
?Axius (Neaxius) glyptocercus — Borradaile, 1903: 537.
Axius acanthus var. mauritianus Bouvier, 1915: 196, fig. 7; Bouvier, 1914: 704; Fourmanoir,
1955: 31, fig. 4. [Type locality. — Mauritius.]
Axius (Neaxius) acanthus var. mauritianus — Bouvier, 1915b: 196, fig. 7.
Axius (Neaxius) glyptocercus — De Man, 1925a: 50, fig. 1; De Man, 1925d: 4 (list), 9, 11-13
(key); Poore & Griffin, 1979: 236, fig. 8.
Axius (Neaxius) acanthus var. mauritiana — De Man, 1925a: 50; De Man, 1925d: 3 (list), 10,
14 (key).
Neaxius glyptocercus — Sakai & De Saint Laurent, 1989: 30; Sakai, 1994: 176, 200 (key);
Tudge & Cunningham, 2002: 844; Tsang et al., 2008: 218, 219.
Neaxius acanthus — Sakai & De Saint Laurent, 1989: 30; Mukai & Sakai, 1992c: 47, figs. 1-3;
Sakai, 1994: 200 (key); Kensley et al., 2000: 212, figs. 5, 7F; Kensley, 2003: 383; Sakai &
Sawada, 2006: 1357; Tsang et al., 2008: 218, 219; Robles et al., 2009: 316.
Neaxius trondlei Ngoc-Ho, 2005: 59, figs. 6, 7. [Type locality. — Ua Huka, Hane Bay,
Marquesas Islands.]
Material examined. — ZMB 2973, holotype, male (TL/CL, 82.0/ 26.4 mm), Cape York, Australia, leg. Daemel; ZMH K 41226 (Museum Godeffroy 16193), 1 male (TL/CL, 88.0/27.3 mm),
Tahiti; ZMH K 8392 (Godeffroy No. 7430), 1 male (TL/CL, 44.0/14.3 mm), Viti-Levu Island.
Determined as E. acanthus: MNHN Th 812 (dry specimen), lectotype, female (TL/CL,
75.0/27.0 mm), New Caledonia; MNHN Th 190, paralectotype, 1 female (CL, 29.0; abdomen
and telson 48.0 mm), New Caledonia; MNHN Th 458, 1 female (TL/CL, 92.0/27.5 mm), Tuléar,
Madagascar, leg. et don. B. Thomassin; RMNH L 42446, 1 female (TL/CL, 95.0/26.1 mm);
RMNH L 42447, 1 male (TL/CL, 73.0/23.8 mm); RMNH L 42448, 1 male (TL/CL,
94.0/30.6 mm); RMNH L 42449, 1 male (TL/CL, 91.0/29.0 mm, rostrum missing); RMNH
L 42450, 1 male (TL/CL, 84.0/28.2 mm); RMNH L 42451, 1 female (TL/CL, 89.0/28.5 mm),
RMNH L 42452, 1 female (TL/CL, 90.0/26.8 mm), Island of Barang Lompo, off Ujung Pandang, Sulawesi, Indonesia, reefflat, in sea grass bed of Thalassia hemprichii, 16.xi.1992, leg.
P.L.A. Erfterneijer; ZMH K 8411, 1 female (TL/CL, 55.0/17.1 mm), Palau Island; SMF 31836
(BLT 18243), 4 males (TL/CL, 64.0/20.4-70.0/24.0 mm), Iriomote-jima, 13.vi.2003, leg. Y. Hirano et al., det. K. Sakai; SAM A10614, 1 male (TL/CL, 71.0/25.0 mm), Nosi Bé, Madagascar;
MNHN (?), 1 male (TL/CL, 41.0/14.0 mm), Lon-Rumphius 1973, Sta. Na-1, 19.i.1973, leg. D.
Sapri, Cat. No. Ca. 798, det. De Saint Laurent.
Determined as Neaxius trondlei Ngoc-Ho, 2005: MNHN Th 1419, holotype, male (TL/CL,
90.0/29.0 mm); MNHN Th, paratype, female (TL/CL, 66.0/22.0 mm), Hane bay, Ua Huka
Island, Marquesas Islands, MUSORSTOM 9, Sta. 19, with algae, corals, pebbles, 5.x.1997, leg.
R. von Cosel, J. Trondl & J. Tardy.
Diagnosis. — Rostrum bifurcate anteriorly, and dentate on lateral margins.
Anterolateral margin of carapace with or without spines between rostral base
and supraorbital spine. Dorsal margin of A2 segment 2 armed with one proximal spine or unarmed, and armed with 2-6 spines or unarmed laterally; scaphocerite with 1-5 ventral spines, A2 penultimate segment unarmed or armed with
three spines ventrally (fig. 62C, D). P2 fixed finger unarmed (fig. 62F) or den-
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
329
TABLE VII
Branchial formula of Neaxius glyptocercus (Von Martens)
Maxillipeds
Podobranch
Arthrobranch
Pleurobranch
Pereiopods
1
2
3
1
2
3
4
5
—
—
—
—
—
—
1
2
—
1
2
—
1
2
1
1
2
1
1
2
1
—
—
1
ticulate (fig. 62E) on the cutting edge. Abdominal somites 2-6, Plps2-5, and
tail-fan beset with thick setae. Male Plp1 absent; male Plp2 biramous, endopod with appendix interna, but without appendix masculina; male Plps3-5 endopods with appendix interna. Female Plp1 uniramous, consisting of proximal
segment and flagellum; Plps2-5 biramous, endopods with appendix interna.
Branchial formula as shown in table VII. Telson trapezoid, posterior margin
with median tooth, dorsal surface with two transverse carinae in anterior half.
Uropodal endopod broad with convex mesial margin, bearing three distinct
teeth on distolateral margin and median carina with two teeth including one on
distal margin. Uropodal exopod armed with a series of distinct teeth on broad
distal margin.
Remarks. — Poore & Griffin (1979: 235) compared N. acanthus with N.
glyptocercus, and mentioned that those two species are different from each
other on four points: (1) N. glyptocercus bears a spine on the anterior border of
the carapace between the rostrum and the supra-antennal spine (missing in N.
acanthus), (2) N. acanthus bears 2-4 spines along the cervical groove (missing
in N. glyptocercus), (3) the dorsal ridge with spine on article 2 of antenna 2 of
N. acanthus is armed near its base, mesially with one spine and laterally with
2-5 spines (N. glyptocercus is unarmed there), and (4) the lateral spination
on the meri of Ps1-2 is much more pronounced in N. acanthus than in N.
glyptocercus. However, the present examination of the specimens regarded as
N. acanthus from the Ryukyu Islands (SMF 31836) shows that the anterolateral
margin of the carapace between the rostral base and the supraorbital spine
is armed with a spine, though the dorsal margin of A2 segment 2 is armed
with one proximal spine and 2-5 lateral spines as mentioned by Poore &
Griffin (1979), and it was also found in the other specimens examined from
various localities that the anterolateral margin of the carapace between the
rostral base and the supraorbital spine is armed with one spine (fig. 62A,
male, ZMH K 8392; fig. 62B, male, ZMH 41226; fig. 62D, female, ZMH
K 8411), though usually unarmed; the dorsal margin of A2 segment 2 is
330
CRM 013 – Katsushi Sakai
armed with one proximal spine (fig. 62C, male, RMNH L 42448; fig. 62D,
female, ZMH K 8411) or unarmed proximally (fig. 61A, male, ZMB 2973, N.
glyptocercus; fig. 62A, male, ZMH 41226; fig. 62B, male, ZMH K 41226),
and with 2-6 spines laterally (fig. 62C, D) or unarmed laterally (fig. 62A, B);
the scaphocerite is armed with one ventral spine (fig. 61A, male, ZMB 2973,
N. glyptocercus), two ventral spines (fig. 61A, ZMB 8392; fig. 62B, ZMH K
41226), or five ventral spines (fig. 62C, male, RMNH 42448; fig. 62D, female,
ZMH K 8411); the A2 penultimate segment is smooth ventrally (fig. 61A,
male, ZMB 2973, N. glyptocercus; fig. 62A, male, ZMB 2972; fig. 62B, male,
ZMH K 41226), or armed with three ventral spines (fig. 62C, male, RMNH
42448; fig. 62D, female, ZMH K 8411); the P2 fixed finger is finely denticulate
on the cutting edge (fig. 61B, male, ZMB 2973, N. glyptocercus; fig. 62E,
male, ZMH K 8392), or unarmed (fig. 62F, ZMH K 8411), and the P2 merus is
denticulate on the ventral margin (fig. 62E, ZMH K 8392) or unarmed (fig. 62F,
female, ZMH K 8411), which suggests that the difference in spinulation is not
to be considered a specific characteristic, but as mere intraspecific variation.
Mukai & Sakai (1992: 47) confirmed, after examining 50 specimens of N.
acanthus from Papua New-Guinea, that the spinulation on the rostrum and
scaphocerite behind the cervical groove, and on the ventral margins of the
P1 merus is varied in morphology and number, and there are also a number
of variations in the burrow diameter of their dwelling holes, carapace length,
body length, and ash-free dry weight, so that N. acanthus (= N. acanthus var.
N. mauritiana Bouvier, 1914) is not different from N. glyptocercus, namely N.
acanthus is synonymous with N. glyptocercus.
Ngoc-Ho (2005) described Neaxius trondlei Ngoc-Ho, 2005 from the Marquesas Islands, mentioning that though N. trondlei is similar to N. glyptocercus
from Cape York, Australia in the morphology and spinulation of the rostrum
and gastric region, as well as in the morphology of the telson and uropods, it
differs from N. acanthus as follows: “(1) the second article of A2 peduncle is
armed near its base with one or two upper spines and unarmed laterally (same
article armed with one upper spine, four or five lateral spines in N. acanthus,
fig. 7L, M); (2) the third article of A2 peduncle bears a lower spine (unarmed
in N. acanthus fig. 7L); (3) one spine on the lower border of Mxp3 carpus
(three to five spines in N. acanthus); (4) the distal half of the lower border
of P1 merus and all lower border of P2 merus are unarmed (spines present
in N. acanthus); and (5) the telson with three well marked transverse carinae
equally distant from one another (distal carina often faint and apart from the
others in N. acanthus)”. However, the differences summed up by that author
are regarded as variations permissible at the species level. Regarding diffe-
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
331
rence (5), it would be better to consider as follows: one of those transverse
carinae developed very close along the posterior margin, is, though it is faint
or marked, to be regarded as a carinated posterior margin rather than as the
distal carina of the three carinae of these two species, because it is apart from
the others, and bear a median spine, so that their telsons are safely considered
to bear two transverse carinae on the dorsal surface.
Type locality. — Cape York, Australia (Von Martens, 1869).
Distribution. — Kenya — Mombasa (Sakai & De Saint Laurent, 1989);
Comoros — Mayotte, Moheli, and Anjouan (Fourmanoir, 1955); Madagascar
— Îles Glorieuses, Nosy Iranja, Nosy Bé, and Tuléar (Sakai & De Saint
Laurent, 1989); Mauritius — Port Louis and Le Chaland (Bouvier, 1914;
Sakai & De Saint Laurent, 1989); Indonesia — west Celebes (De Man,
1896, 1898); Papua New Guinea — Talili Bay, New Britain (Borradaile,
1900, as E. taliliensis), Motupore Island near Port Moresby (Mukai & Sakai,
1992c); Australia — Cape York, Queensland (Von Martens, 1869 as E.
glyptocercus), Fly Point, Cape York, Magnetic Island, Townsville, Bowen,
Norwest Island, Moreton Bay, Amity Point, and Stradbroke Island (Poore &
Griffin, 1979), Murray Island, Torres Strait (Poore & Griffin, 1979); Darwin,
Northern Territory (Sakai, 1994); Queensland (Poore & Griffin, 1979 as
E. glyptocercus); New Caledonia (A. Milne-Edwards, 1878; Sakai & De
Saint Laurent, 1989); Tahiti; Viti-Levu Islands, Republic of the Fiji Islands
(as E. glyptocercus by Godeffroy); Hane Bay, Ua Huka Island, and west
of Haamamao Bay, Marquesas Islands (Ngoc-Ho, 2005); Cebu Channel,
Philippines (Kensley et al., 2000); Mariana Islands — W. Susupe, Saipan
(Holthuis, 1953b); Pago Bay, Guam (Kensley, 2003); Kenting, Ping-Tong
County, southern Taiwan (Kensley et al., 2000); Japan — Ishigaki-jima (Sakai
& De Saint Laurent, 1989, as E. acanthus).
Neaxius mclaughlinae Ngoc-Ho, 2006
Neaxius mclaughlinae Ngoc-Ho, 2006a: 410, figs. 1, 2.
Material examined. — MNHN Th 1196, holotype, male (TL/CL, 74.0/24.0 mm), Gulf of
Guinea, Principe Island, Praia Pequena, 3.vii.1956, CALYPSO 1956, Sta. P 18, diving, 3-4 m;
MNHN Th 1195, paratype, male (TL/CL, 55.0/18.0 mm), 28.vi.1956, ibidem, Sta. P 13, diving,
3 m.
Remarks. — Neaxius mclaughlinae Ngoc-Ho, 2006a from the Gulf of
Guinea, and N. glyptocercus (Von Martens, 1869) from the Indo-West Pacific
are similar, but they differ in that in N. mclaughlinae the telson bears three
transverse carinae on the dorsal surface, whereas in N. glyptocercus it bears
two transverse carinae.
332
CRM 013 – Katsushi Sakai
Type locality. — Gulf of Guinea; Principe Island, Praia Pequena.
Distribution. — Only known from the type locality.
Neaxius vivesi (Bouvier, 1895)
Eiconaxius Vivesi Bouvier, 1895: 7.
Axius (Neaxius) Vivesi — De Man, 1925a: 56, fig. 2; De Man, 1925d: 4 (list), 14 (key).
Neaxius vivesi — Sakai & De Saint Laurent, 1989: 31; Hendrickx, 1995: 390 (list), 393, 397,
figs; Hendrickx et al., 2005: 171.
Remarks. — Neaxius vivesi from the eastern Pacific coast is closely similar
to N. frankeae from Gorgona Island, Pacific coast of Colombia, in that the
dorsal margins of the carpus and palm of the larger cheliped are armed with
spines, but they are different in that in N. vivesi the dorsal surface of the
telson bears three transverse carinae, of which the anterior and median ones
are straight but the posterior one is interrupted medially, and each half with
an anteriorly curved median end is also curved posteriorly at its midpoint (De
Man, 1925a, fig. 2), whereas in N. frankeae the dorsal surface of the telson
bears three transverse carinae, of which the anterior one is, however, straight
but posteriorly curved at both ends, and the median one is curved posteriorly
to surround halfway the posterior one, which is short and posteriorly curved at
both ends (Lemaitre & Ramos, 1992: 346, fig. 1).
Type locality. — Lower California.
Distribution. — Lower California (Bouvier, 1895; De Man, 1925a; Sakai &
De Saint Laurent, 1989); Gorgona Island, Pacific coast of Colombia (Lemaitre
& Ramos, 1992).
Genus Strahlaxius Sakai & De Saint Laurent, 1989
Strahlaxius Sakai & De Saint Laurent, 1989: 24; Poore, 1994: 100 (key); Sakai, 1994: 200
(key).
Diagnosis. — Rostrum bifurcate anteriorly, lateral margins denticulate, extending posteriorly onto gastric region. Anterolateral margins of carapace unarmed. Gastric region convex. Cervical groove distinct, but unarmed. Eyestalks subglobose; cornea pigmented. A2 scaphocerite simple and distinct. P1
unequal. P3 propodus setose, bearing no transverse rows of setae or spines.
Maxilla 2 scaphognathite with posterior whip. No pleurobranchs. Male Plp1
uniramous and unsegmented, with distal setae; male Plp2 biramous, endopod
bearing appendix interna with patch of hooklets distally, but no appendix masculina; male Plps3-5 biramous, endopods with appendix interna. Female Plp1
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
333
uniramous, consisting of proximal segment and flagellum; female Plps2-5 biramous, endopods with appendix interna. Telson wider than long, trapezoid,
and unarmed on lateral margin, tapering toward posterior margin, with median
spine or convexity, dorsal surface smooth. Uropodal endopod wider than long
and trapezoid, with acute distolateral angle. Uropodal exopod largely rounded
on distolateral margin, bearing proximolateral tooth, but no transverse suture
dorsally. [Adapted from Sakai & de Saint Laurent, 1989.]
Type species. — Axius plectrorhynchus Strahl, 1862a, designated by Sakai
& De Saint Laurent (1989). The gender of the generic name, Strahlaxius, is
masculine.
Species included. — Strahlaxius plectrorhynchus (Strahl, 1862a); S. waroona (Poore & Griffin, 1979).
K EY
TO THE
S PECIES
OF THE
G ENUS STRAHLAXIUS
1 – Rostrum triangular and bifurcate anteriorly, and armed irregularly with obtuse denticles
on lateral margins; gastric median carina short and marked. . . . . . . . .S. plectrorhynchus
– Rostrum stoutly triangular and rounded on anterior margin, and denticulate on lateral
margins; gastric median carina indistinct and shorter than in S. plectrorhynchus . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. waroona
Strahlaxius plectrorhynchus (Strahl, 1862)
Axius plectrorhynchus Strahl, 1862a: 1060, figs. 2-4, 11; Strahl, 1862b: 387; Miers, 1884b: 282;
De Man, 1888a: 251; De Man, 1888b: 463, pl. 19 fig. 5; Dworschak, 1992: 214.
Axius plectorhynchus [sic!] — Zehntner, 1894: 195.
Axius (Neaxius) plectorhynchus [sic!] — Borradaile, 1903: 537.
Axius (Neaxius) plectrorhynchus — De Man, 1925: 4 (list), 13 (key); McNeill, 1926: 304;
Estampador, 1937: 500; Poore & Griffin, 1979: 238, fig. 9; Sakai, 1984: 96.
Strahlaxius plectrorhynchus — Sakai & De Saint Laurent, 1989: 24; Sakai, 1994: 200 (key);
Poore, 2004: 178, fig. 48a, b, c.
Axius plectrohynchus [sic!] — Coleman, 1977: 135.
Not: Axius plectrorhynchus — Fulton & Grant, 1902: 60, pl. 5 figs. 7, 8 [= Strahlaxius waroona
(Poore & Griffin, 1979)].
Not: Axius plectorhynchus [sic!] — Hale, 1927a: 84, fig. 81; Hale, 1927b: 309 [= Strahlaxius
waroona (Poore & Griffin, 1979)].
Material examined. — ZMH K 41227, 1 male (TL/CL, 40.0/13.2 mm), 1 ovig. female
(TL/CL, 34.0/11.1 mm), Tahiti (Museum Godeffroy 16193); ZMB 1130 lectotype, ovig. female
(TL/CL, 21.0/7.2 mm), Luzon, Albay, leg. Jagor; MB 5617 (TL/CL, 16.0/5.2 mm), Luzon,
Philippines, leg. Jagor; ZMG 211, 1 male (TL/CL, 26.0/7.8 mm), 1 ovig. female (TL/CL,
29.0/8.7 mm), Bohol, Philippines, 4.vi.1863, leg. C. Semper; NHM 81.31, 1 female (TL/CL,
40.0/11.5 mm), Queensland, Portmolle Beach, between tide marks, v.1881, leg. De Coppingee.
Diagnosis. — Male Plp1 uniramous and unsegmented, with distal setae;
male Plp2 biramous, endopod with appendix interna, but without appendix
334
CRM 013 – Katsushi Sakai
masculina, male Plps3-5 biramous, endopods with appendix interna. Female
Plp1 uniramous, consisting of proximal segment and flagellum; Plps2-5 biramous, endopods with appendix interna. Telson wider than long and trapezoid, tapering toward posterior margin. Uropodal endopod trapezoid and wider
than long. Uropodal exopod without transverse suture. Eggs few in number but
large in size, measuring 0.6-0.7 mm in diameter.
Type locality. — Luzon, Philippines.
Distribution. — Philippines — (Dworschak, 1992), Luzon (Strahl, 1862b;
Estampador, 1937); Indonesia — Ambon (De Man, 1888b), Yapen Island,
Irian Jaya (Sakai & De Saint Laurent, 1989); New Caledonia (Sakai & De
Saint Laurent, 1989); Australia — Fly Point, Hayman Island, Norwest Islet,
Heron Island, Port Curtis, Sellicks Beach, Great Francis Island, and Kingscote
(Poore & Griffin, 1979), Heron Island, Queensland (Sakai, 1984; Sakai & De
Saint Laurent, 1989), Yorketown, S. Australia (Dworschak, 1992), Port Molle,
New South Wales, Beaumaris, Victoria, South Australia, Western Australia.
Strahlaxius waroona (Poore & Griffin, 1979)
Axius plectrorhynchus — Fulton & Grant, 1902: 60, pl. 5 figs. 7, 8. [Not Strahl, 1862b.]
Axius plectorhynchus [sic!] — Hale, 1927a: 84, fig. 81; Hale, 1927b: 309. [Not Strahl, 1862b.]
Axius (Neaxius) waroona Poore & Griffin, 1979: 240, fig. 10.
Strahlaxius waroona — Sakai & De Saint Laurent, 1989: 25; Sakai, 1994: 200 (key); Poore,
2004: 175, fig. 48d, e, f.
Type locality. — North side of Cape Naturaliste, Western Australia, mud
tube under intertidal zone.
Distribution. — Australia — Cape Naturaliste, Murry Island, Beaumaris,
and Flinders (Poore & Griffin, 1979); Tasmania — Jacobs Boat Harbour
(Poore & Griffin, 1979).
SUPERFAMILY CALLIANASSOIDEA DANA, 1852
Callianassidae Dana, 1852a: 12, 14, 19.
Callianassoidea — Sakai, 2005a: 1125; Sakai & Sawada, 2006: 1357.
Diagnosis. — Rostral carina present or not. Carapace with or without
cardiac sulcus(i), cardiac prominence, and dorsal oval. Linea thalassinica
present. Maxilla 2 scaphognathite with or without posterior whip. Uropodal
exopod without distal flap, and dorsal plate and lateral notch present or not.
Remarks. — Manning & Felder (1991: 766) included two families, Callianassidae and Ctenochelidae in the superfamily Callianassoidea, and distinguished them from each other by their key: male Plp2 with an appendix
masculina; uropodal exopod longitudinally carinate dorsally, lacking dorsal
plate; carapace usually with cardiac prominence (in Ctenochelidae); and
male Plp2 without an appendix masculina; uropodal exopod not longitudinally carinate dorsally, bearing dorsal plate; carapace without cardiac prominence (in Callianassidae). In their classification, Callianassidae included 4
subfamilies and 15 genera: Callianassinae (Biffarius, Callianassa, Calliapagurops, Neotrypaea, Notiax, and Trypaea); Cheraminae (Cheramus and
Scallasis); Callichirinae sensu nov. (Callichirus, Corallianassa, Glypturus,
Lepidophthalmus, and Neocallichirus); and Eucalliacinae (Calliax and Eucalliax); and Ctenochelidae included three subfamilies and 6 genera: Ctenochelinae (Ctenocheles, Gourretia, Paracalliax, and Dawsonius); Anacalliacinae
(Anacalliax); and Callianopsinae (Callianopsis).
However, it has turned out that their taxa are not in accordance with the
definitions of their characteristics. For example, Gourretia bears no cardiac
prominence, but was included in one of the subfamilies of the Ctenochelidae,
in which the cardiac prominence is present, and Callianopsis bears a dorsal
plate on the uropodal exopod, but was included in one of the subfamilies of
the Ctenochelidae, in which the dorsal plate is absent on the uropodal exopod.
Though Manning & Felder’s (1991) taxa comprise two families, Callianassidae (Callianassinae, Cheraminae, Callichirinae sensu nov., Eucalliacinae) and Ctenochelidae (Ctenochelinae, Anacalliacinae, and Callianopsinae),
and 7 subfamilies, the present system proposed here comprises 10 families and 8 subfamilies: Anacalliacidae, Bathycalliacidae (Bathycalliacinae and
336
CRM 013 – Katsushi Sakai
Vulcanocalliacinae), Callianassidae (Callianassinae and Callichirinae sensu
nov.), Callianopsidae (Callianopsinae and Neocallianopsinae subfam. nov.),
Ctenochelidae, Eucalliacidae (Calliapaguropinae and Eucalliacinae), Gourretiidae, Lipkecallianassidae (new status), Pseudogourretiidae (new status), and
Thomassiniidae.
The subfamilies Anacalliacinae Manning & Felder, 1991 and Callianopsinae Manning & Felder, 1991 were included in the Ctenochelidae by Manning
& Felder (1991), but in the present revision, the Anacalliacinae Manning &
Felder, 1991 are not included in the Ctenochelidae Manning & Felder, 1991,
because the carapace bears a dorsal oval, so that this taxon is elevated to family
level as Anacalliacidae.
The Callianopsinae Manning & Felder, 1991 are characterized as follows:
the carapace bears a rostral carina and a cardiac prominence as in Ctenochelidae, but the Mxp3 dactylus is oval, and the uropodal exopod bears neither
a dorsal plate nor a lateral notch, not as in the Ctenochelidae, in which the
Mxp3 dactylus is digitiform, and the uropodal exopod bears a lateral notch, so
that the Callianopsinae are elevated to family level and called Callianopsidae
now. In the Callianopsidae, however, two species, Callanopsis goniophthalma
(Rathbun, 1902) and C. anovalis Lin et al., 2007b are fundamentally different
from each other, because the carapace bears a dorsal oval in C. goniophthalma,
but lacks this in C. anovalis, so that C. anovalis is reclassified and included in
the genus Neocallianopsis gen. nov. in Neocallianopsinae subfam. nov., distinguished from C. goniophthalma in the Callianopsinae, so that those two
subfamilies are integrated under the Callianopsidae Manning & Felder, 1991.
The Callianassinae Dana, 1852a and the Callichirinae Manning & Felder,
1991, are included in the family Callianassidae, because the carapace bears
a dorsal oval, and the uropodal exopod bears a dorsal plate, though Mxp3
propodus and dactylus are slender in the Callianassinae, while the Mxp3
propodus is broadened and the dactylus is slender in the Callichirinae.
The Calliapaguropinae Sakai, 1999c and the Eucalliacinae Manning &
Felder, 1991 are characterized as follows: the carapace lacks a dorsal oval and
the uropodal exopod bears a dorsal plate, so that both subfamilies are included
in the family Eucalliacidae Manning & Felder, 1991, but they differ from each
other in that in the Calliapaguropinae the Mxp3 dactylus is digitiform, while
in the Eucalliacinae it is ovate.
The Bathycalliacinae Sakai & Türkay, 1999 and Vulcanocalliacinae
Dworschak & Cunha, 2007, are integrated under Bathycalliacidae Sakai &
Türkay, 1999, because the carapace lacks a dorsal oval, but bears a cardiac
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
337
prominence, the Mxp3 propodus is broadened, the dactylus is subtriangular,
and the uropodal exopod bears neither a dorsal plate nor a lateral notch, though
they are different from each other in that in the Bathycalliacinae the carapace
bears two transverse cardiac sulci, while in the Vulcanocalliacinae it lacks
transverse cardiac sulci.
Poore (1994) included 6 families (Laomediidae, Upogebiidae, Callianideidae, Thomassiniidae, Ctenochelidae, and Callianassidae) in the superfamily
Callianassoidea, but the Laomediidae and Upogebiidae are excluded from the
Callianassoidea, because the morphology of the propyloric ossicle of the gastric mill is different from that of the Callianassoidea and, therefore, they are
included in the superfamily Thalassinoidea. The Callianideidae are also excluded from the Callianassoidea and included in the Axioidea, because the
linea thalassinica is absent in Callianideidae, though it is present over a short
length in Paracallianidea.
In the present revision, the taxa of Callianassoidea and Axioidea are grouped
by the presence or absence of the dorsal oval on the carapace, and the dorsal
plate and lateral notch on the uropodal exopod, and also by the shapes of the
male Plps1-2 and Mxp3 propodus and dactylus.
In consequence, the historical taxa of the Callianassidae and Ctenochelidae
as established by Manning & Felder (1991) must fundamentally be changed
in this new way of classification. This holds especially for the genera of
the subfamily Callianassinae, because little attention had been paid before
to the male Plps1-2 and the uropodal exopod, which are now considered
very important and, therefore, helpful for the classification of the genera and
species, because of their distinct characteristics.
As a consequence, I have also had to reconsider my own earlier classification (Sakai, 2005b) and for that reason I am here giving the new scheme for
the superfamily Callianassoidea, in the following chapters.
Families included. — Anacalliacidae Manning & Felder, 1991; Bathycalliacidae Sakai & Türkay, 1999; Callianassidae Dana, 1852a; Callianopsidae
Manning & Felder, 1991; Ctenochelidae Manning & Felder, 1991; Eucalliacidae Manning & Felder, 1991; Gourretiidae Sakai, 1999a; Lipkecallianassidae
Sakai, 2005b (new status); Pseudogourretiidae Sakai, 2005b (new status); and
Thomassiniidae De Saint Laurent, 1979.
The new classification as here proposed in families, subfamilies, and genera
is as follows:
1. Anacalliacidae Manning & Felder, 1991 (genera Anacalliaopsis gen.
nov.; Anacalliax De Saint Laurent, 1973; Capecalliax gen. nov.)
Anacalliacidae
Bathycalliacidae
Bathycalliacinae
Dorsal
oval
+
Mxp3
propodus
subrectangular
Mxp3
dactylus
digitiform
Uropodal exopod
without dorsal plate, but with lateral notch
subrectangular
subtriangular
without dorsal plate or lateral notch
subrectangular
subtriangular
with dorsal plate, but without lateral notch
Callianassidae
Callianassinae
+
subovate or subrectangular
digitiform
with dorsal plate, but without lateral notch
Callichirinae
+
subquadrate
digitiform or subovate
with dorsal plate, but without lateral notch
Callianopsidae
Callianopsinae
+
subrectangular
ovate
without dorsal plate or lateral notch
−
subrectangular
ovate
without dorsal plate or lateral notch
Ctenochelidae
−
rectangular
digitiform
without dorsal plate, but with lateral notch
Eucalliacidae
Calliapaguropinae
−
subtriangular
digitiform
with dorsal plate; without lateral notch
subrectangular or subquadrate
subovate or subtriangular
with dorsal plate; without lateral notch
Neocallianopsinae
Eucalliacinae
−/+
Gourretiidae
−
rectangular
digitiform
without dorsal plate, with or without lateral notch
Lipkecallianassidae
−
rectangular
digitiform
without dorsal plate or lateral notch
Pseudogourretiidae
−
rectangular
digitiform
unknown
Thomassiniidae
−
rectangular
ovate
with or without dorsal plate, without lateral notch
CRM 013 – Katsushi Sakai
−
−
Vulcanocalliacinae
338
TABLE VIII
Four major characteristics of the families/subfamilies of the superfamily Callianassoidea
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
339
2. Bathycalliacidae Sakai & Türkay, 1999
i. Bathycalliacinae Sakai & Türkay, 1999 (genus Bathycalliax Sakai
& Türkay, 1999)
ii. Vulcanocalliacinae Dworschak & Cunha, 2007 (genus Vulcanocalliax Dworschak & Cunha, 2007)
3. Callianassidae Dana, 1852a
i. Callianassinae Dana, 1852a (genera Callianassa Leach, 1814;
Cheramoides gen. nov.; Cheramus Bate, 1888; Gilvossius Manning
& Felder, 1992; Notiax Manning & Felder, 1991; Trypaea Dana,
1852a)
ii. Callichirinae Manning & Felder, 1991 (genera Balsscallichirus gen.
nov.; Barnardcallichirus gen. nov.; Callichirus Stimpson, 1866;
Corallichirus Manning, 1992; Forestcallichirus gen. nov.; Glypturoides gen. nov.; Glypturus Stimpson, 1866; Grynaminna Poore,
2000; Lepidophthalmoides gen. nov.; Lepidophthalmus Holmes,
1904; Michaelcallianassa Sakai, 2002; Neocallichirus Sakai, 1988;
Podocallichirus Sakai, 1999c; Sergio Manning & Lemaitre, 1994
[sensu nov.]; Thailandcallichirus gen. nov.; Tirmizicallichirus gen.
nov.)
4. Callianopsidae Manning & Felder, 1991
i. Callianopsinae Manning & Felder, 1991 (genera Callianopsis De
Saint Laurent, 1973; Pleurocalliax gen. nov.)
ii. Neocallianopsinae subfam. nov. (genus Neocallianopsis gen. nov.)
5. Ctenochelidae Manning & Felder, 1991 (genus Ctenocheles Kishinouye, 1926)
6. Eucalliacidae Manning & Felder, 1991
i. Calliapaguropinae Sakai, 1999c (genus Calliapagurops De Saint
Laurent, 1973)
ii. Eucalliacinae Manning & Felder, 1991 (genera Andamancalliax
gen. nov.; Calliax De Saint Laurent, 1973; Calliaxina Ngoc-Ho,
2003 [sensu nov.]; Eucalliax Manning & Felder, 1991; Eucalliaxiopsis gen. nov.; Paraglypturus Türkay & Sakai, 1995; Pseudocalliax gen. nov.)
7. Gourretiidae Sakai, 1999a (genera Dawsonius Manning & Felder,
1991; Gourretia De Saint Laurent, 1973; Laurentgourretia Sakai,
340
CRM 013 – Katsushi Sakai
2004b; Paracalliax De Saint Laurent, 1979; Paragourretia Sakai,
2004b)
8. Lipkecallianassidae Sakai, 2005b (new status) (genus Lipkecallianassa
Sakai, 2002)
9. Pseudogourretiidae Sakai, 2005b (new status) (genus Pseudogourretia
Sakai, 2005b)
10. Thomassiniidae De Saint Laurent, 1979 (genera Crosniera Kensley
& Heard, 1991; Garyia gen. nov.; Heardaxius gen. nov.; Mictaxius
Kensley & Heard, 1991; Thomassinia De Saint Laurent, 1979)
K EY
TO THE
FAMILIES
OF THE
S UPERFAMILY C ALLIANASSOIDEA
1 – Maxilla 2 scaphognathite with posterior whip . . . . . . . . . . . . . . . . . . . . . . . Thomassiniidae
– Maxilla 2 scaphognathite without posterior whip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Carapace with cardiac sulcus(i) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Carapace without cardiac sulcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3 – Mxp3 propodus slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anacalliacidae
– Mxp3 propodus broadened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – Uropodal exopod without dorsal plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bathycalliacidae
– Uropodal exopod with dorsal plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucalliacidae
5 – Fingers of larger cheliped elongate and pectinate . . . . . . . . . . . . . . . . . . . . . Ctenochelidae
– Fingers of larger cheliped neither elongate nor pectinate . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – P4 elongate, overreaching P2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lipkecallianassidae
– P4 elongate, not overreaching P2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 – Pleurobranchs present on Ps2-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudogourretiidae
– Pleurobranchs absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 – Uropodal exopod with dorsal plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callianassidae
– Uropodal exopod without dorsal plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 – Mxp3 dactylus ovate or subovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callianopsidae
– Mxp3 dactylus digitiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gourretiidae
FAMILY ANACALLIACIDAE MANNING & FELDER, 1991
Anacalliinae Manning & Felder, 1991: 786 [misspelling for Anacalliacinae].
Diagnosis. — Rostrum triangular, with rostral carina. Carapace with dorsal oval, cardiac prominence, and linea thalassinica, but without cardiac sulcus. Eyestalks dorsoventrally flattened and contiguous. A2 scaphocerite small.
Mxp3 ischium-merus pediform, without meral distal spine; propodus subrectangular; and dactylus digitiform; exopod absent. P1 chelate, unequal in size
and dissimilar in shape, lacking meral hook in both chelipeds. P2 chelate. P3
propodus broadened. P4 subchelate. P5 chelate. Abdominal somite 6 lacking lateral projections. Male Plp1 uniramous and bisegmented. Male Plp2
(fig. 63A, B, C) biramous, endopod distally bearing distinct appendix masculina and small appendix, or only appendix interna. Uropodal exopod with
lateral notch.
Remarks. — The family Anacalliacidae Manning & Felder, 1991 is similar
to Callianassidae Dana, 1852a, because in both families the carapace bears a
dorsal oval and Mxp3 propodus and dactylus are slender. However, they are
different in that in Anacalliacidae the carapace bears a rostral carina, whereas
in the Callianassidae the carapace bears no rostral carina.
The Anacalliacidae include three genera, Anacalliax, Anacalliaopsis gen.
nov., and Capecalliax gen. nov. In Anacalliax the telson is wider than long and
slightly convergent posteriorly, bearing a median concavity on the rounded
posterior margin, the male Plp1 is uniramous and bisegmented, and the distal
segment is expanded distally with the complex distal margin as described and
figured by Biffar (1971b: 232, fig. 2l), though my present figure (fig. 63A) is
drawn from another direction. In Anacalliaopsis, the telson is slightly wider
than long, trapezoid, and convergent toward the almost straight posterior margin, and the male Plp1 is uniramous and bisegmented and chelate distally. In
Capecalliax the telson is wider than long, lateral margins convex, and posterior
margin broadly concave, the male Plp1 is uniramous and bisegmented, and the
distal segment is tapering distally.
Type genus. — Anacalliax De Saint Laurent, 1973.
Genera included. — Anacalliaopsis gen. nov.; Anacalliax De Saint Laurent,
1973; Capecalliax gen. nov.
342
CRM 013 – Katsushi Sakai
K EY
TO THE
G ENERA
OF THE
FAMILY A NACALLIACIDAE
1 – Telson wider than long, posterior margin straight; male Plp1 uniramous and bisegmented, distal segment chelate distally . . . . . . . . . . . . . . . . . . . . . Anacalliaopsis gen. nov.
– Telson wider than long, posterior margin concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Male Plp1 uniramous and bisegmented, distal segment expanded distally . . . Anacalliax
– Male Plp1 uniramous and bisegmented, distal segment tapering distally . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capecalliax gen. nov.
Genus Anacalliaopsis gen. nov.
Diagnosis. — Rostrum with rostral carina; carapace with dorsal oval. A1
peduncle reaching distal end of A2 penultimate segment. Mxp3 ischiummerus pediform, propodus subrectangular, and dactylus digitiform; exopod
absent. Male Plp1 uniramous and bisegmented, distal segment chelate distally;
male Plp2 biramous, endopod bearing mesiodistally appendices interna and
masculina; male Plps3-5 similar, and larger than and different from Plp2,
endopods with stubby appendix interna. Telson slightly wider than long,
trapezoid, and almost straight on posterior margin. Uropodal exopod broad,
with lateral notch.
Type species. — Callianassa agassizi Biffar, 1971b, by present designation
and monotypy. Gender of generic name, Anacalliaopsis, feminine.
Species included. — Anacalliaopsis agassizi (Biffar, 1971b).
Etymology. — A Greek suffix “opsis”, meaning “appearance” is added to
the generic name Anacalliax, from which the new genus is different, though in
gross appearance it is similar to that genus.
Anacalliaopsis agassizi (Biffar, 1971)
Callianassa agassizi Biffar, 1971b: 233, fig. 3.
Anacalliax agassizi — Manning, 1987: 397; Manning & Felder, 1991: 787 (list); Blanco
Rambla, 1995: 61-62, figs. 9-10; Sakai, 1999c: 127; Sakai, 2005b: 211; Tudge et al., 2000:
142.
Diagnosis. — Rostrum spinous, with rostral carina. Carapace with dorsal oval and cardiac prominence. Mxp3 ischium-merus pediform; propodus
subrectangular; dactylus digitiform; exopod absent. P1 unequal. Male Plp1
uniramous and bisegmented, distal segment chelate distally; male Plp2 biramous, endopod bearing mesiodistally appendices interna and masculina. Telson slightly wider than long, widest proximally, and trapezoid, bearing small
proximal lobe on lateral margin. Uropodal exopod with lateral notch. [Cf. Biffar, 1971b: 233, fig. 3f, k, l, m.]
Type locality. — Cumana, Venezuela.
Distribution. — Only known from the type locality.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
343
Genus Anacalliax De Saint Laurent, 1973
Anacalliax De Saint Laurent, 1973: 515; Manning, 1987: 397; Manning & Felder, 1991: 786,
figs. 2-3, 17; Sakai, 1999c: 126; Sakai, 2005b: 208.
Diagnosis. — Rostrum with rostral carina; carapace with dorsal oval. A1
peduncle slightly overreaching A2 penultimate segment. Mxp3 ischium-merus
pediform; propodus subrectangular; and dactylus digitiform; exopod absent.
Male Plp1 uniramous and bisegmented, distal segment expanded distally, with
complex distal margin; male Plp2 biramous, endopod bearing mesiodistally
appendices interna and masculina; male Plps3-5 similar, and larger than
and different from Plp2, endopods with stubby appendix interna. Telson
slightly wider than long, convergent posteriorly, posterior margin with median
concavity. Uropodal exopod with lateral notch.
Type species. — Callianassa argentinensis Biffar, 1971b, by original designation and monotypy. The gender of the generic name, Anacalliax, is feminine.
Species included. — Anacalliax argentinensis (Biffar, 1971b).
Anacalliax argentinensis (Biffar, 1971)
(fig. 63A-C)
Callianassa argentinensis Biffar, 1971b: 229, fig. 2.
Anacalliax argentinensis — Boschi, 1979: 137; Manning, 1987: 397 (list); Manning & Felder,
1991: 787 (list), figs. 2, 3, 17; Sakai, 1999c: 127; Tudge et al., 2000: 142 (list); Sakai, 2005b:
211.
Material examined. — USNM 133056, paratype, male (TL/CL, 12.0/4.3 mm), “Enfondedoarena” [= En fondo de arena = deep [in] sand bottom], Bajo de Los Huesos, Argentina,
South Atlantic, 43◦ 15′ S 64◦ 54′ W, 13 m, 26.vi.1963, det T. Biffar.
Diagnosis. — Rostrum with rostral carina. Carapace with dorsal oval and
cardiac prominence. Mxp3 ischium-merus pediform; propodus subrectangular; dactylus digitiform; exopod absent. Male Plp1 uniramous and bisegmented, distal segment expanded distally, with complex distal margin; male
Plp2 (fig. 63A-C) biramous, endopod elongate, bearing mesiodistally pestleshaped appendix masculina and small appendix interna. Telson slightly wider
than long, convex on lateral margins, and convergent toward posterior margin
with median concavity. Uropodal exopod with lateral notch. [Cf. Biffar, 1971b:
229, fig. 2g, k, I, m.]
Type locality. — Sandy beach on the north coast of Isla del Rey, Rio
Deseado, Province of Santa Cruz, Argentina, 47◦ 42′ S 66◦ W.
Distribution. — Only known from the type locality.
344
CRM 013 – Katsushi Sakai
Fig. 63. Anacalliax argentinensis (Biffar, 1971), Cheramus gaucho (Rodrigues & Manning,
1992) (syn. Poti gaucho), Notiax brachyophthalma (A. Milne-Edwards, 1870), and Callianopsis
goniophthalma (Rathbun, 1902). A, D, F, H, male Plp1; B, C, E, G, I, male Plp2, J, male
Plp3. A-C, Anacalliax argentinensis, USNM 133056, paratype, male (TL/CL, 12.0/4.3 mm),
“En fondo de arena” [= in/on sandy bottom], Bajo de Los Huesos, Argentina, South Atlantic,
43◦ 15′ S 64◦ 54′ W, 13 m; D, E, Cheramus gaucho, USNM 256376, paratype, male (CL, 4.0
mm), off Chui, near the border between Brazil and Uruguay, 33◦ 43′ S 51◦ 13′ W, 150 m; F,
G, Notiax brachyophthalma, USNM 252309, 1 male (TL/CL, 67.0/19.0 mm), River Maullin
estuary, Lalanguithue, Chili; H, I, J, Callianopsis goniophthalma (Rathbun, 1902), USNM
25238, type, male, damaged, off Point Conception, California, 278 fms [508 m].
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Genus Capecalliax gen. nov.
Diagnosis. — Rostrum triangular with rostral carina. Carapace with dorsal
oval and cardiac prominence; cardiac sulcus absent. Eyestalks flattened and
contiguous. Abdominal somite 6 lacking lateral projections. A2 scaphocerite
small. Mxp3 ischium-merus pediform; propodus subrectangular; and dactylus
digitiform; exopod absent. P1 chelate, unequal in size and dissimilar in shape,
lacking meral hook in both chelipeds. P2 chelate. P3 propodus broadened. P4
subchelate. P5 chelate. Male Plp1 uniramous and bisegmented, distal segment
tapering distally to form slender triangle; male Plp2 biramous, endopod
bearing mesiodistally appendices interna and masculina. Telson wider than
long, convex on lateral margins, and broadly concave on posterior margin.
Uropodal exopod with lateral notch.
Remarks. — Sakai (1999c, 2005b) redescribed Anacalliax pixii (Kensley,
1975), which had been included in Callianassa as C. pixii Kensley, 1975.
However, it has turned out that A. pixii is different from the type species of
Anacalliax, A. argentinensis, because in A. pixii the male Plp1 is uniramous
and bisegmented, and the distal segment is tapering distally to form a slender
triangle, the telson is distinctly wider than long, the lateral margins are convex,
and the posterior margin is broadly concave; whereas in A. argentinensis the
male Plp1 is uniramous and bisegmented, but the distal segment is expanded
distally with a complex distal margin, the telson is slightly wider than long,
the lateral margins are convergent posteriorly, and the posterior margin bears
a small median concavity. Therefore, A. pixii is excluded from Anacalliax
and reclassified under the new genus Capecalliax gen. nov. in the family
Anacalliacidae, as Capecalliax pixii.
Type species. — Callianassa pixii Kensley, 1975, by present designation
and monotypy. The gender of the generic name, Capecalliax, is feminine.
Species included. — Capecalliax pixii (Kensley, 1975).
Etymology. — The new generic name is a combination of Calliax and the
type locality, the Cape Province, South Africa.
Capecalliax pixii (Kensley, 1975)
Callianassa pixii Kensley, 1975: 53, figs. 4A-H, 5A-K; Tudge et al., 2000: 143.
Anacalliax pixii — Sakai, 1999c: 127; Sakai, 2005b: 210.
Diagnosis. — Rostrum triangular, with rostral carina. Carapace with dorsal oval. A1 peduncle about as long as A2 peduncle. Mxp3 ischium-merus
pediform; propodus subrectangular; dactylus digitiform; exopod absent. Male
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Plp1 uniramous and bisegmented, distal segment tapering to form slender triangle; male Plp2 biramous, endopod bearing mesiodistally appendices interna
and masculina. Telson wider than long, lateral margins convex, and posterior
margin broadly concave. Uropodal exopod with lateral notch. [Adapted from
Kensley, 1975.]
Type locality. — Kowie River estuary, Cape Province, South Africa.
Distribution. — Only known from the type locality.
FAMILY BATHYCALLIACIDAE SAKAI & TÜRKAY, 1999
Bathycalliacinae Sakai & Türkay, 1999: 204.
Diagnosis. — Rostrum triangular, without rostral carina. Carapace with cardiac prominence and linea thalassinica, with or without narrow dorsomedian
carina and two transverse cardiac sulci, and without dorsal oval. Eyestalks flattened, rhombic, and contiguous; cornea absent. Abdominal somite 6 without
lateral projections. A2 peduncle longer than A1 peduncle, scaphocerite developed as a small process. Maxilla 2 scaphognathite without posterior whip.
Mxp3 ischium-merus subpediform; propodus broadened; dactylus subtriangular; exopod rudimentary, or absent. P1 chelate, unequal in size, and dissimilar in shape, lacking meral hook in both chelipeds. P2 chelate, fingers longer
than palm. P3 propodus broadened; epipods present on Mxp3 and Ps1-4. Male
Plp1 present or absent. Male Plp2 biramous, endopod bearing distally appendix masculina with marginal setae, and mesiodistally small and slender appendix interna, or only with appendix interna; and exopod reaching distal end
of appendix masculina. Male Plps3-5 biramous and foliaceous, endopods with
appendix interna. Telson longer than wide, and rounded or straight on posterior
margin without median spine. Uropodal exopod with or without dorsal plate
in proximal half.
Subfamilies included. — Bathycalliacinae Sakai & Türkay, 1999; Vulcanocalliacinae Dworschak & Cunha, 2007.
K EY
TO THE
S UBFAMILIES
OF THE
FAMILY BATHYCALLIACIDAE
1 – Two transverse cardiac sulci and postcervical carina present . . . . . . . . . Bathycalliacinae
– Transverse cardiac sulcus and postcervical carina absent . . . . . . . . . . Vulcanocalliacinae
Subfamily BATHYCALLIACINAE Sakai & Türkay, 1999
Bathycalliacinae Sakai & Türkay, 1999: 204; Sakai, 2005b: 213.
Diagnosis. — Rostrum triangular, without rostral carina. Carapace with narrow dorsomedian carina, cardiac prominence, two transverse cardiac sulci,
and linea thalassinica, but without dorsal oval. Eyestalks flattened, rhombic,
and contiguous; cornea absent. Abdominal somite 6 without lateral projections. A2 peduncle longer than A1 peduncle, scaphocerite developed as a
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small process. Mxp3 ischium-merus subpediform; propodus broadened; dactylus subtriangular; exopod rudimentary. P1 chelate, unequal in size, and dissimilar in shape, lacking meral hook in both chelipeds. P2 chelate, fingers longer
than palm. P3 propodus broadened; epipods present on Mxp3 and Ps1-4. Male
Plp1 absent. Male Plp2 biramous, endopod bearing distally appendix masculina with marginal setae, and mesiodistally small and narrow appendix interna, and exopod reaching distal end of appendix masculina. Plps3-5 foliaceous and biramous, endopods with appendix interna. Uropodal exopod bearing neither dorsal plate nor lateral notch.
Remarks. — Dworschak & Cunha (2007) mentioned that “Vulcanocalliax
arutyunovi n. gen. & n. sp. is most closely related to Bathycalliax geomar” and
“It should be noted that only the male holotype of B. geomar (SMF 23866) has
Plp1 as descibed and figured by Sakai & Türkay (1999: 208, fig. 1). The male
paratype (SMF 23867) lacks any signs of Plp1.” However, it has already turned
out by the present re-examination of the type specimens of B. geomar Sakai
& Türkay, 1999 (SMF 23866, holotype, female (not male), TL/CL 57.0/13.0;
44◦ 40.146′ N 125◦ 06.685′ E, 625 m, R/V “Sonne”; SMF 23867, allotype (not
paratype), male, Aleutian subduction-zone off Oregon, U.S.A., 44◦ 40.193′ N
125◦ 06.605′ W, 627 m) that the holotype of B. geomar is not a male, but a
female, and the other male specimen is not the paratype but the allotype.
Considering that they described, as cited above, that the male holotype of B.
geomar has the male Plp1, one of the authors may have failed to make a careful
examination of the type specimens of B. geomar, and erroneously defined the
female holotype as a male, and the other male specimen as the paratype. The
fact is, that there are distinct differences between the two species: in B. geomar
the male Plp1 is absent; the male Plp2 is biramous, the endopod bears distally
an appendix masculina with marginal setae, and mesiodistally a small and
narrow appendix interna; and the exopod reaches the distal end of the appendix
masculina; whereas in V. arutyunovi the male Plp1 is present, vestigial and bisegmented; the male Plp2 is biramous, and the endopod bears an appendix
interna mesiomedially, but no appendix masculina.
They (Dworschak & Cunha, 2007) also mentioned that the subfamily Bathycalliacinae shares unique characteristics with the subfamily Vulcanocalliacinae as follows: epipods are present on Mxp3 and Ps1-4, and abdominal somite
1 bears anterolateral projections. However, those two subfamilies are clearly
different from each other as follows: in the type species of Bathycalliax, B.
geomar, the carapace bears two cardiac sulci and a dorsomedian carina, the
male Plp1 is absent, and the uropodal exopod lacks a dorsal plate; whereas in
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349
the type species of Vulcanocalliax, V. arutyunovi, the carapace bears neither
cardiac sulci nor a dorsomedian carina, the male Plp1 is vestigial and bisegmented, and the uropodal exopod bears a dorsal plate in the proximal half. [Cf.
Dworschak & Cunha, 2007, fig. 1g.]
Type genus. — Bathycalliax Sakai & Türkay, 1999.
Genus included. — Bathycalliax Sakai & Türkay, 1999.
Genus Bathycalliax Sakai & Türkay, 1999
Bathycalliax Sakai & Türkay, 1999: 204.
Diagnosis. — Carapace without dorsal oval, bearing narrow dorsomedian carina originating from gastric region and extending to posterior margin of carapace; cardiac prominence with median pit; two transverse cardiac
sulci present. Eyestalks rhombic and contiguous, without corneas, dorsal surfaces depressed. Maxilla 2 scaphognathite without long posterior whip. Mxp3
ischium-merus subpediform; propodus and dactylus broadened; exopod rudimentary. P1 unequal in size and dissimilar in shape. P2 chelate. P3 propodus
broadened. Ps4-5 subchelate. Epipods present at each base of Mxp3 to P4. Abdominal somite 6 without lateral projections. Male Plp1 absent; male Plp2 biramous, endopod bearing distally appendix masculina with marginal setae, and
mesiodistally small and slender appendix interna, and exopod reaching distal
end of appendix masculina; male Plps3-5 biramous, endopod with appendix
interna. Uropodal exopod oval, lacking dorsal plate. [Adapted from Sakai &
Türkay, 1999.]
Type species. — Bathycalliax geomar Sakai & Türkay, 1999, by original
designation. The gender of the name, Bathycalliax, is feminine.
Species included. — Bathycalliax geomar Sakai & Türkay, 1999.
Bathycalliax geomar Sakai & Türkay, 1999
Bathycalliax geomar Sakai & Türkay, 1999: 204, 1-3 figs., 1 tab.
Material examined. — SFM 23867, allotype, male (TL/CL, 48.0/11.5 mm) So-109/119
(44◦ 4.193′ N 125◦ 06.605′ W), Aleutian subduction-zone off Oregon, U.S.A., 627 m, GTV,
25.vi.1996, leg. R/V “Sonne”.
Diagnosis. — Carapace without dorsal oval, but with narrow dorsomedian
carina originating from gastric region and extending to posterior margin of
carapace, cardiac prominence with median pit, and two cardiac sulci. Eyestalks
flattened, cornea absent. Mxp3 ischium-merus subrectangular; propodus subrectangular; dactylus subtriangular; exopod rudimentary. P1 unequal. Telson
subquadrate, and posterior margin truncate. Uropodal exopod ovate, lacking
dorsal plate. [Cf. Sakai & Türkay, 1999: 203, figs. 1f, g.]
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Remarks. — The holotype, SFM 23866, of the present species, Bathycalliax
geomar, which was described in 1999 (Sakai & Türkay, 1999) as a male,
has turned out to be a female, because the Plp1 is uniramous, consisting of
a proximal segment and a flagellum.
Type locality. — Aleutian subduction-zone off Oregon, U.S.A.,
44◦ 40.146′ N 125◦ 06.685′ W, 625 m.
Distribution. — Only known from the type locality.
Subfamily V ULCANOCALLIACINAE Dworschak & Cunha, 2007
Vulcanocalliacinae Dworschak & Cunha, 2007: 37.
Diagnosis. — Rostrum triangular. Carapace with cardiac promincence and
linea thalassinica, but without dorsal oval, low dorsomedian carina, and transverse cardiac sulcus. Eyestalks flattened and contiguous; cornea absent. Mxp3
ischium-merus subrectangular, lacking meral tooth; propodus subrectangular;
dactylus subtriangular; exopod absent. P1 chelate, unequal in size, and dissimilar in shape, lacking meral hook in both chelipeds. Epipods present on
Mxp3 and Ps1-4. Male Plp1 vestigial and bisegmented. Male Plp2 biramous,
endopod bearing appendix interna with hooklets distally, but without appendix
masculina. Male Plps3-5 biramous and foliaceous, endopods bearing appendix
interna. Telson longer than wide, slightly narrowing posteriorly, and almost
straight on posterior margin. Uropodal exopod elevated to form dorsal plate in
lateral half. [Adapted from Dworschak & Cunha, 2007.]
Type genus. — Vulcanocalliax Dworschak & Cunha, 2007, as the nominate
genus.
Genus included. — Only the genus Vulcanocalliax Dworschak & Cunha,
2007.
Genus Vulcanocalliax Dworschak & Cunha, 2007
Vulcanocalliax Dworschak & Cunha, 2007: 37.
Diagnosis. — Rostrum triangular. Carapace with cardiac prominence, but
without dorsal oval, low dorsal carina, cardiac prominence, and transverse
cardiac sulci. Eyestalks flattened and contiguous; cornea absent. Abdominal
somite 6 without lateral projections. A2 peduncle longer than that of A1,
scaphocerite developed as a small process. Mxp3 ischium-merus subrectangular, lacking meral tooth; propodus subrectangular; dactylus subtriangular.
P1 chelate, unequal in size, and dissimilar in shape, lacking meral hook in both
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351
chelipeds. P2 chelate. P3 propodus broadened. Male Plp1 vestigial and bisegmented. Male Plp2 biramous, endopod bearing appendix interna with small
hooklets distally, but no appendix masculina. Male Plps3-5 biramous and foliaceous, endopods bearing appendix interna. Uropodal endopod oval. Uropodal exopod oval, and elevated in lateral half to form dorsal plate, the edge
of which is lined submarginally with short, thick spiniform setae and simple
setae, curved and descending to distal margin, with anterior part of plate depressed, and lacking lateral notch. [Adapted from Dworschak & Cunha, 2007.]
Type species. — Vulcanocalliax arutyunovi Dworschak & Cunha, 2007.
The gender of the generic name, Vulcanocalliax, is feminine.
Species included. — Only the type species, Vulcanocalliax arutyunovi
Dworschak & Cunha, 2007.
Vulcanocalliax arutyunovi Dworschak & Cunha, 2007
Vulcanocalliax arutyunovi Dworschak & Cunha, 2007: 37, figs. 1-5, table 1; Robles et al., 2009:
317.
Type locality. — Atlantic Ocean, Gulf of Cádiz, Captain Arutyuno mud
volcano, 35◦ 39.805′ N 07◦ 19.997′ W, 1339 m.
Distribution. — Only known from the type locality.
FAMILY CALLIANASSIDAE DANA, 1852
Callianassidae Dana, 1852a: 12, 14, 19; Dana, 1852b: 508; ICZN, 1987: 9; Manning & Felder,
1991: 766; Poore, 1994: 101; Sakai, 1999c: 7; Sakai, 2005b: 9; Sakai & Sawada, 2006: 1357;
Poore, 2007: 168.
Cheraminae Manning & Felder, 1991: 780; Tudge et al., 2000: 136.
Diagnosis. — Rostrum either weakly or well developed, and unarmed laterally, lacking rostral carina. Carapace with dorsal oval and linea thalassinica;
cardiac prominence and cardiac sulcus present or absent. Eyestalks usually
flattened. A2 scaphocerite reduced. Median tooth of urocardiac ossicle simple, with smooth posterior surface. Maxilla 2 scaphognathite without posterior
whip. Mxp3 ischium-merus pediform, subpediform, suboperculiform, or operculiform; propodus slender or broadened; and dactylus digitiform or ovate.
P1 chelate, equal, subequal, or unequal in size, and similar or dissimilar in
shape; larger cheliped with or without proximal meral hook, palm rectangular; and fingers moderate in length and not pectinate. P2 chelate. P3 propodus
often broadened to form heel-shape, or rectangular. Abdominal somites 3-5
with tuft of setae laterally. Male Plp1 present or absent; male Plp2 present
or absent, when present, endopod bearing both appendices interna and masculina, either appendix interna or appendix masculina [or neither appendices
interna nor masculina]. Male Plps1-2, when present, smaller than Plps3-5. Female Plp1 uniramous; female Plps2-5 biramous and similar in shape, endopods
with or without appendix interna. Uropodal exopod with dorsal plate.
Remarks. — In Manning & Felder’s (1991) classification, three subfamilies
(Anacalliacinae, Callianopsinae, and Ctenochelinae, in which the genus Gourretia was included) were included in the Ctenochelidae, though Gourretia was
later elevated to subfamily level as Gourretiinae by Sakai (1999a). However,
the Anacalliacinae and Callianopsinae are not included in the Ctenochelidae,
because they bear a dorsal oval on the carapace, whereas the Ctenochelinae
bear no dorsal oval on the carapace. In the present study the Anacalliacinae are
elevated to family level as Anacalliacidae, the Callianopsinae as Callianopsidae, and the Gourretiinae as Gourretiidae.
The subfamily Cheraminae was defined as “Carapace with dorsal oval.
Mxp3 pediform, propodus and dactylus slender. Male Plps3-5 with finger-like
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appendices internae” (Manning & Felder, 1991: 780), and the subfamily Callianassinae was defined as “Carapace with dorsal oval. Mxp3 pediform, subpediform, or operculiform, propodus and dactylus slender, Plps3-5 with stubby
appendices internae” (Manning & Felder, 1991: 767), which means that the
difference between Cheraminae and Callianassinae is only whether or not the
Plps3-5 appendices internae are finger-like (in Cheraminae) or stubby (in Callianassinae). However, such a difference is not characteristic enough to make
a clear distinction between the two subfamilies, because the present examination of the specimens of those subfamilies shows that Plps3-5 appendices internae vary in form from stubby to finger-like, and are not always distinguished
by the definition of finger-like or stubby, for example, in Trypaea costaricensis (Sakai, 2005b), Trypaea brevirostris (Sakai, 2002), Trypaea tabogensis (Sakai, 2005b), Trypaea lewtonae (Ngoc-Ho, 1994), Trypaea amplimaxilla
(Sakai, 2002), Trypaea nieli (Sakai, 2002), and Trypaea thailandica (Sakai,
2005b), all of which are included in the Callianassinae, the Plps3-5 endopods
bear a finger-like, projecting appendix interna, though Trypaea was defined
by Manning & Felder as “Plps3-5 are foliaceous and biramous in both sexes,
with stubby, projecting appendices internae” (Manning & Felder, 1991: 774).
Therefore, the Cheraminae are not differentiated from the Callianassinae, so
that Cheraminae is safely considered synonymous with Callianassinae.
According to Manning & Felder (1991: 767-781), the subfamily Callianassinae is based on the type genus Callianassa Leach, 1814, which is characterized by “Carapace with dorsal oval. Mxp3 pediform, sub-pediform, or
operculiform, propodus and dactylus slender. Plps3-5 with stubby appendices
internae”. The subfamily Cheraminae is based on the type genus Cheramus
Bate, 1888, which is characterized by “Carapace with dorsal oval. Mxp3 pediform, propodus and dactylus slender. Plps3-5 with finger-like appendices internae”. The subfamily Callichirinae sensu nov. is based on the type genus
Callichirus Stimpson, 1886, which is characterized by “Carapace with dorsal oval. Mxp3 subpediform or operculiform, propodus ovate, dactylus slender. Plps3-5 with stubby appendices internae”. The subfamily Eucalliacinae,
finally, is based on the type genus Eucalliax, which is characterized by “Carapace lacking dorsal oval. Mxp3 propodus and dactylus ovate. Plps3-5 with
finger-like appendices internae”. However, it is hardly possible to distinguish
the four subfamilies from one another by the characteristics shown by Manning & Felder (1991), because such characteristics are not always decisive in
the classification, as follows. (1) Mxp3 ischium-merus is pediform, subpediform, or operculiform (subfamilies Callianassinae, Callichirinae sensu nov.,
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
355
and Eucalliacinae); (2) Mxp3 propodus and dactylus are slender (subfamily
Callianassinae); (3) Mxp3 propodus is broadened, and Mxp3 dactylus is subtriangular or ovate (subfamilies Callichirinae sensu nov. and Eucalliacinae). This
confused system was analysed in the world-wide revision of the Callianassidae by Sakai (Sakai, 1999c, 2005b), who divided the Callianassidae into
two groups. The first group, in which the carapace bears a dorsal oval, comprised five subfamilies (Callianassinae, Callichirinae sensu nov., Eucalliacinae,
Calliapaguropinae, and Anacalliacinae), and the second group, in which the
carapace lacks a dorsal oval, included three subfamilies (Lipkecallianassinae,
Bathycalliacinae, and Paracalliacinae). The Callianassinae included only one
genus, Callianassa, because in the Callianassinae the Mxp3 ischium-merus, P3
propodus, Plps3-5 appendices internae, and uropodal exopod have a variety of
intermediate forms, which made it difficult to determine the taxonomic positions of these callianassid species. In the present revision, the Callianassinae
are classified into 6 genera by the types of Plps1-2. In the first group, Plps1-2
are absent (genus Gilvossius); in the second group Plp1 is present, but Plp2
is absent (genus Trypaea); in the third group Plps1-2 are present (genera Callianassa, Cheramus, and Notiax); and in the fourth group Plp1 is present and
Plp2 is rudimentary (Cheramoides gen. nov.). As I mentioned, male Plps1-2
are considered morphologically important in the classification of the callianassid genera, and are also considered to have no relation with other morphological characteristics such as the forms of Mxp3 and telson (Sakai, 2005b:
24-25).
The genus Callianassa Dana, 1852a is distinctly different from the other
genera included in the subfamily Callianassinae (table IX). In the type species
TABLE IX
Features of male Plps1-2 in the genera of the subfamily Callianassinae
Male Plp1
Male Plp2
Callianassa sensu nov.
uniramous, bisegmented
uniramous, protopod rarely protruded
short distolaterally
Cheramus sensu nov.
absent or uniramous,
bisegmented
uniramous, bisegmented
absent
uniramous, unsegmented, or bisegmented
uniramous, unsegmented, bisegmented, or
trisegmented
biramous, slender exopod and endopod
Cheramoides gen. nov.
Gilvossius sensu nov.
Notiax sensu nov.
Trypaea sensu nov.
rudimentary and hooklet-shaped
absent
uniramous, unsegmented, or bisegmented
absent
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of Callianassa, C. subterranea (Montagu, 1808), the male Plp1 is uniramous
and bisegmented, the male Plp2 is uniramous, the protopod is rarely protruded
short distolaterally, whereas in the other genera of the Callianassinae, male
Plps1-2 are varied in shape, and are different from those of Callianassa as
follows. In Cheramus the male Plp1 is absent or uniramous and bisegmented;
male Plp2 is biramous, bearing a slender exopod and endopod. In Notiax the
male Plp1 is uni- or biramous; the male Plp2 is uniramous, and unsegmented
or bisegmented. In Cheramoides gen. nov. the male Plp1 is uniramous and
bisegmented; the male Plp2 is rudimentary. In Trypaea the male Plp1 is uni-,
bi-, or tri-segmented, and the male Plp2 is absent. In Gilvossius male Plps1-2
are absent.
The genus Podocallichirus Sakai, 1999c was established on the base of
the type species Podocallichirus madagassus (Lenz & Richters, 1881), and
included P. balssi (Monod, 1933); P. foresti (Le Loeuff & Intès, 1974);
P. guineensis (De Man, 1928); P. tenuimanus (De Saint Laurent & Le Loeuff,
1979); P. gilchristi (Barnard, 1946); P. madagassus (Lenz & Richters, 1881);
P. masoomi (Tirmizi, 1970); and P. tamakii (Poore, 2000). However, their male
Plps1-2 are too varied in form to be integrated in one genus Podocallichirus,
so that in the present revision, those species which were included in Podocallichirus are grouped into 6 genera, and reclassified as follows. Podocallichirus,
based on P. madagassus, in which the male Plp1 is uniramous and bisegmented, and the distal segment is short and chelate distally; the male Plp2
is biramous, and the endopod bears mesiodistally a fusiform appendix masculina with a small appendix interna in its middle. Balsscallichirus gen. nov.,
based on P. balssi, in which the male Plp1 is uniramous and bisegmented;
and the male Plp2 is uniramous, lacking appendices masculina and interna.
Barnardcallichirus gen. nov., based on P. gilchristi, in which the male Plp1
is uniramous and bisegmented, and the distal segment is longitudinally furrowed, mesial part rounded distally, and lateral part sharply protruded distally
over the mesial part; and the male Plp2 is uniramous and unsegmented, lacking appendices masculina and interna. Forestcallichirus gen. nov. based on
P. foresti, in which the male Plp1 is uniramous and bisegmented, and the distal
segment is longer than the proximal and w-shaped distally; and the male Plp2
is uniramous and obscurely bisegmented. Grynaminna Poore, 2000, based on
G. tamakii Poore, 2000, in which the male Plp1 is uniramous and bisegmented,
and the distal segment is chelate distally; the male Plp2 is biramous, and the
endopod distally bears a subsquare appendix masculina with setae distally and
a small appendix interna mesiomedially. Tirmizicallichirus gen. nov. based on
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357
P. masoomi, in which the male Plp1 is uniramous and bisegmented, and the
distal segment is obtuse distally; the male Plp2 is biramous, and the endopod
is obtuse distally, bearing no appendices interna or masculina.
In this revision, the new way of classification by applying the difference
in shape of the Mxp3 propodus, and the status of male Plps1-2, has made
it possible to distinguish the subfamilies from each other more clearly than
ever. In the present classification, the family Callianassidae comprises two
subfamilies (Callianassinae and Callichirinae sensu nov.), the Callianassinae
includes 6 genera with 89 species and 3 spp. inq., and Callichirinae sensu nov.
includes 16 genera with 78 species and 1 sp. inquirenda.
Nominate subfamily. — Callianassinae Dana, 1852a.
Subfamilies included. — Callianassinae Dana, 1852a; Callichirinae Manning & Felder, 1991 (sensu nov.).
K EY
TO THE
S UBFAMILIES
OF THE
FAMILY C ALLIANASSIDAE
1 – Mxp3 propodus slender, subovate in shape; male Plp1 uniramous and simple distally,
or absent; Plp2 uniramous or biramous, endopod without appendix interna or appendix
masculina, or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callianassinae
– Mxp3 propodus broadened, subquadrate in shape; male Plp1 uniramous and usually
chelate distally, or absent; Plp2 uniramous or biramous, endopod bearing both appendices interna and masculina, either appendix interna or appendix masulina, or neither
appendix interna nor appendix masculina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callichirinae
Subfamily C ALLIANASSINAE Dana, 1852 (sensu nov.)
Callianassinae Dana, 1852a: 10; Bouvier, 1940: 100; Balss, 1957: 1582; Băcescu, 1967: 227;
De Saint Laurent, 1973: 514; De Saint Laurent, 1979: 1395; Manning & Felder, 1991: 767;
Sakai, 1999c: 10.
Cheraminae Manning & Felder, 1991: 780; Tudge et al., 2000: 136. [Type genus, Cheramus
Bate, 1888.]
Diagnosis. — Rostrum triangular, small, or obsolete, lacking rostral carina.
Carapace with dorsal oval, but without cardiac prominence or cardiac sulcus.
Eyestalks flattened and contiguous. Abdominal somite 6 lacking lateral projections. A2 scaphocerite developed as a small process. Mxp3 ischium-merus
varying from pediform to operculiform; propodus slender, convex or straight
on ventral margin; dactylus digitiform. P1 chelate, unequal or subequal in size,
and similar or dissimilar in shape; larger cheliped with or without meral hook.
P2 chelate. P3 propodus rhombic or rectangular. Male Plp1 present or not,
when present, slender and uniramous and unsegmented, bisegmented, or trisegmented. Male Plp2 present or absent, when present, uniramous or biramous, and slender; lacking appendices interna and masculina. Plps3-5 bira-
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CRM 013 – Katsushi Sakai
mous and foliaceous, endopods bearing appendix interna in both sexes. Uropodal exopod with dorsal plate.
Remarks. — In the previous revision of the Callianassidae (cf. Sakai,
2005b), 10 genera from Callianassinae Dana, 1852a: Callianassa Leach,
1814; Trypaea Dana, 1852a; Biffarius Manning & Felder, 1991; Neotrypaea
Manning & Felder, 1991; Notiax Manning & Felder, 1991; Poti Rodrigues &
Manning, 1992; Gilvossius Manning & Felder, 1992; Nihonotrypaea Manning
& Tamaki, 1998; Necallianassa Heard & Manning, 1998; and Pseudobiffarius
Heard & Manning, 2000; and two genera from Cheraminae Manning & Felder,
1991: Cheramus Bate, 1888 and Scallasis Bate, 1888; were integrated into
one subfamily Callianassinae, because Mxp3 ischium-merus, rostrum, and
Plps1-2/3-5 (regardless of sex) are variable and often intermedate in form,
which prevented a clear distinction among the genera. In the present revision,
however, those genera are reclassified mainly by the types of their male
Plps1-2, which are very characteristic in the genera of the Callianassinae. The
classification of those genera is revised by the features of male Plps1-2, after
examining the following 16 genera and one subgenus:
Callianassa Leach, 1814; Gebios Risso, 1822; Gebius Agassiz, 1846;
Mesostylus Bronn & Roemer, 1852; Trypaea Dana, 1852a; Cheramus Bate,
1888; Scallasis Bate, 1888; Trypaea s. str. Borradaile, 1903; Biffarius Manning
& Felder, 1991; Gilvossius Manning & Felder, 1991; Neotrypaea Manning &
Felder, 1991; Notiax Manning & Felder, 1991; Poti Rodrigues & Manning,
1992b; Nihonotrypaea Manning & Tamaki, 1998; Necallianassa Heard &
Manning, 1998; Pseudobiffarius Heard & Manning, 2000 (cf. Sakai, 1999c:
11; Sakai, 2005b: 25). As a result, the following 6 genera: Callianassa;
Cheramus; Gilvossius; Notiax; Trypaea; and Cheramoides gen. nov., are
admitted as valid members of the subfamily Callianassinae.
Type genus. — Callianassa Leach, 1814.
Genera included. — Callianassa sensu nov. (not Leach, 1814); Cheramoides gen. nov.; Cheramus sensu nov. (not Bate, 1888); Gilvossius sensu
nov. (not Manning & Felder, 1992); Notiax sensu nov. (not Manning & Felder,
1991); and Trypaea sensu nov. (not Dana, 1852a).
K EY
TO THE
G ENERA
OF THE
S UBFAMILY C ALLIANASSINAE
1 – Male Plp2 absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Male Plp2 present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 – Male Plp1 absent . . . . . . . . . . . . . . . . . . Gilvossius (= Pestarella; Paratrypaea) sensu nov.
– Male Plp1 present, and uniramous, unsegmented, bisegmented, or tri-segmented . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trypaea (= Biffarius) sensu nov.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
359
3 – Male Plp2 rudimentary and hooklet-shaped . . . . . . . . . . . . . . . . . . Cheramoides gen. nov.
– Male Plp2 uniramous or biramous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – Mxp3 ischium-merus oval . . . . . . . . . . . . . . . . . . . . . . . Notiax (= Rayllianassa) sensu nov.
– Mxp3 ischium-merus subpediform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Male Plp2 biramous, endopod and exopod slender . . . . . . . . . . . . . . Cheramus sensu nov.
– Male Plp2 uniramous, protopod rarely protruded short distolaterally . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callianassa sensu nov.
Genus Callianassa Leach, 1814 (sensu nov.)
Callianassa Leach, 1814: 386, 400; Leach, 1816: 116; Latreille, 1817: 34; Desmarest, 1825:
205; H. Milne Edwards, 1837c: 130; H. Milne Edwards, 1838: 386; Bell, 1846: 217; Nicolet,
1849: 206; Dana, 1852a: 14; Dana, 1852b: 509; Heller, 1863: 201; Stalio, 1877: 105; Boas,
1880: 84; Ortmann, 1899: 1142; Alcock, 1901: 197; Borradaile, 1903: 544; Balss, 1914: 91;
Selbie, 1914: 100; Kemp, 1915: 252; Balss, 1916: 33; Pesta, 1918: 201; Schmitt, 1921: 116;
De Man, 1928b: 91; Stevens, 1928: 324; Bouvier, 1940: 100; Edmondson, 1944: 44; Barnard,
1950: 505; Vinogradov, 1950: 223; Holthuis, 1954b: 334; Hemming, 1958: 142; Williams,
1965: 100; Băcescu, 1967: 227; Biffar, 1971a: 648, figs. 1, 2; De Saint Laurent, 1973: 514; Le
Loeuff & Intès, 1974: 32; De Saint Laurent & Le Loeuff, 1979: 48; ICZN, 1987: 60; Sakai,
1987a: 303; Sakai, 1988: 57; Holthuis, 1991: 239; Manning & Felder, 1991: 767, figs. 14, 6, 8; Poore, 1994: 102; Sakai, 1999c: 11; Felder, 2001: 441; Sakai, 2002: 488; Ngoc-Ho,
2003: 465; Sakai, 2005b: 25; Poore, 2008: 172.
Callianassa s. str., Felder, 2001: 440.
Gebios Risso, 1822: 243. [Type species: Gebios davianus Risso, 1822 (= junior subjective
synonym of Cancer candidus Olivi, 1792, by monotypy). Gender of generic name, Gebios,
feminine.]
Gebius Agassiz, 1846: 160. [Emendation of Gebios Risso, 1822. Gender of generic name,
Gebius, masculine.]
Mesostylus Bronn & Roemer, 1852: 353. [Type species: Pagurus faujasi Desmarest, 1822, by
monotypy. Gender of generic name, Mesostylus, masculine.]
Diagnosis. — Eyestalks depressed. Mxp3 propodus subovate and dactylus
digitiform. Male Plp1 uniramous and bisegmented, and male Plp2 uniramous,
protopod rarely protruded short distolaterally, lacking appendices interna and
masculina; male Plps3-5 biramous, endopods with stubby, embedded appendix
interna. Uropodal exopod with dorsal plate.
Remarks. — The present genus, Callianassa is defined by the forms of male
Plps1-2.
Type species. — Cancer (Astacus) subterraneus Montagu, 1808, by original designation and monotypy. Gender of the generic name, Callianassa, feminine.
Species included. — Callianassa subterranea (Montagu, 1808) currently is
the only known species.
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CRM 013 – Katsushi Sakai
Callianassa subterranea (Montagu, 1808)
Cancer (Astacus) subterraneus Montagu, 1808: 88, pl. 3 figs. 1, 2.
Callianassa subterranea — Leach, 1814: 400; Leach, 1815: 343; Leach, 1816, pl. 32;
Desmarest, 1825, tab. 36, fig. 2; H. Milne Edwards, 1838: 388; Thompson, 1844: 268; Bell,
1846: 219 (not 217, misprint), fig.; White, 1847: 70; A. Milne-Edwards, 1870: 80, 101;
Stalio, 1877: 106; Neumann, 1878: 34; Barrois, 1882: 24; Czerniavsky, 1884: 76, 80; Carus,
1885: 489; Giard & Bonnier, 1890: 362 (= Gilvossius tyrrhenus (Petagna, 1792); Ortmann,
1891: 55, pl. 1 fig. 10 (partim?) (= Gilvossius tyrrhenus (Petagna, 1792)); Adensamer, 1898:
620; Lagerberg, 1908: 53, pl. 1 fig. 15; Bianco, 1909: 603; Blohm, 1913: 17 (partim);
Selbie, 1914: 101; Athanassopoulos, 1917: 32; Wiliamson, 1927: 445, figs. 209, 210; Perrier,
R., 1929: 197; Gustafson, 1934: 14; Nobre, 1936: 121, pl. 40 fig. 101; Lutze, 1938: 170,
figs. 28-51; Makarov, 1938: 62, 63 (partim); Poulsen, 1940: 229, figs. 10-12; Lutze, 1941:
34; Gordon, 1957: 240; Holthuis & Gottlieb, 1958: 115; Holme, 1961: 397; Bourdon, 1965:
1; Holme, 1966: 401; Allen, 1967: 18, 57, fig. 90; Christiansen, 1972: 41, fig. 49; Pastore,
1976: 107; De Saint Laurent & Božić, 1976: 17, figs. 1, 9, 17, 28; Moncharmont, 1979: 71;
Števčić, 1979: 282, fig. 1; Domenech et al., 1981: 150; Adema et al., 1982: 23, fig. 6a-c;
Christiansen & Greve, 1982: 213; Thessalou-Legaki, 1985: 457; Thessalou-Legaki, 1986:
182; García Socias & Massuti Jaume, 1987: 75; Witbaard & Duineveld, 1989: 209-219,
fig. 1; Moyse & Smaldon, 1990: 520, fig. 10-13; Števčić, 1990: 217; Dworschak, 1992: 203;
Hayward et al., 1995: 432, fig. 8.52; Stamhuis et al., 1997: 155; Christiansen & Stene, 1998:
75; Nickell et al., 1998: 733; Simboura et al., 1998: 300; Števčić, 1998: 652; Sakai, 1999c:
14 (key), 19; d’Udekem d’Acoz, 1999: 155; Christiansen, 2000: 234; Hughes et al., 2000:
189; Tudge et al., 2000: 143; Costello et al., 2001: 289; Ngoc-Ho, 2003: 468, figs. 9, 10;
Atkinson & Taylor, 2004: 45, tabs. 1, 2; Ingle & Christiansen, 2004: 93, figs. 69, 71; Sakai,
2005b: 33; Sakai & Sawada, 2006: 1357, figs. 7, 8.
Cancer (Astacus) subterraneus — Desmarest, 1825: 205, fig. 2, 2a-c.
Callianassa (Cheramus) subterranea — Borradaile, 1903: 545; De Man, 1928a: 6, pl. 1 fig. 11h; De Man, 1928b: 27, 91, 92, 94, 97; Makarov, 1938: 63, fig. 21; Bouvier, 1940: 101
(partim), fig. 67; Gurney, 1942: 243-244, fig. 98; Gurney, 1944: 83; Gordon, 1957: 249;
O’Céidigh, 1962: 163.
Cheramus subterraneus — Colosi, 1923: 6.
Callianassa pestae — Lutze, 1938, figs. 10-21 (= C. subterranea).
Callianassa algerica Lutze, 1938: 168 (part.), figs. 22-26 [not fig. 22, eyestalks = C. whitei],
26a, smaller Mxp3 = C. subterranea [not larger Mxp3 = C. candida], 26b, smaller
Mxp3 = C. subterranea [not larger Mxp3 = C. candida], [not fig. 27 = C. candida] [type
locality: Castiglione near Algiers, 36◦ 37′ N 02◦ 40′ E]; Lutze, 1941: 34 [nomen dubium].
Callianassa helgolandica Lutze, 1938: 174, figs. 52-61; Lutze, 1941: 34. [Type locality. —
Helgoland.]
Callianassa stebbingi — Schellenberg, 1928: 78, fig. 59; Lutze, 1938, figs. 10-21 [= Callianassa subterranea]; Abel et al., 1963: 279, fig. 94.
Callianassa hertlingi Lutze, 1941: 34 [a junior synonym of Callianassa helgolandica Lutze,
1938].
Callianassa tyrrhena [not Callianassa tyrrhena Petagna] — Holthuis & Gottlieb, 1958: 62
(partim), fig. 13 (= C. subterranea); Pérez, 1995: 136, fig. 78.
Callianassa (Callichirus) tyrrhena — Zariquiey Alvarez, 1968: 230.
Callianassa (Callianassa) subterranea — Zariquiey Alvarez, 1968: 229; Lagardère, 1973: 88,
94.
Callianassa subterranea? — Robles et al., 2009: 316.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
361
Callianassa subterranea australis Kensley, 1974: 271, figs. 3-5. [Type locality. — South Africa,
Lüderitzbucht, 180 m.]
Callianassa australis — De Saint Laurent & Le Loeuff, 1979: 51; Sakai, 1999c: 40.
Biffarius australis — Tudge et al., 2000: 42.
Not: Callianassa subterranea — H. Milne Edwards, 1837a: 309 [= Gilvossius tyrrhenus
(Petagna, 1792)]; H. Milne Edwards, 1837c: 130, pl. 48 fig. 3-3e; Heller, 1863: 202, pl. 6
figs. 9, 11; Ortmann, 1891: 55, pl. 1 fig. 10 [= Gilvossius tyrrhena (Petagna, 1792)];
Czerniavsky, 1868: 122; Giard & Bonnier, 1890: 362, figs. 1-3 [= C. pontica Czerniavsky,
1884 = Gilvossius candidus (Olivi, 1792)]; Adensamer, 1898: 620 [= Gourretia denticulata
(Lutze, 1937)].
Not: Callianassa — Kinahan, 1859: 266.
Not: Callianassa subterranea forma pontica Czerniavsky, 1884: 81 [nomen dubium] [= C. pontica Czerniavsky, 1884 = Gilvossius candidus (Olivi, 1792)].
Not: Callianassa subterranea var. minor Gourret, 1887: 1034; Gourret, 1888: 96, pl. 8 figs. 1-15
[= Gourretia denticulata (Lutze, 1937)].
Not: Callianassa subterranea var. japonica Ortmann, 1891: 56 [= Trypaea japonica].
Material examined. — SMF-36910, 3 males (TL/CL: 49.0/10.5-12.0/2.9 mm), 1 female
(20.0/4.6 mm), North Sea, German Bight, S. of Helgoland Trench (54◦ 7.4′ N 7◦ 56′ E), 38 m, Van
Veen grab, 8.ix.2004, R/V “Uthörn”; SMF 21879, 1 male (36.0/8.5 mm), North Sea, German
Bight (54◦ 39.90′ N 6◦ 00.00′ E), 42.5 m, 24.v.1987, F/C Senckenberg.
Diagnosis. — Mxp3 ischium-merus subpediform, merus obliquely convex
on distal margin. Male Plp1 uniramous and bisegmented; male Plp2 uniramous, protopod rarely protruded short distolaterally, lacking appendix interna.
Telson truncate on posterior margin with median spine. Uropodal exopod with
dorsal plate.
Remarks. — The male Plp2 can be described as “endopod fused with
protopod” as in the male specimens SMF 25800 from the Canary Islands
(not SMF 27250 from the Adriatic Sea as in Sakai, 2005b, fig. 7) and SMF
21879 from the North Sea, but possibly also as “endopod absent and hence
uniramous” because of the earlier stages of development as in the male
specimens in SMF 36910 from the North Sea, as shown by Ngoc-Ho (2003,
fig. 9G).
It is here confirmed from examining Kensley’s (1974) figures and description that “the close affinity between the European and the South African
species cannot be denied”, which implies indeed that Callianassa subterranea
australis Kensley, 1974 from South Africa, Lüderitzbucht, 180 m, is safely
synonymized with C. subterranea. So, the distribution of the present species,
C. subterranea distributed mostly in the North Atlantic Ocean, the Mediterranen Sea, and the central Atlantic Ocean, has now expanded further to South
Africa through the record of C. subterranea australis Kensley, 1974, just as in
the case of Calocaris barnardi Stebbing, 1914 from Cape Castle, South Africa,
362
CRM 013 – Katsushi Sakai
163 m, which is synonymized with Calocaris macandreae Bell, 1846 from the
North Atlantic in the present study (see p. 241).
Type locality. — Devon, England.
Distribution. — Eastern Atlantic from Scandinavia to Canary Islands and
the eastern Mediterranean; usually subtidal, 10-270 m. Most probably also off
the west coast of South Africa.
Genus Cheramoides gen. nov.
Diagnosis. — Eyestalks flattened. Mxp3 propodus subovate, and dactylus
digitiform. Male Plp1 uniramous, slender, and usually bisegmented; male Plp2
rudimentary and hooklet-shaped, lacking appendices interna and masculina;
Plps3-5 biramous, endopods with projecting, finger-like appendix interna.
Telson subsquare, slightly tapering toward posterior margin, and bearing
median concavity with spine. Uropodal exopod elongate, with dorsal plate.
Remarks. — It was found through the present examination of the specimens
that Cheramus marginatus Manning & Felder, 1991 is different from the type
species of Cheramus, C. profundus (Biffar, 1973) as follows: in C. marginatus
the male Plp2 is rudimentary and hooklet-shaped, whereas in C. profundus it
is biramous, bearing a slender exopod and endopod. Therefore, a new genus,
Cheramoides, is here established.
Type species. — Callianassa marginata Rathbun, 1901, by present designation and monotypy. Gender of the new generic name, Cheramoides, feminine.
Species included. — Cheramoides marginata (Rathbun, 1901).
Etymology. — The new generic name Cheramoides is composed of a
generic name, Cheramus and a Latin suffix, -oides, meaning “similar”.
Cheramoides marginata (Rathbun, 1901)
Callianassa marginata Rathbun, 1901: 92, fig. 15; Schmitt, 1935b: 4; Biffar, 1971a: 654, 689,
figs. 15, 16; Coelho & Ramos, 1973: 161; Rabalais et al., 1981: 100; Manning, 1987: 397;
Sakai, 1999c: 28; Sakai, 2005b: 46.
Callianassa (Callichirus) marginata — Borradaile, 1903: 547; Bouvier, 1925: 472; De Man,
1928b: 29, 94, 113; Schmitt, 1935a: 195, fig. 56.
?Callichirus marginatus — Bouvier, 1905: 804.
Cheramus marginatus — Manning & Felder, 1991: 780, figs. 6, 14; Blanco Rambla et al., 1994:
18, figs. 2, 3; Coelho, 1997: 150; Melo, 1999: 360, figs. 239-240; Tudge et al., 2000: 145;
Coelho et al., 2007: 5 (tab. 2); Heard et al., 2007: 24, fig. 18.
Material examined. — USNM 1071632, male (TL/CL, 19.0/4.4 mm), N.E. of Tintipan Island, Colombia, Caribbean Sea, 108-125 m, 9◦ 56.26′ N 75◦ 53.52′ W – 9◦ 56.35′ N 75◦′ 57.04′ W,
108-126 m, 7.viii.1995, leg. R. Lemaitre & N.H. Campos.
Diagnosis. — Mxp3 ischium-merus subsquare, merus straight on distal margin, and rectangular at distomesial corner. Male Plp1 uniramous, slender, and
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
363
bisegmented; male Plp2 rudimentary and hooklet-shaped; male Plps3-5 biramous, endopods with projecting, finger-like appendix interna. Telson subsquare, slightly tapering toward posterior margin, bearing median concavity
with spine. [Cf. Manning & Felder, 1991: 780, fig. 14b, g, e.]
Remarks. — Callianassa marginata was included in Cheramus by Manning
& Felder (1991), who described that the male Plp2 is absent. However, it was
found through the present examination that it is present, though rudimentary
and hooklet-shaped, which suggests that Callianassa marginata is not to
be included in Cheramus, but instead should be included in a new genus,
Cheramoides as here established.
Type locality. — Puerto Rico, Mayagüez harbour, 315 m.
Distribution. — Northeastern Gulf of Mexico; Arrowsmith Bank, Barbados
to Puerto Rico; Colombia (in Caribbean Sea); Amapá, Maranhão, Ceará,
Alagoas, Rio de Janeiro, Brazil; 55-640 m.
Genus Cheramus Bate, 1888 (sensu nov.)
Cheramus Bate, 1888: 30; Borradaile, 1903: 545 (partim); De Man, 1928b: 95; Gurney, 1944:
83; Manning & Felder, 1991 [18 Dec.]: 780 (partim) [not figs. 2, 4-6, 14 = Callianassa);
Holthuis, 1991 [19 Dec.]: 239; Poore, 1994: 101; Davie, 2002: 459.
Scallasis Bate, 1888: 34; Manning & Felder, 1991: 780.
Poti Rodrigues & Manning, 1992: 9. [Type species. — Poti gaucho Rodrigues & Manning,
1992.]
Diagnosis. — Eyestalks flattened. Mxp3 ischium-merus subpediform; propodus subovate, and dactylus digitiform. Male Plp1 absent or uniramous, slender, and usually bisegmented; male Plp2 biramous, endopod and exopod elongate, lacking appendices interna and masculina; male Plps3-5 biramous, endopods with projecting, finger-like appendix interna. Telson quadrate, subquadrate, trapezoid, or subovate. Uropodal exopod with dorsal plate.
Remarks. — Cheramus Bate, 1888 was based on the type species Callianassa (Cheramus) batei Borradaile, 1903, a replacement name for the preoccupied Cheramus occidentalis Bate, 1888 (cf. Manning & Felder, 1991:
780), but Callianassa batei is a subjective junior homonym of a fossil species,
Callianassa batei Woodward, 1868. Therefore, the type species of Cheramus
is Callianassa profunda Biffar, 1973 (a new name for Callianassa batei Borradaile, 1903).
Biffar (l973: 229) included five species of Callianassa in Cheramus: Callianassa sibogae De Man, 1905; C. propinqua De Man, 1905; C. modesta
De Man, 1905; C. intermedia De Man, 1905, and C. minima Rathbun, 1901.
Manning & Felder (1991) established the subfamily Cheraminae, and Tudge
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CRM 013 – Katsushi Sakai
et al. (2000: 136) showed that the Cheraminae are composed of three genera:
Poti including one species (P. gaucho Rodrigues & Manning, 1992b); Callianassa including 4 species (C. praedatrix (De Man, 1905), C. propinqua,
C. rectangularis Ngoc-Ho, 1991, C. sibogae); and Cheramus including two
species (C. profundus and C. marginatus (Rathbun, 1901)).
However, of the present taxa, Callianassa sibogae (cf. Trypaea) and C. intermedia De Man, 1905 (cf. Trypaea) are reclassified under the genus Trypaea,
because the male Plp2 is absent. Callianassa marginatus is now under Cheramoides gen. nov., because the male Plp2 is different from that in Callianassa
and Cheramus. C. minima, finally, is reclassified as Crosniera minima (Rathbun, 1901) under the family Thomassiniidae, because in Crosniera minima the
antennal region of the anterior carapace bears a vertical row of setal pits; the
male Plp1 is uniramous and bisegmented; and the distal segment is slightly expanded into a leaf-like extension; the male Plp2 is biramous, and the endopod
bears mesiomedially an appendix masculina articulating mesiomedially with a
small appendix interna. Poti gaucho is reclassified as a species of Cheramus,
because male Plps1-2 are the same as those of Cheramus.
The following two species, Callianassa (Calliactites) longicauda Sakai,
1967 and Callianassa (Cheramus) spinophthalma Sakai, 1970a, were included
for the first time in Cheramus by Tudge et al. (2000).
The following two species of Callianassa, Callianassa contipes Sakai, 2002
and Callianassa (Cheramus) lobetobensis De Man, 1905, are also included in
Cheramus, though male Plps1-2 are unknown, because the Mxp3 ischium and
merus and the P3 propodus of those two species, are similar in morphological
features to those of the species of Cheramus, such as C. nigroculata (Sakai,
2002) and C. malaccaensis (Sakai, 2002).
Manning & Felder (1991: 780) defined Scallasis in their diagnosis as
“Carapace with rostral spine, apparently lacking orbit. Cornea well-formed,
terminal, subglobular. Mxp3 with exopod, Plp elongate and slender, with
finger-like appendices internae.”, and they described in the remarks that “the
type specimen of S. amboinae was examined by one of us, who verified that it
is a member of the Cheraminae”. However, Sakai (1999c: 12) also examined
the type specimen of Scallasis amboinae (NHM 1888: 22), which is preserved
at the Natural History Museum, London, and confirmed that the Mxp3 exopod
is absent, though the figure of the type specimen drawn by Bate shows that an
Mxp3 exopod is present (Bate, 1888, pl. 2 fig. 3a). It turned out later that the
type specimen of Scallasis amboinae examined by Sakai is different from the
type specimen of Scallasis amboinae drawn by Bate, because they are different
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
365
from each other in the following features. In the type specimen drawn by Bate,
the eyestalks are globular (Bate, 1888: 34, pl. 2 fig. 3i), and the Mxp3 exopod
is present, whereas in the specimen preserved in the Natural History Museum,
London and examined by Sakai, the eyestalks are triangular and depressed
(Sakai, 1999c, fig. 5d), and the Mxp3 exopod is absent (Sakai, 1999c: 37,
fig. 5f). Those features shown by Sakai’s examination suggested that Scallasis
amboinae should be included in Callianassa, though the preserved specimen
in London is labelled Scallasis amboinae (Bate, 1888). However, it was also
noted that C. amboinae is different from C. subterranea, the type species
of Callianassa, in the shape of the telson: in C. amboinae the telson is
concave medially on the posterior margin, without a median spine, whereas
in C. subterranea the telson is convex medially on the posterior margin, with
a median spine. In the present study, C. amboinae (Bate, 1888) is reclassified
under the genus Trypaea, because the male Plp2 is absent (Sakai, 1999c: 37).
Type species. — Callianassa profunda Biffar, 1973, new name for Cheramus occidentalis Bate, 1888, by subsequent designation. Gender of generic
name, Cheramus, masculine.
Species included. — Cheramus anoplourus (Sakai, 2002); C. aqabaensis
(Dworschak, 2003); C. contipes Sakai, 2002; C. gaucho (Rodrigues & Manning, 1992); C. lobetobensis (De Man, 1905); C. longicauda (Sakai, 1967);
C. malaccaensis (Sakai, 2002); C. modestus (De Man, 1905); C. nigroculatus (Sakai, 2002); C. oblongus (Le Loeff & Intès, 1974); C. orientalis Bate,
1888; C. parvulus (Sakai, 1988); C. praedatrix (De Man, 1905); C. profundus
(Biffar, 1973); C. propinquus (De Man, 1905); and C. spinophthalmus (Sakai,
1970a); C. sp. Robles et al., 2009.
K EY
TO THE
S PECIES
OF THE
G ENUS CHERAMUS
1 – Rostrum reaching or overreaching distal end of A1 proximal segment . . . . . . . . . . . . . 2
– Rostrum failing to reach distal end of A1 proximal segment . . . . . . . . . . . . . . . . . . . . . . 4
2 – P3 propodus oval. From North West Shelf, Western Australia, 83 m . . . . . . C. parvulus
– P3 propodus rectangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Telson bearing distinct proximal lobe. From Kepulauan Kai, Indonesia, 27 m . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. modestus
– Telson bearing small proximal lobe. From East China Sea . . . . . . . . . . . . C. longicauda
4 – Mxp3 merus bearing distal spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Mxp3 merus lacking distal spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 – Uropodal exopod distinctly longer than uropodal endopod; abdominal somite 6 distinctly longer than wide. From Teluk Kwandang, Sulawesi, Indonesia, 75 m . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. propinquus
366
CRM 013 – Katsushi Sakai
– Uropodal exopod slightly longer than uropodal endopod; abdominal somite 6 slightly
longer than wide. From between Bowoni and Boetoeng (= Butung = Buton), Indonesia, 75-94 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. praedatrix
6 – Eyestalks pointed apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Eyestalks triangular or obtuse distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7 – Mxp3 merus slanting on distal margin; telson slightly concave medially on posterior
margin. From Arafura Sea, 28 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. orientalis
– Mxp3 ischium convex on distomesial angle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 – Mxp3 ischium-merus about 1.4 times as long as wide; meral hook of male larger
cheliped with sharp proximal spine on ventral margin. From Murjan, Aqaba, 10-14 m
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aqabaensis
– Mxp3 ischium-merus about 2.1 times as long as wide; meral hook of male larger
cheliped with trifurcate proximal spine on ventral margin. From Tsushima Island,
Japan, 210 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. spinophthalmus
9 – Mxp3 ischium-merus about 1.8-2.3 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . 10
– Mxp3 ischium-merus about 3.0-3.3 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . 13
10 – Telson wider than long, posterior margin concave with median spine . . . . C. oblongus
– Telson longer than wide, posterior margin convex or straight . . . . . . . . . . . . . . . . . . . . 11
11 – Posterior margin of telson slightly convex, lacking median spine. From off Chui, near
border between Brazil and Uruguay, 150 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. gaucho
– Posterior margin of telson bearing median spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 – Telson largely convex on posterior margin. From Andaman Sea, 60.7 m . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. anoplourus
– Telson slightly convex on posterior margin. From off Sombrero Is., West Indies, 686724 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. profundus
13 – Rostrum short, reaching about proximal third of eyestalks . . . . . . . . . . . C. nigroculatus
– Rostrum distinct, reaching to near distal margin of eyestalks . . . . . . . . . . . . . . . . . . . . . 14
14 – Uropodal endopod broadened, uropodal exopod straight on distal margin . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. malaccaensis
– Uropodal endopod slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 – Uropodal exopod straight on distal margin . . . . . . . . . . . . . . . . . . . . . . . . . C. lobetobensis
– Uropodal exopod rounded on distal margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. contipes
Cheramus anoplourus (Sakai, 2002)
Callianassa anoploura Sakai, 2002: 490, fig. 17A-H; Sakai, 2005b: 72.
Diagnosis. — Mxp3 ischium-merus suboperculiform, merus rounded on
mesiodistal margin. Male Plp1 uniramous, tri-segmented; male Plp2 small and
biramous, endopod and exopod almost equal in length, lacking appendices
interna and masculina; male Plps3-5 endopods with projecting, finger-like
appendix interna. Telson subquadrate, rounded on posterior margin, with
median spine.
Type locality. — Andaman Sea, 9◦ 30.351′ N 97◦ 57.168′ E, 60.7 m, sandy
mud, fine sand, and shell fragments.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
367
Distribution. — Known only from the type locality.
Cheramus aqabaensis (Dworschak, 2003)
Callianassa aqabaensis Dworschak, 2003: 416, figs. 2-36; Robles et al., 2009: 316.
Diagnosis. — Plp1 uniramous and bisegmented (Dworschak, 2006: 424,
fig. 26), and male Plp2 biramous with slender, bisegmented endopod and slender unsegmented exopod; and Plps3-5 endopods bearing projecting, finger-like
appendix interna. [Cf. Dworschak, 2006: 424, figs. 30, 31.]
Type locality. — Murjan, Aqaba, 10-14 m.
Distribution. — Aqaba to Saudi Border, Jordania.
Cheramus contipes (Sakai, 2002)
Callianassa contipes Sakai, 2002: 523, fig. 35A-E; Sakai, 2005b: 80.
Diagnosis. — Mxp3 ischium-merus pediform, merus straight distally, continuous to convex mesial margin. Male Plps1-2 unknown. Telson quadrate,
convex on posterior margin, bearing median concavity with spine.
Type locality. — Andaman Sea, 7◦ 30.006′ N 98◦ 56.633′ E, 37.5 m, mud.
Distribution. — Andaman Sea, 7◦ 29.921′ N 99◦ 00.977′ E, 7◦ 44.638′ N
98◦ 16.496′ E; 20.5-30.5 m, mud.
Cheramus gaucho (Rodrigues & Manning, 1992)
(fig. 63D-E)
Poti gaucho Rodrigues & Manning, 1992b: 9, fig. 1; Rodrigues et al., 1998: 380; Melo, 1999:
380, figs. 255-256; Tudge et al., 2000: 145.
Callianassa gaucho — Sakai, 1999c: 28, fig. 3d-e; Sakai, 2005b: 46.
Material examined. — USNM 256376, paratype, male (CL 4.0 mm), off Chui, near the
border between Brazil and Uruguay, 33◦ 43′ S 51◦ 13′ W, 150 m, Fundacão Universidade do Rio
Grande, Cruise Sta. 3-11, leg. F. Dincao.
Diagnosis. — Mxp3 ischium-merus subpediform, merus straight distally,
with rounded distolateral angle. Male Plp1 uniramous and bisegmented
(fig. 63D); male Plp2 biramous, both endopod and exopod elongate, endopod
lacking appendices interna and masculina (fig. 63E). Telson trapezoid, slightly
convex on posterior margin, without median spine. [Cf. Rodrigues & Manning,
1992b: 9, fig. 1p, q, j, u.]
Type locality. — Off Chui, near the border between Brazil and Uruguay
(33◦ 43′ S 51◦ 13′ W), 150 m.
Distribution. — Known only from the type locality.
368
CRM 013 – Katsushi Sakai
Cheramus lobetobensis (De Man, 1905)
Callianassa lobetobensis De Man, 1905: 607; Sakai, 1999c: 48; Tudge et al., 2000: 143; Sakai,
2005b: 91.
Callianassa (Cheramus) lobetobensis — De Man, 1928b: 26, 93, 98, 137, pl. 13 fig. 20, pl. 14
fig. 20a-d.
Diagnosis. — Mxp3 ischium-merus subpediform, merus declined straight
on distal margin. Male Plps1-2 unknown. Telson subovate, distinctly notched
medially on posterior margin, with median spine. Male specimens are not
reported. [Cf. De Man, 1928b: 138, 140, pl. 14 fig. 20, 20a.]
Type locality. — Indonesia, Banda Sea, Lobetobi Strait between Flores and
Solor (8◦ 27′ S 122◦ 54.5′ E), 247 m.
Distribution. — Banda Sea, Lobetobi Strait between Flores and Solor,
Indonesia (8◦ 27′ S 122◦ 54.5′ E); 247 m.
Cheramus longicauda (Sakai, 1967)
Callianassa (Calliactites) longicauda Sakai, 1967: 324, figs. 3, 4 pl. 11C.
Callianassa longicauda — Sakai, 1987a: 303; Sakai, 1999c: 48.
Cheramus longicaudatus — Tudge et al., 2000: 145.
Diagnosis. — Mxp3 ischium-merus subsquare, merus straight with distinct median tooth on distal margin. Male Plp1 uniramous, slender, and bisegmented; male Plp2 biramous, with short endopod and exopod. Telson subquadrate, convex on posterior margin, with median spine. [Cf. Sakai, 1967:
324-326, figs. 3C, F, 4A, B.]
Type locality. — East China Sea (32◦ N 122◦ 30′ E).
Distribution. — Known only from the type locality.
Cheramus malaccaensis (Sakai, 2002)
Callianassa malaccaensis Sakai, 2002: 492, figs. 18A-D, 19A-K; Sakai, 2005b: 91-92.
Diagnosis. — Mxp3 ischium-merus subpediform, merus obliquely truncate
on distal margin. Male Plp1 uniramous and bisegmented; male Plp2 biramous;
male Plps3-5 endopods with projecting, finger-like appendix interna. Telson
subquadrate, longer than wide, and slightly tapering toward posterior margin,
bearing median concavity with spine.
Type locality. — Andaman Sea, 6◦ 45.045′ N 99◦ 20.766′ E, 38.2 m.
Distribution. — Andaman Sea, 6◦ 45.045′ N 99◦ 20.766′ E, 38.2 m;
◦
6 45.019′ N 98◦ 44.832′ E, 82.7 m; 9◦ 30.351′ N 97◦ 57.168′ E, 60.7 m; muddy
sand, sandy mud, fine sand and shell fragments.
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369
Cheramus modestus (De Man, 1905)
Callianassa (Calliactites) modesta De Man, 1905: 604 (partim); Balss, 1925: 212; De Man,
1928b: 26, 97, 118, pl. 10 fig. 16-16b, pl. 11 fig. 16c-e.
Callianassa modesta — Liu & Zhong, 1994: 562; Sakai, 1999c: 48; Komai, 2000b: 345 (list);
Tudge et al., 2000: 143.
Diagnosis. — Mxp3 ischium-merus subpediform, merus straight with median tooth on distal margin. Male Plp1 uniramous and bisegmented; Plp2 biramous. Telson subovate, convex on posterior margin, with median spine.
Type locality. — Elat, Kepulauan Kai, Indonesia, 27 m.
Distribution. — N. South China Sea (Liu & Zhong, 1994); west of Kwandang Bay; Bay of Bima (0◦ 58.5′ N 122◦ 42.5′ E); Great Kei Island, Indonesia
(5◦ 40′ S 132◦ 26′ E); 27-310 m.
Cheramus nigroculatus (Sakai, 2002)
Callianassa nigroculata Sakai, 2002: 525, figs. 36A-D, 37A-H.; Sakai, 2005b: 94.
Diagnosis. — Mxp3 ischium-merus subpediform, merus rounded at distomesial angle. P3 propodus oval. Male Plps1-2 undescribed; Plps3-5 endopods with projecting, finger-like appendix interna. Telson trapezoid, narrowing and slightly concave in posterior two-thirds, bearing median spine on
straight posterior margin, and dorsal surface slightly concave medially. [Cf.
Sakai, 2002: 525-528, figs. 36D, 37A.]
Type locality. — Andaman Sea, 7◦ 30.221′ N 98◦ 22.005′ E – 7◦ 35.932′ N
◦
98 25.348′ N, 43.2-74.8 m.
Distribution. — Andaman Sea, 7◦ 29.786′ N 98◦ 17.348′ E; 7◦ 15.061′ N
◦
98 34.181′ E; 7◦ 35.932′ N 98◦ 25.348′ N; 60.4-78.7 m, muddy sand.
Cheramus oblongus (Le Loeuff & Intès, 1974)
Callianassa oblonga Le Loeuff & Intès, 1974: 38, fig. 9a-r; De Saint Laurent & Le Loeuff,
1979: 55; Sakai, 1999c: 18; Sakai, 2005b: 32.
Cheramus oblongus — Tudge et al., 2000: 145.
Diagnosis. — Rostrum acutely triangular and slightly shorter than eyestalks.
Eyestalks obtuse distally; cornea distinct and centred. A1 peduncle slightly
longer than A2 peduncle. Mxp3 ischium-merus subsquare and more than
twice as long as wide; merus straight on distal margin. P3 propodus short
and subsquare. Male Plp1 uniramous and bisegmented; male Plp2 slightly
developed (De Saint Laurent & Le Loeuff, 1979: 55). Telson trapezoid, bearing
proximal lobe on lateral margin, and median concavity with spine on posterior
margin. [Cf. Le Loeuff & Intès, 1974: 32, fig. 9k, q.]
370
CRM 013 – Katsushi Sakai
Type locality. — Ivory coast, Grand Bassam (4◦ 59′ N 3◦ 48′ W), 200 m.
Distribution. — Only known from the type locality.
Cheramus orientalis Bate, 1888
Cheramus orientalis Bate, 1888: 30, pl. 1 fig. 2.
Callianassa (Cheramus) orientalis — Borradaile, 1903: 546; De Man, 1928a: 9, fig. 2, 2a; De
Man, 1928b: 26, 93, 98, 119, 132, 137.
Callianassa orientalis — Sakai, 1999c: 49, fig. 5a-c; Sakai, 2005b: 20, 95.
Cheramus orientalis — Tudge et al., 2000: 145.
Diagnosis. — Mxp3 ischium-merus subpediform, merus obliquely convex
on distal margin. Male Plps1-2 unknown; Plps3-5 endopods with projecting,
finger-like appendix interna. Telson trapezoid, slightly longer than wide,
tapering toward posterior margin, with small and shallow median concavity.
[Cf. Bate, 1888: 31, fig. 2.]
Type locality. — Arafura Sea (9◦ 59′ S 139◦ 42′ E), 28 m.
Distribution. — Known only from the type locality.
Cheramus parvulus (Sakai, 1988)
Callianassa parvula Sakai, 1988: 59, fig. 3; Sakai, 1999c: 50; Tudge et al., 2000: 143; Davie,
2002: 458; Sakai, 2005b: 95-96.
Diagnosis. — Rostrum acutely triangular, overreaching eyestalks. Mxp3
ischium-merus unknown. Male Plp1 uniramous and bisegmented; Plp2 biramous. Telson trapezoid, wider than long, tapering toward posterior margin,
with median concavity. [Cf. Sakai, 1988: 59, fig. 3D, E, F.]
Type locality. — North West Shelf, Western Australia (19◦ 04.4′ S
118◦ 47.35′ E); 83 m.
Distribution. — Known only from the type locality.
Cheramus praedatrix (De Man, 1905)
Callianassa praedatrix De Man, 1905: 607; Sakai, 1988: 59, fig. 4; Ngoc-Ho, 1994: 54, fig. 2;
Sakai, 1999c: 51; Sakai, 2005b: 100.
Callianassa (Cheramus) praedatrix — De Man, 1928b: 26, 97, 99, 146, pl. 15 fig. 22-22d.
Cheramus praedatrix — Tudge et al., 2000: 145; Davie, 2002: 459.
Diagnosis. — Mxp3 ischium-merus subsquare; merus armed with sharp
spine on distal margin, and rounded on distomesial margin. Male Plp1 uniramous and bisegmented; male Plp2 biramous; male Plps3-5 endopods with projecting, finger-like appendix interna. Telson subsquare, bearing shallow notch
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371
with median spine on straight posterior margin. [Cf. De Man, 1928b: 147, 148,
149, pl. 15 fig. 22-22a, b.]
Type locality. — Between Bowoni and Boetoeng (= Butung = Buton),
Indonesia (4◦ 20′ S 122◦ 58′ E); 75-94 m.
Distribution. — Between Bowoni and Boetoeng (= Butung = Buton), Indonesia; 75-94 m; North West Shelf, Western Australia; sublittoral, benthic,
continental shelf, 41-51 m.
Cheramus profundus (Biffar, 1973)
Callianassa occidentalis Bate, 1888: 29, pl. 2 fig. 2k; Yong, 1900: 425; Borradaile, 1903:
548; Balss, 1925: 212; De Man, 1928b: 115; Schmitt, 1935b: 3; Biffar, 1971a: 649.
[Type locality. — 18◦ 29′ 3N 63◦ 24.6′ W, off Sombrero Is., West Indies, 686-724 m.] [Not:
Callianassa occidentalis Stimpson, 1856.]
Cheramus occidentalis Bate, 1888: 32, pl. 2 fig. 1; Yong, 1900: 246. [Type locality. — 18◦ 29′ 3N
63◦ 24.6′ W, off Sombrero Is., West Indies, 686-724 m.] [Not: Callianassa occidentalis
Stimpson, 1856 (= subjective junior synonym of Callianassa californiensis Dana, 1854).]
Callianassa (Cheramus) Batei Borradaile, 1903: 546; De Man, 1928a: 10, pl. 1 fig. 3; De Man,
1928b: 26, 98. [New name for Cheramus occidentalis Bate, 1888.]
Callianassa batei — Schmitt, 1935b: 5; Biffar, 1971a: 649, 654; Manning, 1987: 398.
Callianassa profunda Biffar, 1973: 225, figs. 1, 2 [New name for Callianassa batei Borradaile,
1903.]; Sakai, 1999c: 28; Sakai, 2005b: 46.
Cheramus batei — Manning & Felder, 1991: 780; Coelho, 1997: 150; Melo, 1999: 358, figs.
237-238.
Cheramus profundus — Tudge et al., 2000: 145.
Diagnosis. — Mxp3 ischium-merus subsquare; merus straight distally, with
obtuse distomesial angle. Male Plp1 uniramous and bisegmented; male Plp2
biramous; male Plps3-5 biramous, endopods bearing finger-like appendix interna. Telson trapezoid, longer than wide, widest proximally, slightly narrowing toward posterior margin with distinct median spine. [Adapted from Biffar,
1973: 227.]
Type locality. — Off Sombrero Is., West Indies, 18◦ 29′ 3N 63◦ 24.6′ W, 686724 m.
Distribution. — Off Sombrero Is., West Indies, 686-724 m; Rio Grande do
Sul, Brazil (Coelho, 1997).
Cheramus propinquus (De Man, 1905)
Callianassa propinqua De Man, 1905: 609; Ngoc-Ho, 1991: 290, fig. 4; Ngoc-Ho, 1994: 54,
fig. 2; Sakai, 1999c: 51; Sakai, 2005b: 100-101.
Callianassa (Cheramus) propinqua — De Man, 1928b: 27, 98, 127, pl. 12 fig. 18-18d.
372
CRM 013 – Katsushi Sakai
Cheramus propinquus — Tudge et al., 2000: 145; Davie, 2002: 459.
Diagnosis. — Mxp3 ischium-merus subpediform; merus armed with sharp
spine on distal margin, and rounded at distomesial corner. Male Plp1 uniramous and bisegmented; male Plp2 biramous; and male Plps3-5 endopods with
projecting, finger-like appendix interna. Telson quadrate, longer than wide,
bearing proximal lobe laterally, two spinules at posterolateral corner, and median spine on convex posterior margin. Uropodal endopod oval and rounded
distally; uropodal exopod longer than endopod, and rectangular. [Cf. Ngoc-Ho,
1991: 290, 291, fig. 4d, f, k.]
Type locality. — Teluk Kwandang, Sulawesi, Indonesia (0◦ 58.5′ N
122◦ 55′ E), 75 m.
Distribution. — Kwandang Bay, Indonesia; New Caledonia; North-West
Shelf, Australia; benthic, continental shelf, continental slope; shelly sand to
mud substrates; 75-300 m.
Cheramus spinophthalmus (Sakai, 1970)
Callianassa (Cheramus) spinophthalma Sakai, 1970a: 40, figs. 2, 3, 4a-b; Sakai, 1987a: 306.
Callianassa spinophthalma — Sakai, 1999c: 52; Sakai, 2005b: 103.
Cheramus spinophthalmus — Tudge et al., 2000: 145; Komai et al., 2002: 39, fig. 7.
Diagnosis. — Mxp3 ischium-merus subsquare, merus truncate on distal
margin, and convex on mesial margin. Male Plp1 uniramous and bisegmented;
male Plp2 biramous; male Plps3-5 endopods with projecting, finger-like
appendix interna. Telson trapezoid, slightly concave on posterior margin, with
median spine. [Cf. Sakai, 1970a: 40, figs. 3b, 4a; Komai, 2002, fig. 11.]
Type locality. — Tsushima Island, Japan (34◦ 58′ N 129◦ 26′ 9E), 210 m.
Distribution. — Known only from the type locality.
Cheramus sp. Robles et al., 2009
Cheramus sp., Robles et al., 2009: 317.
Genus Gilvossius Manning & Felder, 1992 (sensu nov.)
Gilvossius Manning & Felder, 1992: 558.
Pestarella Ngoc-Ho, 2003: 475 [type species: Astacus tyrrhenus Petagna, 1792].
Paratrypaea Komai & Tachikawa, 2008: 36. [Type species. — Callianassa (Trypaea) bouvieri
(Nobili, 1904).]
Diagnosis. — Eyestalks depressed. Mxp3 propodus subovate or subrectangular, and dactylus digitiform. Male Plps1-2 absent. Male Plps3-5 biramous,
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
373
endopods with projecting, finger-like or embedded appendix interna. Uropodal
exopod with dorsal plate.
Remarks. — The genus Pestarella Ngoc-Ho, 2003 was established because
of the absence of male Plps1-2, but it is synonymous with Gilvossius Manning
& Felder, 1992, because the type species of Pestarella, Astacus tyrrhenus
Petagna, 1792 has no male Plps1-2 as in the type species of Gilvossius,
Gonodactylus setimanus DeKay, 1844.
Komai & Tachikawa (2008) established a new genus Paratrypaea Komai
& Tachikawa, 2008 on the basis of the type species, Callianassa (Trypaea)
bouvieri Nobili, 1904. However, their new genus is synonymous with the genus
Gilvossius, because Paratrypaea and Gilvossius have the same type of male
Plps1-2 (Komai & Tachikawa, 2008, fig. 10E).
Type species. — Gonodactylus setimanus DeKay, 1844, by original designation of Manning & Felder (1992). The gender of the name, Gilvossius, is
masculine.
Species included. — Gilvossius bouvieri (Nobili, 1904); G. brachytelson
(Sakai, 2002); G. candidus (Olivi, 1792); G. convexus (De Saint Laurent
& Le Loeuff, 1979); G. diaphorus (Le Loeuff & Intès, 1974); G. gravieri
(Nobili, 1905); G. marchali (Le Loeuff & Intès, 1974); G. persicus (Sakai,
2005b); G. rotundicaudatus (Stebbing, 1902); G. setimanus (DeKay, 1844);
G. tyrrhenus (Petagna, 1792); G. whitei (Sakai, 1999c); G. sp., Robles et al.,
2009: 316.
K EY
TO THE
S PECIES
OF THE
G ENUS GILVOSSIUS
1 – Mxp3 ischium-merus pediform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Mxp3 ischium-merus broadened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 – A2 peduncle distinctly longer than A1 peduncle . . . . . . . . . . . . . . . . . . . . . . . G. diaphorus
– A2 peduncle slightly longer than A1 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Telson distinctly convex on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. persicus
– Telson slightly convex on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. marchali
4 – A2 peduncle about as long as A1 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– A2 peduncle shorter than A1 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 – Telson rounded distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. rotundicaudatus
– Telson trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. candidus
6 – Uropodal endopod narrow, and longer than telson; uropodal exopod without dorsal plate
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. brachytelson
– Uropodal endopod broadened; uropodal exopod with dorsal plate . . . . . . . . . . . . . . . . . . 7
7 – Telson subsquare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– Telson semicircular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
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CRM 013 – Katsushi Sakai
8 – Larger cheliped setose on dactylus, bearing serrate and convex meral hook . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. bouvieri
– Larger cheliped not setose on dactylus, bearing slender meral hook . . . . . . . . . . . . . . . . . 9
9 – Rostrum acutely triangular, cornea small and located medially . . . . . . . . . . . . G. gravieri
– Rostrum obsolete, cornea distinct and occupying definite area medially . . . G. pacificus
10 – Meral hook of male larger cheliped convex on ventral margin . . . . . . . . . . . . G. tyrrhenus
– Meral hook of male larger cheliped sharp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 – Mxp3 merus convex at distomesial corner . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. whitei
– Mxp3 merus rounded at distomesial corner . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 – Mxp3 ischium-merus 1.2 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . G. setimanus
– Mxp3 ischium-merus more than 1.5 times as long as wide . . . . . . . . . . . . . . . G. convexus
Gilvossius bouvieri (Nobili, 1904)
Callianassa (Trypaea) Bouvieri Nobili, 1904: 236; Nobili, 1906b: 105, pl. 6 fig. 3; De Man,
1928b: 27, 107, 146.
Callianassa maldivensis Borradaile; 1904: 753, pl. 58 fig. 3a, 3b; Tudge et al., 2000: 143. [Type
locality. — Hulule, Male Atoll, Maldives.]
Callianassa (Trypaea) maldivensis — Borradaile, 1903: 546 (nomen nudum); Borradaile, 1904:
753, pl. 58 fig. 3a, 3b; Pearson, 1905: 90; De Man, 1928a: 22; De Man, 1928b: 28, 107, 134,
146. [Type locality. — Hulule, Male Atoll, Maldives.]
Callianassa bouvieri — Holthuis, 1958: 37, 38, fig. 15; Sakai, 1970a: 46; Sakai, 1987a: 303;
Dworschak & Pervesler, 1988: 3, fig. 3; Dworschak, 1992: 192; Sakai, 1999c: 40, fig. 6a-c;
Tudge et al., 2000: 143; Sakai & Apel, 2002: 276; Itani, 2004, fig. 3; Sakai, 2005b: 78; Itani,
2007: 203.
Callianassa rectangularis Ngoc-Ho, 1991: 292, fig. 5. [Type locality. — Atoll de Surprise, New
Caledonia, 36 m.]
Cheramus rectangularis — Tudge et al., 2000: 145.
Paratrypaea bouvieri — Komai, 2008: 36, figs. 10-12. [Comb. with Callianassa (Trypaea)
maldivensis Borradaile, 1904.]
Material examined. — SMF 32729, 7 males, 1 female, Maheno-hama, Akajima Island,
Kerama Islands, Ryukyu Archipelago, fine sandy beach, rec. 11.i.1992, leg. A. Fukushima.
Diagnosis. — Rostrum acutely triangular, almost reaching distal margin of
eyestalks. Mxp3 ischium-merus suboperculiform; merus rounded at distomesial corner. In larger cheliped dactylus setose on lateral surface. Male Plps1-2
absent; male Plps3-5 endopods with projecting, finger-like appendix interna.
Telson convex on posterior margin, with small median concavity.
Type locality. — Djibouti.
Distribution. — Egypt, Red Sea; Djibouti, Gulf of Aden; Gulf of Mannar,
India; Maldives; Sri Lanka; Mindanao, Philippines; Bali and the Kei Islands,
Indonesia; Amami-Ohshima, Amakusa Island, and Tsushima, Japan. Here
recorded from Mindanao, the Philippines, and Bali and the Kei Islands,
Indonesia.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
375
Gilvossius brachytelson (Sakai, 2002)
Callianassa brachytelson Sakai, 2002: 499, fig. 21A-E; Sakai, 2005b: 79.
Diagnosis. — Rostrum acutely triangular, reaching distal margin of eyestalks. Mxp3 ischium-merus subsquare; merus straight distally, and convex on
mesial margin. Male Plps1-2 absent; male Plps3-5 endopods with projecting,
finger-like appendix interna. Telson subsquare, and slightly rounded on posterior margin, with distinct median spine. Uropods longer than telson. Uropodal
exopod subsquare, and much longer than endopod; dorsal surface carinate medially.
Type locality. — Andaman Sea, 07◦ 45.452′ N 97◦ 57.951′ E, 70.0 m, coarse
and fine sand.
Distribution. — Andaman Sea, 7◦ 45.452′ N 97◦ 57.951′ E; 08◦ 00.016′ N
◦
97 53.927′ E; 70.0-76.1 m, muddy sand, coarse and fine sand.
Gilvossius candidus (Olivi, 1792)
[Abbreviated list of synonymy.]
Cancer candidus Olivi, 1792: 51, pl. 3 fig. 3.
Callianassa subterranea forma pontica Czerniavsky, 1884: 81 (partim). [Type locality. — Black
Sea.]
Callianassa pontica — Cano, 1891: 5-30; Caroli, 1940: 76; Lutze, 1941: 34; Caroli, 1946: 71;
Vatova, 1949, table 5; Caroli, 1950: 190; Dolgopol’skaia, 1954: 179, figs. 3, 4; De Saint
Laurent & Božić, 1976: 24, figs. 5, 13, 21, 32; Beaubrun, 1979: 84, figs. 58, 59, 68-70;
Moncharmont, 1979: 71; García Raso, 1983: 318, fig. 3; Thessalou-Legaki & Zenetos, 1985:
311; Thessalou-Legaki, 1986: 182; Tudge et al., 2000: 143; Makarov, 2004: 289, figs. 131134, tab. 17.
Callianassa (Callichirus) Pestae De Man, 1928a: 34, pl. 9 figs. 16-16e; De Man, 1928b: 111.
[Type locality. — Mediterranean.]
Callianassa pestae — Lutze, 1937: 6; Lutze, 1938: 167 (partim) (not figs. 10-21 = Callianassa
subterranea); Vatova, 1949, tables, 16, 18, 20, 24, 31; Marcuzzi, 1972: 193; Manning &
Števčić, 1982: 295; Froglia & Grippa, 1986: 261.
Callianassa sp., Băcescu, 1949: 9, figs. 20-22.
Callianassa pestai — Holthuis, 1953a: 95, fig. 3; Doldopolski, 1969: 316, pls. 32-34; Kobyakova & Dolgopol’skaia, 1969: 286, pl. 5 fig. 1a-c; Števčić, 1971: 529; Mazmanidi et al., 1998:
129.
Callianassa algerica Lutze, 1938: 168 (partim) [not: fig. 22, eyestalks = C. whitei], figs. 26a,
larger Mxp3 [not smaller Mxp3 = C. subterranea], 26b, larger Mxp3 [not smaller Mxp3 =
C. subterranea], 27 [nomen dubium]. [Type locality. — Castiglione, near Algiers.]
Callianassa (Callichirus) pontica — Makarov, 1938: 73, 297, figs. 27, 28; Băcescu, 1967: 231,
fig. 105.
Callianassa candida — Giordani Soika, 1943: 83 (partim); Giordani Soika, 1945: 994;
Mikashavidze, 1981: 1415; Lewinsohn & Holthuis, 1986: 20; Števčić, 1990: 217; Dworschak, 1992: 194 (partim); Koukouras et al., 1992: 223; d’Udekem d’Acoz, 1996: 54; Sakai,
1999c: 14, fig. 2a-d; d’Udekem d’Acoz, 1999: 154; Tudge et al., 2000: 143; Costello et al.,
2001: 289; Dworschak, 2002: 63, figs. 1, 2, table 1; d’Udekem d’Acoz, 2003, fig. website.
376
CRM 013 – Katsushi Sakai
Callianassa (Callichirus) pestae — Zariquiey Alvarez, 1968: 230; Kattoulas & Koukouras,
1974: 344, fig. 1.
Callianassa candida — Sakai, 1999c: 14, fig. 2a-d; Dworschak, 2002: 63, figs. 1, 2, table 1;
Sakai, 2005b: 29.
Callianassa (Callianassa) candida — Felder, 2001: 441.
Pestarella candida — Ngoc-Ho, 2003: 476, fig. 12; Dworschak, 2004: 15, figs. 1, 3.
Diagnosis. — Male Plps1-2 absent; Plps3-5 endopods with projecting,
finger-like appendix interna.
Type locality. — Northern Adriatic lagoons: “nelle lagune sprofondato”
(Olivi, 1792).
Distribution. — From Golfo de Cádiz, S.W. Spain, eastern Atlantic, to
Alupka, Black Sea; 10 m.
Gilvossius convexus (De Saint Laurent & Le Loeuff, 1979)
Callianassa convexa De Saint Laurent & Le Loeuff, 1979: 53, fig. 10a-e; Sakai, 1999c: 17;
Tudge et al., 2000: 143; Sakai, 2005b: 31.
Diagnosis. — Mxp3 ischium-merus subsquare, straight distally, and rounded at distomesial corner. Male Plps1-2 absent. Telson semicircular in form.
[Cf. De Saint Laurent & Le Loeuff, 1979: 53, fig. 10d, e.]
Type locality. — South of Cape Bald, Gambia, 18 m.
Distribution. — Gambia; Mauritania.
Gilvossius diaphorus (Le Loeuff & Intès, 1974)
Callianassa diaphora Le Loeuff & Intès, 1974: 32, fig. 7a-v; De Saint Laurent & Le Loeuff,
1979: 49, fig. 9a, b, e, g; Sakai, 1999c: 18; Sakai, 2005b: 32.
Callianassa guineensis — Longhurst, 1958: 31 (partim).
Biffarius diaphora — Tudge et al., 2000: 143.
Diagnosis. — Mxp3 ischium-merus subpediform, with oblique distal margin. Male Plp1 absent; male Plp2 absent or present as feeble papilla; male
Plps3-5 endopods with finger-like appendix interna. Telson trapezoid. [Cf. Le
Loeuff & Intès, 1974: 32, fig. 7k, q, u.]
Remarks. — Le Loeuff & Intès (1974: 35) described the male Plps1-2 of
their species Callianassa diaphora with “les pléopodes 1 et 2 sont le plus
souvent absents mais le pl2 peut subsister à l’état de faible diverticule”.
[= Male Plps1-2 are absent in most cases, but Plp2 may be present as a
feeble papilla], so that this species may safely be reclassified under the genus
Gilvossius by the features of male Plps1-2.
Type locality. — Grand Lahou, Côte d’Ivoire, 5◦ 07′ N 5◦ 04.5′ W, 22 m.
Distribution. — Siera Leone to Ivory Coast; 10-60 m.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
377
Gilvossius gravieri (Nobili, 1905)
Callianassa (Trypaea) Gravieri Nobili, 1905: 396 (not 395); Nobili, 1906b: 107, pl. 6 fig. 4;
Balss, 1915: 2; De Man, 1928a: 23, pl. 6 fig. 11-11e; De Man, 1928b: 27, 107.
Callianassa (Trypaea) cristata Borradaile, 1910: 263, pl. 16 fig. 7; De Man, 1928b: 27, 107.
[Type locality. — Salomon Atoll; Chagos Archipelago.]
Callianassa (Trypaea) gravieri — Holthuis, 1953b: 51.
Callianassa gravieri — Sakai, 1999c: 43, fig. 6d-f; Tudge et al., 2000: 143; Sakai, 2005b: 8586.
Callianassa cristata — Tudge et al., 2000: 143.
Diagnosis. — Rostrum acutely triangular, almost reaching distal margin of
eyestalks. Mxp3 ischium-merus subsquare, merus straight distally and rounded
at distomesial corner. Male Plps1-2 probably absent (De Man, 1928a: 24).
Telson subsquare, and straight on posterior margin, with median spine.
Remarks. — This species probably belongs to the genus Gilvossius by the
absence of male Plps1-2, as shown above.
Type locality. — Obock, Gulf of Aden.
Distribution. — Harmil Is., Red Sea; Obock and Djibouti, Gulf of Aden;
Salomon Atoll; Chagos Archipelago.
Gilvossius marchali (Le Loeuff & Intès, 1974)
Callianassa marchali Le Loeuff & Intès, 1974: 35, fig. 8a-r; De Saint Laurent & Le Loeuff,
1979: 51, fig. 8c, d, f, h; Sakai, 1999c: 18; Tudge et al., 2000: 143.
Callianassa guineensis — Longhurst, 1958: 31 (partim).
Diagnosis. — Mxp3 ischium-merus subpediform, with oblique distal margin. Male Plps1-2 absent. Telson longer than wide and trapezoid, tapering toward posterior margin, with median convexity. [Cf. Le Loeuff & Intès, 1974:
32, fig. 8j, p.]
Type locality. — Cote d’Ivoire, between Vridi and Jacqueville (5◦ 09.5′ N
4◦ 09′ W), 70 m.
Distribution. — Pointe-Noire, Congo; Ivory Coast; Sierra Leone; Senegal;
70-250 m.
Gilvossius pacificus (Guzmán & Thatje, 2003)
Biffarius pacificus Guzmán & Thatje, 2003: 294, figs. 1, 2.
Diagnosis. — Rostrum obsolete. Mxp3 ischium-merus operculiform. Male
Plp1 absent; male Plp2 undescribed, probably absent; Plps3-5 biramous,
endopods with stubby appendix interna in both sexes. [Adapted from Guzmán
& Thatje, 2003.]
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CRM 013 – Katsushi Sakai
Remarks. — In Biffarius pacificus, the male Plp1 is absent, but the male
Plp2 is undescribed, so this species is to be reclassified as a species of
Gilvossius.
Type locality. — Off northern coast of Chile, 22◦ 12′ S to 22◦ 45′ S, 17-30 m.
Distribution. — Only known from the type locality.
Gilvossius persicus (Sakai, 2005)
Callianassa persica Sakai, 2005b: 97, fig. 21.
Diagnosis. — Male Plps1-2 absent.
Type locality. — Persian Gulf, 28◦ 54′ N 50◦ 11′ E, 56 m.
Distribution. — Only known from the type locality.
Gilvossius rotundicaudatus (Stebbing, 1902)
Callianassa rotundicaudata Stebbing, 1902: 41, pl. 8; Pearson, 1905: 90; Stebbing, 1910: 369;
Kensley, 1974: 277 (key); Dworschak, 1992: 202, fig. 11a-f; Sakai, 1999c: 52; Tudge et al.,
2000: 143; Sakai, 2005b: 18, 24, 28, 102.
Callianassa (Calliactites) rotundicaudata — Borradaile, 1903: 545; De Man, 1928b: 26, 92,
94, 97, 123, 136, 149, 150; Barnard, 1950: 512, fig. 95i-l.
Callianassa (Calliactites) rotundicaudata — Stebbing, 1910: 369.
Type locality. — St. Francis Bay, South Africa, 34◦ 02.45′ S 25◦ 10′ E.
Distribution. — South Africa — Orange River mouth, Saldanha Bay; St.
Francis Bay; Algoa Bay; Kowie, Port Alfred; 10-35 m; Cheval Paar, Sri Lanka.
Gilvossius setimanus (DeKay, 1844)
Gonodactylus setimanus DeKay, 1844: 34, pl. 8 fig. 23; Manning, 1987: 386.
Callianassa Stimpsoni Smith, 1873: 549, pl. 2 fig. 8; Kingsley, 1878: 327; Borradaile, 1903:
548; Fowler, 1911: 570; Hay & Shore, 1917: 406, pl. 29 fig. 5. [Type locality. — “Our
species ranges from the coast of the Southern [United] States north to Long Island Sound”
(Smith, 1873).] [Junior homonym of Callianassa stimpsoni Gabb, 1864, a fossil species.]
Callianassa atlantica Rathbun, 1926: 107; Schmitt, 1935b: 4; Biffar, 1971a: 654; Rabalais et
al., 1981: 101, fig. 2; Manning, 1987: 397; Squires, 1990: 334, figs. 181-183. [New name for
C. stimpsoni.]
Callianassa (Callichirus) atlantica — De Man, 1928a: 37, pl. 9 fig. 17-17d; De Man, 1928b:
28, 94, 112; Williams, 1965: 102, fig. 79.
Callianassa stimpsoni — Manning, 1987: 397.
Gilvossius setimanus — Manning & Felder, 1992: 558, fig. 1; Tudge et al., 2000: 143; Sakai,
2005b: 47-48; Heard et al., 2007: 21, fig. 24.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
379
Callianassa setimana — Sakai, 1999c: 28.
Material examined. — USNM 10497, 1 male (TL/CL, 77.0/18.0 mm), New Port, Rhode
Island, 1885, leg. R/V “Albatross”.
Diagnosis. — Mxp3 ischium-merus broadened and subsquare; merus rounded at distomesial corner. Male Plps1-2 absent. Telson semicircular, lacking
median spine on posterior margin. [Cf. Manning & Felder, 1992, fig. 1c, g.]
Remarks. — It is here confirmed that the male Plps1-2 are absent.
Type locality. — New York.
Distribution. — Bass River, Nova Scotia to South Carolina; Franklin
County, Florida, 38 m; northwestern Gulf of Mexico, 134 m; Colombia
(Caribbean side).
Gilvossius tyrrhenus (Petagna, 1792)
Astacus tyrrhenus Petagna, 1792: 418, pl. 5 fig. 3.
Alpheus Tyrhenus Risso, 1816: 94 (only reference), not pl. 2 fig. 2.
Callianassa laticauda Otto, 1821: 11; Otto, 1828: 345, pl. 21 fig. 3; Heller, 1863: 203; Stalio,
1877: 107; Carus, 1885: 489; Caroli, 1940: 73; Lutze, 1941: 34; Gurney, 1942: 245; Caroli,
1946: 71; Zariquiey Alvarez, 1946: 106; Caroli, 1950: 189; Holthuis, 1953a: 91; Forest &
Gantès, 1960: 348. [Type locality. — Nice, S. France, Mediterranean.]
Gebios Davyanus Risso, 1822: 243; Risso, 1827: 52. [Type locality. — Nice, Mediterranean.]
Callianassa tyrrhena — Risso, 1827: 54 (partim); Holthuis, 1947: 320, fig. 1; Holthuis, 1950:
113 (partim), fig. 40; Holthuis, 1953a: 91, 93, fig. 1; Picard, 1957: 48; Forest & Guinot, 1958:
10; Holthuis & Gottlieb, 1958: 62 (partim), not fig. 13 (= C. subterranea); Bourdon, 1965:
1; Forest, 1967: 6 (partim); Glaçon, 1971: 1; Števčić, 1971: 529; Neves, 1974: 13, fig. 4;
Pastore, 1976: 107, 111; De Saint Laurent & Božić, 1976: 22, figs. 4, 12, 20, 31; Holthuis,
1977: 57; Zarkanellas & Bogdanos, 1977: 155; Beaubrun, 1979: 90, figs. 55, 60, 61, 66, 67;
De Saint Laurent & Le Loeuff, 1979: 53; Domenech et al., 1981: 149; Adema et al., 1982: 26,
fig. 7a-c; Manning & Števčić, 1982: 295; Abel et al. 1983: 483, pl. 177; García Raso, 1983:
323, fig. 2; Thessalou-Legaki, 1986: 182; d’Udekem d’Acoz, 1986: 101; Türkay et al., 1987;
d’Udekem d’Acoz, 1989: 176; Moyse & Smaldon, 1990: 520, fig. 10.13; Števčić, 1990: 217;
Thessalou-Legaki, 1990: 659; Holthuis, 1991: 252, 264, fig. 457; Pérez Sánchez & Moreno
Batet, 1991: 1; Dworschak, 1992: 206; Koukouras et al., 1992: 223; Gruner, 1993: 997;
Mayoral et al., 1994: 236; Hayward et al., 1995: 432, figs. 8-52; d’Udekem d’Acoz, 1996:
53; Hadjichristophorou et al., 1997: 22; Ingle, 1997: 78, fig. 7.3; Thessalou-Legaki et al.,
1997a: 439; Thessalou-Legaki et al., 1997b: 435; Thessalou-Legaki & Kiortsis, 1997: 435;
Sakai, 1999c: 21, fig. 3a-c; Thessalou-Legaki et al., 1999: 635; d’Udekem d’Acoz, 1999:
155; Dworschak, 2000b: 155, figs.; Dworschak et al., 2000: 313; Tudge et al., 2000: 143;
Costello et al., 2001: 289; Vanhaelen, 2001: 147; d’Udekem d’Acoz, 2003, fig. on website;
Ingle & Christiansen, 2004: 96, figs. 70, 72; Sakai, 2005b: 40-42.
Callianassa subterranea — H. Milne Edwards, 1837a: 309; H. Milne Edwards, 1837c: 130,
pl. 48 fig. 3-3e; Heller, 1863: 202, pl. 6 fig. 9, 11; Ortmann, 1891: 55, pl. 1 fig. 10.
Gebios davianii — Risso, 1844: 94.
Callianassa Laticauda — Hope, 1851: 14.
Callianassa Candida — Hope, 1851: 14.
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CRM 013 – Katsushi Sakai
Gebia Daviana — Hope, 1851: 15.
Callianassa (Callichirus) laticauda — Stalio, 1877: 664; Carus, 1885: 489; Giard & Bonnier,
1890: 366; Borradaile, 1903: 547; Pesta, 1912: 105; Pesta, 1918: 204; De Man, 1928a: 33,
pl. 8 fig. 15-15d; De Man, 1928b: 28, 91, 92, 111; Bouvier, 1940: 102, fig. 69 (partim);
O’Céidigh, 1962: 164; O’Riordan, 1983: 72. [Not: Callianassa laticauda Otto, 1821.]
Callianassa subterranea laticauda var. — Czerniavsky, 1884: 76, 80.
Callianassa sp., Stebbing, 1893: 184.
Callianassa (Callichirus) Stebbingi Borradaile, 1903: 547. [Type locality. — Jersey, N.E.
Atlantic.]
Callianassa Stebbingi — Selbie, 1914: 100, pl. 14 figs. 8-10.
Callianassa stebbingi — Caroli, 1921a: 245 (not C. subterranea after Thessalou-Legaki, 1990);
Schellenberg, 1928: 78, figs. 59, 60; Steinitz, 1933: 147; Bodenheimer, 1937: 281; Lutze,
1937: 6, fig.; Lutze, 1938: 165, figs. 1-9; Gottlieb, 1953: 441; Števčić, 1969b: 347.
Astacus tyrrhenus — Monod, 1931: 123.
Callianassa (Cheramus) subterranea — Bouvier, 1940: 102 (partim).
Callianassa (Callichirus) tyrrhena — Zariquiey Alvarez, 1968: 230; Andaloro et al., 1979: 86;
Moncharmont, 1979: 72.
Callianassa (Callianassa) tyrrhena — Felder, 2001: 441.
Pestarella tyrrhena — Ngoc-Ho, 2003: 479, figs. 13, 14; Dworschak, 2004: 15, figs. 2, 3;
Dworschak et al., 2006: 1369, figs. 1, 4, 6, tab. 2; Koller et al., 2006: 113; Robles et al.,
2009: 316.
Not: Alpheus Tyrhenus Risso, 1816: 94 (only references), not pl. 2 fig. 2 (= Pontonia pinnophylax (Otto, 1821).
Remarks. — Dworschak et al. (2000) observed that this species was
collected by yabby pump together with Salmoneus erasimorum Dworschak
et al., 2000 (Alpheidae) in Opatja, Kvarner, Adria.
Type locality. — Naples, Italy, Mediterranean.
Distribution. — Eastern Atlantic Ocean, from Isle of Man to Canary Islands;
and Zarzis, Tunisia, to Tantura, Israel in the Mediterranean Sea; intertidal, 520 m.
Gilvossius whitei (Sakai, 1999)
Callianassa Davyana White, 1847: 70 [not: Gebios Davyana Risso, 1822]; De Man, 1928a: 5,
37.
Callianassa (Callichirus) stebbingi — Pesta, 1918: 201, fig. 63.
Callianassa algerica Lutze, 1938: 168 (partim), fig. 22 (eyestalks = C. whitei) [not: figs. 26a
(smaller Mxp3, not larger Mxp3), 26b (smaller Mxp3, not larger Mxp3), 27] [nomen
dubium].
Callianassa candida — Dworschak, 1992: 194 (partim).
Callianassa whitei Sakai, 1999c: 14 (key), 23, fig. 4a-d; Costello et al., 2001: 289; Dworschak,
2002: 63.
Pestarella whitei — Ngoc-Ho, 2003: 484, fig. 15; Robles et al., 2009: 316.
Diagnosis. — Mxp3 ischium-merus operculiform; merus rounded at distomesial corner. Male Plps1-2 absent. Telson semicircular. [Cf. Sakai, 1999c,
fig. 4b.]
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
381
Type locality. — Cape Monsena, S. Val Salina, Istria near Rovinj, Croatia,
Adriatic Sea.
Distribution. — Mediterranean; rarely found in the Adriatic Sea, Croatia.
Gilvossius sp., Robles et al., 2009
Gilvossius sp. Robles et al., 2009: 316.
Genus Notiax Manning & Felder, 1991 (sensu nov.)
Notiax Manning & Felder, 1991: 772.
Rayllianassa Komai & Tachikawa, 2008: 42. [Type species. — Callianassa amboinensis De
Man, 1888 (= Neaxius ngochoae (Sakai, 1999).]
Diagnosis. — Eyestalks flattened. Mxp3 ischium-merus operculiform, suboperculiform, or subpediform; propodus subovate or subrectangular, and dactylus digitiform. Male Plp1 uniramous, and unsegmented or bisegmented; male
Plp2 uniramous, and unsegmented or bisegmented, lacking appendices interna
and masculina; male Plps3-5 biramous and foliaceous, endopods with stubby
appendix interna. Telson trapezoid or subquadrate. Uropodal exopod with dorsal plate.
Remarks. — Rayllianassa Komai & Tachikawa, 2008 was established on the
basis of Callianassa amboinensis De Man, 1888, but is, however, considered
a synonym of Notiax Manning & Felder, 1991, because the type species
of Rayllianassa is included in Notiax that was established on the basis of
Callianassa brachyophthalma A. Milne-Edwards, 1870, which has the same
type of male Plps1-2 as found in C. amboinensis.
Type species. — Callianassa brachyophthalma A. Milne-Edwards, 1870.
The gender of the generic name, Notiax, is feminine.
Species included. — Notiax amboinensis (De Man, 1888b); N. brachyophthalma (A. Milne-Edwards, 1870); N. ceramica (Fulton & Grant, 1906); N.
delicatula (Rodrigues & Manning, 1992); N. limosa (Poore, 1975).
K EY
TO THE
S PECIES
OF THE
G ENUS NOTIAX
1 – Mxp3 ischium-merus rectangular, 2.0 times as long as wide, or suboperculiform, 1.7-1.8
times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Mxp3 ischium-merus operculiform, 1.2-1.3 times as long as wide . . . . . . . . . . . . . . . . . . 3
2 – A1 peduncle reaching distal margin of A2 penultimate segment. From Port Phillip Bay,
Victoria, Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. limosa
– A1 peduncle reaching distal third of A2 terminal segment. From Chiloé Is., Chile . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. brachyophthalma
3 – Male larger cheliped without meral hook . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
382
CRM 013 – Katsushi Sakai
– Male larger cheliped with meral hook . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4 – Distal segment of A1 peduncle three times as long as penultimate segment . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. amboinensis
– Distal segment of A1 peduncle 1.5-2.0 times as long as penultimate segment . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. ngochoae
5 – Male Plp2 uniramous and unsegmented, or absent. From Port Phillip and Western Port,
Victoria, Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. ceramica
– Male Plp2 uniramous and bisegmented. From Praia do Araça, São Sebastião, Brazil . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. delicatula
Notiax amboinensis (De Man, 1888)
Callianassa amboinensis De Man, 1888b: 480, pl. 20 fig. 4; Zehntner, 1894: 194; Borradaile,
1903: 545.
Callianassa (Calliactites) amboinensis — Borradaile, 1903: 545.
Diagnosis. — Cornea located distolaterally on eyestalk. Distal segment 3
of A1 peduncle 3 times as long as segment 2. Mxp3 ischium-merus rounded
in shape; merus rounded on distomesial margin. Merus of larger cheliped
rounded and serrated on ventral margin, lacking ventroproximal lobe. Male
Plps1-2 unknown. Telson subquadrate.
Remarks. — The present species, Notiax amboinensis (De Man, 1888)
was described as Callianassa amboinensis on the basis of a single damaged
female, and can be characterized as follows, though the type specimen was
lost (M. Türkay, in litt.). The A1 distal segment is distinctly elongate as shown
by De Man (1888, pl. 20 fig. 4); the P1 merus is convex ventrally with a
ventroproximal lobe; and the telson is subsquare. The species, Callianassa
ngochoae Sakai, 1999c was earlier synonymized with C. amboinensis, but
the recent survey has indicated that C. amboinensis is to be distinguished
from C. ngochoae, which is here reclassified under the same genus, Notiax
as N. ngochoae.
Type locality. — Ambon (= Amboina), Indonesia.
Distribution. — Only known from the type locality.
Notiax brachyophthalma (A. Milne-Edwards, 1870)
(fig. 63F-G)
Callianassa brachyophthalma A. Milne-Edwards, 1870: 85; De Man, 1928b: 115; Sakai, 1999c:
31.
Callianassa (Trypaea) brachyophthalma — Borradaile, 1903: 546; De Man, 1928b: 27, 94,
115, 134; Holthuis, 1952: 92, fig. 19; Ferrari, 1981: 16, fig. 2.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
383
Notiax brachyophthalma — Manning & Felder, 1991: 772, figs. 6, 11; Tudge et al., 2000: 143.
Material examined. — USNM 252309, 1 male (TL/CL, 67.0/19.0 mm), River Maullin
Estuary, Lalanguithue, Chile, 26.iv.1990, leg. A. Bravo.
Diagnosis. — Mxp3 ischium-merus operculiform; merus rounded at mesiodistal corner. Male Plp1 uniramous and unsegmented (fig. 63F); male Plp2
uniramous and bisegmented (fig. 63G). Telson trapezoid, truncate on posterior
margin, with median spine. [Cf. Ferrari, 1981, figs. 2-7, 16-18.]
Type locality. — Chiloé Is., Chile.
Distribution. — Only known from the type locality.
Notiax ceramica (Fulton & Grant, 1906)
Callianassa ceramica Fulton & Grant, 1906: 12, pl. 5; Hale, 1927a: 86; Poore, 1975: 205; Poore
& Griffin, 1979: 257, figs. 22, 23; Sakai, 1988: 57; Sakai, 1999c: 41.
Callianassa (Trypaea) ceramica — De Man, 1928b: 27, 93, 104.
Biffarius ceramica — Tudge et al., 2000: 143.
Biffarius ceramicus — Davie, 2002: 457; Poore, 2004: 181, fig. 49d, e.
Diagnosis. — Mxp3 ischium-merus suboperculiform; merus rounded on
mesiodistal margin. Male Plp1 uniramous and bisegmented; male Plp2 uniramous and unsegmented. Telson subquadrate, slightly convex on posterior
margin, with distinct median spine.
Type locality. — Port Phillip and Western Port, Victoria, Australia.
Distribution. — Victoria to the south of Western Australia, intertidal to
shallow subtidal (Poore & Griffin, 1979); Plimmerton, North Island, New
Zealand.
Notiax delicatula Rodrigues & Manning, 1992
Biffarius delicatulus Rodrigues & Manning, 1992a: 324, fig. 1; Rodrigues et al., 1998: 380;
Melo, 1999: 352, figs. 233-234; Tudge et al., 2000: 143; Robles et al., 2009: 316.
Callianassa delicatula — Sakai, 1999c: 27.
Diagnosis. — Mxp3 ischium-merus operculiform; merus rounded on distomesial margin. Male Plp1 uniramous and bisegmented; male Plp2 uniramous
and bisegmented. Telson subquadrate, truncate on posterior margin, without
median spine. [Cf. Rodrigues & Manning, 1992a, fig. 1j, r, s, w.]
Type locality. — Praia do Araça, São Sebastião, Brazil.
Distribution. — São Sebastião, State of São Paulo, Brazil.
384
CRM 013 – Katsushi Sakai
Notiax limosa (Poore, 1975)
Callianassa limosa Poore, 1975: 201-205, figs. 4, 5; Poore & Griffin, 1979: 270, figs. 32, 33.
Neocallichirus limnosa — Sakai, 1988: 61.
Neocallichirus limosus — Sakai, 1999c: 103; Sakai, 2005b: 90.
Biffarius limosa — Tudge et al., 2000: 143.
Biffarius limosus — Davie, 2002: 457; Poore, 2004: 181, fig. 49f, g pl. 12b.
Diagnosis. — Mxp3 ischium-merus subsquare; merus rounded on mesiodistal margin; propodus subovate; dactylus digitiform. Male Plp1 uniramous and
bisegmented; male Plp2 uniramous, minute, and lobed medially. Telson trapezoid, convex on posterior margin, without median spine. [Cf. Poore, 1975:
201-204, figs. 4b, 5g, i, j.]
Remarks. — Poore (1975) described the present species as Callianassa
limosa, stating that “Pleopod 1 (male) uniramous, 2-articles, distal article
tapering. Pleopod 2 (male) a minute medially lobed tapered papilla.” However,
the species was later included in the genus Biffarius by Tudge et al. (2000), and
Davie (2002).
Type locality. — Port Phillip Bay, Victoria, Australia.
Distribution. — Australia — central New South Wales to Tasmania; shallow
water to 100 m.
Notiax ngochoae (Sakai, 1999)
Callianassa amboinensis — Poore & Griffin, 1979: 248, fig. 14; Sakai, 1984: 96, figs. 1, 2;
Sakai, 1988: 53, 57, fig. 1; Ngoc-Ho, 1991: 283, fig. 1; Sakai, 1999c: 38; Tudge et al., 2000:
143; Davie, 2002: 458; Sakai, 2005b: 71; Poore, 2008: 172.
Callianassa (Trypaea) amboinensis — De Man, 1928b: 27, 93, 107, 165, pl. 18 fig. 28-28c.
Callianassa ngochoae Sakai, 1999c: 49 (partim).
Rayllianassa amboinensis — Komai & Tachikawa, 2008: 43, figs. 13-15.
Diagnosis. — Cornea located distolaterally on eyestalks. The distal segment
of A1 peduncle 1.5-2 times as long as segment 2. Mxp3 ischium-merus
rounded in shape; merus rounded on distomesial margin. Merus of larger
cheliped rounded and serrated on ventral margin, lacking ventroproximal
lobe. Male Plp1 uniramous and unsegmented; male Plp2 uniramous and
bisegmented. Telson subquadrate, bearing median spinule on posterior margin.
Remarks. — The genus Rayllianassa Komai & Tachikawa, 2008 was established for Callianassa amboinensis De Man, 1888. However, the type species
of Rayllianassa, C. amboinensis is now referred to as Notiax amboinensis and
is distinguished from C. ngochoae (= N. ngochoae). In Notiax amboinensis
(De Man, 1888), the terminal segment of the A1 peduncle is elongate, and
three times as long as the penultimate segment, as shown in the figure of De
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
385
Man (1888, pl. 20 fig. 4), whereas in N. ngochoae the terminal segment of the
A1 peduncle is 1.5-2 times as long as the penultimate segment, as described
by Poore & Griffin (1979, 2007), Sakai (1984, 1988, 1999c, 2005b), Ngoc-Ho
(1991), and Komai & Tachikawa (2008).
Komai & Tachikawa (2008) described the male Plps1-2 of Callianassa
amboinensis, i.e., in their text but without figures, under their new genus
Rayllianassa as follows: “Male Plp1 is greatly reduced to minute papilla;
Plp2 very small, exopod greatly reduced to rudimentary bud, endopod slightly
shorter than protopod.” This description was based on their examination of
the male specimen of C. amboinensis from New Caledonia with the same
catalogue number (MNHN-Th 1071) as that of the male specimen which
Ngoc-Ho (1991: 283, fig. 1) had examined and figured. However, Komai &
Tachkawa’s (2007) description of the male pleopod 2 is, in spite of examining
the specimen with the same catalogue number, quite different from NgocHo’s (1999) description, who had clearly described, with reference to her
figures, as follows: “Premier pléopode du male (fig. 1kl) à un seul article
arrondi, très petit. Deuxième pléopode du mâle (fig. 1k2) à deux articles
courts et grêles. [Male Plp1 a single segment, rounded, very small. Plp2 with
two short and slender segments]”. It cannot be possible, however, that two
different descriptions were made by examining the male specimen(s) with
the same catalogue number (MNHN-Th 1071), which thus can only indicate
that Komai & Tachikawa (2007) most probably made a misdescription. The
characteristic form of Plp2 observed in the male specimen (MNHN-Th 1071),
shown by Ngoc-Ho’s (1999) description and figures, suggests that NgocHo’s C. amboinensis is included in the genus Notiax, thus in the present
classification to be referred to as N. ngochoae Sakai, 1999, and that the
generic classification under Rayllianassa is not permissible. Therefore, the
type species of Rayllianassa, Callianassa amboinensis is also included in the
genus Notiax, as a synonym of N. ngochoae Sakai, 1999.
Type locality. — Ambon (= Amboina), Indonesia.
Distribution. — Chichi-Jima, Ogasawara Islands (Komai & Tachikawa,
2008: 47); Philippines, 91-183 m. Amboina, off Lirung, Salibabu Island;
Java, reef to 18 m, Indonesia. Northern Territory, Table Head, Port Essington;
Dampier Island, Dampier Archipelago, northern Western Australia, reef to
19 m (Poore, 2008); Heron Island, Queensland, Australia. New Caledonia,
18-80 m off Tombeau Bay, Mauritius. Elat, Israel, Gulf of Aqaba.
Genus Trypaea Dana, 1852 (sensu nov.)
Trypaea Dana, 1852a: 14.
386
CRM 013 – Katsushi Sakai
Biffarius Manning & Felder, 1991: 769; Poore, 2004: 181. [Type species. — Callianassa
biformis Biffar, 1971b.]
Neotrypaea Manning & Felder, 1991: 771. [Type species. — Callianassa californiensis Dana,
1854.]
Nihonotrypaea Manning & Tamaki, 1998: 89. [Type species. — Callianassa japonica Ortmann,
1891.]
Necallianassa Heard & Manning, 1998: 883. [Type species. — Necallianassa berylae Heard &
Manning, 1998.]
Pseudobiffarius Heard & Manning, 2000: 70. [Type species. — Pseudobiffarius caesari Heard
& Manning, 2000.]
Diagnosis. — Mxp3 propodus subovate or subquadrate, and dactylus digitiform. Male Plp1 uniramous, slender, and un-, bi- or trisegmented; male Plp2
absent; male Plps3-5 biramous, endopods with projecting, finger-like to stubby
appendix interna. Uropodal exopod with dorsal plate.
Remarks. — Tudge et al. (2000: 142) and Davie (2002: 457) listed 10
species of Biffarius. However, the type species of Biffarius, Callianassa
biformis Biffar, 1971b has the same features of Plps1-2 as the type species
of Trypaea, T. australiensis Dana, 1852b, so that Biffarius is synonymized
here with Trypaea. Those species included in Biffarius by those authors, are
reclassified by the features of the male Plps1-2; five of these species (B.
arenosus (Poore, 1975); B. biformis (Biffar, 1971b); B. fragilis (Biffar, 1970);
B. lewtonae (Ngoc-Ho, 1994); B. poorei Sakai, 1999b) are transferred to
Trypaea; three species (B. ceramicus (Fulton & Grant, 1906); B. delicatulus
Rodrigues & Manning, 1992a; and B. limosa (Poore, 1975)) are transferred to
Notiax; and one species (B. diaphora (Le Loeuff & Intès, 1974)) is transferred
to Gilvossius.
Type species. — Trypaea australiensis Dana, 1852a, by present designation
and original monotypy. Gender of generic name, Trypaea, feminine.
Species included. — Trypaea acanthura (Caroli, 1946); T. amboinae (Bate,
1888); T. amplimaxilla (Sakai, 2002); T. arenosa Poore, 1975; T. australiensis
Dana, 1852a; T. berylae (Heard & Manning, 1998); T. biffari (Holthuis,
1991); T. biformis (Biffar, 1971b); T. brevirostris (Sakai, 2002); T. caesari
(Heard & Manning, 2000); T. caledonica (Ngoc-Ho, 1991); T. californiensis
(Dana, 1854); T. costaricensis (Sakai, 2005b); T. debilis (Hernández-Aguilera,
1998); T. exilimaxilla (Sakai, 2005b); T. filholi (A. Milne-Edwards, 1878);
T. fragilis (Biffar, 1970); T. gigas (Dana, 1852b); T. intermedia (De Man,
1905); T. japonica (Ortmann, 1891); T. joculatrix (De Man, 1905); T. matzi
(Sakai, 2002); T. melissae (Poore, 2008); T. mocambiquensis (Sakai, 2004b);
T. petalura (Stimpson, 1860); T. plantei (Sakai, 2004b); T. poorei (Sakai,
1999b); T. pugnatrix (De Man, 1905); T. pygmaea (De Man, 1928b); T.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
387
sahul (Poore, 2008); T. santarita (Thatje, 2000); T. sibogae (De Man, 1905);
T. tabogensis (Sakai, 2005b); T. tenuipes (Sakai, 2002); T. thailandica (Sakai,
2005b); T. thermophila (Lin, Komai & Chan, 2007); T. thorsoni (Sakai,
2005b); T. tonkinae (Grebenjuk, 1975); T. truncata (Giard & Bonnier, 1890);
T. uncinata (H. Milne Edwards, 1837a).
Also the following 12 species, which were included in Callianassa, are
tentatively reclassified under Trypaea in the present study, because they are
different in morphological features, though the male Plps1-2 are unknown,
from the type species of Callianassa, Callianassa subterranea, which is
peculiar and, therefore, the only species now retained in the genus Callianassa:
T. acutirostella (Sakai, 1988); T. bangensis (Sakai, 2005b); T. chakratongae
(Sakai, 2002); T. gruneri (Sakai, 1999c); T. lewtonae (Ngoc-Ho, 1994);
T. lignicola (Alcock & Anderson, 1899); T. nieli (Sakai, 2002); T. parva
(Edmondson, 1944); T. propriopedis (Sakai, 2002); T. rochei (Bouvier, 1895);
T. spinoculata (Sakai, 2005b); T. stenomastaxa (Sakai, 2002).
The following key to the species of the genus Trypaea has been split-up
into three sections, according to the species’ currently known geographical
distributions.
K EY
TO THE
S PECIES
OF THE
G ENUS TRYPAEA IN THE
EASTERN
PACIFIC O CEAN
1 – Rostrum triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Rostrum indistinctly developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 – A1 peduncle longer than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– A1 peduncle shorter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 – Telson with posteromedian spine and two pairs of subdistal spines on each lateral margin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. tabogensis
– Telson unarmed on lateral and posterior margins . . . . . . . . . . . . . . . . . . . . . . . . . . T. debilis
4 – P3 propodus square at proximoventral corner . . . . . . . . . . . . . . . . . . . . . . . . . . . T. santarita
– P3 propodus rounded at proximoventral corner . . . . . . . . . . . . . . . . . . . . . . . . . . T. uncinata
5 – Eyestalk reaching distal fifth of A1 proximal segment . . . . . . . . . . . . . . . . . . . . . . T. rochei
– Eyestalk overreaching distal margin of A1 proximal segment . . . . . . . . . . . . . . . . . . . . . . 6
6 – Telson without posteromedian spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. biffari
– Telson with posteromedian spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 – Male larger cheliped with subsquare carpus . . . . . . . . . . . . . . . . . . . . . . . . . T. californiensis
– Male larger cheliped with elongate carpus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. gigas
K EY
TO THE
S PECIES
OF THE
G ENUS TRYPAEA IN THE N ORTH ATLANTIC
ATLANTIC O CEAN
AND IN
THE WESTERN
1 – Rostrum sharply triangular and more than half as long as eyestalks; telson with two
sharp spines on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. berylae
388
CRM 013 – Katsushi Sakai
– Rostrum short and less than half as long as eyestalks; telson unarmed on lateral margin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2 – A1 peduncle longer than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– A1 peduncle about as long as, or shorter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . 4
3 – Telson with median spine on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. caesari
– Telson without median spine on posterior margin . . . . . . . . . . . . . . . . . . . . T. costaricensis
4 – A1 peduncle about as long as A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. fragilis
– A1 peduncle shorter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. biformis
K EY
TO THE
S PECIES
G ENUS TRYPAEA WITH THE EXCEPTION OF THOSE
PACIFIC O CEAN AND IN THE N ORTH ATLANTIC
AND WESTERN ATLANTIC O CEAN
OF THE
OCCURRING IN THE EASTERN
1 – Mxp3 ischium-merus broad, 1.3-1.8 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . 2
– Mxp3 ischium-merus slender, more than twice as long as wide . . . . . . . . . . . . . . . . . . 22
2 – Rostrum acutely triangular, and more than half as long as eyestalks . . . . . . . . . . . . . . . . 3
– Rostrum short and triangular, and less than half as long as eyestalks . . . . . . . . . . . . . . . 7
3 – A1 peduncle as long as, or longer than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– A1 peduncle shorter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. bangensis
4 – A1 peduncle as long as A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. gruneri
– A1 peduncle longer than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Telson with posteromedian spine, from North West Shelf, Western Australia, 82 m . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. acutirostella
– Telson without posteromedian spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – Abdominal somite 6 wider than long, from Britomart Reef, Queensland, reef front,
15 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. lewtonae
– Abdominal somite 6 longer than wide, from Andaman Sea, 9◦ 00′ N 98◦ 03′ E, 40.058.0 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. amplimaxilla
7 – A1 peduncle longer than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– A1 peduncle about as long as, or shorter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . 14
8 – Eyestalks armed with distal spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
– Eyestalks unarmed distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9 – Distal spine of eyestalk distinct; Mxp3 ischium-merus subquadrate . . . . . . . T. thorsoni
– Distal spine of eyestalk short; Mxp3 ischium-merus oval . . . . . . . . . . . . . . . . . . . T. filholi
10 – Eyestalks overreaching distal margin of A1 proximal segment . . . . . . . . . . . . . . . . . . . 11
– Eyestalks not reaching distal margin of A1 proximal segment . . . . . . . . . . . . . . . . . . . . 12
11 – Telson oval in dorsal view, and Mxp3 also oval . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. sahul
– Telson subquadrate in dorsal view, and Mxp3 also subquadrate . . . . . . . . . . . . T. poorei
12 – A1 peduncle extremely elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. australiensis
– A1 peduncle slightly longer than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 – P3 propodus rounded on ventral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. arenosa
– P3 propodus convex on ventral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. melissae
14 – A1 peduncle about as long as A2 peduncle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
– A1 peduncle shorter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
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15 – Uropodal exopod with subdistal spine on lateral margin . . . . . . . . . . . . . . . . . T. truncata
– Uropodal exopod unarmed on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 – Rostrum triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. brevirostris
– Rostrum indistinctly developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 – Carpus of male larger cheliped incurvate on dorsal margin, from Bay of Tokyo . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. japonica
– Carpus of male larger cheliped straight on dorsal margin . . . . . . . . . . . . . . . . T. petalura
18 – Eyestalks armed with distal spine, from W. Malay Peninsula . . . . . . . . . . T. spinoculata
– Eyestalks unarmed distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 – Uropodal endopod and telson each with distinct distolateral spine, from Bay of Naples
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. acanthura
– Uropodal endopod and telson each unarmed distolaterally . . . . . . . . . . . . . . . . . . . . . . . 20
20 – Meral hook of male larger cheliped absent, from Hanauma Bay, Oahu . . . . . . . T. parva
– Meral hook of male larger cheliped present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 – Mxp3 ischium divergent distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. thermophila
– Mxp3 ischium subsquare, from Andaman Sea, 185-244 m . . . . . . . . . . . . . . . T. lignicola
22 – Rostrum short and obtusely triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. thailandica
– Rostrum sharply triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 – Rostrum less than half as long as eyestalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
– Rostrum elongate, reaching almost tip of eyestalk, or overreaching proximal half of
eyestalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
24 – Telson armed with median spine on posterior margin . . . . . . . . . . . . . . . . . . . . . . . T. nieli
– Telson unarmed on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 – P3 propodus convex on distal margin. . . . . . . . . . . . . . . . . . . . . . . . . . . T. mocambiquensis
– P3 propodus distinctly concave on distal margin . . . . . . . . . . . . . . . . . . . . . . . T. joculatrix
26 – Rostrum almost reaching tip of eyestalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
– Rostrum overreaching proximal half of eyestalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
27 – P3 propodus subsquare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
– P3 propodus elongate and oval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
28 – Telson medially concave on posterior margin . . . . . . . . . . . . . . . . . . . . . . T. stenomastaxa
– Telson straight on posterior margin, bearing median spine . . . . . . . . . . . T. propriopedis
29 – Mxp3 ischium-merus narrowing distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. exilimaxilla
– Mxp3 ischium-merus obliquely truncate at distal end . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30 – A1 peduncle shorter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
– A1 peduncle subequal to A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
31 – Uropodal endopod armed with subdistal spine on lateral margin . . . . . . . . T. amboinae
– Uropodal endopod unarmed on anterior margin . . . . . . . . . . . . . . . . . . . . . . . . . T. sibogae
32 – A1 peduncle shorter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. intermedia∗ )
– A1 peduncle reaching distal end of A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33 – Uropodal endopod bearing subdistal spine on lateral margin . . . . . . . . . . . . . T. pygmaea
– Uropodal endopod smooth on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . T. pugnatrix
34 – A1 peduncle longer than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
– A1 peduncle about as long as A2 peduncle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36
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35 – Mxp3 ischium-merus narrowing distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. tenuipes
– Mxp3 ischium-merus obliquely truncate at distal end . . . . . . . . . . . . . . . . . T. caledonica
36 – Mxp3 ischium-merus obliquely truncate at distal end . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
– Mxp3 ischium-merus narrowing distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
37 – Telson subsquare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. sp. (Rabalais et al., 1981)
– Telson trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. tonkinae
38 – Uropodal exopod concave on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. matzi
– Uropodal exopod straight on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39 – Carpus of male larger cheliped 0.8-1.0 times as broad as long . . . . . . . T. chakratongae
– Carpus of male larger cheliped 1.4-1.8 times as broad as long . . . . . . . . . . . . . T. plantei
∗ ) In T. intermedia (De Man, 1905) the description of the Mxp3 is missing, as shown in the
original descriptions by De Man (1905, 1928b), which causes great difficulty in composing the
key to the species. However, the key can still be made by adopting, as a clue, the similarity of
T. intermedia to T. pugnatrix (De Man, 1905) and to T. pygmaea (De Man, 1928).
Trypaea acanthura (Caroli, 1946)
Callianassa acanthura Caroli, 1946: 66, figs. 1a, 2; Holthuis, 1953a, fig. 3; De Saint Laurent
& Božić, 1976: 21, figs. 3, 11, 19, 25, 30; Türkay, 1982: 225; d’Udekem d’Acoz, 1996:
54; Abed-Navandi & Dworschak, 1998: 605; Sakai, 1999c: 14; Costello et al., 2001: 289;
Ngoc-Ho, 2003: 466, fig. 8.
Callianassa (Trypaea) acanthura — Zariquiey Alvarez, 1968: 229.
Neocallianassa acanthura — Heard & Manning, 1998: 884; d’Udekem d’Acoz, 1999: 155;
Tudge et al., 2000: 143.
Type locality. — Bay of Naples.
Distribution. — Bay of Naples; Adriatic Sea: Kornati Archipelago and
Vestar Bay, near Rovinj, Croatia; 3-6 m; Ionian Sea; Aegean Sea.
Trypaea acutirostella (Sakai, 1988)
Callianassa acutirostella Sakai, 1988: 57, fig. 2; Sakai, 1999c: 37; Tudge et al., 2000: 143;
Davie, 2002: 458; Sakai, 2005b: 64, figs. 13, 14.
Diagnosis. — Rostrum acutely pointed, almost reaching distal end of eyestalks. Mxp3 ischium-merus suboperculiform, merus rounded on distomesial
margin. Male Plps1-2 unknown. Uropodal endopod without distolateral spine.
Telson almost straight on posterior margin, with median spine.
Type locality. — Western Australia, North West Shelf (19◦ 05.1′ S
118◦ 53.7′ E), 82 m.
Distribution. — North West Shelf of Western Australia, and Arafura Sea.
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391
Trypaea amboinae (Bate, 1888)
Scallasis amboinae Bate, 1888: 34, pl. 2 figs. 3, 4; Manning & Felder, 1991: 780 (list); Tudge
et al., 2000: 145.
Callianassa (Scallasis) amboinae — Borradaile, 1903: 547; De Man, 1928a: 93.
Callianassa (Scallasis) Amboinae — De Man, 1928b: 30.
Callianassa amboinae — Sakai, 1999c: 37, fig. 5d-f; Sakai, 2005b: 68, figs. 15, 16.
Diagnosis. — Mxp3 ischium-merus subpediform; merus obliquely truncate
with rounded mesiodistal corner. Male Plp1 uniramous and bisegmented; male
Plp2 absent. Telson concave on posterior margin, without median spine.
Remarks. — Scallasis amboinae was reclassified under the genus Callianassa by Sakai (1999c: 37; 2005b: 68), because it was found by examining
the type specimen (MNH 1888.22) of Scallasis amboinae that it is different
from Bate’s (1888) figure as follows: the eyestalks are not globose but triangular, the Mxp3 ischium-merus is not slender but rectangular, and lacking an
exopod (Sakai, 1999c, fig. 5f). However, the species is reclassified now under
Trypaea, because the male Plp2 is absent.
Type locality. — Ambon (= Amboina), Indonesia.
Distribution. — Bay of Nhatrang, Vietnam (Sakai, 1997); Ambon, Indonesia (Bate, 1888; De Man, 1928); East Lagoon, New Caledonia (Ngoc-Ho,
1991).
Trypaea amplimaxilla (Sakai, 2002)
Callianassa amplimaxilla Sakai, 2002: 501, figs. 22A-D, 23A-D; Sakai, 2005b: 72.
Type locality. — Andaman Sea, 9◦ 00′ N 98◦ 03′ E, 40.0 m.
Distribution. — Andaman Sea, 7◦ 35.995′ N 98◦ 25.119′ E; 7◦ 59.839′ N
◦
98 13.625′ E; 9◦ 00.009′ N 98◦ 02.962′ E; 9◦ 14.939′ N 97◦ 54.212′ E; 40.0-58.0 m;
muddy sand, sandy mud.
Trypaea arenosa Poore, 1975
Callianassa arenosa Poore, 1975: 197-201, figs. 1, 2; Poore & Griffin, 1979: 250, figs. 15-17;
Sakai, 1988: 57; Sakai, 1999c: 39; Sakai, 2005b: 73.
Biffarius arenosa — Tudge et al., 2000: 142; Davie, 2002: 457; Poore, 2004: 181, fig. 49b, d
pl. 12a.
Diagnosis. — Mxp3 ischium-merus suboperculiform; merus rounded mesiodistally. P3 propodus largely convex on ventral margin. Male Plp1 uniramous and bisegmented; male Plp2 absent. Telson almost straight on posterior
margin, without median spine. [Cf. Poore & Griffin, 1979, fig. 15b.]
Type locality. — Port Phillip Bay, Victoria, Australia.
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Distribution. — Moreton Bay, Queensland; New South Wales; Victoria;
Tasmania; intertidal to shallow water.
Trypaea australiensis Dana, 1852
Trypaea australiensis Dana, 1852a: 19; Dana, 1852b: 513; Dana, 1855, pl. 32 fig. 4; Fulton &
Grant, 1906: 14; Manning & Felder, 1991: 774, figs. 1, 3, 12; Tudge et al., 2000: 143; Davie,
2002: 461.
Trypaea porcellana Kinahan, 1856: 130, pl. 4 fig. 2. [Type locality. — Port Phillip, Victoria,
Australia.]
Callianassa (Trypaea) porcellana — Borradaile, 1903: 546.
Callianassa (Trypaea) australiensis — Borradaile, 1903: 546; De Man, 1928b: 27, 93, 104,
134; Stephenson et al., 1931: 56; Dakin & Colefax, 1940: 182-184, figs. 270, 271; Gurney,
1944: 83, figs. 8, 9; Dakin et al., 1952: 199, pl. 44; Hailstone & Stephenson, 1961: 259-285,
figs. 1-15 pls. 1-3; Hailstone, 1962: 29-31, 2 figs.; McNeill, 1968: 26; Healy & Yaldwyn,
1970, pl. 30.
Callianassa australiensis — Poore & Griffin, 1979: 250, figs. 18-20; Sakai, 1988: 57; Holthuis,
1991: 241, 264, figs. 441, 442; Debelius, 1999: 227, figs.; Sakai, 1999c: 39.
Diagnosis. — A1 distal segment distinctly long, overreaching A2 peduncle
at proximal fifth. Mxp3 ischium-merus suboperculiform; merus distinctly
protruded at mesiodistal angle. Male Plp1 uniramous and bisegmented; male
Plp2 absent. Telson almost straight on posterior margin, without median spine.
Type locality. — Illawarra District, New South Wales, Australia.
Distribution. — Townsville to Port Phillip Bay, eastern Australia; intertidal
(Poore & Griffin, 1979).
Trypaea bangensis (Sakai, 2005)
Callianassa bangensis Sakai, 2005b: 74, figs. 17, 18.
Diagnosis. — Rostrum triangular and pointed apically in dorsal view;
frontal margin of carapace smooth. Eyestalks rounded distally, shorter than
rostrum, and directed downward distally, overreaching basal segment of A1
peduncle. A1 peduncle shorter than A2 peduncle. Mxp3 merus-ischium broad
and oval; merus rounded on distomesial margin; propodus subovate and
dactylus digitiform. Male larger cheliped massive; ischium denticulate on
ventral margin, but unarmed on dorsal margin; merus oval, dorsal margin
arched and smooth, ventral margin arched and denticulate; carpus short, broad,
and rounded on proximoventral margin; chela stout. P3 propodus ovate and
rounded at ventroproximal corner. Plps1-2 unknown. Telson slightly longer
than broad and shield-like, unarmed on convergent lateral margins, and lacking
median spine on straight posterior margin. Uropodal endopod subquadrate,
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393
longer than telson, and rounded distally; uropodal exopod broad, longer than
endopod, and obliquely truncate distally. [Adapted from Sakai, 2005b.]
Type locality. — Six miles of Port Banga, Mindanao, Philippines, 50 m.
Distribution. — Known only from the type locality.
Trypaea berylae (Heard & Manning, 1998)
Neocallianassa berylae Heard & Manning, 1998: 884, figs. 1-3; Tudge et al., 2000: 143; Heard
et al., 2007: 22, fig. 20.
Callianassa berylae — Sakai, 1999c: 129; Sakai, 2005b: 44.
Material examined. — USNM 260881, holotype, male (TL/CL, 18.0/4.5 mm), South
Carolina, U.S.[A.], 43 m, viii.1977, leg. G.W. Pierce.
Diagnosis. — Rostrum sharply triangular, not reaching distal margin of
eyestalks. Mxp3 ischium-merus operculiform; merus broadly rounded at mesiodistal angle (Heard & Manning, 1998, fig. 2f). Male Plp1 uniramous and
bisegmented; male Plp2 uncertain, but probably absent. Telson almost straight
on posterior margin, with median spine, bearing two distinct, posteriorlydirected subdistal spines on lateral margin. [Adapted from Heard & Manning,
1998: 888.]
Type locality. — South Carolina, 32◦ 00′ 57′′ N 79◦ 31′ 03′′ W, 43 m, sand.
Distribution. — Only known from the type locality.
Trypaea biffari Holthuis, 1991
Callianassa affinis Holmes, 1900: 162, pl. 2 figs. 29-30; Rathbun, 1904: 154; Schmitt, 1921:
119, fig. 81; Stevens, 1928: 341, fig. 18; Haig & Abbott, 1980: 580, pl. 166, fig. 24.3.
[A junior primary homonym of Callianassa affinis A. Milne-Edwards, 1861, a fossil species.]
Callianassa (Callichirus) affinis — Borradaile, 1903: 547.
Callianassa (Trypaea) affinis — De Man, 1928b: 27, 101.
Neotrypaea affinis — Manning & Felder, 1991: 771.
Callianassa biffari Holthuis, 1991: 242, fig. 243 (partim, C. tabogensis sp. nov.). [New name
for Callianassa affinis Holmes, 1900.]
Neotrypaea biffari — Tudge et al., 2000: 143.
Type locality. — Point Loma, California, U.S.A.
Distribution. — Santa Monica Bay, Los Angeles, San Diego and San
Clemente Is., U.S.A.; Ensenada and San Quantin Bay, Baja California, Mexico; Panama; on beaches.
Trypaea biformis (Biffar, 1971)
Callianassa biformis Biffar, 1971b: 225, fig. 1; Rabalais et al., 1981: 101; Manning, 1987: 397;
Squires, 1990: 349, figs. 184-186; Sakai, 1999c: 26; Sakai, 2005b: 44.
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Biffarius biformis — Coelho, 1997: 147; Melo, 1999: 348, figs. 230-231; Tudge et al., 2000:
143; Coelho et al., 2007: 5 (table); Heard et al., 2007: 20, fig. 26.
Material examined. — USNM 135053, paratype, male (TL/CL, 20.0/4.5 mm), south end,
mouth of Doboy Sound, Sapelo Island, Georgia, U.S.[A.], 01.i.1969, leg. H. Heard, det.
T. Biffar.
Diagnosis. — Mxp3 ischium-merus operculiform; merus rounded at mesiodistal angle. Male Plp1 uniramous and bisegmented; male Plp2 absent.
Telson slightly rounded on posterior margin, without median spine.
Type locality. — Mouth of Doboy Sound, south end of Sapelo Island,
McIntosh County, Georgia, sandy lower intertidal.
Distribution. — Bass River, Yarmouth, Massachusetts to Florida, western
Atlantic; Gulf of Mexico; sandy lower intertidal; Itamaracá and Casa Caiada,
Brazil (Coelho, 1997).
Trypaea brevirostris (Sakai, 2002)
Callianassa brevirostris Sakai, 2002: 514, figs. 30A-E, 31A-I.
Diagnosis. — Male Plp1 uniramous and bisegmented; male Plp2 absent.
Telson longer than wide, bearing median concavity, with spine on posterior
margin.
Type locality. — Andaman Sea, 7◦ 52′ N 98◦ 42′ E, 21 m, sand with shell
fragments.
Distribution. — Andaman Sea, 6◦ 59.913′ N 99◦ 23.822′ E; 9◦ 30.351′ N
97◦ 57.168′ E; 17.0-60.7 m, mud, fine sand, sand with shell fragments.
Trypaea caesari (Heard & Manning, 2000)
Pseudobiffarius caesari Heard & Manning, 2000: 71, figs. 1, 2, 3a-k, n-o, 4, 5a.
Callianassa caesari — Sakai, 2005b: 44.
Diagnosis. — Rostrum small and triangular. Eyestalks flattened and contiguous along median line, slightly overreaching A1 proximal segment. A1
peduncle slightly longer than A2 peduncle. Mxp3 ischium-merus operculiform
and 1.3-1.8 times as long as wide; merus rounded at distomesial angle. Male
Plp1 uniramous and bisegmented; male Plp2 absent. Telson subquadrate, bearing shallow median concavity, with spine on posterior margin. [From Heard &
Manning, 2000, figs. 3i, k, 4g.]
Type locality. — Buccoo Reef, 11◦ 11′ N 60◦ 49′ W, Tobago.
Distribution. — Northwest corner of Man O’War Bay (11◦ 19′ N 60◦ 34′ W),
Lover’s Beach, Tobago, ca. 2 m; Pirate’s Cove, east side of Man O’War
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
395
Bay; Buccoo Reef (11◦ 11′ N 60◦ 49′ W); Coral Gardens, Buccoo Reef; Pigeon
Point (11◦ 10′ N 60◦ 51′ W); Lowlands Lagoon (= Petit Trou; 13◦ 50′ N 61◦ 05′ W,
1-3 m).
Trypaea caledonica (Ngoc-Ho, 1991)
Callianassa caledonica Ngoc-Ho, 1991: 285, fig. 2.
Diagnosis. — Mxp3 ischium-merus rectangular; merus slightly concave
on distal margin, with rounded mesiodistal angle. P3 propodus oval, with
protruded ventroproximal angle. Male Plp1 uniramous and small; male Plp2
absent. Telson wider than long, bearing median concavity with small spine on
posterior margin.
Remarks. — Callianassa caledonica Ngoc-Ho, 1991 is characterized as
follows. The A1 peduncle is longer than the A2 peduncle, the Mxp3 ischiummerus is subpediform, abdominal somite 6 is elongate, the uropodal endopod
bears a spine on the lateral margin, and the uropodal exopod is broadened and
longer than the endopod, bearing a spine on the lateral margin. However, those
morphological features mentioned above are not decisive in determining the
genus to which C. caledonica should belong. What is more important is that
in C. caledonica the male Plp1 is present, and uniramous and small, and the
male Plp2 is absent, as in the type species of Trypaea, T. australiensis, which
suggests that C. caledonica may safely be included in Trypaea.
Type locality. — East Lagoon, New Caledonia, 21 m.
Distribution. — Only known from the type locality.
Trypaea californiensis (Dana, 1854)
Callianassa californiensis Dana, 1854: 175; Stimpson, 1857: 4, 89, pl. 21 fig. 4; Stimpson,
1860: 24; A. Milne-Edwards, 1870: 82, 101; Lockington, 1878: 301; Bouvier, 1895: 8;
Holmes, 1900: 159, pl. 2 fig. 27; Rathbun, 1904: 154; Hilton, 1916: 63; Schmitt, 1921: 116,
117, fig. 78; Stevens, 1928: 325, 333, figs. 10-13, 16-17, 55-71; MacGinitie, 1934: 166-176,
pls. 5, 6; MacGinitie, 1935: 709, fig. 14; Haig & Abbott, 1980: 579, pl. 166 fig. 24.2; Hart,
1982: 58, fig. 15; Holthuis, 1991: 244, 264, figs. 445, 446; Dworschak, 1992: 192, fig. 2a, c,
e; Sakai, 1999c: 32.
Callianassa occidentalis Stimpson, 1856: 88; Manning, 1987: 399.
Callianassa (Trypaea) californiensis — Borradaile, 1903: 546; De Man, 1928b: 27, 105.
Neotrypaea californiensis — Manning & Felder, 1991: 771, fig. 10; Feldman et al., 2000: 141;
Tudge et al., 2000: 143; Atkinson & Taylor, 2004: 45, tab. 2.
Not: Callianassa (Trypaea) californiensis — Parisi, 1917: 23 (= Trypaea japonica).
Diagnosis. — Rostrum inconspicuous. Eyestalks elongate and triangular,
overreaching A1 proximal segment. Mxp3 ischium-merus operculiform;
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merus rounded at mesiodistal corner. Male Plp1 uniramous and bisegmented;
male Plp2 absent. Telson almost straight on posterior margin, with or without
median spine.
Type locality. — California.
Distribution. — Mutiny Bay, Alaska; Vancouver Is., British Columbia, to
mouth of Tia Juana River, San Diego, California; and Bahia de San Quintin,
Mexico. Inhabits sandy sediments in the upper tidal zone (Dworschak, 1992:
196).
Trypaea chakratongae (Sakai, 2002)
Callianassa chakratongae Sakai, 2002: 513, figs. 28A-D, 29A-G; Sakai, 2004b: 574-581,
figs. 9-12; Sakai, 2005b: 80.
Diagnosis. — Rostrum pointed apically. Eyestalks rounded distally, and
directed downward distally, corneae dispersed on eyestalks. A2 peduncle about
as long as A1 peduncle. Anterolateral margin of carapace without spines.
Mxp3 ischium-merus elongate and pediform, more than twice as long as wide.
P3 propodus straight on ventral margin, and convex at posteroventral angle.
Male Plps1-2 unknown. Telson convergent posteriorly.
Type locality. — Andaman Sea, 9◦ 00.062′ N 97◦ 53.366′ E, 65.4 m, muddy
sand.
Distribution. — Only known from the type locality.
Trypaea costaricensis (Sakai, 2005)
Callianassa costaricensis Sakai, 2005b: 51, figs. 5-10.
Diagnosis. — P3 propodus protruded proximoventrally and rectangular.
Male Plp1 uniramous and bisegmented; male Plp2 absent.
Type locality. — Golfo de Nicoya, Bahia Herradura, Costa Rica, GN-50KG
(09◦ 32.38′ N 84◦ 32.41′ W), 45 m.
Distribution. — Only known from the type locality.
Trypaea debilis (Hernández-Aguilera, 1998)
Biffarius debilis Hernández-Aguilera, 1998: 303, fig. 1; Tudge et al., 2000: 143; Hendrickx et
al., 2005: 170.
Callianassa debilis — Sakai, 2005b: 57.
Diagnosis. — Mxp3 ischium-merus subsquare; merus rounded at mesiodistal angle. Male Plp1 uniramous and bisegmented; male Plp2 absent. Telson
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397
concave on posterior margin, without median spine. [Cf. Hernández-Aguilera,
1998, fig. 1h, i, l.]
Type locality. — Revillagigedo Archipelago, 18◦ 20′ 52′′ N 114◦ 43′ 43′′ W,
Azufre Bay, Clarión Island.
Distribution. — Revillagigedo Archipelago, Pacific side of Mexico.
Trypaea exilimaxilla (Sakai, 2005)
Callianassa exilimaxilla Sakai, 2005b: 80, figs. 19, 20, tab. IV.
Diagnosis. — Mxp3 ischium-merus narrow and pediform. Male Plp1 uniramous and bisegmented; male Plp2 absent; male Plp3 endopod with projecting,
finger-like appendix interna. Telson convergent distally, and almost straight on
posterior margin, with median spine.
Type locality. — South China Sea, 05◦ 09′ N 106◦ 47′ E.
Distribution. — Only known from the type locality.
Trypaea filholi (A. Milne-Edwards, 1878)
Callianassa Filholi A. Milne-Edwards, 1878: 112; Filhol, 1886: 491, pl. 53 figs. 10-12;
Borradaile, 1903: 548; Robles et al., 2009: 316.
Callianassa filholi — Chilton, 1907: 461-464, pl. 16 figs. 1-5; Miller & Batt, 1973: 110;
Berkenbusch et al., 1998: 55; Sakai, 1999c: 43, fig. 7a-c; Berkenbusch et al., 2000: 397;
Tudge et al., 2000: 143; Sakai, 2005b: 85.
Callianassa (Trypaea) Filholi — De Man, 1928b: 27, 101, 104.
Type locality. — Stewart Is.
Distribution. — Timaru; Oamaru; Stewart Is., New Zealand.
Trypaea fragilis (Biffar, 1970)
Callianassa fragilis Biffar, 1970: 45, fig. 3; Biffar, 1971a: 667, figs. 7, 8; Manning, 1987: 397;
Sakai, 1999c: 27; Sakai, 2005b: 45.
Biffarius fragilis — Manning & Felder, 1991: 769; Coelho, 1997: 147; Melo, 1999: 352, figs.
233-234; Tudge et al., 2000: 143; Coelho et al., 2007: 5 (table).
Biffarius cf. fragilis — Heard et al., 2007: 21, fig. 27.
Diagnosis. — Mxp3 ischium-merus subsquare; merus straight distally and
rounded at mesiodistal angle. P3 propodus protruded proximoventrally and
rectangular. Male Plp1 uniramous and bisegmented; male Plp2 absent. Telson
almost straight but slightly concave medially on posterior margin, without
median spine. [Cf. Biffar, 1971a: 667, figs. 7, 8.]
Type locality. — Puerto Rico, Punta Arenas, sandy flat.
Distribution. — Miami, southeastern Florida; Puerto Rico; Antigua; Venezuela, sandy flat; Itamaracá, Brazil (Coelho, 1997).
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CRM 013 – Katsushi Sakai
Trypaea gigas (Dana, 1852)
Callianassa gigas Dana, 1852a: 19; Dana, 1852b: 512; Dana, 1855, pl. 32 fig. 3; Stimpson,
1857b: 489, pl. 21 fig. 3; A. Milne-Edwards, 1870: 81, 101; Lockington, 1878: 302; Holmes,
1900: 162; Rathbun, 1904: 154; Schmitt, 1921: 116, 119, fig. 80; Stevens, 1928: 325,
figs. 6, 9, 14-15, 38-54; MacGinitie, 1935: 712; Haig & Abbott, 1980: 579; Hart, 1982:
56, fig. 14; Holthuis, 1991: 245, 264, figs. 447, 448; Dworschak, 1992: 194, fig. 2b, d, f;
Sakai, 1999c: 32; Sakai, 2005b: 57.
Callianassa longimana Stimpson, 1857a: 86; Stimpson, 1857b: 490, pl. 21 fig. 5; A. MilneEdwards, 1870: 83, 101; Lockington, 1878: 302; Neumann, 1878: 34; Holmes, 1900: 161,
pl. 2 fig. 28; Rathbun, 1904: 154; Hilton, 1916: 63, fig. 14; Schmitt, 1921: 116, 117, fig. 79.
[Type locality. — Puget Sound, Washington State, U.S.A.]
Callianassa (Trypaea) gigas — Borradaile, 1903: 546; De Man, 1928b: 27, 101, 134, 180, 181.
Callianassa (Trypaea) longimana — Borradaile, 1903: 546; De Man, 1928b: 27, 102, 106, 134.
Neotrypaea gigas — Tudge et al., 2000: 143.
Diagnosis. — Rostrum reduced. Eyestalks shaped in elongate triangle. A1
peduncle about as long as A2 peduncle. Mxp3 ischium-merus subsquare, and
1.3-1.8 times as long as wide; merus rounded on distomesial angle. Meral
hook of male larger cheliped convex on ventral margin. Carpus of male larger
cheliped straight on dorsal margin. Male Plp1 uniramous and bisegmented;
male Plp2 absent. Telson square, bearing small median spine on posterior
margin.
Type locality. — Puget Sound.
Distribution. — Vancouver Island, British Columbia; Puget Sound; Elkhorn
Slough, Monterey Bay, San Juan Island, Poulsbo, Allyn, Washington; San
Francisco, California to San Quentin Bay and Gulf of Farallones. Lower tidal
in muddy sediments.
Trypaea gruneri (Sakai, 1999)
Callianassa mucronata — Tirmizi, 1977 (part.): 21, fig. 1b. [Not: Callianassa mucronata
Strahl, 1862.]
Callianassa gruneri — Sakai, 1999c: 44, fig. 8a-g; Sakai, 2005b: 86.
Diagnosis. — Mxp3 ischium-merus subsquare; merus straight, and slanting
mesially on distal margin. P3 propodus convex proximoventrally. Male Plps1-2
uncertain. Telson straight on posterior margin, without median spine.
Type locality. — Philippines, Luzon.
Distribution. — Only known from the type locality.
Trypaea intermedia De Man, 1905
Callianassa intermedia De Man, 1905: 609; Sakai, 1999c: 46; Tudge et al., 2000: 143; Sakai,
2005b: 86.
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399
Callianassa (Cheramus) intermedia — De Man, 1928b: 26, 98, 143, pl. 14 figs. 21-21d.
Not: Callianassa subterranea intermedia forma; Czerniavsky, 1884: 80 [nomen dubium].
Diagnosis. — Rostrum sharply pointed, slightly overreaching eyestalks.
A1 peduncle much shorter than A2 peduncle. Mxp3 ischium-merus narrow
and more than twice as long as wide. P3 propodus subquadrate, swelling at
proximoventral angle. Male Plp1 uniramous and bisegmented; Plp2 absent.
Telson bearing median concavity with spine on posterior margin.
Type locality. — Indonesia, Bali Sea (7◦ 46′ S 114◦ 30.5′ E), 330 m.
Distribution. — Bali Sea (7◦ 46′ S 114◦ 30.5′ E), Indonesia; 330 m.
Trypaea japonica (Ortmann, 1891)
Callianassa subterranea var. japonica Ortmann, 1891: 56, pl. 1 fig. 10a; Bouvier, 1901: 332334; Doflein, 1902: 644; Balss, 1914: 91; Nakazawa, 1927: 1039, fig. 1999; Yokoya, 1930:
543; Kikuchi, 1932: 7; Yokoya, 1933: 52; Miyazaki, 1936: 317-320, figs. 1-3; Kamita, 1957:
107-109, fig. 49.
Callianassa californiensis var. japonica Bouvier, 1901: 332. [Type locality. — Japan.]
Callianassa Harmandi Bouvier, 1901: 332-334. [Type locality. — Japan.]
Callianassa (Trypaea) Harmandi — Borradaile, 1903: 546; Parisi, 1917: 24, fig. 7; De Man,
1928a: 13-15, fig. 6-6j; De Man, 1928b: 27, 102-103; Yü, 1931: 92-93, fig. 3.
Callianassa (Trypaea) japonica — Borradaile, 1903: 546; De Man, 1928a: 19-22, pl. 5 figs. 1010a; De Man, 1928b: 27, 93, 106; Yü, 1931: 95-96, fig. 5; Makarov, 1938: 69-71, fig. 25.
Callianassa (Trypaea) californiensis — Parisi, 1917: 23. [Not: Callianassa californiensis Dana,
1854.]
Callianassa hermandi — Nakazawa, 1927: 1039, fig. 2000.
Callianassa (Trypaea) californiensis var. japonica — De Man, 1928a: 18-19, pl. 4 fig. 9-9e; De
Man, 1928b: 27, 105; Yü, 1931: 94, fig. 4.
Callianassa (Trypaea) harmandi — Makarov, 1938: 66-67, figs. 22, 23.
Callianassa (Trypaea) californiensis var. bouvieri Makarov, 1938: 71-72, fig. 26. [Replacement
name for C. californiensis var. japonica.]
Callianassa japonica — Nakazawa & Kubo, 1947: 754, fig. 2174; Vinogradov, 1950: 224;
Shiino, 1951: 87; Miyake, 1965: 633, fig. 1037 (illustrated by K. Sakai); Sakai, 1968: 23, fig. 8; Sakai, 1969: 232, pls. 9-12; Miyake, 1982: 92, pl. 31 fig. 4; Sakai, 1987a: 303;
Holthuis, 1991: 246, figs. 449, 450; Dworschak, 1992: 198; Liu & Zhong, 1994: 562 (list);
Tamaki et al., 1996: 675, tabs.; Tamaki et al., 1997: 223; Miyabi et al., 1998: 101; Sakai,
1999c: 46; Sakai, 2001: 937-948, figs. 1-4, table 1; Sakai, 2002, fig. 15D-F; Sakai, 2004:
113, fig. 4; Sakai, 2005b: 87; Sakai & Sawada, 2006: 1357.
Callianassa harmandi — Nakazawa & Kubo, 1947: 754, fig. 2173; Vinogradov, 1950: 224;
Shiino, 1951: 87; Liu, 1955: 63, pl. 23 figs. 1-5; Utinomi, 1956: 63, pl. 32 fig. 2; Miyake et
al., 1962: 124.
Callianassa petalura — Liu, 1955, pl. 23 fig. 6-9; Holthuis, 1991, fig. 453 (larger cheliped of
male and female); Debelius, 1999: 277, fig. [Not: Callianassa petalura Stimpson, 1860.]
Nihonotrypaea japonica — Manning & Tamaki, 1998: 889, fig. 1; Tamaki et al., 1999: 37;
Tamaki & Miyabe, 2000: 182; Tudge et al., 2000: 133, 143; Sakai, 2001: 946; Tamaki, 2003:
116; Atkinson & Taylor, 2004: 45, tab. 2; Itani, 2004, fig. 3; Tamaki, 2004a: 87; Tamaki &
Harada, 2006: 65; Itani, 2007: 202; Lin et al., 2007: 153.
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CRM 013 – Katsushi Sakai
Callianassa ?japonica — Yamaguchi & Holthuis, 2001: 112-113, referring to Suiken-figs. 2, 3.
Nihonotrypaea harmandi — Tudge et al., 2000: 143; Tamaki, 2003: 116; Fuji-ie, W. et al.,
2004: 77; Itani, 2004, fig. 3; Tamaki, 2004a: 87; Tamaki & Harada, 2006: 65; Itani, 2007:
202.
Material examined. — SMF 32732 (7 males with the larger cheliped of harmandi-form,
29.0/6.3-34.0/7.7 mm; 11 males with the larger cheliped of japonica- and C. californiensis
var. japonica-types, 19.0/4.5-34.0/8.0 mm; 2 ovig. females, 22.0/5.1 mm; 20 females, 21.0/4.837.0/7.7 mm; 9 damaged males and females), Warabi-I, Izumi-city, Kagoshima Pref., Japan,
2.vi.1989, leg. H. Mukai; SMF 32733, 2 males (35.0/7.7-35.0/4.4 mm with larger chelipeds of
harmandi-form), Okinosu beach, Tokushima, Japan, 30.x.2005, leg. H. Sora.
Diagnosis. — Male Plp1 uniramous and bisegmented; male Plp2 absent.
Telson straight on posterior margin with median spine.
Remarks. — Specimens of the present species were found abundantly in
the sandy-mud tidal zone, and their larger chelipeds are variable in shape,
which has caused numerous synonymous species as shown in the list of
synonyms. It was found through the examination of the specimens that
each collection includes individuals with dactyli in varied forms and also
with different proportions in length of the carpus to the palm, and they
are almost the same morphologically with one another, except for those
differences found in the male larger cheliped. Those specimens were grouped
into three types by the morphological features of their male larger chelipeds
(Sakai, 1969, table 9): an A-group represented by C. japonica; a B-group
by Bouvier’s C. californiensis var. japonica; and a C-group by Bouvier’s
C. harmandi, and they were safely considered to belong to one species,
C. japonica, with morphological variations in the features of the male larger
cheliped, because it seems improbable that each group of those individuals
collected together at each locality and grouped into three types has its
own habitat. This fact was proven morphologically by re-examining the
syntypes (MNHN Th 70, registered as C. californiensis var. japonica Bouvier,
1901, from the surroundings of Tokyo Bay, Japan; MNHN Th 80, registered
as C. Harmandi Bouvier, 1901, from Tokyo Bay, Japan; MNHN Th 496,
registered as C. japonica Ortmann, 1891, leg. Franck, 1895, from Japan;
MNHN Th 507, registered as C. japonica Ortmann, 1891, leg. Franck, from
Japan) (Sakai, 2001: 938). Though Wardiatino & Tamaki (2001) described
C. japonica and C. harmandi as two valid species by the difference in size
of their cornea alone (Wardiatino & Tamaki, 2001, fig. 1), this has turned
out to be incorrect, because it is confirmed by the present examination of the
material (SMF32732, from Warabi I. Izumi-city, Kagoshima) that the cornea
of C. japonica is the same as that of C. harmandi, namely the cornea is round
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401
in both these species. Itani (2007: 202, in Japanese) is skeptical about the
statistical results of measurement of morphological features by Wardiatono
& Tamaki (2001), mentioning that it is not always easy to make a clear
distinction in morphology between Nihonotrypaea (= Callianassa) japonica
and Nihonotrypaea (= Callianassa) harmandi, which are found distributed
all over Japan, though each of them has its own locality, because their eyes
and rostrums change in width (the former) and in shape (the latter) according
to their growth, and the lively dispute over Nihonotrypaea (= Calianassa)
harmandi, that is, whether it is a valid species or not, will be settled if genetic
exchanges (hybrids) will be confirmed between those two species collected in
the same locality.
After my rebuttal (Sakai, 2001, 2004) against Manning & Tamaki (1998),
Tamaki & Miyabe (2000), and Wardiatino & Tamaki (2001), yet Tamaki
(2004), and Tamaki et al. (2006) reported two species, C. japonica and C. harmandi with their data, without paying attention to my earlier paper (Sakai,
1969). They classified in vain the specimens collected together at each locality into two species, because each collection includes individuals whose male
larger chelipeds are morphologically intermediate, just as Bouvier’ collection
from Tokyo Bay (Bouvier, 1901: 332) preserved in one bottle, whose specimens were confirmed to belong to one species, C. japonica (cf. Sakai, 2001:
938), though they had once been classified into three species, Callianassa subterranea var. japonica Ortmann, 1891; Callianassa californiensis var. japonica Bouvier, 1901; and Callianassa Harmandi Bouvier, 1901 (cf. Sakai, 2001:
938).
Type locality. — Bay of Tokyo, Japan.
Distribution. — Peter the Great Bay, intertidal mud flats in bays and
estuaries to 192 m. Nemuro, Funka Bay, Hokkaido to Kagoshima, Kyushu Is.,
on both Pacific side and Japan Sea. Hou Hai, Yellow Sea (Shantung Peninsula
to southwestern coast of Korean peninsula).
Trypaea joculatrix (De Man, 1905)
Callianassa joculatrix De Man, 1905: 610; Poore & Griffin, 1979: 266, figs. 28, 29; Sakai,
1988: 53 (key); Ngoc-Ho, 1991: 287, fig. 3; Liu & Zhong, 1994: 562; Ngoc-Ho, 1994: 51;
Sakai, 1999c: 47; Komai, 2000b: 345 (list); Tudge et al., 2000: 143; Davie, 2002: 458; Sakai,
2005b: 88; Komai & Tachikawa, 2008: 30, figs. 7-9.
Callianassa (Cheramus) joculatrix — De Man, 1928b: 18, 26, 93, 95, 98, 130-137, 141, 146,
148, 151, 153, pl. 12 fig. 19, 19c, pl. 13 fig. 19a, d-m (non pl. 12 fig. 19b); McNeil, 1968: 26.
Diagnosis. — Rostrum acutely triangular. Eyestalks protruded mesiodistally; corneae dispersed on eyestalks. A2 peduncle about as long as A1 peduncle. Mxp3 ischium-merus narrow and more than twice as long as wide;
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CRM 013 – Katsushi Sakai
merus not protruded at distomesial angle. Merus of larger cheliped with proximal spine on dorsal margin. P3 propodus concave on ventral margin. Male
Plp1 uniramous and bisegmented; Plp2 absent. Telson trapezoid and straight
on posterior margin, without median spine.
Type locality. — Bay of Labuan Tring (8◦ 44.5′ S 116◦ 02.5′ E), west coast of
Lombok, Indonesia, 18-27 m.
Distribution. — N. South China Sea; Taiwan; Philippines; Bonin Islands,
Chichi-jima (Komai & Tachikawa, 2008); Indonesia (Lombok, Java Sea);
Arafura Sea, northern Queensland; north-west and north-east Australia; New
Caledonia; South Vietnam; Mombasa; 15-300 m.
Trypaea lewtonae (Ngoc-Ho, 1994)
Callianassa lewtonae Ngoc-Ho, 1994: 52, fig. 1; Sakai, 1999c: 47; Sakai, 2005b: 90.
Biffarius lewtonae — Tudge et al., 2000: 143; Davie, 2002: 457.
Diagnosis. — Rostrum pointed apically, overreaching proximal half of
eyestalks. A1 peduncle slightly longer than A2 peduncle. Mxp3 ischium-merus
subsquare; merus rounded on distomesial margin. Male Plps1-2 unknown.
Abdominal somite 6 wider than long. Telson almost straight on posterior
margin, without median spine.
Type locality. — Queensland, Britomart Reef, reef front (18◦ 17′ S 146◦ 38′ E),
15 m.
Distribution. — Only known from the type locality.
Trypaea lignicola (Alcock & Anderson, 1899)
Callianassa lignicola Alcock & Anderson, 1899: 288; Alcock, 1900, pl. 42 fig. 2, 2a, 2b;
Alcock, 1901: 200; Borradaile, 1903: 545; Balss, 1925: 212; Sakai, 1999c: 48; Tudge et
al., 2000: 143; Sakai, 2005b: 90.
Callianassa (Calliactites) lignicola — De Man, 1928b: 25, 97.
Diagnosis. — Rostrum small and triangular, less than half as long as eyestalks. A1 peduncle shorter than A2 peduncle. Mxp3 ischium-merus suboperculiform; merus rounded at distomesial angle. Mxp3 ischium-merus subsquare
and 1.3-1.8 times as long as wide. Meral hook of male larger cheliped small.
Male Plps1-2 unknown. Telson straight on posterior margin, without median
spine. Uropodal endopod and telson without distinct distolateral spine, respectively. [Adapted from Alcock & Anderson, 1899: 289; Alcock, 1900, pl. 42
fig. 2, 2b.]
Type locality. — Andaman Sea, 185-244 m.
Distribution. — Only known from the type locality.
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Trypaea matzi (Sakai, 2002)
Callianassa matzi Sakai, 2002: 506, figs. 26A-C, 27A-I; Sakai, 2005b: 92.
Diagnosis. — Rostrum triangular and pointed apically. Eyestalks rounded
distally and directed downward distally; corneae dispersed on eyestalks. A2
peduncle about as long as A1 peduncle. Mxp3 ischium-merus slender. Merus
of larger cheliped unarmed on dorsal margin. P3 propodus concave on ventral
margin. Male Plp1 uniramous and bisegmented; Plp2 absent. Telson trapezoid
and almost straight on posterior margin, with minute median spine.
Type locality. — Andaman Sea, 7◦ 59.839′ N 98◦ 13.625′ E, 41.8 m, sandy
mud.
Distribution. — Andaman Sea, 6◦ 43.303′ N 99◦ 03.304′ E; 6◦ 45.961′ N
◦
99 20.968′ E; 9◦ 14.939′ N 97◦ 54.212′ E; 9◦ 00.091′ N 97◦ 43.147′ E; 20.5-83.3 m,
mud, fine sand, muddy sand, with shell fragments.
Trypaea melissae (Poore, 2008)
Biffarius melissae Poore, 2008: 170, fig. 4.
Diagnosis. — Rostrum triangular. Eyestalks with broadly triangular apical
lobes; cornea pigmented and domed. A1 peduncle slightly longer than A2
peduncle. Mxp3 ischium-merus operculiform and two times as long as greatest
width, crista dentata with 18 teeth; carpus, propodus subovate, and dactylus
digitiform. P1 (female) unequal and dissimilar. In larger cheliped, ischium
finely denticulate on ventral margin; merus with denticulate distally-directed
proximal tooth on ventral margin; carpus, propodus, and dactylus unarmed. In
smaller cheliped, merus with small tooth on ventral margin. Male Plp1 simple
and bisegmented. Female Plp1 trisegmented, segment 1 curved, segment 2
one-third as long as segment 1, segment 3 tapering, shorter than segment
2; female Plp2 biramous with curved basis, endopod bisegmented, as long
as exopod. Telson subquadrate. Uropodal endopod oval, with straight lateral
margin; uropodal exopod almost semicircular, with rounded distal and mesial
margins, and barely concave on lateral margin, bearing dorsal plate.
Remarks. — Poore (2008) selected a female as the type specimen and
described the female Plps1-2. However, he did not describe the Plp2 of the
male paratype, which is an important element in the present classification. It
is thus supposed that the male Plp2 is absent in Poore’s Biffarius melissae.
Therefore, his species is safely reclassified under the genus Trypaea (in the
present classification), in which the male Plp2 is absent.
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CRM 013 – Katsushi Sakai
Type locality. — Western Australia, Dampier Archipelago; N.W. of Cohen
Is. (20◦ 22.14′ S 116◦ 46.13′ E), 32-35 m.
Distribution. — Only known from the type locality.
Trypaea mocambiquensis (Sakai, 2004)
Callianassa mocambiquensis Sakai, 2004b: 585-592; Sakai, 2005b: 93.
Diagnosis. — Rostrum short and obtusely triangular in dorsal view. Eyestalks with small cornea. A1 peduncle about as long as A2 peduncle. Mxp3
ischium-merus slender and more than twice as long as wide. P1 unequal; larger
cheliped unarmed, fingers slightly longer than palm and crossed with each
other distally; smaller cheliped also unarmed, fingers longer than palm and
crossed with each other distally. P3 propodus oval, swollen at posteroventral
angle, and straight on ventral margin. Male Plp1 uniramous and bisegmented;
Plp2 absent. Telson subtrapezoid.
Type locality. — Mocambique Channel, 26.0 m.
Distribution. — Only known from the type locality.
Trypaea nieli (Sakai, 2002)
Callianassa nieli Sakai, 2002: 518, fig. 32A-E; Sakai, 2005b: 94.
Diagnosis. — Rostrum obtusely triangular and less than half as long as
eyestalks. Eyestalks rounded distally. A1 peduncle about as long as A2
peduncle. Mxp3 ischium-merus more than twice as long as wide; merus
rounded on mesiodistal margin. Male Plps1-2 undescribed. Telson almost
straight on posterior margin, with median spine. [Cf. Sakai, 2002: 518,
fig. 32C, D.]
Type locality. — Andaman Sea, 7◦ 36.100′ N 98◦ 19.257′ E, 55.4 m.
Distribrution. — Andaman Sea, 7◦ 34.598′ N 98◦ 16.250′ E; 7◦ 36.100′ N
◦
98 19.257′ E; 7◦ 35.995′ N 98◦ 25.119′ E; muddy sand, 48.9-68.8 m.
Trypaea parva (Edmondson, 1944)
Callianassa (Calliactites) parva Edmondson, 1944: 45, fig. 5a-j; Edmondson, 1946: 261.
Callianassa parva — Sakai, 1999c: 50; Tudge et al., 2000: 143; Sakai, 2005: 95.
Diagnosis. — Male Plp1 uniramous and bisegmented, Plp2 undescribed.
[Cf. Edmondson, 1944: 45, fig. 5b, g, i.]
Remarks. — Edmondson (1944) described the male Plp1 as uniramous and
bisegmented, but the Plp2 is undescribed, probably absent, so that this species
is included in Trypaea.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
405
Type locality. — Hanauma Bay, Oahu.
Distribution. — Hanauma Bay and Palos Bay, Hawaii; Celebes (= Sulawesi),
Indonesia.
Trypaea petalura (Stimpson, 1860)
Callianassa petalura Stimpson, 1860: 23; A. Milne-Edwards, 1870: 88, 101; Bouvier, 1901:
332-334; De Man, 1928b: 115; Yokoya, 1939: 277-278; Liu, 1955: 65, pl. 23 figs. 6-9
(= Callianassa japonica); Miyake et al., 1962: 124; Miyake, 1965: 633, fig. 1036 (illustrated
by K. Sakai); Sakai, 1968: 2-3, fig. 9; Sakai, 1969: 233, pls. 13-15; Miyake, 1982: 91, pl. 31
fig. 3; Sakai, 1987a: 303; Holthuis, 1991: 249, 264 (not figs. 453, 454 = C. japonica); Liu
& Zhong, 1994: 562 (list); Sakai, 1999c: 50.
Callianassa (Trypaea) petalura — Borradaile, 1903: 546; De Man, 1928b: 28, 115.
Callianassa subterranea var. japonica — Balss, 1914: 91.
Callianassa subterranea japonica — Kikuchi, 1932: 7.
Callianassa (Trypaea) gigas var. japonica Makarov, 1935: 323-324, fig. 4. [Type locality. —
Patrocle Bay, Peter the Great Bay.]
Callianassa (Trypaea) gigas var. eoa Makarov, 1938: 67-69, fig. 24; Vinogradov, 1950: 223.
[New name for Callianassa (Trypaea) gigas var. japonica Makarov, 1938.]
Nihonotrypaea petalura — Tudge et al., 2000: 143; Itani, 2004, fig. 3; Shimoda & Tamaki,
2004: 23, figs. 1-6, tabs. 1, 2.
Not: Callianassa petalura — Debelius, 1999: 227, fig. (= Trypaea japonica).
Diagnosis. — Male Plp1 uniramous and bisegmented; Plp2 absent.
Type locality. — Japan, Shimoda.
Distribution. — Japan, from Hokkaido to Kyushu Is. on both Pacific side
and Japan Sea, on sandy mud flats, facing open sea; Ho Hai, Yellow Sea,
shallow water (Liu & Zhong, 1994).
Trypaea plantei (Sakai, 2004)
Callianassa plantei Sakai, 2004b: 592-599, figs. 18-23; Sakai, 2005b: 99.
Diagnosis. — Male Plp1 uniramous and bisegmented; Plp2 absent; Plps3-5
endopods with projecting, finger-like appendix interna.
Type locality. — Mozambique Channel, 30 m.
Distribution. — Mozambique Channel; 16-64 m.
Trypaea poorei (Sakai, 1999)
Callianassa poorei Sakai, 1999b: 373, figs. 1, 2; Sakai, 2005b: 100.
Biffarius poorei — Davie, 2002: 457.
Diagnosis. — Rostrum obtusely triangular and less than half as long
as eyestalks. A1 peduncle longer than A2 peduncle. Eyestalks flattened
and contiguous along median line; cornea distinct. Mxp3 ischium-merus
406
CRM 013 – Katsushi Sakai
suboperculiform and 1.6 times as long as wide; merus rounded on mesiodistal
margin. In male larger cheliped, ischium denticulate on ventral margin; meral
hook acutely triangular. P3 propodus convex on ventral margin. Male Plp1
uniramous and bisegmented; Plp2 absent. Telson subquadrate, and truncate on
posterior margin, without a median spine. [Cf. Sakai, 1999b: 373, figs. 1C,
2A.]
Type locality. — East coast of Tasmania (42◦ 09′ S 148◦ 10′ E), Great Oyster
Bay, Tasmania, Australia.
Distribution. — Only known from the type locality.
Trypaea propriopedis (Sakai, 2002)
Callianassa propriopedis Sakai, 2002: 522, fig. 34A-F; Sakai, 2005b: 101.
Diagnosis. — Rostrum acutely triangular and as long as eyestalks. Eyestalks obtusely angulate distally, bearing rounded cornea in centre. A1 peduncle about as long as A2 peduncle. Mxp3 ischium-merus slender and more
than twice as long as wide; merus rounded on distomesial margin. P3 propodus short, subsquare, or subtriangular. Male Plps1-2 undescribed. Telson subsquare and straight on posterior margin, with median spine. [Cf. Sakai, 2002:
522, fig. 34C, D.]
Type locality. — Andaman Sea, 7◦ 59.956′ N 98◦ 38.587′ E, 17.0 m, sand with
shell fragments.
Distribution. — Only known from the type locality.
Trypaea pugnatrix (De Man, 1905)
Callianassa pugnatrix De Man, 1905: 611; Sakai, 1999c: 51; Tudge et al., 2000: 143;
Sakai, 2005b: 101.
Callianassa (Cheramus) pugnatrix — De Man, 1928b: 27, 93, 99, 138, 146, 151, pl. 15 fig. 2323a, pl. 16 fig. 23b-e.
Diagnosis. — Rostrum narrow and protruded, not reaching distal end of
eyestalks. Eyestalks obtuse distally; cornea small and located medially. A1
peduncle about as long as A2 peduncle. Mxp3 ischium-merus subpediform
and more than twice as long as wide; merus slightly concave on distal margin
and rounded on mesial margin. Merus of larger cheliped with sharp spine
proximoventrally. P3 propodus oval. Male Plp1 uniramous and bisegmented;
Plp2 absent. Telson bearing median concavity with acute spine on posterior
margin.
Type locality. — Anchorage off Djangkar, Java, Indonesia, 7◦ 46′ S
114◦ 30.5′ E, “Siboga” Sta. 5, 330 m.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
407
Distribution. — Indonesia — off Djangkar, Java (De Man, 1928b); 330 m;
off Tranquebar, India; 75 m.
Trypaea pygmaea (De Man, 1928)
Callianassa (Cheramus) pygmaea De Man, 1928b: 1, 18, 27, 93, 99, 155, pl. 16 fig. 24-24g.
[Type locality. — Ambon Anchorage, 54 m.]
Callianassa amboinae — Sakai, 1999c: 37 [not C. pygmaea, misidentified].
Callianassa pygmaea — Tudge et al., 2000: 143; Sakai, 2005b: 102.
Diagnosis. — Rostrum acutely triangular, not reaching distal end of eyestalks. Eyestalks pointed distally; cornea distinct and located distolaterally. A1
peduncle about as long as A2 peduncle. Mxp3 ischium-merus narrow and more
than twice as long as wide; merus concave on distal margin and convex on lateral margin. P3 propodus shaped as elongate oval. Male Plp1 uniramous and
bisegmented; male Plp2 absent. Telson subsquare, posterior margin bearing
median concavity, with minute spine. Uropodal exopod and endopod with a
spine on each lateral margin. [Cf. De Man, 1928b, pl. 16 fig. 24-24b, c, g.]
Type locality. — Ambon anchorage, 54 m.
Distribution. — Only known from the type locality.
Trypaea rochei (Bouvier, 1895)
Callianassa Rochei Bouvier, 1895: 7.
Callianassa (Trypaea) Rochei — De Man, 1928a: 17, fig. 8-8d; De Man, 1928b: 28, 104.
Callianassa rochei — Sakai, 1999c: 33.
Neotrypaea rochei — Tudge et al., 2000: 143.
Diagnosis. — A1 peduncle slightly longer than A2 peduncle. Mxp3
ischium-merus suboperculiform; merus rounded at distomesial angle. Male
Plps1-2 undescribed. Telson undescribed.
Type locality. — Baja California.
Distribution. — Only known from the type locality.
Trypaea sahul (Poore, 2008)
Callianassa sahul Poore, 2008: 172, fig. 5.
Diagnosis. — Rostrum broadly triangular. Eyestalks with triangular apical
lobes; cornea pigmented and domed. A1 peduncle longer than A2 peduncle.
Mxp3 ischium-merus operculiform, and 1.5 times as long as greatest width;
ischium with crista dentata with 25 teeth; carpus and propodus subovate (not
expanded); dactylus short and digitiform; exopod absent. P1 (female) equal
and similar; ischium finely denticulate on ventral margin; merus straight and
408
CRM 013 – Katsushi Sakai
unarmed on ventral margin; carpus, propodus, and dactylus unarmed; fixed
finger with proximal blade ending in distally-directed tooth, separated from
apex of finger by notch. P3 propodus with posteriorly-produced rounded lobe.
Abdominal somite 1, 1.5 times length of abdominal somite 2; somites 35 with small, inconspicuous lateral tufts of setae. Female Plp1 uniramous
and bisegmented; female Plp2 biramous, endopod bisegmented; Plps3-5 with
appendix interna. Telson as long as wide, and rounded on posterolateral
angles. Uropodal endopod oval with convex lateral margin. Uropodal exopod
subcircular with rounded distomesial margin, bearing dorsal plate. [Adapted
from Poore, 2008.]
Remarks. — Unfortunately, male specimens of the present species are not
reported, which makes it difficult to classify this species exactly. However, the
present species is safely reclassified under the genus Trypaea, as most of the
species in Callianassa are transferred into Trypaea in the present classification,
because the present species with its equal and similar chelipeds is different
from C. subterranea, the type species of the genus Callianassa, in which the
chelipeds are unequal and dissimilar.
Type locality. — Timor Sea, Sahul Bank (10◦ 40.26′ S, 125◦ 04.84′ E), 18 m.
Distribution. — Only known from the type locality.
Trypaea santarita (Thatje, 2000)
Notiax santarita Thatje, 2000: 289, figs. 1-6, table 1.
Callianassa santarita — Sakai, 2005b: 58.
Diagnosis. — Rostrum triangular and less than half as long as eyestalks.
Eyestalks almost contiguous along median line; cornea distinct. A1 peduncle
shorter than A2 peduncle. Mxp3 ischium-merus oval and 1.8 times as long as
wide. P3 propodus protruded at ventroproximal corner. Male Plp1 uniramous
and bisegmented; Plp2 absent. Telson square and straight on posterior margin,
with median spine. [Cf. Thatje, 2000: 295, 297, figs. 3B, 6C.]
Type locality. — Seno Ponsonby, 55◦ 12.0′ S 68◦ 87.03′ W, Chile, sublittoral,
65 m, mud to sandy-mud.
Distribution. — Seno Ponsonby, Chile.
Trypaea sibogae (De Man, 1905)
Callianassa Sibogae De Man, 1905: 613
Callianassa (?Cheramus) sibogae — De Man, 1928b: 124.
Callianassa (Cheramus) sibogae — De Man, 1928b: 27, 98, pl. 11 fig. 17-17e.
Callianassa sibogae — Ngoc-Ho, 1994: 54, fig. 3; Sakai, 1999c: 52; Sakai, 2005b: 102.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
409
Cheramus sibogae — Tudge et al., 2000: 145; Davie, 2002: 460.
Diagnosis. — Male Plp1 uniramous and bisegmented; male Plp2 absent.
[Cf. Ngoc-Ho, 1994: 56, fig. 3c, h.]
Type locality. — Indonesia (7◦ 46′ S 114◦ 30′ E), 330 m.
Distribution. — Indonesia (7◦ 46′ S 114◦ 30.5′ E), 330 m; North-West Shelf,
Australia; benthic, continental shelf, continental slope, 134-330 m.
Trypaea spinoculata Sakai, 2005
Callianassa spinoculata Sakai, 2005b: 103, figs. 22, 23.
Type locality. — W. Malay Peninsula, 07◦ 00′ N 99◦ 22′ E, 27 m.
Distribution. — Only known from the type locality.
Trypaea stenomastaxa (Sakai, 2002)
Callianassa stenomastaxa Sakai, 2002: 496, fig. 20A-G; Sakai, 2005b: 108.
Remarks. — Mxp3 and P3 propodus are similar to those of Trypaea
thailandica Sakai, 2005b. Males unknown.
Type locality. — Andaman Sea, 8◦ 00.046′ N 98◦ 29.383′ E, 19.0 m, sand with
shell fragments.
Distribution. — Only known from the type locality.
Trypaea tabogensis (Sakai, 2005)
Trypaea tabogensis Sakai, 2005b: 58, figs. 11, 12, tab. III.
Diagnosis. — Male Plp1 uniramous and bisegmented; male Plp2 absent;
male Plps3-5 endopods with projecting, finger-like appendix interna.
Type locality. — Tobaga, Bahia de Panama, Panama.
Distribution. — Only known from the type locality.
Trypaea tenuipes Sakai, 2002
Callianassa tenuipes Sakai, 2002: 528, fig. 38A-D; Sakai, 2005b: 108.
Diagnosis. — Mxp3 ischium-merus narrow and pediform. Male Plp1 uniramous, as thick protrusion; male Plp2 absent. Telson convergent posteriorly,
bearing median spine on straight posterior margin. [Cf. Sakai, 2002: 528,
fig. 38C, D.]
Type locality. — Andaman Sea, 41-61 m.
Distribution. — Only known from the type locality.
410
CRM 013 – Katsushi Sakai
Trypaea thailandica (Sakai, 2005)
Callianassa thailandica Sakai, 2005b: 108, figs. 24, 25.
Diagnosis. — Male Plp1 uniramous and trisegmented; male Plp2 absent;
Plps3-5 endopods with projecting, finger-like appendix interna.
Remarks. — In the original description by Sakai (2005b), male Plps3-5
endopods would bear a stubby appendix interna, but as shown in fig. 25G of
Sakai (2005b), they actually bear a projecting, finger-like appendix interna.
Type locality. — Gulf of Thailand, 13◦ 13′ N 100◦ 34′ E.
Distribution. — Only known from the type locality.
Trypaea thermophila (Lin, Komai & Chan, 2007)
Nihonotrypaea thermophila Lin et al., 2007a: 143, figs. 1-5A, B.
Remarks. — Lin et al. (2007a) described Nihonotrypaea thermophila as a
new species under the genus Nihonotrypaea Manning & Tamaki, 1998, though
Nihonotrypaea was already synonymized with Callianassa (cf. Sakai, 1999c,
2004, 2005) because of its variety of intermediate forms of the Mxp3 ischium
and merus. In the present revision, however, their N. thermophila is reclassified
under Trypaea by the features of the male Plps1-2, as T. thermophila.
Trypaea thermophila is very similar to Trypaea japonica (Ortmann, 1891),
but different geographically and morphologically, because T. thermophila was
collected around the hydrothermal vents off northeastern Taiwan at depths of
128-320 m; the A1 peduncle is distinctly shorter than the A2 peduncle, and
the Mxp3 merus and ischium are the same in length; while T. japonica is
distributed in the shallow waters around Japan and the East China Sea, its A1
peduncle is as long as the A2 peduncle, and the Mxp3 merus is shorter than
the Mxp3 ischium.
Type locality. — Dasi fishing ground, off Kueishan Island, northeastern
Taiwan, 24◦ 52.33′ N 121◦ 59.52′ E, 320 m.
Distribution. — Only known from the type locality.
Trypaea thorsoni (Sakai, 2005)
Callianassa thorsoni Sakai, 2005b: 114, figs. 26, 27.
Diagnosis. — Male Plp1 uniramous and bisegmented; male Plp2 absent.
Type locality. — Bahrein Lightship, Persian Gulf, 27◦ 03′ N 51◦ 02′ E, 71 m.
Distribution. — Only known from the type locality.
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411
Trypaea tonkinae (Grebenjuk, 1975)
Callianassa (Scallasis) tonkinae Grebenjuk, 1975: 302, fig. 3.
Callianassa tonkinae — Sakai, 1999c: 52; Tudge et al., 2000: 143; Sakai, 2002: 503, fig. 24A-C,
25A-G; Sakai, 2005b: 118.
Diagnosis. — Rostrum acutely triangular, slightly overreaching proximal
half of eyestalks. Eyestalks obtuse on distal margin and directed downward
distally. A1 peduncle about as long as A2 peduncle. Mxp3 ischium-merus
subpediform and more than twice as long as wide; merus straight on distal
margin and slightly convex on mesial margin. Merus of larger cheliped with
proximoventral tooth followed distally by denticles. Male Plp1 uniramous and
bisegmented; Plp2 absent. Telson trapezoid, bearing median concavity with
spine on posterior margin.
Type locality. — Tonkin Bay.
Distribution. — Tonkin Bay, South China Sea; Andaman Sea, 7◦ 45.452′ N
97◦ 57.951′ E; 9◦ 00.062′ N 97◦ 53.366′ E; 65.4-70.0 m, muddy sand, coarse and
fine sand; East Lagoon, New Caledonia, 21 m.
Trypaea truncata Giard & Bonnier, 1890
Callianassa truncata Giard & Bonnier, 1890: 362, figs. 2, 4; Caroli, 1939: 73; Caroli, 1940: 73;
Lutze, 1941: 34; Reverberi, 1942a: 89; Reverberi, 1942b: 225; Caroli, 1946: 66, fig. 1b,
3; Holthuis, 1953a: 91, fig. 4; Lagardère, 1966: 195, pls. 2-5; Doldopolski, 1969: 286;
Faure, 1970: 751; Lagardère, 1973: 94; De Saint Laurent & Božić, 1976: 19, figs. 2, 10,
18, 29; Beaubrun, 1979: 84, figs. 56, 57, 62-64; Zavodnik, 1979: 313; Kocatas, 1981: 161162; Mikashavidze, 1981: 1415; Thessalou-Legaki & Zenetos, 1985: 309; Thessalou-Legaki,
1986: 183; Petrescu & Balasescu, 1995: 99; d’Udekem d’Acoz, 1996: 54; Ziebis et al., 1996a:
619; Ziebis et al., 1996b: 227; Abed-Navandi & Dworschak, 1997: 565, figs. 1-9; Sakai,
1999c: 20; Costello et al., 2001: 289; Ngoc-Ho, 2003: 473, fig. 11; d’Udekem d’Acoz, 2003,
fig. website; Makarov, 2004: 294, fig. 137, table 18; Sakai, 2005b: 39.
Callianassa (Trypaea) truncata — Borradaile, 1903: 546; Bouvier, 1940: 102, fig. 68; Gurney,
1942: 244; Zariquiey Alvarez, 1950: 81, fig. 1, pl. 2 figs. 1-6, pl. 3 fig. 2; Dolgopol’skaia,
1954: 186, figs. 5-9; Zariquiey Alvarez, 1968: 229.
Callianassa italica Parisi, 1915: 64, figs. 1, 2. [Type locality. — Italy.]
Callianassa (Trypaea) italica — De Man, 1928a: 11, figs. 5-5h; De Man, 1928b: 27, 101.
?Callianassa truncata — Dolgopol’skaia, 1954: 186, figs. 5, 6; Dolgopol’skaia, 1969: 316, pls.
35-38; Kobyakova & Dolgopol’skaia, 1969: 286.
Callianassa (Trypaeta) truncata — Băcescu, 1967: 229, fig. 104A, D, d, E (B, C after Giard &
Bonnier, 1890) [misspelling].
Necallianassa truncata — d’Udekem d’Acoz, 1999: 156; Tudge et al., 2000: 143.
Diagnosis. — Rostrum triangular and less than half as long as eyestalks.
Eyestalks reaching distal end of A1 proximal segment; cornea distinct. A1
peduncle about as long as A2 peduncle. Mxp3 ischium-merus oval; merus
412
CRM 013 – Katsushi Sakai
rounded on distal margin. Male Plp1 uniramous and bisegmented; Plp2 absent.
Telson trapezoid, bearing small median spine on posterior margin. Uropodal
endopod with distolateral spine. [Cf. Ngoc-Ho, 2003, fig. 11D, J, L.]
Type locality. — Gulf of Naples, Italy.
Distribution. — From Atlantic Ocean: Bay of Biscay, France, 44-57 m
depth, to Romanian coast, south-eastern Black Sea, 15-40 m depth.
Trypaea uncinata (H. Milne Edwards, 1837)
Callianassa uncinata H. Milne Edwards, 1837a: 310, pl. 25 fig. 1; Nicolet, 1849: 208; GuérinMéneville, 1857: 43; A. Milne-Edwards, 1870: 83, 101; Czerniavsky, 1884: 76; Solar et al.,
1970: 20; Sakai, 1999c: 33, fig. 3f.
Callianassa chilensis A. Milne-Edwards, 1860: 302, pl. 16 fig. 2a, 2A; A. Milne-Edwards,
1870: 84, 101; Tudge et al., 2000: 143. [Type locality. — Chile.]
Callianassa (Trypaea) uncinata — Borradaile, 1903: 546.
Callianassa (Trypaea) chilensis — Borradaile, 1903: 546; De Man, 1928a: 15, fig. 7-7c; De
Man, 1928b: 27, 94, 103.
Neotrypaea uncinata — Manning & Felder, 1991: 771; Tudge et al., 2000: 143.
Diagnosis. — A1 peduncle longer than A2 peduncle. Mxp3 ischium-merus
suboperculiform; merus rounded on mesiodistal angle. Male Plp1 uniramous
and bisegmented; male Plp2 absent. Telson concave on posterior margin, with
median spine.
Type locality. — Coast of Chile.
Distribution. — Only known from the type locality.
Trypaea sp. (Rabalais, Holt & Flint, 1981)
Callianassa sp., Rabalais et al., 1981: 106, fig. 4; Manning, 1987: 397; Sakai, 1999c: 30.
Diagnosis. — Rostrum triangular. Mxp3 ischium-merus subpediform. Male
Plps1-2 unknown. Telson convex proximally and rounded on posterior margin,
bearing broad median concavity with spine. [Cf. Rabalais et al., 1981, fig. 4E,
B.]
Distribution. — N.W. Gulf of Mexico.
Subfamily C ALLICHIRINAE Manning & Felder, 1991 (sensu nov.)
Callichirinae Manning & Felder, 1991: 775; Tudge et al., 2000: 136; Ngoc-Ho, 2002a: 540;
Ngoc-Ho, 2003: 486 (partim); Sakai, 2005b: 120.
Diagnosis. — Rostrum developed or reduced, lacking rostral carina. Carapace with dorsal oval; cardiac prominence present or not; cardiac sulcus(i)
present or not. Eyestalks flattened and contiguous. A2 scaphocerite developed
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
413
as a small process. Mxp3 ischium-merus operculiform, rectangular, or pediform; propodus broadened; dactylus subovate or digitiform; exopod present or
not. P1 chelate, unequal or subequal in size, and similar or dissimilar in shape,
lacking meral hook in both chelipeds. P2 chelate. P3 propodus broadened. P4
simple. P5 chelate. Abdominal somite 6 lacking lateral projections. Male Plp1
present or not; male Plp2 uniramous or biramous, endopod with appendices
interna and masculina, with either appendix interna or appendix masculina, or
without appendices interna and masculina; male Plps3-5 biramous and foliaceous, and endopods with appendix interna in both sexes. Uropodal exopod
with dorsal plate. [Adapted from Sakai, 2005b.]
Type genus. — Callichirus Stimpson, 1866.
Genera included. — Balsscallichirus gen. nov., Barnardcallichirus gen.
nov., Callichirus Stimpson, 1866; Corallichirus sensu nov.; Forestcallichirus
gen. nov.; Glypturoides gen. nov.; Glypturus Stimpson, 1866; Grynaminna
Poore, 2000; Lepidophthalmoides gen. nov.; Lepidophthalmus Holmes, 1904;
Michaelcallianassa Sakai, 2002; Neocallichirus Sakai, 1988; Podocallichirus
Sakai, 1999c; Sergio sensu nov.; Thailandcallichirus gen. nov.; and Tirmizicallichirus gen. nov.
K EY
TO THE
G ENERA
OF THE
S UBFAMILY C ALLICHIRINAE
1 – Abdominal somites 3-5 ornamented dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Abdominal somites 3-5 not ornamented dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 – Male and female Plp2 biramous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callichirus
– Male and female Plp2 uniramous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Michaelcallianassa
3 – Male Plp2 uniramous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Male Plp2 biramous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4 – Mxp3 propodus straight on ventral margin . . . . . . . . . . . . . . . Forestcallichirus gen. nov.
– Mxp3 propodus rounded on ventral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Male Plp1 uniramous and un- or bisegmented, distal segment simple . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balsscallichirus gen. nov.
– Male Plp1 uniramous and bisegmented, distal segment subchelate distally . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Barnardcallichirus gen. nov.
6 – A1 peduncle equal to, or shorter than, A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– A1 peduncle distinctly longer and stouter than A2 peduncle . . . . . . . . . . . . . . . . . . . . . 12
7 – Male Plp1 uniramous and bisegmented, distal segment bilobed by median longitudinal
furrow, and subchelate distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sergio
– Male Plp1 uniramous and bisegmented, distal segment simple, subchelate, or w-shaped
distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 – Male Plp2 endopod with appendices interna and masculina . . . . . . . . . . . . . . . . . . . . . . . 9
– Male Plp2 endopod with either appendix interna or appendix masculina . . . . . . . . . . 11
9 – Mxp3 dactylus rounded distally . . . . . . . . . . . . . . . . . . . . . . Thailandcallichirus gen. nov.
414
CRM 013 – Katsushi Sakai
– Mxp3 dactylus pointed distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 – Plp2 biramous, endopod bearing distally triangular appendix masculina with marginal
setae, and mesiodistally small appendix interna . . . . . . . . . . . . . . Glypturoides gen. nov.
– Male Plp2 biramous, endopod bearing embedded appendix masculina distally and
embedded appendix interna distolaterally . . . . . . . . . . . . . . . . . . . . . . . . . . . Neocallichirus
11 – Male Plp2 endopod with appendix interna distally, but without appendix masculina . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corallichirus sensu nov.
– Male Plp2 endopod with appendix masculina mesiodistally, but with or without
appendix interna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glypturus
12 – Male Plp2 endopod without appendices interna and masculina . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tirmizicallichirus gen. nov.
– Male Plp2 endopod with appendix masculina, but with or without appendix interna . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 – P1 unequal, smaller cheliped with spoon-shaped chela, dactylus shaped as elongateoval, dorsal and ventral margins spinous, fixed finger concave on cutting surface, with
spinous lateral and mesial margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podocallichirus
– P1 unequal, smaller cheliped usually compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 – Male Plp2 endopod distally bearing proximally-fused broad appendix masculina with
marginal setae, and mesiomedially small appendix interna with hooklets . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grynaminna
– Male Plp2 endopod bearing mesiodistally appendix masculina with marginal setae,
and with or without appendix interna with hooklets mesiomedially . . . . . . . . . . . . . . . 15
15 – Male Plp1 uniramous and bisegmented, distal segment simple . . . . . . Lepidophthalmus
– Male Plp1 uniramous and bisegmented, distal segment chelate distally . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidophthalmoides gen. nov.
Genus Balsscallichirus gen. nov.
Diagnosis. — Carapace with dorsal oval, rostrum short or reduced, and triangular. A1 peduncle longer than A2 peduncle. Mxp3 ischium-merus elongate
and pediform; carpus longer than wide and longer than merus; propodus broadened, shorter than carpus, and largely convex on ventral margin; and dactylus
digitiform; exopod absent. P1 chelate and unequal; in larger cheliped ischium
denticulate distally on ventral margin; merus with lower triangular lobe on
ventral margin. In smaller cheliped ischium and merus unarmed. P2 chelate.
P3 simple. P4 subchelate. P5 chelate. Male Plp1 uniramous and un- or bisegmented; male Plp2 uniramous and unsegmented, lacking appendices interna
and masculina. Telson wider than long, and slightly convex on posterior margin. Uropodal endopod oval in shape; uropodal exopod broadly convex on lateral margin, with marginal setae, bearing dorsal plate.
Remarks. — The present genus, Balsscallichirus gen. nov. is erected based
on Callianassa balssi, which was included in the genus Podocallichirus Sakai,
1999c, but is distinguished from Podocallichirus by the forms of male Plps1-2.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
415
Podocallichirus guineensis (De Man, 1928a) is included in the present genus,
because Plps1-2 are uniramous, as in B. balssi, although they are small simple
protrusions only in B. guineensis (cf. De Saint Laurent & Le Loeuff, 1979, fig.
15m-n), and the Plp1 is unsegmented.
Type species. — Callianassa (Callichirus) balssi Monod, 1933, by present
designation. The gender of the new generic name, Balsscallichirus, is masculine.
Species included. — Balsscallichirus balssi (Monod, 1933); B. guineensis
(De Man, 1928a).
Etymology. — The new generic name is dedicated to Dr. H. Balss, by adding
his name as a prefix to Callichirus.
K EY
TO THE
S PECIES
OF THE
G ENUS BALSSCALLICHIRUS
1 – P3 propodus distinctly protruded posteriorly at ventroproximal corner . . . . . . . B. balssi
– P3 propodus not protruded posteriorly at ventroproximal corner . . . . . . . . . B. guineensis
Balsscallichirus balssi (Monod, 1933)
[Abbreviated list of synonymy.]
Callianassa (Callichirus) balssi Monod, 1933: 13, fig. 2A-F.
Callianassa balssi — Longhurst, 1958: 45 (partim); Sakai, 1999c: 54; Sakai, 2005b: 189.
Callichirus balssi — Le Loeuff & Intès, 1974: 46, fig. 13a-o; De Saint Laurent & Le Loeuff,
1979: 58, figs. 11c-e, 12d, 13a-b, 14a, 15a-d, 19a.
Podocallichirus balssi — Sakai, 1999c: 54; Sakai, 2005b: 189.
Diagnosis. — Mxp3 ischium-merus pediform; merus convex at mesiodistal
corner; propodus subquadrate, and dactylus digitiform. Male Plp1 uniramous
and bisegmented; male Plp2 uniramous and unsegmented, lacking appendices
interna and masculina. Telson subquadrate, wider than long, and convex on
lateral margins, but slightly concave medially on posterior margin, without
median spine. [Cf. Le Loeuff & Intès, 1974: 46, fig. 13g, m; De Saint Laurent
& Le Loeuff, 1979: 58, figs. 12d, 15a-d, 19a.]
Type locality. — Bay of Repos, Nouadhibou, Mauritania.
Distribution. — Bay of Repos, Nouadhibou, Mauritania; Dakar, Senegal;
Gambia; Pointe-Noire, Congo; 6-10 m.
Balsscallichirus guineensis (De Man, 1928)
[Abbreviated list of synonymy.]
Callianassa (Callichirus) guineensis De Man, 1928a: 45, pl. 10 fig. 19-l9e, pl. 11 fig. 19f-19r.
Callichirus guineensis — Le Loeuff & Intès, 1974: 46, fig. 12a-o; De Saint Laurent & Le
Loeuff, 1979: 64, figs. 11h, 12e, 13g-h, 14d, 15m-p, 19d.
416
CRM 013 – Katsushi Sakai
Podocallichirus guineensis — Sakai, 1999c: 55; Sakai, 2005b: 190.
Diagnosis. — Mxp3 ischium-merus pediform; merus angulated at mesiodistal corner; propodus subovate, and dactylus digitiform. Male Plp1 uniramous
and unsegmented; male Plp2 uniramous and unsegmented. [Cf. De Man,
1928a: 45, pl. 10 fig. 19a, b; De Saint Laurent & Le Loeuff, 1979: 64,
figs. 15m, n, 19d).
Type locality. — Prampram, Ghana, 9-10 m.
Distribution. — Ghana; Nigeria; 9-15 m.
Genus Barnardcallichirus gen. nov.
Diagnosis. — Carapace with dorsal oval. Rostrum short and acutely triangular. A1 peduncle longer than A2 peduncle. Mxp3 ischium-merus pediform;
carpus longer than wide; propodus broadened, shorter than carpus, and largely
convex on ventral margin; and dactylus digitiform; exopod absent. P1 chelate
and unequal; in larger cheliped, merus with lower triangular lobe on ventral
margin; in smaller cheliped, ischium and merus unarmed. P2 chelate. P3 simple. P4 subchelate. P5 chelate. Male Plp1 uniramous and bisegmented, distal segment longitudinally furrowed, mesial part rounded distally and lateral
part sharply protruded distally over mesial part; male Plp2 uniramous and unsegmented, lacking appendices interna and masculina; Plps3-5 with appendix
interna. Telson wider than long, almost straight on posterior margin, without
median spine. Uropodal endopod oval; uropodal exopod broadly convex on
distal margin, bearing dorsal plate.
Remarks. — The present genus, Barnardcallichirus gen. nov. is established
on the basis of Callichirus tenuimanus De Saint Laurent & Le Loeuff,
1979, which was included in the genus Podocallichirus Sakai, 1999c, and
is distinguished from Podocallichirus by the form of the male Plp2. In
Barnardcallichirus, the male Plp2 is uniramous and unsegmented, lacking
appendices interna and masculina, whereas in Podocallichirus, the male Plp2 is
biramous, and the endopod bears mesiodistally a fusiform appendix masculina
provided mesiomedially with a proximally-fused appendix interna with a patch
of hooklets.
Type species. — Callichirus tenuimanus De Saint Laurent & Le Loeuff,
1979, by present designation. The gender of the new generic name, Barnardcallichirus, is masculine.
Species included. — Barnardcallichirus gilchristi (Barnard, 1946);
B. tenuimanus (De Saint Laurent & Le Loeuff, 1979).
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
417
Etymology. — The new generic name is dedicated to Dr. K.H. Barnard by
adding his name as a prefix to the existing name Callichirus; Dr. Barnard devoted his life to extensive carcinological studies in South Africa by describing
many species, including Callianassa gilchristi.
K EY
TO THE
S PECIES
OF THE
G ENUS BARNARDCALLICHIRUS
1 – Mxp3 propodus subovate, distinctly expanded ventrally . . . . . . . . . . . . . . . . . . B. gilchristi
– Mxp3 propodus subovate, slightly rounded ventrally . . . . . . . . . . . . . . . . . . B. tenuimanus
Barnardcallichirus gilchristi (Barnard, 1946)
Callianassa gilchristi Barnard, 1946: 379; Barnard, 1950: 509, fig. 95a-e; Kensley, 1974: 277;
Tudge et al., 2000: 143.
Podocallichirus gilchristi — Sakai, 1999c: 56, fig. 9a-d; Sakai, 2005b: 191.
Diagnosis. — Mxp3 ischium-merus pediform; merus angulated at distomesial corner; propodus subquadrate, and dactylus digitiform. Male Plp1 uniramous and bisegmented, distal segment longitudinally furrowed, mesial part
rounded distally and lateral part sharply protruded distally over mesial part;
male Plp2 undescribed; Plps3-5 with appendix interna. Telson subquadrate,
wider than long, and almost straight on posterior margin, without median
spine. [Cf. Barnard, 1950, fig. 95; Sakai, 1999c, fig. 9b.]
Type locality. — False Bay and Durban Bay, South Africa.
Distribution. — South Africa — Saldanha Bay, False Bay to Port Elizabeth;
Durban Bay and off Zululand coast; 37 m.
Barnardcallichirus tenuimanus (De Saint Laurent & Le Loeuff, 1979)
[Abbreviated list of synonymy.]
Callianassa balssi — Longhurst, 1958: 31 (partim). [Not: Callianassa balssi Monod, 1933.]
Callichirus tenuimanus De Saint Laurent & Le Loeuff, 1979: 61, figs. 11a-b, 12a-b, 13e-f, 14c,
15e-h, 19c.
Podocallichirus tenuimanus — Sakai, 1999c: 55; Sakai, 2005b: 190.
Diagnosis. — Mxp3 ischium-merus pediform; merus angulated at mesiodistal corner; propodus subovate, and dactylus digitiform. Male Plp1 uniramous
and bisegmented, distal segment longitudinally furrowed, mesial part rounded
distally and lateral part sharply protruded distally over mesial part; male Plp2
uniramous and unsegmented. Telson subquadrate, wider than long, and slightly
convex on posterior margin, without median spine. [Cf. De Saint Laurent & Le
Loeuff, 1979: 61, figs. 12b, 15e-f, 19c.]
Type locality. — Sierra Leone, 7 m.
Distribution. — Only known from the type locality.
418
CRM 013 – Katsushi Sakai
Genus Callichirus Stimpson, 1866
[Abbreviated list of synonymy.]
Callichirus Stimpson, 1866: 61, figs. 11a-b, 12a-b, 13e-f, 14c, 15e-h, 19c; Manning & Felder,
1991: 775; Sakai, 1999c: 59; Sakai, 2005b: 126.
Diagnosis. — Carapace with dorsal oval. Rostrum reduced to a small triangle; rostral carina, hepatic sulcus, and hepatic prominence undeveloped. A1
peduncle longer and more robust than A2 peduncle; distal segment elongate.
Mxp3 ischium-merus broadened and operculiform; propodus broadened and
rounded on ventral margin; and dactylus digitiform; exopod usually absent.
P1 chelate, equal or unequal; in male larger cheliped ischium very elongate,
merus normal in length or elongate, bearing meral hook or not, and carpus
much longer than chela. P2 chelate. P3 propodus broadened. P4 subchelate.
P5 subchelate or chelate. Abdominal somites 3-5 ornamented dorsally. Male
Plp1 uniramous, blade-like, and un-, bi- or trisegmented, distal segment simple
distally; male Plp2 biramous and blade-like, endopod without appendices interna and masculina, or with appendix interna but without appendix masculina;
Plps3-5 with appendix interna. Telson wider than long, and almost straight on
posterior margin. Uropodal endopod strap-shaped; uropodal exopod convex on
distal margin, bearing dorsal plate. [Adapted from Sakai, 1999c: 59.]
Remarks. — Callichirus major Say, 1818, the type species of Callichirus,
belonging to the family Callichiridae, is similar to Callianassa subterranea,
the type species of Callianassa, in that the uropodal exopod bears a dorsal
plate, but Callichirus is different from Callianassa in the proportion in length
of A1 peduncle to A2 peduncle, in the forms of Mxp3 propodus and dactylus,
in telson and uropodal endopod, and also in the presence or absence of a
transverse row of setae on the dorsal surface of abdominal somites 3-5.
Type species. — Callianassa major Say, 1818, by original designation. The
gender of the generic name, Callichirus, is masculine.
Species included. — Callichirus adamas (Kensley, 1974); C. islagrande
(Schmitt, 1935b); C. kraussi (Stebbing, 1900); C. major (Say, 1818);
C. seilacheri (Bott, 1955).
K EY
TO THE
S PECIES
OF THE
G ENUS CALLICHIRUS
1 – Eyestalks obtuse or rounded distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Eyestalks narrowly protruded distally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
2 – Telson subsquare, and largely concave on posterior margin . . . . . . . . . . . . . . . . C. kraussi
– Telson wider than long, bearing concavity between proximal and median lobes on lateral
margin, converging toward rounded posterior margin, with median notch without spine
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. major
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
419
3 – Telson subcircular, with broad median concavity, without spine on posterior margin . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. seilacheri
– Telson trapezoid or subsquare, broader than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – Telson trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. islagrande
– Telson subsquare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. adamas
Callichirus adamas (Kensley, 1974)
Callianassa adamas Kensley, 1974: 266, 277, figs. 1, 2; De Saint Laurent & Le Loeuff, 1979:
67, figs. 14f, 16a, 17a, 19f, 20e-g, 23f-i.
Callichirus adamas — Manning & Felder, 1986: 439; Sakai, 1999c: 63; Tudge et al., 2000:
144; Sakai, 2005b: 127.
Diagnosis. — Rostrum inconspicuous. Eyestalks narrowly protruded distally, reaching middle of A1 penultimate segment. Mxp3 ischium-merus oval;
propodus subtriangular, and dactylus digitiform. Ischium of male larger cheliped elongate, bearing median protrusion on mesial margin. Male Plp1 uniramous and bisegmented; male Plp2 biramous, exopod small, endopod without
appendices interna or masculina; Plp3 with pad of rounded hooks. Telson subquadrate, slightly broader than long, and lobulated at posterolateral corners;
posterior margin short. [Adapted from Kensley, 1974; and from De Saint Laurent & Le Loeuff, 1979.]
Type locality. — Orange River mouth, South Africa.
Distribution. — Orange River mouth and Lambert’s Bay, South Africa;
Senegal; Cape Verde Islands.
Callichirus islagrande (Schmitt, 1935)
[Abbreviated list of synonymy.]
Callianassa (Callichirus) islagrande Schmitt, 1935b: 5, pl. 1 fig. 3, pl. 2 fig. 1, pl. 3 fig. 2, pl. 4
fig. 5.
Callianassa (Calichirus) islagtande [sic!] — Solar et al., 1970: 20.
Callianassa islagrande — Biffar, 1971a: 654; Manning & Felder, 1986: 438, fig. 2.
Callichirus islagrande — De Saint Laurent & Le Loeuff, 1979: 79; Manning & Felder, 1991:
775; Sakai, 1999c: 60, fig. 11b-f; Sakai, 2005b: 128.
Material examined. — USNM 70792, paratype, male (TL/CL, 67.0/10.5 mm), Grand Isle,
Louisiana, 1930, leg. N.N. Anderson.
Diagnosis. — Mxp3 ischium-merus subsquare; merus obliquely straight on
distal margin. Male Plp1 uniramous and trisegmented, and rounded distally;
male Plp2 biramous, endopod bearing neither appendix interna nor appendix
masculina, and exopod shorter than endopod (Sakai, 1999c: 60, fig. 11e-f).
Telson trapezoid, wider than long and trapezoid, with convex posterolateral
420
CRM 013 – Katsushi Sakai
corners, and slightly concave medially on posterior margin, without median
spine.
Type locality. — Grand Isle, Louisiana, U.S.A.
Distribution. — Northern and western Gulf of Mexico (Manning & Felder,
1986); common in the shallow subtidal of sandy beaches.
Callichirus kraussi (Stebbing, 1900)
(fig. 64A-C)
[Abbreviated list of synonymy.]
Callianassa kraussi Stebbing, 1900: 39, pls. 2, 3; Kensley, 1974: 277 (key); Kensley, 1975: 57;
Ngoc-Ho, 2002b: 224.
Callianassa (Callichirus) kraussi — De Man, 1928b: 28, 94, 95, 113, 179, 182, 183; Barnard,
1950: 506, fig. 94; Holthuis, 1991: 248, figs. 451, 452, 264.
Callichirus kraussi — Sakai, 1999c: 64, fig. 13a-e; Sakai, 2005b: 130.
Callianassa (Callianassa) kraussi — Felder, 2001: 441.
Material examined. — USNM 105736, 1 male (TL/CL, 83.0/19.0 mm), Langebaan, South
Africa, 5.vi.1957, det. L.B. Holthuis.
Diagnosis. — Mxp3 ischium-merus operculiform; merus straight on distal
margin; propodus subquadrate, and dactylus digitiform. Abdominal somites 35 distinctively ornamented. Male Plp1 (fig. 64A) uniramous and trisegmented;
male Plp2 (fig. 64B, C) biramous, endopod distinct, bearing mesiodistally
embedded appendix interna with patch of hooklets. Telson subquadrate, wider
than long, and protruded at posterolateral angles to form median concavity
with tubercle on posterior margin.
Remarks. — In the male specimen examined (USNM 105736) the Plp2
exopod is rather longer than the endopod, though in the male specimen
(RMNH D 12002) the Plp2 exopod is much smaller and shorter than the
endopod (Sakai, 1999c: 64, fig. 13d).
Type locality. — Cape of Good Hope, Gordon’s Bay.
Distribution. — South Africa — Saldanha Bay; False Bay to Zululand; a
little below high water mark, sandy littoral zone in bays and estuaries.
Callichirus major (Say, 1818)
(fig. 64D-E)
Callianassa major Say, 1818: 238; White, 1847: 70; Gibbes, 1850: 194; A. Milne-Edwards,
1870: 86, 101; Stimpson, 1871: 122; Yong, 1900: 425; Schmitt, 1935b: 3; Lutze, 1937: 1-15,
figs. 1-3; Pearse et al., 1942: 153, 155, 156, 185, figs. 10, 14; Willis, 1942: 2; Gurney, 1944:
83; Pohl, 1946: 71-80, figs. 7-28; Hoyt & Weimer, 1963: 10; Frankenberg et al., 1967: 113120; Holthuis, 1969: 12; Biffar, 1971a: 651-653; Coelho & Ramos, 1973: 161; Rabalais et
al., 1981: 105; Williams, 1984: 183, fig. 127.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
421
Fig. 64. Callichirus kraussi (Stebbing, 1900), Callichirus major (Say, 1818), and Callichirus
seilacheri (Bott, 1955). A, D, F, male Plp1; B, C, E, G, H, male Plp2. A, B, C, Callichirus
kraussi, USNM 105736, male (TL/CL, 83.0/19.0 mm), Langebaan, South Africa; D, E,
Callichirus major, USNM, male (TL/CL, 100.0/19.0 mm); F, G, H, Callichirus seilacheri,
USNM 142540, male. Scales, 1 mm.
Callichirus major — Stimpson, 1866: 47; Stimpson, 1871: 122; Kingsley, 1878: 327; Fowler,
1911: 571; Hay & Shore, 1917: 407, pl. 29 fig. 10; De Saint Laurent, 1973: 514; Rodrigues,
1983: 25, figs. 23-52; Manning & Felder, 1986: 439, fig. 1; Manning, 1987: 397; Rodrigues
& Hödl, 1990: 50, fig. 1; Manning & Felder, 1991: 775, figs. 1, 3-6; Dworschak, 1992: 208;
Blanco-Rambla, 1997: 7-12, figs. 1-3; Coelho, 1997: 147; Rodrigues et al., 1998: 380; Melo,
1999: 354, figs. 235-236; Sakai, 1999c: 61, fig. 11a; Strasser et al., 1999a: 211; Strasser et
al., 1999b: 844; Tudge et al., 2000: 144; Coelho et al., 2007: 5; Robles et al., 2009: 317.
Callianassa (Callichirus) major — Borradaile, 1903: 547; De Man, 1928a: 30, pl. 7 fig. 1414b, pl. 8 fig. 14c, d; De Man, 1928b: 29, 91, 94, 111 (key); Williams, 1965: 100, fig. 78;
Rodrigues, 1971: 191, figs. 1-20.
422
CRM 013 – Katsushi Sakai
Material examined. — USNM, male (TL/CL, 100.0/19.0 mm).
Diagnosis. — Mxp3 ischium-merus rhombic; merus obliquely concave
on distal margin; propodus subquadrate, and dactylus digitiform. Male Plp1
uniramous and trisegmented (fig. 64D), distal segment obtuse distally; male
Plp2 (fig. 64E) biramous, endopod without appendices interna and masculina.
Telson subovate, wider than long, and convergent toward rounded posterior
margin with median notch without spine, bearing concavity between proximal
and median lobes on lateral margin.
Type locality. — Coast of southern [United] States and east Florida, St.
Johns River.
Distribution. — North Carolina — Cape Lookout, Barden Inlet, Ferry
Landing; Gulf of Mexico; Venezuela — Guaracayal, Tunantal, and Carenero;
Brazil — Rio Grande do Norte; Pernambuco, São Paulo, Santos; intertidal.
Callichirus seilacheri (Bott, 1955)
(fig. 64F-H)
Callianassa seilacheri Bott, 1955: 47, fig. 7a-g.
Callichirus seilacheri — Manning & Felder, 1986: 439, fig. 3; Manning & Felder, 1991: 775;
Hendrickx, 1995: 390; Sakai, 1999c: 62, fig. 12a-f; Tudge et al., 2000: 144; Sakai, 2005b:
129; Hernáez et al., 2008: 351.
Callianassa garthi Retamal, 1975: 178, figs. 1-8. [Type locality. — Playa Negra, Chile, 36◦ 45′ S
73◦ 10′ W.]
Callichirus garthi — Tudge et al., 2000: 144.
Material examined. — USNM 142540, male.
Diagnosis. — Mxp3 ischium-merus rhombic; merus obliquely concave on
distal margin; propodus subtriangular, and dactylus digitiform. Male Plp1 uniramous and trisegmented (fig. 64F); male Plp2 (fig. 64G, H) biramous, endopod bearing distally embedded appendix interna, but no appendix masculina.
Abdominal somites 3-5 with transverse row of setae. Telson subovate, wider
than long, convex medially on lateral margin, and bearing broad median concavity without spine on rounded posterior margin.
Type locality. — El Salvador.
Distribution. — El Salvador; Chile, Tubul, south of Talcahuano to Playa
Negra near Viña del Mar.
Genus Corallichirus Manning, 1992 (sensu nov.)
Corallichirus Manning, 1992: 571, figs. 1b, 2.
Diagnosis. — Rostrum acutely triangular. Carapace with dorsal oval, and
proximally non-calcified anterolateral projections. A1 peduncle shorter than
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
423
A2 peduncle. Mxp3 ischium-merus subpediform; propodus broadened and
ovate on ventral margin; dactylus digitiform; exopod absent. P1 chelate,
unequal, and dissimilar; male larger cheliped with or without meral hook.
P2 chelate. P3 propodus broadened. P4 subchelate. P5 chelate. Abdominal
somites 3-5 not ornamented dorsally. Male Plp1 uniramous and bisegmented,
distal segment chelate distally; male Plp2 biramous, endopod bearing small
appendix interna distally, but no appendix masculina; male Plps3-5 biramous
and foliaceous, endopods with stubby, embedded appendix interna. Telson
wider than long, and varied in shape. Uropodal endopod wider than long
and rhombic, overreaching telson. Uropodal exopod convex on lateral margin,
bearing dorsal plate.
Remarks. — Manning (1987: 392) established a new genus, Corallianassa
including two species, C. longiventris (A. Milne-Edwards, 1870) and C. borradailei (De Man, 1928), and Manning & Felder (1991: 776) showed that
the genus Corallianassa includes C. longiventris (A. Milne-Edwards, 1870),
C. hartmeyeri (Schmitt, 1935b), and C. xutha Manning, 1988. However, later,
in 1992, Manning erected a new genus Corallichirus based on Corallianassa
xutha as the type species, excluded from the genus Corallianassa, because the
relative length of abdominal somites 2 to 6 is different between the two genera.
In the type species of Corallichirus, C. xuthus, abdominal somite 2 is subequal
in length to abdominal somite 6, whereas in the type species of Corallianassa,
C. longiventris, abdominal somite 2 is much longer than abdominal somite 6.
However, such a difference is not to be regarded as a vital feature that determines their generic status, because it is a difference at the species level rather
than on the generic level. Sakai (2005b: 134) reclassified Corallianassa hartmeyeri, Corallianassa longiventris, and Corallichirus xuthus under the genus
Glypturus (cf. Sakai, 2005b: 134). However, in the present revision G. hartmeyeri is synonymized with G. longiventris, and G. xuthus is reclassified under Corallichirus sensu nov., because G. xuthus is different from the type
species of Glypturus, G. acanthochirus Stimpson, 1866, in the features of male
Plps1-2.
Type species. — Corallianassa xutha Manning, 1988, by original designation. The gender of the generic name, Corallichirus, is masculine.
Species included. — Corallichirus bayeri Kensley, 2001b; C. intesi (De
Saint Laurent & Le Loeuff, 1979); C. xuthus (Manning, 1988).
K EY
TO THE
S PECIES
OF THE
G ENUS CORALLICHIRUS
1 – Uropodal endopod narrow and fusiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bayeri
424
CRM 013 – Katsushi Sakai
– Uropodal endopod broad and fusiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Carpus of larger cheliped serrated on ventral margin; eyestalks narrowly protruded
anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. intesi
– Carpus of larger cheliped slightly cristate ventrally; eyestalks rounded anteriorly . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. xuthus
Corallichirus bayeri Kensley, 2001
Corallichirus bayeri Kensley, 2001b: 328, figs. 1, 2.
Remarks. — The male and female are similar in the structures of their
pereiopods, pleopods, and uropods (Kensley, 2001: 231); however it is necessary to examine the structures of their male Plps1-2, and to report and interpret
those observations.
Type locality. — Agat Bay, north of Alutom Island, Guam.
Distribution. — Known only from the type locality.
Corallichirus intesi (De Saint Laurent & Le Loeuff, 1979)
(fig. 65C-D)
Callichirus intesi De Saint Laurent & Le Loeuff, 1979: 69, figs. 14g, 16c, 17b, 18b, 19b, 21a-c,
23 j-m; Tudge et al., 2000: 144.
Glypturus intesi — Manning, 1987: 392; Sakai, 1999c: 73; Sakai, 2005b: 133.
Corallianassa intesi — Abed-Navandi, 2000: 294-295.
Material examined. — MNHN Th 371, Goree Island, Republic of Senegal, 5 m. MNHN Th
371, type specimens of Callichirus intesi De Saint Laurent & Le Loeuff, 1979, 4 males (TL,
68-97 mm), 5 females (TL, 71-98 mm), Goree Island, Republic of Senegal, 5 m, in front of Port
of Dakar.
Diagnosis. — Mxp3 ischium-merus operculiform; merus curved on distomesial margin; propodus subquadrate, and dactylus digitiform. Male Plp1
uniramous and bisegmented, and distal segment chelate distally (fig. 65C);
male Plp2 endopod bearing small appendix interna distally, but no appendix
masculina (fig. 65D; De Saint Laurent & Le Loeuff, 1979, fig. 23J, k). Telson
slightly concave on posterior margin, with low median convexity.
Type locality. — Senegal, Port Dakar.
Distribution. — Goree and Dakar, Senegal; Ilha da Boavista; Ilha do Maio
and Ilha de Santiago, CapeVerde Is.
Corallichirus xuthus (Manning, 1988)
(fig. 65G-H)
Callianassa hartmeyeri — Hult, 1938: 7, figs. 1-4, pl. 1; Schmitt, 1939: 15. [Not: Callianassa
hartmeyeri Schmitt, 1935b.]
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
425
Fig. 65. Glypturus acanthochirus Stimpson, 1866, Corallichirus intesi (De Saint Laurent &
Le Loeuff, 1979), Glypturus longiventris (A. Milne-Edwards, 1870), Corallichirus xuthus
(Manning, 1988), and Glypturoides trilobata (Biffar, 1970). A, C, E, G, I, male Plp1; B, D, F, H,
J, male Plp2. A, B, Glypturus acanthochirus, USNM 266220, 1 male (TL/CL, 47.0/13.0 mm),
Parguera Bay, Lajas, Puerto Rico, Caribbean Sea; C, D, Corallichirus intesi, MNHN Th 371,
Goree Island, Republic of Senegal, 5 m; E, F, Glypturus longiventris, USNM 122449, 1 male,
S.W. reef of Somerset, 25 fms (45.7 m); G, H, Corallichirus xuthus, USNM 99861, holotype
male (TL/CL/ 43.0/10.0 mm), Clipperton Is. (10◦ 18′ N 106◦ 33′ W); I, J, Glypturoides trilobata,
USNM 205642, 1 male (TL/CL, 48.0/10.0 mm), in front of FSU Marine Laboratory, Florida.
Scales 1 mm.
426
CRM 013 – Katsushi Sakai
Callianassa (Callichirus) placida — Chace, 1962: 617 (partim). [Not: Callianassa placida De
Man, 1905.]
Callianassa placida — Hernández Aguilera et al., 1986: 206. [Not: Callianassa placida De
Man, 1905.]
Corallianassa xutha Manning, 1988: 885, fig. 3; Manning & Felder, 1991: 777, figs. 1, 2, 5;
Lemaitre & León, 1992: 44; Lemaitre & Ramos, 1992: 347; Hendrickx, 1995: 390.
Corallichirus xuthus — Manning, 1992: 571, figs. 1, 2; Hernández-Aguilera, 1998: 304; Tudge
et al., 2000: 144; Hendrickx et al., 2005: 170.
Glypturus xuthus — Sakai, 1999c: 75; Sakai, 2005b: 135.
Material examined. — USNM 99861, holotype, male (TL/CL/43.0/10.0 mm), Clipperton Is.
(10◦ 18′ N 106◦ 33′ W).
Diagnosis. — Mxp3 ischium-merus subsquare; merus subtriangular; propodus subsquare; and dactylus digitiform. Male Plp1 uniramous and bisegmented, and chelate distally (fig. 65G); male Plp2 biramous, endopod bearing
appendix interna distally, but without appendix masculina (fig. 65H). Telson
wider than long and trapezoid, and almost straight on posterior margin, with
low median convexity. [Cf. Manning, 1988, fig. 3J.]
Remarks. — The present species, Corallichirus xuthus (Manning, 1988) is
similar to the type species of Glypturus, G. acanthochirus in the shape of
its male Plp1 and other characteristics. However, the male Plp2 is different
in structure between the two species: in Corallichirus xuthus the male Plp2
endopod bears a small appendix interna distally, but not an appendix masculina
(fig. 65H), whereas in Glypturus acanthochirus it bears distally an appendix
masculina with a small appendix interna mesiomedially (fig. 65B).
Type locality. — Clipperton Is. (10◦ 18′ N 106◦ 33′ W), eastern Pacific Ocean,
southwest of Mexico.
Distribution. — Eastern Pacific: María Madre Island, Mexico; Bahía Azufre,
Isla Clarión, Mexico, Tres Marias Islands, Baja California; 7-18 m; Clipperton
Island; Port Utria, Colombia; Socorro island, Gorgona Island, and Indefatigable Island, Galapagos Islands; Ecuador, lagoon.
Genus Forestcallichirus gen. nov.
Diagnosis. — Carapace with dorsal oval. Rostrum small and obtusely triangular. A1 peduncle longer than A2 peduncle. Mxp3 ischium-merus pediform,
keeping same width from proximal to distal; propodus broadened, convex on
ventral margin; dactylus digitiform; exopod absent. P1 chelate; in larger cheliped, ischium and merus minutely denticulate on ventral margin; chela smooth
on dorsal and ventral margins; in smaller cheliped, ischium, merus, and chela
unarmed on dorsal and ventral margins. P2 chelate. P3 simple. P4 subchelate.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
427
P5 chelate. Male Plp1 uniramous and bisegmented, distal segment longer than
proximal one and w-shaped distally; male Plp2 uniramous and bisegmented,
lacking appendices interna and masculina; Plps3-5 with appendix interna. Telson wider than long, slightly convergent in posterior two-thirds, and straight on
posterior margin. Uropodal endopod oval in shape; uropodal exopod broadly
convex on distal margin, with setae, bearing dorsal plate.
Remarks. — The present new genus Forestcallichirus gen. nov. is separated
from the genus Podocallichirus Sakai, 1999c, because the male Plps1-2 are
different in structure between the two genera. In Forestcallichirus the male
Plp1 is uniramous and bisegmented, the distal segment is w-shaped distally,
and the male Plp2 is uniramous; whereas in Podocallichirus the male Plp1
is uniramous and bisegmented, but the distal segment is chelate, and Plp2 is
biramous, and the endopod bears mesiodistally a fusiform appendix masculina
with marginal setae and a small, proximally-fused appendix interna with
hooklets mesiomedially.
Type species. — Callichirus foresti Le Loeuff & Intès, 1974, by present
designation and monotypy. The gender of the new generic name, Forestcallichirus, is masculine.
Species included. — Forestcallichirus foresti (Le Loeuff & Intès, 1974).
Etymology. — The new generic name Forestcallichirus is dedicated to Prof.
J. Forest, who has played a leading role in the carcinological world as one of
the great carcinologists of the National Museum of Natural History, Paris, by
adding his name as a prefix to the generic name Callichirus.
Forestcallichirus foresti (Le Loeuff & Intès, 1974)
[Abbreviated list of synonymy.]
Callianassa guineensis — Longhurst, 1958: 31 (partim). [Not: Callianassa (Callichirus)
guineensis De Man, 1928a.]
Callichirus foresti Le Loeuff & Intès, 1974: 46, fig. 14a-x; De Saint Laurent & Le Loeuff, 1979:
58, figs. 11f-g, 12c, 13c-d, 14b, 15i-l, 19b.
Podocallichirus foresti — Sakai, 1999c: 55; Sakai, 2000: 189; Sakai, 2005b: 189.
Material examined. — Males, Côte d’Ivoire, leg. R/V “Reine Pokou”, det. Le Loeuff &
Intès, 1974.
Diagnosis. — Mxp3 ischium-merus pediform; merus angulated on distomesial corner; propodus subovate, and dactylus digitiform. Male Plp1 uniramous
and bisegmented, distal segment w-shaped distally; male Plp2 uniramous and
bisegmented, lacking appendices interna and masculina; Plps3-5 with appendix interna. Telson subquadrate, wider than long, slightly convergent in posterior two-thirds, and straight on posterior margin. [Cf. Le Loeuff & Intès, 1974:
428
CRM 013 – Katsushi Sakai
46, fig. 14m, s, t, u; De Saint Laurent & Le Loeuff, 1979: 58, figs. 12c, 15i-l,
19b.]
Type locality. — Grand Lahou (5◦ 11′ N 3◦ 49.05′ W), Ivory Coast, 25 m.
Distribution. — Senegal; Guinea; Sierra Leone; Ivory Coast; Pointe-Noire,
Congo; 5-30 m.
Genus Glypturoides gen. nov.
Diagnosis. — Carapace with dorsal oval, but without rostral carina, cardiac
prominence, and transverse cardiac sulci. A1 peduncle shorter than A2 peduncle. Mxp3 ischium-merus ovate; propodus broadened and truncate on ventral
margin; dactylus digitiform; exopod absent. P1 chelate, unequal, and dissimilar; in male larger cheliped meral hook triangular. P2 chelate. P3 propodus
broadened. P4 subchelate. P5 chelate. Abdominal somites 3-5 not ornamented
dorsally. Male Plp1 uniramous and bisegmented, distal segment subchelate
with acute tip mesiodistally and rounded prominence distolaterally (fig. 65I;
Biffar, 1970, fig. 1m); male Plp2 biramous, endopod bearing distally triangular appendix masculina with marginal setae, and mesiodistally small appendix
interna (fig. 65J); male Plps3-5 endopods foliaceous, bearing appendix interna.
Telson wider than long. Uropodal endopod longer than wide, and rhombic in
shape, overreaching telson; uropodal exopod convex on lateral margin, bearing
distal flap. [Adapted from Sakai, 1999c: 72.]
Remarks. — Callianassa trilobata Biffar, 1970, was classified by Manning & Lemaitre (1994) as Sergio trilobatus, and later reclassified by Sakai
(1999, 2005b) under Neocallichirus Sakai, 1988 as N. trilobatus. However,
this species does not belong in Neocallichirus either, because it is also different from the type species of the genus Neocallichirus, Neocallichirus horneri
Sakai, 1988, in the form of the male Plp2. In N. trilobatus the Plp2 endopod
bears an appendix masculina distally and a small appendix interna mesiodistally (Biffar, 1973, fig. 1m, o); whereas in N. horneri the male Plp2 endopod
bears an embedded appendix masculina distally and an embedded appendix
interna distolaterally (Sakai, 1988, fig. 8I). Thus, a new genus, Glypturoides
gen. nov. is established for N. trilobatus, which is reclassified under the new
genus as Glypturoides trilobata.
Type species. — Callianassa trilobata Biffar, 1970, by present designation
and monotypy. The gender of the new generic name, Glypturoides, is feminine.
Species included. — Glypturoides trilobata (Biffar, 1970).
Etymology. — The new generic name, Glypturoides is composed of a
generic name, Glypturus and a Latin suffix, -oides, meaning “like” or “similar
to”.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
429
Glypturoides trilobata (Biffar, 1970)
(fig. 65I-J)
Callianassa trilobata Biffar, 1970: 36, fig. 1; Biffar, 1971a: 653, 654, 704, figs. 21, 22;
Manning, 1987: 397.
Neocallichirus trilobata — Manning & Felder, 1991: 779.
Sergio trilobatus — Manning & Lemaitre, 1994: 41; Tudge et al., 2000: 145; Robles et al.,
2009: 317.
Neocallichirus trilobatus — Sakai, 1999c: 93, fig. 20e-f; Sakai, 2005b: 169.
Material examined. — USNM 205642, 1 male (TL/CL, 48.0/10.0 mm), in front of FSU
Marine Laboratory, Florida, 06.iv.1985, leg. D.L. Felder & R.B. Manning.
Type locality. — Off Pinellas Point, Tampa Bay, Florida, 2-3 m.
Distribution. — Tampa Bay, Miami, Lemon Bay, Florida, U.S.A.
Genus Glypturus Stimpson, 1866
Glypturus Stimpson, 1866: 46; Manning, 1987: 390, 398; Manning & Felder, 1991: 778; Sakai,
1999c: 72; Sakai, 2005b: 130.
Corallianassa Manning, 1987: 392, 397; Manning & Felder, 1991 (partim): 776, figs. 1, 2, 5.
[Type species. — Callianassa longiventris A. Milne-Edwards, 1870.]
Corallianassa s. str., Felder, 2001: 440
Glypturus s. str., Felder, 2001: 440.
Diagnosis. — Rostrum acutely or obtusely triangular. Anterolateral projections on carapace calcified or non-calcified proximally. Carapace with dorsal
oval, but without rostral carina, cardiac prominence, and transverse cardiac sulcus(i). A1 peduncle shorter than A2 peduncle. Mxp3 ischium-merus subpediform; propodus broadened and rounded on ventral margin; dactylus digitiform;
exopod absent. P1 chelate, unequal, and dissimilar; male larger cheliped with
or without meral hook. P2 chelate. P3 propodus broadened. P4 subchelate.
P5 chelate. Abdominal somites 3-5 not ornamented dorsally. Male Plp1 uniramous and bisegmented, distal segment subchelate or w-shaped distally; male
Plp2 biramous, endopod bearing mesiodistally a proximally-fused appendix
masculina with slender, proximally-fused appendix interna mesiomedially, or
lacking appendices interna and masculina; male Plps3-5 biramous and foliaceous, endopods with stubby, embedded appendix interna. Telson wider than
long, and varied in shape. Uropodal endopod wider than long, and rhombic or
subquadrate, overreaching telson; uropodal exopod convex on lateral margin,
bearing dorsal plate. [Adapted from Sakai, 1999c: 72.]
Remarks. — Manning (1987) established a new genus, Corallianassa for
Callianassa longiventris A. Milne-Edwards, 1870, including Callianassa longiventris var. borradailei De Man, 1928, and later in 1988 added Corallianassa
430
CRM 013 – Katsushi Sakai
hartmeyeri (Schmitt, 1935b) and Corallianassa xutha Manning, 1988. In 1992,
however, he (Manning, 1992) erected a new genus, Corallichirus for Corallianassa xutha, because Corallianassa xutha is different from Corallianassa
longiventris. In Corallianassa longiventris abdominal somite 2 is longer than
abdominal somite 6, while in Corallianassa xutha abdominal somite 2 is about
as long as somite 6. The three species included in Corallianassa were later reclassified under Glypturus by Sakai (2005b: 134-135). However, in the present
revision, Glypturus hartmeyeri (Schmidt, 1935b) is synonymized with G. longiventris, because in the type species of Corallianassa, C. longiventris (A.
Milne-Edwards, 1870), the male Plps1-2 are the same in features as those of
the type species of Glypturus, G. acanthochirus Stimpson, 1866, and G. xuthus
is reclassified under Corallichirus sensu nov.
Type species. — Glypturus acanthochirus Stimpson, 1866: 46, by original
designation. The gender of the generic name, Glypturus, is masculine.
Species included. — G. acanthochirus Stimpson, 1866; G. armatus (A.
Milne-Edwards, 1870) (= G. motupore Poore & Suchanek, 1988); G. articulatus (Rathbun, 1906); G. assimilis (De Man, 1928b); G. borradailei (De Man,
1928a); G. collaroy (Poore & Griffin, 1979); G. haswelli (Poore & Griffin,
1979); G. lanceolatus (Edmondson, 1944) [= G. nakasonei (Sakai, 1967)];
G. longiventris (A. Milne-Edwards, 1870) [= Callianassa placida De Man,
1905; G. hartmeyeri (Schmitt, 1935b); Callianassa (Callichirus) oahuensis
Edmondson, 1944]; G. martensi (Miers, 1884a); G. rabalaisae sp. nov.
K EY
TO THE
S PECIES
OF THE
G ENUS GLYPTURUS
1 – Telson wider than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Telson trapezoid or pentagonal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 – Eyestalks rounded distally; cornea distinct, occupying distal half of eyestalk . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. articulatus
– Eyestalks with tubercle mesiodistally; cornea located distolaterally . . . . . . . G. collaroy
3 – Uropodal exopod strongly curved mesially in its posterior half . . . . . . . . . . . . . . . . . . . . 4
– Uropodal exopod bilobate and broadened in its posterior half . . . . . . . . . . . . . . . . . . . . . 7
4 – A1 peduncle reaching to distal end of A2 penultimate segment . . . . . . . . . . G. haswelli
– A1 peduncle reaching to middle of A2 distal segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Telson rounded on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. assimilis
– Telson angulate at both posterolateral angles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – Eyestalks not reaching distal end of A1 proximal segment . . . . . . . . . . . . . . G. martensi
– Eyestalks reaching distal end of A1 proximal segment . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. lanceolatus (= G. nakasonei)
7 – Telson pentagonal, widest at middle part . . . . . . . . . . . . . . . G. armatus (= G. motupore)
– Telson trapezoid, widest at proximal part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
431
8 – Telson with median spine on posterior margin . . . . . . . . . . . . . . . . G. rabalaisae sp. nov.
– Telson without median spine on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 – Abdominal somite 2 elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. borradailei
– Abdominal somite 2 not elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 – Cornea distinct and located apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. longiventris
– Cornea distinct and located distolaterally . . . . . . . . . . . . . . . . . . . . . . . . . G. acanthochirus
Glypturus acanthochirus Stimpson, 1866
(fig. 65A-B)
Glypturus acanthochirus Stimpson, 1866: 46; Stimpson, 1871: 121; Kingsley, 1899: 821
(footnote); De Man, 1928b: 19, 25, 180; Manning, 1987 (part.): 390, 398, fig. 3 (not figs. 4,
5 = G. armata); Poore & Suchanek, 1988: 201, fig. 4d; Manning & Felder, 1991: 778, figs. 2,
4; Dworschak, 1992: 209; Dworschak & Ott, 1993: 282; Blanco Rambla, 1995: 61, figs. 7,
8; Sakai, 1999c: 73, fig. 14i; Tudge et al., 2000: 144; Sakai, 2005b: 133.
? Glypturus acanthochirus — Schmitt, 1924: 93.
Callianassa (Callichirus) acanthochirus — Schmitt, 1935b: 4, 20, pl. 1 fig. 6, pl. 2 fig. 5, pl. 3
fig. 4, pl. 4 fig. 6; Heard & Reames, 1979: 52.
Callianassa acanthochirus — Gurney, 1944: 84; Biffar, 1971a: 655, figs. 3, 4; Heard & Reames,
1979: 52, fig. 1.
Callichirus acanthochirus — De Saint Laurent & Le Loeuff, 1979: 96.
Not: Callianassa acanthochirus — Rabalais et al., 1981: 103, fig. 3 [= Glypturus rabalaisae
Sakai, 2005b].
Material examined. — SMF 23508, 1 female (TL/CL 89.0/21.5), Key Largo, Florida Bay,
Florida, U.S.A., 16.v.1971, leg. J. Doerjes; USNM 266220, 1 male (TL/CL, 47.0/13.0 mm),
Parguera Bay, Lajas, Puerto Rico, Caribbean Sea, 06.i.1955, leg. B. Padilla, det. R.B. Manning.
Diagnosis. — Carapace with distinct anterolateral spines. Rostrum sharply
triangular. Anterolateral spines on carapace non-calcified proximally. A1 peduncle reaching to distal margin of A2 penultimate segment. Mxp3 ischiummerus operculiform; propodus subquadrate, and dactylus digitiform. Male
Plp1 uniramous, bisegmented, and subchelate distally (fig. 65A); male Plp2
biramous, endopod bearing mesiodistally a proximally-fused appendix masculina provided mesiomedially with slender, proximally-fused appendix interna (fig. 65B); male Plps3-5 endopods with narrow, triangular appendix interna. Telson wider than long, parallel-sided in proximal half, and curving toward rounded posterior margin. Uropodal endopod oval.
Remarks. — Biffar (1971) showed by his figure that the male Plp2 endopod
bears mesiodistally an articulate appendix masculina provided mesiomediallly
with an articulate appendix interna (Biffar, 1971a: 655, fig. 3h, i). However,
it has turned out through the present examination of the holotype (USNM
266220) that the male Plp2 endopod bears mesiodistally a proximally-fused
appendix masculina provided mesiomedially with a proximally-fused appendix interna (fig. 65B).
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CRM 013 – Katsushi Sakai
Type locality. — Florida Keys.
Distribution. — Atlantic coast of Florida, Gulf of Mexico, West Indies
(Dry Tortugas; Puerto Rico; Jamaica; Barbados; Antigua); Caribbean coast of
Colombia and Venezuela.
Glypturus armatus (A. Milne-Edwards, 1870)
Callianassa armata A. Milne-Edwards, 1870: 90, 101, pl. 1; De Man, 1902: 754; Kensley,
1975: 48, fig. 1A-H; Manning, 1987: 392 (partim).
Callianassa (Callichirus) armata — Borradaile, 1903: 547; De Man, 1928b: 28, 93, 109.
Callichirus laurae De Saint Laurent (in: Vaugelas & De Saint Laurent), 1984: 147, pl. 1
figs. A-D; Abu-Hilal et al., 1988: 233, 4 figs. [Type locality. — Gulf of Aqaba, Red Sea.]
Glypturus accanthochirus — Manning, 1987: 390 (partim), figs. 4, 5 (= G. armata, fig. 4 from
A. Milne-Edwards, 1870; fig. 5 from Kensley, 1975).
Glypturus motupore Poore & Suchanek, 1988: 198, figs. 1-3, 4a, table 1; Tudge et al., 2000:
144. [Type locality. — Bootless Inlet, Motupore Is., Papua New Guinea.]
Glypturus laurae — Poore & Suchanek, 1988: 201, fig. 4c; Dworschak, 1992: 209; Tudge et
al., 2000: 144; Robles et al., 2009: 317.
Glypturus armatus — Poore & Suchanek, 1988: 201; Sakai, 1999c: 76; Tudge et al., 2000: 144;
Sakai, 2005b: 137, table VIII.
Neocallichirus motupore — Sakai, 1999c: 105; Sakai, 2005b: 182.
Callianassa (Callianassa) laurae — Felder, 2001: 441.
Remarks. — The present species, Glypturus armatus, which was included
in Callianassa, and later in Glypturus by Manning (1987) as a synonym
of G. acanthochirus Stimpson, 1866, is different, however, from G. acanthochirus, because in G. armatus the telson is pentagonal with convex lateral
margins (Kensley, 1975, fig. 1f), while in G. acanthochirus the telson is wider
than long, and the lateral margins are parallel-sided in the proximal half, curving toward the rounded posterior margin. Poore & Suchanek (1988) mentioned
that “Glypturus motupore shares with G. laurae a broadly-based upturned rostrum, tapering eyestalks with a small cornea but differs most importantly in
the shape of the telson (narrower proximally in G. motupore) in the uropodal endopod (broader and more truncate in G. laurae). The larger chelipeds of
the two species differ in the shape of the cutting edge of the dactylus (without a notch in G. laurae), and in that lateral granules at the base of the fixed
finger are absent in G. motupore except in one individual, but are a constant
prominent feature of G. laurae”. However, those differences pointed out by
Poore & Suchanek (1988) are not decisive to separate those two species, so
in the present paper G. laurae as well as G. motupore are synonymized with
G. armatus. This is because, those three species are almost the same in the
following morphological features: larger cheliped, tail fan, spination on carapace, P3 propodus, relative length of A1 to A2, and Mxp3, i.e., features that
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433
are considered taxonomically important in the classification as proposed in the
present revision.
Type locality. — Fiji.
Distribution. — Mataiva, Tuamotu Arch.; Mauritius; Ternate, Indonesia;
Bootless Inlet, Motupore Is., Papua New Guinea; Fiji; Djibouti; Gulf of Aden;
Aqaba; Red Sea; intertidal, 5 to 45 m.
Glypturus articulatus (Rathbun, 1906)
Callianassa articulata Rathbun, 1906: 892, fig. 47; Chilton, 1911: 511.
Callianassa (Callichirus) articulata — De Man, 1928b: 28, 94, 108; Edmondson, 1944: 54,
fig. 9a-j.
Corallianassa articulata — Dworschak, 1992: 210, fig. 14a-e; Tudge et al., 2000: 144.
Glypturus articulatus — Sakai, 1999c: 76, fig. 15a-f; Sakai, 2005b: 138.
Diagnosis. — Mxp3 ischium-merus subsquare, merus rounded on distomesial margin; propodus subquadrate, and dactylus subovate. Male Plp1 bladeshaped, uniramous, and bisegmented; male Plp2 unknown. Telson wider than
long and subsquare, convex on lateral margins, and almost straight on posterior
margin.
Type locality. — Vicinity of Modu Manu, Hawaii, 23-33 fathoms [ca. 4260 m].
Distribution. — Honolulu, Oahu, Hawaii; Kermadec Islands; Gilbert Island.
Glypturus assimilis (De Man, 1928b)
Callianassa Martensi — De Man, 1888: 482-483, pl. 21 fig. 1 (Not: Callianassa Martensi
Miers, 1884).
Callianassa (Callichirus) assimilis De Man, 1928b: 28, 109.
Glypturus assimilis — Sakai, 1999c: 78, fig. 16a-f; Sakai, 2005b: 138.
Corallichirus bayeri Kensley, 2001: 328, figs. 1, 2. [Type locality. — Agat Bay, north of Alutom
Island, Guam, among rocks, 2.5-6 m.]
Callianassa assimilis — Tudge et al., 2000: 143.
Diagnosis. — Rostrum sharply triangular. Anterolateral spines of carapace non-calcified proximally. A1 peduncle shorter than A2 peduncle. Mxp3
ischium-merus rectangular; merus obliquely curved distally, and rounded on
mesial margin; propodus subquadrate, and dactylus digitiform. Male Plps1-2
unknown. Telson semicircular; uropodal endopod fusiform.
Type locality. — Indonesia, Ambon.
Distribution. — Ambon, Indonesia; Gilbert Is.; Agat Bay, north of Alutom
Island, Guam.
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CRM 013 – Katsushi Sakai
Glypturus borradailei (De Man, 1928a)
Callianassa (Callichirus) longiventris var. borradailei De Man, 1928a: 27; De Man, 1928b: 29,
108.
Callianassa (Callichirus) borradailei — Ward, 1942: 62.
Corallianassa borradailei — Manning, 1987: 394, figs. 7 (from Borradaile, 1902), 8; Manning,
1992, fig. 1a; Tudge et al., 2000: 144; Ngoc-Ho, 2002b: 224.
Diagnosis. — Rostrum sharply pointed apically, overreaching middle of
eyestalks. Anterolateral margins of carapace with sharp teeth bearing noncalcified proximal area. Eyestalks oval; cornea distinct and located apically. A1
peduncle reaching middle of A2 distal segment. Mxp3 propodus subquadrate,
and dactylus digitiform. P1 ischium and merus armed with sharp teeth on
ventral margin. Abdominal somite 2 longer than somite 6. Telson shaped as
inverted trapezoid. Uropodal exopod bilobate and broadened in distal half.
Type locality. — Goidu, Goifurfehendu Atoll, Maldive Archipelago.
Distribution. — Only known from the type locality.
Glypturus collaroy (Poore & Griffin, 1979)
Callianassa collaroy Poore & Griffin, 1979: 260, figs. 24, 25.
Glypturus collaroy — Sakai, 1988: 61; Sakai, 2005b: 139, fig. 29.
Corallianassa collaroy — Sakai, 1992b: 212, fig. 1; Tudge et al., 2000: 144; Davie, 2002: 460.
Neocallichirus collaroy — Sakai, 1999c: 94 (key), 98.
Diagnosis. — Mxp3 ischium-merus oval; merus rounded, with spine on
distomesial margin; propodus subquadrate, and dactylus subovate. Male Plp1
uniramous, bisegmented, and w-shaped distally; male Plp2 biramous, endopod
with appendices interna and masculina. Telson wider than long and subsquare,
convex on lateral margins, and concave medially on posterior margin.
Type locality. — Collaroy, Long Reef, New South Wales, Australia.
Distribution. — Collaroy, New South Wales; Society Islands — Maharepa,
Moorea, French-Polynesia; coral reef, sand bottom, low intertidal among
boulders.
Glypturus haswelli (Poore & Griffin, 1979)
Callianassa haswelli Poore & Griffin, 1979: 263, figs. 26, 27.
Callianassa nakasonei — Sakai, 1987a: 306; Tudge et al., 2000: 143.
Glypturus haswelli — Sakai, 1999c: 82; Davie, 2002: 460.
Diagnosis. — Mxp3 ischium-merus subsquare; merus rounded on mesiodistal margin; propodus subquadrate, and dactylus digitiform. Male Plp1 bladeshaped, uniramous, bisegmented, and subchelate; male Plp2 biramous, endopod without appendices interna and masculina (Poore & Griffin, 1979,
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435
fig. 27m, n). Telson shaped in inverted trapezoid, bearing low median convexity on posterior margin.
Remarks. — Callianassa (Callichirus) nakasonei Sakai, 1967 was synonymized with Callianassa haswelli Poore & Griffin, 1979, by Manning
(1987: 392), and later with Glypturus martensi from Mauritius by Sakai
(2005b: 18). However, C. nakasonei is different from G. martensi, because in
G. martensi the palm of the larger cheliped is denticulate on the distal margin,
and the telson is trapezoid, bearing no median convexity on the posterior margin; whereas in C. nakasonei the palm of the larger cheliped is not denticulate
on the distal margin and the telson is trapezoid, bearing a median convexity on
the posterior margin.
Type locality. — Queensland, Whitsunday Group, Australia.
Distribution. — Torres Strait; Queensland, Islands off northern and central
Queensland coast.
Glypturus lanceolatus (Edmondson, 1944)
Callianassa (Callichirus) lanceolata Edmondson, 1944: 52, fig. 8a-i.
Callianassa (Callichirus) nakasonei Sakai, 1967: 46, fig. 3. [Type locality. — East coast of
Tonaki Island, Okinawa-Island, Ryukyu Archipelago.]
Glypturus lanceolatus — Sakai, 1999c: 83; Sakai, 2005b: 142.
Diagnosis. — Mxp3 ischium-merus subsquare; merus rounded on distomesial margin; propodus subtriangular, and dactylus digitiform. Male Plps1-2 unknown. Telson wider than long and trapezoid, bearing low median convexity
on posterior margin.
Type locality. — Hanauma Bay, Oahu, in shallow water on the reef.
Distribution. — Oahu, Hawaii; Queensland, Whitsunday Group, Australia
(Poore & Griffin, 1979); and Okinawa, Japan (Sakai, 1967).
Glypturus longiventris (A. Milne-Edwards, 1870)
(fig. 65E-F)
Callianassa longiventris A. Milne-Edwards, 1870: 92, 101; Biffar, 1971a: 651, 653, 685,
figs. 13, 14.
Callianassa (Callichirus) longiventris — Borradaile, 1903: 547; De Man, 1928b: 19, 29 (list),
94, 108 (key); Schmitt, 1935b: 4, pl. 1 fig. 4, pl. 2 fig. 3, pl. 3 fig. 3, pl. 4 fig. 3.
Callianassa (Callichirus) Coutierei Nobili, 1904: 237; Nobili, 1906a: 60; Nobili, 1906b: 110,
pl. 7 fig. 1; De Man, 1928b: 28, 109, 174, 179. [Type locality. — Djibouti.]
?Callianassa (Callichirus) longiventris — Borradaile, 1904: 752, pl. 58 fig. 2. [Type locality.
— Hulue, Male Atoll, Maldives.]
Callianassa placida De Man, 1905: 612; Hernández-Aguilera et al., 1986: 206. [Type locality.
— Off Laiwui, coast of Obi Major, Indonesia.]
Glypturus grandimanus — Balss, 1924: 179, figs. 3, 4; Biffar, 1971a: 640, 649; Manning, 1987:
399. [Not: Callianassa grandimana Gibbes, 1850.]
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CRM 013 – Katsushi Sakai
Callianassa (Callichirus) placida — De Man, 1928b: 29, 93, 108, 171, pl. 18 fig. 29-29b, pl. 19
fig. 29c-e; Chace, 1962: 617.
Callianassa (Callichirus) hartmeyeri Schmitt, 1935b: 3, 4. [Replacement name for Glypturus
grandimanus sensu Balss, 1924.] [Type locality. — Jamaica Is.: Kingston (Kingston Harbor,
17◦ 57′ N 76◦ 47′ W).]
Callianassa (Callichirus) oahuensis Edmondson, 1944: 56, fig. 10a-h. [Type locality. —
Hanauma Bay, Oahu.]
Callianassa hartmeyeri — Biffar, 1971a: 640, 641, 649, 651, 653; Manning, 1987: 388, 399.
Callichirus longiventris — De Saint Laurent & Le Loeuff, 1979: 97.
Callichirus sp. aff. placidus — De Saint Laurent & Le Loeuff, 1979: 97.
Corallianassa longiventris — Manning, 1987: 392, fig. 6; Manning & Felder, 1991: 777;
Coelho, 1997: 148; Melo, 1999: 366, figs. 243-244; Dworschak et al., 2006: 1369, figs. 2, 5,
6, tab. 2; Coelho et al., 2007: 5 (table).
Corallianassa hartmeyeri — Manning, 1988: 884, figs. 1, 2; Manning & Chace, 1990: 34,
figs. 18, 19; Manning & Felder, 1991: 777; Manning, 1992: 571; Coelho, 1997: 148; Melo,
1999: 364, figs. 241-242; Tudge et al., 2000: 144; Coelho et al., 2007: 5.
Corallichirus placidus — Manning, 1992: 571; Tudge et al., 2000: 144.
Glypturus coutierei — Sakai, 1999c: 78, figs. 17a-f, 18a-d.
Glypturus longiventris — Sakai, 1999c: 74; Sakai, 2005b: 134.
Glypturus hartmeyeri — Sakai, 1999c: 74; Sakai, 2005b: 134.
Callianassa coutierei — Tudge et al., 2000: 143.
Material examined. — SMF 23509, 1 ovig. female (TL/CL, 11.0/2.4 mm), Bahia Gairaca,
ca. 20 km N.E. of Santa Marta, Magdalena, Colombia, 16.iv.1980; USNM 122449, 1 male, S.W.
reef of Somerset, Bermuda, 25 fm (45.7 m), 15.vi.1964, leg. T. Tucker, det. T.A. Biffar; SMNH
16239 1 female (lacking posterior to abdominal somite 2), Viti Levu, Namuka, Fiji Is., barrier
reef, 18.vi.1917, leg. S. Bock.
Diagnosis. — Mxp3 ischium-merus operculiform; propodus subquadrate,
and dactylus digitiform. Male Plp1 uniramous, bisegmented, and w-shaped
distally (fig. 65E); male Plp2 biramous, endopod bearing distally proximallyfused appendix masculina, and mesiodistally slender, proximally-fused appendix interna (fig. 65F). Telson wider than long and trapezoid, bearing median convexity on posterior margin. [Cf. Biffar, 1971a: 686, 688, figs. 13e, g;
14a, e.]
Remarks. — The present species, Glypturus longiventris (A. MilneEdwards, 1870) is similar to G. acanthochirus Stimpson, 1866 in that the male
Plp1 is uniramous and bisegmented, and the distal segment is chelate distally;
the male Plp2 is biramous, and the endopod bears a proximally-fused appendix
masculina and a small, proximally-fused appendix interna. However, they are
different from each other in the features of the telson and anterolateral spines
on the carapace. In G. longiventris the telson is wider than long and trapezoid,
and the anterolateral spines on the carapace are calcified proximally; whereas
in G. acanthochirus the telson is wider than long, and parallel-sided in the
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437
proximal half, curving toward the rounded posterior margin, and the anterolateral spines on the carapace are not calcified proximally.
Four species in Glypturus, G. coutierei (Nobili, 1904) from Djibouti;
Mindanao, Philippines; Tahiti; Fiji; Goidu, Goifurfehendu Atoll, Maldive
Archipelago; Indonesia; Tuléar, S.W. Madagascar; G. placida (De Man, 1905)
from Indonesia; G. hartmeyeri (Schmitt, 1935b) from Jamaica Is.: Kingston;
and G. oahuensis (Edmondson, 1944) from Hanauma Bay, Oahu, Hawaii,
show almost the same features as those of G. longiventris (A. Milne-Edwards,
1870) in the eyestalks, the anterolateral spines of the carapace, the denticulation on the ventral margins of P1 ischium and merus, and also in the telson
and uropodal endopod, so that those four species are here synonymized with
G. longiventris (A. Milne-Edwards, 1870) from Martinique, though in G. hartmeyeri and G. oahuensis no males are known. Therefore, it is safely concluded
that G. longiventris is distributed in the Atlantic and Indo-Pacific oceans.
Type locality. — Martinique.
Distribution. — Bermuda; Carrie Bow Cay; southeastern Florida; Belize;
Caribbean Sea (Jamaica; Martinique; Virgin Is.), Pará and Pernambuco, Brazil
(as G. longiventris). — Jamaica, Caribbean Sea, and Ascension, Alagoas,
Brazil, South Atlantic (as G. hartmeyeri). — Hawaii (Hanauma Bay, Oahu) (as
G. oahuensis). — Mindanao, Philippines; Tahiti; Fiji; Goidu, Goifurfehendu
Atoll, Maldive Archipelago; Indonesia (off Seba, Savu; off Laiwui, coast of
Obi Major); Gulf of Aden (Perim; Djibouti; Aden); Tuléar, S.W. Madagascar
(as G. coutierei). — East coast of Tonaki Island, Okinawa-Island, Ryukyu
Archipelago; Mauritius; Ambon, Indonesia; north and central Queensland,
Australia; Sri Lanka; Belligom, West Pakistan (as G. martensi).
Glypturus martensi (Miers, 1884)
Callianassa Martensi Miers, 1884a: 13-15, pl. 1 fig. 1; Lanchester, 1900: 261, pl. 12 fig. 4, 4a;
Nobili, 1906b: 111, fig. 7.
Callianassa (Callichirus) Martensi — Borradaile, 1903: 547; De Man, 1928b: 29, 109, 171.
Callianassa (Callichirus) martensi — Tirmizi, 1974: 286, figs. 1-4.
Callichirus martensi — De Saint Laurent & Le Loeuff, 1979: 97.
Callianassa martensi — Sakai, 1984: 99, fig. 3; Dworschak, 1992: 200, fig. 8a-e.
Glypturus martensi — Sakai, 1988: 61; Tudge et al., 2000: 144; Davie, 2002: 460; Sakai, 2005b:
142.
Diagnosis. — Mxp3 ischium-merus subsquare; propodus subtriangular,
and dactylus digitiform. In larger cheliped palm denticulate on distal margin
(Sakai, 1984, fig. 3B). Male Plp1 uniramous, bisegmented, and subchelate distally; male Plp2 biramous, endopod bearing distally proximally-fused appendix masculina with appendix interna mesioproximally. Telson wider than long
438
CRM 013 – Katsushi Sakai
and trapezoid, and almost straight on posterior margin, with median convexity.
[Cf. Tirmizi, 1974, figs. 1, 4C, D.]
Type locality. — Mauritius.
Distribution. — East coast of Tonaki Island, Okinawa-Island, Ryukyu
Archipelago; Ambon, Indonesia; north and central Queensland, Australia; Sri
Lanka; Belligom, West Pakistan; Mauritius.
Glypturus rabalaisae sp. nov.
Callianassa acanthochirus — Rabalais et al., 1981: 103, fig. 3.
Glypturus rabalaisae Sakai, 2005b: 135 [invalid, as no type specimen was designated].
Material. — Holotype, male (collected on 21.i.1976); paratypes, females (17.ix.1975),
(11.x.1976), South Texas outer continental shelf, northwestern Gulf of Mexico (Rabalais et
al., 1981).
Remarks. — The present species was described without designating any
types by Sakai (2005b), so the species name is invalid according to Art. 72.3
of the ICZN. In the present revision this species is redescribed by designating
the type.
The present species is different from G. acanthochirus Stimpson, 1866,
because in G. acanthochirus the A1 peduncle reaches to the distal margin
of the A2 penultimate segment, and the telson bears no median spine on the
posterior margin; whereas in G. rabalaisae sp. nov. the A1 peduncle reaches
to the middle of the A2 distal segment, and the telson bears a median spine on
the posterior margin.
Type locality. — Off Galveston, Texas, 15-65 to 90 m.
Distribution. — Only known from the type locality.
Genus Grynaminna Poore, 2000
Grynaminna Poore, 2000: 150.
Diagnosis. — Rostrum low and obtusely triangular. Carapace with dorsal
oval, but without rostral carina, hepatic sulcus, or cardiac prominence. A1
peduncle longer and stouter than A2 peduncle; distal segment elongate. Mxp3
ischium-merus broadly subpediform; propodus broadened and truncate on
ventral margin; dactylus slender and digitiform; exopod rudimentary. P1
unequal, in larger cheliped merus convex and denticulate on ventral margin.
P2 chelate. P3 propodus broad and pediform. P4 subchelate. P5 chelate.
Abdominal somites 3-5 without dorsal ornament. Male Plp1 uniramous and
bisegmented, and distal segment subchelate distally; male Plp2 biramous,
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439
endopod bearing distally subsquare appendix masculina with distal setae and
small appendix interna mesiomedially (fig. 66A, B); male Plps3-5 biramous
and foliaceous, endopods with appendix interna. Telson wider than long and
hexagonal. Uropodal endopod elongate and ovate; uropodal exopod broadly
triangular, bearing dorsal plate.
Remarks. — Poore (2000: 150) established Grynaminna Poore, 2000 and
mentioned in the remarks that his genus resembles Callichirus, but differs from
all the callichirine genera in the digitiform appendix interna of male Plp2.
Poore’s (2000) original designation of the present genus made at that time,
about ten years ago, should not have been rejected (cf. Sakai, 2005b: 126, 187188), because he also indicated the importance of the features of male Plps1-2,
though his designation was made without examining all the genera included in
the present Callichirinae. The present classification was made systematically
by applying the different features in the structures of male Plps1-2.
Type species. — Grynaminna tamakii Poore, 2000, by original designation
and monotypy. The gender of the generic name, Grynaminna, is feminine.
Species included. — Only the type species, Grynaminna tamakii Poore,
2000.
Grynaminna tamakii Poore, 2000
(fig. 66A-B)
Grynaminna tamakii Poore, 2000: 150, figs. 1-4.
Podocallichirus tamakii — Sakai, 2005b: 193, figs. 39, 40.
Material examined. — NSMT-Cr. 12532, paratype, 1 male (TL/CL, 66.0/13.5 mm, chelipeds
missing); NSMT-Cr. 12533, paratype, 1 male (TL/CL, 65.0/13.0 mm, chelipeds missing);
NSMT-Cr. 12534, paratype, 1 male (TL/CL, 68.0/15.3 mm, chelipeds missing); S. end of
Shimabara Peninsula, Nagasaki Pref., Kyushu, Japan, leg. A. Tamaki, 8.vii.1998.
Remarks. — The present species was included in Podocallichirus Sakai,
1999c, but it has turned out to be different from the type species of Podocallichirus, P. madagassa Lenz & Richters, 1881, in the form of the male Plps1-2.
In P. madagassa, the male Plp1 is uniramous and bisegmented, and the distal
segment is bilobed distally and curved inward distally (fig. 66C); the male Plp2
is biramous, and the endopod bears mesiodistally an oval appendix masculina
with a proximally-fused appendix interna (fig. 66D). In P. tamakii, in contrast,
the male Plp1 is uniramous, bisegmented, and its distal segment is subchelate
distally; the male Plp2 is biramous, and the endopod bears distally a subsquare
appendix masculina with distal setae and a small appendix interna mesiomedially (fig. 66A, B).
440
CRM 013 – Katsushi Sakai
Fig. 66. Grynaminna tamakii Poore, 2000 and Podocallichirus madagassus (Lenz & Richters,
1881). A, B, D, male Plp2, C, male Plp1. A, B, Grynaminna tamakii, NSMT-Cr. 12532,
paratype, male (TL/CL, 66.0/13.5 mm, chelipeds missing), south end of Shimabara peninsula,
Nagasaki Pref., Kyusyu, Japan; C, D, Podocallichirus madagassus, SMF 4953, paratype, 1 male
(TL/CL, 49.0/10.7 mm), Madagascar. Scales 1 mm.
Therefore, based on the type species, P. tamakii, the original concept of
the genus Grynaminna is reinstated in the present classification, in which the
Callichirinae are classified mainly on the basis of the characters of the male
pleopods.
Type locality. — Minami-Arima-cho, Shimabara Peninsula, Nagasaki Prefecture, Japan, 32◦ 37′ N 130◦ 13′ E.
Distribution. — Known only from the type locality.
Genus Lepidophthalmoides gen. nov.
Diagnosis. — Rostrum small and acutely triangular, lacking rostral carina.
Carapace with dorsal oval, but without hepatic sulcus and prominence, and
anterolateral margin of carapace with or without spine. A1 peduncle longer
and stouter than A2 peduncle; distal segment elongate. Mxp3 ischium-merus
broadly rectangular; propodus broadened and rounded on ventral margin;
dactylus digitiform or subovate; exopod rudimentary or absent. P1 unequal; in
larger cheliped merus with meral crest ventroproximally. P2 chelate. P3 propodus broadened. P4 subchelate. P5 chelate. Abdominal somites 3-5 without dor-
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441
sal ornament. Male Plp1 uniramous and unsegmented or bisegmented, distal
segment subchelate or chelate distally; male Plp2 biramous, endopod with appendix masculina, but without appendix interna; or endopod with appendices
masculina and interna; male Plps3-5 biramous and foliaceous, endopods with
appendix interna. Telson wider than long and varied in shape. Uropodal endopod oval or subrhombic, overreaching telson; uropodal exopod broadened on
distolateral corner, bearing dorsal plate. [Adapted from Sakai, 1999c, 2002.]
Remarks. — Manning & Felder (1991: 778) showed that Lepidophthalmus
eiseni Holmes, 1904 is a junior synonym of L. bocourti (A. Milne-Edwards,
1870), the type species of the genus Lepidophthalmus. However, Felder (2003:
436) subsequently distinguished L. eiseni from L. bocourti, because they
are different from each other: in L. eiseni the male Plp1 is chelate distally
(Felder, 2003, fig. 24), whereas in L. bocourti it is subchelate (Felder, 2003,
figs. 7, 15). It has turned out, however, through the present examination of
the male holotype of L. bocourti, that the male Plp1 distal segment is not
subchelate, but simple (fig. 67A), so that Lepidophthalmus eiseni is to be
excluded from Lepidophthalmus and is here reclassified under the new genus
Lepidophthalmoides gen. nov. as the type species, Lepidophthalmoides eiseni.
In Lepidophthalmoides, the Plp2 endopod can have two types of shape. In
Lepidophthalmoides eiseni, the male Plp2 endopod bears mesiodistally an appendix masculina with distal setae, but no appendix interna; in Lepidophthalmoides jamaicense (Schmitt, 1935b) it bears mesiodistally an appendix masculina provided mesiomedially with a proximally-fused appendix interna.
Type species. — Lepidophthalmus eiseni Holmes, 1904, by present designation. The gender of the new name, Lepidophthalmoides, is feminine.
Species included. — Lepidophthalmoides eiseni (Holmes, 1904); L. jamaicense (Schmitt, 1935b); L. louisianensis (Schmitt, 1935b); L. richardi
(Felder & Manning, 1997); L. rosae (Nobili, 1904); L. siriboia (Felder &
Rodrigues, 1993); L. socotrensis (Sakai & Apel, 2002); L. tridentata (Von
Martens, 1869).
Etymology. — The generic name Lepidophthalmoides is composed of a
generic name, Lepidophthalmus, with the Latin suffix -oides, meaning “alike”,
or “similar to”.
K EY
TO THE
S PECIES
OF THE
G ENUS LEPIDOPHTHALMOIDES
1 – Anterolateral spines on carapace absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Anterolateral spines on carapace present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 – Dactylus of smaller cheliped denticulate on dorsal margin . . . . . . . . . . . . . . L. socotrensis
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CRM 013 – Katsushi Sakai
– Dactylus of smaller cheliped smooth on dorsal margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Merus of larger cheliped not concave subproximally on dorsal margin . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. louisianensis
– Merus of larger cheliped distinctly concave subproximally on dorsal margin . . . . . . . . . 4
4 – Uropodal endopod broadly rhombic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. jamaicense
– Uropodal endopod narrowly rhombic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. eiseni
5 – Merus of larger cheliped distinctly concave subproximally on dorsal margin . . . . . . . . . 6
– Merus of larger cheliped not concave subproximally on dorsal margin . . . . . . . . . . . . . . 7
6 – Telson rounded on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. tridentata
– Telson straight on posterior margin, bearing median convexity . . . . . . . . . . . . . L. richardi
7 – Fixed finger of male larger cheliped unarmed on cutting edge . . . . . . . . . . . . . . . . L. rosae
– Fixed finger of male larger cheliped armed with median tooth on cutting edge . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. siriboia
Lepidophthalmoides eiseni (Holmes, 1904)
Lepidophthalmus Eiseni Holmes, 1904: 311, pl. 35 figs. 6-13.
Callianassa (Callichirus) Eiseni — De Man, 1928b: 28.
Callianassa Eiseni — De Man, 1928b: 110.
Callianassa (Callichirus) eiseni — Schmitt, 1935b: 9.
Callianassa eiseni — Holthuis, 1954a: 12-15, fig. 3; Holthuis, 1954c: 160; Bott, 1955: 47,
fig. 6a-g.
Lepidophthalmus eiseni — Felder & Rodrigues, 1993: 373; Tudge et al., 2000: 144; Felder,
2003: 429, figs. 20-29.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment chelate
distally; male Plp2 biramous, endopod bearing mesiodistally appendix masculina with distal setae, but without appendix interna (cf. Holthuis, 1954,
fig. 3i); Plps3-5 endopods with small, triangular appendix interna.
Possible type locality. — San Jose del Cabo, Lower California (Felder,
2003).
Distribution. — Mexico, Guatemala, Nicaragua, Costa Rica (Felder, 2003).
Lepidophthalmoides jamaicense (Schmitt, 1935)
Callianassa jamaicense Schmitt, 1935b: 1, 9, pl. 1 fig. 1, pl. 2 figs. 6, 8, pl. 4 fig. 1; Biffar,
1971a: 650, 654; Coelho & Ramos, 1973: 162 (partim); Abel & Kim, 1986: 27, 295, 296,
302-303, figs. j, k, l (partim); Manning, 1987: 397; Dworschak, 1992: 196 (partim) (not
fig. 4a-d = L. siriboia).
Callianassa jamaicensis — Holthuis, 1974: 231.
Callichirus jamaicensis — De Saint Laurent & Le Loeuff, 1979: 67, 69 (only species name);
Coelho & Ramos-Porto, 1987: 30.
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443
Lepidophthalmus jamaicense — Felder et al., 1991: 101A (part.); Manning & Felder, 1991: 778,
fig. 13a-e (not f, which is defined as L. jamaicense var. louisianensis); Felder & Rodrigues,
1993: 357, 358, 367, 373; Sakai, 1999c: 66; Tudge et al., 2000: 144; Sakai, 2005b: 145.
Diagnosis. — Mxp3 ischium-merus rectangular; propodus subtriangular,
and dactylus digitiform; exopod rudimentary. Male Plp1 uniramous and bisegmented, distal segment chelate distally; male Plp2 biramous, endopod bearing
mesiodistally oval appendix masculina with distal setae, provided mesiomedially with proximally-fused appendix interna with hooklets (cf. Felder & Manning, 1997, fig. 1h). Telson wider than long and subrectangular, and weakly
trilobate on posterior margin (cf. Schmitt, 1935b: 9-12, pl. 4 fig. 1).
Type locality. — Jamaica, Montego Bay.
Distribution. — Caribbean Sea — Dangriga and Sao Luiz-MA, Rio Anil,
Belize; Honduras, sandy beach between stones; Jamaica, Montego Bay, blackish pond.
Lepidophthalmoides louisianensis (Schmitt, 1935)
[Abbreviated list of synonymy.]
Callianassa jamaicense var. louisianensis Schmitt, 1935b: 1, 12-15, pl. 1 fig. 2, pl. 2 figs. 2, 4,
7, pl. 4 fig. 4; Hedgpeth, 1950: 113-114; Biffar, 1971a: 641-642, 650; Phillips, 1971: 165196, figs. 1-3a, c, e, 6, 7b, c, d, 8b, c, d, tabs. 1-3, 5-7; Fotheringham, 1980: 63, fig.; Felder
et al., 1986: 91; Manning & Felder, 1989: 9.
Lepidophthalmus louisianensis — Felder & Staton, 1990: 137A; Felder et al., 1991: 101A
(partim); Lemaitre & Rodrigues, 1991: 629; Manning & Felder, 1991: 778 (partim); Manning
& Felder, 1992: 560; Felder & Felgenhauer, 1993: 263-276, figs.; Felder & Manning, 1997:
309-331, figs. 1-7; Sakai, 1999c: 67, fig. 14a-b; Dworschak, 2000b: 99; Tudge et al., 2000:
144; Atkinson & Taylor, 2004: 45, tab. 2; Robles et al., 2009: 317.
Diagnosis. — Male Plp1 uniramous and bisegmented, and subchelate
distally; male Plp2 biramous, endopod bearing distally appendix masculina
with proximally-fused appendix interna mesiomedially (cf. Sakai, 1999c,
fig. 14b). Telson wider than long and subrectangular, bearing weak median
convexity on posterior margin (cf. Felder & Rodrigues, 1993, fig. 1i).
Type locality. — Grand Isle, Louisiana.
Distribution. — Florida (Perdido Key, Big Lagoon; 20-50 m); Alabama
(Nobile Bay and Dauphin Is. near airport; intertidal); Mississippi (Bay St.
Louis; intertidal); and Louisiana.
Lepidophthalmoides richardi (Felder & Manning, 1997)
Lepidophthalmus jamaicense complex — Felder et al., 1991: 101A (part.).
Lepidophthalmus richardi Felder & Manning, 1997: 320-329, figs. 4a-j, 5a-f, 6a-i, 7a; Sakai,
1999c: 68; Tudge et al., 2000: 144.
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CRM 013 – Katsushi Sakai
Lepidophthalmus sp. “a” — Staton et al., 2000: 161, figs. 1-3, tabs. 3-5.
Lepidophthalmus manningi Felder & Staton, 2000: 170, figs. 1, 2. [Type locality. — Laguna
San Augustin, near village of Palma Sola, 19◦ 55.23′ N 96◦ 31.85W, Veracruz, Mexico.]
Diagnosis. — Male Plp1 uniramous, bisegmented, and chelate distally; male
Plp2 biramous, endopod bearing distolaterally a proximally-fused appendix
masculina with marginal setae, and a small appendix interna mesiodistally.
[Cf. Felder & Manning, 1997, figs. 6e-f, i.]
Remarks. — Felder & Staton (2000) described the new species Lepidophthalmus manningi, and differentiated it from L. richardi Felder & Manning,
1997, mentioning the reason that “L. manningi typically has a deeper proximal
meral notch on the superior margin of the major cheliped, a larger lobe on the
proximal article of the female first pleopod, a larger appendix interna on the
male second pleopodal endopod, a narrower distal lobe on the female second
pleopodal endopod, and a less strongly developed carination on mesial margins
of the lateral ventral plates on the second abdominal somite of mature male”. It
seems difficult, however, to distinguish L. manningi from L. richardi by those
minor features, so that L. manningi is here synonymized with L. richardi. In
the present revision, Lepidophthalmus richardi is reclassified under the new
genus Lepidophthalmoides, because the male Plp1 distal segment is chelate
distally as in Lepidophthalmoides eiseni, the type species of the genus Lepidophthalmoides.
Type locality. — Intertidal shoreline at Pelican Beach Hotel, near Dangriga,
Belize.
Distribution. — Belize; Veracruz, Tabasco, and western Campeche, Mexico.
Lepidophthalmoides rosae (Nobili, 1904)
Callianassa (Callichirus) Rosae Nobili, 1904: 237; Nobili, 1906b: 108, pl. 7 fig. 2; De Man,
1928b: 29, 110; Balss, 1933: 88.
Lepidophthalmus rosae — Sakai, 1999c: 71, fig. 14g-h; Sakai, 2005b: 151, figs. 30-32.
Callianassa rosae — Tudge et al., 2000: 143.
Diagnosis. — Male Plpl unsegmented and chelate distally; Plp2 biramous,
endopod bearing mesiodistally appendix masculina with a few long distal
setae, provided mesiomedially with proximally-fused appendix interna with
distal hooklets. [Cf. Sakai, 1999c, fig. 14g.]
Type locality. — Red Sea.
Distribution. — Madagascar; Red Sea; Lombok, Indonesia; Mindanao,
Philippines.
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445
Lepidophthalmoides siriboia (Felder & Rodrigues, 1993)
Callianassa jamaicense — Biffar, 1971a: 650, 654 (partim); Abele & Kim, 1986: 27 (part.);
Griffis & Suchanek, 1991, tab. 2; Dworschak, 1992: 196 (partim), fig. 4a-d. [Not: L. jamaicense (Schmitt, 1935b).]
Callianassa (Callichirus) jamaicensis — Rodrigues, 1971: 198 (partim).
Callianassa jamaicensis — Rodrigues, 1971: 202-204, figs. 21-40, tab. 2 (partim); Coelho &
Ramos, 1973: 162 (part.); Tiefenbacher, 1976: 314 (part.), fig. 1c, d (not: fig. 1a, 1b =
L. louisianensis (Schmitt, 1935b)); Griffis & Suchanek, 1991, tab. 2.
Callichirus jamaicensis — De Saint Laurent & Le Loeuff, 1979: 67, 96 (partim); Coelho &
Ramos-Porto, 1987: 30 (partim).
Lepidophthalmus siriboia Felder & Rodrigues, 1993: 367, figs. 2e-h, 4a-f, 6a-l; Coelho, 1997:
148; Rodrigues et al., 1998: 380; Melo, 1999: 370, figs. 247-248; Sakai, 1999c: 69; Coelho
et al., 2007: 5 (tab.).
Lepidophthalmus siriboius — Tudge et al., 2000: 144.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment chelate; male Plp2 biramous, endopod bearing mesiodistally ovoid appendix
masculina with distal setae, and proximally-fused appendix interna with
hooklets mesiomedially. [Cf. Felder & Rodrigues, 1993, fig. 6c, d, e, f, g.]
Type locality. — Brazil: Maranhão, São Luís, mouth of Rio Anil.
Distribution. — Brazil — Marapanin, Pará; mouth of Rio Anil, São Luís,
Maranhão; Pernambuco; mouth of Rio Gramame, Joáo Pessoa, Paraiba; mouth
of Rio Caravelas, Bahia.
Lepidophthalmoides socotrensis (Sakai & Apel, 2002)
Lepidophthalmus socotrensis Sakai & Apel, 2002: 278, figs. 3-7.
Diagnosis. — Mxp3 ischium-merus subsquare; merus rounded on distomesial angle; propodus subquadrate, and dactylus digitiform. Male Plp1 uniramous and bisegmented, distal segment subchelate distally; male Plp2 biramous, endopod bearing mesiodistally appendix masculina with distal setae and
proximally-fused appendix interna. Telson wider than long and subrectangular,
weakly trilobate on posterior margin.
Type locality. — Khawr Girmah, Socotra Island, Republic of Yemen,
intertidal mudflat.
Distribution. — Khawr Girmah; Khawr Qariyah; Qalansiyah Lagoon; Ras
Kharmah, Socotra Island, Socotra Archipelago, Yemen; intertidal mud, sand/
mud in eulittoral, open lagoon, eulittoral.
Lepidophthalmoides tridentata (Von Martens, 1869)
Callianassa tridentata Von Martens, 1869: 614; A. Milne-Edwards, 1870: 94, 101.
446
CRM 013 – Katsushi Sakai
Callianassa (Callichirus) tridentate — Borradaile, 1903: 547; De Man, 1928a: 27, pl. 7 fig. 1313h; De Man, 1928b: 30, 93, 110, 171, 175; Sakai, 1970b: 393, figs. 1-3.
Lepidophthalmus tridentatus — Sakai, 1999c: 71, fig. 14e-f; Sakai, 2005b: 144; Dworschak,
2007: 122; Robles et al., 2009: 317.
Corallichirus tridentatus — Tudge et al., 2000: 144; Itani, 2007: 203.
Diagnosis. — Male Plp1 uniramous, bisegmented, and chelate distally. Male
Plp2 unknown.
Type locality. — Java, Indonesia.
Distribution. — Java, Indonesia; Okusu-gawa, Okinawa Island; MiyaraBay, Ishigaki Is., Ryukyu Archipelago (Itani, 2007: 203).
Genus Lepidophthalmus Holmes, 1904
Lepidophthalmus Holmes, 1904: 310; Manning & Felder, 1991: 778, figs. 5, 13; Poore, 1994:
102; Sakai, 1999c: 64; Sakai & Apel, 2002: 278; Sakai, 2005b: 143.
Lepidophthalmus s. str. — Felder, 2001: 440.
Diagnosis. — Rostrum spinous, single-pointed or trispinose apically, lacking rostral carina. Carapace with dorsal oval, but without hepatic sulcus and
cardiac prominence; anterolateral margin of carapace with or without spine.
A1 peduncle longer and stouter than A2 peduncle; distal segment elongate.
Mxp3 ischium-merus broadly rectangular; propodus broadened and rounded
on ventral margin; dactylus digitiform with obtuse tip; exopod rudimentary
or absent. P1 unequal, larger cheliped with ventral meral hook. P2 chelate.
P3 propodus broadened. P4 subchelate. P5 chelate. Abdominal somites 3-5
without dorsal ornament. Male Plp1 uniramous and bi- or tri-segmented, distal
segment simple (fig. 67A); male Plp2 biramous, endopod bearing only appendix interna, or mesiodistally appendix masculina with marginal setae and
proximally-fused appendix interna mesiomedially (fig. 67B); male Plps3-5 biramous and foliaceous, endopod with appendix interna. Telson wider than
long, posterior margin usually trilobate. Uropodal endopod subrhombic or
leaf-like, overreaching telson; uropodal exopod broadened on distolateral corner, bearing dorsal plate. [Adapted from Sakai, 1999c; Sakai & Apel, 2002.]
Remarks. — The genus Lepidophthalmus is distinguished from the genera
Glypturus Stimpson, 1866 and Neocallichirus Sakai, 1988, because in Lepidophthalmus the A1 peduncle is distinctly longer than the A2 peduncle, and
the male Plp1 distal segment is simple.
Type species. — Callianassa bocourti A. Milne-Edwards, 1870, by monotypy. Gender of generic name, Lepidophthalmus, masculine.
Species included. — Lepidophthalmus bocourti (A. Milne-Edwards, 1870);
L. grandidieri (Coutière, 1899); L. rafai Felder & Manning, 1998; L. sinuensis
Lemaitre & Rodrigues, 1991; L. turneranus (White, 1861).
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K EY
TO THE
S PECIES
OF THE
447
G ENUS LEPIDOPHTHALMUS
1 – Anterolateral projections on carapace indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Anterolateral projections on carapace distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 – Male Plp1 distal segment simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. bocourti
– Male Plp1 distal segment simple, with subterminal tooth . . . . . . . . . . . . . . . . . . . . . L. rafai
4 – In larger cheliped dorsal margins of dactylus and palm, anteroventral margin of palm,
and dorso- and ventrodistal angles of carpus, spinous . . . . . . . . . . . . . . . . . . L. grandidieri
– In larger cheliped dactylus, palm, and carpus without spines . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Male Plp1 distal segment simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. turneranus
– Male Plp1 distal segment simple, with subterminal tooth . . . . . . . . . . . . . . . . . L. sinuensis
Lepidophthalmus bocourti (A. Milne-Edwards, 1870)
(fig. 67A-B)
Callianassa bocourti A. Milne-Edwards, 1870: 95, 101. [Type locality. — San Jose del Cabo,
Lower California.]
Callianassa (Callichirus) Bocourti — Borradaile, 1903: 547; De Man, 1928b: 28, 94, 115.
Callianassa bocourti — De Saint Laurent & Le Loeuff, 1979: 96 [in partim = Lepidophthalmoides eiseni].
Lepidophthalmus bocourti — Manning & Felder, 1991: 778 (partim); Felder & Rodrigues,
1993: 373; Hendrickx, 1995: 390; Felder & Manning, 1997: 319; Felder & Manning, 1998:
398, 406; Nates & Felder, 1998: 188, 190, 194-196, 204 (partim); Sakai, 1999c: 70, fig. 14cd; Felder & Staton, 2000: 171, 179; Staton et al., 2000: 158, 167; Tudge et al., 2000: 144;
Felder, 2001: 442; Felder et al., 2002, fig. 3C, D; Felder, 2003: 436, figs. 20-29.
Lepidophthalmus bocourti aff. — Staton et al., 2000: 158, 167, fig. 2 [designated as Lepidophthalmoides aff. bocourti].
Lepidophthalmus bocourti — Manning & Felder, 1991: 778 [in partim = Lepidophthalmoides
eiseni]; Hendrickx, 1995: 390; Sakai, 1999c: 70 [partim], fig. 14c-d; Felder, 2001: 442;
Felder, 2003: 431, figs. 1-19; Sakai, 2005b: 149.
Not: Lepidophthalmus bocourti — Biffar, 1972, excluded as published lit. by ICZN Article 8.4.
Not: Lepidophthalmus bocourti — Lemaitre & Ramos, 1992: 349, fig. 4; Lemaitre & León,
1992: 44. [Not assigned to L. bocourti by Felder, 2003: 431.]
Material examined. — SMF 2186, 1 male, 1 female, Estero near La Playa de las Flores
near La Libhertad, El Salvador, 18.ix.1952, leg. O. Schuster; USNM 1002607, 1 male (TL/CL,
69.0/17.0 mm), Paso Caballo, Nicaragua, North Pacific Ocean, 11.iv.2000, leg. D.L. Felder and
S. Nates.
Diagnosis. — Mxp3 ischium-merus subsquare; propodus subtriangular, and
dactylus digitiform. Male Plp1 uniramous and bisegmented, distal segment
simple distally (fig. 67A); male Plp2 biramous, endopod with appendix masculina with distal setae and proximally-fused appendix interna mesiomedially
(fig. 67B) (cf. Sakai, 1999c: 70, fig. 14d); Plps3-5 endopods with small triangular appendix interna. Telson wider than long, subrectangular, and trilobate
on posterior margin.
Type locality. — San Jose del Cabo, Lower California.
448
CRM 013 – Katsushi Sakai
Fig. 67. Lepidophthalmus bocourti (A. Milne-Edwards, 1870), Lepidophthalmus sinuensis
Lemaitre & Rodrigues, 1991, Neocallichirus grandimanus (Gibbes, 1850), and Sergio guassutinga (Rodrigues, 1971). A, C, E, H, male Plp1; B, D, F, G, I, male Plp2. A, B, Lepidophthalmus bocourti, USNM 1002607, 1 male (TL/CL, 69.0/17.0 mm), Paso Caballo, Nicaragua,
North Pacific Ocean; C, D, Lepidophthalmus sinuensis, USNM 252403, paratype, male (TL/CL,
60.0/12.0 mm), on mouth of brackish creek draining to beach, bungalow beach near town of
San Bernardo del Ciento, 09◦ 17′ N 76◦ 99′ W, Caribbean coast of Colombia; E, F, G, Neocallichirus grandimanus, USNM 205630, neotype, male (TL/CL, 67.0/18.5 mm), S. Roosevelt
Blvd, Key West, Florida, 0.5 m; H, I, Sergio guassutinga, USNM 256886, paratype, male
(TL/CL, 100.6/26.6 mm), Praia do Araç, São Sebastião, São Paulo, Brazil. Scales, 1 mm.
Distribution. — Eastern Pacific, from San Jose del Cabo, Baja California,
Mexico, and El Salvador to Colombia; intertidal (Lemaitre & Ramos, 1992).
Lepidophthalmus grandidieri (Coutière, 1899)
Callianassa Grandidieri Coutière, 1899: 285, figs. 1-5; Tudge et al., 2000: 143.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
449
Callianassa (Callichirus) Grandidieri — Borradaile, 1903: 547; De Man, 1928b: 28, 92, 110.
Lepidophthalmus grandidieri — Sakai, 1999c: 71; Sakai, 2005b: 151.
Diagnosis. — Mxp3 and male Plps1-2 unknown. Telson slightly wider than
long, subrectangular, and rounded on posterior margin.
Type locality. — Northeast coast of Madagascar, River Mahanara.
Distribution. — Only known from the type locality.
Lepidophthalmus rafai Felder & Manning, 1998
Lepidophthalmus rafai Felder & Manning, 1998: 398, figs. 1-3; Tudge et al., 2000: 144; Sakai,
2005b: 150.
Type locality. — Beach at Playa Basura, Bahia de Buenaventura, Pacific
coast of Colombia.
Distribution. — Only known from the type locality.
Lepidophthalmus sinuensis Lemaitre & Rodrigues, 1991
(fig. 67C-D)
Lepidophthalmus sinuensis Lemaitre & Rodrigues, 1991: 623, figs. 1-4; Manning & Felder,
1991: 778; Nates et al., 1999: 526-541; Sakai, 1999c: 68; Tudge et al., 2000: 144; Sakai,
2005b: 148.
Material examined. — USNM 252403, paratype, male (TL/CL, 6.0/12.0 mm), on mouth
of brackish creek draining to beach, Bungalow beach near town of San Bernardo del Ciento,
09◦ 17′ N 76◦ 99′ W, Caribbean coast of Colombia, leg. Departamento de Córdoba.
Diagnosis. — Frontal margin of carapace with rostrum, and truncate tooth
on both sides of rostrum. Mxp3 ischium-merus rectangular; propodus subtriangular, and dactylus digitiform. Male Plp1 uniramous, trisegmented and simple,
with subdistal tooth (fig. 67C) (cf. Lemaitre & Rodrigues, 1991, fig. 4c); male
Plp2 biramous, endopod bearing distally appendix masculina with distal setae
and proximally-fused appendix interna (fig. 67D) (cf. Lemaitre & Rodrigues,
1991, fig. 4d).
Type locality. — Colombia, mouth of Rio Sinú, 9◦ 07′ N 75◦ 0′ W.
Distribution. — Caribbean coast of Colombia; intertidal, 1.5 m.
Lepidophthalmus turneranus (White, 1861)
[Abbreviated list of synonymy.]
Callianassa turnerana White, 1861a: 42, pl. 6; White, 1861b: 479; A. Milne-Edwards, 1870:
89, 101; Nobili, 1900: 3; Rathbun, 1900a: 308; Lenz, 1911: 316, figs. 1-11; Vanhöffen, 1911:
105, fig. a, d; Balss, 1916: 33; Monod, 1927: 595; Holthuis, 1991: 250, 246, fig. 455; Clark
& Presswell, 2001: 155.
450
CRM 013 – Katsushi Sakai
Callichirus turneranus — Le Loeuff & Intès, 1974: 40, fig. 10a-s; De Saint Laurent & Le
Loeuff, 1979: 64, figs. 14e, 19e, 20a-d, 23a-e.
Lepidophthalmus turneranus — Sakai, 1999c: 65; Sakai, 2005: 144.
Diagnosis. — Mxp3 ischium-merus rectangular; propodus subquadrate;
dactylus digitiform with obtuse tip. Male Plp1 uniramous and bisegmented,
and simple distally; male Plp2 biramous, endopod with fused appendix interna
(Le Loeuff & Intès, 1974, fig. 10r). Telson ovate, and trilobate on posterior
margin (Le Loeuff & Intès, 1974, fig. 10s).
Type locality. — Cameroon, fresh water.
Distribution. — Togo; Cameroon to Congo, W. Africa; lagoons and estuaries to practically fresh water.
Genus Michaelcallianassa Sakai, 2002
Michaelcallianassa Sakai, 2002: 480; Sakai, 2005b: 156.
Diagnosis. — Rostrum spinous. Carapace with dorsal oval, linea thalassinica entire. Eyestalks flattened, contiguous; cornea located dorsally. A1
peduncle slightly longer than A2 peduncle. Maxilla 2 scaphognathite without
elongate posterior whip. Mxp3 ischium-merus subpediform; propodus broadened, and rounded on ventral margin; dactylus digitiform; no exopod. P1 unequal and chelate. P2 chelate. P3 propodus distinctly protruded on posterior
side; dactylus trilobed on dorsal margin. P4 subchelate. P5 chelate. Abdominal
somites 4-5 bearing transverse row of setae; pleura 3-5 with rounded integumental patches. Male Plp1 uniramous and bisegmented; male Plp2 uniramous
and bisegmented, distal segment bilobed distally, lacking appendices interna
and masculina; male Plps3-5 biramous and foliaceous, endopods with embedded appendix interna with hooklets on mesial margin. Telson subquadrate,
wider than long and convergent in posterior two-thirds, and concave medially on posterior margin. Uropodal endopod broadly oval; uropodal exopod
distinctly longer than endopod and rounded distally, bearing dorsal plate.
[Adapted from Sakai, 2002: 480.]
Type species. — Michaelcallianassa indica Sakai, 2002, by original designation and monotypy. The gender of the generic name, Michaelcallianassa, is
feminine.
Species included. — Michaelcallianassa indica Sakai, 2002.
Michaelcallianassa indica Sakai, 2002
Michaelcallianassa indica Sakai, 2002: 481, figs. 11A-C, 12A-D, 13A-G, 14A-J; Sakai, 2005b:
157, fig. 33.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
451
Type locality. — Persian (Arabian) Gulf, 29◦ 29.897′ N 49◦ 54.108′ E, 32 m
depth.
Distribution. — Persian (= Arabian) Gulf (29◦ 29.897′ N 49◦ 54.108′ E;
32 m); Andaman Sea (7◦ 15.087′ N 99◦ 02.918′ E; 7◦ 00.082′ N 99◦ 15.660′ E –
9◦ 00.062′ N 97◦ 53.366′ E; 9◦ 30.991′ N 97◦ 57.706′ E; 17.0-65.4 m); mud, muddy
sand, and sand with shell fragments.
Genus Neocallichirus Sakai, 1988
Neocallichirus Sakai, 1988: 61; Manning & Felder, 1991: 779, figs. 1, 3, 4; Poore, 1994: 102;
Sakai, 1999c: 84; Sakai, 2000: 92; Davie, 2002: 461; Sakai, 2005b: 160.
Neocallichirus s. str. — Felder, 2001: 440.
Diagnosis. — Front of carapace with low rostrum and paired lateral,
subtriangular projections. Carapace with dorsal oval, lacking rostral carina,
cardiac prominence, and transverse cardiac sulcus(i). A1 peduncle shorter
than A2 peduncle. Mxp3 ischium-merus subpediform; propodus broadened,
rounded, or truncate on ventral margin; dactylus digitiform and obtuse distally;
exopod absent. P1 chelate, unequal, and dissimilar; in male larger cheliped
merus with or without ventral convexity. P2 chelate. P3 propodus broadened.
P4 subchelate. P5 chelate. Abdominal somites 3-5 not ornamented dorsally.
Male Plp1 uniramous and bisegmented, distal segment subchelate or wshaped distally; male Plp2 biramous, endopod bearing appendices interna
and masculina, or only appendix masculina; male Plps3-5 biramous and
foliaceous, endopods with appendix interna. Telson wider than long, and
varied in shape. Uropodal endopod wider than long, and ovate or subquadrate,
overreaching telson. Uropodal exopod convex on lateral margin, bearing dorsal
plate.
Type species. — Neocallichirus horneri Sakai, 1988 by original designation. The gender of the generic name, Neocallichirus, is masculine.
Species included. — N. angelikae Sakai, 2000; N. auchenorhynchus Sakai,
2005b; N. audax (De Man, 1911); N. branneri (Rathbun, 1900b) [= Glypturus
siguanensis Boone, 1927]; N. caechabitator Sakai, 1988; N. calmani (Nobili, 1904); N. darwinensis Sakai, 1988; N. denticulatus (Ngoc-Ho, 1994);
N. grandimanus (Gibbes, 1850); N. guaiqueri (Blanco Rambla & Liñero
Arana, 1994) [= G. sp., Blanco Rambla et al., 1995]; N. guarus (Rodrigues,
1971); N. horneri Sakai, 1988; N. indicus (De Man, 1905) [= Neocallichirus
taiaro Ngoc-Ho, 1995; Callianassa natalensis Barnard, 1946; Callianassa
(Cheramus) variabilis Edmondson, 1944]; N. jousseaumei (Nobili, 1904); N.
karumba (Poore & Griffin, 1979); N. kempi (Sakai, 1999c) [= C. maxima
452
CRM 013 – Katsushi Sakai
A. Milne-Edwards, 1870]; N. manningi Kazmi & Kazmi, 1992; N. mauritianus (Miers, 1882); N. mirim (Rodrigues, 1971); N. moluccensis (De Man,
1905); N. mortenseni (Sakai, 2005b); N. mucronatus (Strahl, 1862a); N. nickellae Manning, 1993; N. pentagonocephalus (Rossignol, 1962); N. rathbunae
(Schmitt, 1935b); N. sassandrensis (Le Loeuff & Intès, 1974); N. vigilax (De
Man, 1916); N. winslowi (Miers, 1884a); and N. sp. (Rathbun, 1906).
K EY
1–
–
2–
–
3–
–
4–
–
5–
–
6–
–
7–
–
8–
–
9–
–
10 –
–
11 –
–
12 –
–
13 –
–
14 –
–
15 –
–
16 –
–
17 –
–
18 –
–
TO THE
S PECIES
OF THE
G ENUS NEOCALLICHIRUS∗ )
Eyestalks located on necklike frontal margin of carapace . . . . . . . N. auchenorhynchus
Eyestalks located on frontal margin of carapace . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Frontal margin of carapace with spinous anterolateral projections . . . . . . . . . . . . . . . . . 3
Frontal margin of carapace with low or inconspicuous convexities . . . . . . . . . . . . . . . . . 8
Telson trapezoid. A1 peduncle almost reaching distal end of A2 penultimate segment
............................................................................ 4
Telson broadened proximally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
A1 peduncle almost reaching middle of A2 distal segment . . . . . . . . . . . N. denticulatus
A1 peduncle almost reaching distal end of A2 penultimate segment . . . . . . . . N. vigilax
Telson unarmed on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. rathbunae
Telson with spine on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Uropodal endopod fusiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. winslowi
Uropodal endopod trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Mxp3 ischium 1.8 times as long as wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .N. angelikae
Mxp3 ischium 1.2 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. mortenseni
Uropodal sympod and uropodal exopod, respectively, with sharp spine . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. sassandrensis
Uropodal sympod and uropodal exopod, respectively, without spine . . . . . . . . . . . . . . . 9
Uropodal endopod slenderly or broadly fusiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Uropodal endopod broadened distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Telson rounded on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Telson concave or w-shaped medially on posterior margin . . . . . . . . . . . . . . . . . . . . . . . 13
Mxp3 without exopod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. mucronatus
Mxp3 with rudimentary exopod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Telson rounded on lateral margins . . . . . . . . . . . . . . . . . . . . . . . . N. kempi [= C. maxima]
Telson parallel-sided proximally and tapering toward posterior margin . . . N. karumba
Telson w-shaped medially on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . N. mirim
Telson straight or concave on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Uropodal endopod broadly lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Uropodal endopod slenderly lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Denticulate meral tooth of male larger cheliped squarish . . . . . . . . . . . . . . . . . N. guarus
Denticulate meral tooth of male larger cheliped rounded . . . . . . . . . . . . . . . . . . N. audax
Male Plp1 w-shaped distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. pentagonocephalus
Male Plp1 chelate distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. guaiqueri
Uropodal endopod oval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Uropodal endopod trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Merus of larger cheliped with triangular meral hook . . . . . . . . . . . . . . . . . N. mauritianus
Merus of larger cheliped denticulate proximally on ventral margin . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. caechabitator
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
453
19 – Eyestalks denticulate distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
– Eyestalks not denticulate distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
20 – Telson semicircular . . . . . N. indicus [= Neocallichirus taiaro; Callianassa natalensis;
Callianassa (Cheramus) variabilis]
– Telson wider than long and trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. manningi
21 – Eyestalks separated from each other and triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
– Eyestalks contiguous and triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22 – Eyestalks long and pointed apically, with cornea centrally . . . . . . . . . . . N. jousseaumei
– Eyestalks short, with cornea mediolaterally. . . . . . . . . . . . . . . . . . . . . . . . .N. moluccensis
23 – Eyestalks deformed distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
– Eyestalks simple distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
24 – Eyestalks truncate distally, and cornea small . . . . . . . . . . . . . . . . . . . . . . . . N. darwinensis
– Eyestalks hook-like distally, and cornea distinct . . . . . . . . . . . . . . . . . . . . N. grandimanus
25 – Cutting edge of dactylus of larger cheliped with distinct proximal protrusion . . . . . . 26
– Cutting edge of dactylus of larger cheliped without distinct proximal protrusion . . . 27
26 – Ischium of larger cheliped with denticulate protrusion on ventral margin . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. calmani
– Ischium of larger cheliped without denticulate protrusion on ventral margin . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. nickellae
27 – In female larger cheliped dactylus with triangular median concavity on cutting edge .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. horneri
– In female larger cheliped dactylus smooth on cutting edge . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. branneri [= G. siguanensis]
∗ ) Callianassa sp. (Rathbun, 1906) not listed in this key.
Neocallichirus angelikae Sakai, 2000
Neocallichirus angelikae Sakai, 2000: 91, figs. 1-3; Davie, 2002: 461; Sakai, 2005b: 176.
Diagnosis. — Rostrum inconspicuous. Frontal margin of carapace with
spinous anterolateral projections. Mxp3 ischium-merus subrectangular; merus
obliquely declined distally and rounded on distomesial margin; propodus subquadrate, and dactylus digitiform. Male Plp1 blade-shaped, uniramous, and
bisegmented, distal segment slightly bilobed distally; Plp2 endopod bearing
distally stout appendix masculina provided mesiomedially with minute appendix interna with hooklets. Telson subrectangular, posterior margin narrow and
straight, with a median spine. Uropodal endopod trapezoid. [Cf. Sakai, 2000:
91, figs. 1E, 2A, 3G, H, I.]
Type locality. — Murat Bay, Great Australian Bight, South Australia, stony
flat (32◦ 07′ S 133◦ 40′ E).
Distribution. — Only known from the type locality.
Neocallichirus auchenorhynchus Sakai, 2005
Neocallichirus auchenorhynchus Sakai, 2005b: 184, fig. 37.
Type locality. — Unknown.
454
CRM 013 – Katsushi Sakai
Neocallichirus audax (De Man, 1911)
Callianassa audax De Man, 1911: 223; Dworschak, 1992: 190, fig. 1a-d; Tudge et al., 2000:
143.
Callianassa (Callichirus) audax — De Man, 1928b: 1, 18, 28, 113, 179, pl. 20 fig. 31-31i; Rao
& Kartha, 1967: 279, figs. 1, 2; ?Tirmizi, 1967: 151-154, figs. 1, 2.
Callichirus audax — De Saint Laurent & Le Loeuff, 1979: 97.
Neocallichirus audax — Sakai, 1999c: 95, fig. 21d-f; Sakai, 2005b: 176.
Type locality. — Straits of Malacca.
Distribution. — Straits of Malacca, Malay Peninsula; West Pakistan; Ratnagiri, India; east coast of India.
Neocallichirus branneri (Rathbun, 1900)
Glypturus branneri Rathbun, 1900b: 150, pl. 8 figs. 5-8; Rathbun, 1901: 93; Rathbun, 1920:
328, fig. 3; Verrill, 1922: 33, pl. 1 fig. 2, pl. 8 fig. 1a-e; Schmitt, 1924: 93; Schmitt, 1935a:
194, fig. 55; Manning, 1987: 397.
Callianassa branneri — Schmitt, 1935b: 4; Gurney, 1944: 82, figs. 16, 17; Weimer & Hoyt,
1964: 764; Biffar, 1971a: 652, 654, 661, 661-667, figs. 5, 6; Coelho & Ramos, 1973: 161;
Manning, 1987: 398.
Neocallichirus branneri — Rodrigues et al., 1998: 380; Melo, 1999: 374, figs. 249-250.
Remarks. — The present species, Neocallichirus branneri was synonymized
with Neocallichirus grandimana (Gibbes, 1890) in previous papers (Sakai,
1999c, 2005b). However, it has turned out that N. branneri is not conspecific
with N. grandimana, because in N. branneri the A1 peduncle barely reaches
the middle of the A2 distal segment, in the male P1 the cutting edge of the
dactylus is triangularly concave at its distal third, and the cutting edge of the
fixed finger bears a low median tooth; whereas in N. grandimana the A1 peduncle reaches the distal third of the A2 distal segment, in the male P1 the
cutting edge of the dactylus is distinctly concave submedially, and the cutting
edge of the fixed finger is unarmed (cf. Manning & Felder, 1991, fig. 2), so
that those two species are considered valid.
Type locality. — Mamanguape Stone Reef, Brazil.
Distribution. — Only known from the type locality.
Neocallichirus caechabitator Sakai, 1988
Neocallichirus caechabitator Sakai, 1988: 67, figs. 9, 10; Sakai, 1999c: 96; Tudge et al., 2000:
144; Davie, 2002: 461; Sakai, 2005b: 177.
Diagnosis. — Rostrum small. Eyestalks contiguous and triangular in distal
third; cornea small. Frontal margin of carapace with low convexities anterolaterally. Mxp3 ischium-merus rectangular; merus obliquely declined distally
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
455
and rounded on mesiodistal margin; propodus subquadrate, and dactylus digitiform. Merus of larger cheliped denticulate proximally on ventral margin.
Male Plps1-2 unknown. Telson suboval, bearing no median spine but median
concavity on short and straight posterior margin. Uropodal endopod broadened
distally. [Adapted from Sakai, 1988: 67, figs. 9E, 10A.]
Type locality. — Shoal Bay, False Creek Point, Darwin, Northern Territory,
Australia, 1.0 m.
Distribution. — Only known from the type locality.
Neocallichirus calmani (Nobili, 1904)
Callianassa (Cheramus) Calmani Nobili, 1904: 236; Nobili, 1906b: 104, pl. 5 fig. 2; De Man,
1928b: 26, 100.
Callichirus calmani — De Saint Laurent & Le Loeuff, 1979: 97.
Callianassa calmani — Dworschak, 1992: 192, fig. 3a-f; Tudge et al., 2000: 143.
Neocallichirus calmani — Sakai, 1999c: 96, fig. 22a-d; Sakai, 2005b: 177; Robles et al., 2009:
317.
Type locality. — Obock, Djibouti.
Distribution. — Obock, Djibouti, Bay of Aden; in sandy intertidal and
shallow subtidal; Aqaba, Red Sea.
Neocallichirus darwinensis Sakai, 1988
Neocallichirus darwinensis Sakai, 1988: 62, figs. 5, 6; Sakai, 1999c: 98; Tudge et al., 2000:
144; Davie, 2002: 461; Sakai, 2005b: 177.
Diagnosis. — Rostrum small. Eyestalks contiguous and triangular with
truncate apex; cornea small. Frontal margin of carapace with low convexities
anterolaterally. Mxp3 ischium-merus rectangular; merus obliquely declined
distally and convex on distomesial margin; propodus subquadrate, and dactylus digitiform. Male Plps1-2 unknown. Telson shaped as inverted trapezoid,
bearing no median spine on posterior margin. Uropodal endopod trapezoid.
[Adapted from Sakai, 1988, figs. 5G, 6G.]
Type locality. — Mindil Beach, Darwin, Northern Territory, Australia.
Distribution. — Only known from the type locality.
Neocallichirus denticulatus Ngoc-Ho, 1994
Neocallichirus denticulatus Ngoc-Ho, 1994: 56, fig. 4; Sakai, 1999c: 98; Tudge et al., 2000:
144; Davie, 2002: 461; Sakai, 2005b: 178; Robles et al., 2009: 317.
Diagnosis. — Rostrum triangular, with acute apex, reaching middle of eyestalks; cornea distinct. Frontal margin of carapace with anterolateral spinous
456
CRM 013 – Katsushi Sakai
projections. A1 peduncle almost reaching middle of A2 distal segment. Mxp3
ischium-merus subrectangular; merus obliquely declined distally and convex
on distomesial margin. Male Plps1-2 unknown. Telson shaped as inverted
trapezoid with mediolateral convexity, lacking median spine on slightly convex
posterior margin. [Adapted from Ngoc-Ho, 1994: 56, fig. 4h, m.]
Type locality. — Queensland, N.W. of Townsville (18◦ 56′ S 146◦ 50′ E),
24 m, muddy sand.
Distribution. — Only known from the type locality.
Neocallichirus grandimanus (Gibbes, 1850)
(fig. 67E-G)
Callianassa grandimana Gibbes, 1850: 194; Stimpson, 1866: 47; Stimpson, 1871: 122;
Kingsley, 1899: 823; De Man, 1928b: 19; Schmitt, 1935b: 2; Biffar, 1971a: 649, 671-674;
Manning, 1987: 388, 397, fig. 2; Dworschak, 1992: 196.
Glypturus grandimanus — Rathbun, 1900b: 151.
Glypturus grandimana — Borradaile, 1903: 548; De Man, 1928b: 25.
Glypturus siguanensis Boone, 1927: 85, fig. 17; Manning, 1987: 397. [Type locality. — Siguana
Bay, Isle of Pines near Cuba, Gulf of Mexico.]
Callianassa siguanensis — Biffar, 1971a: 649.
Neocallichirus grandimana — Sakai, 1988: 61; Manning & Felder, 1991: 779, figs. 3, 4;
Lemaitre & León, 1992: 44; Lemaitre & Ramos, 1992: 349, fig. 5; Dworschak & Ott, 1993:
281; Felder & Manning, 1995: 478, figs. 1d-f, 3f, 4e, g-h; Coelho, 1997: 148; Melo, 1999:
376, figs. 251-252; Blanco-Rambla, 2000: 73, fig. 2; Dworschak, 2004: 15, fig. 4; Coelho et
al., 2007: 5 (table).
Neocallichirus grandimanus — Manning, 1993: 113; Sakai, 1999c: 89; Tudge et al., 2000: 144;
Sakai, 2005b: 164.
Not: Glypturus grandimanus — Balss, 1924: 179, figs. 3, 4; Manning, 1987: 399. [= C. hartmeyeri Schmitt.]
Material examined. — USNM 205630, neotype, male (TL/CL, 67.0/18.5 mm), S. Roosevelt
Blvd, Key West, Florida, 0.5 m, 08.viii.1986, leg. et det. R. Manning.
Diagnosis. — Mxp3 propodus subquadrate, and dactylus digitiform. Male
Plp1 (fig. 67E) uniramous and bisegmented, distal segment w-shaped distally
(cf. Felder & Manning, 1995, fig. 1f); male Plp2 (fig. 67F, G) biramous,
endopod bearing mesiodistally appendix masculina with distal setae and small
conical appendix interna (cf. Felder & Manning, 1995, fig. 4e-f).
Type locality. — Florida, Key West.
Distribution. — Atlantic side: Bermuda; southeast of Florida; west coast of
Florida, including Keys and Dry Tortugas; Tobago; Venezuela; Bimini, Little
San Salvador, Bahamas; Cuba; Puerto Rico; Barbados; Curaçao; South Water
Cay (lagoon side, intertidal to 0.5 m), and Belize; Mamanguape and Pernambuco, Brazil. Pacific side: Panama, Ecuador; Gorgona Island, Colombia.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
457
Neocallichirus guaiqueri (Blanco Rambla & Liñero Arana, 1995)
Neocallichirus sp. Blanco Rambla & Liñero Arana, 1994: 20-22, figs. 4, 5.
Sergio guaiqueri — Blanco Rambla et al., 1995: 102, text-figs. 1-3.
Sergio guaiqueri — Lemaitre & Felder, 1996: 453; Tudge et al., 2000: 144.
Neocallichirus guaiqueri — Sakai, 1999c: 90; Sakai, 2005b: 165.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subovate,
and dactylus digitiform. Male Plp1 uniramous, bisegmented, and subchelate
distally; male Plp2 biramous, and endopod bilobed distally, but uncertain
about appendices interna and masculina. Telson wider than long and trapezoid,
and concave medially on posterior margin, without median spine. Uropodal
endopod longer than wide and slenderly lanceolate. [Cf. Blanco Rambla &
Liñero Arana, 1994: 20-22, figs. 4, 5.]
Type locality. — Venezuela, Anzoategui State, north of Jose, Petersen grab
(10◦ 08.40′ N 64◦ 50.10′ W).
Distribution. — Venezuela.
Neocallichirus guarus (Rodrigues, 1971)
Callianassa (Callichirus) guara Rodrigues, 1971: 210, figs. 61-76; Biffar, 1971a: 654; Coelho
& Ramos, 1973: 162; Manning, 1987: 397.
Neocallichirus guarua — Manning & Felder, 1991: 779.
Sergio guara — Manning & Lemaitre, 1994: 41; Lemaitre & Felder, 1996: 453; Coelho, 1997:
149; Rodrigues et al., 1998: 381; Melo, 1999: 386, figs. 259-260; Tudge et al., 2000: 144;
Coelho et al., 2007: 5 (tab. 2).
Neocallichirus guarus — Sakai, 1999c: 90; Sakai, 2005b: 166.
Material examined. — USNM 256885, male (CL 23.0; abdomen + telson, 50.0 mm), Praia
do Araça, São Sebastião, São Paulo, Brazil, vii.1966, leg. S.A. Rodrigues.
Diagnosis. — Mxp3 ischium-merus subrectangular; merus subtriangular;
propodus subquadrate, and dactylus digitiform. Male Plp1 uniramous and
bisegmented, distal segment subchelate distally; male Plp2 biramous, endopod
bearing distally proximally-fused appendix masculina with proximally-fused
appendix interna distolaterally. Telson wider than long and trapezoid, and concave medially on posterior margin, without median spine. Uropodal endopod
broadly lanceolate. [Cf. Rodrigues, 1971: 210, figs. 67, 74-76.]
Type locality. — Brazil, São Sebastião, São Paulo, beach.
Distribution. — Tampa Bay, Miami; Lemon Bay, Florida; Pará, Maranháo,
Ceará, and Pernambuco, Brazil.
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CRM 013 – Katsushi Sakai
Neocallichirus horneri Sakai, 1988
Neocallichirus horneri Sakai, 1988: 65, figs. 7, 8; Manning & Felder, 1991: 779; Sakai, 1999c:
99; Tudge et al., 2000: 144; Davie, 2002: 461; Sakai, 2005b: 178.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subquadrate,
and dactylus digitiform. Male Plp1 uniramous and bisegmented, distal segment
w-shaped distally; male Plp2 biramous, endopod bearing distally proximallyfused appendix masculina with distal setae, and proximally-fused appendix
interna distolaterally. Telson wider than long and trapezoid, and convex on
posterior margin, without median spine. [Cf. Sakai, 1988: 65, figs. 7F, 8A, H,
I.]
Type locality. — Australia, Northern Territory, Darwin, Nightcliff.
Distribution. — Only known from the type locality.
Neocallichirus indicus (De Man, 1905)
Callianassa (Cheramus) indica De Man, 1905: 605; De Man, 1928b: 26, 93, 100, 159, 160,
pl. 17 fig. 26-26g.
Callianassa (Cheramus) variabilis Edmondson, 1944: 47, figs. 1, 6a-i, 7a-j, l, p. [Type locality.
— Hanauma Bay, Oahu, in a gravel bed of the intertidal zone.]
Callianassa natalensis Barnard, 1946: 379; Barnard, 1950: 506, 511, fig. 95f-h; Kensley, 1974:
277. [Type locality. — Zululand, South Africa.]
Callianassa indica — Kensley, 1975: 50, fig. 2A-E; Sakai, 1987a: 302, 306.
Neocallichirus taiaro Ngoc-Ho, 1995: 212, figs. 1, 2; Tudge et al., 2000: 144. [Type locality. —
Taiaro atoll, Tuamotu Isle, French Polynesia.]
Neocallichirus indicus — Sakai, 1999c: 99, fig. 23a-e; Tudge et al., 2000: 144; Sakai & Apel,
2002: 277, fig. 2; Sakai, 2005b: 178; Robles et al., 2009: 317.
Callianassa variabilis — Tudge et al., 2000: 143.
Neocallichirus natalensis — Tudge et al., 2000: 144.
Type locality. — Indonesia, south coast of Kangeang, Bay of Kankamaran
(6 59′ S 115◦ 24′ 7E).
Distribution. — Hanauma Bay, Oahu, Hawaii; Tuamotu Isle, French Polynesia; Tonaki Is., Ryukyu Is.; Flores, Indonesia; Mauritius; Kangeang Reef,
Bay of Kankamaran, East Indies; Sandspit, Karachi; Djibouti, Gulf of Aden;
Red Sea; Zululand, South Africa.
◦
Neocallichirus jousseaumei (Nobili, 1904)
Callianassa (Cheramus) Jousseaumei Nobili, 1904: 236; Nobili, 1906b: 101, pl. 6 fig. 2; De
Man, 1928b: 26 (list), 100 (key), pl. 18 fig. 27-27a.
Callichirus jousseaumei — De Saint Laurent & Le Loeuff, 1979: 97.
Callianassa jousseaumei — Dworschak, 1992: 198, figs. 5a-d, 6a-c.
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459
Neocallichirus jousseaumei — Sakai, 1999c: 100, fig. 22e-g; Tudge et al., 2000: 144; Sakai,
2005b: 179.
Diagnosis. — Rostrum inconspicuous. Eyestalks triangular and separated
from each other; cornea located medially. Frontal margin of carapace with low
convexities anterolaterally. Mxp3 ischium-merus rectangular, merus obliquely
declined distally and convex on distomesial margin; propodus subovate, and
dactylus digitiform. Male Plp1 uniramous, bisegmented, and subchelate distally; male Plp2 biramous, endopod simple, bearing appendix masculina distally, but no appendix interna. Telson shaped as inverted trapezoid, bearing no
median spine on slightly convex posterior margin. Uropodal endopod trapezoid. [Adapted from De Man, 1928b, pl. 18 fig. 27a, c; Sakai, 1999c: 100,
fig. 22f, g.]
Type locality. — Djibouti, Perim, Gulf of Tadjourah, Red Sea.
Distribution. — Djibouti and Perim, Gulf of Aden; Safaga, Tubaya AlKabir, Red Sea; Aqaba, Gulf of Aqaba, 9 m.
Neocallichirus karumba (Poore & Griffin, 1979)
Callianassa karumba Poore & Griffin, 1979: 266, figs. 30, 31.
Glypturus karumba — Sakai, 1988: 61; Tudge et al., 2000: 144; Davie, 2002: 460.
Neocallichirus karumba — Sakai, 1999c: 101; Sakai, 2005b: 180; Dworschak, 2008: 75
(partim), fig. 1 (not figs. 2-6 = Neocallichirus kempi Sakai, 1999c); Robles et al., 2009:
317.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subtriangular, and dactylus digitiform; exopod small. Male Plp1 uniramous and
bisegmented, and w-shaped distally; male Plp2 biramous, endopod bearing
mesiodistally appendix masculina with small, proximally-fused appendix interna mesiodistally. Telson wider than long and trapezoid, and slightly rounded
on posterior margin, without median spine. [Cf. Poore & Griffin, 1979: 266,
figs. 30c, 31f, g, h; Dworschak, 2008, fig. 1i.]
Type locality. — Norman River, Karumba, Queensland, Australia.
Distribution. — Only known from the type locality.
Neocallichirus kempi Sakai, 1999
Callianassa (Callichirus) maxima — Kemp, 1915: 252, pl. 13 figs. 1-5; De Man, 1928b: 29,
92, 112. [= Callianassa maxima A. Milne-Edwards, 1870.]
Callianassa maxima — Pillai, 1954: 23-26; Daniel, 1981: 193, pl. 6 figs. 1, 2 (larvae); Tudge
et al., 2000: 143.
Neocallichirus kempi Sakai, 1999c: 101, fig. 24a-e; Sakai, 2005b: 180.
460
CRM 013 – Katsushi Sakai
Neocallichirus karumba — Dworschak, 2008: 75 (partim), figs. 2-6.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subquadrate,
and dactylus digitiform; exopod small. Male Plp1 uniramous and bisegmented,
distal segment subchelate distally (cf. Dworschak, 2008, figs. 3h, 4d, 4k,
5h); male Plp2 biramous, endopod bearing mesiodistally appendix masculina
with small, proximally-fused appendix interna with hooklets mesiomedially
(cf. Dworschak, 2008, figs. 3i, 3p, 4e). Telson wider than long and suboval,
rounded laterally, and convex medially on posterior margin, without median
spine. [Cf. Kemp, 1915: 252, pl. 13 fig. 5; Sakai, 1999c: 101, fig. 24b, c;
Dworschak, 2008, fig. 3a.]
Remarks. — Dworschak (2008) regarded the present species Neocallichirus
kempi Sakai, 1999c as a junior synonym of Neocallichirus karumba (Poore &
Griffin, 1979). However, those two species are different, because in N. kempi
the male Plp1 distal segment is subchelate distally, and the male Plp2 endopod
bears mesiodistally an appendix masculina with a small, proximally-fused appendix interna with hooklets mesiomedially, and the telson is suboval; whereas
in G. karumba the male Plp1 distal segment is w-shaped distally (Poore &
Griffin, 1979, fig. 31g), and the male Plp2 endopod bears mesiodistally an appendix masculina with a small appendix interna with hooklets mesiodistally
(Poore & Griffin, 1979, fig. 31h; Dworschak, 2008, fig. 1j), and the telson is
trapezoid (Poore & Griffin, 1979, fig. 28c; Dworschak, 2008, fig. 1a).
Type locality. — Bangka, Indonesia.
Distribution. — Papua New Guinea (Dworschak, 2008); Bangka, Indonesia; Nalbano Island and Barhampur Island, Chilka Lake near Madras, India; Kayamkulam Lake, central Travancore, India (Pillai, 1954); Singapore
(Dworschak, 2008); Taiwan (Dworschak, 2008).
Neocallichirus manningi Kazmi & Kazmi, 1992
Neocallichirus manningi Kazmi & Kazmi, 1992: 296, fig. 1; Tudge et al., 2000: 144; Sakai,
2005b: 180-181.
Neocallichirus indicus — Sakai, 1999c: 99; Sakai, 2000: 277.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subquadrate,
and dactylus digitiform. Male Plps1-2 unknown. Telson trapezoid, and convex
on posterior margin, without median spine. [Cf. Kazmi & Kazmi, 1992: 296,
fig. 1D, C.]
Type locality. — Sandspit, Karachi, 24◦ 48′ N 66◦ 58′ E, lower tidal region.
Distribution. — Only known from the type locality.
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Neocallichirus mauritianus (Miers, 1882)
Callianassa mauritiana Miers, 1882: 341; Miers, 1884a: 15, pl. 1 fig. 2; Nobili, 1906b: 106,
figs. 5, 6; Michel, 1974: 256; Kensley, 1975: 51, fig. 3A-H; Tudge et al., 2000: 143.
Callianassa (Trypaea) mauritiana — Borradaile, 1903: 546.
Callianassa (Cheramus) mauritiana — De Man, 1928a: 10, pl. 2 fig. 4; De Man, 1928b: 26,
99, 160.
Neocallichirus mauritianus — Sakai, 1999c: 103, fig. 21a-c; Sakai, 2005b: 181.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subovate, and
dactylus digitiform. Male Plp1 uniramous, bisegmented, and subchelate distally; male Plp2 endopod biramous, endopod bearing mesiodistally embedded
appendix masculina, and distolaterally an embedded appendix interna with
hooklets. Telson subovate and convex on posterior margin, with median spine.
[Cf. Kensley, 1975: 51, fig. 3B, C, H; Sakai, 1999c: 103, fig. 21b, c.]
Type locality. — Mauritius.
Distribution. — Mauritius; Red Sea.
Neocallichirus mirim (Rodrigues, 1971)
Callianassa (Callichirus) mirim Rodrigues, 1971: 214, figs. 77-98.
Callianassa mirim — Biffar, 1971a: 654; Coelho & Ramos, 1973: 162; Manning, 1987: 397;
Dworschak, 1992: 202.
Callichirus mirim — Ferrari, 1981: 12, fig. 1; Rodrigues, 1983: 31, figs. 53-60; Rodrigues,
1984a: 239, figs. 1-39 [larval stage]; Rodrigues, 1984b: 914 [larval stage]; Rodrigues &
Hödl, 1990: 50, fig. 1.
Neocallichirus mirim — Manning & Felder, 1991: 779.
Sergio mirim — Manning & Lemaitre, 1994: 41; Rodrigues et al., 1998: 381; Melo, 1999: 388,
figs. 261-262; Tudge et al., 2000: 144; Coelho et al., 2007: 5 (tab. 2).
Neocallichirus mirim — Sakai, 1999c: 91, fig. 20d; Sakai, 2005b: 167.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subquadrate,
and dactylus digitiform. Male Plp1 uniramous, bisegmented, and w-shaped
distally; male Plp2 bearing distally proximally-fused appendix masculina with
setae, and proximally-fused appendix interna mesioproximally. Telson wider
than long, convex laterally, and curving toward rounded posterior margin with
median concavity with sharp spine. [Cf. Rodrigues, 1971: 214, figs. 84, 94, 95,
98.]
Type locality. — São Paulo, São Sebastião, Brazil.
Distribution. — From Santos, State of São Paulo, Brazil, to Argentina;
common in the lower intertidal and shallow subtidal zone of sandy beaches.
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CRM 013 – Katsushi Sakai
Neocallichirus moluccensis (De Man, 1905)
Callianassa (Cheramus) moluccensis De Man, 1905: 606; De Man, 1928b: 26, 93, 99, 159,
pl. 16 fig. 25-25a, pl. 17 fig. 25b-c.
Neocallichirus moluccensis — Sakai, 1999c: 104, fig. 25a-f; Tudge et al., 2000: 144; Sakai,
2005b: 181.
Diagnosis. — Rostrum inconspicuous. Eyestalks short, triangular, with truncate apex, and separated from each other; cornea distinct and located mediolaterally. Frontal margin of carapace with low convexities anterolaterally. Mxp3
ischium-merus rectangular; merus obliquely declined distally and convex on
mesiodistal margin; propodus subquadrate, and dactylus digitiform. Male Plp1
uniramous, bisegmented, and subchelate distally; male Plp2 biramous, endopod bearing distally appendix masculina with small appendix interna with distal hooklets. Telson shaped as inverted trapezoid, bearing no median spine on
posterior margin. [Cf. De Man, 1928b, pl. 17 fig. 25b-c; Sakai, 1999c: 104,
fig. 25c, d, e.]
Type locality. — Amboina, reef.
Distribution. — Amboina [= Ambon], Indonesia.
Neocallichirus mortenseni Sakai, 2005
Neocallichirus mortenseni Sakai, 2005b: 170, figs. 34-36.
Diagnosis. — Rostrum triangular, with acute apex. Eyestalks triangular in
distal third. Frontal margin of carapace with anterolateral spinous projections.
A1 peduncle slightly shorter than A2 peduncle. Mxp3 ischium-merus rectangular; propodus subquadrate, and dactylus digitiform. Male Plps1-2 unknown.
Telson shaped as inverted trapezoid, bearing median spine on posterior margin.
Uropodal endopod trapezoid and armed with spinule on lateral margin.
Type locality. — Off N. coast of Taboga, Panama.
Distribution. — Only known from the type locality.
Neocallichirus mucronatus (Strahl, 1862)
[Abbreviated list of synonymy.]
Callianassa mucronata Strahl, 1862a: 1056; Strahl, 1862b: 383; Tirmizi, 1977: 21 (part.),
figs. 1-3 (not fig. 1b = Callianassa gruneri Sakai, 1999c); Poore & Griffin, 1979: 273,
figs. 34, 35; Dworschak, 1992: 202, fig. 9a-f.
Callianassa brevicaudata A. Milne-Edwards, 1870: 91, 101, pl. 2 figs. 2, 2a, 2b; Tudge et al.,
2000: 143. [Type locality. — Zanzibar.]
Callianassa (Cheramus) novaeguineae Thallwitz, 1891: 31, pl. 1 fig. 9. [Type locality. — Papua
New Guinea.]
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463
Callianassa (Callichirus) mucronata — Borradaile, 1903: 547; Nobili, 1906b: 108; De Man,
1928b: 112, 175, pl. 19 fig. 30-30e.
Glypturus mucronatus — Sakai, 1988: 61; Tudge et al., 2000: 144.
Neocallichirus mucronatus — Sakai, 1999c: 105, fig. 26a-I; Sakai & Apel, 2002: 278; Sakai,
2005b: 182; Robles et al., 2009: 317.
Glypturus novaeguineae — Davie, 2002: 461.
Diagnosis. — Rostrum sharply triangular, reaching middle of eyestalks.
Eyestalks triangular in distal third. Frontal margin of carapace with low
convexities anterolaterally. Mxp3 ischium-merus oblong, merus obliquely
declined distally and convex on distomesial margin; propodus subovate, and
dactylus digitiform. Male Plp1 uniramous, bisegmented, and subchelate distally; male Plp2 biramous, endopod bearing distally fused appendix masculina,
but uncertain about appendix interna. Telson shaped as inverted trapezoid,
bearing no median spine on slightly convex posterior margin. Uropodal endopod fusiform. [Adapted from Sakai, 1999c, 2005b.]
Type locality. — Luzon, Philippines.
Distribution. — Papua New Guinea; Queensland, Australia, central reef;
Luzon, Philippines; Maldives; Amboina; N. Ceram; northwest coast of New
Guinea; Siau Island and Karakelong Island, Indonesia; Djibouti and Perim,
Gulf of Aden; Zanzibar; Madagascar, Baie de Pasandava.
Neocallichirus nickellae Manning, 1993
Neocallichirus nickellae Manning, 1993: 110-113, figs. 4-6; Sakai, 1999c: 92; Blanco Rambla,
2000: 75, fig. 4; Tudge et al., 2000: 144; Sakai, 2005b: 168.
Type locality. — Republic of Trinidad and Tobago, Tobago, Buccoo Reef,
Coral Garden (11◦ 11′ N 60◦ 49′ W).
Distribution. — Only known from the type locality.
Neocallichirus pentagonocephalus (Rossignol, 1962)
Callianassa pentagonocephala Rossignol, 1962: 139, fig. 1a-c.
Callichirus pentagonocephala — De Saint Laurent & Le Loeuff, 1979: 78, figs. 14k, 16b, 18d,
19i, 21d, 23w-x; Tudge et al., 2000: 144.
Neocallichirus pentagonocephala — Sakai, 1999c: 87.
Neocallichirus pentagonocephalus — Sakai, 2005b: 163.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subquadrate,
and dactylus digitiform. Male Plp1 uniramous, bisegmented, and shallowly
w-shaped distally; male Plp2 biramous, endopod bearing distally proximallyfused appendices masculina and interna. Telson hexagonal with shallow
464
CRM 013 – Katsushi Sakai
median concavity without spine on posterior margin. [Cf. De Saint Laurent
& Le Loeuff, 1979: 78, figs. 18d, 19i, 23w-x.]
Type locality. — Congo, Bay of Pointe Noire.
Distribution. — Only known from the type locality.
Neocallichirus rathbunae (Schmitt, 1935)
Callianassa (Callichirus) rathbunae Schmitt, 1935b: 4, 15, pl. 1 fig. 5, pl. 2 fig. 2, pl. 3 fig. 1,
pl. 4 fig. 2.
Callianassa rathbunae — Biffar, 1971a: 699-704, figs. 19, 20; Suchanek, 1983: 281; Manning
& Heard, 1986: 347-349, fig. 1; Manning, 1987: 397; Dworschak, 1992: 202, fig. 10.
Callichirus rathbunae — De Saint Laurent & Le Loeuff, 1979: 97.
Neocallichirus rathbunae — Manning & Felder, 1991: 779; Manning, 1993: 113; Felder &
Manning, 1995: 487; Coelho, 1997: 149; Melo, 1999: 378, figs. 253-254; Sakai, 1999c: 92;
Abed-Navandi, 2000: 294; Tudge et al., 2000: 144; Sakai, 2005b: 168.
Neocallichirus raymanningi Blanco Rambla & Lemaitre, 1999: 768, 4 figs. [Type locality. —
Playa Cristal, south coast of Gulf of Cariaco, Sucre State, Venezuela.]
Neocallichirus maryae Karasawa, 2004: 87; Coelho et al., 2007: 5 (table); Robles et al., 2009:
317. [Replacement name for Neocallichirus rathbunae (Schmitt, 1935b).]
Type locality. — Bluefields, Jamaica.
Distribution. — Indian River, Miami, Florida; west coast of Florida; Bimini,
Bahamas; Jamaica; St. Croix, Virgin Islands; Twin Cays, Cassiopeia Cove,
Belize; N. of Santa Marta, Colombia; Venezuela; Pernambuco, Brazil.
Neocallichirus sassandrensis (Le Loeuff & Intès, 1974)
Callichirus sassandrensis Le Loeuff & Intès, 1974: 43, fig. 11a-t; De Saint Laurent & Le Loeuff,
1979: 71, figs. 14i, 18c, 19i.
Neocallichirus sassandrensis — Sakai, 1999c: 87; Tudge et al., 2000: 144; Sakai, 2005b: 164.
Diagnosis. — Mxp3 ischium-merus subrectangular; propodus subquadrate,
and dactylus digitiform. Male Plps1-2 unknown. Telson trapezoid with straight
posterior margin, bearing median spine. Uropodal protopod and uropodal
exopod, respectively, with spine. [Cf. Le Loeuff & Intès, 1974: 43, fig. 11k,
q-t.]
Type locality. — Ivory Coast, Sassandra (4◦ 58.8′ N 6◦ 01′ W); 10 m.
Distribution. — Only known from the type locality.
Neocallichirus vigilax (De Man, 1916)
Callianassa (Callichirus) vigilax De Man, 1916: 57, pl. 1 figs. 1-6; De Man, 1928b: 30, 93,
109.
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Neocallichirus vigilax — Sakai, 1999c: 108; Sakai, 2005b: 183.
Diagnosis. — Rostrum triangular, reaching middle of eyestalks. Eyestalks
contiguous and obtuse apically. A1 peduncle almost reaching distal end of A2
penultimate segment. Frontal margin of carapace with anterolateral spinous
projections. Mxp3 ischium-merus rectangular; merus obliquely declined distally and rounded on mesiodistal margin; propodus subquadrate, and dactylus
digitiform. Male Plps1-2 unknown. Telson shaped as inverted trapezoid, bearing no median spine on slightly convex posterior margin. [Adapted from De
Man, 1916: 57, pl. 1 figs. 1-4.]
Type locality. — Ambon, Maluku Is., Indonesia.
Distribution. — Only known from the type locality.
Neocallichirus winslowi (Edmondson, 1944)
Callianassa (Callichirus) winslowi Edmondson, 1944: 59, fig. 11a-g.
Glypturus winslowi — Sakai, 1999c: 84.
Callianassa winslowi — Tudge et al., 2000: 143.
Diagnosis. — Rostrum sharply triangular, not reaching distal margin of
eyestalks. Frontal margin of carapace with anterolateral spinous projections.
Mxp3 propodus subquadrate, and dactylus digitiform. Male Plp1 blade-shaped,
uniramous, bisegmented, and subchelate distally; male Plp2 unknown. Telson
shaped as inverted trapezoid, bearing small median spine on posterior margin.
Uropodal endopod fusiform. [Adapted from Edmondson, 1944, fig. 11f.]
Type locality. — Maui, Hawaii.
Distribution. — Only known from the type locality.
Neocallichirus sp. (Rathbun, 1906)
Callianassa sp., Rathbun, 1906: 893, fig. 48.
Callianassa (Trypaea) sp. — De Man, 1928b: 28.
Neocallichirus sp. — Sakai, 1999c: 108.
Remarks. — Rathbun (1906) gave a figure of her uncertain species, showing
that the Mxp3 ischium-merus is subrectangular, the merus is subtriangular, the
propodus is broadened, and the dactylus is slender, but no figure of the tailfan, which makes it very difficult to determine the genus to which her species
belongs. So, it is tentatively classified under Neocallichirus. [Cf. Rathbun,
1906: 893, fig. 48.]
Distribution. — [Honolulu, Hawaii?]
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CRM 013 – Katsushi Sakai
Genus Podocallichirus Sakai, 1999
Podocallichirus Sakai, 1999c: 53; Sakai, 2005b: 187.
Diagnosis. — Carapace with dorsal oval. Rostrum shaped as elongate
triangle, with acute apex. A1 peduncle longer and stouter than A2 peduncle.
Mxp3 ischium-merus rectangular; merus about as long as wide, lateral margin
convex; carpus longer than wide, and longer than merus; propodus broadened,
shorter than carpus, and largely convex on ventral margin; dactylus digitiform
and obtuse distally; exopod absent. P1 unequal and chelate; in male merus of
larger cheliped proximally convex on ventral margin; dactylus spoon-like, and
spinose on dorsal and ventral margins; fixed finger broadly concave on cutting
surface, armed with spines on both mesial and lateral sides; in male merus
of smaller cheliped unarmed on ventral margin; dactylus unarmed on both
dorsal and ventral margins; in female meri of both larger and smaller chelipeds
bearing proximal hook on ventral margin; chelae as in males. P2 chelate. P3
simple. P4 subchelate. P5 chelate. Male Plp1 uniramous and bisegmented,
distal segment short and chelate distally; Plp2 biramous, endopod bearing
mesiodistally fusiform appendix masculina with marginal setae and a small,
proximally-fused appendix interna with hooklets mesiomedially. Telson wider
than long, and slightly convex on posterior margin. Uropodal endopod rhombic
in shape; uropodal exopod broadly convex on lateral margin, bearing dorsal
plate.
Remarks. — Sakai (2005b: 187) included 8 species in the genus Podocallichirus Sakai, 1999c: P. balssi (Monod, 1933); P. foresti (Le Loeuff & Intès,
1974); P. guineensis (De Man, 1928a); P. tenuimanus (De Saint Laurent & Le
Loeuff, 1979) from the eastern Atlantic-Mediterranean region; and P. gilchristi
(Barnard, 1946); P. madagassus (Lenz & Richters, 1881); P. masoomi (Tirmizi,
1970); P. tamakii (Poore, 2000) from the Indo-West Pacific region.
In the present revision, however, only P. madagassus is retained in the genus
Podocallichirus as the type species of the genus, while the other species are
reclassified by the forms of the male Plps1-2; P. balssi and P. guineensis
are reclassifed under the new genus Balsscallichirus gen. nov.; P. foresti
is reclassified under the new genus Forestcallichirus gen. nov.; P. gilchristi
(Barnard, 1946) and P. tenuimanus (De Saint Laurent & Le Loeuff, 1979)
under the new genus Barnardcallichirus gen. nov.; P. masoomi under the new
genus Tirmizicallichirus gen. nov., and P. tamakii under Grynaminna Poore,
2000.
Type species. — Callianassa madagassa Lenz & Richters, 1881. The
gender of the generic name, Podocallichirus, is masculine.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
467
Species included. — Only the type species, Podocallichirus madagassus
(Lenz & Richters, 1881).
Podocallichirus madagassus (Lenz & Richters, 1881)
(fig. 66C-D)
Callianassa madagassa Lenz & Richters, 1881: 427, figs. 20-23; Tudge et al., 2000: 143.
Callianassa (Callichirus) madagassa — Borradaile, 1903: 547; De Man, 1928a: 42, fig. 18-18e;
De Man, 1928b: 29, 92, 113.
Podocallichirus madagassus — Sakai, 1999c: 56, fig. 10a-d; Sakai, 2005b: 191, fig. 38.
Material. — SMF 7938, lectotype, 1 female (TL/CL, 52.0/11.2 mm), SMF 4953, paralectotype, 1 male (TL/CL, 49.0/10.7 mm), Nosi Bé, Madagascar, 1880, leg. Ebenau.
Diagnosis. — Mxp3 ischium-merus subpediform; merus convex on
mesiodistal margin. Male Plp1 uniramous and bisegmented, distal segment
short, chelate distally, with distal emargination (fig. 66C); male Plp2 biramous,
endopod bearing mesiodistally fusiform appendix masculina with proximallyfused appendix interna mesiomedially (fig. 66D). Telson wider than long and
subrectangular, and posterior margin slightly convex medially, without median
spine.
Type locality. — Nosi Bé, N.W. Madagascar.
Distribution. — Only known from the type locality.
Genus Sergio Manning & Lemaitre, 1994 (sensu nov.)
Sergio Manning & Lemaitre, 1994 (not 1993): 39 (partim).
Sergio s. str. — Felder, 2001: 440.
Diagnosis. — Carapace with dorsal oval, but without rostral carina, cardiac
prominence and transverse cardiac sulci. Rostrum small and spinous. Carapace
with anterolateral projections calcified proximally. A1 peduncle shorter than
A2 peduncle. Mxp3 ischium-merus subquadrate; merus triangular; propodus
broadened and rounded on ventral margin; dactylus narrow and digitiform; exopod absent. P1 chelate, unequal, and dissimilar; in male larger cheliped merus
with small meral hook. P2 chelate. P3 propodus broadened. P4 subchelate. P5
chelate. Abdominal somites 3-5 not ornamented dorsally. Male Plp1 uniramous and bisegmented, distal segment bilobed by median longitudinal furrow,
and subchelate distally; male Plp2 biramous, endopod bearing a proximallyfused or not fused appendix masculina, with or without proximally-fused appendix interna. Male Plps3-5 biramous and foliaceous, endopods with appendix interna in both sexes. Telson wider than long and varied in shape. Uropodal
endopod longer than wide and rhombic, overreaching telson; uropodal exopod
468
CRM 013 – Katsushi Sakai
convex on lateral margin, bearing dorsal plate. [Adapted from Sakai, 1999c:
72.]
Remarks. — The genus Sergio Manning & Lemaitre, 1994 was established for Callianassa guassutinga Rodrigues, 1971, and included three other
species, S. guarus (Rodrigues, 1971), S. mirim (Rodrigues, 1971), and S. trilobatus (Biffar, 1970). Though in the previous revisions (Sakai, 1999c, 2005b)
Sergio was synonymized with Neocallichirus Sakai, 1988 (cf. Sakai, 1999c,
2005b), in the present revision it has turned out by comparing Sergio with
Neocallichirus on the male Plp1, that Sergio is different from Neocallichirus.
In S. guassutinga, the type species of Sergio, the male Plp1 distal segment is
bilobed by a median, longitudinal furrow, and subchelate distally; whereas in
N. horneri Sakai, 1988, the type species of Neocallichirus, the male Plp1 distal
segment is w-shaped distally.
The three species (S. guarus (Rodrigues, 1971), S. mirim (Rodrigues, 1971),
and S. trilobata (Biffar, 1970)), included in Sergio by Manning & Lemaitre
(1994) are thus reclassified by the forms of the male Plps1-2. S. trilobatus
(Biffar, 1970) is included in Glypturoides gen. nov., because the male Plp1
distal segment is subchelate distally, and the male Plp2 endopod bears distally
a triangular appendix masculina with marginal setae, and mesiodistally a small
appendix interna (fig. 65J). The other two species, S. guarus and S. mirim are
reclassified under Neocallichirus.
Type species. — Callianassa guassutinga Rodrigues, 1966, by original
designation. The gender of the generic name, Sergio, is masculine.
Species included. — Sergio cacahuate (Felder & Manning, 1995); S.
guassutinga (Rodrigues, 1971); S. lemaitrei (Manning, 1993); S. mericeae
Manning & Felder, 1995; S. monodi (De Saint Laurent & Le Loeuff, 1979);
S. pachydactylus (A. Milne-Edwards, 1870); S. sulfureus Lemaitre & Felder,
1996.
K EY
TO THE
S PECIES
OF THE
G ENUS SERGIO
1 – Telson convex on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Telson concave on posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 – Eyestalks with truncate apex with denticles . . . . . . . . . . . . . . . . . . . . Sergio pachydactylus
– Eyestalks with obtuse apex without denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – In male larger cheliped carpus and palm denticulate on ventral margin . . . . . S. lemaitrei
– In male larger cheliped carpus and palm smooth on ventral margin . . . . . . . S. cacahuate
4 – Posterior margin of telson bearing concavity with triangular median protrusion . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. monodi
– Posterior margin of telson concave and smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
469
5 – Male Plp2 endopod bearing appendix masculina distally . . . . . . . . . . . . . . S. guassutinga
– Male Plp2 endopod bearing appendix masculina mesiodistally . . . . . . . . . . . . . . . . . . . . . 6
6 – In male larger cheliped, dactylus bearing two teeth on proximal half of cutting edge, and
is distal to these tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sulfureus
– In male larger cheliped, dactylus bearing truncate tooth and sharp tooth on proximal
half of cutting edge, and is distal to these serrated . . . . . . . . . . . . . . . . . . . . . . . S. mericeae
Sergio cacahuate (Felder & Manning, 1995)
Neocallichirus cacahuate Felder & Manning, 1995: 478, figs. 1a-c, 2, 3a-e, 4a-c, 5; Sakai,
1999c: 88; Blanco Rambla, 2000: 74-75, fig. 3; Tudge et al., 2000: 144; Sakai, 2005b: 164.
Diagnosis. — Mxp3 ischium-merus subrectangular; merus subtriangular.
Male Plp1 uniramous and bisegmented, distal segment shorter than proximal,
and divided into two lobes by weak longitudinal furrow, and subchelate distally
(cf. Felder & Manning, 1995: 484, fig. 1c); male Plp2 biramous, endopod
bearing mesiodistally a proximally-fused appendix masculina with long distal
setae, provided mesiomedially with a proximally-fused small, grain-shaped
appendix interna (cf. Felder & Manning, 1995, figs. 1c, 4a). Telson wider than
long and trapezoid, and slightly sinuous on posterior margin (cf. Felder &
Manning, 1995, fig. 4c).
Remarks. — Felder & Manning (1995) described the present species as
Neocallichirus cacahuate, mentioning that “N. cacahuate can be distinguished
from the members of the closely related genus Sergio on the basis of its
relatively longer, posteriorly rounded to truncate telson, and its broader, more
rectangular uropodal endopods.” However, this species belongs to Sergio by
the form of the male Plp1, which is the same as that of the type species of
Sergio, S. guassutinga, as described in the diagnosis.
Type locality. — Florida, West Palm Beach County, Lake Worth, north
side of Peanut Island, sparsely vegetated sandy to shelly sand intertidal flats
(26◦ 46.7′ N 80◦ 2.9′ W).
Distribution. — Only known from the type locality.
Sergio guassutinga (Rodrigues, 1971)
(figs. 67H-I, 68)
[Abbreviated list of synonymy.]
Callianassa (Callichirus) guassutinga Rodrigues, 1971: 204, figs. 41-60.
Callianassa guassutinga — Biffar, 1971a: 651, 653, 674, figs. 9, 10.
Sergio guassutinga — Manning & Lemaitre, 1994: 40; Coelho, 1997: 149; Rodrigues et al.,
1998: 381; Melo, 1999: 378, figs. 253-254; Coelho et al., 2007: 5 (tab. 2).
Neocallichirus guassutingus — Sakai, 1999c: 90.
470
CRM 013 – Katsushi Sakai
Fig. 68. Sergio guassutinga (Rodrigues, 1971). A, male Plp1; B, male Plp2. SMF 32727, 1 male
(TL/CL, 90.0/22.3 mm), Cabo Branco, Paraíba, Brazil. Scales 1 mm.
Neocallichirus guassutinga — Sakai, 2005b: 166.
Material examined. — USNM 256886, paratype, male (TL/CL, 100.6/26.6 mm), Praia do
Araç, São Sebastião, São Paulo, Brazil, 06.i.1966, leg. S.A. Rodrigues; SMF 32727, 1 male
(TL/CL, 90.0/22.3 mm), Cabo Branco, Paraíba, Brazil, 16.ix.1982, leg. P.M. Dijck, det. S.A.
Rodrigues, presented by R.M. Shimizu.
Diagnosis. — Mxp3 ischium-merus subrectangular; merus subtriangular.
Male Plp1 uniramous, blade-shaped, and bisegmented, distal segment bilobed
by median longitudinal furrow, and subchelate distally (figs. 67H, 68A); male
Plp2 biramous, endopod bearing distally appendix masculina (figs. 67I, 68B).
Telson trapezoid, concave on posterior margin, without median spine.
Type locality. — Brazil, São Sebastião.
Distribution. — North side of Fort Pierce Inlet, Florida; Indian River
Lagoon, St. Lucie County, Florida; Louisiana; 12-13 m; Texas; Barra del
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
471
Tordo, Tamaulipas, Mexico; Gulf of Panama; Pernambuco and São Paulo,
Brazil.
Sergio lemaitrei (Manning, 1993)
Neocallichirus lemaitrei Manning, 1993: 107, figs. 1-3; Felder & Manning, 1995: 488, fig. 6;
Blanco-Rambla, 1998: 20, figs. 1-4; Sakai, 1999c: 91; Tudge et al., 2000: 144; Sakai, 2005b:
167.
Diagnosis. — Rostrum inconspicuous. Eyestalks contiguous and pointed
apically. Frontal margin of carapace with inconspicuous anterolateral convexities. Mxp3 ischium-merus rectangular; merus obliquely declined on mesiodistal margin; propodus subquadrate, and dactylus digitiform. In male larger cheliped carpus and palm denticulate on ventral margin. Male Plp1 blade-shaped,
uniramous, bisegmented, and subchelate distally; male Plp2 biramous, endopod bearing fused appendix masculina distally, and fused with appendix interna mesiodistally. Telson shaped as inverted trapezoid, lacking median spine
on posterior margin. [Adapted from Manning, 1993: 107, figs. 1h, 3a, c; Felder
& Manning, 1995, fig. 6b, c.]
Remarks. — The present species, which was described by Manning (1993)
as Neocallichirus lemaitrei, is characterized as follows: the male Plp1 is uniramous and bisegmented, and the distal segment is bilobed by a weak longitudinal furrow, and subchelate distally; the male Plp2 endopod bears an appendix
masculina provided mesiomedially with a proximally-fused appendix interna
(cf. Felder & Manning, 1995, fig. 6b). The male Plp1 of N. lemaitrei shows the
typical form of that of S. guassutinga, so that Neocallichirus lemaitrei is now
reclassifed under Sergio, as S. lemaitrei.
Type locality. — Isla del Rosario (10◦ 10′ N 75◦ 46′ W), Colombia, beach on
south side.
Distribution. — Isla del Rosario, Colombia, Caribbean side.
Sergio mericeae Manning & Felder, 1995
Sergio mericeae Manning & Felder, 1995: 267, figs. 1a-f, 2a-f, 3a-f, 4a-f, 5a-g; Robles et al.,
2009: 317.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment divided into two lobes by weak longitudinal furrow, and subchelate distally;
male Plp2 biramous, endopod bearing mesiodistally a proximally-fused appendix masculina with distal setae.
Type locality. — North side of Fort Pierce Inlet, Florida, Indian River
Lagoon, intertidal sandflat, 27◦ 28.2′ N 80◦ 18.8′ W.
472
CRM 013 – Katsushi Sakai
Distribution. — Florida, Louisiana, Texas; Mexico (Manning & Felder,
1995).
Sergio monodi (De Saint Laurent & Le Loeuff, 1979)
Callichirus monodi De Saint Laurent & Le Loeuff, 1979: 71, figs. 14h, 16f, 17c, 18a, 19h,
22a-b, 23n-r; Tudge et al., 2000: 144.
Neocallichirus monodi — Sakai, 1999c: 86; Sakai, 2005b: 162.
Diagnosis. — Mxp3 ischium-merus subsquare; merus obliquely curved distally and rounded on distomesial margin; propodus subquadrate, and dactylus digitiform. Male Plp1 uniramous, bisegmented, distal segment bilobed by
weak longitudinal furrow, and subchelate distally; male Plp2 biramous, endopod bearing mesiodistally appendix masculina with proximally-fused appendix interna laterally (cf. De Saint Laurent & Le Loeuff, 1979, fig. 23o). Telson
wider than long, bearing broad concavity with triangular median protrusion on
posterior margin. [Cf. De Saint Laurent & Le Loeuff, 1979: 71, figs. 17c, 18a,
19h, 23n-r.]
Type locality. — Senegal.
Distribution. — Only known from the type locality.
Sergio pachydactylus (A. Milne-Edwards, 1870)
Callianassa pachydactyla A. Milne-Edwards, 1870: 86, pl. 2 fig. 1a-d.
Callianassa (Cheramus) pachydactyla — Borradaile, 1903: 545; De Man, 1928b: 19, 26, 94,
100, 121, 160-164.
Callichirus pachydactyla — De Saint Laurent & Le Loeuff, 1979: 76, figs. 14j, 16d-e, 18e, 19k,
22c-d, 23s-v.
Neocallichirus pachydactylus — Sakai, 1999c: 86, fig. 20a-c; Dworschak et al., 2000: 309;
Tudge et al., 2000: 144; Sakai, 2005b: 163.
Not: Callianassa pachydactyla — Longhurst, 1958: 42 [= Forestcallichirus foresti (Le Loeuff
& Intès, 1974)].
Diagnosis. — Rostrum triangular, reaching proximal third of eyestalks. Eyestalks with truncate apex with denticles. Male Plp1 uniramous and bisegmented, distal segment bilobed by short longitudinal furrow and subchelate
distally; male Plp2 biramous, endopod bearing appendix masculina distally,
but no appendix interna. Telson shaped as inverted trapezoid, lacking median
spine on posterior margin. [Adapted from De Saint Laurent & Le Loeuff, 1979,
23s-t.]
Remarks. — Dworschak et al. (2000) stated that this species was collected
by yabby pump together with Salmoneus caboverdensis Dworschak, Anker &
Abed-Navandi 2000 (Alpheidae) in the Cape Verde Islands.
Type locality. — Cape Verde Islands.
Distribution. — Cape Verde Is.; Senegal; Ghana; Île Príncipe; intertidal.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
473
Sergio sulfureus Lemaitre & Felder, 1996
Sergio sulfureus Lemaitre & Felder, 1996: 453-463, text-figs. 1-6; Tudge et al., 2000: 145.
Neocallichirus sulfureus — Sakai, 1999c: 92; Sakai, 2005b: 169.
Diagnosis. — Mxp3 ischium-merus subrectangular; merus subtriangular;
propodus subquadrate, and dactylus digitiform. Male Plp1 uniramous, bladeshaped, and bisegmented, distal segment bilobed by weak longitudinal furrow,
and subchelate distally; male Plp2 biramous, endopod bearing mesiodistally
a proximally-fused appendix masculina with distal setae, provided mesiolaterally with a proximally-fused appendix interna. Telson wider than long and
trapezoid, posterior margin concave, without median spine. Uropodal endopod
longer than broad, and with tuft of setae dorsally near posterodistal corner. [Cf.
Lemaitre & Felder, 1996, figs. 6b, d, e, j.]
Type locality. — Caribbean coast of Colombia, S.W. shoreline of Barú,
beach facing Rosario Islands.
Distribution. — Only known from the type locality.
Genus Thailandcallichirus gen. nov.
Diagnosis. — Front of carapace with three low, subtriangular projections.
Carapace with dorsal oval, but without rostral carina, cardiac prominence, or
transverse cardiac sulcus(i). A1 peduncle shorter than A2 peduncle. Mxp3
ischium-merus operculiform; propodus subovate, broadly rounded on ventral margin; dactylus subovate, rounded distally; exopod absent. P1 chelate,
unequal, and dissimilar; in male larger cheliped merus with meral hook.
P2 chelate. P3 propodus broadened. P4 subchelate. P5 chelate. Abdominal
somites 3-5 not ornamented dorsally. Male Plp1 uniramous and bisegmented,
distal segment subchelate distally; male Plp2 biramous, endopod bearing
mesiodistally appendix masculina with distal setae, provided with appendix
interna mesiomedially (cf. Sakai, 1983, fig. 1); male Plps3-5 biramous and
foliaceous, endopods with stubby, embedded appendix interna. Telson wider
than long and trapezoid. Uropodal endopod rhombic. Uropodal exopod convex
on lateral margin, bearing dorsal plate.
Remarks. — In the previous revision, Callianassa ranongensis Sakai,
1983 was included by Sakai (1999c) in Neocallichirus. However, it is found
that N. ranongensis is different from the type species of Neocallichirus, N.
horneri Sakai, 1988. In N. ranongensis the Mxp3 dactylus is rounded distally,
and the male Plp2 endopod bears mesiodistally an appendix masculina with
distal setae, provided with an appendix interna mesiomedially, whereas in
474
CRM 013 – Katsushi Sakai
N. horneri the Mxp3 dactylus is obtuse distally, and the male Plp2 endopod
bears distally a proximally-fused appendix masculina with distal setae, and a
proximally-fused appendix interna distolaterally. Therefore, N. ranongensis is
excluded from Neocallichirus, and reclassified under the present new genus
Thailandcallichirus as T. ranongensis.
Type species. — Callianassa ranongensis Sakai, 1983, by present designation and monotypy. The gender of the generic name, Thailandcallichirus, is
masculine.
Species included. — Thailandcallichirus ranongensis (Sakai, 1983).
Etymology. — The new generic name Thailandcallichirus is composed of
a generic name, Callichirus and the type locality, Thailand.
Thailandcallichirus ranongensis (Sakai, 1983)
Callianassa ranongensis Sakai, 1983: 111-115, figs. 1, 2; Sakai, 1987b: 45, figs. 1, 2.
Neocallichirus ranongensis — Sakai, 1999c: 107; Sakai, 2005b: 183.
Lepidophthalmus ranongensis — Tudge et al., 2000: 144.
Type locality. — Hatsaikhao, Ranong Province, Thailand, muddy mangrove
swamp.
Distribution. — Ranong Province, Thailand; Halmahera, Indonesia.
Genus Tirmizicallichirus gen. nov.
Diagnosis. — Carapace with dorsal oval. A1 peduncle longer than A2 peduncle. Mxp3 ischium-merus pediform, with parallel mesial and lateral margins; propodus subovate, broadened and strongly convex on ventral margin;
dactylus digitiform and obtuse distally. Male Plp1 uniramous and bisegmented,
distal segment obtuse distally; male Plp2 biramous, endopod obtuse distally,
lacking appendices interna and masculina; Plps3-5 with inconspicuous appendix interna. Telson subquadrate, wider than long, with low median convexity on posterior margin. Uropodal endopod oval; uropodal exopod broadly
rounded on lateral margin, bearing dorsal plate.
Remarks. — Podocallichirus masoomi (Tirmizi, 1970) (cf. Sakai, 1999c),
has turned out to be different from the type species of Podocallichirus,
Podocallichirus madagassus (Lenz & Richters, 1881), in the forms of male
Plps1-2. In P. masoomi the male Plp1 is uniramous and bisegmented, and the
distal segment is simple and obtuse distally, the male Plp2 is biramous, and
the endopod bears neither an appendix interna nor an appendix masculina.
Whereas in P. madagassus the male Plp1 is uniramous and bisegmented, but
the distal segment is short and chelate distally, the male Plp2 is biramous, and
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
475
the endopod bears mesiodistally a fusiform appendix masculina, fused mesiomedially with an appendix interna with hooklets. Therefore, Podocallichirus
masoomi is excluded from Podocallichirus and reclassified under the present
new genus, Tirmizicallichirus gen. nov. as Tirmizicallichirus masoomi (Tirmizi, 1970).
Type species. — Callianassa (Callichirus) masoomi Tirmizi, 1970, by
present designation and monotypy. The gender of the generic name, Tirmizicallichirus, is masculine.
Species included. — Tirmizicallichirus masoomi (Tirmizi, 1970).
Etymology. — The new generic name is dedicated to the late Prof. Dr. N.M.
Tirmizi, who was a leading carcinologist in Pakistan, by adding her name as a
prefix to Callichirus.
Tirmizicallichirus masoomi (Tirmizi, 1970)
[Abbreviated list of synonymy.]
Callianassa (Callichirus) masoomi Tirmizi, 1970: 245, figs. 1-3.
Callianassa (Callichirus) kewalramanii Sankolli, 1971: 94, figs. 5, 8. [Type locality. —
Ratnagiri, Bombay, India.]
Podocallichirus masoomi — Sakai, 1999c: 58; Sakai, 2005b: 192.
Material examined. — Invertebrate Reference Museum, Karachi Univ., holotype, female
(TL 22.5 mm), Karachi, sandy shore.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment obtuse distally; male Plp2 biramous, endopod obtuse distally, bearing neither appendix interna nor appendix masculina; Plps3-5 with inconspicuous appendix
interna. Telson subquadrate, wider than long, with low median convexity on
posterior margin. [Cf. Sankolli, 1971: 94, figs. 5c, 6h, 8a, b.]
Type locality. — Pakistan, Bholegi, W. of Karachi, intertidal zone of muddy
sand beach with loose stones.
Distribution. — Bholegi, W. of Karachi, Pakistan; Ratnagiri, Bombay, India.
FAMILY CALLIANOPSIDAE MANNING & FELDER, 1991
(SENSU NOV.)
Diagnosis. — Rostrum sharply triangular, bearing rostral carina. Carapace
with or without dorsal oval, and without cardiac sulcus. Eyestalks flattened
and contiguous. A2 scaphocerite present as a small process. Maxilla 2 scaphognathite without posterior whip. Mxp3 ischium-merus subrectangular; propodus broadened; dactylus ovate or subovate; exopod absent. P1 chelate, unequal
in size, and dissimilar in shape; in larger cheliped merus with small hook
ventroproximally; palm oblong; fingers as long as or shorter than palm; in
smaller cheliped merus with hook ventroproximally; fingers longer than palm.
P2 chelate. P3 propodus broadened. P4 simple. P5 subchelate. Abdominal
pleuron 2 unarmed ventrally, pleura 4-5 with or without spine-like projection
posteroventrally; and abdominal somite 6 with spine-like projection medioventrally. Male Plp1 uniramous and bisegmented, distal segment sickle-shaped;
male Plp2 endopod bearing appendix masculina with distal setae and small
appendix interna; male Plps3-5 similar, endopods with finger-like appendix
interna. Telson subsquare, bearing proximal lobe on lateral margin. Uropodal
exopod without dorsal plate or lateral notch.
Remarks. — The present family Callianopsidae is different from the Callianassidae by the form of the uropodal exopod; in the Callianopsidae the
uropodal exopod lacks both dorsal plate and lateral notch, while in the Callianassidae the uropodal exopod bears a dorsal plate but no lateral notch.
Subfamilies included. — Callianopsinae Manning & Felder, 1991; Neocallianopsinae subfam. nov.
K EY
TO THE
S UBFAMILIES
OF THE
FAMILY C ALLIANOPSIDAE
1 – Carapace with dorsal oval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callianopsinae
– Carapace without dorsal oval . . . . . . . . . . . . . . . . . . . . . . . Neocallianopsinae subfam. nov.
Subfamily C ALLIANOPSINAE Manning & Felder, 1991
Callianopsinae Manning & Felder, 1991: 787; Hopkins & Feldmann, 1997: 237; Sakai, 2005b:
226.
Diagnosis. — Rostrum sharply triangular, bearing rostral carina. Carapace
with dorsal oval and cardiac prominence, but without cardiac sulcus. Eye-
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CRM 013 – Katsushi Sakai
stalks flattened and contiguous. Linea thalassinica present. A2 scaphocerite
present as a small process. Mxp3 ischium-merus rectangular or subrectangular; propodus subrectangular; dactylus ovate; exopod absent. Male Plp1 uniramous, bisegmented; male Plp2 biramous. Uropodal exopod bearing neither
dorsal plate nor lateral notch.
Remarks. — C. goniophthalma (Rathbun, 1902), the type species of Callianopsis, the type genus of the Callianopsinae, was described by Hopkins &
Feldmann (1997: 238) with their figures (1997, fig. 1j, k) as follows: “The
first pleopod in the male of the species is sickle-shaped and second pleopod
possesses an ovate exopod that is longer than wide and a longer than wide
obovate endopod that possesses a bifurcate appendix interna”. However, it has
turned out by the present examination of the type specimen of C. goniophthalma (USNM 25238, type, male), that the male Plp2 endopod bears mesiomedially an appendix masculina with distal setae and a small appendix interna
proximally.
Type genus. — Callianopsis De Saint Laurent, 1973.
Genera included. — Callianopsis De Saint Laurent, 1973; Pleurocalliax
gen. nov.
K EY
TO THE
G ENERA
OF THE
S UBFAMILY C ALLIANOPSINAE
1 – Pleura 2-6 with spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pleurocalliax gen. nov.
– Pleura 5-6 with spine-like projection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callianopsis
Genus Callianopsis De Saint Laurent, 1973
Callianopsis De Saint Laurent, 1973: 515; Hopkins & Feldmann, 1997: 237; Sakai, 2005b: 227.
Diagnosis. — Rostrum small and triangular, bearing rostral carina. Carapace with dorsal oval and cardiac prominence, but without cardiac sulcus.
TABLE X
Major features of Callianopsinae
Dorsal oval
on carapace
Dorsal plate on
uropodal exopod
Mxp3
exopod
Spine or spine-like projection
on abdominal pleura
Callianopsis
present
absent
absent
present on
pleura 5-6
Pleurocalliax
gen. nov.
present
absent
absent
present on
pleura 2-6
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
479
Eyestalks flattened and contiguous. Linea thalassinica present. A2 scaphocerite present as a small process. Mxp3 ischium-merus subrectangular; propodus subrectangular; dactylus subovate; exopod absent. P1 chelate, unequal in
size, and dissimilar in shape; in larger cheliped merus with small meral hook
ventroproximally; palm rectangular; fingers almost as long as palm; in smaller
cheliped merus with small meral hook ventroproximally; fingers longer than
palm. P2 chelate. P3 propodus broadened. P4 simple. P5 subchelate. Abdominal pleura 2-4 unarmed ventrally; pleuron 5 with spine-like projection posteroventrally; and abdominal pleuron 6 with spine-like projection anteroventrally. Male Plp1 uniramous and bisegmented, distal segment sickle-shaped;
male Plp2 endopod bearing mesiomedially appendix masculina with distal setae and small appendix interna proximally; male Plps3-5 similar, endopods
with finger-like appendix interna. Telson subsquare. Uropodal exopod bearing
neither dorsal plate nor lateral notch.
Type species. — Callianassa goniophthalma Rathbun, 1902, by original
designation and monotypy. The gender of the generic name, Callianopsis, is
feminine.
Species included. — Callianopsis goniophthalma (Rathbun, 1902).
Callianopsis goniophthalma (Rathbun, 1902)
(fig. 63H-J)
Callianassa goniophthalma Rathbun, 1902: 886; Rathbun, 1904: 154, pl. 8; Schmitt, 1921: 121,
fig. 82; Balss, 1925: 211; Stevens, 1928: 342, fig. 19; Williams et al., 1989: 28.
?Callianassa (Calliactites) goniophthalma — Borradaile, 1903: 545; De Man, 1928b: 25 (list),
95, 96.
Callianopsis goniophthalma — Hart, 1982: 54, fig. 13; Manning & Felder, 1991: 789 (list),
figs. 7, 18; Hendrickx, 1995: 390 (list); Hopkins & Feldmann, 1997: 238, figs. 1-4; Tudge et
al., 2000: 142; Hendrickx et al., 2005: 170 [in the family Ctenochelidae]; Sakai, 2005b: 229.
Material examined. — USNM 25238, type, male, damaged, off Point Conception, California, 278 fms, Sta. 3198, R/V “Albatross”.
Diagnosis. — Maxilla 2 scaphognathite without posterior whip (cf. Hopkins
& Feldmann, 1997, fig. iC). Mxp3 ischium-merus subrectangular, propodus
subrectangular, and dactylus subovate; exopod absent. Male Plp1 uniramous
and bisegmented, distal segment sickle-shaped (fig. 63H); male Plp2 biramous,
endopod bearing mesiomedially appendix masculina with distal setae and
small appendix interna proximally (fig. 63I); male Plps3-5 endopods with
appendix interna (fig. 63J). Uropodal exopod bearing neither dorsal plate nor
lateral notch.
Remarks. — Rathbun (1902) established Callianassa goniophthalma Rathbun, 1902, which was later described as ?Callianassa (Calliactites) gonioph-
480
CRM 013 – Katsushi Sakai
thalma by Borradaile (1903: 545) and De Man (1928: 25). However, the type
species of Calliactites, C. secura Lanchester, 1902 is a synonym of Callianidea typa H. Milne Edwards, 1837, now included in the Callianideidae, and is,
therefore, not any more a member of the Callianassidae. So, Manning & Felder
(1991: 787) established a new subfamily Callianopsinae in the Ctenochelidae,
based on Callianopsis as the type genus. However, the type species of Callianopsis, C. goniophthalma, bears a dorsal oval, which suggests that C. goniophthalma seems not to be included in the Ctenochelidae but probably in the
Callianassidae. However, C. goniophthalma is not included in the Callianassidae either, in which the Mxp3 dactylus is digitiform, but in the Callianopsidae
sensu nov., in which the Mxp3 dactylus is oval.
Type locality. — Off Point Conception, California, “Albatross” Sta. 3198,
513 m.
Distribution. — Alaska — Clarence Strait (Rathbun, 1902; Hopkins & Feldmann, 1997), Funter Bay (Stevens, 1928), Yes Bay (Hopkins & Feldmann,
1997). California — off Point Conception (Rathbun, 1902; Hopkins & Feldmann, 1997), off Harris Point, San Miguel Island (Schmitt, 1921). Baja California — San Cristobal Bay, Mexico (Hopkins & Feldmann, 1997); 351595 m.
Genus Pleurocalliax gen. nov.
Diagnosis. — Carapace with dorsal oval, cardiac prominence, and linea thalassinica, but without transverse cardiac sulcus. Mxp3 ischium-merus broadened and pediform; dactylus ovate. P1 unequal and dissimilar; in larger cheliped merus with hook ventroproximally. Abdominal pleuron 2 with laterallydirected acute projection posteroventrally; pleura 3-4 with laterally-directed
spine medioventrally; and pleura 5-6 with laterally-directed spine anteroventrally. Male Plps1-2 unknown. Telson convergent toward posterior margin.
Uropodal exopod bearing neither dorsal plate nor lateral notch. [Described by
Alcock & Anderson, 1894: 163; Alcock & Anderson, 1896, pl. 26 fig. 2.]
Remarks. — Callianassa coecigena Lanchester, 1902 was included by
Borradaile (1903: 545) and De Man (1928: 26) in the genus Calliactites s.
str. Borradaile, 1903 (syn. of Callianidea H. Milne Edwards, 1837a). However,
Calliactites coecigena, which is differrent from the type species of Calliactites,
C. secura Lanchester, 1902, is rather similar to the type species of the genus
Callianopsis, C. goniophthalma, in that the carapace bears a dorsal oval and
a cardiac prominence (Alcock & Anderson, 1896, pl. 26 fig. 2), but differs in
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
481
that in Calliactites coecigena abdominal pleuron 2 is armed with a laterallydirected acute projection posteroventrally, pleura 3-4 with a laterally-directed
acute projection medioventrally, and pleura 5-6 with a laterally-directed acute
projection anteroventrally; whereas in Callianopsis goniophthalma abdominal
pleura 2-5 are unarmed and pleuron 6 is armed with a laterally-directed acute
projection anteroventrally; which means that Calliactites coecigena is not to
be included in the genus Callianopsis, either. Therefore, Calliactites coecigena
is reclassified under the present new genus, Pleurocalliax gen. nov.
Type species. — Callianassa coecigena Alcock & Anderson, 1894. The
gender of the generic name, Pleurocalliax, is feminine.
Species included. — Pleurocalliax coecigena (Alcock & Anderson, 1894).
Etymology. — The type species of the present new genus Pleurocalliax is
characterized by the lateral projections on the abdominal pleura, so that the
new generic name is composed of a Greek word “pleura” meaning “side”, and
the generic name Calliax.
Pleurocalliax coecigena (Alcock & Anderson, 1894)
Callianassa coecigena Alcock & Anderson, 1894: 163; Alcock & Anderson, 1896, pl. 26 fig. 2,
2a-b; Alcock, 1901: 198; Balss, 1925: 211.
(?) Callianassa (Calliactites) coecigena — Borradaile, 1903: 545.
Callianassa (Calliactites) coecigena — De Man, 1928b: 25 (list), 96 (key).
Callianopsis coecigena — De Saint Laurent & Le Loeuff, 1979: 95.
Callianopsis coecigena — Sakai, 2005b: 229.
Diagnosis. — Rostrum acutely triangular, reaching distal margin of eyestalks. Carapace with dorsal oval, cardiac prominence, and linea thalassinica,
but without cardiac sulcus. Eyestalks petaloid; cornea absent. Mxp3 ischiummerus rectangular; propodus rectangular dactylus ovate. P1 unequal; in larger
cheliped palm setose on lateral surface. Abdominal pleuron 2 with laterallydirected acute projection posteroventrally; pleura 3-4 with laterally-directed
spine medioventrally; and pleura 5-6 with laterally-directed spine anteroventrally. Male Plps1-2 unknown. Telson convergent toward convex posterior margin. Uropodal exopod bearing neither dorsal plate nor lateral notch. [Cf. Alcock & Anderson, 1894: 163; Alcock & Anderson, 1896, pl. 26 fig. 2a.]
Remarks. — The type specimen is inaccessible. However, this species is
very characteristic in that abdominal pleuron 2 is armed with a laterallydirected acute projection posteroventrally, pleura 3-5 with a laterally-directed
acute projection medioventrally, and pleuron 6 with a laterally-directed acute
projection anteroventrally, so that Callianassa coecigena is reclassified under
the new genus Pleurocalliax as P. coecigena.
482
CRM 013 – Katsushi Sakai
Type locality. — Off Sri Lanka, Bay of Bengal, 365-690 m.
Distribution. — Off Sri Lanka (Alcock & Anderson, 1894); 365-690 m.
Subfamily N EOCALLIANOPSINAE subfam. nov.
Diagnosis. — Rostrum sharply triangular, bearing rostral carina. Carapace
bearing median longitudinal carina with cardiac prominence, and linea thalassinica, but without dorsal oval and cardiac sulcus. Mxp3 ischium-merus
rectangular; propodus rectangular; dactylus ovate; exopod absent. Abdominal
pleura 2-4 unarmed ventrally; pleuron 5 armed with spine-like projection posteroventrally; abdominal somite 6 with spine-like projection anteroventrally.
Uropodal exopod bearing neither dorsal plate nor lateral notch.
Remarks. — The present new subfamily Neocallianopsinae is distinguished
from the Callianopsinae Manning & Felder, 1991 by the absence of a dorsal
oval on the carapace.
Genus Neocallianopsis gen. nov.
Diagnosis. — Rostrum sharply triangular, bearing rostral carina. Carapace
bearing median longitudinal carina with cardiac prominence, extending from
rostral carina to posterior margin of carapace, and also bearing linea thalassinica, but without dorsal oval or cardiac sulcus. Eyestalks flattened and
contiguous; lacking cornea. A2 scaphocerite present as a small process. Mxp3
ischium-merus rectangular; propodus rectangular; dactylus ovate; exopod absent. P1 chelate and unequal in size, and dissimilar in shape; in larger cheliped
merus with small hook ventroproximally; palm rectangular; fingers shorter
than palm; in smaller cheliped merus usually with small hook ventroproximally; fingers longer than palm. P2 chelate. P3 propodus broadened. P4 simple. P5 subchelate. Abdominal pleura 2-4 unarmed ventrally; pleuron 5 with
spine-like projection posteroventrally; and abdominal somite 6 with spine-like
projection anteroventrally. Male Plp1 uniramous and bisegmented, distal segment sickle-shaped; male Plp2 endopod bearing mesiomedially appendix masculina with distal setae and small appendix interna proximally; male Plps3-5
similar, endopods with finger-like appendix interna. Telson subsquare. Uropodal exopod bearing neither dorsal plate nor lateral notch.
Type species. — Callianopsis anovalis Lin, Komai & Chan, 2007, by
present designation and monotypy. The gender of the generic name, Neocallianopsis, is feminine.
Species included. — Neocallianopsis anovalis (Lin, Komai & Chan, 2007).
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
483
Neocallianopsis anovalis (Lin, Komai & Chan, 2007)
Callianopsis anovalis Lin, Komai & Chan, 2007b: 1195, figs. 1, 2.
Material examined. — NTOU A00162, holotype, 1 male (TL/CL, 35.0/9.4 mm), Dasi fishing
port, Yilan County, Taiwan.
Remarks. — Lin et al. (2007b) described Callianopsis anovalis and included the species in the Ctenochelidae Manning & Felder, 1991, mentioning
that the carapace bears no dorsal oval, though Manning & Felder (1991) described in the definition of Callianopsis, of the Callianopsinae, that the carapace has a dorsal oval, as earlier described by Hopkins & Feldmann (1997:
237). It thus seems that things were confused and complicated, because apparently there has been some kind of misunderstanding. However, according to
the result of the present examination, in the genus Callianopsis the carapace
bears a dorsal oval, and is therefore included in the subfamily Callianopsinae,
in which the carapace bears a dorsal oval, although Lin et al. (2007b) described
that the carapace bears no dorsal oval in their Callianopsis anovalis; and in the
family Ctenochelidae the carapace bears no dorsal oval, though Manning &
Felder (1991) included the Callianopsinae (bearing a dorsal oval on the carapace) in the Ctenochelidae (which have no dorsal oval on the carapace). Therefore, Lin et al.’s (2007b) Callianopsis anovalis is not to be included in the
genus Callianopsis, for lack of a dorsal oval on the carapace, but is reclassified under the present new genus Neocallianopsis, which does not belong to
the Callianopsinae, but to the new subfamily Neocallianopsinae subfam. nov.
under the family Callianopsidae, instead of under the family Ctenochelidae.
Neocallianopsis anovalis from Taiwan is similar to Callianopsis goniophthalma (Rathbun, 1902) from off Point Conception, California, 513 m, in that
the Mxp3 dactylus is ovate, the male Plp1 is uniramous and bisegmented, with
the distal segment sickle-shaped; the male Plp2 is biramous, the endopod bears
an appendix masculina with distal setae and a small appendix interna proximally; and the uropodal exopod bears neither a dorsal plate nor a lateral notch.
However, they differ from each other in that in N. anovalis the carapace bears
no dorsal oval, but bears a median longitudinal carina with a cardiac prominence, extending from the rostral carina to the posterior margin of the carapace,
and the fixed finger of the larger cheliped bears no median tooth on the cutting
edge. While in C. goniophthalma the carapace bears a dorsal oval and a cardiac
prominence, but lacks a median longitudinal carina; and the fixed finger of the
larger cheliped bears a distinct triangular median tooth on the cutting edge.
Type locality. — Dasi fishing port, Yilan County, Taiwan.
Distribution. — Only known from the type locality.
FAMILY CTENOCHELIDAE MANNING & FELDER, 1991
Ctenochelinae Manning & Felder, 1991: 784; Poore, 1994: 103; Hendrickx, 1995: 387 (key),
fig. 8; Tudge et al., 2000: 135; Davie, 2002: 463.
Ctenochelidae — Sakai, 2005b: 235; Sakai & Sawada, 2006: 1358.
Diagnosis. — Rostrum acutely triangular and unarmed on lateral margins,
bearing rostral carina extending posteriorly to gastric region. Carapace bearing neither dorsal oval nor cardiac sulcus; cardiac prominence present or absent; and linea thalassinica present. Eyestalks flattened and contiguous; cornea
unpigmented. A2 scaphocerite strong. Maxilla 2 scaphognathite without posterior whip. Mxp3 ischium-merus rectangular; propodus rectangular; dactylus
digitiform; exopod present or not. P1 chelate and unequal in size, dissimilar in
shape; in larger cheliped merus without hook ventroproximally, palm subglobular, fingers elongate and pectinate; in smaller cheliped merus without hook
ventroproximally, fingers elongate but not pectinate. P2 chelate. P3 propodus
rectangular. Male Plp1 uniramous and bisegmented or four-segmented; male
Plp2 biramous and foliaceous, endopod with appendices interna and masculina
mesiodistally, and male Plp2 smaller than Plps3-5. Abdominal somites 3-5
with a tuft of setae. Telson subquadrate to trapezoid. Uropodal exopod with
lateral notch, but without dorsal plate.
Remarks. — Manning & Felder (1991: 784) established the family Ctenochelidae Manning & Felder, 1991, including three subfamilies, Ctenochelinae
Manning & Felder, 1991 (including 4 genera, Ctenocheles Kishinouye, 1926;
Dawsonius Manning & Felder, 1991; Gourretia De Saint Laurent, 1973;
Paracalliax De Saint Laurent, 1979), Anacalliacinae Manning & Felder, 1991
(including one genus Anacalliax De Saint Laurent, 1973), and Callianopsinae
Manning & Felder, 1991 (including one genus Callianopsis De Saint Laurent,
1973). However, it is found that the type genus Ctenocheles Kishinouye, 1926
in the subfamily Ctenochelinae Manning & Felder, is clearly different from
the other two type genera, i.e., Anacalliax of the subfamily Anacalliacinae
and Callianopsis of the subfamily Callianopsinae, in that the carapace bears
no dorsal oval and the male larger cheliped bears pectinate chelae. Therefore,
the Anacalliacinae and the Callianopsinae, which bear a dorsal oval on the
carapace and non-pectinate chelae in the male larger cheliped, are excluded
from the family Ctenochelidae.
486
CRM 013 – Katsushi Sakai
Type genus. — Ctenocheles Kishinouye, 1926.
Genera included. — Ctenocheles Kishinouye, 1926.
Genus Ctenocheles Kishinouye, 1926
?Pentacheles Balss, 1914: 75.
Thaumastocheles — Doflein, 1906: 522 (partim).
Ctenocheles Kishinouye, 1926: 63; Powell, 1949b: 369; Holthuis, 1967: 377; De Saint Laurent,
1973: 514; Le Loeuff & Intès, 1974: 24; Poore & Griffin, 1979: 277; De Saint Laurent & Le
Loeuff, 1979: 47 (key), 81; Coelho & Ramos-Porto, 1987: 29 (key), 31; Manning, 1987: 397
(list); Sakai, 1987a: 306 (list); Manning & Felder, 1991: 784, figs. 2, 7; Poore, 1994: 103
(key); Sakai, 1999a: 88; Davie, 2002: 464; Sakai, 2005b: 237.
Diagnosis. — Carapace with rostral carina with or without cardiac prominence, but without dorsal oval and transverse cardiac sulcus. Eyestalks flattened. Mxp3 usually without exopod, merus usually with spine on distal margin; propodus rectangular; and dactylus digitiform. P1 chelate, unequal, and
dissimilar; in larger cheliped merus with or without median tooth; palm subglobular; fingers elongate and pectinate. P2 chelate. P3 propodus rectangular. P4 simple. P5 subchelate. Male Plp1 uniramous and bisegmented or foursegmented; male Plp2 biramous and foliaceous, endopod with appendices interna and masculina. Female Plp1 uniramous and bisegmented; female Plp2
biramous, endopod with appendix interna. Plps3-5 different from Plps1-2 in
size and shape, bearing appendices internae. Telson subquadrate. Uropodal
exopod with lateral notch, lacking dorsal plate. [Adapted from Sakai, 2005b.]
Type species. — Ctenocheles balssi Kishinouye, 1926, by original designation and monotypy. Gender of generic name, Ctenocheles, masculine.
Species included. — Ctenocheles balssi Kishinouye, 1926; C. collini Ward,
1945; C. holthuisi Rodrigues, 1978; C. leviceps Rabalais, 1979; C. serrifrons
Le Loeuff & Intès, 1974; C. maorianus Powell, 1949a; C. “A”, Holthuis, 1967;
C. “B”, Holthuis, 1967; C. sp., De Saint Laurent & Le Loeuff, 1979; C. sp.
Coelho, 1997.
K EY
TO THE
S PECIES
OF THE
G ENUS CTENOCHELES
1 – Uropods slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. maorianus
– Uropods broadened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Cardiac prominence present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Cardiac prominence absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 – Male Plp1 bisegmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. serrifrons
– Male Plp1 four-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. balssi
4 – Merus of larger cheliped with median tooth on ventral margin . . . . . . . . . . . . C. holthuisi
– Merus of larger cheliped unarmed on ventral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
487
5 – Ischium of larger cheliped smooth on ventral margin . . . . . . . . . . . . . . . . . . . . . C. leviceps
– Ischium of larger cheliped serrated on ventral margin . . . . . . . . . . . . . . . . . . . . . . C. collini
Ctenocheles balssi Kishinouye, 1926
Thaumastocheles Doflein, 1906: 522.
?Pentacheles sp. nov.? — Balss, 1914: 75, fig. 43.
Ctenocheles balssi Kishinouye, 1926: 63, fig. 1; Yokoya, 1933: 55; Makarov, 1938: 76, fig. 29;
Holthuis, 1967: 377; Suzuki, 1979: 296, pl. 18 fig. 234; Sakai, 1987a: 306 (list); Manning
& Felder, 1991: 784 (list); Noguchi & Akamine, 1992: 25, fig. 1; Liu & Zhong, 1994: 562
(list); Matsuzawa & Hayashi, 1997: 39, 42 (in key), figs. 1-3; Sakai, 1999a: 88, figs. 1a-e,
2a-g, 3a-g; Komai, 2000b: 346 (list); Tudge et al., 2000: 142; Sakai, 2005b: 240; Sakai &
Sawada, 2006: 1358, figs. 13, 14.
Ctenocheles Balssi — De Man, 1928b: 25 (list).
Diagnosis. — Cardiac prominence present. Mxp3 exopod absent. In larger
cheliped ischium and merus both smooth on ventral margin. Male Plp1 foursegmented. Uropods broadened.
Type locality. — Ohsu near Kashiwasaki, Niigata Prefecture, Japan.
Distribution. — Japan, East China Sea to South China Sea (Liu & Zhong,
1994: 562).
Ctenocheles collini Ward, 1945
Ctenocheles collini Ward, 1945: 134, pl. 13; Holthuis, 1967: 377; Poore & Griffin, 1979: 277,
fig. 37; Manning & Felder, 1991: 784 (list); Matsuzawa & Hayashi, 1997: 45 (in key); Sakai,
1999a: 88 (list); Tudge et al., 2000: 142; Sakai, 2005b: 241, figs. 43, 44.
Diagnosis. — Cardiac prominence absent. Mxp3 exopod absent. In larger
cheliped ischium finely serrated on ventral margin; merus unarmed on ventral
margin. Uropods broadened.
Type locality. — Moreton Bay, Queensland, Australia.
Distribution. — Moreton Bay, Queensland, Australia (Ward, 1945; Poore &
Griffin, 1979); 15-40 m.
Ctenocheles holthuisi Rodrigues, 1978
Ctenocheles holthuisi Rodrigues, 1978: 113, figs. 1-21; Coelho & Ramos-Porto, 1987: 31; Manning, 1987: 397 (list); Manning & Felder, 1991: 784 (list); Coelho, 1997: 150; Matsuzawa
& Hayashi, 1997: 45 (in key); Rodrigues et al., 1998: 382; Melo, 1999: 392, figs. 263-264;
Sakai, 1999a: 88 (list); Tudge et al., 2000: 142; Sakai, 2005b: 239; Coelho et al., 2007: 5
(tab. 2).
Diagnosis. — Cardiac prominence absent. Mxp3 ischium-merus pediform;
merus armed with a strong distomedian spine; exopod vestigial. In larger
cheliped ischium serrated on ventral margin; merus with median tooth on
488
CRM 013 – Katsushi Sakai
ventral margin. Plps1-2 unknown. Telson pentagonal, and convex on posterior
margin; uropods broadened. [Cf. Rodrigues, 1978: 113, figs. 1-10, 1-21.]
Type locality. — Off mouth of Rio São Francisco, Brazil, 10◦ 37′ 09′′ S
36◦ 14′ 00′′ W, 75 m.
Distribution. — Brazil — Alagoas (Coelho, 1997); off mouth of Rio São
Francisco (Rodrigues, 1978; Coelho & Ramos-Porto, 1987); 75 m.
Ctenocheles leviceps Rabalais, 1979
Ctenocheles leviceps Rabalais, 1979: 295, figs. 2-29; Rabalais et al., 1981: 100; Manning, 1987:
397 (list); Williams et al., 1989: 28; Manning & Felder, 1991: 784 (list), fig. 7; Matsuzawa
& Hayashi, 1997: 45 (in key); Sakai, 1999a: 88 (list); Tudge et al., 2000: 142; Sakai, 2005b:
239.
Diagnosis. — Cardiac prominence absent. Mxp3 exopod vestigial. In larger
cheliped ischium and merus unarmed on ventral margin, respectively.
Uropods broadened.
Type locality. — Port Aransas, Texas, U.S.A., 27◦ 38′ N 96◦ 41′ W.
Distribution. — Texas — Port Aransas (Rabalais, 1979); Gulf of Mexico
(Rabalais et al., 1981); 10-49 m.
Ctenocheles maorianus Powell, 1949
Ctenocheles maorianus Powell, 1949a: 408; Powell, 1949b: 369, pl. 68 figs. 3-7; Dell, 1956:
149; Holthuis, 1967: 378; Manning & Felder, 1991: 784 (list); Dworschak, 1992: 209, figs.
12a-b, 13a-c; Matsuzawa & Hayashi, 1997: 45 (in key); Sakai, 1999a: 88 (list); Tudge et al.,
2000: 142; Sakai, 2005b: 243.
Diagnosis. — Rostrum very short and acute, reaching proximal third of
eyestalks. Uropods slender. [Adapted from Dworschak, 1992.]
Type locality. — Hauraki Gulf, New Zealand.
Distribution. — New Zealand — Hauraki Gulf, Tiritiri Island, off Plate
Island, Bay of Plenty and Tasman Bay (Powell, 1949a); off Cape Runaway,
off Kaikoura, Marlborough, Castlecliffian (upper Pleistocene) near Castlecliff
(Holthuis, 1967); Little Town (Dworschak, 1992); 35-73 m.
Ctenocheles serrifrons Le Loeuff & Intès, 1974
Ctenocheles sp. Crosnier, 1969: 536, fig. 18.
Ctenocheles serrifrons Le Loeuff & Intès, 1974: 24, fig. 3a-u; De Saint Laurent & Le Loeuff,
1979: 83; Manning & Felder, 1991: 784 (list), fig. 2; Matsuzawa & Hayashi, 1997: 42 (in
key); Sakai, 1999a: 88 (list); Tudge et al., 2000: 142; Sakai, 2005b: 238.
Diagnosis. — Cardiac prominence present. Mxp3 exopod vestigial. Larger
cheliped unknown. Male Plp1 bisegmented. Uropods broadened.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
489
Type locality. — Ivory Coast, 5◦ 2′ N 5◦ 4.5′ W; 50 m.
Distribution. — Only known from the type locality.
Ctenocheles “A”, Holthuis, 1967
Ctenocheles “A”, Holthuis, 1967: 379, figs. 1, 2a; Manning, 1987: 397 (list); Manning & Felder,
1991: 784 (list); Sakai, 1999a: 88; Sakai, 2005b: 240.
Distribution. — Strait of Florida; 109-406 m (Holthuis, 1967).
Ctenocheles “B”, Holthuis, 1967
Ctenocheles “B”, Holthuis, 1967: 382, fig. 2b; Manning, 1987: 398 (list); Manning & Felder,
1991: 784 (list); Sakai, 1999a: 88; Sakai, 2005b: 240.
Distribution. — Colombia (Holthuis, 1967).
Ctenocheles sp. De Saint Laurent & Le Loeuff, 1979
Ctenocheles sp., De Saint Laurent & Le Loeuff, 1979: 83, fig. 25; Sakai, 1999a: 88 (list); Sakai,
2005b: 239.
Distribution. — Gabon; Benin; 48-110 m (De Saint Laurent, 1979).
Ctenocheles sp. Coelho, 1997
Ctenocheles sp., Coelho, 1997: 150.
Distribution. — Amapá and Rio Grande do Sul, Brazil.
FAMILY EUCALLIACIDAE MANNING & FELDER, 1991
(SENSU NOV.)
Eucalliinae Manning & Felder, 1991: 781 [misspelling].
Eucalliacinae — Sakai, 1999c: 108; Ngoc-Ho, 2003: 487; Sakai, 2005b: 195.
Diagnosis. — Carapace with or without dorsal oval. Maxilla 2 scaphognathite without posterior whip. Mxp3 propodus broadened; dactylus subtriangular or digitiform. P1 chelate, unequal, and dissimilar, lacking meral hook
in both chelipeds. Uropodal exopod with dorsal plate but without lateral notch.
Subfamilies included. — Calliapaguropinae Sakai, 1999c; Eucalliacinae
Manning & Felder, 1991.
K EY
TO THE
S UBFAMILIES
OF THE
FAMILY E UCALLIACIDAE
1 – Mxp3 dactylus subovate or subtriangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucalliacinae
– Mxp3 dactylus digitiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calliapaguropinae
Subfamily C ALLIAPAGUROPINAE Sakai, 1999
Calliapaguropinae Sakai, 1999c: 7; Sakai, 2005b: 205.
Diagnosis. — Rostrum narrowly elongate, lacking rostral carina. Carapace
without dorsal oval (in sensu Ngoc-Ho, 2002a, with faint dorsal oval), cardiac prominence, or hepatic sulcus. Eyestalks cylindrical and set apart. Abdominal somite 6 lacking lateral projections. A2 peduncle remarkably thick;
A2 scaphocerite present as a small process. Maxilla 2 scaphognathite without
posterior whip. Mxp3 ischium-merus suboperculiform, bearing meral teeth;
propodus subtriargular; dactylus digitiform. P1 chelate, unequal, and dissimilar, lacking meral hook in both chelipeds. P2 chelate. P3 propodus broadened. Ps4-5 chelate. Male Plp1 uniramous, slender, and bisegmented. Male
Plp2 biramous and blade-like, endopod with appendix interna. Female Plp2
biramous and blade-like, endopod with appendix interna. Male and female
Plps1-2 smaller than Plps3-5. Plps3-5 biramous and foliaceous, endopods with
appendix interna in both sexes. Uropodal exopod with dorsal plate. [Cf. Sakai,
2005b: 206.]
Remarks. — Ngoc-Ho (2003: 486) included Calliapagurops in the Callichirinae sensu Tudge et al. (2000). However, Calliapagurops is different
492
CRM 013 – Katsushi Sakai
from Callichirus, type genus of the Callichirinae. In the type species of Calliapagurops, C. charcoti De Saint Laurent, 1973, the carapace bears no dorsal
oval, the eyestalks are cylindrical and set apart, and the corneae are present
distally; the male Plp2 endopod bears mesiodistally a small appendix interna.
Whereas in the type species of Callichirus, C. major (Say, 1818), the carapace
bears a dorsal oval, the eyestalks are flattened and contiguous; and the male
Plp2 endopod bears neither an appendix interna nor an appendix masculina.
As a result, Calliapagurops is not included in the Callichirinae, but in the Calliapaguropinae.
Type genus. — Calliapagurops De Saint Laurent, 1973.
Genus included. — Calliapagurops De Saint Laurent, 1973 is the only
known genus so far.
Genus Calliapagurops De Saint Laurent, 1973
Calliapagurops De Saint Laurent, 1973: 515; Sakai, 1999c: 8; Ngoc-Ho, 2002a: 540; Sakai,
2005b: 207.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment
pointed distally; male Plp2 biramous and blade-like, endopod with appendix
interna mesiodistally, but without appendix masculina.
Type species. — Calliapagurops charcoti De Saint Laurent, 1973, by
original designation. Gender of the generic name, Calliapagurops, masculine.
Species included. — Calliapagurops charcoti De Saint Laurent, 1973;
C. foresti Ngoc-Ho, 2002a.
K EY
TO THE
S PECIES
OF THE
G ENUS CALLIAPAGUROPS
1 – Mxp3 merus with 3-4 spines on distal margin; eyestalks not narrowing distally, corneas
as wide as proximal part of eyestalks, overreaching distal end of A1 penultimate
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. charcoti
– Mxp3 merus with 6-10 spines on distal margin; eyestalks narrowing distally, corneas
small, not reaching distal end of A1 penultimate segment . . . . . . . . . . . . . . . . . . C. foresti
Calliapagurops charcoti De Saint Laurent, 1973
Calliapagurops charcoti De Saint Laurent, 1973: 515; Manning & Felder, 1991: 771; Sakai,
1999c: 8, fig. 1a-e; d’Udekem d’Acoz, 1999: 155; Tudge et al., 2000: 143; Costello et al.,
2001: 289; Türkay, 2001: 289; Ngoc-Ho, 2002a: 540; Ngoc-Ho, 2003: 487, fig. 16; Sakai,
2005b: 207.
Diagnosis. — Eyestalks not narrowing distally; corneas as wide as proximal
part of eyestalks, overreaching distal end of A1 penultimate segment. Mxp3
merus with 3-4 spines on distal margin.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
493
Type locality. — Off Azores Is. near Flores Islands, Sta. 109, 39◦ 33′ N
31◦ 17′ W, 190-230 m; Expedition “Biacores”, 1971, R/V “Jean Charcot”.
Distribution. — Only known from the type locality.
Calliapagurops foresti Ngoc-Ho, 2002
Calliapagurops foresti Ngoc-Ho, 2002a: 539, figs. 1-3; Sakai, 2005b: 208.
Diagnosis. — Eyestalks narrowing distally; corneas small, not reaching
distal end of A1 penultimate segment. Mxp3 merus with 6-10 spines on distal
margin.
Remarks. — Ngoc-Ho (2002a: 546) mentioned two differentiating characters: (1) the eyestalks are narrowing distally, and the corneas are small in
C. foresti (vs. the eyestalks are not narrowing distally, and the corneas are as
wide as the proximal part of the eyestalks in C. charcoti); (2) the Mxp3 merus
bears 3-10 spines on the distal margin in C. foresti (vs. 3-4 spines on that margin in C. charcoti).
Type locality. — Philippines, MUSORSTOM 2, Sta. 71, 14◦ 00.1′ N
120◦ 17.8′ E, 192 m.
Distribution. — Only known from the type locality.
Subfamily E UCALLIACINAE Manning & Felder, 1991 (sensu nov.)
Eucalliinae Manning & Felder, 1991: 781 (misspelling).
Eucalliacinae — Sakai, 1999c: 108; Ngoc-Ho, 2003: 487; Sakai, 2005b: 195.
Diagnosis. — Rostrum developed or reduced, lacking rostral carina. Carapace with or without dorsal oval, and with or without cardiac prominence and
cardiac sulcus(i). Eyestalks flattened and contiguous. A2 scaphocerite developed as a small process. Maxilla 2 scaphognathite without posterior whip.
Mxp3 ischium-merus subpediform; propodus subrectangular or subquadrate;
dactylus subtriangular or subovate; exopod present or not. P1 chelate, unequal
or subequal in size, and similar or dissimilar in shape, lacking meral hook
in both chelipeds. P2 chelate. P3 propodus broadened. P4 simple. P5 chelate.
Abdominal somite 6 without lateral projections. Male Plp1 present; male Plp2
biramous, endopod with appendices masculina and interna, or with either appendix interna, or appendix masculina. Plps3-5 biramous and foliaceous, endopods bearing appendix interna in both sexes. Uropodal endopod oval or subtriangular; uropodal exopod with dorsal plate. [Adapted from Sakai, 2005b.]
Remarks. — In the previous revision (Sakai, 2005b), Calliax De Saint
Laurent, 1973 was stated to be the type genus of the Eucalliacinae, because
494
CRM 013 – Katsushi Sakai
the type genus of the Eucalliacinae, Eucalliax Manning & Felder, 1991 was
considered to be a junior synonym of Calliax on account of its similarity to
Calliax in the shapes of male Plp2 and Mxp3. However, it has turned out
that they are different from each other in the shapes of male Plp1, telson, and
uropodal endopod, so that Eucalliax is safely defined again as the type genus
of the Eucalliacinae.
The genus Calliaxina Ngoc-Ho, 2003 was established by designating
C. punica as the type species, which is characterized by the presence of the
Mxp3 exopod, and including two other species. However, such a characteristic is not of vital importance in distinguishing the genera, but, in contrast,
the presence of (a) cardiac sulcus(i) on the carapace is considered to be much
more important in the classification of the genera. In the present revision, the
classification made by applying the presence of (a) sulcus(i) proposes that Calliaxina Ngoc-Ho, 2003, until now including three species, should be expanded,
now including 8 species and thus presenting a new concept.
Type genus. — Eucalliax Manning & Felder, 1991, by original designation,
and as the nominate genus.
Genera included. — Andamancalliax gen. nov.; Calliax De Saint Laurent, 1973; Calliaxina Ngoc-Ho, 2003 (sensu nov.); Eucalliax Manning &
Felder, 1991; Eucalliaxiopsis gen. nov.; Paraglypturus Türkay & Sakai, 1995;
Pseudocalliax gen. nov.
K EY
TO THE
G ENERA
OF THE
S UBFAMILY E UCALLIACINAE
1 – Cardiac sulcus present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calliaxina sensu nov.
– Cardiac sulcus absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Mxp3 exopod present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Mxp3 exopod absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 – Male Plp1 small and simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudocalliax gen. nov.
– Male Plp1 uniramous and bisegmented, distal segment chelate . . . . . . . . . Paraglypturus
4 – Rostrum obscurely present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Rostrum triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 – Male Plp1 distal segment simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucalliax
– Male Plp1 distal segment sickle-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calliax
6 – Male Plp1 distal segment emarginate distally . . . . . . . . . . . . . . Andamancalliax gen. nov.
– Malpe Plp1 distal segment strongly incurvate distally . . . . . . . . Eucalliaxiopsis gen. nov.
Genus Andamancalliax gen. nov.
Diagnosis. — Carapace without dorsal oval. Cardiac sulcus absent. Mxp3
ischium-merus rectangular; propodus subquadrate, and rounded ventrally;
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
495
dactylus subtriangular and truncate distally, with thick setae; exopod absent.
Male Plp1 uniramous and bisegmented, distal segment shorter than proximal
one, and emarginate distally; male Plp2 biramous, endopod bearing mesiodistally appendix masculina with long apical setae, but no appendix interna. Telson subsquare. Uropodal exopod with dorsal plate.
Type species. — Calliax andamanica Sakai, 2002, by present designation
and monotypy. The gender of the generic name, Andamancalliax, is feminine.
Species included. — Andamancalliax andamanica (Sakai, 2002).
Etymology. — The new genus is named after the type locality of the type
species by adding Andaman as a prefix to Calliax.
Andamancalliax andamanica (Sakai, 2002)
Calliax andamanica Sakai, 2002: 463, figs. 1A-G, 2A-D; Sakai, 2005b: 201.
Diagnosis. — Rostrum triangular in dorsal view, reaching distal part of
eyestalks. Eyestalks with cornea distally. Mxp3 ischium-merus rectangular;
propodus subquadrate, and rounded ventrally; dactylus subtriangular and
longer than propodus, and truncate distally, with thick setae; exopod absent.
P1 unequal. Male Plp1 uniramous and bisegmented, distal segment shorter
than proximal one, and emarginate distally; male Plp2 biramous, endopod
bearing mesiodistally distinct appendix masculina with long apical setae, but
no appendix interna. Telson subsquare. Uropodal exopod with dorsal plate.
[Cf. Sakai, 2002: 463, figs. 1D, C, 2A.]
Type locality. — Andaman Sea, 9◦ 30.364′ N 97◦ 57.346′ E, 42.5 m, sand with
shell fragments.
Distribution. — Andaman Sea (7◦ 44.638′ N 98◦ 16.496′ E; 7◦ 45.002′ N
98◦ 15.103′ E; 9◦ 30.364′ N 97◦ 57.346′ E, 30.5-70.0 m, coarse and fine sand,
mud, sandy mud, with shell fragments).
Genus Calliax De Saint Laurent, 1973
Calliax De Saint Laurent, 1973: 514; Manning, 1987: 397; Sakai, 1987a: 306; Sakai, 1988: 61;
Manning & Felder, 1991: 783, figs. 3, 15f-j; Poore, 1994: 101; Sakai, 1999c: 109; Davie,
2002: 459; Sakai, 2005b: 196.
Diagnosis. — Carapace without dorsal oval and cardiac sulcus. Maxilla 2
scaphognathite without posterior whip. Mxp3 ischium-merus rectangular;
propodus subtriangular or subquadrate dactylus subtriangular and truncate
distally, with thick setae; exopod absent. P1 unequal in size and dissimilar in
shape. Male Plp1 uniramous and bisegmented, distal segment sickle-shaped;
496
CRM 013 – Katsushi Sakai
male Plp2 biramous, endopod mesiodistally with stout appendix masculina
exceeding distal margin of endopod; appendix interna present or absent. Telson
convex distally. Uropodal endopod oval and much longer than telson; uropodal
exopod with dorsal plate. [Cf. Ngoc-Ho, 2003: 490, figs. 17, 18.]
Type species. — Calliax lobata De Gaillande & Lagardère, 1966, by
original designation. The gender of the generic name, Calliax, is feminine.
Species included. — Calliax doerjesti Sakai, 1999c; Calliax lobata De
Gaillande & Lagardère, 1966.
K EY
TO THE
S PECIES
OF THE
G ENUS CALLIAX
1 – Eyestalks rounded distally; telson broader than long, narrowing in posterior half toward
straight posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. doerjesti
– Eyestalks triangular distally; telson broader than long, slightly narrowing laterally
toward rounded posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lobata
Calliax doerjesti Sakai, 1999
Calliax doerjesti Sakai, 1999c: 112, figs. 27d-g, 28, 29a-f; Sakai, 2005b: 199.
Diagnosis. — Carapace without dorsal oval or cardiac sulci. Mxp3 ischiummerus subrectangular; merus subtriangular; propodus subquadrate; dactylus
subtriangular and truncate distally, with thick setae; exopod absent. P1 unequal. Male Plp1 uniramous and bisegmented, distal segment curved distally,
with pointed tip; male Plp2 biramous, endopod mesiomedially with stout appendix masculina exceeding distal margin of endopod; appendix interna absent. Telson subrectangular, posterior margin broadly convex, lacking median
spine. Uropodal exopod with dorsal plate. [Cf. Sakai, 1999c: 112, figs. 27g,
28, 29e.]
Type locality. — Florida Bay, Key Largo, U.S.A.
Distribution. — Only known from the type locality.
Calliax lobata (De Gaillande & Lagardère, 1966)
[Abbreviated list of synonymy.]
Callianassa (Callichirus) lobata De Gaillande & Lagardère, 1966: 259, pls. 1-4.
Calliax lobata — De Saint Laurent & Božić, 1976: 28, figs. 7, 15, 23, 27, 34; Sakai, 1999c:
110, fig. 27a-c; Sakai, 2000: 110, figs. 27a-c; Costello et al., 2001: 289; Ngoc-Ho, 2003: 490,
figs. 17, 18; Sakai, 2005b: 197.
Diagnosis. — Carapace without dorsal oval or cardiac sulcus. Maxilla 2
scaphognathite without posterior whip. Mxp3 ischium-merus subrectangular;
merus subtriangular; propodus subtriangular; dactylus subtriangular; exopod
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
497
absent. Male Plp1 uniramous and bisegmented, distal segment curved distally,
with pointed tip; male Plp2 biramous, endopod bearing mesiomedially distinct
appendix masculina extending over distal margin of endopod, and small and
slender appendix interna. Telson slightly wider than long and rounded on
posterior margin. Uropodal exopod with dorsal plate. [Cf. Sakai, 1999c, fig.
27b.]
Type locality. — Port Miou, Toulon, France, Mediterranean; 2-8 m.
Distribution. — Port Miou, Toulon, France, Mediterranean, 2-8 m (de
Gaillande & Lagardère, 1966); Patraikos Gulf, Greece, 15 m, sandy mud
(Thessalou-Legaki, 1985, 1986).
Genus Calliaxina Ngoc-Ho, 2003 (sensu nov.)
Calliaxina Ngoc-Ho, 2003: 493.
Diagnosis. — Carapace without dorsal oval, but with cardiac sulcus (or
sulci). Mxp3 ischium-merus subrectangular; propodus subtriangular or approximately so; dactylus subovate or subtriangular; exopod present or absent.
P1 subequal. Male Plp1 uniramous and bisegmented, distal segment simple
or subchelate distally, with or without appendix inerna; male Plp2 biramous,
endopod bearing mesiomedially appendix masculina with apical setae, and a
small appendix interna. Telson straight on posterior margin. Uropodal endopod
oval and much longer than telson; uropodal exopod with dorsal plate.
Remarks. — Ngoc-Ho (2003: 487) established the genus Calliaxina NgocHo, 2003 by designating C. punica De Saint Laurent & Manning, 1982, as the
type species, mentioning that “This work considers the presence of an exopod
on the Mxp3 an important taxonomic character”, and included two other
species (Calliaxina novaebritanniae and C. sakaii) in Calliaxina. However,
in the present revision 5 species (Calliax jonesi Heard, 1989, C. aequimana
(Baker, 1907), C. bulimba (Poore & Griffin, 1979), Eucalliax mcilhennyi
Felder & Manning, 1994, and the recent species, E. panglaoensis (Dworschak,
2006)) are also included in Calliaxina, because they have in common with one
another such characteristics as the presence of a cardiac sulcus or sulci, and the
typical forms of Mxp3 ischium-merus, propodus, and dactylus, though four of
them have no Mxp3 exopod.
Type species. — Calliax punica De Saint Laurent & Manning, 1982, by
original designation. The gender of the generic name, Calliaxina, is feminine.
Species included. — Calliaxina aequimana (Baker, 1907); C. bulimba
(Poore & Griffin, 1979); C. jonesi (Heard, 1989); C. mcilhennyi (Felder
& Manning, 1994); C. novaebritanniae (Borradaile, 1900); C. panglaoensis
498
CRM 013 – Katsushi Sakai
(Dworschak, 2006); C. punica De Saint Laurent & Manning, 1982; C. sakaii
De Saint Laurent & Le Loeuff, 1979.
K EY
TO THE
S PECIES
OF THE
G ENUS CALLIAXINA
1 – Mxp3 exopod absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Mxp3 exopod present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 – Male Plp1 uniramous and bisegmented, distal segment with median lobe . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aequimana
– Male Plp1 uniramous and bisegmented, distal segment subchelate . . . . . . . . . . . . . . . . . . 3
3 – Male Plp1 longer branch of subchela incurvate distally and shorter one bilobed distally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. jonesi
– Male Plp1 longer branch of subchela incurvate distally and shorter one simple distally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4 – Male Plp1 longer branch of subchelate distal segment obtuse distally . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. panglaoensis
– Male Plp1 longer branch of subchelate distal segment pointed distally . . . C. mcilhennyi
5 – Mxp3 exopod rudimentary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bulimba
– Mxp3 exopod distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 – A1 peduncle reaching distal margin of A2 penultimate segment . . . . . . . . . . . . C. punica
– A1 peduncle overreaching distal margin of A2 penultimate segment . . . . . . . . . . . . . . . . 7
7 – Mxp3 exopod elongate, overreaching ischium . . . . . . . . . . . . . . . . . . . . C. novaebritanniae
– Mxp3 exopod short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sakaii
Calliaxina aequimana (Baker, 1907)
Callianassa aequimana Baker, 1907: 182-185, pl. 24 figs. 1-8; Hale, 1927a: 87, fig. 83; Poore
& Griffin, 1979: 245, figs. 12, 13.
Callianassa (Callichirus) aequimana — De Man, 1928b: 28, 93, 114.
Callianassa (Callichirus) Novae-britanniae — De Man, 1928b: 29 (partim). [Not Callianassa
novae-britanniae Borradaile, 1900.]
Callianassa (Callichirus) novae-britanniae var. — De Man, 1928b: 114.
Calliax aequimana — De Saint Laurent & Manning, 1982: 222; Sakai, 1988: 222; Sakai, 1999c:
118, fig. 31a-e; Tudge et al., 2000: 145; Davie, 2002: 459; Sakai, 2005b: 202.
Diagnosis. — Carapace without dorsal oval, but with cardiac sulcus. Mxp3
ischium-merus subovate; propodus subtriangular and broader proximally than
distally; dactylus subovate; exopod absent. P1 subequal, in larger cheliped
ischium denticulate on ventral margin. Male Plp1 uniramous and bisegmented,
distal segment with median lobe, and obtuse distally; male Plp2 biramous,
endopod bearing mesiomedially distinct appendix masculina with a small
appendix interna subproximally. Telson wider than long and slightly concave
on posterior margin. Uropodal exopod with dorsal plate. [Cf. Poore & Griffin,
1979: 245, figs. 12c, 13f, i, j.]
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
499
Remarks. — Poore & Griffin (1979: 257) mentioned that “This species (Callianassa bulimba Poore, 1979) is closely related to C. aequimana common in
southern Australia”. However, those two species are different. In C. aequimana
the rostrum is reduced, the cardiac sulcus is incomplete, and Mxp3 lacks an
exopod; whereas in C. bulimba the rostrum is triangular, the cardiac sulcus is
complete, and Mxp3 bears a rudimentary exopod.
Type locality. — Kingston, in the south of Western Australia.
Distribution. — Intertidal and subtidal mudflats down to 9 m, often in
estuaries; southern Queensland through N.S.W., Victoria, South Australia, to
the south of Western Australia (Poore & Griffin, 1979).
Calliaxina bulimba (Poore & Griffin, 1979)
Callianassa bulimba Poore & Griffin, 1979: 257, fig. 21.
Calliax bulimba — De Saint Laurent & Manning, 1982: 222; Sakai, 1999c: 119, fig. 32a-c;
Sakai, 2005b: 202.
Diagnosis. — Rostrum triangular, reaching proximal third of eyestalks.
Eyestalks contiguous except for distal part shaped as triangle; cornea distinct
and located at distal third. Mxp3 propodus subquadrate, with three proximal
teeth; dactylus subovate; exopod rudimentary. Telson widest on proximal
margin, slightly narrowing and curving to straight posterior margin. [Adapted
from Poore & Griffin, 1979.]
Type locality. — Mud Island, Moreton Bay, Queensland Australia.
Distribution. — Only known from the type locality.
Calliaxina jonesi (Heard, 1989)
Calliax jonesi Heard, 1989: 129, figs. 1-5; Sakai, 1999c: 116; Sakai, 2005b: 200.
Eucalliax jonesi — Tudge et al., 2000: 145.
Diagnosis. — Carapace without dorsal oval, but with cardiac sulcus. Mxp3
ischium-merus subrectangular; merus triangular; propodus subtriangular;
dactylus subtriangular; exopod absent. Male Plp1 uniramous and bisegmented,
distal segment subchelate distally, longer branch of subchela incurvate distally and shorter one bilobed distally; male Plp2 biramous, endopod bearing
mesiodistally distinct appendix masculina with a small, proximally-fused appendix interna subproximally. Telson subrectangular, weakly trilobate on posterior margin. Uropodal exopod with dorsal plate. [Cf. Heard, 1989: 129, figs.
3E, 4F, 5A, C.]
Remarks. — The present species was included in Calliax, but it belongs to
Calliaxina, because in Calliax the carapace bears no cardiac sulcus, and Plp1
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is not subchelate but sickle-shaped distally; whereas in Calliaxina the carapace
bears a cardiac sulcus, and Plp1 is subchelate distally.
Type locality. — Bimini Harbor, Bahamas, 25◦ 44′ N 79◦ 15′ W, 3-5 m.
Distribution. — Only known from the type locality.
Calliaxina mcilhennyi (Felder & Manning, 1994)
Eucalliax mcilhennyi Felder & Manning, 1994: 341, figs. 1-6.
Calliax mcilhennyi — Sakai, 1999c: 116; Sakai, 2005b: 200.
Diagnosis. — Carapace without dorsal oval, but with cardiac sulcus. Mxp3
ischium-merus subovate; merus subtriangular; propodus subtriangular; dactylus subtriangular; exopod absent. Male Plp1 uniramous and bisegmented, distal segment subchelate distally; longer branch incurvate and truncate distally,
and shorter one simple distally; male Plp2 biramous, endopod bearing mesiomedially a distinct appendix masculina with a small appendix interna subproximally. Telson wider than long, posterior margin slightly concave. Uropodal
exopod with dorsal plate. [Cf. Felder & Manning, 1994: 341, figs. 1-6.]
Remarks. — The present species, Calliaxina mcilhennyi, which was included in Calliax, is reclassified here under Calliaxina, because it bears a cardiac sulcus on the carapace.
Type locality. — Indian River lagoon, St. Lucie County (27◦ 27.7′ N
◦
80 18.7′ W), Florida, sandflat with sparse seagrass, south side of Fort Pierce
Inlet.
Distribution. — Santa Marta, Colombia; Indian River lagoon, St. Lucie
County, Florida.
Calliaxina novaebritanniae (Borradaile, 1900)
[Abbreviated list of synonymy.]
Callianassa novae-britanniae Borradaile, 1900: 419, pl. 39 fig. 14a-d.
Calliax novaebritanniae — De Saint Laurent & Manning, 1982: 211-224, figs. 1c, 2b, 6c; Sakai,
2005b: 202.
Paraglypturus novaebritanniae — Sakai, 1999c: 123, fig. 32d-f.
Calliaxina novaebritanniae — Ngoc-Ho, 2003: 493.
Diagnosis. — Carapace without dorsal oval, but with cardiac sulcus. Mxp3
ischium-merus subovate; merus subquadrate; propodus subtriangular; dactylus
subtriangular, distal margin truncate, with setae; exopod long, overreaching
ischium. Male Plp1 uniramous and bisegmented, distal segment subchelate,
with appendix interna; Plp2 biramous, endopod bearing appendix masculina
with a small appendix interna. Telson wider than long, and posterior margin
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
501
almost straight. Uropodal exopod with dorsal plate. [Cf. De Man, 1928a: 49;
Sakai, 1999c: 123, fig. 32c, f.]
Remarks. — The present species is included in Calliaxina by the form of
the male Plps1-2 and the presence of a cardiac sulcus.
Type locality. — New Britain.
Distribution. — New Britain, Papua New Guinea.
Calliaxina panglaoensis (Dworschak, 2006)
Eucalliax panglaoensis Dworschak, 2006: 349, figs. 1-7.
Diagnosis. — Rostrum without rostral carina. Carapace without dorsal
oval, but with cardiac sulcus. Mxp3 ischium-merus rectangular; propodus
subtriangular; dactylus subovate; exopod absent. Male Plp1 uniramous and
bisegmented, distal segment subchelate, longer branch and shorter branch both
obtuse distally; male Plp2 biramous, endopod bearing mesiomedially a distinct
appendix masculina and a small appendix interna subproximally. Telson about
1.6 times as wide as long, and posterior margin straight. Uropodal exopod with
dorsal plate. [Quoted from Dworschak, 2006.]
Remarks. — The present species Calliaxina panglaoensis (Dworschak,
2006) was described as a species of the genus Eucalliax. However, it is not
included in Eucalliax, but in Calliaxina, because in Eucalliax the carapace
bears no cardiac sulcus, whereas in Calliaxina it bears a cardiac sulcus.
Type locality. — Alona Beach, Panglao Isl., Bohol, Philippines, 09◦ 32.9′ N
123◦ 46.6′ E, intertidal.
Distribution. — Only known from the type locality.
Calliaxina punica (De Saint Laurent & Manning, 1982)
[Abbreviated list of synonymy.]
Calliax punica De Saint Laurent & Manning, 1982: 211-224, figs. 1-6; Sakai, 2005b: 198; Sakai
& Sawada, 2006: 1357.
Paraglypturus punica — Sakai, 1999c: 122; Costello et al., 2001: 289; Sakai, 2005b: 205.
Calliaxina punica — Ngoc-Ho, 2003: 496, figs. 19, 20.
Diagnosis. — Carapace without dorsal oval. Cardiac sulcus present. Mxp3
ischium-merus subovate; merus subquadrate; propodus subquadrate; dactylus
subquadrate; exopod distinct. Male Plp1 uniramous and bisegmented, distal
segment subchelate distally, exceptionally bearing appendix interna mesiomedially; male Plp2 biramous, endopod bearing mesiomedially a distinct appendix masculina with a small appendix interna subproximally. Telson wider than
long. Uropodal exopod with dorsal plate.
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Remarks. — The present species is included neither in the genus Paraglypturus, nor in the genus Calliax, in neither of which the carapace bears any
sulcus, but instead in Calliaxina, in which the carapace does bear a sulcus.
Type locality. — Gulf of Tunisia.
Distribution. — Salammbô, Tunisia (De Saint Laurent & Božić, 1976);
Port of Carthago, Tunisia (De Saint Laurent & Manning, 1982); Sardegna
(= Sardinia), and Naples, Italy (De Saint Laurent & Manning, 1982); southern
Euboikos Gulf, Messolongi lagoon, Salamina, Saronikos Gulf, Crete, Greece
(Thessalou-Legaki, 1986).
Calliaxina sakaii (De Saint Laurent & Le Loeuff, 1979)
[Abbreviated list of synonymy.]
Callianassa (Callichirus) novaebritanniae — Sakai, 1966: 161, figs. 1-4. [Not Borradaile,
1900.]
Calliax sakaii De Saint Laurent & Le Loeuff, 1979: 95; Sakai, 2005b: 2003.
Paraglypturus sakaii — Sakai, 1999c: 124, fig. 33d, e.
Calliaxina sakaii — Ngoc-Ho, 2003: 493; Itani, 2007: 203.
Diagnosis. — Carapace without dorsal oval but cardiac sulcus present.
Mxp3 ischium-merus subovate; merus subquadrate; propodus subtriangular;
dactylus subtriangular; exopod short. Male Plp1 uniramous and bisegmented,
distal segment subchelate distally, longer branch incurvate and shorter one
bilobed distally, bearing appendix interna; male Plp2 biramous, endopod
bearing mesiomedially a distinct appendix masculina with a small appendix
interna subproximally. Telson wider than long and posterior margin straight.
Uropodal exopod with dorsal plate. [Cf. Sakai, 1966: 161, figs. 1c, 2g, 4a, c.]
Type locality. — Tomioka, Amakusa, Japan, intertidal under Zostera vegetation.
Distribution. — Urauch-gawa, Iriomote Is., Ryukyu Archipelago (Itani,
2007); Amakusa, East China Sea (Sakai, 1966).
Genus Eucalliax Manning & Felder, 1991
Eucalliax Manning & Felder, 1991: 781, figs. 7, 15a-e; Poore, 1994: 101.
Eucalliax s. str. — Felder, 2001: 440.
Diagnosis. — Carapace without dorsal oval and cardiac sulcus. Mxp3
ischium-merus rectangular; propodus subsquare; dactylus subtriangular; exopod absent. Male Plp1 uniramous and bisegmented, distal segment simple
and obtuse distally; male Plp2 biramous, endopod bearing distally a distinct
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
503
appendix masculina with setae, and an appendix interna subdistally on the lateral margin (Biffar, 1970, fig. 2m). Telson distinctly wider than long. Uropodal
exopod with dorsal plate. [Cf. Sakai, 1999c: 117, fig. 30C.]
Remarks. — The genus Eucalliax Manning & Felder, 1991 is very similar
to the genus Eucalliaxiopsis gen. nov., but they differ in that in Eucalliaxiopsis the male Plp1 is uniramous and bisegmented, but the distal segment is
broadened medially and strongly incurvate distally, and the male Plp2 endopod bears mesiomedially a distinct appendix masculina with setae, provided
subproximally with a small, proximally-fused protrusion as appendix interna.
Type species. — Callianassa quadracuta Biffar, 1970, by original designation and monotypy. The gender of the generic name, Eucalliax, is feminine.
Species included. — Eucalliax quadracuta (Biffar, 1970).
Eucalliax quadracuta (Biffar, 1970)
Callianassa quadracuta Biffar, 1970: 40, fig. 2; Biffar, 1971a: 694, figs. 17, 18. [Type species
of Eucalliax Manning & Felder, 1991.]
Calliax quadracuta — De Saint Laurent & Manning, 1982: 222; Manning, 1987: 397; Sakai,
1999c: 117, fig. 30a-c; Sakai, 2005b: 200.
Eucalliax quadracuta — Manning & Felder, 1991: 781; Tudge et al., 2000: 145.
Diagnosis. — Cardiac sulci absent. Mxp3 ischium-merus rectangular; merus
subquadrate; propodus subsquare; dactylus subtriangular; exopod absent. Male
Plp1 uniramous and bisegmented, distal segment simple and obtuse distally;
male Plp2 biramous, endopod distally bearing a distinct appendix masculina
with setae, and an appendix interna subdistally on the lateral margin; Plps3-5
with appendix interna. Telson wider than long, posterior margin convex medially. Uropodal exopod with dorsal plate. [Cf. Biffar, 1970: 40, fig. 2f, i, k, m;
Sakai, 1999c: 117, fig. 30b.]
Type locality. — Venezuela, Cumaná.
Distribution. — Gulf of Uraba, Colombia, Caribbean coast; Cumaná, Venezuela.
Genus Eucalliaxiopsis gen. nov.
Diagnosis. — Carapace without dorsal oval or cardiac sulci. Mxp3 ischiummerus suboperculiform; propodus subtriangular; dactylus subtriangular; exopod absent. Male Plp1 uniramous and bisegmented, distal segment broadened
medially and incurvate distally, bearing long marginal setae; male Plp2 biramous, endopod bearing mesiomedially a distinct appendix masculina overreaching top of endopod, provided subproximally with small, proximallyfused protrusion as appendix interna. Telson wider than long, posterior margin
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CRM 013 – Katsushi Sakai
almost straight. Uropodal exopod with dorsal plate. [Adapted from Rodrigues
& Manning, 1992, fig. 2.]
Remarks. — The type species of the new genus Eucalliaxiopsis, E. cearaensis, is very similar to that of the genus Eucalliax, E. quadracuta, but
they are different from each other in the forms of Plps1-2. In E. cearaensis
the male Plp1 is uniramous and bisegmented; the distal segment is broadened
medially and incurvate distally, bearing long marginal setae; the male Plp2 is
biramous and the endopod bears mesiomedially a distinct appendix masculina
overreaching the top of the endopod, provided subproximally with a small
subproximally-fused protrusion as an appendix interna (cf. Rodrigues &
Manning, 1992: 329, fig. 2t); whereas in E. quadracuta the male Plp1 is
uniramous and bisegmented, but the distal segment is simple and obtuse
distally; the male Plp2 is biramous, but the endopod bears distally a distinct
appendix masculina and a small appendix interna; Plps3-5 with finger-like
appendix interna.
Type species. — Eucalliax cearaensis Rodrigues & Manning, 1992, by
present designation and monotypy. The gender of the generic name, Eucalliaxiopsis, is feminine.
Species included. — Eucalliaxiopsis cearaensis (Rodrigues & Manning,
1992).
Eucalliaxiopsis cearaensis (Rodrigues & Manning, 1992)
Eucalliax cearaensis Rodrigues & Manning, 1992a: 327, fig. 2; Tudge et al., 2000: 145; Coelho
et al., 2007: 5 (tab. 2).
Eucalliax cearensis – Rodrigues et al., 1998: 380; Melo, 1999: 368, figs. 245-246.
Calliax cearaensis — Sakai, 1999c: 112; Sakai, 2005b: 199.
Type locality. — Barro de Ceará, Fortaleza, right bank of river mouth
(3 45′ S 38◦ 35′ W), Brazil.
Distribution. — Only known from the type locality.
◦
Genus Paraglypturus Türkay & Sakai, 1995
Paraglypturus Türkay & Sakai, 1995: 26; Sakai, 2005b: 204.
Definition. — Rostrum poorly developed, lacking rostral carina. Carapace
without dorsal oval, cardiac sulcus, and hepatic prominence. Mxp3 ischiummerus rectangular; merus bullet-headed in shape; propodus broadened and
subquadrate; dactylus subtriangular, bearing setae on distal margin; exopod
distinct. P1 unequal, larger cheliped without meral hook. P2 chelate. P3
propodus subtrapezoid; dactylus triangular. P4 subchelate. P5 chelate. Male
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
505
Plp1 uniramous and bisegmented, distal segment subchelate distally; male
Plp2 biramous and foliaceous, endopod bearing mesiomedially appendix
masculina with distal setae and shorter appendix interna with hooklets; male
Plps3-5 biramous and foliaceous, endopods bearing appendix interna. Telson
wider than long and subtrapezoid. Uropodal endopod oval and much longer
than telson; uropodal exopod with dorsal plate and lateral notch, bearing
proximolateral portion with thick setae, and transparent yellow circle in middle
of mesial half. [Adapted from Sakai, 1999c: 122.]
Type species. — Paraglypturus calderus Türkay & Sakai, 1995, by original
designation and monotypy. Gender of generic name, Paraglypturus, masculine.
Species included. — Only the type species, Paraglypturus calderus Türkay
& Sakai, 1995.
Paraglypturus calderus Türkay & Sakai, 1995
Paraglypturus calderus Türkay & Sakai, 1995: 27, figs. 2-6; Sakai, 2005b: 200.
Type locality. — North rim of Esmeralda Caldera, Marianas, 14◦ 58.03′ N
145◦ 15.14′ E; 63 m.
Distribution. — Only known from the type locality.
Genus Pseudocalliax gen. nov.
Diagnosis. — Carapace with dorsal oval and cardiac sulcus. Mxp3 ischiummerus rectangular; propodus subtriangular; dactylus subovate; exopod present.
P1 subequal, larger cheliped without meral hook. Male Plp1 uniramous and
simple; male Plp2 biramous, endopod with appendix interna mediolaterally,
but without appendix masculina. Telson wider than long and subtrapezoid.
Uropodal endopod oval and much longer than telson; uropodal exopod with
dorsal plate.
Remarks. — Callianassa tooradin Poore & Griffin, 1979 was included in
the genus Calliax De Saint Laurent, 1973 by Sakai (1988: 61), because in
Callianassa tooradin the male Plp1 is absent (Poore & Griffin, 1979: 227),
and therefore it differs from the type species of the genus Callianassa, C.
subterranea (Montagu, 1808), in which the male Plp1 is present. Later, the
species was reclassified under the genus Paraglypturus Türkay & Sakai, 1995
by Sakai (1999c), and then once again under the genus Calliax by Sakai
(2005b). The present examination of the male specimen preserved in the
USNM shows, however, that in Callianassa tooradin the carapace bears a
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CRM 013 – Katsushi Sakai
cardiac sulcus and a dorsal oval as shown by Poore & Griffin (1979: 275),
and the male Plp1 is small and simple as informed by G.C.B. Poore (in litt.),
so that Calliax tooradin is reclassified under the new genus Pseudocalliax gen.
nov., belonging to the family Eucalliacidae.
Type species. — Callianassa tooradin Poore & Griffin, 1979, by present
designation and monotypy. Gender of the generic name, Pseudocalliax, feminine.
Species included. — Pseudocalliax tooradin (Poore & Griffin, 1979).
Pseudocalliax tooradin (Poore & Griffin, 1979)
Callianassa tooradin Poore & Griffin, 1979: 275, fig. 36.
Calliax tooradin — Sakai, 1988: 61; Davie, 2002: 459; Sakai, 2005b: 204.
Paraglypturus tooradin — Sakai, 1999c: 124, figs. 33a-c; Sakai, 2005b: 204.
Material examined. — USNM, 1 male, Victoria Australia.
Diagnosis. — Rostrum obtusely triangular, lacking rostral carina. Carapace with dorsal oval, cardiac prominence, cardiac sulcus, and linea thalassinica. Eyestalks flattened and contiguous. A2 scaphocerite present as a
small process. Mxp3 ischium-merus rectangular; propodus subrectangular;
dactylus subovate; exopod present. P1 chelate, unequal in size, and dissimilar in shape; larger and smaller chelipeds without meral hook ventroproximally. P2 chelate. P3 propodus broadened. P4 simple. P5 subchelate. Abdominal pleura 2-5 unarmed ventrally. Abdominal somite 6 lacking lateral projections. Male Plp1 small and simple; male Plp2 biramous, endopod bearing
appendix interna mediolaterally, but no appendix masculina. Telson wider than
long and subtrapezoid; parallel-sided in proximal half and narrowing toward
slightly rounded posterior margin. Uropodal exopod with dorsal plate.
Type locality. — Australia, Victoria, Crib Point, Western Port, fine sand
sediment, 5 m.
Distribution. — Only known from the type locality.
FAMILY GOURRETIIDAE SAKAI, 1999
Gourretiinae Sakai, 1999a: 95; Ngoc-Ho, 2003: 498.
Ctenochelidae — Ngoc-Ho, 2003: 498.
Gourretiidae — Sakai, 2005b: 217; Sakai & Sawada, 2006: 1357.
Dawsoniinae Sakai, 2006: 1276.
Diagnosis. — Rostrum small and unarmed on lateral margins, bearing no
rostral carina. Carapace with linea thalassinica, but without dorsal oval, and
cardiac sulcus incomplete or absent; vertical row of setae rarely present on
branchial region. Eyestalks flattened and contiguous. A2 scaphocerite present
as a small process. Maxilla 2 scaphognathite without posterior whip. Mxp3
ischium-merus rectangular; merus with mesial spine; propodus rectangular;
dactylus digitiform; exopod present or absent. P1 chelate, unequal in size, and
dissimilar in shape; in larger cheliped merus with hook ventroproximally, palm
rectangular, fingers elongate; in smaller cheliped merus usually with hook
ventroproximally (not in Gourretia lahouensis), fingers elongate. P2 chelate.
P3 propodus subpediform. P4 simple. P5 chelate. Male Plp1 uniramous and
bisegmented, distal segment simple, subchelate, or chelate distally; male Plp2
biramous and slender, endopod bearing appendix masculina provided with
appendix interna, or bearing only appendix interna. Abdominal somite 6 with
or without lateral projection. Telson subtriangular or subsquare. Uropodal
exopod without dorsal plate, and with or without lateral notch.
Remarks. — The subfamily Pseudogourretiinae Sakai, 2005b was established and tentatively classified under the family Gourretiidae. However, it has
turned out that the Pseudogourretiinae is to be included neither in the family Gourretiidae because of the presence of pleurobranchs, nor in the family
Axiidae Huxley, 1878 because of its distinct characteristics as follows. The
eyestalks are flattened and contiguous, the linea thalassinica is present incompletely, and the maxilla 2 scaphognathite bears no posterior whip, which
features are not observed in the Axiidae, though the Pseudogourretiinae are
closely similar to the Axioidea in that the pleurobranchs are present as in
the genera of the Axioidea: Axius Leach, 1815; Bouvieraxius Sakai & De
Saint Laurent, 1989; Calaxius Sakai & De Saint Laurent, 1989; Calocaris
Bell, 1846; Eiconaxius Bate, 1888; Neaxius Borradaile, 1903; Parascytoleptus Sakai & De Saint Laurent, 1989; Scytoleptus Gerstaecker, 1856; Spongiax-
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CRM 013 – Katsushi Sakai
ius Sakai & De Saint Laurent, 1989; and Strahlaxius Sakai & De Saint Laurent, 1989. Whereas they are absent in the axioid genera Acanthaxius Sakai
& De Saint Laurent, 1989; Allaxius Sakai & De Saint Laurent, 1989; Ambiaxius Sakai & De Saint Laurent, 1989; Axiopsis Borradaile, 1903; Axiorygma
Kensley & Simmons, 1988; Calastacus Faxon, 1893; Calaxiopsis Sakai & De
Saint Laurent, 1989; Calocarides Wollenbaeck, 1908; Dorphinaxius Sakai &
De Saint Laurent, 1989; Eutrichocheles Wood-Mason, 1876b; Oxyrhynchaxius Parisi, 1917; and Paraxius Bate, 1888. In the present revision, therefore, the
Pseudogourretiinae are elevated to family-level, and are distinguished from the
Axiidae and Gourretiidae.
Type genus. — Gourretia De Saint Laurent, 1973.
Genera included. — Dawsonius sensu Sakai, 2005 (not Manning & Felder,
1991); Gourretia Sakai, 1999c; Laurentgourretia Sakai, 2004b; Paracalliax
De Saint Laurent, 1979; Paragourretia Sakai, 2004b.
K EY
TO THE
G ENERA
OF THE
FAMILY G OURRETIIDAE
1 – Eyestalks elongate and rectangular, without cornea . . . . . . . . . . . . . . . . . . . . . . Paracalliax
– Eyestalks shorter, with cornea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Abdominal somite 6 with lateral projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dawsonius
– Abdominal somite 6 without lateral projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Uropodal exopod without lateral notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gourretia
– Uropodal exopod with lateral notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – A2 scaphocerite strong, Mxp3 exopod absent . . . . . . . . . . . . . . . . . . . . . . Laurentgourretia
– A2 scaphocerite obtuse, Mxp3 exopod present . . . . . . . . . . . . . . . . . . . . . . . . Paragourretia
Genus Dawsonius Manning & Felder, 1991 sensu Sakai, 2005
Dawsonius Manning & Felder, 1991 sensu Sakai, 2005b.
Dawsonius — Sakai, 2005b: 245.
Diagnosis. — Rostrum triangular. Carapace without dorsal oval and cardiac
sulcus, but with low dorsal carina in anterior half. A2 scaphocerite small
and pointed. Mxp3 propodus rectangular; dactylus digitiform; exopod present.
Abdominal somite 6 bearing lateral projection anteriorly on each side. Male
Plp1 uniramous and bisegmented, distal segment subchelate distally; male
Plp2 biramous, endopod bearing mesiodistally appendix masculina provided
mesiomedially with proximally-fused appendix interna. Telson subquadrate,
posterior margin rounded. Uropodal exopod without lateral notch.
Type species. — Callianassa latispina Dawson, 1967, by original designation and monotypy. The gender of the generic name, Dawsonius, is masculine.
Species included. — Dawsonius latispina (Dawson, 1967).
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
509
Dawsonius latispina (Dawson, 1967)
(fig. 69E)
Callianassa latispina Dawson, 1967: 190, fig. 1; Biffar, 1971a: 651 (key), 654 (key), 679, figs.
11-12; Herper, 1975: 619; Rabalais et al., 1981: 105, fig. 3g-k.
Gourretia latispina — Manning, 1987: 398 (list); Williams et al., 1989: 28.
Dawsonius latispina — Manning & Felder, 1991: 785 (list), figs. 4, 16; Poore, 1994: 103 (key);
Coelho, 1997: 150; Melo, 1999: 394, figs. 265-266; Tudge et al., 2000: 142; Sakai, 2005b:
245; Coelho et al., 2007: 5 (tab. 2); Robles et al., 2009: 317.
Callianopsis latispina — Sakai, 2005b: 228.
Fig. 69. Gourretia denticulata (Lutze, 1937) and Dawsonius latispina (Dawson, 1967). A, carapace lateral view, B, male Plp1, C, male Plp2; D, male Plp3, E, distal end of male Plp2 endopod.
A-D, Gourretia denticulata, SMF 28053, neotype, 1 male (20.0/4.4, left minor cheliped detached, lacking left P/4), in front of Sotto Castello, Limski Canal, Istria, Croatia, 45◦ 08.020′ N
013◦ 39.030′ E, 29 m; E, Dawsonius latispina, USNM 103755, paratype, male (TL/CL, 30.0/
9.0 mm), S. of Grand Isle, Louisiana, 13.7 m. Scales 1 mm.
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CRM 013 – Katsushi Sakai
Material examined. — USNM 103755, paratype, male (TL/CL, 30.0/9.0 mm), S. of Grand
Isle, Louisiana, 7.5 fms [ca. 13.5 m], 4.ii.1959, leg. et don. C.E. Dawson; USNM 172310,
lectotype, 1 female (CL, 6.0 mm), Sta. I-4, 28◦ 14′ N 96◦ 29′ W, 10 m, 13.vi.1976.
Diagnosis. — Rostrum triangular. Carapace lacking dorsal oval, but bearing
low dorsal carina in anterior half. A2 scaphocerite pointed and 0.6 mm
long. Mxp3 ischium-merus pediform; merus with mesiodistal spine; propodus
rectangular; dactylus digitiform; exopod present. P1 unequal, both chelipeds
with meral hook ventroproximally. Abdominal somite 6 with lateral projection
anteriorly on each side. Male Plp1 uniramous and bisegmented, distal segment
subchelate distally; male Plp2 (fig. 69E) biramous, endopod longer than
exopod, bearing mesiodistally appendix masculina with apical setae, provided
mesiomedially with proximally-fused appendix interna; Plps3-5 with appendix
interna. Telson subquadrate, posterior margin rounded. Uropodal exopod
without lateral notch. [Adapted from Dawson, 1967: 190; Biffar, 1971a: 679,
figs. 11b, I, 12b, e.]
Remarks. — The type species of Dawsonius, Dawsonius latispina was
examined in order to confirm the presence or absence of the Mxp3 exopod,
because the Mxp3 exopod is shown and described in the figure and description
of Dawsonius latispina by Biffar (1971a, fig. 11b), though it is shown neither
in the figure and description by Dawson (1967, fig. 1f, h), nor in those of
Manning & Felder (1991, fig. 16c, d). It was found through the present
examination of the type specimens that the Mxp3 exopod is present.
Type locality. — Off Grand Isle, Louisiana, 135 m.
Distribution. — Louisiana; Texas; off south-western Florida; Honduras;
Amapá, Pará, and Alagoas, Brazil (Coelho, 1997).
Genus Gourretia De Saint Laurent, 1973
Gourretia De Saint Laurent, 1973: 514; Le Loeuff & Intès, 1974: 26; Poore & Griffin, 1979:
278; De Saint Laurent & Le Loeuff, 1979: 48 (key), 78; Manning, 1987: 398 (list); Manning
& Felder, 1991: 785, fig. 3; Poore, 1994: 103 (key); Davie, 2002: 464; Sakai, 2002: 468;
Ngoc-Ho, 2003: 498; Sakai, 2005b: 218.
Diagnosis. — Rostum triangular, without rostral carina. Carapace lacking dorsal oval, transverse cardiac sulcus, and cardiac prominence; linea thalassinica present (fig. 67A). A2 scaphocerite rudimentary. Dorsal surfaces of
eyestalks flattened. Mxp3 ischium-merus subpediform; merus usually with
mesiodistal spine; propodus rectangular; dactylus digitiform; exopod present.
P1 with proximal meral hook. P3 propodus broadened. P4 simple. P5 chelate.
Abdominal somite 6 lacking lateral projection. Male Plp1 uniramous and
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511
bisegmented, distal segment chelate distally; male Plp2 biramous, endopod
bearing mesiodistally appendix masculina with distal setae, provided proximally with proximally-fused appendix interna; Plps3-5 with triangular appendix interna. Telson subtriangular or subsquare. Uropodal exopod lacking lateral notch. [Adapted from Sakai, 2005b.]
Type species. — Callianassa denticulata Lutze, 1937, by original designation and monotypy. Gender of the generic name, Gourretia, feminine.
Species included. — Gourretia barracuda Le Loeuff & Intès, 1974; G. denticulata (Lutze, 1937); G. laresi Blanco Rambla & Liñero Arana, 1994;
G. loeuffintesi Sakai, 2005b; G. manihinae Sakai, 1984; G. nosybeensis Sakai,
2004b; Gourretia sp. Robles et al., 2009: 317.
K EY
TO THE
S PECIES
OF THE
G ENUS GOURRETIA
1 – Telson subquadrate, recorded from Nosy Bé, Moçambique Channel . . . . G. nosybeensis
– Telson trapezoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – P3 propodus rectangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– P3 propodus ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 – P1 chela of larger cheliped much wider than carpus, from Ivory Coast, 100-250 m . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. barracuda
– P1 chela of larger cheliped as wide as carpus, from the Gulf of Guinea, Benin, 39 m . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. loeuffintesi
4 – P3 propodus wide on distal margin, from Tanzania, 35 m . . . . . . . . . . . . . . . G. manihinae
– P3 propodus narrow on distal margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – Male Plp1 with stout proximal and distal segments, from eastern Atlantic Ocean and
Mediterranean Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. denticulata
– Male Plp1 with elongate, slender proximal and distal segments, from northwestern
Chimana Island, Venezuela, 71 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. laresi
Gourretia barracuda Le Loeuff & Intès, 1974
Gourretia barracuda Le Loeuff & Intès, 1974: 30, fig. 6a-t; De Saint Laurent & Le Loeuff,
1979: 80, fig. 24a; Manning & Felder, 1991: 785 (list); Tudge et al., 2000: 142; Sakai, 2005b:
219.
Type locality. — Abidjan, Ivory Coast, 100-250 m.
Distribution. — Abidjan, Ivory Coast (Le Loeuff & Intès, 1974; De Saint
Laurent & Le Loeuff, 1979); 100-250 m.
Gourretia denticulata (Lutze, 1937)
(fig. 69A-D)
[Abbreviated list of synonymy.]
Callianassa subterranea var. minor Gourret, 1887: 1034; Gourret, 1888: 96, pl. 8 figs. 1-15.
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CRM 013 – Katsushi Sakai
Callianassa denticulata Lutze, 1937: 6, figs. 1-7; Lutze, 1938: 170; Vatova, 1949: 65, tabs. 22,
31, 32.
Callianassa minor — Holthuis & Gottlieb, 1958: 56, figs. 11-12; Harmelin, 1964: 68, 95;
Picard, 1965: 59, 104 (list); Števčić, 1969a: 128 (list); Štjepčević & Parenzan, 1980: 47.
Gourretia serrata De Saint Laurent & Le Loeuff, 1979: 79, 80 (key), fig. 24c; Thessalou-Legaki
& Zenetos, 1985: 311; Thessalou-Legaki, 1986: 182, 184 (list); Dounas et al., 1993: 49.
Gourretia denticulata — Lewinsohn & Holthuis, 1986: 24; Števčić, 1990: 218; Manning &
Felder, 1991: 785 (list), fig. 3; Dworschak, 1992: 210; Koukouras et al., 1992: 223; López
de la Rosa et al., 1998: 394, fig. 1; Sakai, 1999c: 128; d’Udekem d’Acoz, 1999: 156; Tudge
et al., 2000: 142; Costello et al., 2001: 289; Ngoc-Ho, 2003: 499, fig. 21; Sakai, 2005b: 220;
Sakai & Türkay, 2005: 323, figs. 1-3, tabl. 1; Sakai & Sawada, 2006: 1357.
Not: Gourretia minor — Le Loeuff & Intès, 1974: 26, fig. 4a-k (= Gourretia sp. De Saint
Laurent & Le Loeuff, 1979) (Gourretia loeuffintesi Sakai, 2005b).
Material examined. — SMF 28053, neotype, 1 male (20.0/4.4, left minor cheliped detached, lacking left P4), in front of Sotto Castello, Limski Canal, Istria, Croatia, 45◦ 08.020′ N
013◦ 39.030′ E, 29 m, soft mud, R/V “Burin”, 1.ix.1999.
Diagnosis. — Male Plp1 uniramous and bisegmented, distal segment chelate
distally (fig. 69B); male Plp2 biramous, endopod bearing mesiodistally appendix masculina with distal setae, provided proximally with proximally-fused
appendix interna (fig. 69C); male Plps3-5 biramous and broad, endopods with
appendix interna mesiomedially (fig. 69D); Plps3-5 with small triangular appendix interna. Telson subsquare.
Remarks. — The male neotype specimen from the Limski Canal, Istria,
Croatia, is probably an immature one, because the male Plp1 distal segment is
emarginate distally, as shown by López de la Rosa et al. (1998: 394, fig. 1F),
which suggests that it has not yet developed into a chela.
Type locality. — Limski-Kanal, Istria, Croatia, 29 m, Adriatic Sea.
Distribution. — Eastern Atlantic Ocean — Bay of Cadiz, Spain (López
de la Rosa et al., 1998). Mediterranean — Gulf of Marseille, Ionian Sea,
Cyprus, Malta, and along the coast of Israel; Baie de Kotor (De Saint Laurent
& Božić, 1976); Piran Gulf (Manning & Števcic, 1982). Adriatic Sea (Lutze,
1937; Dworschak, 1992). Tyrrhenian Sea — Ischia (Dworschak, 1992), Malta;
Ionian Sea. Aegean Sea — N. Euboikos Gulf, Patraikos, and Rhodos, 5-15 m,
in pure sand (Thessalou-Legaki, 1986; Koukouras et al., 1992). Cyprus; Israel
(Lewinsohn & Holthuis, 1986); 2.5-146 m.
Gourretia laresi Blanco Rambla & Liñero Arana, 1994
Gourretia laresi Blanco Rambla & Liñero Arana, 1994: 25, fig. 8; Tudge et al., 2000: 142;
Sakai, 2005b: 222.
Diagnosis. — Rostrum sharply triangular, not reaching distal end of eyestalks. Eyestalks separated at distal part, shaped in triangle. Mxp3 ischium
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
513
with strong, curved projection on ventral margin. P1 unequal. Male Plp1 uniramous and bisegmented, distal segment slightly subchelate distally; male Plp2
biramous, endopod bearing mesiodistally appendix masculina with small appendix interna mesiodistally. [Adapted from Blanco Rambla & Liñero Arana,
1994.]
Type locality. — Northwest Chimana Islands, 10◦ 18.40′ N 64◦ 47.40′ W,
Venezuela.
Distribution. — Only known from the type locality.
Gourretia loeuffintesi sp. nov.
Gourretia minor — Le Loeuff & Intès, 1974: 26, fig. 4a-k.
Gourretia sp., De Saint Laurent & Le Loeuff, 1979: 81, fig. 24b.
Gourretia loeuffintesi Sakai, 2005b: 221 (name invalid).
Material examined. — Holotype, male, Dahomey, 6◦ 11.5′ N 2◦ 12.5′ W, Republic of Benin,
Gulf of Guinea, 3.x.1963 (cf. Le Loeuff & Intès, 1974: 26).
Diagnosis. — Rostrum acutely triangular, reaching middle of eyestalks.
Eyestalks contiguous and rounded distally. Mxp3 ischium-merus pediform. P3
propodus oval. Telson subtriargular, narrowing in posterior half and rounded
on posterior margin. [Adapted from Sakai, 2005b: 222.]
Remarks. — The present species was described by Sakai (2005b), without
designating a type specimen, so that the species name was invalid according to
Art. 72.3 of the ICZN. In the present revision, this species is properly named
G. loeuffintesi by designating a type.
Type locality. — Benin, 6◦ 11.5′ N 2◦ 12.5′ W, 39 m, Gulf of Guinea.
Distribution. — Only known from the type locality.
Gourretia manihinae Sakai, 1984
Gourretia manihinae Sakai, 1984: 101, figs. 4-5; Tudge et al., 2000: 142.
Type locality. — Pangani Bay, Tanzania, 5◦ 29.2′ S 39◦ 03.2′ E, 35 m.
Distribution. — Tanzania — Pangani Bay (Sakai, 1984); 35 m.
Gourretia nosybeensis Sakai, 2004
Gourretia nosybeensis Sakai, 2004b: 563, figs. 4-6; Sakai, 2005b: 224.
Diagnosis. — Rostrum shaped as slender triangle, reaching distal end of
eyestalks. A1 peduncle as long as A2 peduncle. Mxp3 merus with distinct
subdistal tooth. P3 propodus rounded. Male Plp1 uniramous, bisegmented;
distal segment subchelate distally; male Plp2 biramous, exopod shorter than
514
CRM 013 – Katsushi Sakai
endopod; endopod bilobed distally, mesial lobe bearing distally appendix
masculina with long setae distally and fused appendix interna mesiodistally.
Telson subsquare. Uropodal exopod without lateral notch. [Adapted from
Sakai, 2004b.]
Type locality. — Nosy Bé, Madagascar, and near Île d’Ampasindava, 47 m.
Distribution. — Only known from the type locality.
Gourretia sp. Robles et al., 2009
Gourretia sp. Robles et al., 2009: 317.
Genus Laurentgourretia Sakai, 2004
Laurentgourretia Sakai, 2004b: 558; Sakai, 2005b: 224.
Diagnosis. — Rostrum sharply triangular, without rostral carina, and slightly
overreaching middle of eyestalks. Eyestalks obtuse distally; cornea small and
located mesiodistally. Carapace lacking dorsal oval, transverse cardiac sulcus, and cardiac prominence. A2 scaphocerite distinct, overreaching middle
of P2 penultimate segment. Mxp3 ischium-merus subpediform; merus with
three mesial spines; propodus rectangular; dactylus digitiform; exopod absent.
P3 propodus ovate. P4 simple. P5 subchelate. Abdominal somite 6 lacking triangular anteroventral projections. Male Plps1-2 unknown. Telson subsquare
and convex on proximal third of lateral margin. Uropodal exopod with lateral
notch. [Adapted from Sakai, 2004b.]
Remarks. — The type-species of the genus Laurentgourretia, L. rhopalommata Sakai, 2004b from Nosy Mitsio, Mozambique channel, is closely similar
to Paragourretia phuketensis Sakai, 2002 from the Andaman Sea. However,
they differ from each other in that in L. rhopalommata the Mxp3 exopod is
absent, whereas in P. phuketensis the Mxp3 exopod is present.
Type species. — Laurentgourretia rhopalommata Sakai, 2004b, by original
designation and monotypy. Gender of generic name, Laurentgourretia, feminine.
Species included. — Laurentgourretia rhopalommata Sakai, 2004b.
Laurentgourretia rhopalommata Sakai, 2004
Laurentgourretia rhopalommata Sakai, 2004b: 557, figs. 1-3.
Diagnosis. — Rostrum sharply triangular, slightly overreaching middle of
eyestalks. Eyestalks obtuse distally; cornea small and located mesiodistally.
A2 scaphocerite strongly protruded. Mxp3 merus with three strong mesial
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
515
teeth. Telson subsquare and convex on proximal third of lateral margin.
Uropodal exopod with lateral notch.
Type locality. — Nosy Mitsio, Mozambique channel, 12◦ 38.400′ S
48◦ 33.200′ E, 50 m.
Distribution. — Only known from the type locality.
Genus Paracalliax De Saint Laurent, 1979
Paracalliax De Saint Laurent, 1979: 1396; De Saint Laurent & Le Loeuff, 1979: 47 (key), 84;
Manning & Felder, 1991: 785, figs. 1, 7; Poore, 1994: 103 (key): Sakai, 2005b: 215.
Diagnosis. — Rostrum small and triangular, bearing obscure rostral carina.
Carapace without dorsal oval and transverse cardiac sulcus, but with cardiac
prominence. Eyestalks distinct, elongate, flattened, and contiguous. A2 peduncle about as long as A1 peduncle; A2 scaphocerite small. Mxp 3 ischiummerus pediform, bearing meral spine; propodus rectangular; dactylus digitiform; exopod present. P1 chelate, unequal in size and dissimilar in shape,
lacking proximal meral hook in both chelipeds. P2 chelate. P3 propodus rectangular. Abdominal somite 6 with lateral projections. Male Plps1-2 unknown;
female Plp1 uniramous, bisegmented, distal segment foliaceous; female Plp2
biramous and foliaceous, endopod with appendix interna; Plps3-5 biramous
and foliaceous, endopods with appendix interna (cf. De Saint Laurent & Le
Loeuff, 1979: 87). Telson subquadrate, posterior margin with median spine.
Uropodal exopod lacking lateral notch.
Remarks. — The present genus Paracalliax was included in the family
Ctenochelidae by Manning & Felder (1991: 784). However it was subsequently included in the family Gourretiidae Sakai, 1999a by Sakai (2005b),
because in the Gourretiidae the rostral carina is absent; the fingers of the larger
cheliped are not pectinate and the palm is subquadrate; and the uropodal exopod bears no lateral notch. In the Ctenochelidae, however, the rostral carina
is present; the fingers of the larger cheliped are pectinate and the palm is oval;
and the uropodal exopod bears a lateral notch; so that Paracalliax is safely
included in the Gourretiidae.
Type species. — Paracalliax bollorei De Saint Laurent, 1979, by original
designation and monotypy. Gender of generic name, Paracalliax, feminine.
Species included. — Paracalliax bollorei De Saint Laurent, 1979.
Paracalliax bollorei De Saint Laurent, 1979
Paracalliax bollorei De Saint Laurent, 1979: 1396; De Saint Laurent & Le Loeuff, 1979: 86,
figs. 26a-I, 27a-c, 28a-h; Manning & Felder, 1991: 785 (list), figs. 1, 7; Tudge et al., 2000:
142; Sakai, 2005b: 216.
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CRM 013 – Katsushi Sakai
Diagnosis. — Rostrum triangular, not reaching proximal fifth of eyestalks. Eyestalks elongate and compressed; cornea absent. P1 unequal. Mxp3
ischium-merus pediform; merus with mesiodistal spine; exopod present. Male
Plps1-2 unknown. Telson subquadrate, and convex on proximal third of lateral
margin, bearing median spine on posterior margin. [Adapted from De Saint
Laurent & Le Loeuff, 1979: 86, figs. 26-28.]
Type locality. — Banc d’Arguin, Mauritania, 20-100 m?
Distribution. — Only known from the type locality.
Genus Paragourretia Sakai, 2004
Paragourretia Sakai, 2004b: 568; Sakai, 2005b: 225.
Diagnosis. — Rostrum triangular, without rostral carina. Carapace large and
thick, lacking dorsal oval, but bearing cardiac prominence and incomplete,
transverse cardiac sulcus; row of transverse setae present in anterior branchial
region. Eyestalks flattened. A2 scaphocerite obtuse or reduced. Mxp3 ischiummerus subpediform; merus with or without subdistal spine on mesial margin;
propodus rectangular; dactylus digitiform; exopod present. Larger cheliped
with proximal meral hook. P3 simple, propodus rectangular. P4 subchelate. P5
chelate. Abdominal somite 6 without lateral projections. Male Plp1 uniramous
and bisegmented, distal segment subchelate distally; male Plp2 biramous,
endopod bearing appendix masculina with distal setae, provided with elongate
appendix interna, or bearing only appendix interna; Plps3-5 with triangular
appendix interna. Telson subtriangular or subsquare. Uropodal exopod with
lateral notch.
Remarks. — The present genus Paragourretia is closely similar to Gourretia, but they differ in that in Paragourretia the uropodal exopod has a lateral
notch, whereas in Gourretia it does not have a lateral notch, although this
was described (Sakai, 2002) for the type-species of the genus Paragourretia,
P. phuketensis, i.e., that the uropodal exopod bears no notch on the lateral
margin, which may have been caused by misobservation, because the notch
is present on the lateral margin of the uropodal exopod as later described by
Sakai (2005b).
Type species. — Gourretia phuketensis Sakai, 2002, by original designation. Gender of generic name, Paragourretia, feminine.
Species included. — Paragourretia aungtonyai (Sakai, 2002); P. biffari
Blanco Rambla & Liñero Arana, 1994; P. coolibah Poore & Griffin, 1979;
P. crosnieri Ngoc-Ho, 1991; P. lahouensis Le Loeuff & Intès, 1974; P. phuketensis (Sakai, 2002).
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
K EY
TO THE
S PECIES
OF THE
517
G ENUS PARAGOURRETIA
1 – Mxp3 merus with subdistal spine on mesial margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Mxp3 merus without subdistal spine on mesial margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 – Telson subtriangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. biffari
– Telson subquadrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. phuketensis
3 – Uropodal exopod without lateral notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lahouensis
– Uropodal exopod with lateral notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 – Mxp3 exopod overreaching ischium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. coolibah
– Mxp3 exopod not reaching distal margin of ischium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 – A1 peduncle reaching distal part of A2 distal segment . . . . . . . . . . . . . . . . . . . P. crosnieri
– A1 peduncle reaching middle of A2 distal segment . . . . . . . . . . . . . . . . . . . . P. aungtonyai
Paragourretia aungtonyai (Sakai, 2002)
Gourretia aungtonyai Sakai, 2002: 476, figs. 6A-H, 7A-F, 8A-G.
Paragourretia aungtonyai — Sakai, 2005b: 225.
Diagnosis. — Mxp3 ischium-merus pediform; merus unarmed on mesial
margin; propodus rectangular; and dactylus digitiform; exopod present. Male
Plp1 absent (probably just missing); male Plp2 biramous, endopod only with
appendix interna distally. Telson subtriangular, though deformed (see fig. 6G
by Sakai, 2002).
Type locality. — Andaman Sea, 9◦ 30.351′ N 97◦ 57.168′ E, 60.7 m, sandy
mud, fine sand and shell fragments.
Distribution. — Andaman Sea, 9◦ 30.351′ N 97◦ 57.168′ E; 7◦ 29.921′ N
◦
99 00.977′ E; 20.5 m.
Paragourretia biffari (Blanco Rambla & Liñero Arana, 1994)
Gourretia biffari Blanco Rambla & Liñero Arana, 1994: 22, figs. 6, 7; Tudge et al., 2000: 142;
Sakai, 2005b: 222.
Diagnosis. — Rostrum short and triangular, with acute and upturned apex,
reaching middle of eyestalks. Eyestalks flattened, overreaching proximal half
length of A1 segment 1. Mxp3 ischium-merus pediform; merus with mesiodistal spine; exopod present. Male Plps1-2 undescribed. Telson narrowing toward
convex posterior margin, bearing low median convexity on lateral margin. [Cf.
Blanco Rambla & Lin̄ero Arana, 1994: 22, figs. 6, 7.]
Type locality. — Northwest Barcelona, 10◦ 15.30′ N 64◦ 42.30′ W, 50 m,
Venezuela.
Distribution. — Northwest Barcelona, Venezuela; off Trujillo, Honduras.
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CRM 013 – Katsushi Sakai
Paragourretia coolibah (Poore & Griffin, 1979)
Gourretia coolibah Poore & Griffin, 1979: 278, figs. 38, 39; Manning & Felder, 1991: 785
(list); Tudge et al., 2000: 142; Davie, 2002: 465; Sakai, 2005b: 223; Dworschak, 2009: 130,
figs. 1-34, tab. 1.
Diagnosis. — Rostrum acutely triangular, reaching middle of eyestalks.
Eyestalks compressed, tapering to broad mesiodistal lobe, and bearing a few
minute pigment spots subdistally. P1 unequal. Mxp3 ischium-merus pediform;
merus lacking mesiodistal spine; exopod present. Male Plps1-2 unknown.
Telson slightly broader than long, parallel-sided in proximal half, then tapering
to broadly rounded posterior margin. Uropodal exopod with notch on distal
margin. [Adapted from Poore & Griffin, 1979: 278, figs. 38c, 39f.]
Type locality. — 350 km ENE of Troughton Is., Joseph Bonaparte Gulf,
Western Australia, 58 m.
Distribution. — Only known from the type locality.
Paragourretia crosnieri (Ngoc-Ho, 1991)
Gourretia crosnieri Ngoc-Ho, 1991: 294, fig. 6; Tudge et al., 2000: 142; Sakai, 2005b: 223.
Diagnosis. — Rostrum acutely triangular, reaching middle of eyestalks.
Eyestalks rounded distally; cornea distinct and located centrally. Mxp3
ischium-merus pediform; merus lacking mesiodistal spine; exopod present.
Male Plps1-2 unknown. Telson longer than broad, broadened in proximal third,
then convergent to almost straight posterior margin. [Adapted from Ngoc-Ho,
1991: 294, fig. 6e, n.]
Type locality. — Prony Bay, Île Ouen, New Caledonia, 29 m.
Distribution. — Only known from the type locality.
Paragourretia lahouensis (Le Loeuff & Intès, 1974)
Gourretia lahouensis Le Loeuff & Intès, 1974: 28, fig. 5a-v; De Saint Laurent & Le Loeuff,
1979: 79 (key), 80; Manning & Felder, 1991: 785 (list); Tudge et al., 2000: 142; Sakai,
2005b: 221.
Diagnosis. — Rostrum triangular, reaching proximal third of eyestalks.
Eyestalks depressed and pointed mesiodistally; cornea located centrally. Mxp3
ischium-merus pediform; exopod present. Male Plp1 uniramous and bisegmented, distal segment subchelate distally; male Plp2 biramous, endopod
bearing at distal third on mesial margin appendix masculina with small
appendix interna mesiomedially. Telson longer than broad, broadened in
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
519
proximal third, then convergent to slightly rounded posterior margin. [Adapted
from Le Loeuff & Intès, 1974: 28, fig. 5j, p, r, r′ , s.]
Type locality. — Grand Lahou, Ivory Coast, 5◦ 7.4′ N 5◦ 4.5′ W, 15 m.
Distribution. — Only known from the type locality.
Paragourretia phuketensis (Sakai, 2002)
Gourretia phuketensis Sakai, 2002: 469, figs. 4A-H, 5A-H.
Paragourretia phuketensis — Sakai, 2005b: 226.
Diagnosis. – Rostrum sharply triangular, almost reaching distal end of
eyestalks. Eyestalks rounded distally and directed downward from rostrum,
cornea spotted and located centrally, in distal half. Mxp3 ischium-merus
pediform; merus with mesiodistal spine; exopod present. Male Plp1 uniramous
and bisegmented; male Plp2 biramous, endopod bearing at distal third of
mesial margin appendix masculina with appendix interna mesiomedially.
Telson longer than broad, broadened in proximal third, then convergent to
almost straight posterior margin. [Cf. Sakai, 2002: 469, figs. 4E, 5B, G.]
Type locality. — Andaman Sea, 8◦ 29.993′ N 98◦ 06.162′ E, 42.0 m, muddy
sand.
Distribution. — Only known from the type locality.
FAMILY LIPKECALLIANASSIDAE SAKAI, 2005 (NEW STATUS)
Lipkecallianassinae Sakai, 2005b: 212.
Diagnosis. — Rostrum acutely triangular and longer than eyestalks, lacking
rostral carina. Carapace without dorsal oval, cardiac prominence, and transverse hepatic sulcus. Eyestalks short, contiguous, and semicircular on lateral margins. Linea thalassinica present. A2 scaphocerite present as a small
process. Maxilla 2 scaphognathite without posterior whip. Mxp3 ischiummerus subpediform; merus with mesiodistal tooth; propodus rectangular;
dactylus digitiform. P1 uncertain. P2 chelate. P3 propodus rectangular. P4
elongate, overreaching P2. Abdominal somite 6 lacking lateral projections.
Male Plp1 uniramous and slender; male Plp3 biramous and foliaceous, endopod with appendix interna. Male Plps2, 4-5 and female Plps1-5 unknown. Telson longer than wide and slightly convergent toward w-shaped posterior margin with median spine. Uropodal endopod shaped as elongate triangle, about
twice as long as wide, and longer than telson. Uropodal exopod without dorsal
plate.
Remarks. — Lipkecallianassa Sakai, 2002, type genus of the family Lipkecallianassidae (new status) was originally included in the subfamily Callianassinae under the family Callianassidae. Subsequently (Sakai, 2005b) the
genus was distinguished from the other callianassid taxa into a separate subfamily, Lipkecallianassinae Sakai, 2005. However, the present revision has
made it possible to distinguish this genus even further from the other genera
and families included in the Callianassoidea, because Lipkecallianassa is very
characteristic and distinguishable; in the type-species of the genus Lipkecallianassa, Lipkecallianassa abyssa Sakai, 2002 the uropodal exopod lacks a
dorsal plate, and P4 is elongate, overreaching P2, whereas in the type species of
Callianassa, type genus of the Callianassidae, Callianassa subterranea (Montagu, 1808) the uropodal exopod bears a dorsal plate, and P4 does not overreach P2. Therefore, the genus Lipkecallianassa Sakai, 2002 was reclassified
and included into a new subfamily Lipkecallianassinae as the type genus, by
Sakai (2005b); this subfamily is now raised to family level (new status).
Type genus. — Lipkecallianassa Sakai, 2002.
Genera included. — Lipkecallianassa Sakai, 2002.
522
CRM 013 – Katsushi Sakai
Genus Lipkecallianassa Sakai, 2002
Lipkecallianassa Sakai, 2002: 477; Sakai, 2005b: 212.
Diagnosis. — Rostrum acutely triangular, overreaching eyestalks. Carapace
without dorsal oval and cardiac prominence, but with linea thalassinica. A1 peduncle shorter than A2 peduncle. Mxp3 ischium-merus subrectangular, with
parallel sides; merus with mesiodistal tooth; propodus rectangular; dactylus
digitiform; exopod absent. P1 uncertain. P3 propodus rectangular. P4 elongate,
overreaching P2. Male Plps1-2 unknown; male Plp3 biramous and foliaceous,
endopod with appendix interna. Telson longer than wide and slightly convergent toward w-shaped posterior margin with median spine. Uropodal exopod about 2.2 times as long as wide, much longer than endopod, and broadly
rounded on posterior margin, with setae, lacking dorsal plate.
Type species. — Lipkecallianassa abyssa Sakai, 2002, by original designation and monotypy. Gender of generic name, Lipkecallianassa, feminine.
Species included. — Only the type-species, Lipkecallianassa abyssa Sakai,
2002.
Lipkecallianassa abyssa Sakai, 2002
Lipkecallianassa abyssa Sakai, 2002: 477, figs. 9A-D, 10A-H; Sakai, 2005b: 213.
Type locality. — Andaman Sea, 7◦ 45.179′ N 97◦ 19.907′ E, 493.0 m.
Distribution. — Only known from the type locality.
FAMILY PSEUDOGOURRETIIDAE SAKAI, 2005 (NEW STATUS)
Pseudogourretiinae Sakai, 2005b: 230.
Diagnosis. — Rostrum triangular, lacking rostral carina. Carapace without
dorsal oval and cardiac sulcus, but with cardiac prominence. Linea thalassinica
incompletely present. Eyestalks flattened and contiguous. A2 scaphocerite
small. Maxilla 2 scaphognathite without posterior whip. Mxp3 ischiummerus rectangular; merus with subdistal tooth at mesiodistal angle; propodus
rectangular; dactylus digitiform; exopod present. Ps1-2 unknown. P3 propodus
rectangular. Pleurobranchs present on Ps2-4. Abdominal somite 6 unknown.
Male Plp1 uniramous and bisegmented, distal segment chelate. Male Plps2-5
and female Plps1-5 unknown. Uropodal exopod unknown.
Type genus. — Pseudogourretia Sakai, 2005b.
Genera included. — Pseudogourretia Sakai, 2005b.
Genus Pseudogourretia Sakai, 2005
Pseudogourretia Sakai, 2005b: 230.
Diagnosis. — Male Plp1 chelate; male Plp2 unknown. Pleurobranchs
present on Ps2-4.
Remarks. — Pseudogourretia is similar to Gourretia De Saint Laurent,
1973, Laurentgourretia Sakai, 2004b, and Paragourretia Sakai, 2004b, in that
the rostral carina is absent; the maxilla 2 scaphognathite bears no posterior
whip; and the male Plp1 is chelate distally, but differs from these, in that in
Pseudogourretia pleurobranchs are present on Ps2-4, whereas in the genera
Gourretia, Laurentgourretia, and Paragourretia these pleurobranchs are absent on Ps2-4. Therefore, Pseudogourretia was earlier assigned to its own subfamily (Sakai, 2005b), which is here reclassified as the family Pseudogourretiidae (new status).
Type species. — Pseudogourretia portsudanensis Sakai, 2005b by original
designation and monotypy. Gender of generic name, Pseudogourretia, feminine.
Species included. — Pseudogourretia portsudanensis Sakai, 2005b.
524
CRM 013 – Katsushi Sakai
Pseudogourretia portsudanensis Sakai, 2005
Pseudogourretia portsudanensis Sakai, 2005b: 231.
Diagnosis. — Rostrum triangular, pointed apically, reaching middle of eyestalks. Eyestalks oval, and obtuse distally; cornea small and located centrally.
Mxp3 ischium-merus pediform; merus with mesiodistal spine; exopod present.
P1 unknown. Male Plp1 uniramous, bisegmented, and chelate distally; male
Plp2 unknown. Telson unknown. [Adapted from Sakai, 2005b.]
Type locality. — Port Sudan (19◦ 43.3′ N 37◦ 40.5′ E – 19◦ 42.5′ N 37◦ 40.2′ E),
Red Sea, 517-583 m.
Distribution. — Only known from the type locality.
FAMILY THOMASSINIIDAE DE SAINT LAURENT, 1979
Thomassiniinae De Saint Laurent, 1979: 1396; De Saint Laurent & Le Loeuff, 1979: 95; Sakai,
1992c: 18.
Callianideidae — Kensley & Heard, 1991: 497 (partim).
Thomassiniidae — Poore, 1994: 104; Poore, 1997: 398.
Diagnosis. — Rostrum obsolete or triangular. Carapace without dorsal oval,
cardiac sulcus, cardiac prominence, and posterolateral lobes; linea thalassinica
completely or incompletely present. Cervical groove present, barely present, or
absent. Eyestalks flattened and contiguous, either completely, or except for distal part; cornea medially or distally present, or absent. A2 scaphocerite small
or absent. Maxilla 2 scaphognathite with posterior whip. Mxp3 ischial crista
dentata present or reduced to smooth carina; ischium and merus with or without brush of stiff setae; propodus rectangular; dactylus ovate with marginal setae; exopod present or absent. P1 subequal. P3 propodus rectangular or ovate.
P5 simple or subchelate. Epipods present on Mxps2-3 and Ps1-3, or Mxps23 and Ps1-4, those occasionally rudimentary. Abdominal somite 1 with row
of setae. Male Plp1 present or absent; male Plp2 biramous, endopod bearing appendices interna and masculina, or bearing only appendix interna; male
Plps3-5 endopods with appendix interna. Telson wider than long, and trapezoid. Uropodal endopod oval, bearing transverse row of setae at proximolateral angle. Uropodal exopod with or without dorsal plate, bearing neither distal
flap nor lateral notch. [Adapted from Kensley & Heard, 1991: 528.]
Remarks. — Kensley & Heard (1991: 497) included 7 genera (Callianidea,
Michelea, Mictaxius, Thomassinia, Crosniera, Marcusiaxius, and Meticonaxius) in the family Callianideidae. Later Poore (1997) classified eight genera
under three families, Callianideidae (Callianidea), and Thomassiniidae (Crosniera, Mictaxius, and Thomassinia) in the superfamily Callianassoidea, and
Micheleidae (Michelea, Tethisea, Meticonaxius, and Marcusiaxius) in the superfamily Axioidea.
In the Thomassiniidae the linea thalassinica shows a variety of patterns; it
is complete in Garyia gen. nov.; Mictaxius; and Heardaxius gen. nov.; and
incomplete in Crosniera, Ngochoaxius gen. nov., and Thomassinia, while in
the Callianideidae the linea thalassinica is absent as in Callianidea H. Milne
526
CRM 013 – Katsushi Sakai
Edwards, 1837a, though it can be present as a very short one, as in Paracallianidea Sakai, 1992c, so that the Callianideidae are safely reclassified under
the Axioidea.
Type genus. — Thomassinia De Saint Laurent, 1979, by being the nominate
genus.
Genera included. — Crosniera Kensley & Heard, 1991; Garyia gen. nov.;
Heardaxius gen. nov.; Mictaxius Kensley & Heard, 1991; and Thomassinia De
Saint Laurent, 1979.
K EY
TO THE
G ENERA
OF THE
FAMILY T HOMASSINIIDAE
1 – Uropodal exopod with dorsal plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mictaxius
– Uropodal exopod without dorsal plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – Linea thalassinica complete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Linea thalassinica incomplete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 – Male Plp1 bisegmented; male Plp2 appendices masculina and interna fused . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heardaxius gen. nov.
– Male Plp1 absent; male Plp2 appendix masculina with appendix interna. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Garyia gen. nov.
4 – Male Plp1 absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thomassinia
– Male Plp1 present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crosniera
TABLE XI
Species characters in the Thomassiniidae
Crosniera corindon
Crosniera dayrati
Crosniera minima
Crosniera panie
Crosniera sp. 1
Crosniera sp. 2
Garyia arno
Garyia moorea
Heardaxius salvati
Mictaxius thalassicola
Thomassinia aimsae
Thomassinia gebioides
Thomassinia sp. 1
Dorsal plate Linea
Male Plp1
of uropodal thalassinica
exopod
Male Plp2
absent
absent
absent
absent
absent
unknown
absent
absent
absent
present
absent
absent
absent
masculina and interna fused
unknown
masculina with interna
masculina and interna fused
masculina and interna fused
unknown
masculina and interna fused
masculina with interna
masculina and interna fused
masculina and interna fused
unknown
masculina and interna fused
masculina and interna fused
incomplete
incomplete
incomplete
incomplete
incomplete
unknown
complete
complete
complete
complete
incomplete
incomplete
incomplete
unsegmented
unknown
bisegmented
unsegmented
unsegmented
unknown
absent
absent
bisegmented
absent
unknown
absent
absent
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
527
Genus Crosniera Kensley & Heard, 1991
Crosniera Kensley & Heard, 1991: 500, fig. 2B; Poore, 1994: 104 (key); Poore, 1997: 399.
Diagnosis. — Rostrum triangular. Cervical groove incomplete. Linea thalassinica incompletely present. Eyestalk flattened, and pointed or obtuse distally; cornea medially present or reduced. A2 scaphocerite vestigial. Mxp3 ischial crista dentata present; ischium and merus without brush of stiff setae; exopod present. P1 subequal. P3 propodus rectangular. Male Plp1 uniramous, unsegmented and obtuse distally, or bisegmented, proximal segment elongate and
distal one short and leaf-like or obtuse distally. Male Plp2 biramous, elongate
and leaf-like, endopod bearing mesiomedially appendix masculina with stiff
distal setae, provided mesiomedially with appendix interna, or fused appendices masculina and interna. Male Plps3-5 endopods with appendix interna.
Telson with proximal lobe without spine on lateral margin, posterior margin
rounded. Uropodal exopod ovate, lacking dorsal plate. Setal rows present on
carapace and abdominal somite 6, and present or absent on abdominal somites
1-5. [Adapted from Kensley & Heard, 1991; Poore, 1997.]
Type species. — Callianassa minima Rathbun, 1901, by original designation. Gender of generic name, Crosniera, feminine.
Species included. — Crosniera corindon Poore, 1997; C. dayrati NgocHo, 2005; C. minima (Rathbun, 1901); C. panie Poore, 1997; Crosniera sp. 1,
Poore, 1997; Crosniera sp. 2, Poore, 1997.
K EY
TO THE
S PECIES
OF THE
G ENUS CROSNIERA
1 – Telson with distinct median spine on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . C. panie
– Telson without distinct spine on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 – A1 peduncle reaching distal end of A2 penultimate segment . . . . . . . . . . . . . . . C. minima
– A1 peduncle slightly longer than A2 peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 – Rostrum shaped as elongate triangle, reaching distal end of eyestalks . . . . . C. corindon
– Rostrum triangular, overreaching distal end of eyestalks . . . . . . . . . . . . . . . . . . . C. dayrati
Crosniera corindon Poore, 1997
Crosniera corindon Poore, 1997: 400, figs. 28, 29A-H.
Material examined. — MNHN Th 1217, holotype, 1 male (TL/CL, 15.0/4.0 mm), Makassar
Strait, 0◦ 14.9′ S 117◦ 51.7′ E, 150 m, 31.x.1980.
Diagnosis. — Scaphocerite short and drop-shaped. Male Plp1 uniramous
and unsegmented, and curved and cylindrical; male Plp2 endopod bearing
mesiomedially fused appendices interna and masculina, represented by distal
528
CRM 013 – Katsushi Sakai
hooklets and subdistal long setae on lateral margin. [Cf. Poore, 1997: 402,
fig. 29-M, O.]
Remarks. — It was confirmed by the present re-examination that the holotype regarded as a female by Poore is a male, the male Plp1 is uniramous and
cyrindrical, and the male Plp2 endopod bears mesiomedially fused appendices
interna and masculina, though the figures of Plps1-2 are shown as those of a
female by Poore, who considered the holotype to be a female.
Type locality. — Makassar Strait, Indonesia, 0◦ 14.9′ S 117◦ 51.7′ E, 150 m.
Distribution. — Indonesia, Makassar Strait (Poore, 1997); 150 m.
Crosniera dayrati Ngoc-Ho, 2005
(fig. 70)
Crosniera dayrati Ngoc-Ho, 2005: 63, figs. 8, 9.
Material examined. — MNHN Th 418, holotype, 1 female (TL/CL, ca. 20.0/5.2 mm,
posterior part of telson missing), Marquesas Islands, Nuku Hiva, MUSORSTOM 9, R/V “Alis”
Sta. 1306, 283-448 m, 10.ix.1997, leg. P. Bouchet, B. Dayrat, and B. Richer de Forges.
Diagnosis. — Eyestalks rounded, cornea pale. Linea thalassinica incomplete. Mxp3 merus with subdistal spine on mesial margin (fig. 70A). P2
chelate (fig. 70B). Ps3-4 propodi simple, lacking setal rows on ventrolateral
surfaces. Female Plp1 uniramous and bisegmented, distal segment simple; female Plps2-5 biramous, endopods with appendix interna.
Crosniera minima (Rathbun, 1901)
Callianassa minima Rathbun, 1901: 92, fig. 16; Schmitt, 1935b: 5 (key); Biffar, 1971a: 651
(key), 654 (key); De Saint Laurent & Le Loeuff, 1979: 95; Manning, 1987: 397 (list);
Manning & Felder, 1991: 765.
Callianassa (Cheramus) minima — Borradaile, 1903: 546; De Man, 1928b: 26 (list), 94, 97;
Schmitt, 1935a: 195.
Crosniera minima — Kensley & Heard, 1991: 503, figs. 5, 6, table 1B.
Material examined. — USNM 23779, syntypes, 1 male, carapace and abdomen separated,
lacking Plp1; 1 female, one carapace with abdomen; and pair of P1 (shown by Kensley & Heard,
1991, fig. 5E, F); USNM 266258, 1 female (TL/CL, 14.0/4.0 mm), Puerto Rico, Caribbean Sea,
11.vi.1980, leg. B.F. Kensley; USNM 253452, 1 male (TL/CL. 11.0/3.6 mm), S. of Mobile,
Alabama, United States, Gulf of Mexico, 12 m, 29.vi.1980, leg. B. Kensley; USNM 253543,
1 male (TL/CL, 10.0/3.5 mm, rostrum broken), San Juan, Puerto Rico, Caribbean Sea, 282
m, 10.vi.1980, leg. B.F. Kensley; USNM 24668, 1 female (no carapace), Fish Hawk Sta.
6062, Mayagüez Harbour, Puerto Rico, 45-55 m; USNM 1100476, 2 males (TL/CL, 12.0/4.0;
6.0/2.8 mm); 1 male (18.0/4.8 mm), A-141 (M2 A-1), Puerto Rico, deep water; 1 female
(15.0/5.3 mm), M2, A-2, Puerto Rico, deep water.
Diagnosis. — Rostrum acutely triangular, reaching beyond eyestalks. Linea
thalassinica incomplete, reaching posterior fourth of carapace. P1 dissimilar.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
529
Fig. 70. Crosniera dayrati (Ngoc-Ho, 2005). A, Mxp3, lateral view; B, P2, lateral view. A,
B, MNHN Th 418, holotype, 1 female (TL/CL, ca. 20.0/5.2 mm, posterior part of telson
missing), Marquesas Islands, Nuku Hiva, MUSORSTOM 9, R/V “Alis” Sta. 1306, 283-448 m.
Scale 1 mm.
Male Plp1 uniramous and bisegmented, distal segment leaf-like; male Plp2 biramous, endopod bearing mesiomedially appendix masculina with long stiff
setae, provided mesiomedially with slender appendix interna. Male Plps3-5
biramous, endopods with appendix interna. Female Plp1 uniramous and trisegmented. Telson with proximal lobe followed posteriorly by three denticles on
lateral margin, bearing setae on posterior margin. Uropodal endopod bearing
subdistal spine on anterior margin, and median carina on dorsal surface; uropodal exopod with two longitudinal carinae.
Type locality. — Mayagüez Harbour, Puerto Rico.
Distribution. — Alabama — off Mobile (Kensley & Heard, 1991); Puerto
Rico — (Kensley & Heard, 1991), Mayagüez Harbour (Rathbun, 1901; Poore,
1997); Venezuela; 35-75 m.
530
CRM 013 – Katsushi Sakai
Crosniera panie Poore, 1997
Crosniera panie Poore, 1997: 402, figs. 29I-N, 30.
Diagnosis. — Male Plp1 simple, curved, and cylindrical. Male Plp2 biramous, endopod with fused appendices interna and masculina, represented only
by long stiff setae.
Type locality. — East Lagoon, near Mt Panié, New Caledonia, 20◦ 33.25′ S
164◦ 49.3′ E, 40 m.
Distribution. — New Caledonia — East Lagoon (Poore, 1997); 40 m.
Crosniera sp. 1, Poore, 1997
Crosniera sp. 1, Poore, 1997: 402, fig. 31.
Distribution. — Sulu Archipelago (Poore, 1997); 2570 m.
Crosniera sp. 2, Poore, 1997
Crosniera sp. 2, Poore, 1997: 406.
Distribution. — Angeles Bay, Gulf of California (Poore, 1997); 4 m.
Genus Garyia gen. nov.
Diagnosis. — Rostrum obsolete. Cervical groove present. Linea thalassinica
complete. Eyestalks closely depressed and contiguous, with acute mesial apex,
cornea present apically. A2 scaphocerite absent. P1 subequal. P3 propodus rectangular. P5 simple. Male Plp1 absent; male Plp2 biramous, endopod bearing
subproximally appendix masculina with distal setae on mesial margin, provided mesiomedially with slender appendix interna; male Plps3-5 endopods
with appendix interna. Telson wider than long and trapezoid. Uropodal exopod ovate, lacking dorsal plate. Setal rows present on carapace and uropodal
endopod.
Remarks. — Thomassinia moorea Poore, 1997 is excluded from the genus
Thomassinia, because T. moorea is different from the type species of Thomassinia, T. gebioides De Saint Laurent, 1979 as follows. In T. moorea
the linea thalassinica is complete, and the Plp2 endopod bears an appendix
masculina with appendix interna; whereas in T. gebioides the linea thalassinica
is incomplete, running on the anterior three-fourths of the carapace only, and
the male Plp2 endopod bears fused appendices masculina and interna. Those
differences are to be considered as important characteristics at the generic level
rather than on the species level, and those two species are thus distinguishable
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
531
from each other at the generic level, so that T. moorea is reclassified under the
new genus Garyia gen. nov. as G. moorea.
Type species. — Thomassinia moorea Poore, 1997, by present designation.
The gender of the generic name, Garyia, is feminine.
Species included. — Garyia arno (Poore, 1997); G. moorea (Poore, 1997).
Etymology. — The new genus is named Garyia in honour of Dr. Gary C.
B. Poore, Museum of Victoria, who described the present species through his
study on Thalassinidea.
Garyia arno (Poore, 1997)
Mictaxius arno Poore, 1997: 407, figs. 32, 33A-G.
Material examined. — Mictaxius arno Poore, 1997, USNM 95570, holotype, 1 male
(TL/CL, 14.5/4.1 mm), S.E. of Arno lagoon, Arno Atoll, Marshall Islands, 22.vii.1950, leg.
J.W. Wells, det. G. Poore (L.B. Holthuis, 1957 as Callianassa).
Diagnosis. — Rostrum short and triangular. Eyestalks tapering to narrowly
protruded mediodistal apex; cornea spotted and located distally. Linea thalassinica complete. A1 peduncle as long as A2 peduncle. Mxp3 ischium without crista dentata; exopod small and digitiform. Male Plp1 absent; male Plp2
with totally fused appendices interna and masculina. Telson as long as basal
width, broadened at proximal third, then converging to slightly rounded posterior margin. [Adapted from Poore, 1997.]
Type locality. — Arno Atoll, Marshall Islands, 36 m.
Distribution. — Only known from the type locality.
Garyia moorea (Poore, 1997)
(figs. 71, 72)
Thomassinia moorea Poore, 1997: 416, fig. 38.
Fig. 71. Garyia moorea (Poore, 1997). Whole body, lateral view. MNHN Th 1220, holotype,
Tiahura, Moorea Is., Tahiti, 17◦ 30′ S 149◦ 50′ E. Scale 1 mm.
532
CRM 013 – Katsushi Sakai
Fig. 72. Garyia moorea (Poore, 1997). A, male Plp2; B, telson and uropod on left side; C, telson
and uropod on right side. A-C, MNHN Th 1220, holotype, Tiahura, Moorea Is., Tahiti, 17◦ 30′ S
149◦ 50′ E. Scale 1 mm.
Material examined. — MNHN Th 1220, holotype of Thomassinia moorea Poore, 1997,
1 male (TL/CL, 12.0/3.5 mm), Tahiti, Moorea Is., 17◦ 30′ S 149◦ 50′ E, leg. B.A. Thomassin.
Diagnosis. — Cephalothorax with submarginal setal row behind anterolateral margin (fig. 71). Linea thalassinica complete. Abdominal somites convex
or straight ventrally. Male Plp1 absent. Male Plp2 biramous, endopod bearing
mesially subproximal appendix masculina with distal setae, provided mesiomedially with slender appendix interna (fig. 72A). Telson wider than long and
trapezoid (fig. 72B, C). Uropodal endopod with transverse row of spiniform
setae. Uropodal exopod broadly rounded on distal margin, bearing no dorsal
plate.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
533
Type locality. — Moorea Is., Tiahura, 17◦ 30′ S 149◦ 50′ E, Tahiti.
Distribution. — Only known from the type locality.
Genus Heardaxius gen. nov.
Diagnosis. — Rostrum reduced to small triangle. Cervical groove weakly
present. Linea thalassinica complete. Eyestalks flattened and pointed distally;
cornea pigmented and situated subapically. A2 scaphocerite vestigial. Maxilla
2 scaphognathite with posterior whip. Mxp3 ischial crest moderately dentate;
ischium and merus without brush of stiff setae; dactylus ovate; exopod
vestigial. P1 subequal. P3 propodus subcircular. P5 simple or subchelate.
Male Plp1 uniramous and bisegmented; male Plp2 biramous, endopod bearing
mesiomedially fused appendices interna and masculina, represented by distal
hooklets and subdistal long setae on lateral margin; male Plps3-5 biramous,
endopods with appendix interna. Telson wider than long, and convergent
toward convex posterior margin without median spine, bearing proximal
lobe on lateral margin. Uropodal exopod ovate, lacking dorsal plate. Setal
rows present on carapace, Ps2-4, and abdominal somites 1-6. [Adapted from
Kensley & Heard, 1991; Poore, 1997.]
Remarks. — The type species of the genus Mictaxius, M. thalassicola Poore
& Heard, 1991 is similar to M. salvati, because in those two species the linea
thalassinica is complete, and the male Plp2 endopod bears mesiomedially
fused appendices interna and masculina, represented by distal hooklets and
subdistal long setae on the lateral margin. However, they are different from
each other at the generic level: in M. thalassicola the uropodal exopod bears
a dorsal plate, while in M. salvati the uropodal exopod lacks a dorsal plate,
so that M. salvati is excluded from Mictaxius, and reclassified under the new
genus Heardaxius as H. salvati.
Type species. — Mictaxius salvati Ngoc-Ho, 2005, by present designation
and monotypy. Gender of generic name, Heardaxius, masculine.
Species included. — Heardaxius salvati (Ngoc-Ho, 2005).
Etymology. — The new genus is named Heardaxius in honour of Dr.
Richard W. Heard, Gulf Coast Research Laboratory, Ocean Springs, MS, who
is one of the leading carcinologists in the U.S.A.
Heardaxius salvati (Ngoc-Ho, 2005)
Mictaxius salvati Ngoc-Ho, 2005: 66, figs. 10, 11.
Material examined. — MNHN Th 1417, holotype, hermaphroditic with ova (TL/CL,
14.0/4.0 mm), French Polynesia, Mururoa, Mission II Tuamotu, 40 m, 1965, B. Salvat coll.
534
CRM 013 – Katsushi Sakai
Diagnosis. — Rostrum weakly developed. Eyestalks slightly flattened,
and acute mesiodistally; cornea distinct and located distolaterally. Linea
thalassinica complete. Mxp3 ischium and merus rectangular; ischium with
weak crista dentata; bearing small exopod. P1 subequal. Telson slightly
broader than long, and broadened in proximal half, then tapering to slightly
rounded posterior margin. Uropodal endopod and exopod rounded, and about
as long as telson. Hermaphrodite. [Adapted from Ngoc-Ho, 2005.]
Remarks. — The hermaphroditic type-specimen is broken and in poor
condition.
Type locality. — Mururoa, French Polynesia, 40 m.
Distribution. — Only known from the type locality.
Genus Mictaxius Kensley & Heard, 1991
Mictaxius Kensley & Heard, 1991: 527; Poore, 1994: 104 (key); Poore, 1997: 399 (key), 406.
Diagnosis. — Rostrum obsolete. Carapace with cervical groove. Linea thalassinica complete. Eyestalks moderately flattened and pointed distally; cornea
moderately pigmented and situated subapically. A2 scaphocerite vestigial.
Maxilla 2 scaphognathite with posterior whip. Mxp3 ischial crest moderately
dentate; ischium and merus without brush of stiff setae; dactylus oval; exopod
vestigial. P1 subequal. P3 propodus subcircular. P5 simple or subchelate. Male
Plp1 absent; male Plp2 biramous, endopod bearing fused appendices interna
and masculina, represented by stiff distal setae and hooklets; male Plps3-5
biramous, endopods with appendix interna. Telson wider than long and semicircular. Uropodal exopod ovate, bearing dorsal plate. Setal rows present on
carapace, Ps2-4, and abdominal somites 1-6. [Adapted from Kensley & Heard,
1991; Poore, 1997.]
Remarks. — Kensley & Heard (1991: 527) established Mictaxius based
on the type species, M. thalassicola Kensley & Heard, 1991 from Pico Feo,
Atlantic Panama, in a shallow water Thalassia patch, as a new genus of the
family Callianideidae. Later Poore (1994: 104) excluded Mictaxius from the
Callianideidae, and included it in the Thomassiniidae De Saint Laurent, 1979.
The type species of the genus Mictaxius, M. thalassicola is distinguished
from the other three species (M. arno, M. salvati, and M. sp. 1, Poore, 1997)
that were hitherto classified under Mictaxius, because in M. thalassicola the
uropodal exopod bears a dorsal plate, whereas in the other three species the
uropodal exopod lacks a dorsal plate. Hence, these three last mentioned species
are herein reclassified, i.e., M. arno under Garyia gen. nov., M. salvati under
Heardaxius gen. nov., and M. sp. 1, Poore, 1997 under Thomassinia.
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
535
Type species. — Mictaxius thalassicola Kensley & Heard, 1991, by original
designation and monotypy. Gender of generic name, Mictaxius, masculine.
Species included. — M. thalassicola Kensley & Heard, 1991.
Mictaxius thalassicola Kensley & Heard, 1991
Mictaxius thalassicola Kensley & Heard, 1991: 527, figs. 22-24, table 1 L.; Poore, 1997: 408.
Material examined. — USNM 1011798, 1 male (TL/CL, 15.0/4.0 mm) and 2 females
(TL/CL, 15.0/4.0-19.0/4.5 mm), Klein Lagoen, Aruba, Netherlands Antilles, Caribbean Sea,
1.5 m, 10.i.1987, leg. R. Lemaitre.
Diagnosis. — Male Plp1 absent; male Plp2 biramous, endopod with fused
appendices masculina and interna; male Plps3-5 endopods with appendix
interna. Uropodal endopod without longitudinal carina; uropodal exopod with
longitudinal carina, bearing dorsal plate.
Type locality. — Pico Feo, Atlantic Panama.
Distribution. — Atlantic Panama — Pico Feo and Isla Mira, San Blas
Islands (Kensley & Heard, 1991).
Genus Thomassinia De Saint Laurent, 1979
Thomassinia De Saint Laurent, 1979: 1396; Kensley & Heard, 1991: 528; Poore, 1994: 104
(key); Poore, 1997: 399 (key), 410.
Diagnosis. — Rostrum obsolete. Cervical groove present dorsally. Linea
thalassinica incomplete. Eyestalks slightly flattened and contiguous; cornea
present apically. A2 scaphocerite absent. Mxp3 ischium without crista dentata;
ischium and merus with brush of stiff setae; dactylus ovate and setose;
exopod absent. P1 subequal. P3 propodus oval. P5 simple or subchelate. Male
Plp1 absent; male Plp2 biramous, endopod with fused appendices interna
and masculina; male Plps3-5 endopods with appendix interna. Telson wider
than long, evenly tapering toward straight posterior margin. Uropodal exopod
subcircular, and rounded at anterolateral corner, with dense marginal setae,
bearing no dorsal plate. Setal rows present on carapace and uropodal endopod.
[Adapted from Kensley & Heard, 1991; Poore, 1997.]
Remarks. — The type species of the genus Thomassinia, T. gebioides De
Saint Laurent, 1979 was described by Poore (1997: 413) as follows; “linea
thalassinica beginning at lateral margin of eyes, longitudinal and reaching
three-quarters along cephalothorax”. In T. aimsae Poore, 1997 the linea
thalassinica is as incomplete as in T. gebioides. However, it has turned out
through the present examination that in T. moorea Poore, 1997 included in
536
CRM 013 – Katsushi Sakai
the genus Thomassinia by Poore (1997), the linea thalassinica is complete.
Therefore, T. moorea is reclassified under the new genus Garyia.
Type species. — Thomassinia gebioides De Saint Laurent, 1979, by original
designation and monotypy. Gender of generic name, Thomassinia, feminine.
Species included. — Thomassinia aimsae Poore, 1997; T. gebioides De
Saint Laurent, 1979; Thomassinia sp. 1 (Poore, 1997).
Thomassinia aimsae Poore, 1997
Thomassinia aimsae Poore, 1997: 410, figs. 34, 35.
Diagnosis. — Rostrum weakly developed. Eyestalks flattened, contiguous,
and acute distally; cornea located distally. Linea thalassinica beginning at
lateral margin of eyestalks, and reaching posterior third of carapace. Mxp3
ischium with crista dentata. Telson shaped as inverted trapezoid. Uropodal
endopod triangular, bearing transverse row of 9 spiniform setae at anterolateral
corner. [Adapted from Poore, 1997.]
Type locality. — Davies Reef, Queensland, Australia, 18◦ 50′ S 147◦ 39′ E,
5 m.
Distribution. — Australia — Davies Reef, Queensland (Poore, 1997); 5 m.
Thomassinia gebioides De Saint Laurent, 1979
Thomassinia gebioides De Saint Laurent, 1979: 1396; Poore, 1997: 413, figs. 36, 37.
Material examined. — MNHN Th 818, paratype of Thomassinia gebioides, 1 female
(TL/CL, 14.0/3.2 mm); MNHN Th 822, paratypes of Thomassinia gebioides, 3 males, 2
females, Madagascar, Tuléar, leg. B.A. Thomassin.
MNHN Th 1220, Tiahura, Moorea Is., Tahiti, 17◦ 30′ S 149◦ 50′ E (= Thomassinia moorea
Poore, 1997, holotype).
Diagnosis. — Small species. Rostrum short and triangular in dorsal view.
Dorsal oval absent. Cervical groove present in middle of carapace; linea
thalassinica incomplete. Eyestalks slightly flattened and contiguous, cornea
distal, ommatidia scattered. A2 scaphocerite absent. Setal row present along
anterolateral margin of carapace. Mxp3 ischial crista dentata absent; ischium
and merus with brush of stiff setae, but different in shape. P1 unequal; in larger
cheliped fingers shorter than palm; in smaller cheliped fingers longer than
palm, cutting edge of fixed finger with distinct tooth. P2 chelate, fingers longer
than palm. P3 propodus subsquare. P4 propodus rectangular. P5 subchelate.
Abdominal somites 2-6 bearing tufts of fine setae along ventral margin. Male
Plp1 absent; male Plp2 biramous, endopod bearing elongate, fused appendices
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
537
masculina and interna, represented only by long setae and distal hooklets; male
Plps3-5 biramous, endopods with elongate appendix interna.
Type locality. — Tuléar, Madagascar.
Distribution. — Madagascar, Tuléar (De Saint Laurent, 1979; Poore, 1997).
Thomassinia sp. 1 (Poore, 1997)
Mictaxius sp. 1; Poore, 1997: 408, figs. 33H-J.
Diagnosis. — Linea thalassinica incomplete. Male Plp1 absent. Male Plp2
biramous, endopod with fused appendices interna and masculina. Uropodal
exopod without dorsal plate. [Cf. Poore, 1997: 408, fig. H, I, J.]
Remarks. — The present species, which was defined as a species of the
genus Mictaxius, is different, however, from the type species of Mictaxius, M.
thalassicola, in that the linea thalassinica is incomplete, so that Mictaxius sp.
1 Poore is here reclassified under the genus Thomassinia, as T. sp. 1 (Poore,
1997).
Distribution. — Tahiti — Tiahura, Moorea Island (Poore, 1997).
DATA ON BIOGEOGRAPHY
In order to facilitate references to the ranges of various species, the known
distribution of the species of the family Axiidae as well as of those of the
subfamilies Callianassinae and Callichirinae of the family Callianassidae, are
here summarized in three tables, i.e., tables XII, XIII, and XIV.
TABLE XII
The distribution of the species in the family Axiidae as currently known
Atlantic Ocean and Mediterranean Sea
Mediterranean Sea
Axius mediterraneus
Axius stirhynchus
Levantocaris hornungae
West Africa
Calocarides capensis
Calocarides macphersoni
Calocarides rudolfi
Manaxius kensleyi
North and western Atlantic
Axiopsis serratifrons
Axiorygma nethertoni
Axius armatus
Axius serratus
Axius stirhynchus
Bouvieraxius longipes
Calocarides coronatus
Calocarides laevis
Coelhocalaxius spinosus
Eutrichocheles defensus
Eutrichocheles foveolatus
Eutrichocheles granulimanus
Eutrichocheles spinipleurus
Guyanacaris hirsutimana
Heterocalaxius carneyi
Manaxius angulatus
Manaxius inaequalis
Paraxiopsis gracilimana
Paraxiopsis hispida
Paraxiopsis pindatyba
Paraxiopsis vicina
Pillsburyaxius kirkmilleri
540
CRM 013 – Katsushi Sakai
TABLE XII
(Continued)
Indo-Pacific
Eastern Pacific
Axiopsis baronai
Calocarides lev
Calocarides quinqueseriatus
Calocarides rostriserratus
Guyanacaris caespitosa
Leonardsaxius spinulicauda
Manaxius galapagensis
Manaxius jenneri
Hawaii, central and southern Pacific,
and eastern Australia
Acanthaxius clevai
Acanthaxius garawa
Acanthaxius gadaletae
Acanthaxius miyazakiensis
Acanthaxius spinosissimus
Albatrossaxius bythos
Alienaxiopsis lizardensis
Allaxiopsis bougainvillensis
Axiopsis consobrina
Axiopsis serratifrons
Bouvieraxius rudis
Bruceaxius polychaetes
Calaxius acutirostris
Calocarides vigilus
Dorphinaxius kermadecensis
Eutrichocheles modestus
Eutrichocheles tuamotu
Leonardsaxius werribee
Manaxius aganaensis
Manaxius euophthalmus
Manaxius pailoloensis
Marianaxius kroppi
Michelaxiopsis australiensis
Newzealandaxius novaezealandiae
Oxyrhynchaxius tricarinatus
Parascytoleptus tridens
Paraxiopsis brocki
Paraxiopsis johnstoni
Paraxiopsis majuro
Paraxiopsis paulayi
Paraxiopsis plumosimanus
Paraxius altus
Planaxius brevifrons
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
541
TABLE XII
(Continued)
Indo-Pacific
Platyaxiopsis odontorhyncha
Pseudoaxiopsis carpentariensis
Scytoleptus barbatus
Spongiaxius brucei
Japan, eastern China, and Taiwan
Acanthaxius miyazakiensis
Acanthaxius pilocheirus
Amakusaxius amakusanus
Axiopsis consobrina
Balssaxius habereri
Boasaxius princeps
Calaxius acutirostris
Calocarides chani
Calocarides okhotskensis
Calocarides soyoi
Leonardsaxius amurensis
Litoraxius boshu
Manaxius mimasensis
Manaxius ohsumiensis
Oxyrhynchaxius japonicus
Paraxiopsis brocki
Planaxius brevifrons
Sarawak, Gulf of Thailand
Manaxius thailandensis
Indonesia
Acanthaxius miyazakiensis
Acanthaxius pilocheirus
Acanthaxius spinosissimus
Allaxiopsis picteti
Allaxiopsis bougainvillensis
Allaxiopsis spinimana
Anophthalmaxius eccoptodactylus
Axiopsis consobrina
Axiopsis serratifrons
Bouvieraxius michelae
Bouvieraxius springeri
Calocarides tenuicornis
Colemanaxius pitatucensis
Manaxius euophthalmus
Manaxius sibogae
Paraxiopsis brocki
Ralumcaris bisquamosa
542
CRM 013 – Katsushi Sakai
TABLE XII
(Continued)
Indo-Pacific
Northern and Western Australia
Acanthaxius clevai
Acanthaxius ningaloo
Acanthaxius pilocheirus
Acanthaxius spinosissimus
Australocaris pinjarup
Bouvieraxius keiensis
Bouvieraxius rudis
Eutrichocheles austrinus
Leonardsaxius werribee
Leonardsaxius sp. aff. werribee
Oxyrhynchaxius manningi
Paratrichocheles pumilus
Paraxiopsis brocki
Pilbaraxius kariyarra
Planaxius brevifrons
Platyaxius bardi
Platyaxius brevirostris
Scytoleptus barbatus
Spongiaxius brucei
India
Axiopsis serratifrons
Eutrichocheles modestus
Persian Gulf and Gulf of Oman
Calocarides longispinis
Gulf of Aden
Allaxius aethiopicus
Axiopsis serratifrons
Red Sea
Allaxius aethiopicus
Axiopsis serratifrons
Madagascar
Bouvieraxius rudis
Calaxius acutirostris
Parascytoleptus tridens
Scytoleptus serripes
East and South Africa
Axiopsis serratifrons
Axiopsis sp.
Calaxius acutirostris
Calocarides capensis
Michelaxiopsis australiensis
Scytoleptus serripes
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
543
TABLE XIII
The distribution of the species in the subfamily Callianassinae as currently known
Atlantic Ocean and Mediterranean Sea
Mediterranean Sea
Callianassa subterranea
Gilvossius candidus
Gilvossius tyrrhenus
Gilvossius whitei
Trypaea acanthura
Trypaea truncata
West Africa
Callianassa subterranea
Cheramus oblongus
Gilvossius candidus
Gilvossius convexus
Gilvossius diaphorus
Gilvossius marchali
Gilvossius tyrrhenus
North and western Atlantic
Cheramoides marginata
Cheramus gaucho
Cheramus profundus
Gilvossius setimanus
Notiax delicatulus
Trypaea berylae
Trypaea biformis
Trypaea caesari
Trypaea costaricensis
Trypaea fragilis
Indo-Pacific
Eastern Pacific
Gilvossius pacificus
Notiax brachyophthalma
Trypaea biffari
Trypaea californiensis
Trypaea debilis
Trypaea gigas
Trypaea rochei
Trypaea santarita
Trypaea tabogensis
Trypaea uncinata
Hawaii, central and south Pacific,
and eastern Australia
Cheramus modestus
Gilvossius bouvieri
Notiax ceramica
Notiax limosa
Notiax ngochoae
544
CRM 013 – Katsushi Sakai
TABLE XIII
(Continued)
Indo-Pacific
Trypaea amboinae
Trypaea arenosa
Trypaea australiensis
Trypaea bangensis
Trypaea caledonica
Trypaea filholi
Trypaea gruneri
Trypaea lewtonae
Trypaea parva
Trypaea poorei
Trypaea tonkinae
Japan, eastern China, and Taiwan
Cheramus longicauda
Cheramus spinophthalmus
Gilvossius bouvieri
Trypaea exilimaxilla
Trypaea japonica
Trypaea joculatrix
Trypaea petalura
Trypaea thermophila
Sarawak, Gulf of Thailand
Cheramus anoplourus
Cheramus contipes
Cheramus malaccaensis
Cheramus nigroculatus
Gilvossius brachytelson
Trypaea amplimaxilla
Trypaea brevirostris
Trypaea chakratongae
Trypaea lignicola
Trypaea matzi
Trypaea nieli
Trypaea propriopedis
Trypaea spinoculata
Trypaea stenomastaxa
Trypaea tenuipes
Trypaea thailandica
Trypaea tonkinae
Indonesia
Cheramus lobetobensis
Cheramus modestus
Cheramus praedatrix
Cheramus propinquus
Gilvossius bouvieri
Notiax amboinensis
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
545
TABLE XIII
(Continued)
Indo-Pacific
Notiax ngochoae
Trypaea amboinae
Trypaea intermedia
Trypaea joculatrix
Trypaea parva
Trypaea pugnatrix
Trypaea pygmaea
Trypaea sahul
Trypaea sibogae
Northern and Western Australia
Cheramus orientalis
Cheramus parvulus
Cheramus propinquus
Notiax ceramica
Notiax ngochoae
Trypaea acutirostella
Trypaea joculatrix
Trypaea melissae
Trypaea poorei
Trypaea sibogae
India
Gilvossius gravieri
Trypaea pugnatrix
Persian Gulf and Gulf of Oman
Gilvossius persicus
Trypaea thorsoni
Gulf of Aden
Gilvossius bouvieri
Gilvossius gravieri
Red Sea
Cheramus aqabaensis
Gilvossius bouvieri
Gilvossius gravieri
Notiax ngochoae
Madagascar
–
East and South Africa
Callianassa subterranea
Gilvossius rotundicaudatus
Trypaea joculatrix
Trypaea mocambiquensis
Trypaea plantei
546
CRM 013 – Katsushi Sakai
TABLE XIV
The distribution of the species in the subfamily Callichirinae sensu nov. as currently known
Atlantic Ocean and Mediterranean Sea
Mediterranean Sea
–
West Africa
Balsscallichirus balssi
Balsscallichirus guineensis
Barnardcallichirus tenuimanus
Callichirus adamas
Corallichirus intesi
Forestcallichirus foresti
Neocallichirus pentagonocephalus
Neocallichirus sassandrensis
Sergio monodi
Sergio pachydactylus
Western Atlantic
Callichirus islagrande
Callichirus major
Glypturoides trilobatus
Glypturus acanthochirus
Glypturus longiventris
Glypturus rabalaisae
Lepidophthalmoides eiseni
Lepidophthalmoides jamaicense
Lepidophthalmoides louisianensis
Lepidophthalmoides richardi
Lepidophthalmoides siriboia
Lepidophthalmus sinuensis
Lepidophthalmus turneranus
Neocallichirus branneri
Neocallichirus grandimanus
Neocallichirus guaiqueri
Neocallichirus guarus
Neocallichirus mirim
Neocallichirus nickellae
Neocallichirus rathbunae
Sergio cacahuate
Sergio guassutinga
Sergio lemaitrei
Sergio mericeae
Sergio sulfureus
Indo-Pacific
Eastern Pacific
Callichirus seilacheri
Corallichirus xuthus
Lepidophthalmus bocourti
Lepidophthalmus rafai
Neocallichirus mortenseni
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
547
TABLE XIV
(Continued)
Indo-Pacific
Hawaii, central and south Pacific,
and eastern Australia
Corallichirus bayeri
Glypturus armatus
Glypturus articulatus
Glypturus assimilis
Glypturus collaroy
Glypturus lanceolatus
Glypturus longiventris
Glypturus martensi
Lepidophthalmoides rosae
Neocallichirus denticulatus
Neocallichirus indicus
Neocallichirus karumba
Neocallichirus mucronatus
Neocallichirus winslowi
Japan, eastern China, and Taiwan
Glypturus lanceolatus
Glypturus longiventris
Glypturus martensi
Grynaminna tamakii
Lepidophthalmoides tridentatus
Neocallichirus kempi
Sarawak, Gulf of Thailand
Neocallichirus kempi
Thailandcallichirus ranongensis
Indonesia
Glypturus armatus
Glypturus assimilis
Glypturus longiventris
Glypturus martensi
Lepidophthalmoides rosae
Lepidophthalmoides tridentatus
Michaelcallianassa indica
Neocallichirus indicus
Neocallichirus kempi
Neocallichirus moluccensis
Neocallichirus mucronatus
Neocallichirus vigilax
Thailandcallichirus ranongensis
Northern and Western Australia
Glypturus haswelli
Glypturus lanceolatus
Neocallichirus angelikae
Neocallichirus caechabitator
Neocallichirus darwinensis
Neocallichirus horneri
548
CRM 013 – Katsushi Sakai
TABLE XIV
(Continued)
Indo-Pacific
India
Glypturus armatus
Glypturus borradailei
Glypturus longiventris
Glypturus martensi
Michaelcallianassa indica
Neocallichirus audax
Neocallichirus indicus
Neocallichirus kempi
Neocallichirus manningi
Neocallichirus mauritianus
Tirmizicallichirus masoomi
Persian Gulf and Gulf of Oman
Lepidophthalmoides socotrensis
Michaelcallianassa indica
Neocallichirus calmani
Neocallichirus indicus
Neocallichirus jousseaumei
Neocallichirus mucronatus
Gulf of Aden
Glypturus armatus
Glypturus longiventris
Neocallichirus calmani
Neocallichirus indicus
Neocallichirus jousseaumei
Neocallichirus mucronatus
Red Sea
Glypturus armatus
Lepidophthalmoides rosae
Neocallichirus calmani
Neocallichirus indicus
Neocallichirus jousseaumei
Neocallichirus mauritianus
Madagascar
Glypturus longiventris
Lepidophthalmoides rosae
Lepidophthalmus grandidieri
Neocallichirus mucronatus
Podocallichirus madagassus
East and South Africa
Barnardcallichirus gilchristi
Callichirus adamas
Callichirus kraussi
Neocallichirus indicus
Neocallichirus mucronatus
ACKNOWLEDGEMENTS
I am most grateful to Prof. Dr. Michael Türkay of the Forschungsinstitut
Senckenberg, Frankfurt am Main (SMF) for his support, which enabled me to
pursue the present work on Axioidea and Callianassoidea. I am much indebted
to Mrs. Sonja Spal and Mrs. Gülay Sagirli, the Alexander von HumboldtStiftung, Germany, who provided me with financial support for my stay in
Europe. I am also heartily thankful to Dr. J. Carel von Vaupel Klein, University of Leiden, and editor of the series Crustaceana Monographs, for making inevitable arrangements for the publication of this monograph; to the late
Dr. Lipke B. Holthuis and Dr. C.H.J.M. Fransen of the National Museum of
Natural History, Leiden, for giving me very useful advice on nomenclatorial
problems; to Dr. P. Clark, the Natural History Museum, London, who kindly
showed me the references of Leach, 1815-16; and to Dr. P.A. Coelho, Universidade Federal de Pernambuco, PE, Brazil, who kindly sent me his papers and
the data of his new taxa on my request.
My thanks are extended to Emeritus Prof. S. Ohta, Ocean Research Ecology
Division, University of Tokyo (ORE), who invited me to participate in his
dredging operations by the R/V “Tansei-Maru” in 1993.
I wish to thank the following individuals who entrusted me with the important specimens necessary for my study: M. Türkay, A. Allspach, P. Friesleber,
and K. Pietratus in Frankfurt am Main (SMF); S. Keable, Australian Museum,
Sydney (AM); R.M. Shimizu, Museu Nacional do Rio de Janeiro (MHN,
Rio de Janeiro); A. Crosnier and R. Cleva, Muséum national d’Histoire naturelle, Paris (MNHN); P. Clark and M. Lowe, the Natural History Museum,
London (NHM); P. Dworschak, Naturhistorisches Museum, Wien (NHMW);
S. Ohta, Ocean Research Institute, University of Tokyo, Tokyo (ORIUT);
L.B. Holthuis, C.H.J.M. Fransen, C. Pepermans, and I.J. Smit, Nationaal
Natuurhistorisch Museum — Naturalis (RMNH) [formerly: Rijksmuseum van
Natuurlijke Historie], Leiden; C. Magalhaes, Instituto Nacional de Pesquisas
da Amazonia, Coordenação de Pesquisas em Biologia Aquatica, Manaus,
AM, Brazil; T.-M. Chan, Institute of Marine Biology, National Taiwan Ocean
University, Keelung Taiwan (NTOU); H. Namikawa, National Science Museum, Showa Memorial Institute, Tsukuba, Japan (NSMT R); T. Komai, Natural History Museum and Institute, Chiba, Japan (CBM); R. Lemaitre and
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K. Reed, National Museum of Natural History, Washington, D.C. (USNM);
D. Platvoet, Zoölogisch Museum Amsterdam, Universiteit van Amsterdam
(ZMA); C.O. Coleman, Zoologisches Museum, Humboldt-Universität zu
Berlin (ZMB); T. Wolff and B. Bisballe, Zoologisk Museum, University
of Copenhagen, Copenhagen (ZMC); V. Spiridonov, Zoological Museum of
Moscow, M.V. Lomonosov State University, Moscow (ZMM); and
S. Friedrich, Zoologische Staatssammlung, Munich (ZSM).
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1923-1930. Journ. Col. Agric., Tokyo Imp. Univ., 12(1): 1-226, figs. 1-71, tabs. 1-4.
— —, 1939. Macrura and Anomura of decapod Crustacea found in the neighbourhood of
Onagawa, Miyagi-ken. Sci. Rep. Tohoku Imp. Univ., (4) 14(2-3): 261-289, figs. 1-13.
YONG , C.G., 1900. The stalk-eyed Crustacea of British Guiana, West Indies, and Bermuda.
1-514, colour pl. 1. (J.M. Watkins, London).
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592
CRM 013 – Katsushi Sakai
Z ARIQUIEY C ENARRO , R., 1935. Adiciones al ensayo de un catalogo de los Crustáceos
Decápodos mariños de España y Marruecos Español de D. Alvaro de Miranda y Rivera,
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Z ARKANELLAS , A. & C. B OGDANOS, 1977. Benthic studies of a polluted area in the upper
Saronikos Gulf. Thalassographica, 1(2): 155-177.
Z EHNTNER , L., 1894. Crustacés de l’archipel Malais. Rev. Suisse Zool., 2: 135-214, pls. 7-9.
Z HONG , Z.-R., 2000. A new species of Axiidae from the South China Sea (Crustacea:
Decapoda: Thalassinidea). Act. zootax. Sinica, 25(1): 33-36, figs. 1-11.
Z IMSEN , E., 1964. The type material of I.C. Fabricius: 1-656. (Munksgaard, Copenhagen).
ADDENDUM I
Meanwhile, Liu & Liu (2009) have described a second species in the genus
Michaelcallianassa, i.e., M. sinica Liu & Liu, 2009 from the South China
Sea. The authors mentioned that their species, Michaelcallianassa sinica, is
different from the type species of the genus, M. indica Sakai, 2002.
I here include this additional species as follows.
Michaelcallianassa sinica Liu & Liu, 2009
Michaelcallianassa sinica Liu & Liu, 2009: 40, figs. 1-5.
Remarks. — The present species is added as the second species of the
genus Michaelcallianassa from the Beibu Gulf (Tonkin Gulf) of the northern
South China Sea. M. sinica differs from the type species of Michelcallianassa,
M. indica Sakai, 2002 in the short uropodal endopod and exopod, and the
elongated carpus of the smaller cheliped. In addition, another difference is
observed in the shapes of the eyestalks: in M. indica the eyestalks are tapering
distally, while in M. sinica they are distally obtuse (Liu & Liu, 2009, fig. 2A).
REFERENCE
L IU , W. & L. RUIYU [= J. Y. L IU], 2009. Michaelcallianassa sinica sp. nov. (Crustacea,
Decapoda, Thalassinidea, Callianassidae) from the South China Sea. Zootaxa, 2294: 3946.
ADDENDUM II
Recently, Anker (2010) has reported a new genus and species from Nosi-Bé,
Madagascar, Ctenocheloides attenboroughi Anker, 2010, assigning the new
genus to the family Ctenochelidae Manning & Felder, 1991. However, according to the criteria adopted in the present monograph, this species does not belong in the Ctenochelidae, because of the following considerations. Ctenocheloides attenboroughi bears a moderately developed dorsal oval, and the uropodal exopod bears neither a dorsal plate nor a lateral notch, and is, therefore,
different from the other species currently included in the Ctenochelidae. This
family, viz., is not only characterized by: (1) the rostrum is distinct and acutely
triangular, (2) the P1 chelipeds are unequal in size, and dissimilar in shape,
and (3) the fingers of the larger cheliped being elongate and pectinate, but
also by (4) the dorsal oval being absent and (5) the uropodal exopod lacking
a dorsal plate, but bearing a lateral notch (please refer to table VIII herein).
As a result, the genus Ctenocheloides is here separated at family level from
the Ctenochelidae as a new family, Ctenocheloidae fam. nov., thus constituting the 11th family in the superfamily Callianassoidea Dana, 1852a, according
to the present classification.
Family C TENOCHELOIDAE fam. nov.
Diagnosis. — Carapace with moderately developed dorsal oval; cardiac
prominence present; cervical groove clearly delimited; linea thalassinica running complete length of carapace. Rostrum weakly developed, lacking cardiac
mediodorsal carina with mid-pit. Eyestalks somewhat flattened dorsally. A2
scaphocerite strong. Maxilla 2 scaphognathite without posterior whip. Mxp3
ischiomerus pediform; dactylus digitiform; exopod absent. P1 chelate, and almost equal in size and similar in shape; ischium armed with teeth on ventral margin; merus with small tooth on ventral margin; palm ovoid; finger
elongate and pectinate. Plp2-5 biramous, and endopods with stubby appendix
interna. Telson trapezoid. Uropodal exopod without lateral notch and dorsal
plate. [Modified from Anker, 2010.]
Remarks. — This new family is similar to the Ctenochelidae in that
the fingers of the larger cheliped of P1 are pectinate, but otherwise they
596
CRM 013 – Katsushi Sakai
are fundamentally different from each other, because in the present new
family Ctenocheloidae the rostrum is weakly developed; the carapace bears
a moderately developed dorsal oval; P1 are chelate and almost equal in size
and similar in shape; the ischium is armed with teeth on the ventral margin;
the merus is armed with a small tooth on the ventral margin; the fingers
of both chelipeds are elongate and pectinate; the telson is trapezoid; the
uropodal exopod bears neither a lateral notch nor a dorsal plate. Whereas in
the Ctenochelidae, the rostrum is distinct, acutely triangular, and unarmed
on the lateral margins; the carapace bears no dorsal oval; P1 are chelate,
unequal in size, and dissimilar in shape; in the larger cheliped the ischium
and merus bear no teeth ventroproximally; the palm is subglobular; the fingers
are elongate and pectinate; in smaller cheliped the ischium and merus bear no
teeth ventroproximally; the fingers are elongate but not pectinate; the telson
is subquadrate to trapezoid; the uropodal exopod bears a lateral notch, but no
dorsal plate.
Type genus. — Ctenocheloides Anker, 2010, as the nominate genus.
Genera included. — Only the type genus is known.
Ctenocheloides Anker, 2010
Ctenocheloides Anker, 2010: 1790.
Diagnosis. — Same as for the family, above.
Type species. — Ctenocheloides attenboroughi Anker, 2010, by original
designation, and monotypy. The gender of the generic name is masculine.
Species included. — Only the type species is known.
Ctenocheloides attenboroughi Anker, 2010
Ctenocheloides attenboroughi Anker, 2010: 1789, figs. 1-8, table 1.
Diagnosis. — With the characters of the genus.
Remarks. — Only the female holotype is known.
Type locality. — Madagascar, Nosy-Bé, the large bay east of Hellville, 1.5
m depth.
REFERENCE
A NKER , A., 2010. Ctenocheloides attenboroughi n. gen., n. sp. (Crustacea: Decapoda: Axiidea:
Ctenochelidae), a new ghost shrimp with pectinate claw fingers from Madagascar. Journ.
nat. Hist., London, 44(29-30): 1789-1805, figs. 1-6, table 1.
ERRATA
In:
S AKAI , K., 2005b. Callianassoidea of the world (Decapoda, Thalassinidea). Crustaceana
Monographs, 4: i-vi, 1-285, figs. 1-44.
Please correct:
Error: . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Correction:
p. 17, line 8 from bottom: Pseudotrypaea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucallichirus
p. 31, line 6 from bottom: Alupka — Black Sea [the type locality of Callianassa candida] . . . .
. . . . . . . . . . . . . . . . . . . . Kuvi Bay, ca. 2.5 km South of Rovinj, Istria, Croatia (Sakai, 1999c: 16).
pp. 35, 36, captions of figs. 5, 6: Callianassa subterranea (Montagu, 1808). SMF 25800 . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . SMF 27250
p. 37, caption fig. 7: Callianassa subterranea (Montagu, 1808). SMF 27250 . . . . . SMF 25800
p. 43, line 6 from top: Mediterranean [the type locality of Callianassa whitei] . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rovinj, Adriatic Sea.
p. 236, line 10 from bottom: Eucallichirus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucalliax
Please add in References:
M ELO , G. A. S. D E, 1999. Manuel de identificação dos Crustacea Decapoda do litoral
brasileiro: Anomura, Thalassinidea, Palinuridea, Astacidea: 1-551. (Editora Pleiâde/
FAPESP, São Paulo).
TAXONOMIC INDEX
Acanthaxius, 3, 10, 16, 18–21, 45, 83, 120, 133,
170, 508
Acanthaxius amakusana, 45
Acanthaxius angulatus, 133
Acanthaxius caespitosa, 120
Acanthaxius clevai, 20, 21, 540, 542
Acanthaxius formosa, 20, 21, 24, 25
Acanthaxius gadaletae, 20–22, 540
Acanthaxius garawa, 21, 22, 24, 540
Acanthaxius gathaagudu, 21, 28
Acanthaxius grandis, 20, 21, 28, 29
Acanthaxius hirsutimanus, 121
Acanthaxius kirkmilleri, 20, 170
Acanthaxius miyazakiensis, 20, 21, 24, 25, 540,
541
Acanthaxius ningaloo, 21, 26, 542
Acanthaxius pilocheira, 28
Acanthaxius pilocheirus, 20, 21, 28, 45, 83, 120,
541, 542
Acanthaxius polyacanthus, 20, 21, 24, 25
Acanthaxius polychaetes, 20, 83
Acanthaxius spinosissimus, 20, 21, 29, 540–542
Acanthaxius spinulicaudus, 126
Albatrossaxius, 30, 32
Albatrossaxius gen. nov., 3, 10, 16, 17, 29
Albatrossaxius bythos, 30–32, 540
Albatrossaxius sp., 30, 31
Alienaxiopsis, 32, 33
Alienaxiopsis gen. nov., 3, 10, 16, 17, 32
Alienaxiopsis clypeata, 32, 33, 35, 36
Alienaxiopsis lizardensis, 36, 540
Alienaxiopsis lizardensis sp. nov., 32–35
Allaxiopsis, 36, 37
Allaxiopsis gen. nov., 3, 10, 16, 17, 36
Allaxiopsis bougainvillensis, 37, 540, 541
Allaxiopsis bougainvillensis sp. nov., 36–38
Allaxiopsis picteti, 36, 37, 39, 40, 541
Allaxiopsis spinimana, 36, 37, 40, 541
Allaxius, 3, 10, 16, 17, 32, 36, 37, 40, 41, 76,
508
Allaxius aethiopica, 43
Allaxius aethiopicus, 32, 41, 42, 76, 77, 542
Allaxius clypeatus, 33
Allaxius lizardensis sp. nov., 32
Allaxius picteti, 39
Allaxius princeps, 77
Allaxius sculptimanus, 58
Allaxius spinimanus, 40
Alpheus Tyrhenus, 379, 380
Amakusaxius, 45
Amakusaxius gen. nov., 3, 12, 16, 17, 44
Amakusaxius amakusanus, 45, 541
Ambiaxius, 4, 10, 210, 211, 213, 508
Ambiaxius aberrans, 214
Ambiaxius alcocki, 211, 212
Ambiaxius foveolatus, 214
Ambiaxius franklinae, 211, 212
Ambiaxius japonicus, 213, 214
Ambiaxius sp., 211, 212
Ambiaxius surugaensis, 211, 212
Anacalliacidae, 3, 11, 335–338, 340–342, 353
Anacalliacinae, 335, 336, 341, 353, 355, 485
Anacalliaopsis, 341, 342
Anacalliaopsis gen. nov., 5, 11, 337, 341, 342
Anacalliaopsis agassizi, 342
Anacalliax, 5, 11, 335, 337, 341–343, 345, 485
Anacalliax agassizi, 342
Anacalliax argentinensis, 343–345
Anacalliax pixii, 345
Anacalliinae, 341
Andamancalliax, 495
Andamancalliax gen. nov., 6, 11, 339, 494
Andamancalliax andamanica, 495
Anophthalmaxius, 3, 10, 16, 45
Anophthalmaxius eccoptodactylus, 46, 47, 541
Anophthalmaxius enoptodactylus, 46
Astacus modestus, 112
Astacus Scaber, 112
Astacus scaber, 112, 116
Astacus tyrrhenus, 372, 373, 379, 380
Austinogebia edulis, 2
Australocaris, 3, 11, 16, 18, 48
Australocaris pinjarup, 48, 542
Axia, 65
Axia acantha, 326
Axia acanthus, 325
600
CRM 013 – Katsushi Sakai
Axia serratifrons, 51, 56
Axia stirhynchus, 70
Axiidae, 1–3, 10–13, 15, 16, 104, 116, 129, 130,
209, 210, 253, 267, 319, 507, 508, 539
Axiinae, 15
Axioidea, ix, 1–3, 9, 10, 13, 197, 337, 507, 525,
526, 549
Axiopsis, 3, 10, 16, 18, 30, 33, 41, 50–52, 67,
73, 76, 103, 125, 128, 178, 508
Axiopsis (Axiopsis) amakusana, 45
Axiopsis (Axiopsis) australiensis, 144
Axiopsis (Axiopsis) baronai, 51
Axiopsis (Axiopsis) caespitosa, 120
Axiopsis (Axiopsis) clypeata, 33
Axiopsis (Axiopsis) consobrina, 52, 177, 178
Axiopsis (Axiopsis) Habereri, 74
Axiopsis (Axiopsis) habereri, 74
Axiopsis (Axiopsis) inaequalis, 135
Axiopsis (Axiopsis) irregularis, 52, 54
Axiopsis (Axiopsis) longipes, 80
Axiopsis (Axiopsis) mediterranea, 67
Axiopsis (Axiopsis) miyazakiensis, 24
Axiopsis (Axiopsis) pailoloensis, 141
Axiopsis (Axiopsis) Picteti, 39, 40
Axiopsis (Axiopsis) picteti, 39
Axiopsis (Axiopsis) pilocheira, 20, 21, 28, 170
Axiopsis (Axiopsis) pitatucensis, 103, 104, 131
Axiopsis (Axiopsis) polyacantha, 24
Axiopsis (Axiopsis) princeps, 77
Axiopsis (Axiopsis) rudis, 81
Axiopsis (Axiopsis) serratifrons, 52, 58
Axiopsis (Axiopsis) soyoi, 97
Axiopsis (Axiopsis) sp., 58
Axiopsis (Axiopsis) spinimana, 39, 40
Axiopsis (Axiopsis) spinosissima, 29
Axiopsis (Axiopsis) spinulicauda, 126
Axiopsis (Axiopsis) tenuicornis, 98
Axiopsis (Axiopsis) werribee, 125, 127
Axiopsis (Calocarides), 86
Axiopsis (Calocarides) coronata, 89
Axiopsis (Paraxiopsis), 156
Axiopsis (Paraxiopsis) aethiopica, 43
Axiopsis (Paraxiopsis) appendiculis, 107, 109
Axiopsis (Paraxiopsis) biserrata, 113, 116
Axiopsis (Paraxiopsis) bisquamosa, 182, 183
Axiopsis (Paraxiopsis) Brocki, 158
Axiopsis (Paraxiopsis) brocki, 157, 158
Axiopsis (Paraxiopsis) defensus, 111
Axiopsis (Paraxiopsis) johnstoni, 161
Axiopsis (Paraxiopsis) sp., 255
Axiopsis (Paraxiopsis) vicina, 163
Axiopsis aethiopica, 41
Axiopsis affinis, 58
Axiopsis australiensis, 50, 144, 148
Axiopsis baronai, 51, 52, 540
Axiopsis biserratus, 113
Axiopsis bougainvillensis sp. nov., 39
Axiopsis brasiliensis, 58, 62
Axiopsis braziliensis, 58
Axiopsis Brocki, 158
Axiopsis brucei, 193
Axiopsis bythos, 30, 31
Axiopsis consobrina, 51–55, 58, 148, 177, 178,
181, 182, 540, 541
Axiopsis defensus, 111
Axiopsis eximia, 51
Axiopsis hirsutimana, 121
Axiopsis inaequalis, 135
Axiopsis irregularis, 51, 52
Axiopsis longipes, 80
Axiopsis nethertoni, 144
Axiopsis nodulosus, 261
Axiopsis oxypleura, 101
Axiopsis pailoloensis, 141
Axiopsis pica, 58, 60, 61
Axiopsis Picteti, 39, 40
Axiopsis princeps, 77
Axiopsis sculptimana, 58, 61, 62
Axiopsis serratifrons, 30, 41, 51–57, 59–62,
103, 125, 539–542
Axiopsis soyoi, 97
Axiopsis sp., 51, 58, 63, 99, 255, 542
Axiopsis spinipes, 58
Axiopsis spinulicauda, 126
Axiopsis spinulicauda amurensis, 125
Axiopsis tenuicornis, 98
Axiopsis tsushimaensis, 51, 52, 55
Axiopsis werribee, 50, 125, 127
Axiorygma, 3, 10, 16, 17, 63, 144, 508
Axiorygma nethertoni, 63, 64, 539
Axius, 3, 10, 15, 16, 18, 31, 45, 64–66, 74, 83,
104, 125, 126, 144, 150, 507
Axius (Axius) armatus, 66
Axius (Axius) novae-zealandiae, 149, 150
Axius (Axius) novaezealandiae, 150
Axius (Axius) odontorhynchus, 173
Axius (Axius) serratus, 67
Axius (Axius) stirhynchus, 70
Axius (Eiconaxius), 268
Axius (Eiconaxius) acutifrons, 270, 271, 280
Axius (Eiconaxius) Agassizi, 272
Axius (Eiconaxius) agassizi, 272
Axius (Eiconaxius) andamanensis, 272
Axius (Eiconaxius) asper, 273
Axius (Eiconaxius) Borradailei, 275
Axius (Eiconaxius) borradailei, 275
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Axius (Eiconaxius) caribbaeus, 275
Axius (Eiconaxius) caribbaeus carinatus, 276
Axius (Eiconaxius) caribbaeus rotundifrons,
285
Axius (Eiconaxius) communis, 275
Axius (Eiconaxius) consobrinus, 276, 290
Axius (Eiconaxius) crista-galli, 273, 278, 279
Axius (Eiconaxius) crista-galli antillensis, 273
Axius (Eiconaxius) farreae, 278
Axius (Eiconaxius) indica, 279
Axius (Eiconaxius) Investigatoris, 247
Axius (Eiconaxius) kermadecensis, 107
Axius (Eiconaxius) kermadeci, 281
Axius (Eiconaxius) laccadivensis, 282
Axius (Eiconaxius) parvus, 278, 284
Axius (Eiconaxius) rotundifrons, 285
Axius (Eiconaxius) Sibogae, 295
Axius (Eiconaxius) sibogue, 295
Axius (Eiconaxius) singularis, 285
Axius (Eiconaxius) spiniger, 285
Axius (Eiconaxius) Weberi, 286
Axius (Iconaxiopsis) andamanensis, 272
Axius (Iconaxiopsis) farreae, 278
Axius (Iconaxiopsis) laccadivensis, 282
Axius (Neaxius) acanthus, 328
Axius (Neaxius) euryrhynchus, 321, 323, 324
Axius (Neaxius) glyptocercus, 328
Axius (Neaxius) Gundlachi, 323
Axius (Neaxius) laevis, 90
Axius (Neaxius) mauritiana, 328
Axius (Neaxius) mauritianus, 328
Axius (Neaxius) orientalis, 323
Axius (Neaxius) plectorhynchus, 333
Axius (Neaxius) plectrorhynchus, 333
Axius (Neaxius) Vivesi, 332
Axius (Neaxius) waroona, 334
Axius (Neaxius?) euryrhynchus, 321
Axius (Neaxius?) odontorhynchus, 172, 173
Axius (Paraxius) altus, 165
Axius (Paraxius) armatus, 66
Axius (Paraxius) serratus, 67
Axius (Paraxius) spinulicauda, 126
Axius (Paraxius) tridens, 152
Axius acanthus, 328
Axius acutifrons, 270, 279
Axius affinis, 51, 58
Axius armatus, 66, 67, 539
Axius australiensis, 144
Axius Bisculcata, 70
Axius biserrata, 113
Axius biserratus, 112
Axius Brocki, 156, 158
Axius clypeatus, 33
601
Axius crista-galli, 276
Axius defensus, 111
Axius euryrhynchus, 321
Axius farreae, 278
Axius glyptocercus, 325, 326
Axius gundlachi, 323
Axius Habereri, 74
Axius habereri, 73, 74
Axius inaequalis, 135
Axius laevis, 90
Axius longipes, 80
Axius longispina, 303
Axius mauritianus, 328
Axius mediterraneus, 66–68, 539
Axius miyazakiensis, 24
Axius nodulosus, 255, 261, 265
Axius pailoloensis, 134, 141
Axius plectorhynchus, 333, 334
Axius plectrohynchus, 333
Axius plectrorhynchus, 333, 334
Axius princeps, 76, 77
Axius rudis, 81
Axius serratifrons, 56
Axius serratus, 65–67, 69, 72, 73, 539
Axius soyoi, 97
Axius sp., 31, 128
Axius spinipes, 58
Axius spinosissimus, 29
Axius spinulicauda, 126, 127
Axius spinulicauda amurensis, 125, 126
Axius Stirhynchus, 70
Axius stirhynchus, 15, 65–67, 70–72, 539
Axius stirynchus, 70
Axius werribee, 128
Balssaxius, 74
Balssaxius gen. nov., 3, 12, 16, 18, 73
Balssaxius habereri, 74, 75, 541
Balsscallichirus, 415
Balsscallichirus gen. nov., 5, 11, 339, 356, 413,
414, 466
Balsscallichirus balssi, 415, 546
Balsscallichirus guineensis, 415, 546
Barnardcallichirus, 416, 417
Barnardcallichirus gen. nov., 5, 11, 339, 356,
413, 416, 466
Barnardcallichirus gilchristi, 416, 417, 548
Barnardcallichirus tenuimanus, 416, 417, 546
Bathycalliacidae, 3, 5, 10, 11, 335, 337, 339,
340, 347
Bathycalliacinae, 5, 10, 335–339, 347, 348, 355
Bathycalliax, 5, 10, 339, 348, 349
Bathycalliax geomar, 348–350
602
CRM 013 – Katsushi Sakai
Biffarius, 335, 358, 384, 386
Biffarius arenosa, 391
Biffarius arenosus, 386
Biffarius australis, 361
Biffarius biformis, 386, 394
Biffarius ceramica, 383
Biffarius ceramicus, 383, 386
Biffarius debilis, 396
Biffarius delicatulus, 383, 386
Biffarius diaphora, 376, 386
Biffarius fragilis, 386, 397
Biffarius lewtonae, 386, 402
Biffarius limosa, 384, 386
Biffarius limosus, 384
Biffarius melissae, 403
Biffarius pacificus, 377, 378
Biffarius poorei, 386, 405
Boasaxius, 19, 77
Boasaxius gen. nov., 3, 10, 16, 76
Boasaxius princeps, 75, 77, 541
Bouvieraxius, 3, 10, 11, 16, 18, 79, 80, 209, 507
Bouvieraxius keiensis, 80, 542
Bouvieraxius longipes, 79, 80, 539
Bouvieraxius michelae, 80, 81, 541
Bouvieraxius rudis, 77, 79–81, 540, 542
Bouvieraxius springeri, 79, 80, 82, 541
Briancara foveolatus, 213
Briancaris, 4, 10, 210, 211, 213
Briancaris aberrans, 213
Briancaris foveolata, 214
Briancaris japonica, 213, 214
Bruceaxius, 83
Bruceaxius gen. nov., 3, 10, 16, 19, 20, 82
Bruceaxius polychaetes, 83, 540
Calastacus, 4, 10, 94, 210, 214, 215, 221, 224,
508
Calastacus angulatus, 133
Calastacus colpos, 215, 216, 220
Calastacus crosnieri, 2, 215–217
Calastacus euophthalmus, 134
Calastacus felix, 245, 246
Calastacus inflatus, 215, 218
Calastacus investigatoris, 247, 248
Calastacus laevis, 215, 216, 219, 220
Calastacus longispinis, 88, 91, 92
Calastacus mclaughlinae, 215, 220, 221
Calastacus mexicanus, 215, 216, 220, 221
Calastacus myalup, 215, 221
Calastacus quinqueseriatus, 94, 95
Calastacus rathbunae, 247
Calastacus rostriserratus, 96
Calastacus spinosus, 101, 102
Calastacus stilirostris, 94, 215, 216, 220, 223
Calastacus stylirostris, 223
Calaxiopsis, 4, 10, 210, 215, 221, 223, 224, 245,
249, 250, 508
Calaxiopsis felix, 245, 246
Calaxiopsis manningi, 249, 250
Calaxiopsis mclaughlinae, 221
Calaxiopsis serrata, 85, 224, 225, 227, 245,
249, 250
Calaxiopsis sp., 168, 224, 228
Calaxiopsis manningi, 250
Calaxiopsis serrata, 249
Calaxius, 3, 10, 16, 17, 84, 86, 99, 123, 124,
136, 209, 507
Calaxius acutifrons, 136
Calaxius acutirostis, 86
Calaxius acutirostris, 2, 84, 85, 99, 123, 124,
209, 540–542
Calaxius carneyi, 123, 124
Calaxius euophthalmus, 134
Calaxius galapagensis, 135
Calaxius inaequalis, 135
Calaxius jenneri, 135
Calaxius kensleyi, 136
Calaxius manningi, 2, 85, 86
Calaxius mimasensis, 136
Calaxius oxypleura, 101
Calaxius pailoloensis, 141
Calaxius sibogae, 141
Calaxius sp., 85, 86
Calaxius spinosus, 99, 101
Calaxius tungi, 85, 86
Calianassa harmandi, 401
Calianassa japonica, 401
Callanopsis anovalis, 336
Callanopsis goniophthalma, 336
Calliactites, 198, 480
Calliactites coecigena, 480, 481
Calliactites secura, 480
Callianassa, 5, 11, 335, 339, 345, 354–356, 358,
359, 361, 363–365, 387, 391, 408, 410,
418, 432, 505, 521, 531
Callianassa (Calliactites), 198
Callianassa (?Cheramus) sibogae, 408
Callianassa (?Scallasis) coeca, 308
Callianassa (Calichirus) islagtande, 419
Callianassa (Calliactites) amboinensis, 382
Callianassa (Calliactites) coeca, 308
Callianassa (Calliactites) coecigena, 481
Callianassa (Calliactites) goniophthalma, 479,
480
Callianassa (Calliactites) lignicola, 402
Callianassa (Calliactites) longicauda, 364, 368
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Callianassa (Calliactites) modesta, 369
Callianassa (Calliactites) parva, 404
Callianassa (Calliactites) rotundicaudata, 378
Callianassa (Calliactites) secura, 203
Callianassa (Callianassa) candida, 376
Callianassa (Callianassa) kraussi, 420
Callianassa (Callianassa) laurae, 432
Callianassa (Callianassa) subterranea, 360
Callianassa (Callianassa) tyrrhena, 380
Callianassa (Callichirus) acanthochirus, 431
Callianassa (Callichirus) aequimana, 498
Callianassa (Callichirus) affinis, 393
Callianassa (Callichirus) armata, 432
Callianassa (Callichirus) articulata, 433
Callianassa (Callichirus) assimilis, 433
Callianassa (Callichirus) atlantica, 378
Callianassa (Callichirus) audax, 454
Callianassa (Callichirus) balssi, 415
Callianassa (Callichirus) Bocourti, 447
Callianassa (Callichirus) borradailei, 434
Callianassa (Callichirus) Coutierei, 435
Callianassa (Callichirus) Eiseni, 442
Callianassa (Callichirus) eiseni, 442
Callianassa (Callichirus) Grandidieri, 449
Callianassa (Callichirus) guara, 457
Callianassa (Callichirus) guassutinga, 469
Callianassa (Callichirus) guineensis, 415, 427
Callianassa (Callichirus) hartmeyeri, 436
Callianassa (Callichirus) islagrande, 419
Callianassa (Callichirus) jamaicensis, 445
Callianassa (Callichirus) kewalramanii, 475
Callianassa (Callichirus) kraussi, 420
Callianassa (Callichirus) lanceolata, 435
Callianassa (Callichirus) laticauda, 380
Callianassa (Callichirus) lobata, 496
Callianassa (Callichirus) longiventris, 434, 435
Callianassa (Callichirus) madagassa, 467
Callianassa (Callichirus) major, 421
Callianassa (Callichirus) marginata, 362
Callianassa (Callichirus) Martensi, 437
Callianassa (Callichirus) martensi, 437
Callianassa (Callichirus) masoomi, 475
Callianassa (Callichirus) maxima, 459
Callianassa (Callichirus) mirim, 461
Callianassa (Callichirus) mucronata, 463
Callianassa (Callichirus) nakasonei, 435
Callianassa (Callichirus) Novae-britanniae,
498
Callianassa (Callichirus) novae-britanniae,
498
Callianassa (Callichirus) novaebritanniae, 502
Callianassa (Callichirus) oahuensis, 430, 436
Callianassa (Callichirus) Pestae, 375
603
Callianassa (Callichirus) pestae, 376
Callianassa (Callichirus) placida, 426, 436
Callianassa (Callichirus) pontica, 375
Callianassa (Callichirus) rathbunae, 464
Callianassa (Callichirus) Rosae, 444
Callianassa (Callichirus) Stebbingi, 380
Callianassa (Callichirus) stebbingi, 380
Callianassa (Callichirus) tridentate, 446
Callianassa (Callichirus) tyrrhena, 360, 380
Callianassa (Callichirus) vigilax, 464
Callianassa (Callichirus) winslowi, 465
Callianassa (Cheramus) Batei, 371
Callianassa (Cheramus) batei, 363
Callianassa (Cheramus) Calmani, 455
Callianassa (Cheramus) indica, 458
Callianassa (Cheramus) intermedia, 399
Callianassa (Cheramus) joculatrix, 401
Callianassa (Cheramus) Jousseaumei, 458
Callianassa (Cheramus) lobetobensis, 364, 368
Callianassa (Cheramus) mauritiana, 461
Callianassa (Cheramus) minima, 528
Callianassa (Cheramus) moluccensis, 462
Callianassa (Cheramus) novaeguineae, 462
Callianassa (Cheramus) orientalis, 370
Callianassa (Cheramus) pachydactyla, 472
Callianassa (Cheramus) praedatrix, 370
Callianassa (Cheramus) propinqua, 371
Callianassa (Cheramus) pugnatrix, 406
Callianassa (Cheramus) pygmaea, 407
Callianassa (Cheramus) sibogae, 408
Callianassa (Cheramus) spinophthalma, 364,
372
Callianassa (Cheramus) subterranea, 360, 380
Callianassa (Cheramus) variabilis, 451, 453,
458
Callianassa (Scallasis) Amboinae, 391
Callianassa (Scallasis) amboinae, 391
Callianassa (Scallasis) tonkinae, 411
Callianassa (Trypaea) acanthura, 390
Callianassa (Trypaea) affinis, 393
Callianassa (Trypaea) amboinensis, 384
Callianassa (Trypaea) australiensis, 392
Callianassa (Trypaea) Bouvieri, 374
Callianassa (Trypaea) bouvieri, 372, 373, 399
Callianassa (Trypaea) brachyophthalma, 382
Callianassa (Trypaea) californiensis, 395, 399
Callianassa (Trypaea) ceramica, 383
Callianassa (Trypaea) chilensis, 412
Callianassa (Trypaea) cristata, 377
Callianassa (Trypaea) eoa, 405
Callianassa (Trypaea) Filholi, 397
Callianassa (Trypaea) gigas, 398, 405
Callianassa (Trypaea) Gravieri, 377
604
CRM 013 – Katsushi Sakai
Callianassa (Trypaea) gravieri, 377
Callianassa (Trypaea) Harmandi, 399
Callianassa (Trypaea) harmandi, 399
Callianassa (Trypaea) italica, 411
Callianassa (Trypaea) japonica, 399, 405
Callianassa (Trypaea) longimana, 398
Callianassa (Trypaea) maldivensis, 374
Callianassa (Trypaea) mauritiana, 461
Callianassa (Trypaea) petalura, 405
Callianassa (Trypaea) porcellana, 392
Callianassa (Trypaea) Rochei, 407
Callianassa (Trypaea) sp., 465
Callianassa (Trypaea) truncata, 411
Callianassa (Trypaea) uncinata, 412
Callianassa (Trypaeta) truncata, 411
Callianassa ?japonica, 400
Callianassa acanthochirus, 431, 438
Callianassa acanthura, 390
Callianassa acutirostella, 390
Callianassa adamas, 419
Callianassa aequimana, 498, 499
Callianassa affinis, 393
Callianassa agassizi, 342
Callianassa algerica, 360, 375, 380
Callianassa amboinae, 365, 391, 407
Callianassa amboinensis, 381, 382, 384, 385
Callianassa amplimaxilla, 391
Callianassa anoploura, 366
Callianassa aqabaensis, 367
Callianassa arenosa, 391
Callianassa argentinensis, 343
Callianassa armata, 432
Callianassa articulata, 433
Callianassa assimilis, 433
Callianassa atlantica, 378
Callianassa audax, 454
Callianassa australiensis, 392
Callianassa australis, 361
Callianassa balssi, 414, 415, 417
Callianassa bangensis, 392
Callianassa batei, 363, 371
Callianassa berylae, 393
Callianassa biffari, 393
Callianassa biformis, 386, 393
Callianassa bisulcata, 70
Callianassa bocourti, 446, 447
Callianassa borradailei, 429
Callianassa bouvieri, 374
Callianassa brachyophthalma, 381, 382
Callianassa brachytelson, 375
Callianassa branneri, 454
Callianassa brevicaudata, 462
Callianassa brevirostris, 394
Callianassa bulimba, 499
Callianassa caesari, 394
Callianassa caledonica, 395
Callianassa californiensis, 371, 386, 395, 399–
401
Callianassa calmani, 455
Callianassa Candida, 379
Callianassa candida, 360, 375, 376, 380
Callianassa ceramica, 383
Callianassa chakratongae, 396
Callianassa chilensis, 412
Callianassa coecigena, 480, 481
Callianassa collaroy, 434
Callianassa contipes, 364, 367
Callianassa convexa, 376
Callianassa costaricensis, 396
Callianassa coutierei, 436
Callianassa cristata, 377
Callianassa Davyana, 380
Callianassa debilis, 396
Callianassa delicatula, 383
Callianassa denticulata, 511, 512
Callianassa diaphora, 376
Callianassa Eiseni, 442
Callianassa eiseni, 442
Callianassa exilimaxilla, 397
Callianassa Filholi, 397
Callianassa filholi, 397
Callianassa fragilis, 397
Callianassa garthi, 422
Callianassa gaucho, 367
Callianassa gigas, 398
Callianassa gilchristi, 417
Callianassa goniophthalma, 479
Callianassa Grandidieri, 448
Callianassa grandimana, 435, 456
Callianassa gravieri, 377
Callianassa gruneri, 398, 462
Callianassa guassutinga, 468, 469
Callianassa guineensis, 376, 377, 427
Callianassa Harmandi, 400, 401
Callianassa harmandi, 399–401
Callianassa hartmeyeri, 424, 436, 456
Callianassa haswelli, 434, 435
Callianassa helgolandica, 360
Callianassa hermandi, 399
Callianassa hertlingi, 360
Callianassa indica, 458
Callianassa intermedia, 363, 364, 398
Callianassa islagrande, 419
Callianassa italica, 411
Callianassa jamaicense, 442, 443, 445
Callianassa jamaicensis, 442, 445
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Callianassa japonica, 361, 386, 399–401, 405
Callianassa joculatrix, 401
Callianassa jousseaumei, 458
Callianassa karumba, 459
Callianassa kraussi, 420
Callianassa Laticauda, 379
Callianassa laticauda, 379, 380
Callianassa latispina, 508, 509
Callianassa lewtonae, 402
Callianassa lignicola, 402
Callianassa limosa, 384
Callianassa lobetobensis, 368
Callianassa longicauda, 368
Callianassa longimana, 398
Callianassa longiventris, 429, 435
Callianassa louisianensis, 443
Callianassa madagassa, 466, 467
Callianassa major, 418, 420
Callianassa malaccaensis, 368
Callianassa maldivensis, 374
Callianassa marchali, 377
Callianassa marginata, 362, 363
Callianassa marginatus, 364
Callianassa Martensi, 433, 437
Callianassa martensi, 437
Callianassa matzi, 403
Callianassa mauritiana, 461
Callianassa maxima, 451, 452, 459
Callianassa minima, 363, 527, 528
Callianassa minor, 361, 511, 512
Callianassa mirim, 461
Callianassa mocambiquensis, 404
Callianassa modesta, 363, 369
Callianassa mucronata, 398, 462
Callianassa nakasonei, 434, 435
Callianassa natalensis, 451, 453, 458
Callianassa ngochoae, 382, 384
Callianassa nieli, 404
Callianassa nigroculata, 369
Callianassa novae-britanniae, 500
Callianassa oblonga, 369
Callianassa occidentalis, 371, 395
Callianassa orientalis, 370
Callianassa pachydactyla, 472
Callianassa parva, 404
Callianassa parvula, 370
Callianassa pentagonocephala, 463
Callianassa persica, 378
Callianassa pestae, 360, 375
Callianassa pestai, 375
Callianassa petalura, 399, 405
Callianassa pixii, 345
Callianassa placida, 426, 430, 435
605
Callianassa plantei, 405
Callianassa pontica, 361, 375
Callianassa poorei, 405
Callianassa praedatrix, 364, 370
Callianassa profunda, 363, 365, 371
Callianassa propinqua, 363, 364, 371
Callianassa propriopedis, 406
Callianassa pugnatrix, 406
Callianassa pygmaea, 407
Callianassa quadracuta, 503
Callianassa ranongensis, 473, 474
Callianassa rathbunae, 464
Callianassa rectangularis, 364, 374
Callianassa Rochei, 407
Callianassa rochei, 407
Callianassa rosae, 444
Callianassa rotundicaudata, 378
Callianassa sahul, 407
Callianassa santarita, 408
Callianassa secura, 198, 199, 203
Callianassa seilacheri, 422
Callianassa sensu nov., 355, 358, 359
Callianassa setimana, 379
Callianassa sibogae, 363, 364, 408
Callianassa siguanensis, 456
Callianassa sp., 375, 380, 412, 453, 465
Callianassa spinoculata, 409
Callianassa spinophthalma, 372
Callianassa Stebbingi, 380
Callianassa stebbingi, 360, 380
Callianassa stenomastaxa, 409
Callianassa Stimpsoni, 378
Callianassa stimpsoni, 378
Callianassa subterranea, 356, 359–361, 365,
375, 379, 380, 387, 399, 401, 405, 408,
418, 505, 511, 521, 543, 545
Callianassa subterranea australis, 361
Callianassa subterranea intermedia, 399
Callianassa subterranea japonica, 405
Callianassa subterranea laticauda, 380
Callianassa tabogensis sp. nov., 393
Callianassa tenuipes, 409
Callianassa thailandica, 410
Callianassa thorsoni, 410
Callianassa tonkinae, 411
Callianassa tooradin, 505, 506
Callianassa tridentata, 445
Callianassa trilobata, 428, 429
Callianassa truncata, 411
Callianassa turnerana, 449
Callianassa tyrrhena, 360, 379
Callianassa uncinata, 412
Callianassa variabilis, 458
606
CRM 013 – Katsushi Sakai
Callianassa whitei, 360, 375, 380
Callianassa winslowi, 465
Callianassida, ix, 1–3, 9
Callianassidae, 1–3, 5, 10–13, 335–341, 353,
355, 357, 358, 477, 480, 521, 539
Callianassidea, 1
Callianassinae, 5, 10, 11, 335–339, 354–358,
521, 539, 543
Callianassoidea, ix, 1–3, 5, 9, 10, 197, 335, 337,
338, 340, 521, 525, 549, 595
Callianidea, 4, 10, 197–199, 206, 480, 525
Callianidea elongata, 203
Callianidea Gundlachi, 323
Callianidea gundlachi, 320
Callianidea laevicauda, 206
Callianidea laevicauda occidentalis, 207
Callianidea lepta, 315
Callianidea leura, 315
Callianidea madagascariensis, 199
Callianidea madagascariensis sp. nov., 199
Callianidea mucronata, 203
Callianidea planocula, 203, 205
Callianidea ryukyu, 200–203
Callianidea ryukyu sp. nov., 199
Callianidea sp., 199, 203
Callianidea Steenstrupi, 206
Callianidea Steenstrupii, 206
Callianidea typa, 198, 199, 202–205, 480
Callianidea typus, 203
Callianidea vandoverae, 300, 312, 316
Callianidea typa, 199, 202, 205, 206
Callianideidae, 2–4, 9, 10, 13, 197, 337, 480,
525, 526, 534
Callianideinae, 197
Callianisea, 198
Callianisea elongata, 203
Callianopsidae, 3, 5, 11, 12, 336–340, 353, 477,
483
Callianopsidae sensu nov., 480
Callianopsinae, 5, 335, 336, 338, 339, 353, 477,
478, 480, 482, 483, 485
Callianopsis, 5, 11, 335, 339, 478–481, 483,
485
Callianopsis anovalis, 482, 483
Callianopsis coecigena, 481
Callianopsis goniophthalma, 344, 478–481, 483
Callianopsis latispina, 509
Calliapaguropinae, 6, 336, 338, 339, 355, 491,
492
Calliapagurops, 6, 11, 335, 339, 491, 492
Calliapagurops charcoti, 492, 493
Calliapagurops foresti, 492, 493
Calliax, 6, 11, 335, 339, 345, 481, 493–496,
499, 500, 502, 505
Calliax aequimana, 497, 498
Calliax andamanica, 495
Calliax bulimba, 497, 499
Calliax cearaensis, 504
Calliax doerjesti, 496
Calliax jonesi, 497, 499
Calliax lobata, 496
Calliax mcilhennyi, 500
Calliax novaebritanniae, 500
Calliax punica, 497, 501
Calliax quadracuta, 503
Calliax sakaii, 502
Calliax tooradin, 506
Calliaxina, 339, 494, 497–502
Calliaxina aequimana, 497, 498
Calliaxina bulimba, 497–499
Calliaxina jonesi, 497–499
Calliaxina mcilhennyi, 497, 498, 500
Calliaxina novaebritanniae, 497, 498, 500
Calliaxina panglaoensis, 497, 498, 501
Calliaxina punica, 494, 497, 498, 501
Calliaxina sakaii, 497, 498, 502
Calliaxina sensu nov., 11, 494
Calliaxis adriatica, 116
Callichiridae, 418
Callichirinae, 5, 336, 338, 339, 357, 412, 413,
440, 491, 492, 539, 546
Callichirinae sensu nov., 11, 335, 336, 354, 355,
357
Callichirus, 5, 10, 11, 335, 339, 354, 413, 415,
417, 418, 427, 439, 474, 475, 492
Callichirus acanthochirus, 431
Callichirus adamas, 418, 419, 546, 548
Callichirus audax, 454
Callichirus balssi, 415
Callichirus calmani, 455
Callichirus foresti, 427
Callichirus garthi, 422
Callichirus guineensis, 415
Callichirus intesi, 424
Callichirus islagrande, 418, 419, 546
Callichirus jamaicensis, 442, 445
Callichirus jousseaumei, 458
Callichirus kraussi, 418, 420, 421, 548
Callichirus laurae, 432
Callichirus longiventris, 436
Callichirus major, 418, 420, 421, 492, 546
Callichirus marginatus, 362
Callichirus martensi, 437
Callichirus mirim, 461
Callichirus monodi, 472
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Callichirus pachydactyla, 472
Callichirus pentagonocephala, 463
Callichirus placidus, 436
Callichirus rathbunae, 464
Callichirus seilacheri, 418, 419, 421, 422, 546
Callichirus tenuimanus, 416, 417
Callichirus turneranus, 450
Callisea, 198
Callistocaris, 211, 213
Callocaris, 228
Callocaris macandreae, 234
Calocaridae, 13, 209
Calocarides, 4, 10, 16, 18, 73, 86–88, 91, 95,
97, 126, 127, 131, 132, 508
Calocarides armata, 66
Calocarides capensis, 87–89, 91, 539, 542
Calocarides chani, 2, 87–89, 97, 541
Calocarides coronatus, 87–90, 539
Calocarides crassipes, 87
Calocarides habereri, 74
Calocarides laevis, 87, 88, 90, 539
Calocarides lev, 87, 88, 90, 91, 540
Calocarides longispinis, 87, 88, 91, 92, 542
Calocarides macphersoni, 87, 88, 91–93, 539
Calocarides okhotskensis, 541
Calocarides okhotskensis sp. nov., 87, 88, 94
Calocarides quinqueseriatus, 87, 88, 94, 95,
540
Calocarides rostriserratus, 87, 88, 96, 540
Calocarides rudolfi, 87, 88, 96, 97, 539
Calocarides soyoi, 87, 97, 541
Calocarides spinulicauda, 126
Calocarides tenuicornis, 87, 88, 98, 541
Calocarides vigila, 98
Calocarides vigilus, 87, 88, 98, 540
Calocarides werribee, 127
Calocarididae, 3, 4, 10–13, 129, 130, 209, 210,
232
Calocaris, 4, 10, 97, 129, 130, 210, 211, 228,
229, 232, 242–244, 247, 507
Calocaris (Calastacus) amakusana, 45
Calocaris (Calastacus) euophthalma, 134
Calocaris (Calastacus) felix, 246
Calocaris (Calastacus) hirsutimana, 120, 121
Calocaris (Calastacus) Investigatoris, 247
Calocaris (Calastacus) investigatoris, 247
Calocaris (Calastacus) jenneri, 135
Calocaris (Calastacus) longispinis, 88, 91
Calocaris (Calastacus) mimasenis, 136
Calocaris (Calastacus) oxypleura, 101, 102
Calocaris (Calastacus) quinqueseriata, 95
Calocaris (Calastacus) Sibogae, 141
Calocaris (Calastacus) sibogae, 134
607
Calocaris (Calastacus) sp., 101
Calocaris (Calastacus) stilirostris, 223
Calocaris (Calocarides), 86
Calocaris (Calocarides) coronatus, 89
Calocaris (Calocarides) crassipes, 89
Calocaris (Calocaris) aberrans, 214
Calocaris (Calocaris) Alcocki, 212
Calocaris (Calocaris) alcocki, 212
Calocaris (Calocaris) Barnardi, 234
Calocaris (Calocaris) barnardi, 234
Calocaris (Calocaris) granulosus, 231, 251
Calocaris (Calocaris) Macandreae, 234
Calocaris aberrans, 213
Calocaris Alcocki, 211, 212
Calocaris alcocki, 211, 212
Calocaris barnardi, 229, 234, 236, 237, 239–
241, 361
Calocaris caribbaea, 229, 230
Calocaris caribbaeus, 229
Calocaris granulosa, 229, 231
Calocaris granulosus, 231
Calocaris investigatoris, 247
Calocaris isochela, 229, 232
Calocaris lev, 90
Calocaris longispinis, 88, 91
Calocaris Macandrae, 233, 234
Calocaris Macandreae, 233
Calocaris macandreae, 129, 211, 229, 231,
233–243, 362
Calocaris Macandrei, 233
Calocaris Mc Andreae, 234
Calocaris mimasensis, 136
Calocaris rudolfi, 96
Calocaris sp., 229, 234, 241, 242
Calocaris spinosus, 102
Calocaris templemani, 242–244
Calocarisopsis, 242, 243
Calocarisopsis gen. nov., 4, 11, 209, 210, 242,
244
Calocarisopsis templemani, 234, 242–244
Cancer (Astacus) subterraneus, 359, 360
Cancer candidus, 359, 375
Cancer modestus, 110, 112, 113
Capecalliax, 341, 345
Capecalliax gen. nov., 5, 11, 337, 341, 342, 345
Capecalliax pixii, 345
Calliapagurops foresti, 493
Cheraminae, 335, 353, 354, 357, 358, 363, 364
Cheramoides, 362, 363
Cheramoides gen. nov., 5, 11, 339, 355, 356,
358, 359, 362, 364
Cheramoides marginata, 362, 543
608
CRM 013 – Katsushi Sakai
Cheramus, 5, 11, 335, 339, 354–358, 362–365,
372
Cheramus anoplourus, 365, 366, 544
Cheramus aqabaensis, 365–367, 545
Cheramus batei, 371
Cheramus contipes, 365–367, 544
Cheramus gaucho, 344, 365–367, 543
Cheramus lobetobensis, 365, 366, 368, 544
Cheramus longicauda, 365, 368, 544
Cheramus longicaudatus, 368
Cheramus malaccaensis, 364–366, 368, 544
Cheramus marginatus, 362, 364
Cheramus minima, 364
Cheramus modestus, 365, 369, 543, 544
Cheramus nigroculata, 364
Cheramus nigroculatus, 365, 366, 369, 544
Cheramus oblongus, 365, 366, 369, 543
Cheramus occidentalis, 363, 365, 371
Cheramus orientalis, 365, 366, 370, 545
Cheramus parvulus, 365, 370, 545
Cheramus praedatrix, 365, 366, 370, 544
Cheramus profundus, 362, 364–366, 371, 543
Cheramus propinquus, 365, 371, 372, 544, 545
Cheramus rectangularis, 374
Cheramus sensu nov., 355, 358, 359
Cheramus sibogae, 409
Cheramus sp., 365, 372
Cheramus spinophthalmus, 365, 366, 372, 544
Cheramus subterraneus, 360
Coelhocalaxius, 99, 102
Coelhocalaxius gen. nov., 4, 16, 17, 98, 99
Coelhocalaxius spinosus, 99, 100, 539
Colemanaxius, 103, 104, 131
Colemanaxius gen. nov., 4, 12, 16, 17, 103
Colemanaxius pitatucensis, 103–105, 541
Coralaxiidae, 3, 4, 10, 13, 253, 254
Coralaxiinae, 253
Coralaxius, 4, 10, 253–255, 258, 261
Coralaxius abelei, 255–258, 261, 265
Coralaxius galapagensis, 255, 258
Coralaxius indopacificus, 255, 258, 259
Coralaxius nodulosus, 261, 262
Corallianassa, 335, 423, 429, 430
Corallianassa articulata, 433
Corallianassa borradailei, 423, 434
Corallianassa collaroy, 434
Corallianassa hartmeyeri, 423, 430, 436
Corallianassa intesi, 424
Corallianassa longiventris, 423, 430, 436
Corallianassa xutha, 423, 426, 430
Corallichirus, 339, 422, 423, 430
Corallichirus bayeri, 423, 424, 433, 547
Corallichirus intesi, 423–425, 546
Corallichirus placidus, 436
Corallichirus sensu nov., 11, 413, 414, 423, 430
Corallichirus tridentatus, 446
Corallichirus xuthus, 423–426, 546
Crosniera, 6, 9, 11, 197, 340, 525–527
Crosniera corindon, 526, 527
Crosniera dayrati, 526–529
Crosniera minima, 364, 526–528
Crosniera panie, 526, 527, 530
Crosniera sp., 526, 527, 530
Ctenocheles, 6, 11, 335, 339, 485, 486, 489
Ctenocheles Balssi, 487
Ctenocheles balssi, 486, 487
Ctenocheles collini, 486, 487
Ctenocheles holthuisi, 486, 487
Ctenocheles leviceps, 486–488
Ctenocheles maorianus, 486, 488
Ctenocheles serrifrons, 486, 488
Ctenocheles sp., 486, 488, 489
Ctenochelidae, 1–3, 6, 11, 335–340, 353, 479,
480, 483, 485, 507, 515, 595, 596
Ctenochelinae, 335, 353, 485
Ctenocheloidae, 595, 596
Ctenocheloidae fam. nov., 595
Ctenocheloides, 595, 596
Ctenocheloides attenboroughi, 595, 596
Dawsoniinae, 507
Dawsonius, 6, 11, 335, 339, 485, 508, 510
Dawsonius latispina, 508–510
Dorphinaxius, 4, 10, 16, 17, 106, 107, 109, 508
Dorphinaxius appendiculis, 107
Dorphinaxius kermadecensis, 107–109, 540
Econaxius antillensis, 273
Econaxius sibogae, 275
Eiconaxiidae, 3, 4, 10, 13, 14, 267, 289
Eiconaxiopsididae, 13, 290
Eiconaxiopsididae fam. nov., 3, 5, 11, 14, 289
Eiconaxiopsis, 275, 290, 291
Eiconaxiopsis gen. nov., 5, 11, 267, 289, 290
Eiconaxiopsis heinrichi, 290
Eiconaxiopsis heinrichi sp. nov., 276, 291–295
Eiconaxiopsis sibogae, 281, 291, 295
Eiconaxius, 4, 10, 30, 267–269, 289–291, 507
Eiconaxius acanthus, 326, 328, 331
Eiconaxius acutifrons, 268, 270, 271, 279–281
Eiconaxius Agassizi, 272
Eiconaxius agassizi, 268, 270, 272
Eiconaxius albatrossae, 268, 270, 272–274
Eiconaxius andamanensis, 268, 269, 272
Eiconaxius antillensis, 268–270, 273
Eiconaxius asper, 269, 270, 273, 278
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Eiconaxius baja, 269, 270, 273, 274
Eiconaxius bandaensis sp. nov., 269, 274, 275
Eiconaxius Borradailei, 275
Eiconaxius borradailei, 269, 275
Eiconaxius caribbaeus, 269, 275
Eiconaxius carinatus, 269, 276
Eiconaxius communis, 275
Eiconaxius consobrinus, 268–270, 276, 277,
290, 291
Eiconaxius crista-galli, 273, 278
Eiconaxius cristagalli, 269, 270, 276, 278
Eiconaxius demani, 269, 278, 284
Eiconaxius farreae, 269, 270, 278
Eiconaxius faxoni, 269, 279
Eiconaxius faxoni sp. nov., 269–271, 279
Eiconaxius glyptocercus, 331
Eiconaxius hakuhou, 269, 270, 279
Eiconaxius indica, 279
Eiconaxius indicus, 2, 269, 279, 280
Eiconaxius indonesicus, 281
Eiconaxius indonesicus sp. nov., 269, 271, 280,
281
Eiconaxius kermadecensis, 107
Eiconaxius kermadeci, 268–270, 281, 282
Eiconaxius kimbla, 269, 270, 281, 282
Eiconaxius laccadivensis, 268–270, 272, 282
Eiconaxius laevis, 90
Eiconaxius mallacoota, 269, 270, 283
Eiconaxius mortenseni, 269, 284
Eiconaxius parvus, 269, 278, 284
Eiconaxius rotundifrons, 269, 285
Eiconaxius sibogae, 274, 275, 295, 296
Eiconaxius singularis, 269, 270, 285
Eiconaxius spiniger, 269, 285
Eiconaxius spinigerus, 269
Eiconaxius taliliensis, 328, 331
Eiconaxius Vivesi, 332
Eiconaxius Weberi, 288
Eiconaxius weberi, 269, 270, 286, 287
Eucalastacus, 4, 10, 210, 244
Eucalastacus torbeni, 244
Eucalliacidae, 3, 6, 11, 336–340, 491, 506
Eucalliacinae, 6, 335, 336, 338, 339, 354, 355,
491, 493, 494
Eucalliax, 6, 11, 335, 339, 354, 494, 501–504
Eucalliax cearaensis, 504
Eucalliax cearensis, 504
Eucalliax jonesi, 499
Eucalliax mcilhennyi, 497, 500
Eucalliax panglaoensis, 497, 501
Eucalliax quadracuta, 503
Eucalliaxiopsis, 503, 504
Eucalliaxiopsis gen. nov., 6, 11, 339, 494, 503
609
Eucalliaxiopsis cearaensis, 504
Eucalliaxiopsis quadracuta, 504
Eucalliinae, 491, 493
Euconaxius, 86
Euconaxius coronatus, 87, 89
Euconaxius crassipes, 89
Eutrichocheles, 4, 11, 16, 19, 109–111, 113,
114, 116, 142, 155, 156, 162, 182, 508
Eutrichocheles austrinus, 110, 111, 542
Eutrichocheles biserrata, 114–116, 118
Eutrichocheles bisquamosa, 182, 183
Eutrichocheles brocki, 158
Eutrichocheles crosnieri, 111, 113, 114, 117,
118
Eutrichocheles defensus, 110, 111, 539
Eutrichocheles foveolatus, 110–112, 539
Eutrichocheles granulimanus, 110–112, 539
Eutrichocheles johnstoni, 161
Eutrichocheles modestus, 84, 110–119, 155,
182, 209, 540, 542
Eutrichocheles pindatyba, 162
Eutrichocheles pumilus, 155, 156
Eutrichocheles sp., 142
Eutrichocheles spinipleurus, 111, 119, 539
Eutrichocheles tuamotu, 111, 119, 540
Eutricholeles, 183
Evaxius, 185
Evaxius tricarinatus, 190, 192
Farrea occa, 278
Forestcallichirus, 427
Forestcallichirus gen. nov., 5, 11, 339, 356, 413,
426, 427, 466
Forestcallichirus foresti, 427, 472, 546
Garyia, 531, 536
Garyia gen. nov., 6, 10, 340, 525, 526, 530, 531,
534
Garyia arno, 526, 531
Garyia moorea, 526, 531, 532
Gebia Daviana, 380
Gebios, 358, 359
Gebios davianii, 379
Gebios davianus, 359
Gebios Davyana, 380
Gebios Davyanus, 379
Gebius, 358, 359
Gilvossius, 5, 10, 339, 355, 356, 358, 372, 373,
376–378, 386
Gilvossius bouvieri, 373, 374, 543–545
Gilvossius brachytelson, 373, 375, 544
Gilvossius candidus, 361, 373, 375, 543
Gilvossius convexus, 373, 374, 376, 543
610
CRM 013 – Katsushi Sakai
Gilvossius diaphorus, 373, 376, 543
Gilvossius gravieri, 373, 374, 377, 545
Gilvossius marchali, 373, 377, 543
Gilvossius pacificus, 374, 377, 543
Gilvossius persicus, 373, 378, 545
Gilvossius rotundicaudatus, 373, 378, 545
Gilvossius sensu nov., 355, 358
Gilvossius setimanus, 373, 374, 378, 543
Gilvossius sp., 373, 381
Gilvossius tyrrhena, 361
Gilvossius tyrrhenus, 360, 361, 373, 374, 379,
543
Gilvossius whitei, 373, 374, 380, 543
Glypturoides, 428
Glypturoides gen. nov., 5, 11, 339, 413, 414,
428, 468
Glypturoides trilobata, 425, 428, 429
Glypturoides trilobatus, 546
Glypturus, 5, 11, 335, 339, 413, 414, 423, 426,
428–430, 432, 437, 446
Glypturus acanthochirus, 66, 423, 425, 426,
430–432, 436, 438, 546
Glypturus accanthochirus, 432
Glypturus armata, 66, 431, 432
Glypturus armatus, 430, 432, 547, 548
Glypturus articulatus, 430, 433, 547
Glypturus assimilis, 430, 433, 547
Glypturus borradailei, 430, 431, 434, 548
Glypturus branneri, 454
Glypturus collaroy, 430, 434, 547
Glypturus coutierei, 436, 437
Glypturus grandimana, 456
Glypturus grandimanus, 435, 436, 456
Glypturus hartmeyeri, 423, 430, 436, 437
Glypturus haswelli, 430, 434, 547
Glypturus intesi, 424
Glypturus karumba, 459, 460
Glypturus lanceolatus, 430, 435, 547
Glypturus laurae, 432
Glypturus longiventris, 423, 425, 430, 431,
435–437, 546–548
Glypturus martensi, 430, 435, 437, 547, 548
Glypturus motupore, 430, 432
Glypturus mucronatus, 463
Glypturus nakasonei, 430
Glypturus novaeguineae, 463
Glypturus oahuensis, 437
Glypturus rabalaisae, 431, 438, 546
Glypturus rabalaisae sp. nov., 430, 431, 438
Glypturus siguanensis, 451, 453, 456
Glypturus sp., 451
Glypturus winslowi, 465
Glypturus xuthus, 423, 426, 430
Gonodactylus setimanus, 373, 378
Gourretia, 6, 11, 335, 339, 353, 485, 508, 510,
511, 516
Gourretia aungtonyai, 517
Gourretia barracuda, 511
Gourretia biffari, 517
Gourretia coolibah, 518
Gourretia crosnieri, 518
Gourretia denticulata, 361, 509, 511, 512
Gourretia lahouensis, 507, 518
Gourretia laresi, 511, 512
Gourretia latispina, 509
Gourretia loeuffintesi, 511–513
Gourretia loeuffintesi sp. nov., 513
Gourretia manihinae, 511, 513
Gourretia minor, 512, 513
Gourretia nosybeensis, 511, 513
Gourretia phuketensis, 516, 519
Gourretia serrata, 512
Gourretia sp., 511–514
Gourretiidae, 2, 3, 6, 11, 12, 336–340, 353, 507,
508, 515
Gourretiinae, 353, 507
Grynaminna, 5, 11, 339, 356, 413, 414, 438–
440, 466
Grynaminna tamakii, 356, 439, 440, 547
Guyanacaris, 120, 121
Guyanacaris gen. nov., 4, 10, 16, 19, 119, 120
Guyanacaris caespitosa, 120, 540
Guyanacaris hirsutimana, 120–122, 539
Heardaxius, 533
Heardaxius gen. nov., 6, 11, 340, 525, 526, 533,
534
Heardaxius salvati, 526, 533
Heterocalaxius, 124
Heterocalaxius gen. nov., 4, 10, 16, 17, 123
Heterocalaxius carneyi, 124, 539
Homarus scaber, 112
Iconaxiopsis, 268
Iconaxiopsis andamanensis, 272
Iconaxiopsis consobrina, 276, 290, 291, 294
Iconaxiopsis consobrinus, 291
Iconaxiopsis farreae, 278
Iconaxiopsis kermadecensis, 107, 109
Iconaxiopsis laccadivensis, 282
Iconaxiopsis spiniger, 285
Iconaxius, 268
Iconaxius (Iconaxiopsis?) consobrinus, 276,
290, 291
Iconaxius acutifrons, 270
Iconaxius Agassizi, 272
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Iconaxius asper, 273
Iconaxius Borradailei, 275
Iconaxius caribbaeus, 275
Iconaxius communis, 275
Iconaxius consobrinus, 294
Iconaxius crista-galli, 279
Iconaxius cristagalli, 273, 278, 279
Iconaxius indica, 279
Iconaxius kermadeci, 281
Iconaxius parvus, 284
Iconaxius rotundifrons, 285
Isea, 198
Isea elongata, 203
Jaxea nocturna, 116
Laomediidae, 1, 2, 9, 116, 337
Laurentgourretia, 6, 12, 339, 508, 514
Laurentgourretia rhopalommata, 514
Leonardsaxius, 50, 125–128
Leonardsaxius gen. nov., 4, 10, 16, 18, 50, 51,
124
Leonardsaxius amurensis, 125, 126, 541
Leonardsaxius sp., 125, 128, 542
Leonardsaxius spinulicauda, 125–127, 540
Leonardsaxius werribee, 50, 125, 127, 128, 540,
542
Lepidophthalmoides, 10, 441, 444
Lepidophthalmoides gen. nov., 5, 11, 339, 413,
414, 440, 441
Lepidophthalmoides bocourti, 447
Lepidophthalmoides eiseni, 441, 442, 444, 447,
546
Lepidophthalmoides jamaicense, 441, 442, 445,
546
Lepidophthalmoides louisianensis, 441–443,
445, 546
Lepidophthalmoides richardi, 441–443, 546
Lepidophthalmoides rosae, 441, 442, 444, 547,
548
Lepidophthalmoides siriboia, 441, 442, 445,
546
Lepidophthalmoides socotrensis, 441, 445, 548
Lepidophthalmoides tridentata, 441, 442, 445
Lepidophthalmoides tridentatus, 547
Lepidophthalmus, 5, 11, 335, 339, 413, 414,
441, 446, 447
Lepidophthalmus bocourti, 441, 446–448, 546
Lepidophthalmus Eiseni, 442
Lepidophthalmus eiseni, 441, 442
Lepidophthalmus grandidieri, 446–449, 548
Lepidophthalmus jamaicense, 443
Lepidophthalmus louisianensis, 443
611
Lepidophthalmus manningi, 444
Lepidophthalmus rafai, 446, 447, 449, 546
Lepidophthalmus ranongensis, 474
Lepidophthalmus richardi, 443, 444
Lepidophthalmus rosae, 444
Lepidophthalmus sinuensis, 446–449, 546
Lepidophthalmus siriboia, 445
Lepidophthalmus siriboius, 445
Lepidophthalmus socotrensis, 445
Lepidophthalmus sp., 444
Lepidophthalmus tridentatus, 446
Lepidophthalmus turneranus, 446, 447, 449,
450, 546
Levantocaris, 4, 10, 16, 18, 129, 130, 171, 210
Levantocaris hornungae, 115, 129, 130, 539
Lipkecallianassa, 6, 12, 340, 521, 522
Lipkecallianassa abyssa, 521, 522
Lipkecallianassidae, 3, 6, 12, 336–338, 340, 521
Lipkecallianassinae, 355, 521
Litoraxius, 4, 10, 16, 19, 130
Litoraxius boshu, 130, 541
Lophaxiopsis, 245, 246
Lophaxiopsis gen. nov., 4, 12, 210, 245
Lophaxiopsis felix, 246
Lophaxius, 4, 11, 210, 245–248
Lophaxius investigaroris, 248
Lophaxius investigatoris, 247, 248
Lophaxius rathbunae, 245, 247, 248
Lophaxius rudis, 81
Manaxius, 4, 10, 16, 17, 84, 103, 104, 131–133,
136, 140, 142
Manaxius aganaensis, 131, 133, 540
Manaxius aganaensis nom. nov., 104
Manaxius aganaensis sp. nov., 132–134
Manaxius angulatus, 132, 133, 539
Manaxius euophthalmus, 122, 132, 134, 136,
540, 541
Manaxius galapagensis, 132, 135, 540
Manaxius inaequalis, 132, 135, 539
Manaxius jenneri, 132, 135, 540
Manaxius kensleyi, 132, 133, 136, 539
Manaxius mimasensis, 132, 136, 541
Manaxius ohsumiensis, 541
Manaxius ohsumiensis sp. nov., 132, 133, 137–
139
Manaxius pailoloensis, 132, 133, 141, 540
Manaxius paulayi, 133
Manaxius pitatucensis, 103, 104, 131, 133, 134
Manaxius sibogae, 132, 134, 141, 541
Manaxius thailandensis, 541
Manaxius thailandensis sp. nov., 132, 142
612
CRM 013 – Katsushi Sakai
Marcusiaxius, 5, 11, 197, 198, 297–299, 301,
302, 311, 525
Marcusiaxius colpos, 299, 301, 302
Marcusiaxius lemoscastroi, 299, 301, 302
Marcusiaxius longispina, 301–303
Marcusiaxius microps, 302, 304
Marcusiaxius minutus, 302, 304
Marcusiaxius sp., 302, 305
Marcusiaxius torbeni, 302, 304
Marcusiaxius wamsoi, 302, 305
Marianaxius, 4, 11, 16, 18, 142, 143
Marianaxius kroppi, 143, 540
Meinertaxius, 261, 265
Meinertaxius gen. nov., 4, 10, 254, 255, 260
Meinertaxius nodulosus, 261, 262, 264, 265
Mesostylus, 358, 359
Metaxius, 305
Metaxius microps, 304
Meticonaxiidae, 3, 5, 11, 13, 198, 297, 298
Meticonaxiinae, 297
Meticonaxius, 5, 11, 197, 198, 297, 298, 300,
301, 305, 306, 311, 525
Meticonaxius ? longispina, 303
Meticonaxius bouvieri, 300, 306
Meticonaxius capricorni, 306, 307
Meticonaxius dentatus, 303, 306, 307
Meticonaxius lemoscastroi, 302
Meticonaxius longispina, 300, 301, 303
Meticonaxius microps, 301, 304
Meticonaxius minutus, 304
Meticonaxius monodon, 298, 300, 306, 307
Meticonaxius noumea, 306, 308
Meticonaxius soelae, 306, 308
Meticonaxius sp., 306, 307, 309
Meticonaxius spicatus, 306, 308
Michaelcallianassa, 5, 11, 339, 413, 450, 593
Michaelcallianassa indica, 450, 547, 548
Michaelcallianassa sinica, 593
Michelaxiopsis, 4, 10, 16, 18, 50, 51, 143, 144
Michelaxiopsis australiensis, 50, 144–148, 540,
542
Michelaxiopsis nauo, 146–148
Michelcallianassa, 593
Michelcallianassa indica, 593
Michelcallianassa sinica, 593
Michelea, 5, 11, 197, 198, 297, 298, 300, 311,
312, 525
Michelea abranchiata, 303, 312, 313
Michelea dampieri, 312, 313
Michelea devaneyi, 312–314
Michelea hortus, 312, 315
Michelea lamellosa, 300, 312, 315
Michelea lepta, 312, 315
Michelea leura, 312, 313, 315
Michelea microphylla, 300, 312, 313, 316
Michelea novaecaledoniae, 312, 316
Michelea paraleura, 312, 313, 316
Michelea pillsburyi, 298, 300, 312, 313, 316
Michelea sp., 312, 317
Michelea vandoverae, 298, 300, 312, 316
Micheleidae, 3, 5, 11, 13, 14, 197, 198, 297,
298, 300, 311, 525
Micheleinae, 297, 311
Mictaxius, 6, 10, 197, 340, 525, 526, 533–535,
537
Mictaxius arno, 531, 534
Mictaxius salvati, 533, 534
Mictaxius sp., 534, 537
Mictaxius thalassicola, 526, 533–535, 537
Michaelcallianassa indica, 593
Neaxiopsis, 5, 10, 319, 320
Neaxiopsis euryrhyncha, 320–322, 324
Neaxiopsis euryrhynchus, 321, 323, 324
Neaxiopsis gundlachi, 320–324
Neaxiopsis orientalis, 321, 323, 324
Neaxius, 5, 10, 319, 320, 324, 325, 507
Neaxius acanthus, 325, 327–330
Neaxius frankeae, 325, 332
Neaxius glyptocercus, 325–331
Neaxius mauritiana, 330
Neaxius mauritianus, 325
Neaxius mclaughlinae, 325, 331
Neaxius ngochoae, 381
Neaxius taliliensis, 325
Neaxius trondlei, 325, 328, 330
Neaxius vivesi, 325, 332
Necallianassa, 358, 386
Necallianassa berylae, 386
Necallianassa truncata, 411
Neocallianassa acanthura, 390
Neocallianassa berylae, 393
Neocallianopsinae, 336, 338, 482
Neocallianopsinae subfam. nov., 5, 336, 339,
477, 482, 483
Neocallianopsis, 482, 483
Neocallianopsis gen. nov., 5, 11, 336, 339, 482
Neocallianopsis anovalis, 482, 483
Neocallichirus, 5, 11, 335, 339, 413, 414, 428,
446, 451, 452, 465, 468, 473, 474
Neocallichirus angelikae, 451–453, 547
Neocallichirus auchenorhynchus, 451–453
Neocallichirus audax, 451, 452, 454, 548
Neocallichirus branneri, 451, 453, 454, 546
Neocallichirus cacahuate, 469
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Neocallichirus caechabitator, 451, 452, 454,
547
Neocallichirus calmani, 451, 453, 455, 548
Neocallichirus collaroy, 434
Neocallichirus darwinensis, 451, 453, 455, 547
Neocallichirus denticulatus, 451, 452, 455, 547
Neocallichirus grandimana, 454, 456
Neocallichirus grandimanus, 448, 451, 453,
456, 546
Neocallichirus guaiqueri, 451, 452, 457, 546
Neocallichirus guarua, 457
Neocallichirus guarus, 451, 452, 457, 546
Neocallichirus guassutinga, 470
Neocallichirus guassutingus, 469
Neocallichirus horneri, 428, 451, 453, 458, 468,
473, 474, 547
Neocallichirus indicus, 451, 453, 458, 460, 547,
548
Neocallichirus jousseaumei, 451, 453, 458, 459,
548
Neocallichirus karumba, 451, 452, 459, 460,
547
Neocallichirus kempi, 451, 452, 459, 460, 547,
548
Neocallichirus lemaitrei, 471
Neocallichirus limnosa, 384
Neocallichirus limosus, 384
Neocallichirus manningi, 452, 453, 460, 548
Neocallichirus maryae, 464
Neocallichirus mauritianus, 452, 461, 548
Neocallichirus mirim, 452, 461, 546
Neocallichirus moluccensis, 452, 453, 462, 547
Neocallichirus monodi, 472
Neocallichirus mortenseni, 452, 462, 546
Neocallichirus motupore, 432
Neocallichirus mucronatus, 452, 462, 463, 547,
548
Neocallichirus natalensis, 458
Neocallichirus nickellae, 452, 453, 463, 546
Neocallichirus pachydactylus, 472
Neocallichirus pentagonocephala, 463
Neocallichirus pentagonocephalus, 452, 463,
546
Neocallichirus ranongensis, 473, 474
Neocallichirus rathbunae, 452, 464, 546
Neocallichirus raymanningi, 464
Neocallichirus sassandrensis, 452, 464, 546
Neocallichirus sp., 452, 457, 465
Neocallichirus sulfureus, 473
Neocallichirus taiaro, 451, 453, 458
Neocallichirus trilobata, 429
Neocallichirus trilobatus, 428, 429
Neocallichirus vigilax, 452, 464, 465, 547
613
Neocallichirus winslowi, 452, 465, 547
Neotrypaea, 335, 358, 386
Neotrypaea affinis, 393
Neotrypaea biffari, 393
Neotrypaea californiensis, 395
Neotrypaea gigas, 398
Neotrypaea rochei, 407
Neotrypaea uncinata, 412
Newzealandaxius, 149, 150
Newzealandaxius gen. nov., 4, 10, 16, 148
Newzealandaxius novaezealandiae, 149, 150,
540
Ngochoaxius gen. nov., 525
Nihonotrypaea, 358, 386, 410
Nihonotrypaea harmandi, 400, 401
Nihonotrypaea japonica, 399, 401
Nihonotrypaea petalura, 405
Nihonotrypaea thermophila, 410
Nipponcalaxiopsis, 249, 250
Nipponcalaxiopsis gen. nov., 4, 10, 210, 248
Nipponcalaxiopsis manningi, 227, 249
Notiax, 5, 11, 335, 339, 355, 356, 358, 381, 382,
385, 386
Notiax amboinensis, 381, 382, 384, 544
Notiax brachyophthalma, 344, 381–383, 543
Notiax ceramica, 381–383, 543, 545
Notiax delicatula, 381–383
Notiax delicatulus, 543
Notiax limosa, 381, 384, 543
Notiax ngochoae, 382, 384, 385, 543, 545
Notiax santarita, 408
Notiax sensu nov., 355, 358, 359
Oxyrhynchaxius, 4, 10, 16, 19, 150, 151, 508
Oxyrhynchaxius japonicus, 150, 151, 541
Oxyrhynchaxius manningi, 150, 151, 542
Oxyrhynchaxius tricarinatus, 150, 151, 540
Pagurus faujasi, 359
Paracalliacinae, 355
Paracallianidea, 4, 9, 10, 198, 205, 206, 337,
526
Paracallianidea laevicauda, 204, 206, 207
Paracalliax, 6, 12, 335, 340, 485, 508, 515
Paracalliax bollorei, 515
Paracalocaris, 4, 10, 210, 250
Paracalocaris granulosa, 231
Paracalocaris sagamiensis, 2, 231, 250, 251
Paraglypturus, 6, 11, 339, 494, 502, 504, 505
Paraglypturus calderus, 505
Paraglypturus novaebritanniae, 500
Paraglypturus punica, 501
Paraglypturus sakaii, 502
614
CRM 013 – Katsushi Sakai
Paraglypturus tooradin, 506
Paragourretia, 6, 11, 340, 508, 516, 517
Paragourretia aungtonyai, 516, 517
Paragourretia biffari, 516, 517
Paragourretia coolibah, 516–518
Paragourretia crosnieri, 516, 518
Paragourretia lahouensis, 516–518
Paragourretia phuketensis, 514, 516, 517, 519
Parascytoleptus, 4, 11, 16, 17, 152, 507
Parascytoleptus tridens, 152–154, 540, 542
Paratrichocheles, 156
Paratrichocheles gen. nov., 4, 10, 16, 17, 79,
155, 156
Paratrichocheles pumilus, 156, 542
Paratrypaea, 10, 358, 372, 373
Paratrypaea bouvieri, 374
Paraxiopsis, 4, 11, 16, 18, 110, 119, 143, 156,
157, 162, 209
Paraxiopsis austrinus, 111
Paraxiopsis bisquamosa, 184
Paraxiopsis brocki, 110, 153, 157–159, 162,
164, 210, 540–542
Paraxiopsis crosnieri, 113
Paraxiopsis defensus, 111
Paraxiopsis dianae, 156
Paraxiopsis foveolata, 112
Paraxiopsis gracilimana, 157, 158, 160, 539
Paraxiopsis granulimana, 112
Paraxiopsis hispida, 157, 161, 539
Paraxiopsis hispidus, 158, 210
Paraxiopsis johnstoni, 157, 161, 540
Paraxiopsis majuro, 157, 161, 540
Paraxiopsis paulayi, 157, 161, 162, 210, 540
Paraxiopsis pindatyba, 157, 158, 162, 539
Paraxiopsis plumosimanus, 157, 158, 162, 540
Paraxiopsis pumilus, 156
Paraxiopsis sp., 157, 164
Paraxiopsis spinipleura, 119
Paraxiopsis tuamotu, 119
Paraxiopsis vicina, 157, 163, 164, 539
Paraxius, 4, 11, 16, 18, 164, 165, 171, 508
Paraxius altus, 165–167, 540
Paraxius Picteti, 39
Paraxius tridens, 152, 154
Pentacheles, 486
Pentacheles sp. nov., 487
Pestarella, 10, 358, 372, 373
Pestarella candida, 376
Pestarella tyrrhena, 380
Pestarella whitei, 380
Pilbaraxius, 4, 10, 16, 167, 168
Pilbaraxius kariyarra, 168, 542
Pillsburyaxius, 170
Pillsburyaxius gen. nov., 4, 12, 16, 18, 20, 169
Pillsburyaxius kirkmilleri, 170, 539
Planaxius, 4, 11, 16, 18, 171
Planaxius brevifrons, 171, 540–542
Platyaxiopsis, 173
Platyaxiopsis gen. nov., 4, 10, 16, 172
Platyaxiopsis odontorhyncha, 172, 173, 541
Platyaxiopsis odontorhynchus, 173
Platyaxius, 4, 10, 16, 17, 173, 174
Platyaxius bardi, 174, 175, 542
Platyaxius brevirostris, 173, 174, 176, 542
Pleurocalliax, 481
Pleurocalliax gen. nov., 5, 12, 339, 478, 480,
481
Pleurocalliax coecigena, 481
Podocallichirus, 5, 11, 339, 356, 413, 414, 416,
427, 439, 466, 467, 474, 475
Podocallichirus balssi, 356, 415, 466
Podocallichirus foresti, 356, 427, 466
Podocallichirus gilchristi, 356, 417, 466
Podocallichirus guineensis, 356, 415, 416, 466
Podocallichirus madagassa, 439
Podocallichirus madagassus, 356, 440, 466,
467, 474, 548
Podocallichirus masoomi, 356, 357, 466, 474,
475
Podocallichirus tamakii, 356, 439, 440, 466
Podocallichirus tenuimanus, 356, 417, 466
Pontonia pinnophylax, 380
Posthonocaris, 79
Poti, 358, 363, 364
Poti gaucho, 344, 363, 364, 367
Pseudaxiopsis, 178
Pseudaxiopsis carpentariensis, 178
Pseudaxiopsis carpentariensis gen. et sp. nov.,
178
Pseudaxiopsis carpentariensis sp. nov., 182
Pseudoaxiopsis, 177, 182
Pseudoaxiopsis gen. nov., 4, 10, 16, 17, 177
Pseudoaxiopsis carpentariaensis, 54
Pseudoaxiopsis carpentariaensis sp. nov., 52,
179, 180
Pseudoaxiopsis carpentariensis, 541
Pseudoaxiopsis carpentariensis sp. nov., 177,
178, 182
Pseudobiffarius, 358, 386
Pseudobiffarius caesari, 386, 394
Pseudocalliax, 506
Pseudocalliax gen. nov., 6, 10, 339, 494, 505,
506
Pseudocalliax tooradin, 506
Pseudogourretia, 6, 12, 340
Pseudogourretiidae, 3, 6, 336–338, 340
AXIOIDEA OF THE WORLD AND A RECONSIDERATION OF CALLIANASSOIDEA
Pseudogourretiinae, 507, 508
Ralumcaris, 107, 183
Ralumcaris gen. nov., 4, 10, 16, 17, 107, 182
Ralumcaris bisquamosa, 183, 184, 541
Rayllianassa, 11, 359, 381, 384, 385
Rayllianassa amboinensis, 384
S. stirhynchus, 72
Sakaiocaris, 193
Salmoneus caboverdensis, 472
Salmoneus erasimorum, 380
Scallasis, 335, 358, 363, 364
Scallasis amboinae, 364, 365, 391
Scytoleptus, 4, 11, 16, 17, 152, 185, 186, 507
Scytoleptus barbatus, 189, 192, 541, 542
Scytoleptus barbatus sp. nov., 186–189
Scytoleptus serripes, 186, 189–192, 542
Scytoleptus tricarinatus, 190
Sergio, 5, 10, 339, 413, 467–469, 471
Sergio cacahuate, 468, 469, 546
Sergio guaiqueri, 457
Sergio guara, 457
Sergio guarus, 468
Sergio guassutinga, 448, 468–471, 546
Sergio lemaitrei, 468, 471, 546
Sergio mericeae, 468, 469, 471, 546
Sergio mirim, 461, 468
Sergio monodi, 468, 472, 546
Sergio pachydactylus, 468, 472, 546
Sergio sensu nov., 11, 413
Sergio sulfureus, 468, 469, 473, 546
Sergio trilobata, 468
Sergio trilobatus, 428, 429, 468
Spongiaxius, 4, 10, 16, 18, 149, 172, 193, 508
Spongiaxius brucei, 149, 172, 193, 194, 541,
542
Spongiaxius novaezealandiae, 149, 150
Spongiaxius odontorhynchus, 173
Spongiaxius pitatucensis, 104
Strahlaxiidae, 1, 3, 5, 10, 11, 13, 14, 319, 320
Strahlaxius, 5, 11, 319, 320, 332, 333, 508
Strahlaxius plectrorhynchus, 333
Strahlaxius waroona, 333, 334
Tethisea, 5, 11, 197, 198, 297–299, 301, 309,
311, 525
Tethisea indica, 299, 309
Tethisea mindoro, 309, 310
Thailandcallichirus, 474
Thailandcallichirus gen. nov., 5, 11, 339, 413,
473
Thailandcallichirus ranongensis, 474, 547
615
Thalassia, 207, 534
Thalassia hemprichii, 327, 328
Thalassina anomala, 2
Thalassinida, ix, 1, 2, 9
Thalassinidae, 1, 2, 9
Thalassinidea, ix, 1, 2, 9, 298, 531
Thalassinoidea, ix, 1, 2, 9, 337
Thaumastocheles, 486, 487
Thaumastocheles zaleucus, 116
Thaumastochelidae, 116
Thomassinia, 6, 9, 10, 197, 340, 525, 526, 530,
534–537
Thomassinia aimsae, 526, 535, 536
Thomassinia gebioides, 526, 530, 535, 536
Thomassinia moorea, 530–532, 535, 536
Thomassinia sp., 526, 536, 537
Thomassiniidae, 3, 6, 9–11, 197, 336–338, 340,
364, 525, 526, 534
Thomassiniinae, 525
Tirmizicallichirus, 475
Tirmizicallichirus gen. nov., 5, 11, 339, 356,
413, 414, 466, 474, 475
Tirmizicallichirus masoomi, 475, 548
Trypaea, 5, 10, 12, 335, 339, 354–356, 358, 364,
365, 385–388, 391, 395, 403, 404, 408,
410
Trypaea acanthura, 386, 389, 390, 543
Trypaea acutirostella, 387, 388, 390, 545
Trypaea amboinae, 386, 389, 391, 544, 545
Trypaea amplimaxilla, 354, 386, 388, 391, 544
Trypaea arenosa, 386, 388, 391, 544
Trypaea australiensis, 386, 388, 392, 395, 544
Trypaea bangensis, 387, 388, 392, 544
Trypaea berylae, 386, 387, 393, 543
Trypaea biffari, 386, 387, 393, 543
Trypaea biformis, 386, 388, 393, 543
Trypaea brevirostris, 354, 386, 389, 394, 544
Trypaea caesari, 386, 388, 394, 543
Trypaea caledonica, 386, 390, 395, 544
Trypaea californiensis, 386, 387, 395, 543
Trypaea chakratongae, 387, 390, 396, 544
Trypaea costaricensis, 354, 386, 388, 396, 543
Trypaea debilis, 386, 387, 396, 543
Trypaea exilimaxilla, 386, 389, 397, 544
Trypaea filholi, 386, 388, 397, 544
Trypaea fragilis, 386, 388, 397, 543
Trypaea gigas, 386, 387, 398, 543
Trypaea gruneri, 387, 388, 398, 544
Trypaea intermedia, 386, 389, 390, 398, 545
Trypaea japonica, 361, 386, 389, 395, 399, 405,
410, 544
Trypaea joculatrix, 386, 389, 401, 544, 545
Trypaea lewtonae, 354, 387, 388, 402, 544
616
CRM 013 – Katsushi Sakai
Trypaea lignicola, 387, 389, 402, 544
Trypaea matzi, 386, 390, 403, 544
Trypaea melissae, 386, 388, 403, 545
Trypaea mocambiquensis, 386, 389, 404, 545
Trypaea nieli, 354, 387, 389, 404, 544
Trypaea parva, 387, 389, 404, 544, 545
Trypaea petalura, 386, 389, 405, 544
Trypaea plantei, 386, 390, 405, 545
Trypaea poorei, 386, 388, 405, 544, 545
Trypaea porcellana, 392
Trypaea propriopedis, 387, 389, 406, 544
Trypaea pugnatrix, 386, 389, 390, 406, 545
Trypaea pygmaea, 386, 389, 390, 407, 545
Trypaea rochei, 387, 407, 543
Trypaea sahul, 387, 388, 407, 545
Trypaea santarita, 387, 408, 543
Trypaea sensu nov., 355, 358
Trypaea sibogae, 387, 389, 408, 545
Trypaea sp., 390, 412
Trypaea spinoculata, 387, 389, 409, 544
Trypaea stenomastaxa, 387, 389, 409, 544
Trypaea tabogensis, 354, 387, 409, 543
Trypaea tenuipes, 387, 390, 409, 544
Trypaea thailandica, 354, 387, 389, 409, 410,
544
Trypaea thermophila, 387, 389, 410, 544
Trypaea thorsoni, 387, 388, 410, 545
Trypaea tonkinae, 387, 390, 411, 544
Trypaea truncata, 387, 389, 411, 543
Trypaea uncinata, 387, 412, 543
Upogebia edulis, 2
Upogebiidae, ix, 1–3, 9, 337
Vulcanocalliacinae, 336–339, 347, 348, 350
Vulcanocalliax, 5, 11, 339, 349–351
Vulcanocalliax arutyunovi, 348, 349, 351
Vulcanocalliax arutyunovi n. gen. & n. sp., 348
Zostera, 502
J.c. Vaupel Klein