P r Iad

_

::

_

C 4.

CFZT.M

MEMOIRES

DU MUSEUM

NATIONAL

D’HISTOIRE

NATURELLE

Resultats des Campagnes MUSORSTOM

Volume 18

Coordonne par

Alain CROSNIER

Publie avec le concours de VUniversite Santa Ursula de Rio de Janeiro et VInstitut frangais

de Recherche scientificjue pour le Developpement en Cooperation (ORSTOM)

COM

Source : MNHN. Paris

MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE

Redacteur en chef (Editor-in-Chief) : Jean-Lou Justine

Redacteurs (Editors) : Jean-Marie Betsch, Philippe Bouchet, Christian Erard & Jean-Lou Justine

Assistante de redaction ( Copy editor ) : Bernadette Gottini-Ch arles

Adresse (Address)

Memoires du Museum national d'Histoire naturelle

57, rue Cuvier

F-75005 Paris

Tel. : [33] 01 40 79 34 37

Fax. : [331 01 40 79 38 08

e-mail : memoires@mnhn.fr

Les Memoires du Museum national d'Histoire

naturelle publient des travaux originaux majeurs,

tels que des monographies ou des volumes a au¬

teurs multiples. Les auteurs sont invites, pour

toutes les questions editoriales, a prendre contact

avec le directeur de la publication. Les manuscrits

peuvent etre en fran^ais ou en anglais.

Vente en France :

Museum national d’Histoire naturelle

Service des Publications Scientifiques

Diffusion : Delphine HENRY

57, rue Cuvier

F-75005 Paris

Tel. : [33] 01 40 79 37 00

Fax : [33] 01 40 79 38 40

e-mail : dhenry@mnhn.fr

Parution et prix irreguliers. Les ordres perma¬

nents d'achat et les commandes de volumes sepa-

res sont re?us par le Service des Publications

Scientifiques, Diffusion (pour la France et les DOM-

TOM uniquement), ou par Backhuys Publishers.

Catalogue sur demande. Une liste des derniers

titres parus figure en page 3 de couverture.

The Memoires du Museum national d'Histoire

naturelle publishes major original contributions,

such as monographs or multi-authored volumes.

Prospective authors should contact the Editor-in-

Chief. Manuscripts in French or English will be

considered.

Sales Office:

Universal Book Services

Dr W. Backhuys

P.O. Box 321

NL-2300 AH Leiden

The Netherlands

Tel. : 131] (71) 517 02 08

Fax: [31] (71) 517 18 56

e-mail: backhuys@euronet.nl

Volumes are published at irregular intervals, and

at irregular prices. Standing orders and orders for

single volumes should be directed to the Service

des Publications Scientifiques du Museum,

(France and DOM-TOM only) or Backhuys

Publishers. Price list and catalogues are available

on request. Recently published memoirs are listed

on page 3 of the cover.

Printed on acid-free paper

Imprim6 sur papier non acide

Source :

B i b 1 1 o t hfcque Cen t ra 1 e rtus6um

3 3001 00018401 9

Source : MNHN. Paris

Ce volume des Resultats des Campagnes

MUSORSTOM est dedie a la Mere Fatima Mar on

RAMOS, presidente de VUniversite Santa Ursula, a

Rio de Janeiro. Docteur es Sciences de VUniversite

Pierre et Marie Curie, la Mere Fatima s'interesse

tout particulierement aux recherches oceano-

graphiques qu'elle developpe dans le cadre de

VUniversite qu'elle preside. Elle a participe a de

nombreuses campagnes en mer et, en tant que co¬

chef de mission, a la campagne MD 55, faite avec

le navire oceanographique “ Marion Dufresne ” au

large des cotes du Bresil et consacree,

principalement, a V etude de lafaune bathyale ; elle

suit les travaux effectues sur cette faune par le

Dr Marcos TAVARES, professeur a VUniversite

Santa Ursula, et elle est V artisan de la mise en

place, a cette meme Universite, d'un DEA

d'Oceanographie biologique dont les enseignements

sont assures par des professeurs bresiliens et

frangais. Connaissant de nombreux oceanographes

frangais, la mere Fatima, membre actif de la

cooperation franco-bresilienne en Qceanographie,

s' est egalement interessee aux Resultats des

Campagnes MUSORSTOM et son Universite participe

au financement de la publication de ce volume.

Resultats des Campagnes MUSORSTOM

Volumes deja parus :

Volume 1 : Mem. ORSTOM , 91 : 1-558, 225 fig., 39 pi. (1981). ISBN : 2-7099-0578-7.

Volume 2 : Mem. Mus. natn. Hist, nat., (A), 133 : 1-525, 126 fig., 37 pi. (1986). ISBN : 2-85653-136-9.

Volume 3 : Mem. Mus. natn. Hist, nat., (A), 137 : 1-254, 82 fig., 9 pi. (1987). ISBN : 2-85653-141-5.

Volume 4 : Mem. Mus. natn. Hist, nat., (A), 143 : 1-260, 103 fig., 23 pi. (1989). ISBN : 2-85653-150-4.

Volume 5 : Mem. Mus. natn. Hist, nat., (A), 144 : 1-385, 128 fig., 35 pi. (1989). ISBN : 2-85653-164-4.

Volume 6 : Mem. Mus. natn. Hist, nat., (A), 145 : 1-388, 190 fig., 4 pi. couleur (1990). ISBN : 2-85653-171-7.

Volume 7 : Mem. Mus. natn. Hist, nat., (A), 150 : 1-264, 587 fig. (1991). ISBN : 2-85653-180-6.

Volume 8 : Mem. Mus. natn. Hist, nat., (A), 151 : 1-468, 198 fig. (1991). ISBN : 2-85653-186-5.

Volume 9 : Mem. Mus. natn. Hist, nat., (A), 152 : 1-520, 283 fig., 6 pi. couleur (1992). ISBN : 2-85653-191-1.

Volume 10 : Mem. Mus. natn. Hist, nat., 156 : 1-491, 163 fig., 2 pi. couleur (1993). ISBN : 2-85653-206-3.

Volume 11 : Mem. Mus. natn. Hist, nat., 158 : 1-426, 159 fig., (1993). ISBN : 2-85653-208-X.

Volume 12 : Mem. Mus. natn. Hist, nat., 161 : 1-569, 269 fig., 11 pi. couleur (1994). ISBN : 2-85653-212-8.

Volume 13 : Mem. Mus. natn. Hist, nat., 163 : 1-517, 132 fig., 4 pi. couleur (1995). ISBN : 2-85653-224-1.

Volume 14 : Mem. Mus. natn. Hist, nat., 167 : 1-647, 987 fig., 3 pi. couleur (1995). ISBN : 2-85653-217-9

Volume 15 : Mem. Mus. natn. Hist, nat., 168 : 1-539, 205 fig., 6 pi. couleur (1996). ISBN : 2-85653-501-1.

Volume 16 : Mem. Mus. natn. Hist, nat., 172 : 1-667, 432 fig., 2 pi. couleur (1997). ISBN : 2-85653-506-2.

Volume 17 : Mem. Mus. natn. Hist, nat., 174 : 1-213, 93 fig., ( 1997); ISBN : 2-85653-500-3.

Volume 18 : Mem. Mus. natn. Hist, nat., 176 : 1-570, 458 fig., 7 pi. cquieur (1997). ISBN : 2-85653-511-9.

resultats des campagnes

Volume 18

\MUStuMi

\PARIS

\V

Source

ISBN : 2-85653-511-9

ISSN : 1243-4442

© Editions du Museum national d'Histoire naturelle, Paris, 1997

Photocopies :

Les Publications Scientifiques du Museum adherent au Centre Fran^ais d'Exploitation du Droit de Copie (CFC), 20, rue

des Grands- Augustins, 75006 Paris. Le CFC est membre de V International Federation of Reproduction Rights

Organisations (IFRRO). Aux Etats-Unis d’Am^rique, contacter le Copyright Clearance Center, 27, Congress Street,

Salem, Massachusetts 01970.

Photocopies:

The Scientific Publications of the Museum adhere to the Centre Frangais d’ Exploitation du Droit de Copie (CFC), 20,

rue des Grands-Augustins, 75006 Paris. The CFC is a member of International Federation of Reproduction Rights

Organisations (IFRRO). In USA, contact the Copyright Clearance Center, 27, Congress Street, Salem, Massachusetts

01970.

MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE

TOME 176

ZOOLOGIE

Resultats des Campagnes MUSORSTOM

Volume 18

Coordonne par

Alain CROSNIER

Museum national d'Histoire naturelle

Laboratoire de Biologie des Invert6br6s marins et Malacologie

55 rue Buffon

F-75005 Paris

Publie avec le concours de I'Universite Santa Ursula de Rio de Janeiro et de I'Institut frangais

de Recherche scientifique pour le Developpement en Cooperation (ORSTOM)

EDITIONS

DU MUSEUM

PARIS

1997

Source

Source : MNHN , Paris

SOMMAIRE

Pages

1. Bryozoa: Lepraliomorpha et autres Ascophorina, essentiellement des eaux

neo-caledoniennes (en anglais) . 9

Dennis P. GORDON & Jean-Loup D'HONDT

2. Cirripedia Thoracica: Nouvelles repartitions et nouvelles especes

de Verrucomorpha de I'ocean Indien et du Sud-Ouest Pacifique (en anglais) . 125

John S. BUCKERIDGE

3. Crustacea Isopoda Serolidae: Acutiserolis cidaris et Caecoserolis novaecaledoniae ,

deux especes nouvelles de la mer du Corail (en anglais) . 151

Gary C.B. Poore & Angelika Brandt

4. Crustacea Decapoda : Pseudopandalus curvirostris , genre et espece

nouveaux (Pandalidae) de Nouvelle-Caledonie . 169

Alain Crosnier

5. Crustacea Decapoda : Chelonika macrochela , genre et espece nouveaux appartenant

a la famille des Pandalidae (Caridea) et provenant des eaux neo-caledoniennes

(en anglais) . 177

Charles H. J. M. Fransen

6. Crustacea Decapoda : Crevettes d'eau profonde du genre Plesionika Bate, 1888

(Pandalidae) de la Polynesie franchise. Description de cinq especes nouvelles

(en anglais) . 187

Tin-Yam Chan & Alain Crosnier

7. Crustacea Decapoda : Diacanthurus gen. nov., nouveau genre de Paguridae

represente par des formes actuelles et une forme fossile. Description de deux

especes nouvelles (en anglais) . 235

Patsy A. McLaughlin & Jacques Forest

8. Crustacea Decapoda: Cyclodorippidae recoltes dans I'archipel de Vanuatu

(Brachyura) . 261

Marcos Tavares

9. Crustacea Decapoda : Une revision des especes indo-ouest pacifiques

du genre Calappa Weber, 1795 (Calappidae) (en anglais) . 271

Bella Galil

10. Crustacea Decapoda : Xanthoidea d'eau profonde du Sud-Ouest Pacifique

et de I'ocean Indien occidental (en anglais) . 337

Peter J.F. Davie

11. Pycnogonides recoltes durant ces dernieres annees autour de la Nouvelle-Caledonie

et du Vanuatu (en anglais) . 389

Jan H. Stock

12. Brachiopodes recoltes dans les eaux de la Nouvelle-Caledonie et

des lies Loyaute, Matthew et Chesterfield . 411

Bernard Laurin

13. Pisces Gadiformes : Taxonomie des grenadiers de la region neo-caledonienne

(Pacifique sud-ouest) (en anglais) . 473

Tomio Iwamoto & Nigel R. Merrett

CONTENTS

Pages

1. Bryozoa: Lepraliomorpha and other Ascophorina, mainly from

New Caledonian waters . 9

Dennis P. GORDON & Jean-Loup D'HONDT

2. Cirripedia Thoracica: New ranges and species of Verrucomorpha from the Indian

and Southwest Pacific Oceans . 125

John S. BUCKERIDGE

3. Crustacea Isopoda Serolidae: Acutiserolis cidaris and Caecoserolis novaecaledoniae ,

two new species from the Coral Sea 1^1

Gary C.B. POORE & Angelika BRANDT

4. Crustacea Decapoda : Pseudopandalus curvirostris, new genus and species

(Pandalidae) off New Caledonia (in French) . 169

Alain Crosnier

5. Crustacea Decapoda : Chelonika macrochela , a new genus and

new species of pandalid shrimp (Caridea) from New Caledonian waters . 177

Charles H. J. M. Fransen

6. Crustacea Decapoda: Deep sea shrimps of the genus Plesionika Bate, 1888

(Pandalidae) from French Polynesia, with description of five new species 187

Tin-Yam Chan & Alain Crosnier

7. Crustacea Decapoda: Diacanthurus gen. nov., a new- genus of hermit crabs

(Paguridae) with both Recent and fossil representation, and the descriptions

of two new species . . . . 235

Patsy A. McLaughlin & Jacques Forest

8. Crustacea Decapoda: Cyclodorippidae from the Vanuatu Archipelago

(Brachyura) (in French) . 261

Marcos Tavares

9. Crustacea Decapoda: A revision of the Indo-Pacific species of the genus

Calappa Weber, 1795 (Calappidae) .... 271

Bella Galil

10. Crustacea Decapoda: Deep water Xanthoidea from the South-Western Pacific

and the Western Indian Ocean . 337

Peter J.F. Davie

11. Pycnogonida collected in recent years around New Caledonia and Vanuatu . 389

Jan H. Stock

12. Brachiopods dredged from around New Caledonia and the Loyalty, Matthew

and Chesterfield Islands (in French) . 411

Bernard La UR IN

13. Pisces Gadiformes : Taxonomy of grenadiers of the New Caledonian region,

southwest Pacific . 473

Tomio Iwamoto & Nigel R. Merrett

tSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 18 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULTATS DES C

Bryozoa: Lepraliomorpha and other Ascophorina,

mainly from New Caledonian waters

Dennis P. GORDON

New Zealand Oceanographic Institute

National Institute of Water & Atmospheric Research (NIWA)

P.O. Box 14-901, Kilbirnie

Wellington; New Zealand

&

Jean-Loup D'HONDT

Museum national d'Histoire naturelle

Laboratoire de Biologie des Invertebres marins et Malacologie

57 rue Cuvier

75005 Paris, France

ABSTRACT

This paper describes a fauna of 98 species of ascophorine bryozoans from 1984-89 MUSORSTOM cruises, mainly in the

New Caledonian EEZ. Ten of the species occur solely in the Philippines and some species occur in both regions. The fauna

is noteworthy for its endemism (57 ot the 84 New Caledonian species, i.e., 68%, are endemic) and its high taxonomic

novelty, the latter contributing to a clearer appreciation of the taxonomic limits of some genera and families. Two new

families (Phorioppniidae, Buffonellodidae), 54 new species, and 16 new genera are described, mostly from New

Caledonia; some, from elsewhere, are the consequence of systematic revision. The new genera are: Xynexecha

(Exechonellidae), Parkermavella (Bitectiporidae), Phorioppnia , Oppiphorina , Punctiscutella (Phorioppniidae),

Haswelliporina, Mosaicoporina (Porinidae), Wrigiana, Ijimaia (Calwelliidae), Ipsibuffonella, Maiabuffonella

(Buffonellodidae), Macrocamera (Eminoeciidae), Pseudoplatyglena (Euthyrisellidae), Richbunea (Celleporidae), Lifuella

(Phidoloporidae), and Ptoboroa (Batoporidae). The most speciose family in the collection is the Phidoloporidae,

represented by 7 genera and 19 species. The most speciose genus in the collection is, remarkably, the little-known deep-

sea genus Siphonicytara , with 6 species, all new, which more than doubles the number of species previously described.

Ten of the species in the New Caledonian fauna studied here are shared only with New Zealand, and 4 only with the

Philippines.

Gordon, D.P. & d Hondt, J.-L., 1997. — Bryozoa: Lepraliomorpha and other Ascophorina, mainly from New

Caledonian waters. In: A. Crosnier (ed.), R6sultats des Campagnes MUSORSTOM, Volume 18. Mem. Mus. natn. Hist . nat

176: 9-124. Paris ISBN 2-85653-511-9.

10

D. P. GORDON & J.-L. D HONDT

RESUME

Bryozoa: Lepraliomorpha et autres Ascophorina, essentiellement des eaux neo-caledoniennes.

Ce travail decrit une faune de 98 especes de Bryozoaires Ascophorina, recueillies lors des campagnes Musorstom de

1984 & 1989, essentiellement autour de la Nouvelle-Caledonie. Dix de ces espdces n'ont 6t6 r£coltees qu'aux Philippines,

certaines 6tant communes aux deux regions. Cette faune est remarquable par son enddmisme (57 des 84 esp£ces

n£o-cal6doniennes, soit 68 %, sont endemiques) el son pourcentage 61ev6 en nouveaux taxons, qui ont permis une meil-

leure delimitation taxinomique de certains genres et families. Deux nouvelles families (Phorioppniidae et

Buffonellodidae), 54 espfcces nouvelles et 16 nouveaux genres sont d£crits, pour la plupart de Nouvelle-Caledonie ; la

creation de certains d’entre eux, provenant d’autres regions, est la consequence du travail de revision systematique entre-

pris. Les nouveaux genres sont les suivants : Xynexecha (Exechonellidae), Parkermavella (Bitectiporidae), Phorioppnia ,

Oppiphorina , Punctiscutella (Phorioppniidae), Haswelliporina, Mosaicoporina (Porinidae), Wrigiana , Ijimaia

(Calwelliidae), Ipsibuffonella, Maiabuffonella (Buffonellodidae), Macrocamera (Eminoeciidae), Pseudoplatyglena

(Euthyrisellidae), Richbunea (Celleporidae), Lifuella (Phidoloporidae) et Ptoboroa (Orbituliporidae). La famille

Phidoloporidae presente, dans la collection, la plus riche diversite specifique, avec 7 genres et 19 especes. Le genre le

plus riche en especes dans la collection est paradoxalement un genre profond et peu connu, Siphonicytara, avec 6 especes,

toutes nouvelles, ce qui correspond *1 plus du double du nombre des especes precedemment d6crites. Dix des especes de la

faune neo-caiedoniennc appartiennent aussi e celle de Nouvelle-Zelande, et qualre seulement & celle des Philippines.

INTRODUCTION

This work completes the systematic description of the ascophorine bryozoans collected during recent

MUSORSTOM cruises (1984-1989) mainly in New Caledonian waters. Most of the species dealt with in this account

may be referred to as lepralioid, i.e., having a cryptocystidean shield with an underlying compensation sac (ascus),

but some other ascophorines are also described, not having been previously sorted from the samples. Some of these

were physically attached to lepralioid colonies. Following Gordon (1989a), the Ascophorina are regarded as

including the Cribrilinidae and related families. VOIGT & HlLLMER (1983) have demonstrated the cribrilinid

affinities of certain gymnocystal ascophorines (infraorder Hippothoomorpha) and GORDON and VOIGT (1996) have

presented evidence that the Cribrilinidae (infraorder Cribriomorpha) are the stem group for some (possibly all)

umbonuloid-shielded ascophorines and those lepralioid-shielded taxa that may be derived from them. There is

evidence that, in those Cretaceous Cribrilinidae sensu lato that had a reduced costal shield, there must have been an

ascus proximal to the small area of flexible frontal membrane under the costal area in order for the hydrostatic

mechanism to operate. This ascus is retained in gymnocystal and lepralioid ascophorines and is inferred to occur or

have been present in those umbonuloid species which have a much-reduced area of umbonuloid shield. Thus the

ascus appears to be a symplesiomorphy linking all the ascophorine infraorders.

Some of the species reported on here were collected during the MUSORSTOM 3 cruise to the Philippines, but the

vast majority come from New Caledonian waters. Previous reports in this present series have dealt with 40 species

of Ascophorina from New Caledonia (GORDON & D’HONDT, 1991; GORDON, 1993a; GORDON & BRAGA, 1994).

The present report describes 98 species, of which 84 pertain to the New Caledonian fauna, making a total of 123

New Caledonian ascophorine species taken by these cruises, of which 77 (63%) were new to science. This report

also recognises two new families and 16 new genera. This level of diversity has been helpful in evaluating a

number of generic boundaries and the relationships between certain lepralioid families.

LIST OF STATIONS

Philippines

Estase 2

Station DR 07. — 28.1 1.1984, 05°56.79'N, 126°14.38'E, 890 m: Haswelliporina ?venusta.

Musorstom 3

Station CP 87. — 31.5.1985, 14°00.6’N, 120°19.6'E, 191-197 m: Conescharellina breviconica.

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

1 1

1.6.1985, 14°00.0'N, 120°17.6'E, 189-199 m: Conescharellina breviconica. Conescharellina

Station CP 100.

catella.

Station CP 101. — 1.6.1985, 14°00.15'N, 120°19.25'E, 194-196 m: Conescharellina breviconica.

Station CP 102. - 1.6.1985, 14°00.8'N, 120°17.8’E, 192 m: Characodoma sp., Conescharellina breviconica

Conescharellina catella.

Station CP 106. 2.6.1985, 13°47.0'N, 120°30.3'E, 640-668 m: Conescharellina breviconica , Crucescharellina

japonica.

Station DR 117. - 3.6.1985, 12°31.3'N, 120°39.5'E, 92-97 m: Tetraplaria ventricosa, Actisecos regularis.

Haswelliporina quinaria, Hemismittoidea lennea, Parasmittina serrula, Parasmittina marginata, Parasmittina

sp„ fedora platydiscus, Characodoma areolata. Characodoma biavicularia . Characodoma glabra. Characodoma

parva. Lifuella calyciformis, Reteporellina cruciformis , Triphyllozoon gracile. Conescharellina catella.

Station CP 139. — 6.6.1985, 1 1°53.0’N, 122°14.0’E. 240-267 m: Reteporellina spiramina.

New Caledonia

BlOCAL

Station DW 08. 12.8.1985, 20°34.35'S, 166°53.90’E, 435 m: Haswelliporina multiaviculata, Semihaswellia

umbrella , Siphonicytara vittata.

Station KG 22. 28.8.1985, 22°46.44'S, 166°19.93'E, 2050 m: Terminocella perlucens, Mucropetraliella

philippinensis.

Station DW 33. - 29.8.1985, 23°09.71’S, 167°10.27'E, 675 m: Gen. sp. indet., Haswelliporina multiaviculata.

Siphonicytara armata , Siphonicytara vittata , Buffonellodes crosnieri, Ipsibuffonella repens , Galeopsis mimicus.

Station DW 36. 29.8.1985, 23°08.64'S, 167°10.99'E, 650 m: Siphonicytara vittata , Galeopsis mimicus

Station DW 38. - 30.8.1985, 22°59.74’S, 167015.3PE, 360 m: Tetraplaria sp., Phorioppnia nova , Haswelli-

ponna vaubam , Gigantopora oropiscis , Siphonicytara excentrica , Buffonellaria regenerata , Schedocleido-

chasma sp., Rhynchozoon ligulatum, Reteporella defensa , Reteporellina granulosa.

Station DW 44. - 30.8.1985, 22°47.30'S, 167°14.30’E, 440 m: Siphonicytara vittata, lodictyum blandum,

Reteporellina spiramina.

Station DW 46. - 30.8.1985, 23°00.43'S, 167°28.76'E, 775 m: Emballotheca rare, Lagenipora sp., lodictyum

trochus.

Station DW 51. 31.8.1985, 23°05.27'S, 167°44.95'E, 700 m: Haswelliporina multiaviculata, Siphonicytara

vittata. '

Station DW 65. — 3.9.1985, 24°47.90’S, 168°09.09’E, 275 m: Parkermavella minuta.

Station DW 66. - 3.9.1985, 24°55.43’S, 168°21.67’E, 515 m: Oppiphorina epaxia, Haswelliporina

multiaviculata , Mosaicoporina uniserial is, Mawatarius secundus , Hemismittoidea ennea , Parasmittina glabra,

Parasmittina erecta, Smittina asymmetrica , Yrbozoon ringens , Buffonellaria erecta , Buffonellaria regenerata,

Galeopsis pentagonus, lodictyum trochus , Reteporellina spiramina , Harpago dissidens, Trochosodon sp.

Station CP 67. 3.9.1985, 24°55.44'S, 168°21.55'E, 500 m: Parasmittina erecta, Smittina asymmetrica,

Galeopsis mimicus, Richbunea gracilis.

Station DW 70. - 4.9.1985, 23°24.70'S, 167°53.65'E, 965 m: Haswelliporina multiaviculata. Siphonicytara

glabra, Onchoporoides moseleyi.

Station CP 75. - 4.9.1985, 22°18.65'S, 167°23.30'E, 825 m: Haswelliporina multiaviculata. Galeopsis

mimicus.

Station DW 82. — 6.9.1985, 20°30.65'S, 166°50.30'E, 440 m: Hippothyris caledonica.

Station CP 84. —6.9.1985, 20°43.49'S, 167°00.27'E, 210 m: Hippothoa calciophilia. Smittina abyssicola

Station DS 98. — 7.9.1985, 21°24.10'S. 166°29.76'E, 2365 m: Ichthyaria simplex.

Station KG 101. — 8.9.1985, 21°26.51'S, 166°24.43'E, 1790 m: Ptoboroa gelasina.

Station CP 108. - 9.9.1985, 22°02.55'S, 167°05.68'E, 335 m: Xynexecha pulchra. Phorioppnia cookae.

Haswelliporina multiaviculata. Reteporella concinnoides, Reteporella defensa.

Station CP 109. - 9.9.1985, 22°10.03'S, 167°15.22'E, 495 m: Haswelliporina multiaviculata. Buffonellaria

erecta.

Musorstom 4

Station DW 149. - 14.9.1985, 19°07.60’S, 163°22.70’E, 165 m: Conescharellina atalanta.

Station DW 150. — 14.9.1985, 19°07.50'S, 163°22.10'E, 1 10 m: Conescharellina atalanta.

12

D. P. GORDON & J.-L. D’HONDT

Station DW 151. — 14.9.1985, 19°07.00'S, 163°22.00'E, 200 m: Haswelliporina multiaviculata, Mucro-

petraliella serrata , Iodictyum sp.

Station DW 185. — 18.9.1985, 19°06.20’S, 163°29.50'E, 235 m: Haswelliporina vaubani, Haswelliporina

quinaria, Wrigiana strepsis.

Station DW 187. — 18.9.1985, 19°08.30'S, 163°29.30'E, 65-120 m: Calyptotheca sp., Pseudoplatyglena

mirabilis, Osthimosia sp.

Station DW 220. — 29.9.1985, 22°58.50'S, 167°38.30'E, 505-550 m: Haswelliporina multiaviculata.

Station DW 223. — 30.9.1985, 22°57.00'S, 167°30.00'E, 545-560 m: Rhynchozoon ligulatum.

Station DW 231. — 1.10.1985, 22°33.70'S, 167°10.50'E, 75 m: Hippomenella avicularis, Rhynchozoon

tubulosum , Reteporella concinnoides.

Chalcal 2

Station DW 72. — 28.10.1986, 24°54.50’S, 168°22.30'E, 527 m: Microporella lineata.

Station DW 76. — 30.10.1986, 23°40.50'S, 167°45.20'E, 470 m: Siphonicytara vittata, Galeopsis mimicus.

Reteporella defensa.

Station DW 78. — 30.10.1986, 23°41.30'S, 167°59.60'E, 233-360 m: Macrocamera erecta.

Station DW 81. — 31.10.1986, 23° 19.60'S, 168°03.40’E, 311m: Haswelliporina multiaviculata.

BlOGEOCAL

Station KG 210. — 9.4. 1987, 22°44.00'S, 166°30.97'E, 1 190 m: Domosclerus sp., Crucescharellina aster.

Station KG 21 1. — 9.4.1987, 22°41.80'S, I66°32.53'E, 975 m: Crucescharellina aster.

Station CP 232. — 12.4.1987, 21033.81'S, 166°27.07'E, 760-790 m: Tetraplaria orospinea, Haswelliporina

multiaviculata, Siphonicytara granulosa, Crucescharellina aster.

Station DW 253. — 16.4.1987, 21°31.75'S, 166°28.73'E, 310-315 m: Phorioppnia cookae, Oppiphorina epaxia,

Reteporella sp.

Station CP 260. — 17.4.1987, 21°00.00’S, 167°58.34'E, 1820-1980 m: Bryosartor sp., Semihaswellia ?umbrella ,

Siphonicytara mosaica , Ichthyaria simplex, Crucescharellina aster.

Station KG 261. — 18.4.1987, 21°02.04,S, 167°02.32’E, 1508 m: Ichthyaria simplex .

Station KG 262. — 18.4.1987, 21°02.26'S, 167°02.03’E, 1380 m: Crucescharellina aster.

Station CP 265. — 18.4.1987, 21°04.09'S, 167°00.40'E, 1760-1870 m: Tetraplaria orospinea, Siphonicytara

mosaica, Onchoporoides moseleyi.

Station KG 267. — 18.4.1987, 21°02.20'S, 166°58.76'E, 1935 m: Crucescharellina aster.

Station CP 272. — 19.4.1987, 21°00.04'S, 166°56.94’E, 1615-1710 m: Onchoporoides moseleyi.

Station CP 273. — 20.4.1987, 21°01.53'S, 166°57.4rE, 1920-2040 m: Ichthyaria simplex.

Station KG 275. — 20.4.1987, 21°05.80’S, 166°53.14’E, 1959 m: Fam., gen., sp. indet., Ichthyaria simplex,

Trochosodon sp.

Station KG 287. — 27.4.1987, 20°43.01’S, 166°52.53’E, 1560 m: Ichthyaria simplex.

Station CP 290. — 27.4.1987, 20°36.91'S, 167°03.34'E, 920-760 m: Reteporellina projecta.

Station DW 307. — 1.5.1987, 20°35.38’S, 166°55.25'E, 470-480 m: Kladapheles gammadeka, Richbunea gracilis.

Station DW 313. — 2.5.1987, 20°58.95'S, 166°59.04'E, 1640-1600 m: Ichthyaria simplex.

Station KG 316. — 2.5.1987, 20°48.33'S, 166°53.29'E, 1660 m: Onchoporoides moseleyi, Conescharellina

catella.

Station CP 317. — 2.5.1987, 20°48.12'S, 166°53.16'E, 1630-1620 m: Riscodopa sp., Ichthyaria simplex ,

Trochosodon sp., Crucescharellina aster.

MUSORSTOM 6

Station DW 405. — 14.2.1989, 20°29.75'S, 166°41.00'E, 520 m: Haswelliporina multiaviculata.

Station CP 419. — 16.2.1989, 20°41.65'S, 167°03.70'E, 283 m: Haswelliporina quinaria, Hippothyris caledonica,

Parkermavella fidelis, Smittoidea maunganuiensis multiporosa, Galeopsis pentagonus, Iodictyum

bicuspidatum, Iodictyum trochus, Reteporella defensa , Reteporella ferox.

Station DW 421. — 16.2.1989, 20°26.27'S, 166°40.17'E, 245 m: Haswelliporina quinaria, Galeopsis pentagonus,

Iodictyum illinguum, Reteporella defensa.

Station DW 431. — 18.2.1989, 20°22.25'S, 166°10.00'E, 21 m: Puellina harmeri, Microporella sp., Reteporella

orstomia.

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

13

St3™65- ~ 212 1989’ 21°03.55'S, 167°32.25'E, 480 m: Haswelliporina multiaviculata, Siphonicytara

Station DW 489. - 24.2.1989, 20°48.87'S, 167°05.86'E, 700 m: Galeopsis lageniporoides.

Smib 4

Station DW 37. - 7.3.1989, 24°54.50'S, 168°22.30'E, 540 m: Haswelliporina multiaviculata , Riscodopa parva

Galeopsis mimicus. y r '

Station DW 38. - 7.3.1989, 24°54.50'S, 168°22.00'E, 510 m: Galeopsis pentagons.

SYSTEMATIC ACCOUNT

Suborder ASCOPHORINA Levinsen, 1909

Infraordcr CRIBRIOMORPHA Hanner, 1926

Superfamily CRIBRILINOIDEA Hincks, 1879

Family CRIBRILINIDAE Hincks, 1879

Genus and species indet.

Fig. I

MATERIAL EXAMINED. — New Caledonia. BioCAL: stn DW 33, 675 m.

description. — Colony presumably encrusting, known only from three ancestrulae partially encrusted by

zooids of Buffonellodes crosmeri sp. nov. on a bivalve shell. Ancestrulae relatively large, 0.77-0.81 mm long

0.56-0.72 mm wide, oval, a smooth sloping gymnocyst bordering a field of c. 20 costae; these tapering, with short

pinnae whose tips fuse with those of adjacent pinnae leaving spaces between; no pelmatidia (lumen pores). Orifice

longer than wide, narrowing proximally, the proximal rim rounded; no condyles; a pair of tubercles present on the

Remarks. — No autozooids are present and it is not possible to identify the species

Superfamily BIFAXARIOIDEA Busk, 1884

Family BIFAXARIIDAE Busk, 1884

Genus DOMOSCLERUS Gordon, 1988

Type Species. — Domosclerus piscis Gordon, 1988.

Domosclerus sp.

Figs 2-3

Material EXAMINED. — New Caledonia. Biogeocal: stn KG 210, 1190 m.

SCRIPTION Small ancestrulate colony 3.6 mm long with the bases of two branches arising from the

ronts of zooids. Stem bisenal, the zooids alternating back to back. Zooids 0.66-0.75 mm long, c. 0.25 mm wide

at the orifice. Frontal shield with sparsely scattered pores over the frontal/distal half of each zooid, either side of a

median suture, with fewer pores laterally; peristome not projecting beyond the plane of the frontal shield. Internal

primary zooidal chamber low-walled, entirely lacking costae. Some zooids with an adventitious avicularium on the

14

D. P. GORDON & J.-L. D’HONDT

taolw Mo, ds toed, kenozooidal oriffc. Booked by a poo of ov.clono.

rdmen- The low wa“ “ ,he contli"on m

Aberrodomus Gordon, but that genus has avicularia with cross-bars.

Superfamily CATENICELL01DEA Busk, 1852

Family CATEN1CELLIDAE Busk, 1852

Subfamily DITAXIP0R1NAE Stach, 1935

Genus BRYOSARTOR Gordon & Braga, 1994

TYPE Species. — Bryosartor sulilis Gordon & Braga, 1994.

Bryosartor ? sulilis Gordon & Braga, 1994

Figs 4-6

[7]Bryosartor sutilis Gordon & Braga, 1994: 59, fig. 1 a-i.

MATERIAL EXAMINED. — New Caledonia. Biogeocal: stn CP 260, 1820-1980 m.

Distribution. — Southern New Caledonia and northern Norfolk Ridge, 425-1980 m.

REMARKS - A single incomplete segment was found amongst bryozoan specimens from BIOGEOCAL

Stn CP 260 which may represent a second species of Bryosartor. Comprising only four complete zooids, these are

proportionately slightly longer ,0.58-0.68 mm long, and more slender ,0.26-0.30 mm w.de, wit a branch w^h

of 0.38 mm, than the zooids in the type material. The lateral pore-chambers are more widely separated n the new

specimen, the costal field is shorter in proportion to zooid length, and the proximal gymnocystal pore js more

regularly rounded. The new specimen also represents a depth-range extension. Only more material will determine

thiSSS^r^ e,or in the figure given by Goroon and Braca (1994, for the maximum

number of zooids in a segment of B. sutilis - it should be 16, not 6.

Infraorder HIPPOTHOOMORPHA Gordon, 1989a

Superfamily HIPPOTHOOIDEA Busk, 1859

Family HIPPOTHOIDAE Busk, 1859

Genus HIPPOTHOA Lamouroux, 1821

Type Species. — Hippothoa divaricata Lamouroux, 1821.

Source

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

15

Hippothoa calciophilia Gordon, 1984

Fig. 7

Hippothoa calciophilia Gordon, 1984: 108, fig. 10 A, pi. 42 A-C. — Ryland & Hayward, 1992: 247, fig. 13 a-b.

MATERIAL EXAMINED. — New Caledonia. B10CAL: stn CP 84, 210 m.

DISTRIBUTION. — New Caledonia: northern Norfolk Ridge, 210 m. New Zealand: Raoul Island (Kermadec

Islands), 10-60 m. Australia: Heron Island and Low Isles (Great Barrier Reef), 0-28 m.

REMARKS. — This species characteristically occurs on calcareous substrata. Barnacle plates and oyster valves

have been previously reported; in the present case the sole colony in the collection co-occurred with Smittina

abyssicola on the dorsal surface of an erect fenestrate colony of the bryozoan Smittipora sp. The midfrontal carina

noted by GORDON (1984) in Kermadec H. calciophilia is somewhat more developed in the present material, which

also has predator boreholes in both autozooids and ovicells.

Infraorder UMBONULOMORPHA Gordon, 1989a

Superfamily ARACHNOPUSIOIDEA Jullien, 1888

Family EXECHONELLIDAE Harmer, 1957

Genus XYNEXECHA nov.

Diagnosis. — Colony erect, cylindrical, from an encrusting base. Zooids with a tubular peristome and flared

rim, the frontal shield comprising a central, evenly perforated area separated from an imperforate (save for areolar

pores) periphery by a row of pores that are frequently larger. No avicularia or ovicells. Colony base mostly

comprising kenozooids with perforate frontals.

Type Species. — Xynexecha pulchra sp. nov.

Xynexecha pulchra sp. nov.

Figs 8-9

MATERIAL EXAMINED. — New Caledonia. BlOCAL: stn CP 108, 335 m.

Types. — Holotype: from BlOCAL Stn CP 108, 22°02.55'S, 167°05.68'E, 335 m, MNHN-Bry 20021.

Paratype : same locality as the holotype, MNHN-Bry 19959.

Description. — Colony comprising an erect cylindrical stem from a small encrusting base. Stem -1.3-

1.4 mm diameter, not including projecting peristomes, with zooids arranged more or less in whorls of

4 alternating with the proximal and distal ends of zooids in whorls above and below. Zooids large, 0.84-1.57 mm

long and 0.75-1.03 mm wide, with a tubular peristome 0.37-0.45 mm long that has a slightly flared circular to

subelliptical rim 0.28-0.36 mm across (outer diameter). Frontal shield comprising a slightly convex evenly

perforated umbonuloid field centrally, surrounded by a weakly tubercular imperforate area that has several simple

areolae along its margin; seen laterally, the umbonuloid shield appears supported by small buttresses a little above

the surrounding cryptocyst, the pores between the buttresses generally transversely elongated. Exterior surface of

peristome weakly tubercular like the proximolateral cryptocyst. Primary orifice transversely elliptical (-0.28 mm

wide), with no condyles. No oral spines or avicularia. Dimorphic orifices and/or ovicells not apparent in the sole

specimen. Simple uniporous mural septula occur in the lateral zooidal walls. The base of the colony comprises

16

D P. GORDON & J.-L. D'HONDT

mainly kenozooids. These are smaller in diameter than autozooids; they have a small umbonuloid frontal shield

centrally but lack orifices and peristomes.

Distribution. — New Caledonia: northern Norfolk Ridge, 335 m.

REMARKS. — Xynexecha differs from other Recent exechonellids in its erect mode of growth, the occurrence of

kenozooids in the encrusting colony base, and in the form of the frontal shield with its distinct, slightly raised

umbonuloid area. Some species attributed to Exechonella Duvergier, 1924 have tubular peristomes (e.g.,

Lagenipora tuberculata MacGillivray, 1883, and erect Tubucellaria marginata MacGillivray, 1895 - see GORDON,

1984), but the frontal shield is not zoned and the pores are initially relatively large and rimmed. In contrast, the

type species of Exechonella , E. grandis (Duvergier, 1921), lacks a tubular peristome, though it does have a raised

rim and large frontal pores. It also has an occasional tiny avicularium near the zooidal margin like that in its

congener E. antillea (Osburn) (see COOK, 1985; FRANSEN, 1986).

Superfamily LEPRALIELLOIDEA Vigneaux, 1949

Family LEPRALIELLIDAE Vigneaux, 1949

Genus KLADAPHELES Gordon, 1993b

Type Species. — Kladapheles gammadeka Gordon, 1993b.

Kladapheles gammadeka Gordon, 1993b

Figs 10-11

Kladapheles gammadeka Gordon, 1993b: 208, figs 13-14.

MATERIAL EXAMINED. — New Caledonia. BlOGEOCAL: stn DW 307, 470-480 m.

Norfolk Island. N.Z. Oceanographic Institute, NIWA, unregistered slide of material from NZOI Stn G10, southern

Norfolk Ridge, 970 m depth.

DISTRIBUTION. — New Caledonia between Ouv6a and Lifou, 470-480 m; southern Norfolk Ridge, 970 m.

Remarks. — Ovicells are lacking from both the type and BlOGEOCAL specimens, but a re-examination of

bryozoan material from the type locality has yielded a specimen with a single ovicell. This forms a smooth¬

surfaced bulge distal to the orifice, flush with the surface of the fertile zooid but raised somewhat above the next

zooid in the series and somewhat offset from it. Membranes and opercula are lacking, but it appears that the

ovicellular opening is below the level of the zooidal operculum. In the sole MUSORSTOM specimen one of the

adventitious avicularia is larger than the others in the colony but of the same general form.

Infraorder LEPRALIOMORPHA Gordon, 1989a

Superfamily SMITTINOIDEA Levinsen, 1909

Family BITECTIPORIDAE MacGillivray, 1895

Genus HIPPOTHYRIS Osburn, 1952

Type Species. — Hippothyris emplastra Osburn, 1952.

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

17

Hippothyris caledonica sp. nov.

Fig. 12

Material EXAMINED. — New Caledonia. Biocal: stn DW 82, 440 m.

Musorstom 6: stn CP 419, 283 m.

Types. — Holotype : part of a colony from Musorstom 6 Stn CP 419, 20°41.65’S, 167°03.70’E, 283 m,

MNHN-Bry 20030.

Paratypes : MNHN-Bry 19969 from the same locality as the holotype; MNHN-Bry 19970 and MNHN-Bry

20033, both from Biocal Stn DW 82, 20°30.65’S, 166°50.30'E, 440 m.

DESCRIPTION. — Colony encrusting, forming multiple layers. Zooids large, 0.96-1.18 mm long and 0.66-

0.94 mm wide (periancestrular zooids as small as -0.66 mm x 0.40 mm), subquadrate to subhexagonal, the

frontal shield with 3-4 rows of pseudopores surrounding an extensive imperforate area suborally; ca. 3-4 small

areolae distributed along each lateral margin. Orifice wider than long, attaining 0.25 mm width in the largest

zooids, with a gently concave proximal rim and tiny condyles in the corners. No oral spines. Avicularia paired,

adjacent to the orifice, with a very thin cross-bar and an acute rostrum that is directed distally. Ovicell subglobular,

at least two-thirds of the frontal area with scattered pores and secondary calcification encroaching distally.

Ancestrula resembling later zooids, -0.56 mm long and 0.40 mm wide.

Distribution. — New Caledonia: between Ouvea and Lifou, 283-440 m.

Remarks. — Hippothyris is a small genus, but the number of known species is slowly increasing, especially

from the New Zealand-New Caledonian region. The type species is known from Baja California and north Chile

(Osburn,1952; MOYANO,1991); H. austrinus occurs off central Chile (Moyano, 1991); and H. aganactete and

H. ordinaria are neozelanic (GORDON, 1984, 1989a). All are characterised by large zooids, similar orifices, and an

imperforate suboral area.

Hippothyris caledonica differs from the other species in its paired oral avicularia. The present material encrusts

three-dimensional substrata mostly comprising homotrematid and other Foraminifera, leaving cavities beneath.

Genus PARKERMAVELLA nov.

Diagnosis. — Colony encrusting. Zooids lepralioid, the frontal shield imperforate, with marginal areolar

pores only. Orifice with a proximal sinus. Articulated oral spines occur distally. One or more adventitious

avicularia occur near the orifice or elsewhere on the frontal shield including the lateral margin. Ovicell prominent

or subimmersed, the ectooecial calcification smooth, with numerous perforations that may be rimmed. Secondary

calcification, sometimes incorporating small avicularia may encroach upon the distolateral margin of the ovicell.

Type Species. — Lacerna incurvata Uttley & Bullivant, 1972.

Remarks. — A new genus is necessary for Schizomavella- like species with an imperforate frontal shield and

only marginal areolae - the type species of Schizomavella , Lepralia auriculata Hassall, 1842, and related European

species have evenly perforated shields (Hayward & Ryland, 1979). The late Shane Parker of the South

Australian Museum, before his untimely death in 1992, suggested such a new genus. Parkermavella is named in

honour and recognition of Shane and his work. Apart from the type species, Parkermavella includes, inter alia, the

following Australasian species: Schizoporella punctigera MacGillivray, 1883; Hippomenella curvata Uttley &

Bullivant, 1972 (syn. Schizomavella trachoma Gordon, 1989a); Schizomavella schizoporelloides Gordon, 1984;

and Schizomavella virago Gordon, 1989a.

Parkermavella fidelis sp. nov.

Figs 13-14

Material examined. — New Caledonia. Musorstom 6: stn CP 419, 283 m.

18

D. P. GORDON & J.-L. D'HONDT

TYPE. — Holotype: MUSORSTOM 6: stn CP 419, 20°41.65'S, 167°03.70’E, 283 m, MNHN-Bry 20046. No

separate paratypes.

Description. — Colony encrusting, the zooids variable in size and shape from quadrate to elongate-oval;

0.66-1.13 mm long and 0.37-0.96 mm wide, the calcareous surface smooth to tubercular. Areolar pores 3-7, along

each lateral margin. Orifice with a broadly U-shaped median sinus, flanked by flattened condyles on the shoulders.

Articulated oral spines 5-6. A somewhat columnar, suboral avicularium occurs immediately proximal to the sinus,

the palatal surface slanted to face somewhat distally; the rostrum rounded, with a complete cross-bar. Ovicell

prominent, smooth, with numerous ectooecial pores; apparently closed by the zooidal operculum. At the proximal

end of several zooids is a raised structure with a central concavity; it appears avicularian but in every case it is

sealed frontally and typical avicularian structures are not evident.

Distribution. — New Caledonia: Loyalty Islands, 283 m.

Remarks. — The unique holotype specimen of Parkermavella fidelis is part of a formerly sheet-like

encrustation occurring on a branching stem of Haswelliporina quinaria sp. nov. Thus the underside of the colony is

largely free of any contact with the substratum. The species name fidelis (Latin, faithful) alludes to the geographic

distribution of the species in the Loyalty Islands.

Parkermavella minuta sp. nov.

Fig. 15

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 65, 275 m.

TYPE. — Holotype : a unique, tiny colony, now fragmented into two parts, from BlOCAL Stn DW 65,

24°47.90'S, 168°09.09'E, 275 m, MNHN-Bry 20052. No separate paratypes.

Description. — Colony encrusting, small. Zooids small, 0.18-0.47 mm long and 0.13-0.32 mm wide, the

calcareous frontal shield lightly textured, imperforate, with 3-6 areolar pores along each lateral margin. Orifice with

a small rounded sinus, the entrance to which may be slightly constricted, the condyles flattened on the lateral

shoulders; articulated oral spines 8, extending almost right around the margin of the anter. Avicularia adventitious,

small and oval-shaped, one occurring in the centre of the frontal shield directed proximally and usually another

interzooidally, with a thin, minutely ligulate crossbar, the rostrum with lateral shelves constricting the central

space. Ovicell subimmersed, with small scattered pores and tubercles, the female orifice flanked by 2 pairs of

spines.

DISTRIBUTION. — New Caledonia: northern Norfolk Ridge, 275 m.

Remarks. — Only a single colony 2.2 mm across was found in the collection. Although very small, at least

one mature ovicell is present, indicating that the species probably does not attain a significant size.

Family SMITTINIDAE Levinsen, 1909

Genus SMITTOIDEA Osburn, 1952

Type Species. — Smittoidea prolifica Osburn, 1952.

Smittoidea maunganuiensis multiporosa ssp. nov.

Figs 16-17

Material EXAMINED. — New Caledonia. Musorstom 6: stn CP 419, 283 m.

Source :

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

19

Types. — Holotype : a preserved colony from Musorstom 6 Stn CP 419, 20°41.65'S. 167°03°70'E, 283 m,

MNHN-Bry 20002.

Paratypes : MNHN-Bry 20003, a preserved colony, and MNHN-Bry 20062, a slide, both from the same locality

as the holotype.

Description. — Colony encrusting, producing elevated lamellae beyond the immediate area of substratum.

Zooids 0.84-1.71 mm long and 0.49-1.34 mm wide, the zooids mostly elongate-rectangular in shape. Frontal

shield lightly undular-tubercular, imperforate frontally with numerous marginal areolae. Orifice with a short

peristome laterally and proximally; no oral spines. Lyrula not very broad, the alae short to moderately developed,

flanked by acicular condyles directed toward the alae. Operculum more or less semicircular with a broadly arcuate

proximal edge. Suboral avicularium relatively small, suboval in shape, directed proximally, unequally divided by a

cross-bar with a tiny ligula on the rostral side. Ovicell prominent, subglobular, the ectooecium smooth with

numerous pores.

Distribution. — New Caledonia: north side of Lifou Island, 283 m.

Remarks. — The present material is very similar to the New Zealand species Smittoidea maunganuiensis

(Waters). It differs, however, in a number of minor characters that collectively warrant recognition as a new

subspecies. These comprise a low peristome between the orifice and the avicularium, the more oval shape of the

avicularium, the lack of orificial spines in young marginal zooids, the more numerous ectooecial pores, and the

lack of encroaching secondary calcification around the margin of the ovicell.

Genus HEMISMITTOIDEA Soule & Soule, 1973

Type Species. — Hemismittoidea corallinea Soule & Soule, 1973.

Hemismittoidea ennea sp. nov.

Figs 18-19, ?20

Material EXAMINED. — New Caledonia. Biocal: stn DW 66. 515 m.

?Philippines. Musorstom 3: stn DR 117, 97-92 m.

Type. — Holotype : fragments of the same colony from Biocal Stn DW 66, 24°55.43'S, 168°21.67'E,

515 m, MNHN-Bry 20071. No separate paratypes.

Description. — Colony encrusting. Zooids small, 0.45-0.55 mm long and 0.26-0.40 mm wide, the frontal

shield evenly and lightly textured with small polygonal ridges and depressions, with 7-9 conspicuous areolar pores

along the margins. A thin raised peristome present, with a median U-shaped sinus and 9 small spine bases around

the distal margin (the actual spines lacking from the present material). Primary orifice with a medium-sized alate

lyrula, flanked by acicular condyles directed towards the alae. At the base of the peristome on 1 side is an

avicularium; this is directed proximolaterally towards the adjacent margin; the distal half to two-thirds of the acute

rostral rim minutely serrate, the palate calcified, a crossbar completely lacking. Ovicell prominent, emplaced upon

the frontal shield of the distal zooid, the ectooecium initially smooth, becoming covered with secondary

calcification between the numerous small pores which become sunken; the two most proximal pairs of oral spines

are retained on each side of the fertile orifice.

Distribution. — New Caledonia: northern Norfolk Ridge, 515 m. ?Philippines: Mindoro Strait, 92-97 m.

Remarks. — Hemismittoidea ennea has more oral spines than any other species of the genus, i.e., nine

(greek, ennea). The ovicell differs from other congeners in being evenly covered with secondary calcification. The

unique, fertile holotype colony is unusual in having avicularia that lack a crossbar. It is possible that this is

because of breakage, but they do not appear broken; nevertheless, all other species of Hemismittoidea have

20

D. P. GORDON & J.-L. D'HONDT

crossbars, as do most smittinids. It is this feature that prevents us from including, with certainty, a small infertile

colony of a Hemismittoidea from MUSORSTOM 3 Stn DR 1 17 in the Philippines. It includes periancestrular and

later zooids; there are 8-9 oral spines and greatly resembles zooids of H. entiea but the avicularia have ligulate

crossbars (Fig. 20). If this colony had occurred at the same New Caledonian station as the holotype of H. ennea ,

then we would be more confident of conspecificity.

Genus PARASMITT1NA Osburn, 1952

Type Species. — Lepralia jeffreysi Norman, 1876.

Parasmittina glabra sp. nov.

Figs 21-22, 723-24

Material EXAMINED. — New Caledonia. Biocal: stn DW 66, 515 m.

TYPES. — Holotype : a small colony from BlOCAL Stn DW 66, 24°55.43'S, 168°21.67’E, 515 m, MNHN-Bry

20063.

Paratypes : 3 fragments from the same locality in a separate well on the holotype slide.

Description. — Colony encrusting. Zooids 0.35-0.58 mm long and 0.20-0.52 mm wide, with smooth frontal

shield and 5-9 conspicuous round areolar pores along the margins. Frontal shield rising to a peristomial rim. that

has a median sinus and 7-8 distal spine bases (spines not attached in the present material). Primary orifice with a

medium-sized alate lyrula (-0.041 mm wide) and a pair of subacute non-acicular condyles directed towards the alae.

Avicularia paired, 1 either side of the peristome, each directed lateroproximally towards the adjacent margin; with a

tiny semicircular opesia separated by the crossbar from a minutely and irregularly serrate acute rostrum. Ovicell

prominent, emplaced upon the distal frontal shield, the ectooecial surface almost smooth, with numerous pores

that have slightly raised rims.

Distribution. — New Caledonia: northern Norfolk Ridge, 515 m.

Remarks. — From the same station is additional material (three fragments from another colony) that may

belong to the same species (Figs 23-24). Though infertile, it is extremely similar in general appearance and

orificial characteristics (6-7 oral spines, however), but differs in its slightly larger zooidal size and therefore

proportionately wider lyrula (-0.085 mm) (Fig. 24), and the fact that one of the paired avicularia is frequently

directed distally (Fig. 23).

Parasmittina marginata (Canu & Bassler, 1929)

Figs 25-26

Smittina trispinosa - Okada, 1929: 28. text-fig. 13. Non Johnston, 1838.

Smittina reticulata - Canu & Bassler, 1929: 337, pi. 39, figs 9-10. Non J. MacGillivray, 1842.

Smittina ophidiana var. marginata Canu & Bassler, 1929: 339, pi. 29, figs 4-5.

[?] Smittina trispinosa var. munita Canu & Bassler, 1929: 341, pi. 41, fig. 5.

MATERIAL EXAMINED. — Philippines. MUSORSTOM 3: stn DR 117, 97-92 m.

Distribution. — Japan, Mutsu Bay, Honshu. Philippines, 18-969 m.

Remarks. — This species is distinctively characterised by the long, narrow and curving, near-midfrontal

aviculanum that is proximally directed. It is very similar to Smittoidea paeifica Soule & Soule, 1973; this

species, which has also been illustrated by Ryland and Hayward (1992), has a similar, though much shorter,

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

21

avicularium that is placed somewhat more suborally, and in the median position that is characteristic of species of

Smittoidea. Nevertheless, both species share the characteristically shaped avicularian crossbar which is "angled

slightly forward, and may be thickened at the ends so that the opesial premandibular area of the rostrum is reduced

to a circle" instead of a semicircle.

Parasmittina serrula Soule & Soule, 1973

Fig. 27

Parasmittina serrula Soule & Soule, 1973: 386, fig. 3 D-F. — Gordon, 1984: 96, pi. 35 B-C. — Winston, 1984: 23.

fig. 45. — Ryland & Hayward, 1992: 272, figs 23 e-f, 24 a.

Smittina raigii - Rho & Seo, 1986: 40, pi. 10, figs 3-4. Non Audouin, 1826.

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn DR 117, 97-92 m.

Distribution. — Hawaiian Islands (type locality on Maui), 3-130 m. Belize and Jamaica, -15 m. Kermadec

Ridge, 55-318 m. Heron Island (Great Barrier Reef), < 20 m. Mindoro Strait, Philippines, 92-97 m. Southern

South Korea (no depth given).

Remarks. — The present material, newly recorded from the Philippines, closely matches published

descriptions and illustrations of P. serrula from other localities, though differing in details. There is some variation

in lyrula width throughout its range, being narrower in Caribbean and Hawaiian populations and wider in Great

Barrier Reef and Philippine samples. The present material lacks giant avicularia and the smaller avicularia are not

as high-sided as previously described - the latter possibly a consequence of the degree of secondary calcification.

Parasmittina erecta sp. nov.

Figs 28-29

Parasmittina sp. Gordon, 1985: 171, fig. 19.

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 66, 515 m. — Stn CP 67, 500 m.

New Zealand. NZOI Stn K795, 350 m.

Types. — Holotype : part of a colony from BIOCAL Stn DW 66, 24°55.43'S, 168°21.67’E, 515 m, MNHN-Brv

20069.

Paratypes : MNHN-Bry 20070, from the same location as the holotype; MNHN-Bry 20065, from BIOCAL Stn

CP 67, 24°55.44'S, 168°21.55'E, 500 m.

DESCRIPTION. — Colony erect, dichotomously branching in 1 plane, the zooids facing on 1 side only; branch

width 0.39-0.94 mm, the zooids in 2-4 series; distance between dichotomies 1.20-2.25 mm. Zooids 0.52-0.66 mm

long and 0.24-0.34 mm wide, the frontal shield more or less smooth and imperforate centrally, with -4-7 areolar

pores along each lateral margin. Peristome with a median sinus, bordered distally by 6 oral spines. Primary orifice

with a variable medium to broad alate lyrula flanked by curved condyles. Avicularia single or paired, variable in

length and orientation; generally the rostrum is narrow and acute with small denticulations and directed proximally;

if an avicularium is short (~ half the length of the longer avicularia) and adjacent to the peristome, it may be

directed laterally toward the secondary orifice; if placed more proximally, it may be directed proximally or distally

or obliquely toward the peristome. Ovicell relatively large, emplaced on the distal frontal shield, with numerous

pores in the smooth ectooecium. Dorsal surface faintly textured, with lines marking the zooidal boundaries.

Distribution. — New Caledonia: northern Norfolk Ridge, 500-515 m. New Zealand: southern end of

Kermadec Ridge, 350 m.

Remarks. — Gordon's (1985) record of this species from the southern Kermadec Ridge was based on a

single worn fragment. Nevertheless, close comparison of the Kermadec and Philippine colonies confirms that they

are conspecific.

22

D. P. GORDON & J.-L. D'HONDT

Parasmittina sp.

Figs 30-31

Material EXAMINED. — New Caledonia. Musorstom 3: stn DR 117, 97-92 m.

DESCRIPTION. — A single young colony only 1.26 mm diameter, encrusting, the marginal zooids tending to

suberect, the ancestrula occluded. Largest zooidal dimensions 0.28-0.34 mm long, 0.18-0.26 mm wide. Frontal

shield more or less smooth, rising to a suboral peristome with a median U-shaped pseudosinus. Areolar pores

sparse, usually only 1-2. Primary orifice with a broad alate lyrula; no condyles. Oral spines 6. Adventitious

avicularia single or paired, small and oval with a low, lightly crenulated rostral rim, the mandibular pivots

considerably projecting, approaching each other quite closely; avicularia set near the lateral margins of the zooid

where the peristome starts to rise, directed obliquely laterally.

Remarks. — It has not been possible to determine the species from such a small specimen, but the small

oval avicularia are very distinctive and generally rare in Parasmittina species, so it should be possible to relate the

specimen to larger, fertile colonies should they be discovered.

Genus SMITTINA Norman, 1903

Type Species. — Lepralia landsborovii Johnston, 1 847.

Smittina asymmetrica sp. nov.

Figs 32-34

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 66, 515 m. — Stn CP 67. 500 m.

Types. — Holotype : dome-shaped colony from Biocal Stn CP 67, 24°55.44'S, 168°21.55'E, 500 m, MNHN-

Bry 20005.

Paratypes : MNHN-Bry 20067, and MNHN-Bry 20061, both from BiOCAL Stn DW 66, 515 m.

Description. — Colony encrusting, able to form multiple layers by frontal budding, resulting in a dome¬

shaped colony 9 mm diameter and 5 mm high. Zooids subquadrate to subhexagonal, sometimes wider (0.45-

1.01 mm) than long (0.60-0.98 mm); at the growing margin the frontal shield appears convex and higher than the

lateral margins, rising to an elevated peristomial orifice. Later in ontogeny the margins become raised vertically to

the level of the peristomial rim; as a consequence the frontal shield appears considerably sunken and the hypostegal

coelom is relatively spacious. Frontal shield varies from lightly textured at the colony margin to somewhat

tubercular in the older parts; evenly, but not densely, perforated by relatively large pseudopores. Peristomial orifice

subpyriform with a spatulate suboral avicularium set in the peristomial sinus; beneath is a very broad lyrula whose

alae almost touch the adjacent acicular condyles. No oral spines. Ovicell large, prominent, occupying most of the

frontal area and hypostegal coelom of the next distal zooid, the peristomial orifice angled asymmetrically to one

side, as is the outer of two suboral avicularia set within it; the innermost avicularium is aligned with the zooidal

axis; ovicellular ectooecium regularly and evenly perforated.

Distribution. — New Caledonia: northern Norfolk Ridge, 500-515 m.

Remarks. — Smittina asymmetrica is a very striking species insofar as zooidal characters are concerned,

notably the raised margins and asymmetrical female peristomial orifice with two suboral avicularia, the one above

the other.

Source MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

23

Smittina abyssicola (Harmer, 1957)

Figs 35-36

Porella abyssicola Harmer, 1957: 946, pi. 65, figs 8-10.

MATERIAL EXAMINED. — New Caledonia. BlOCAL: stn CP 84, 210 m.

DISTRIBUTION. — Indonesia: Halmahera, Mollucca Islands, 1089 m. New Caledonia: Loyalty Islands, 210 m.

Remarks. — The present material occurs on the abfrontal side of a fenestrate species of the bryozoan

Smittipora sp. Although infertile, it accords very well with Harmer's (1957) description and illustrations,

including of the elevated peristome, broad lyrula, and ligulate avicularian crossbar.

Family MAWATARIIDAE Gordon, 1990

Genus MAWATARIUS Gordon, 1990

TYPE Specie S. — Prostomaria inexspectabilis Gordon, 1985.

Mawatarius secundus sp. nov.

Figs 37-41

MATERIAL EXAMINED. — New Caledonia. BlOCAL: stn DW 66, 515 m.

Types. — Holotype: a branching, fertile, colony fragment from BlOCAL Stn DW 66, 24°55 43'S

168°21.67'E, 515 m, MNHN-Bry 20043.

Paratype : two fragments, one ancestruiate, from the same locality as the holotype, MNHN-Bry 20055.

Description. — Colony erect, uniserial, dichotomously branching in 1 plane. Zooids 0.66-0.87 mm long and

0.36-0.47 mm wide, mostly all facing in the same frontal direction; from a position adjacent to the proximal end

of the orifice, each zooid produces a daughter zooid (or 2 zooids at bifurcations) at an angle - thus the orientation of

the zooids imparts a zigzag appearance to each branch. Frontal shield densely and minutely granular, with sparse

pseudopores and faint longitudinal ridges. A secondary orifice, composed of a short peristomial rim, is fluted

around the inner face of the rim and has a pseudosinus. At a deeper level, not readily visible frontally, is the

primary orifice, with an anvil-shaped lyrula; the adjacent corners of the poster slightly projecting and flattened,

forming condylar surfaces. Nor oral spines or avicularia. Ovicell appearing as a small bulge externally, the surface

calcification resembling that of the frontal shield, opening into the peristome above the primary orifice.

Ancestrular zooid erect and claviform, proximally tapering.

Distribution. — New Caledonia: northern Norfolk Ridge, 515 m.

Remarks. — Mawatarius secundus is only the second species of the genus (hence the specific name) to be

described. It differs from the type species in having a deeper poster and prominent lyrula and ihe zooidal orifices

mostly facing in the same direction (alternate orifices in M. inexspectabilis in some cases face at right angles to

each other).

Although the ancestrula is present in one of the paratype fragments of M. secundus, it was not attached to any

substratum. It otherwise resembles that of the type species which has a short chitinous ring at the proximal end

attaching it to a flat pad of calcium carbonate. There are no accessory rootlets or encrusting kenozooids.

24

D. P. GORDON & J.-L. D’HONDT

Family PETRALIELLIDAE Harmer, 1957

Genus MUCRO PETRA LI ELLA Stach, 1936

Type Species. — Lepralia ellerii MacGillivray, 1869.

Mucropetraliella philippinensis (Canu & Bassler, 1929)

Fig. 42

Petraliella philippinensis Canu & Bassler, 1929: 261, pi. 35, figs 3-11.

Mucropetraliella philippinensis - HARMER, 1957: 710, pi. 45, figs 19-21. — DUMONT, 1981: 635. — WINSTON, 1986:

21. — D’HONDT, 1986: 705.

Material EXAMINED. — New Caledonia. Biocal: sin KG 22, 2050 m.

Distribution. — Red Sea, < 30 m. Japan, 73-146 m. Philippines, 34-621 m. Sumbawa Island, Indonesia,

0-36 m. New Caledonia, 40-2050 m.

Remarks. — The specimen from Biocal Stn KG 22 was represented by a single tiny infertile colony. The

depth at which it was taken is greater than indicated by previous records. Harmer (1957) described the orificial

sinuses as almost closed, whereas they are rather more open in the present specimen, but this is a character likely

to vary. Certainly the lyrula is characteristically short and wide in the Biocal specimen, conforming to HARMER's

description, but there are also bases of four small oral spines, not previously reported for M. philippinensis . These

spines are extremely short, however, occurring low within the orifice and are easily overlooked. The Biocal

specimen does have the characteristic paired lateral avicularia and the arc of small avicularia distal to the orifice, but

lacks the occasional larger avicularia.

On the underside of the sole colony in the collection is an ancestrula of the catenicellid bryozoan Terminocella

perlucens (Harmer) - it is identical to later zooids in morphology and is separated by a thin chitinous joint from a

short calcified stalk attached to the substratum. This represents a new station record for T. perlucens , reported

earlier by Gordon (1993a) from four other Biocal stations (505-1395 m).

Mucropetraliella serrata (Livingstone, 1926)

Fig. 43

Petralia vultur var. serrata Livingstone, 1926: 95, pi. 6, figs 7-10.

Mucropetraliella serrata - Harmer, 1957: 718, pi. 46, fig. 8, text-fig. 6. — D’HONDT, 1986: 705.

Material EXAMINED. — New Caledonia. Musorstom 4: stn DW 151, 200 m.

Distribution. — New Caledonia, north of Belep Islands, 200 m, and Chesterfield Bank, 15-19 m. Also

Sri Lanka, Philippines, Torres Strait, Queensland, Great Barrier Reef, 0-201 m.

Remarks. — Only one small colony fragment occurred in the collection. It may have been transported from

a shallower depth.

Genus RISCODOPA Gordon, 1989a

Type Species. — Mucropetraliella cotyla Cook & Chimonides. 1981.

Riscodopa parva Gordon, 1989a

Riscodopa parva Gordon, 1989a: 57. pi. 30, figs C-E.

Source :

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

25

MATERIAL EXAMINED. — New Caledonia. Smib 4: stn DW 37, 540 m.

Distribution. — New Zealand: northwest slope of North Island, Challenger Plateau, Bounty Trough, 477-

4059 m. Norfolk Ridge: New Caledonia to Norfolk Island, 540-831 m.

REMARKS. — A single colony 3.19 mm diameter occurred in the collection. This is only the second record of

this abyssal species, which thus ranges from 24°54.5'S to 45°21. 1'S.

Riscodopa sp.

Material EXAMINED. — New Caledonia. BlOGEOCAL: stn CP 317, 1630-1620 m.

Description. — Colony tiny, discoidal, -1.35 mm diameter. Zooids somewhat shield-shaped, with a

U-shaped arrangement of relatively large drop-shaped foramina on the frontal shield, with another foramen

suborally at the open end of the U. Orifice evidently lacking a lyrula or denticles, the operculum high-arched

(0.098-1.30 mm long) with a straight proximal rim. No obvious oral spines. A pair of lateral-oral avicularia

adjacent to the orifice, the mandible similar in shape to the operculum but half the length. Ancestrular zooid

central, the opesial diameter 0.18 mm. Ovicells not present.

Distribution. — New Caledonia: South Loyalty Basin near Lifou, 1620-1630 m.

Remarks. — Only a single tiny colony occurred in the collection. It had become dried and, as a consequence,

was collapsed by surface-tensional effects during dehydration. Thus the zooidal features are somewhat distorted. The

overall form of the colony is as for the other two species of the genus, but the paucity, size, and distribution of

frontal pseudopores is unusual. More specimens are needed to characterise this species adequately.

Superfamily SCHIZOPORELLOIDEA Jullien, 1883

Family SCHIZOPORELLIDAE Jullien, 1883

Genus HIPPOMENELLA Canu & Bassler, 1917

Type Species. — Lepralia mucronelliformis Waters, 1899.

Hippomenella avicularis (Livingstone, 1926)

Fig. 44

Lepralia tuberculata var. avicularis Livingstone, 1926: 93, pi. 5, figs 1-3.

Hippomenella spatulata Harmer, 1957: 1095, pi. 72, figs 27, 31. — Powell, 1967: 378, text-fig. 106 a-f, pi. 17, fig. d.

— D’HONDT, 1986: 735, pi. 8, fig. 6. — Rho & Seo, 1986: 40. pi. 9, figs 1-3.

Hippomenella avicularis - Hayward & COOK, 1983: 81, fig. 19G.

Material EXAMINED. — New Caledonia. MUS0RST0M 4: sin DW 231, 75 m.

Distribution. — Cheju-do, southern South Korea (depth not given); New Caledonia, Loyalty Islands and

Chesterfield Plateau, 0-75 m; Australia: Surprise Shoal, Great Barrier Reef, 51 m (type locality), and Port Jackson,

New South Wales; South Africa: eastern continental shelf and slope, 90-780 m.

Remarks. — The present material comprised one small infertile colony fragment.

26

D. P. GORDON & J.-L. D'HONDT

Family LANCEOPORIDAE Harmer, 1957

Genus EMBALLOTHECA Levinsen, 1909

Type SPECIES. — Eschara quadrata MacGillivray, 1880.

Emballotheca rara sp. nov.

Figs 45-46

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 46, 775 m.

Types. — Holotype : a single preserved colony from BIOCAL Stn DW 46. 23°00.43'S, 167°28.76’E 775 m

MNHN-Bry 19996.

Paratypes: MNHN-Bry 19997, a preserved colony, and MNHN-Bry 20044, three dried colonies on a slide, all

from the same locality as the holotype.

Description. — Colony encrusting, forming a small multilamellate nodule up to -6 mm high and 5 mm

diameter, over the underlying substrate (frequently a small coral fragment), the oldest zooids occurring at the apex

of the nodule. Zooids 0.71-1.00 mm long and 0.47-0.98 mm wide, somewhat quadrate in outline, frequently

parallel-sided, a thin raised line of calcification indicating the boundary between zooids. The presence of long buds

at the colony margin indicates multizooidal budding. Some zooids lack orifices. Zooidal frontal shield evenly

perforated by small pseudopores except for two areas suborally and proximally. Inconspicuous areolar pores, a little

larger than pseudopores, occur along the lateral margins. Orifice wider (0.20-0.32 mm) than long, with a variably

convex proximal rim and a pair of long, horizontal, acicular condyles. No oral spines. Avicularia tiny, borne

singly beside the orifice or on either side, with a thin, complete cross-bar, the rostrum rounded, directed obliquely

towards the centre of the frontal shield. Ovicells relatively large, occupying two-thirds of the frontal shield of the

distal zooid, though not especially conspicuous; the skeletal surface perforated much like the frontal shield except

for an imperforate area midproximally, and, on the most mature ovicell in the colony, at least one of the thin

raised ridges characteristic of the family and genus. Ovicells contain single embryos, 0.28 mm maximum diameter.

distribution. — Northern Norfolk Ridge, 775 m.

Remarks. The nodular form of the colony and the tiny lateral-oral avicularia are distinctive features of this

species.

Genus CALYPTOTHECA Harmer, 1957

Type Species. — Schizoporella nivea var. wasinensis Waters, 1913.

Calyptotheca sp.

Figs 47-48

Material EXAMINED. — New Caledonia. Musorstom 4: stn DW 187, 65-120 m.

DESCRIPTION. — Unique juvenile colony encrusting a segment of the bryozoan Nellia tenella (Lamarck).

Zooids 0.32-0.53 mm long and 0.20-0.42 mm wide, separated by thin raised ridges. Zooidal frontal shield more or

less evenly perforated by pseudopores, with low ridges between the pores; marginal areolae the same size as

pseudopores, sparsely distributed at the corners of the zooid. Orifice with a broad shallow sinus and tiny condyles

No oral spines. Avicularia and ovicells absent from the young colony.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

27

DISTRIBUTION. — Northern New Caledonia, near Belep Islands, 65-120 m.

Remarks. — Without ovicells and avicularia, it is impossible to comment further on the affinities of this

species. On the basis of the morphology of the neanic zooids it could equally be a species of Hippoporina.

Family TETRAPLARIIDAE Harmer, 1957

Genus TETRAPLARIA Tenison-Woods, 1879

TYPE Specie S. — Tetraplaria australis Tenison-Woods, 1879.

Tetraplaria orospinea sp. nov.

Figs 49-52

Material EXAMINED. — New Caledonia. Biogeocal: stn CP 232, 760-790 m. — Stn CP 265, 1760-1870 m.

TYPES. — Holotype : a colony segment from Biogeocal Stn CP 232, 21°33.81'S, 166°27.07'E, 760-790 m,

MNHN-Bry 20028.

Paratype : encrusting base and a detached segment of the same colony from BIOGEOCAL Stn CP 265,

21°04.09'S, 167°00.40'E, 1760-1870 m, MNHN-Bry 20037.

Description. — Colony erect, jointed, from a small encrusting base. Encrusting basal zooids 0.56-0.81 mm

long and 0.28-0.55 mm wide, the frontal shield granular and evenly perforated with pseudopores except suborally.

Orifice a little wider than long, with a short rounded median sinus and inconspicuous flattened condyles at the

corners; bases of 6 oral spines around the distal rim. Erect parts of colony arising from interzooidal kenozooids, the

segments at least up to -5 mm long and 0.85 mm diameter and comprising up to 16 zooids; these not arranged in

distinct longitudinal series but forming a clockwise spiral up the segment. Erect zooids relatively large, 0.81-

1.22 mm long and 0.41-0.74 mm wide, the frontal shield and orifice as for the encrusting zooids, except that oral

spines may be reduced to 4 in number. No avicularia. Ovicells or dimorphic orifices not encountered in the limited

material.

Distribution. — New Caledonia: South Loyalty Basin, 760-1870 m.

Remarks. — Although only limited material of this species occurred in the collection (two segments and an

encrusting base), it is distinctive enough to be recognised as new. It is readily distinguished from other species by

the relatively large size of the zooids and especially the small oral spines, apparently unique in the genus,

occurring both in encrusting and erect zooids.

Tetraplaria ventricosa (Haswell, 1881)

Figs 53-54

Onchopora ventricosa Haswell, 1881: 36, pi. 1, fig. 3.

Tetraplaria ventricosa - Harmer, 1957: 1053, pi. 69, figs 1-4 ( cum syn.). — d'Hondt, 1986: 706. — Hayward, 1988:

318.

Material EXAMINED. — Philippines. MUSORSTOM 3: stn DR 117, 97-92 m.

DISTRIBUTION. — Philippines, Indonesia, Timor, Andaman Islands, Sri Lanka, Seychelles, Mauritius, India,

Queensland, New Caledonia, Fiji, 0-97 m.

28

D. P. GORDON & J.-L. DHONDT

Remarks. — Harmer (1957) included a number of species in the synonymy of T. ventricosa , but these

probably need checking. For example, the present material accords very well with Diploecium simplex

Kirkpatrick, 1888, but the orificial sinus of this species seems narrower, and the zooids less ventricose, than

indicated by Harmer for T. ventricosa . Harmer also suggested that Onchopora mutica Busk, 1855 might be a

senior synonym of T. ventricosa , but regarded the small holotype specimen of O. mutica as being inadequate to

provide clear evidence of conspecificity. Scanning electron micrographs of the uncoated holotype [micrographs

kindly supplied by P.J. CHIMONIDES, The Natural History Museum, London] suggests that O. mutica is indeed a

senior synonym of T. ventricosa but not of D. simplex. Careful SEM comparisons need to be made between all of

the species attributed to T. ventricosa. This is beyond the scope of the present study, so Harmer's synonymy is

provisionally accepted here.

Tetraplaria sp.

Figs 55-56

Material EXAMINED. — New Caledonia. Biocal: stn DW 38, 360 m.

Description. — Colony evidently erect and articulated, the sole specimen in the collection being proximally

tapered; this segment almost 0.45 mm long and -0.91 mm diameter, comprising approximately 30 zooids (some

incomplete) arranged more or less in 5 longitudinal series. Zooids relatively broad, generally transversely diamond¬

shaped, 0.56-0.75 mm long and 0.56-0.81 mm wide. Frontal shield evenly perforated by pseudopores except

immediately suborally and along the lateral margins. Orifice with a wide-V-shaped sinus that may be somewhat

rounded, with condylar processes at the corners. No oral spines per sey but some orifices appear to have had

2-4 tiny ephemeral spines, judging from the tiny holes around the distal margin or some orifices. Avicularia and

ovicells not present.

Distribution. — New Caledonia: northern Norfolk Ridge, 360 m.

Remarks. — The sole specimen almost certainly represents a new species but we hesitate to name it on the

basis of the limited material.

Family PHORIOPPNIIDAE nov.

Diagnosis. Colony erect and branching, with circular stems from an encrusting base or with articulated

segments and basally rooted. Zooids with a pseudoporous lepralioid frontal shield and raised margins. Areolar pores

small, sparse, commonly paired proximally. Zooidal orifice with a wide poster, condyles weak or absent.

Articulated oral spines absent. No avicularia. Female zooids, orifices, and ovicells sometimes enlarged, the

ectooecial surface punctuated like the zooidal frontal shield and covered by the epitheca of the hypostegal coelom.

Remarks. — A new family is proposed for the following four genera: Phorioppnia gen. nov. (Holocene)

Quadriscutella Bock & Cook, 1993 (Oligocene-Holocene), Punctiscutella gen. nov. (Oligocene), and Oppiphorina

gen. nov. (Holocene). Justification for this family is given below.

Genus PHORIOPPNIA nov.

Type Species. — Phorioppnia cookae sp. nov.

Diagnosis. — Colony erect, rod-like, non-articulated, the direction of zooidal budding producing longitudinal

rows trending c ockwise up the stem. Zooidal frontal shield evenly, though sometimes sparsely, pseudoporous

wi h small areolar pores at the proximolateral corners and raised interzooidal ridges. Orifices dimorphic. The

Source MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

29

zooidal orifice lacking obvious condyles, the poster tapering and rounded. No oral spines. No avicularia. Ovicells

prominent though not large, the cctooecial surface resembling the frontal shield, the maternal orifice transversely

broadly elliptic, the ovicell closed by the broad maternal operculum.

Phorioppnia cookae sp. nov.

Figs 57-59

Material EXAMINED. — New Caledonia. Biocal: stn CP 108, 335 m.

Biogeocal: stn DW 253, 310-315 m.

Types. — Holotype: a single fertile stem from Biogeocal Stn DW 253, 21°31.75'S, 166°28.73'E, 310-

315 m, MNHN-Bry 20058.

Paratypes : MNHN-Bry 19961, an ancestrulate colony with three stems, attached to a hydroid, from Biocal

Stn CP 108, 22°02.55'S, 167°05.68’E, 335 m; MNHN-Bry 20059, three stems from the same locality as the

holotype.

DESCRIPTION. — Colony erect, comprising unbranched rod-like stems, 0.67-1. 10 mm diameter. Up to 3 stems

arise from a common periancestrular base; the longest stem in the collection (detached) is 1 cm long. Cross

sections of the stems show them to be quadriserial but the zooids are not arranged in parallel longitudinal series;

rather the series slant slightly obliquely in a clockwise direction around the axis. Ancestrula 0.64 mm long and

0.38 mm wide, longitudinally oval in outline, semierect and attached by ?extrazooidal calcification that envelopes a

hydroid stem. Periancestrular zooids 0.66 mm long and -0.46 mm wide. Postancestrular zooids organised into

stems immediately beyond the periancestrular zooids, large, as small as 0.67 mm long near the ancestrular region,

generally 1.22-1.43 mm long and 0.62-0.75 mm wide; frontal shield lepralioid, evenly, if somewhat sparsely,

perforated by pseudopores, the proximolateral part of the shield appearing somewhat sunken in relation to the

raised zooidal margin. Areolar pores a little larger than the pseudopores are sparse; one pair occurs proximal ly, and

2-3 occur along each lateral margin, usually at the widest part of the zooid and adjacent to the orifice. Orifice

somewhat pyriform, the rounded poster about one-third the orificial length, not clearly delimited from the anter by

any condyles. No oral spines. No avicularia. Female zooids about the same size as autozooids but the orifice

extremely broad and elliptical, with a somewhat thicker, upturned proximal rim. Ovicell somewhat prominent and

recumbent, wider than long, the skeletal surface punctuated like the frontal shield, the opening closed by the

maternal operculum.

Distribution. — New Caledonia: South Loyalty Basin, 310-335 m.

Remarks. — There are a number of ascophorines in the southwestern Pacific that have hippoporine orifices,

i.e., with a broad rounded poster, and evenly pseudoporous frontal shields, several of which have been attributed to

Hippoporina (e.g., Brown, 1952; Powell, 1967; Gordon, 1984, 1989a). As Gordon (1994) has pointed out,

however, Hippoporina , newly included in the Bitectiporidae, has characteristic ‘smittinid’ ovicells, with both

ovicellular layers calcified and unfused and relatively large foramina in the ectooecium. This is in contrast to

‘schizoporellid’ ovicells in which both ovicellular layers may be fused and/or the ectooecial surface resembles the

pseudoporous frontal shield. Accordingly, most, if not all, of the southwest Pacific species previously attributed to

Hippoporina need reclassifying (cf. BOCK & COOK, 1993).

Bock and COOK (1993) included their new genus Quadriscutella in the Euthyrisellidae, noting a resemblance to

Tropidozoum Harmer which also has articulated colonies, raised zooidal margins, and dimorphic orifices. On the

other hand, there are significant differences between these two genera, as BOCK and COOK also pointed out (viz, the

methods of branching and brooding and the presence of an extrazooidal basal coelom in Tropidozoum). The type

genus of Euthyrisellidae, Euthyrisella Bassler, is so very different from Quadriscutella , however, that we are

persuaded to conclude that any similarities to that family are superficial, especially since there are species in the

Musorstom collections that appear even further unrelated to the Euthyrisellidae but have morphological characters

in common with Quadriscutella.

30

D. P. GORDON & J.-L. D’HONDT

One such example is Phorioppnia cookae. It is evidently related to Quadriscutella , which it resembles in

having a sparsely perforated frontal shield, raised zooidal margins, similarly distributed areolar pores (especially the

proximal pair), and dimorphic orifices. By contrast, Quadriscutella typically has enlarged female zooids and ovicells

(not just orifices), composite orificial rims in autozooids, and kenozooidal branch points (not seen in the limited

New Caledonian material). On the basis of these distinctions, we have no hesitation in establishing a new genus,

Phorioppnia. [Phorioppnia is an anagram of Hippoporina.)

Another erect hippoporine species in the MUSORSTOM collection, Hippoporina epaxia Gordon, 1984, though

lacking dimorphic orifices, has raised zooidal margins and appears related. A new genus, Oppiphorina ,

is established for it (see below). There are some similarities between O. epaxia and the articulated species

Quadriscutella punctata Bock & Cook, 1993 which, though it has branch-site kenozooids, lacks the reproductive

dimorphism and characteristic ‘tetra-composite’ orifices of the other species of Quadriscutella. Accordingly,

we propose Punctiscutella gen. nov., with the type species Punctiscutella punctata (Bock & Cook, 1993).

On the basis of these four genera, we propose a new family, Phorioppniidae (Oligocene-Holocene), dis¬

tributed from New Caledonia and the Kermadec Ridge to the Challenger Plateau (New Zealand) and southeastern

Australia.

Phorioppnia cookae is named for Patricia L. COOK (Research Associate, The Museum of Victoria, Melbourne)

in recognition of her outstanding contribution to a knowledge of Bryozoa.

Phorioppnia nova sp. nov.

Figs 60-61

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 38, 360 m.

Types. — Holotype : a single stem from Biocal Stn DW 38, 22°59.74'S, 167°15.3rE, 360 m, MNHN-Bry

20056.

Paratype : MNHN-Bry 20054, from the same locality as the holotype.

Description. — Colony erect, rod-like, 0. 8-0.9 mm diameter, the zooids arranged in 6 series, each series

slanting slightly obliquely in a clockwise direction up the stem. Zooids 0.73-1.07 mm long and 0.39-0.66 mm

wide; frontal shield mostly evenly, though sparsely, pseudoporous, the areolae about the same size as the

pseudopores and distributed as in P. cookae , the lateral rims raised slightly. Orificial characters as for P. cookae ,

except for smaller dimensions. Ovicells not present.

Distribution. — New Caledonia: northern Norfolk Ridge southwest of the lie des Pins, 360 m.

Remarks. — Although infertile, the single fragment of this species very closely resembles P. cookae , even in

the way the longitudinal series of zooids trends gently clockwise up the stem (in contrast to those of Oppiphorina,

for example). The zooids of P. nova are smaller than those of P. cookae , however; postancestrular zooids (the only

ones available in the limited material of P. nova) do not overlap in zooidal length, and scarcely overlap in zooidal

width.

Genus OPPIPHORINA nov.

Type Species. — Hippoporina epaxia Gordon, 1984.

Diagnosis. — Colony erect, rod-like, non-articulated, from an encrusting base; the direction of zooidal

budding producing longitudinal rows trending anticlockwise up the stem. Zooids with an evenly pseudoporous

frontal shield and slightly elevated interzooidal ridges. Orifice with a broad rounded poster barely delimited from the

anter by a pair of weak condyles. No oral spines. No avicularia. Ovicells subimmersed but prominent, more or less

evenly perforated and resembling the zooidal cryptocyst, closed by the zooidal operculum.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

31

Oppiphoritia epaxia (Gordon, 1984)

Figs 62-64

Hippoporina epaxia Gordon, 1984: 76, pi. 25 C-D.

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 66, 515 m.

BlOGEOCAL: stn DW 253, 310-315 m.

DISTRIBUTION. — New Zealand: Raoul and Curtis Islands (Kermadec Ridge), 18-440 m. New Caledonia:

South Loyalty Basin and northern Norfolk Ridge, 310-515 m.

REMARKS. — A new genus is introduced here for Hippoporina epaxia Gordon, 1984, distinguished from

Hippoporina sensu stricto by the ‘pseudoporous’ ectooecium and erect rod-like growth. Oppiphorina epaxia begins

life as an encrusting colony of ramifying biserial ‘runners’; zooids in the encrusting phase can produce ovicells

(Gordon, 1984). In contrast to Phorioppnia , zooidal orifices are not dimorphic and the direction of zooidal

budding is such that the longitudinal rows slant slightly obliquely anticlockwise up the stem. Oppiphorina epaxia

resembles Punctiscutella punctata (see above) in zooidal and ovicellular characters, but lacks the branch-site

kenozooids and articulations of that genus and species.

Hippoporina powelli Gordon, 1989a may possibly be included in Oppiphorina , but until ovicells are found in

this species the generic attribution remains uncertain.

Oppiphorina is an anagram of Hippoporina.

Family, genus and species indet.

Fig. 65

MATERIAL EXAMINED. — New Caledonia. BlOGEOCAL: stn KG 275, 1959 m.

Description. — Colony erect, rod-like, the stem quadriserial, more or less circular in cross section. Zooids

large, weakly calcified and fragile, 1.07-1.13 mm long and 0.32-0.42 mm wide, alternating in back-to-back pairs;

frontal shield convex, the surface somewhat tubercular and imperforate except for a broadly transversely crescentic

band of pores across the middle of the shield. Evidently a pair of similar-sized areolar pores proximally, and 1-2

such pores along each lateral margin. Orifice relatively large, subcircular, the broad poster barely delimited from

the anter by a pair of tiny structures at the sides of the orifice. No oral spines. No avicularia. Ovicells not seen.

Remarks. — The sole material comprises a few tiny stem fragments. The orifice and operculum are

hippoporine but it is not possible to ascribe the specimen to a family with any certainty.

Family PORINIDAE d'Orbigny, 1852

Remarks. — The relationships of the family Porinidae to other lepralioid ascophorines is uncertain. Although

porinids are characterised by erect colonies of spiraminate zooids, the family is currently best accommodated in the

superfamily Schizoporelloidea. Frontal shields of porinids are pseudoporous, with the pores completely penetrating

the shield, as in schizoporellids. The ovicells also appear similar, insofar as the outer layer of ovicellular calcifica¬

tion resembles that of the frontal shield with which it is continuous. Two other families, the Margarettidae and

Siphonicytaridae, have a similar colony form and have ascopores that externally resemble the spiramina of

porinids. The Porinidae appear to be related more closely to the former of these two families, which shares a

similar type of frontal shield. Overall, the Porinidae and Margarettidae have more in common with schizoporel-

loidean families like Tetraplariidae and Phorioppniidae than the Siphonicytaridae, which have only a marginally

areolate frontal shield and, though lepralioid, are constructed more like the umbonuloid family Tessaradomidae.

32

D. F. GORDON & J.-L. D’HONDT

Genus HASWELLIPORINA nov.

TYPE Species. — Haswellina multiaviculata Gordon, 1984.

DIAGNOSIS. — Colony erect, branching, from a small encrusting base that includes at least a few autozooids,

no rootlets. Branches cylindrical, the zooids quincuncial or regularly whorled, with both peristomes and spiramina

tubular and projecting. Frontal shield externally not regularly pseudoporous, the pores tending to be small, and

denser near the zooidal margins. Primary orifice with weak, rounded sinus. Small scattered avicularia associated

with frontal shields and peristomes. Ovicells peristomial, somewhat concealed but detectable externally as bulges.

Pore-chambers tubular, especially axially.

Remarks. — A new genus is established here for Haswellina sensu GORDON (1984). Haswellina Livingstone,

with type species Myriozoum australiensis Haswell, 1881, was interpreted by UTTLEY (1956) to be both

generically and specifically distinct from Spiroporina vertebralis Stoliczka, 1865 and GORDON (1984) followed

him in this. Previously, Canu (1913), BASSLER (1935), and Brown (1952) among others, synonymised

S. vertebralis and M. australiensis. UTTLEY (1956) rejected this synonymy, treating Spiroporina as a junior

synonym of Porina d'Orbigny, 1852, but retaining Haswellina as a genus of Schizoporellidae. Brown's (1952)

concept of Spiroporina was based, not on STOLICZKA's (1865) Miocene species, but on Recent New Zealand

species belonging to Galeopsis Jullien in Jullien & Calvet, 1903 (see Gordon 1984). It is now clear that

UTTLEY (1956) made the same error concerning Haswellina.

Thanks to the courtesy of Dr Norbert Vavra, the opportunity was taken to examine the type specimen of

Spiroporina vertebralis Stoliczka, held at the Naturhistorisches Museum, Vienna. Non-type specimens of Porina

gracilis (Lamarck) (courtesy of the late Shane Parker, South Australian Museum, Adelaide) and of so-called

Haswellina australiensis (Haswell) (courtesy of Mary Spencer Jones, Natural History Museum, London) were also

examined. The conclusions from this examination are:

1. Spiroporina vertebralis and Haswellina australiensis are not conspecific, as UTTLEY (1956) rightly asserted,

but they are congeneric. The latter species differs primarily in that the peristomial avicularia occur only on the

projecting proximal rim; in 5. vertebralis the rim scarcely projects and the avicularia occur around the entire rim.

2. Spiroporina Stoliczka, 1865 may be considered a synonym of Porina d'Orbigny, 1852, as Brown (1958)

rightly concluded. Spiroporina differs only in having zooids in whorls - in every other respect the genera are

identical. Brown (1958) went so far as to reduce S. vertebralis to a subspecies of Porina gracilis. In our opinion,

Porina vertebralis and P. australiensis may be considered full species.

3. There still remain, however, species that resemble Porina but present a distinct "facies". For these species

a new genus, Haswelliporina , is established here. The most immediately obvious, though variable, distinction

is that of pseudopore size and density, so that Haswelliporina zooids appear less obviously porous that those

of Porina in which they are coarser and more evenly distributed. Also, in Haswelliporina both the peristomes

and spiraminal pores are tubular and projecting, with only a few avicularia in the actual peristomial rim, and

the colony base is slightly more extensive, incorporating encrusting autozooids, than we have seen in Porina

species.

Apart from the type species, the following species are here included in Haswelliporina: Haswelliporina quinaria

sp. nov., Haswellina vaubani d'Hondt, 1986, Spiroporina brevitubulata Harmer, 1957, S. venusta Harmer, 1957.

Haswelliporina multiaviculata Gordon, 1984

Fig. 66

Haswellina multiaviculata Gordon, 1984: 74, pi. 25, A; 1985: 177, fig. 27.

Material EXAMINED. — New Zealand. N.Z. Oceanographic Institute: stns K795, southern Kermadec Ridge.

350 m, and K840 ( holotype slide H-273), near Macauley Island, 398-412 m.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

33

New Caledonia. BlOCAL: stn DW 08, 435 m. — Stn DW 33, 675 m. — Stn DW 51, 700 m. — Stn DW 66, 515 m.

— Stn DW 70, 965 m. — Stn CP 75, 825 m. — Stn CP 108, 335 m. — Stn CP 109, 495 m.

MUSORSTOM 4: stn DW 151, 200 m. — Stn DW 220, 505-550 m.

Chalcal 2: stn DW 81, 311 m.

Biogeocal: stn CP 232, 760-790 m.

MUSORSTOM 6: stn DW 405, 520 m. — Sin CP 465, 480 m.

Smib 4: stn DW 37, 540 m.

Description. — Colony erect, dendroid, attaining > 22 mm in height and > 17 mm lateral spread (as indicated

by the largest specimens in the collection). Stems up to 2.5 mm thick at the base; branch tips are 0.5 mm

minimum diameter, with zooids arranged in quincunx in 6 longitudinal series. Zooids -0.74-0.88 mm long and

-0.56 mm wide, the frontal shield relatively smooth, moderately perforated by pseudoporcs, with no clear

boundaries between zooids. Zooidal peristomes up to 0.49 mm long, generally shorter, with an internal

peristomial diameter of 0.094-1.13 mm; the spiraminal opening as a short tube -1.32-0.30 mm proximal to the

base of the peristome where it curves into the frontal shield. Primary orifice suborbicular, a little wider (0.1 1) than

long. Adventitious avicularia common, of 2 kinds - tiny sessile or papilliform subcircular ones (occasionally borne

at the tips of long projections), and larger ones (-0. 13 -0. 15 mm) with long acute rostra; both kinds may occur on

the peristomes, with 2-3 set into or just below the rim. Ovicells peristomial, appearing externally as a bulge in

the frontal shield distal to each fertile peristome.

Distribution. — New Caledonia: D'Entrecasteaux Reefs to Loyalty Islands and northern Norfolk Ridge, 311-

965 m. New Zealand: Kermadec Ridge, 350-412 m; Three Kings Islands, 205-516 m.

REMARKS. — There are numerous colonies and fragments in the MUSORSTOM samples, giving a more

complete picture of the characters of this species than indicated in the limited New Zealand material, hence a full

description is given above. There are some differences in the New Caledonian material - zooids are a little larger

overall with longer peristomes, and the distance between the spiraminal opening and peristome base greater than in

the Kermadec specimens. On the other hand, orificial width, length of the larger avicularia, and internal diameters

of zooids and peristomes are the same in the New Caledonian and Kermadec populations. We therefore conclude

that they are conspecific. Haswelliporina multiaviculata has a similar surface appearance to H. brevitubulata

(Harmer) which, however, lacks acute-mandibled avicularia.

Haswelliporina vaubani (d'Hondt, 1986)

Figs 67-68

Haswellina vaubani d’Hondt, 1986: 729, pi. 1, figs 5-6.

Material EXAMINED. — New Caledonia. BlOCAL: stn DW 38, 360 m.

MUSORSTOM 4: stn DW 185, 235 m.

DESCRIPTION. — Colony erect, branching, the branches in the present material 6-serial, 0.75-0.87 mm

diameter, some curving outwards and upwards. Zooids whorled, the length of each zooid (0.69-0.75 mm) equivalent

to the distance between whorls, the maximum external width of each zooid being the branch circumference divided

by the numbers of whorls* (i.e., 0.39-0.46 mm wide). Frontal shield perforated by numerous small pseudopores,

often in small sulci, that penetrate the innermost thin layer of calcification surrounding the zooidal coelom.

Zooidal peristomes tubular, projecting 0.18-0.35 mm from the zooidal surface, the internal diameter 0.13-

0.15 mm; at the base of the peristome a short tubular spiramen. Peristomial rim incorporating 2-3 tiny acute

avicularia with complete crossbar; similar avicularia occur sparsely on the frontal shield, along with occasional

larger spatulate avicularia directed proximally. Zooidal pore-chambers tubular, most obviously expressed axially in

cross-sections and at growing tips.

Distribution. — New Caledonia: d’Entrecasteaux Reefs, 235-460 m.

34

D. P. GORDON & J.-L. D'HONDT

Remarks. — The present material adds to the description given by d’Hondt (1986), who described material

with larger zooids in only four series. It is superficially similar in external appearance to H. multiaviculata but

may easily be distinguished by its whorled zooids and spatulate adventitious avicularia.

Haswelliporina quinaria sp. nov.

Figs 69-70

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn DR 117, 97-92 m.

New Caledonia. Musorstom 4: stn DW 185, 235 m.

Musorstom 6: stn CP 419, 283 m. — Stn DW 421, 245 m.

TYPES. — Holotype : a single dried colony from MUSORSTOM 6 Stn CP 419, 20°41.65'S, 167°03.70'E,

283 m, MNHN-Bry 20010.

Paratypes : MNHN-Bry 19965, 19966, 20109, all from the same locality as the holotype. — MNHN-Bry

20106, from MUSORSTOM 3 Stn DR 1 17, 12°31.3'N, 120°39.5’E, 97-92 m.

DESCRIPTION. — Colony erect (the largest colony 19 mm high), with cylindrical stems dichotomously

branching at irregular intervals and angles (frequently at right angles), stem diameter between zooidal peristomes

0.45-0.70 mm (depending on the number of zooids in each whorl) in younger parts of branches, thickening to

-1.15 mm near the base of the colony. Zooids generally arranged in whorls of 5, sometimes 4 (especially for

ovicelled zooids), or even 3, the zooids in each whorl alternating with those above and below; zooids -0.84-

0.96 mm long and -0.35 mm wide. Frontal shield lepralioid, the inner face evenly perforated, the pore diameter

greater in the innermost skeletal layer; externally initially relatively smooth and sparsely perforated with tiny

pseudopores, with a single row of slightly larger areolar pores along the common interzooidal furrow; becoming

more densely porous and lacking interzooidal boundaries in older zooids. Primary orifice subpyriform, longer (0.14

mm) than wide (0.10 mm), with no clear distinction between anter and poster owing to a lack of condyles.

Peristome initially a low ridge encircling the primary orifice, with longitudinal ridges on its inner face; becoming

very long (up to 0.49 mm along the dorsal side) and tubular, typically with 3 evenly spaced small acute-mandibled

avicularia, apically directed, in the peristomial rim; maximum rim diameter (including avicularia) 0.25 mm. At the

base of the peristome, on its proximal side is a short-tubed circular spiramen. Tiny adventitious avicularia,

frequently papilliform, occur sparsely on the frontal shield and the side of the peristomial tube, each with an acute

rostrum and complete cross-bar. Ovicells peristomial, visibly externally as densely perforated bulges distal to each

of the zooidal peristomes in the same whorl.

Distribution. — Philippines: Mindoro Strait, 92-97 m. New Caledonia: north of the Belep Islands and

between Ouvea and Lifou, 235-283 m.

Remarks. — Haswelliporina quinaria is very similar to H. vaubani (d'Hondt, 1986) discovered at 450-460 m

south of the d'Entrecasteaux Reefs, but H. vaubani has zooids that attain a larger size and relatively large spatulate

avicularia borne frontally. Such avicularia are absent from all of the many colonies and colony fragments of

H. quinaria in the samples.

Haswelliporina Ivenusta (Harmer, 1957)

Figs 71-73

Spiroporina venusta Harmer, 1957: 851, pi. 56, fig. 12.

Material EXAMINED. — Philippines. Estase 2: stn DR 07, 890 m.

Description. — Sole colony erect, branching, from a small encrusting base 5 mm diameter that is mostly

kenozooidal/aviculiferous with some incorporated autozooids; maximum length of colony 12 mm. Main stem and

branches thick throughout - ranging from 1.96 mm above the base to a minimum thickness of 1.39 mm. Zooids

Source MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

35

not whorled; zooid length and width difficult to determine externally; zooidal internal diameter maximally

0.34 mm. Frontal shield with small, scattered pseudopores of which some are in short, thin sulci. Zooidal

peristomes extending not more than 0.30 mm from frontal surface, the rim relatively thin when new,

with c. 3 tiny oval avicularia set below the rim; internal diameter of peristome 0.15 mm. Spiramen opening

c. 0.15-0.19 mm proximal to the base of the peristome, on a slight eminence, not a tubular projection. Avicularia

of two sizes: small oval avicularia about the same diameter as the spiramen opening (c. 0.047-0.056 mm), and

larger oval/subspatulate avicularia 0.13-0.19 mm long, both kinds with a complete crossbar and well-developed

palate. Ovicells not detected.

DISTRIBUTION. — Indonesia: near Halmahera, 1089 m. Philippines, 890 m.

REMARKS. — Harmer's (1957) description was based on a sole fragment, mostly comprising the small

encrusting base. As a consequence (and because the sole holotype specimen cannot be borrowed), it is not possible

to be certain of conspecificity with the Philippine specimen which is further developed. Importantly, both

specimens have the same two kinds of avicularia, of the same form and dimensions, and the distance between the

spiramen opening and the base of the peristome is the same. In contrast, the internal diameter of the peristome in

the Indonesian specimen, as measured from Harmer's illustration, is only c. two-thirds that of the Philippine one.

Genus MOSAICOPORINA nov.

Type Species. — Porina tricephala Gordon, 1985.

Diagnosis. — Colony branching, erect and cylindrical or repent and uniserial. Zooidal frontal shield perforated

by small pseudopores, the surface minutely mosaic-like and with scattered papillae, some of which may be

aviculilerous. Peristomes long, incorporating the spiraminal tube, the peristomial rim with tiny kenozooids and/or

avicularia. Primary orifice with sinusoid poster. Ovicell presumed to be peristomial.

Mosaicoporina uniserialis sp. nov.

Figs 74-75

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 66, 515 m.

Type. — Holotype : unique colony fragment from BlOCAL Stn DW 66, 24°55.43'S, 168°21.67'E, 515 m,

MNHN-Bry 20107. No separate paratypes.

Description. — Colony repent, branching and uniserial, raised above the substratum by conical extensions of

the basal wall beneath the peristomial area, each zooid producing at least 2 daughter zooids (sometimes 4) by distal

budding. Zooids long (0.94-1.23 mm), nearly parallel-sided and 0.37-0.40 mm wide. Frontal shield sparsely but

evenly perforated by small pseudopores, the surface (including the basal surface) minutely but distinctively mosaic¬

like, with flattened polygonal areas delimited by a shallow network of grooves; scattered papillae occur frontally

and laterally, but not basally. Zooidal peristome fairly long (up to 0.6 mm), flaring a little at the rim, the internal

transverse diameter at the rim up to 0.15-0.18 mm; spiraminal tube incorporated into the peristome and opening at

the rim; the rim edge with up to 8 holes, evidently representing the tiny chambers of kenozooids. Primary orifice

with a sinusoid poster; no condyles. Avicularia not apparent. Ovicells not present.

Distribution. — New Caledonia: northern Norfolk Ridge, 515 m.

Remarks. — A new genus is introduced here for two species of Porinidae that are characterised by the

distinctive minutely mosaic surface of the frontal shield as well as the incorporation of the spiraminal tube into the

peristome. The type species, known only from a single station north of Raoul Island on the Kermadec Ridge, is

36

D. P. GORDON & J.-L. D’HONDT

semi-erect, with triserial branches fusing with the substratum wherever they come into contact with it. The colony

of M. tricephala occurs on small calcareous fragments; early astogeny results in an erect stem attached directly to

the substratum with neither rootlets nor lateral spread of encrusting zooids. The unique holotype colony of

M. uniserialis is not affixed to a substratum, so the method of attachment is unknown.

Genus SEMIHASWELLIA Canu & Bassler, 1917

Type Species. — Porina proboscidea Waters, 1889.

Semihaswellia umbrella sp. nov.

Figs 76-77

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 08, 435 m.

Biogeocal: stn CP 260, 1820-1980 m.

Types. — Holotype : fragment of a fertile colony from Biocal stn DW 08, 20°34.35'S, 166°53.90'E, 435 m,

MNHN-Bry 20111.

Paratype: an infertile colony from the same locality, MNHN-Bry 19964.

DESCRIPTION. — Colony erect and branching (maximally 7-8 mm high in the present material), the single

stem attached directly to the substratum without rootlets or basally encrusting zooids, the top of the stem with

four branches arising near each other, curving outwards in half umbrella-like fashion, the zooids opening on the

lower face. Stem 1.37-1.51 mm diameter, the branches 1.09-1.23 mm diameter, the zooids arranged

in 2-3 longitudinal series. Zooids very large, 1.50-2.76 mm long and 0.66-0.94 mm wide, the frontal shield

lepralioid, the outer surface densely granular with sparse pseudopores and slightly larger areolar pores along the

interzooidal furrows. Zooidal peristomes very prominent, up to 1.36 mm long (generally shorter), with flaring

circular rims 0.47-0.57 mm external diameter, the internal peristomial diameter 0.20-0.34 mm; outer surface of

peristomial tube granular, with low longitudinal ridges that continue onto the rim giving it a slightly corrugated

appearance. Spiraminal tube prominent, extending to 0.38 mm long, with a circular opening 0.094-0.1 13 mm

internal diameter. Primary orifice with a sinusoid poster and no condyles, 0.28-0.34 mm wide. Ovicell fairly

prominent, subglobular, 0.72 mm wide, occurring behind the peristome and opening into it.

Distribution. — New Caledonia: Loyalty Islands, 435-1980 m.

Remarks. — It is not certain if the colony fragment from Biogeocal station CP 260 belongs to this

species. It has the same overall branch and zooidal dimensions; however, the branching is somewhat more open

and there are avicularia, which are lacking in the type specimens. The avicularia are subcircular with a broadly

triangular rostral area and delicate cross-bar. They occur singly adjacent to some of the spiramina.

Semihaswellia is a little-known genus. Apart from the type species (5. proboscidea ), there are currently two

nominal Recent species (5. sinuosa Canu & Bassler, 1928: Gulf of Mexico; S. sulcosa Canu & Bassler, 1930:

Galapagos) and three nominal fossil species (S. exilis , S. tripora , and IS. clara - all species of Canu & BASSLER,

1920 from the Paleogene of North America). With the exception of S. sulcosa , it is likely that none of them is

congeneric with the type species, but all need thorough study using modern techniques. Semihaswellia proboscidea

was collected at 823 m depth off St Thomas in the U.S. Virgin Islands by the "Challenger" Expedition. It has not

been collected since. Scanning electron micrographs (courtesy of Mr P.J. Chimonides, The Natural History

Museum, London) show that the New Caledonian material is unquestionably congeneric. The specimen from

Biogeocal Stn CP 232 resembles the "Challenger" material even more closely in having avicularia; these,

however, do not occur dorsally, as in S. proboscidea (according to Waters, 1899), and occur closer to the

spiramen.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

37

Family ACTISECIDAE Harmer, 1957

Genus ACTISECOS Canu & Bassler, 1927

Type Species. — Actisecos regularis Canu & Bassler, 1927.

Actisecos regularis Canu & Bassler, 1927

Figs 78-81

Actisecos regularis Canu & Bassler, 1927: 11, pi. 1, fig. 13. — Canu & Bassler. 1929: 517, pi. 66, figs 1-4,

text-fig. 215. — Harmer, 1957: 856, pi. 60, figs 12, 16. — Cook, 1966: 208.

MATERIAL EXAMINED. — Philippines. MUSORSTOM 3: stn DR 117, 97-92 m.

Distribution. — Philippines, 84-104 m. Indonesia, 82-88 m.

REMARKS. — The affinities of the Actisecidae are not clear. Superficially, the zooids, their orifices, and

peristomes resemble those of some species of Exechonella , and Harmer (1957) described the operculum as

‘membraniporine’. An ultrastructural examination was made of the interior of the frontal shield in the present

material to ascertain whether or not it is umbonuloid. Apart from a faint line indicating a ring scar, there is no

convincing trace of umbonuloid microstructure. The genus and family are provisionally included in the superfamily

Schizoporelloidea until more information is forthcoming.

Harmer (1957) described the ancestrula as somewhat conical, which it is in the present material. The orifice is

subcircular, with a median cusp on the proximal rim.

Family GIGANTOPORIDAE Bassler, 1935

Genus GIGANTOPORA Ridley, 1881

Type Species. — Gigantopora lyncoides Ridley, 1881.

Gigantopora oropiscis sp. nov.

Figs 82-83

[l]Gigantopora polymorpha - Gordon, 1984: 78, pi. 26, D. Non Busk, 1884.

MATERIAL EXAMINED. — New Caledonia. BlOCAL: stn DW 38, 360 m.

New Zealand. N.Z. Oceanographic Institute, NIWA, unregistered slide of material from NZOI Stns K826, K828,

K840, northern Kermadec Ridge, 390-490 m.

Type. — Holotype : unique specimen from BlOCAL Stn DW 38, 22°59.74' S, 167° 15.31' E, 360 m, MNHN-

Bry 2003 1 . No separate paratypes.

Description. — Colony erect, presumably from an encrusting base, the unique holotype specimen

comprising a single stem 7.8 mm long and 2.09 mm diameter (including projecting peristomes), the zooids

arranged in 4-5 longitudinal series. Zooids large, 0.94-1.34 mm long (measured from one proximal corner to the

proximal edge of the avicularium on the same side) and 0.67-0.96 mm wide, the lateral rims slightly thickened and

raised. Frontal shield evenly perforated by pseudopores in the proximal half, tending to imperforate and minutely

38

D. P. GORDON & J.-L. D'HONDT

tubercular suborally. Primary orifice subcircular, -0.21 mm diameter, the poster about one-third the length,

delimited from the anter by small condyles. Peristomial complex conspicuous, comprising a frontally very broad

bridge with a surface of small tubercles; at each side of the peristomial bridge is an avicularium - each is somewhat

frontally directed, toward the apex of the peristome, with an acute rostrum and complete cross-bar, a narrow palatal

cleft continuing under the cross-bar; externally, the avicularian chamber has several pores frontolaterally; between

the proximal edge of the peristome and the frontal shield is a subhemispherical spiramen that is little seen when

zooids are viewed en face . Ovicell somewhat recumbent on the distal zooid, with a relatively thick bounding rim;

the skeletal surface tubercular and imperforate centrally, but with a number of pores peripherally.

Distribution. — New Caledonia; northern Norfolk Ridge southwest of the lie des Pins, 360 m.

New Zealand: northern Kermadec Ridge, 390-490 m.

Remarks. — The present material is distinguished from other species of Gigantopora in the southwest Pacific

by the broad frontal extent of the peristomial bridge that largely conceals the spiramen when zooids are viewed en

face, as well as by details of the avicularia and ovicell. The New Caledonian specimen resembles infertile

encrusting New Zealand material from the northern Kermadec Ridge mistakenly attributed by GORDON (1984) to

Gigantopora polymorpha (Busk). The Kermadec material resembles G. oropiscis in zooidal and orificial

dimensions, in the lack of a non-protruding spiraminal rim, and in the general appearance of the zooids and

peristomial orifice. The frontal extent of the peristomial bridge is proportionately not as great as in the

New Caledonian specimen; also the avicularian palatal cleft is mostly quite open in the Kermadec zooids, except

for one zooid in which it is as narrow as in G. oropiscis. On balance, we conclude that the Kermadec specimens are

probably conspecific, and reflect differences between zooids in encrusting and erect parts of colonies. Only the

finding or more material from both areas will settle the matter.

The species name, oropiscis , is derived from the Latin os , oris , mouth, and piscis, fish, alluding to the

resemblance of the peristomial orifice to the gaping mouth of a fish.

Family MICROPORELLIDAE Hincks, 1879

Genus MICROPORELLA Hincks, 1877

Type Species. — Eschara ciliata Pallas, 1766.

Microporella lineata Canu & Bassler, 1929

Microporella lineata Canu & Bassler, 1929: 332, pi. 40, fig. 5. — Gordon, 1984: 102, pi. 39, A.

Material EXAMINED. — New Caledonia. Chalcal 2: stn DW 72, 527 m.

Distribution. — Philippines, near Luzon, 192 m. New Caledonia, northern Norfolk Ridge, 527 m.

New Zealand, Kermadec Ridge, 140-500 m.

Remarks. — This is only the third record of this species, which extends from 13°20'N to 33°02rS, and 192-

527 m. It is easily distinguished by its uniserial colony form.

Microporella sp.

Figs 84-88

Material EXAMINED. — New Caledonia. Musorstom 6: stn DW 431, 21 m.

Description. — Colony encrusting. Zooids 0.33-0.42 mm long and 0.22-0.28 mm wide, the frontal shield

evenly distributed with small tubercles and pseudopores. Orifice with a smooth proximal rim and 6 oral spines.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

39

Ascopore denticulate, non-reticulated. Avicularia single, originating adjacent to the ascopore but appearing more

proximally when ontogeny is complete, almost at the half-way point near the lateral margin and directed laterally.

Ovicell not encountered.

Distribution. — New Caledonia: west of Ouv6a Island, 21 m.

REMARKS. — This species is represented by a single tiny ancestrulate colony occurring on Reteporella

orstomia sp. nov. It greatly resembles the specimen illustrated by GORDON (1984) from the Kermadec Ridge that

was attributed to Microporella ciliata. This species, which has been accorded a wide distribution around the world

is, however, unlikely to be found beyond ports and harbours in the Australasian region. Both the Kermadec and

Ouv<5a specimens were infertile. It is likely that, when fertile material of this species is found, it will provide

evidence that this represents a new species.

It is interesting to note that in newly developed zooids the pseudopores are reticulate, as in Fenestrulina. The

proximal rim of the orifice may also have minute irregularities, but these denticulations are smoothed over as

ontogeny is completed.

INCERTAE SEDIS

Family CALWELLIIDAE MacGillivray, 1887

Genus ICHTHYARIA Busk, 1884

Type SPECIES. — Ichthyaria oculata Busk, 1884.

lchthyaria simplex sp. nov.

Figs 228-230

MATERIAL EXAMINED. — New Caledonia. BIOCAL: stn DS 98, 2365 m.

Biogeocal: stn CP 260, 1820-1980 m. — Stn KG 261, 1508 m. — Stn CP 273, 1920-2040 m. — Stn KG 275,

1959 m. — Stn KG 287, 1560 m. — Stn DW 313, 1640-1600 m. — Stn CP 317, 1630-1620 m.

Types. — Holotype : an alcohol-preserved, ovicelled, incomplete colony (ancestrula and rootlets lacking) from

Biogeocal Stn DW 313, 20°58.95’S, 166°59.04'E, 1640-1600 m, MNHN Bry-19991.

Paratypes : two small colonies in the same container as the holotype and from the same locality, MNHN Bry-

19994; other colonies from Biocal Stn DS 98, 21°24.10'S, 166°29.76'E, 2365 m, MNHN Bry-19995;

Biogeocal Stn CP 260, 21°00.00'S, 167°58.34,E, 1820-1980 m, MNHN Bry-19992; Biogeocal Stn KG 287,

20°43.0rS, 166°52.53'E, 1560 m, MNHN Bry-19993.

Description. — Colony erect, unilaminar and biserial, unbranched, exceedingly delicate and flaccid, to 8 mm

high, not including rhizoids, maximum width 0.80 mm and a thickness of c. 0.17 mm. The sides of the colony

more or less parallel except where it tapers proximally towards the ancestrula. Zooids alternating, 0.77-1.16 mm

long and 0.37-0.39 mm wide, increasing in length along the colony distally, with a high-arched orifice that has a

straight proximal rim. Distal to the orifice is a median short stumpy, non-articulated spine flanked by a pair of

septular pores on each side. The septular pores continue along the lateral margins, connecting the visceral and

hypostegal coeloms. Proximal to the orifice is a pair of windows in the frontal calcification; these may be unequal,

or even single in zooids near the ancestrula. Presumed ascopore large, open, off-centre. At the junction where three

zooids abut, 3-4 interzooidal communication pores can be seen in face view in the lateral wall, between two of the

zooids, which slants at a low angle at this point. Ovicell large, recumbent on the frontal face of the distal zooid,

0.54 mm long and 0.41 mm wide, the endooecium lightly calcified, the ectooecium entirely membranous, opening

widely above the zooidal operculum. Rhizoids, originating from septular pores, descend the lateral margins of the

40

D. P. GORDON & J.-L. D HONDT

colony on both sides, converging with those from the ancestrula to form a tangle of dichotomously forking

rootlets.

Distribution. — New Caledonia, Loyalty Basin and around Lifou, 1508-2365 m.

Remarks. — Zooids typically have an oval hole in the frontal shield on the side adjacent to the median axis.

At first it was thought these were predator bore-holes, but they appear to occur in each zooid. It is difficult to be

certain but they appear to be ascopores. Certainly the suboral pores are not ascopores, being of the same

construction as the lateral septular pores and occluded with tissue centrally.

Five species of Ichthyaria are recognised here - the type species and /. profunda d'Hondt, 1981 both occur in

deep water off Uruguay, and /. plana (David & Pouyet, 1986) occurs in the western Indian Ocean. Onchopora

grimaldii Jullien & Calvet, 1903 and O. picoensis Jullien & Cal vet, 1903 each have a toothed crescentic ascopore

but the pattern of budding is characteristic of Ichthyaria. Insofar as the shape of the ascopore is a variable character

in microporellid genera, we regard the pattern of budding as having greater taxonomic significance and so include

the two species of JULLIEN and CALVET in Ichthyaria , but both species need re-examining. Ichthyaria simplex

represents the first record of the genus in the Pacific. Unusually, however, /. simplex has an offset oval ascopore,

whereas /. oculata and /. plana have a circular ascopore (Harmer, 1902; David & Pouyet, 1986) and /. profunda

has a slit-like ascopore.

The Calwelliidae are not very speciose. Five genera have been accepted in recent years (Calwellia W. Thomson,

1858; Ichthyaria Busk, 1884; Malakosaria Goldstein, 1882; Onchoporella Busk, 1884; Onchoporoides Ortmann,

1890) and two more are introduced below. Onchoporoides is monotypic and none of the others has more than five

species. The genera are reasonably well circumscribed with the possible exception of Ichthyaria and Onchoporoides.

The latter could be viewed as a laterally expanded Ichthyaria. Onchopora plana David & Pouyet, 1986 (here

included in Ichthyaria , Onchopora being a junior synonym of Margaretta Gray), links the two genera. It is

2-3-serial and also has a circular ascopore. There has been some doubt over the generic allocation of Calwellia

uniserialis Powell, 1967. GORDON (1989a) pointed out similarities with Ichthyaria - the species is here included in

a new genus introduced below.

One other species, Brettia ijimai Okada, 1921, attributed by Sil£n (1941) and subsequent authors to the

savignyellid genus Halysisis , represents a new, sixth, genus of Calwelliidae. Thanks to the courtesy of Dr Ji Eun

Seo (Woosuk University, Republic of Korea), it has been possible to examine material of this species, which

Okada (1921, p. 26, text-fig. 4) illustrated very well. The species is uniserial and branching, lightly calcified,

with a sinus instead of an ascopore. The presence of a pair of uncalcified windows adjacent to the sinus and a

typical calwelliid ovicell (i.e., calcified endooecium and membranous ectooecium) give evidence of a calwelliid

affinity. For this species, which ranges from Korea through Japan to the Bonin (Ogasawara) Islands, we propose

the new generic name Ijimaia , and the new combination Ijimaia ijimai (Okada).

Genus WRIGIANA nov.

Type Species. — Wrigiana strepsis sp. nov.

DIAGNOSIS. — Colony erect, dichotomously branching, jointed, anchored by rootlets. Zooids with a narrow

caudate portion proximally; strictly biserial in internodes and budded more or less back to back, but the dilated part

of the zooid may be skewed so that the zooids in a branch face more or less in the same direction. Orifice

semicircular, with short oral tubercles, and suboral foramina and an ascopore. Ovicell conspicuous, opening above

the operculum, the ectooecium entirely membranous.

Wrigiana strepsis sp. nov.

Figs 231-233

Material EXAMINED. — New Caledonia. Musorstom 4: stn DW 185, 235 m.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

41

Types. — Holotype : an alcohol-preserved colony 1 1 mm high, from Musorstom 4 Stn DW 185, 19°06.20'S,

163°29.50'E, 235 m, MNHN Bry-19960. No other types.

Description. — Colony erect, dichotomously branching, jointed, anchored by rootlets, attaining 1 1 mm

height and 12 mm lateral spread. Zooids 0.67-1.14 mm long (including cauda, 0.07-0.41 mm long) and 0.20-

0.23 mm wide, somewhat claviform in shape with a tapering caudal portion. Zooids biserial, arranged more or less

back to back, with the proximal half or more of each zooid base attached to the dorsal side of its parent, but the

zooids often skewed so as to face somewhat in the same direction, though this is variable. New branches originate

with a single zooid that has a very short caudal portion and chitinous joint proximally. This zooid originates

behind the orifice or on a distolateral shoulder of the parent zooid. The second zooid of a branch is budded back-to-

back from the first via a long caudal portion attached to almost the entire proximal half of the parent zooid. Both

the parent and first daughter zooids of a branch bud zooids distally, re-establishing branch biseriality. Orifice

semicircular, high-arched, with 4 distal spine-like tubercles; suborally, a pair of foramina and a median ascopore.

Ovicell conspicuous, c. 0.33 mm long and 0.32 mm wide, the entooecium with longitudinal striations, the

endooecium entirely membranous.

Distribution. — New Caledonia, east of the Belep Islands, 235 m.

Remarks. — It is clear that Calwellia uniserialis Powell, 1967 belongs to this genus. As Powell (1967)

pointed out, although the branches appear uniserial, each zooid has a tubular caudal portion that occurs along the

dorsal side of the alternate zooidal dilatation to insert on the zooid below. Zooids in the branches face the same

way, adding to the impression of uniseriality. Wrigiana uniserialis (Powell) comb. nov. is known only from

549 m depth in the vicinity of the Three Kings Islands, New Zealand, thus the genus ranges from the Belep

Islands to the Three Kings Islands.

In having a narrow caudal portion to the zooid, Wrigiana resembles Calwellia which, however, has zooids

arranged back to back in regularly alternating decussate pairs. The name Wrigiana is derived from the Anglo-Saxon

noun wrigian, turn, twist, wry, alluding to the frequent skewing of the zooidal dilatation. The name of the type

species, strepsis, is a Greek feminine noun meaning a turning or twisting.

Genus ONCHOPOROIDES Ortmann, 1890

Type Species. — Carbasea moseleyi Busk, 1884.

Onchoporoides moseleyi (Busk, 1884)

Carbasea moseleyi Busk, 1884: 56, pi. 33, fig. 4. — LEVINSEN, 1909: 75, 264.

Flustra moseleyi - JELLY, 1889: 102.

Onchoporoides moseleyi - Ortmann, 1890: 12. — GORDON, 1989a: 64, pi. 35 A (cum syn.).

Material EXAMINED. — New Caledonia. Biocal: stn DW 70, 965 m.

Biogeocal: stn CP 265, 1760-1870 m. — Stn CP 272, 1615-1710 m. — Stn KG 316, 1660 m.

DISTRIBUTION. — New Caledonia and northern Norfolk Ridge, 965-1870 m; New Zealand (Kermadec Ridge to

Solander Trough), 526-2677 m.

Remarks. — The specimen from BIOGEOCAL Stn KG 316, Loyalty Basin, is the northernmost record of this

monotypic genus, of which the type locality is the Kermadec Ridge northwest of Macauley Island.

Family BUFFONELLODIDAE nom. nov.

DIAGNOSIS. — Colony encrusting. Zooids with an imperforate frontal shield. Orifice with a median sinus or

broad poster; the latter may have a median convexity. Avicularium adventitious, single, borne suborally or

42

D. P. GORDON & J.-L. D'HONDT

frontally on the shield, the crossbar complete. Ovicell with a smooth imperforate endooecium and membranous

ectooecium.

Remarks. — A new family name is introduced here to replace the Buffonellidae Jullien, 1888, based on

Buffonella Jullien, 1888, preoccupied by Buffonella Keferstein, 1868 (Amphibia). The Buffonellodidae is related to

the Lacernidae Jullien, 1888 (see Parker & Gordon, 1992), from which it differs in lacking frontal-shield pores,

and possibly the Eminooeciidae Hayward & Thorpe, 1988, which differs in ovicellular and other details. Included

genera are Aimulosia Jullien, 1888, Buffonellodes Strand, 1928, Ipsibuffonella gen. nov. (see below), and

Maiabuffonella gen. nov. (type species Buffonellodes madrecilla Gordon, 1989a, characterised by a personate

ovicell and laterofrontal avicularium).

Genus BUFFONELLODES Strand, 1928

Type Species. — Buffonella rimosa Jullien, 1888.

Buffonellodes crosnieri sp. nov.

Figs 1, 89-90

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 33, 675 m.

TYPES. — Holotype : an encrusting colony on a piece of bivalve shell from BlOCAL Sin DW 33, 23°09.71'S,

167°10.27'E, 675 m, MNHN-Bry 20191; gold-coated.

Paratypes : parts of the same colony, MNHN-Bry 20179, uncoated.

Description. — Colony encrusting, somewhat linear in form, with a pluriserial lobe extending outward from

the ancestrular region on each side. Ancestrula occluded in present material but evidently had a small opesia

encircled by spines, 0.37 mm long and larger overall than the two immediately succeeding daughter zooids (one

middistal, the other extending laterally to the right of where the ancestrula and other daughter zooid join) which

range from 0.30-0.32 mm long and 0.20-0.26 mm wide; later zooids can attain 0.75 mm long and 0.70 mm wide.

Frontal shield smooth, rising to the peristome that surrounds the orifice. Interrupting the peristomial ridge distally

are 3 articulated spines. Primary orifice with a median, shallow V-shaped sinus flanked by ledge-like condyles.

A median suboral avicularium present, set in the peristomial rim, oval in shape with a complete crossbar,

the chamber symmetrically developed from concealed septular pores on each side. Ovicell somewhat recumbent on

the distal zooid, with a smooth presumed endooecium and membranous ectooecium. Small basal pore-chambers

present around the distolateral margins.

Distribution. — New Caledonia, southwest of lie des Pins, 675 m.

REMARKS. — Unlike B. rimosa , several species, including B. crosnieri, have a low peristome surrounding the

orifice. Buffonellodes crosnieri also differs in its linear colony form, the small shallow sinus, and the triplet of oral

spines.

The species is named for Alain Crosnier, in recognition of his unstinting efforts in bringing the results of the

MUSORSTOM cruises to publication.

Genus IPSIBUFFONELLA nov.

Type Species. — Ipsibuffonella repens sp. nov.

Diagnosis. — Colony encrusting, uni- to pluriserial. Zooidal frontal shield imperforate. Areolar pores

lacking. Orifice with a broad arcuate poster, condyles scarcely developed. No peristome or oral spines. Avicularium

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

43

suboral, median. Oviceil globular, the endooecium smooth, imperforate; ectooecium membranous; ovicell not

closed by zooidal operculum.

REMARKS. — Ipsibuffonella is established for /. repens sp. nov. from the northern part of the Norfolk Ridge

near New Caledonia. The genus differs from Buffonellodes chiefly in lacking articulated oral spines and a distinct

orificial sinus flanked by ledge-like condyles.

Ipsibuffonella repens sp. nov.

Figs 91-92

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 33, 675 m.

TYPES. — Holotype : Preserved colony from BIOCAL Stn DW 33, 23°09.71'S, 167°10.27'E, 675 m, MNHN-

Bry 19971.

Paratypes : MNHN-Bry 19972 (preserved) and MNHN-Bry 20042 (dried), both from the same location as the

holotype.

DESCRIPTION. — Colony encrusting, ramifying, uni- to pluriserial, seldom exceeding more than 4 zooids

across. Zooids relatively long, sometimes claviform with a caudate proximal portion; 0.88-1.4 mm long, and

0.54-0.75 mm wide across the dilated portion of the zooid. Frontal shield smooth, imperforate, quite convex with

the sides sloping to the interzooidal furrow or the substratum where the zooidal margin may flatten a little at the

edge. No marginal areolar pores, but a tiny pair of pseudopores is associated with the avicularian umbo, 1 on either

side; in transparency, the avicularium chamber is seen to originate from a tiny, concealed, uniporous septulum a

little beyond each pseudopore. Orifice a little wider than long, with a broad arcuate poster; no separate condyles per

se, the proximal edges of the distal oral rim serving as foci of articulation where they encounter the lateral corners

of the poster. No oral spines. Suboral avicularium median, occurring on the distal face of an associated umbo, with

a rounded rostrum and thin crossbar. Ovicell prominent, a little longer than wide, the endooecial surface smooth,

imperforate, the ectooecium membranous; not closed by the zooidal operculum.

DISTRIBUTION. — New Caledonia: northern Norfolk Ridge, south of lie des Pins, 675 m.

Remarks. — Ipsibuffonella differs from Buffonellodes sensu lato in the uniseriality of parts of the colony

(hence ip si-, from ipsos, Greek, ivy) and the particular combination of zooidal characters. For example, the type

species of Buffonellodes, B. rimosa Jullien, 1888, has, by contrast, oral spines, marginal areolae, an orificial

sinus, and well-developed condyles (GORDON, 1984; Hayward, 1991).

Several colonies occurred on the inner concave surface of a bivalve shell. Microscopic examination of several

alcohol-preserved zooids by transmitted-light microscopy revealed stages of spermatogenesis in one colony; other

colonies had well-developed embryos in the ovicells. The colonies were collected in the month of August (1985).

Family EMINOOECIIDAE Hayward & Thorpe, 1988

Genus MACROCAMERA nov.

Type Species. — Macrocamera erecta sp. nov.

Diagnosis. — Colony erect and rigid, dichotomously branching, the zooids facing on one side. Zooidal frontal

shield imperforate, with sparse areolar pores. Orifice with a wide V-shaped poster delimited from the anter by

condyles. No oral spines. Suboral avicularium present, the crossbar complete. Female zooid with somewhat wider

orifice; ovicell huge, the ectooecial surface resembling the frontal shield; not closed by the zooidal operculum.

44

D. P. GORDON & J.-L. D'HONDT

Macrocamera erecta sp. nov.

Figs 93-94

MATERIAL EXAMINED. — New Caledonia. Chalcal 2: stn DW 78, 233-360 m.

Types. — Holotype: a fertile colony (preserved) from Chalcal 2 Stn DW 78, 23°41.30'S, 167°59.60'E, 233-

360 m, MNHN-Bry 19968.

Paratypes: a fertile colony (MNHN-Bry 19973) and a slide of dried branch fragments (MNHN-Bry 20029), both

from the same locality as the holotype.

Description. — Colony erect, rigid, dichotomously branching from a short base up to 4.0 mm diameter;

from the base arises a short stem -3-4 mm high and -2.5 mm diameter that branches outwards more than upwards,

dichotomising at intervals of 2. 8-4.0 mm; the branches between axils narrowing from 1.62 mm (near main stem)

to 0.79 mm (outermost) diameter. Zooids in 4-7 (rarely more) longitudinal series, ‘facing on the underside of the

branches, i.e., facing the substratum. Zooids 0.47-0.68 mm long and 0.32-0.40 mm wide (range of maximum

widths), the frontal shield lightly creased with short low ridges and depressions, imperforate, with up to 5 small

areolar pores placed irregularly on the shield between the margins and the centre; newly formed zooids clearly

delineated by thin raised lines along the interzooidal boundaries. Orifice about as wide (0.1 1-0.13 mm) as long,

with a broad V-shaped poster separated from the anter by condyles, each with a transverse furrow. No oral spines.

A median suboral avicularium present, the crossbar complete, the opesia a transverse slit, the rostral area almost an

equilateral triangle the rostral foramen near-circular; directed proximally. No other avicularia. Female zooids always

occurring at the extreme lateral margins of branches and facing laterally; ovicell huge, up to 0.56 mm wide and

long, deep-bodied, displacing the chamber of the next zooid in the series; the calcareous surface like that of the

frontal shield, the secondary calcification and outer epitheca continuous with those of the distal zooid; female

orifice typically a little wider (0.13-0.15 mm) than autozooidal orifices; ovicellular opening not closed by the

zooidal operculum. Abfrontal branch surface marked by thin lines (vibices) delineating zooidal boundaries.

Distribution. — New Caledonia: northern Norfolk Ridge, 233-360 m.

Remarks. — Macrocamera is established for the above new species from New Caledonia and for

Schizomavella pansa from Fiordland, New Zealand (243-253 m). Macrocamera pansa comb, nov., though infertile,

is clearly very similar to the type species, differing in zooidal details (more areolar pores, avicularium a little

further from the orifice) and in the pattern of branching. Though a new genus is clearly required, the family

placement poses some difficulty. The branching habit and huge ovicell are reminiscent of the Phorioppniidae (see

earlier in this paper), but all phorioppniids recognised here have zooids opening all around the stem, none has

avicularia, and all have at least some pseudopores in the frontal shield. The family Eminooeciidae was established

by Hayward and Thorpe (1988) for Eminooecia Hayward & Thorpe, 1988 and Isoschizoporella Rogick, 1960,

together embracing five Antarctic species. Morphological features cited as characteristic of the family included a

non-pseudoporous frontal shield, adventitious avicularia, and the association of the ovicell with "avicularian

polymorphs or inferred homologues". The latter structures appear not to be homologous, however. Although

avicularian chambers may develop concurrently alongside the ovicell in Isoschizoporella , the distal avicularium

chamber is not continuous with the ovicellular coelom. It is evident from their illustrations that the ovicell in

Eminooecia is indeed the ovicellular coelom - "Distally, the two layers of the ovicell are widely separated"

(Hayward & Thorpe, 1988: page 7, fig. 2); the avicularian chamber illustrated in their figure 5 (Ibid., page 13)

is separate from the ovicellular coelom enclosed between the endooecial and ectooecial layers. A further significant

distinction between Eminooecia and Isoschizoporella is that the ectooecium is wholly calcified in the former but

frontally membranous in the latter. In this regard, Isoschizoporella more closely resembles buffonellodids (see

above), even though they lack avicularia associated with the ovicell. Although Macrocamera lacks a capacious

ovicellular coelom, it is here tentatively allied with Eminooecia on the basis of similarities in the frontal shield,

the suboral avicularium, external ovicellular calcification, and erect habit.

The generic name is derived from the Greek makros , long, and kamara (Latin, camera ), a chamber.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

45

Superfamily EUTHYRISELLOIDEA Bassler, 1953

Family EUTHYR1SELLIDAE Bassler, 1953

Genus PSEUDOPLATYGLENA nov.

Type Species. — Pseudoplatyglena mirabilis sp. nov.

DIAGNOSIS. — Colony erect, cellariiform, with chitinous joints, the circular stems with a median abaxial non-

projecting keel. Zooidal frontal shield imperforate with conspicuous marginal areolae. Orifice with broad

subconvex poster and stout blunt condyles, with a broad suboral subhypostegal coelom protected by a stout

crescentic bar supported by buttresses. No oral spines. No avicularia. Ovicell unknown.

Pseudoplatyglena mirabilis sp. nov.

Figs 95-102

MATERIAL EXAMINED. — New Caledonia. Musorstom 4: stn DW 187, 65-120 m.

TYPES. — Holotype : a preserved 10-mm-long colony fragment with a single branch dichotomy, from

Musorstom 4 Stn DW 187, 19°08.30'S, 163°29.30’E, 65-120 m, MNHN-Bry 19962.

Paratype : a dry 6-mm-long fragment with part of a branch dichotomy, from the same locality as the holotype,

MNHN-Bry 20032.

DESCRIPTION. — Colony erect, cellariiform, jointed, with dichotomous branching. Branches circular, 0.56-

0.83 mm diameter, each nearly parallel-sided, with the proximal end abruptly tapering to a single chitinous joint;

proximal to the joint is a calcified zooidal chamber that appears to pertain to the zooid distal to the joint. Zooids

0.73-1.01 mm long and 0.24-0.16 mm wide, arranged more or less in 6 longitudinal series. Frontal shield

lepralioid, imperforate, the calcareous surface minutely granular/papillate frequently with a low continuous or

discontinuous median ridge; lateral margins sloping into an interzooidal furrow in which are variably sized areolae,

generally conspicuous. Abaxially there is a median longitudinal ridge; this is flush with the zooidal surface and

therefore non-projecting, but on either side is a conspicuous furrow with areolae and deep-set septular pores; the

cavity of each furrow is not longitudinally continuous, being crossed by transverse ridges associated with each

zooid. Zooidal orifice longer than wide, laterally indented where stout blunt condyles occur, separating the broad

subconvex poster from the high-arched anter. The proximal rim of the orifice comprises a stout crescentic bar that

is supported by 3-5 buttresses shortly radiating proximally. Beneath this arrangement is a relatively large

subhypostegal coelom. Oral spines lacking. No avicularia. Ovicells not present.

Distribution. — New Caledonia: north of the Belep Islands, 65-120 m.

Remarks. — Pseudoplatyglena is established here for a new genus whose affinities appear to lie with the

Euthyrisellidae. Superficially, P. mirabilis remarkably resembles some of the species of the Cretaceous genus

Platyglena Marsson. This genus was characterised by erect, mostly articulated, colonies; some species were

characterised by a suboral costal field superficially resembling the crescentic buttressed suboral bar in

Pseudoplatyglena mirabilis , and others had a similar median longitudinal ridge on the frontal shield.

Pseudoplatyglena differs significantly, however, in having a strictly lepralioid shield with no trace of costae.

Instead, there are features which would seem to ally the genus with some among the Euthyrisellidae, especially

Tropidozoum Harmer. These include the cellariiform habit, the shape of the orifice (cf. T. cellariiforme Harmer),

the investing coelom and epitheca around the chitinous joints, the abfrontal keel, and the subhypostegal coelom.

The abfrontal keel of Pseudoplatyglena differs insofar as it lacks the separate kenozooidal chambers on either side,

however; instead, the coeloms adjacent to the keel are merely lateral extensions of the frontal hypostegal coeloms.

46

D. P. GORDON & J.-L. D’HONDT

There is also no organic connection, via septula, between visceral and hypostegal coeloms as in several ot the

euthyrisellid genera.

Superfamily SIPHONICYTAROIDEA Harmer, 1957, superfam. nov.

Family SIPHONICYTARIDAE Harmer, 1957

Genus SIPHONICYTARA Busk, 1884

Type Species. — Siphonicytara serrulata Busk, 1884.

Siphonicytara armata sp. nov.

Figs 103-105

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 33, 675 m.

Types. — Holotype : a rooted colony from BiOCAL Stn DW 33, 23°09.7rS, 167°10.27'E, 675 m, MNHN-Bry

19989.

Paratope: part of a colony from the same locality, MNHN-Bry 20114.

DESCRIPTION. — Colony erect, comprising slightly curving, basally rooted, unbranched cylindrical stem to at

least 24 mm high and 1.73 mm diameter (not including zooidal peristomes); no discernible abfrontal side. Zooids

8-1 1 -serial in the thickest part of the stem (as at the apex), 0.96-1.16 mm long and -0.47 mm wide, the zooidal

surface granular. The arrangement of orifices, ascopores, areolae, and ridges are best appreciated by examining the

stem apex. Here the peristomial orifice (internal diameter 0.15-0.17 mm) is surrounded by a more or less triangular

arrangement of ridges and flanked by 3-4 deeply sunken areolar septula. Proximal to the peristome the ascopore,

with an areolar pore either side, is set in the middle of a subcircular to heart-shaped ridge; the centre-to-centre-

distance from ascopore to peristomial orifice 0.39-0.43 mm; distal half of the ascopore with a hooded surround.

Operculum D-shaped, with a small proximal projection at each lateral corner Avicularia single or paired both

distolateral and proximolateral to the orifice (thus 2-4 per zooid, and more or less aligned in longitudinal series up

the stem), circular in outline, 0.103 mm diameter, with a near-median cross-bar. Ancestrula 1.04 mm long,

subtubular, tapering proximally, lacking avicularia and a ridge between the orifice and ascopore.

Distribution. — New Caledonia: south of lie des Pins, 675 m.

Remarks. — Siphonicytara armata is distinguished from the other New Caledonian species described below by

the lack of a clear abfrontal surface and the disposition of the avicularia. It resembles the Philippine species

5. insolita (Canu & Bassler, 1929) in the type of avicularia, but those of S. insolita are directed laterally outward

instead of towards the peristome.

Siphonicytara excentrica sp. nov.

Fig. 106

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 38, 360 m.

Type. — Holotype: Biocal Stn DW 38, 22°59.74'S, 167°15.3rE, 360 m, MNHN-Bry 20122. No separate

paratypes.

DESCRIPTION. — Colony an erect cylindrical stem (incomplete in the unique holotype specimen) 1.2 mm

diameter (not including zooidal peristomes); an abfrontal side weakly discernible. Zooids 8-serial, 0.90-0.98 mm

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

47

long and -0.39 mm wide with a lightly textured non-granular surface. Peristomial orifice with an internal diameter

of 0.15-0.17 mm, mostly placed asymmetrically within the surrounding ridge system owing to the occurrence of a

relatively large avicularium on one side; at least one series of zooids (on the side interpreted to be abfrontal) has

symmetrical peristomes flanked by a pair of avicularia. Conspicuous areolar pores occur lateral (when one

avicularium is present) and proximolateral to the peristome. Ascopore set in a transversely narrow area bounded by

a ridge that is continuous or lacking along one side; the distal side of the ascopore slightly hooded; the centre-to-

centre distance from ascopore to peristomial orifice 0.45-0.57 mm. Avicularia suboval, 0.20-0.28 mm long, on

one or both sides of the peristome, directed to the peristomial orifice and facing obliquely laterally, with a cross-bar

delimiting a small opesia from a relatively large palate.

Distribution. — New Caledonia: south of lie des Pins, 360 m.

REMARKS. — Siphonicytara excentrica is readily distinguished by the occurrence of asymmetrically placed

peristomes on a majority of the zooids.

Siphonicytara glabra sp. nov.

Figs 107-108

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 70, 965 m.

Types. — Holotype: a branching fragment from Biocal Stn DW 70, 23°24.70'S, 167°53.65'E, 965 m,

MNHN-Bry 20124.

Paratype : from same locality as holotype, MNHN-Bry 20117.

DESCRIPTION. — Colony erect, bifurcating at intervals (5.5 mm in the limited material available, which

allows for only 1 measurement), the stem and branches somewhat lensoid in cross section, 0.83-0.98 mm

diameter, with a distinct abfrontal surface that is slightly flattened. Zooids 4-serial, 0.81-0.94 mm long and -0.47-

0.53 mm wide, the surface glabrous, becoming finely granular in the oldest zooids. Peristomial orifice (internal

diameter 0. 13-0. 16 mm) bounded by ridges distally and proximally but not laterally; with lateral areolae along each

side proximally, the most distal ones often replaced by 1-2 avicularia. Ascopore with a small areolar pore on 1 or

both sides, together set between distal and proximal ridges and sometimes a lateral one; the centre-to-centre distance

from ascopore to peristomial orifice 0.35-0.41 mm; distal side of ascopore slightly hooded. Avicularia suboval,

relatively small, 0.094-0.17 mm long and initially subcolumnar, on 1 or both sides of the orifice (not attached to

the peristome) or lacking, directed more or less distally, facing proximofrontally, with a complete crossbar.

Abfrontally, there is a sinuously zigzag median longitudinal ridge to which all frontal transverse ridges connect. In

the adjacent area between each peristome and the median ridge is an avicularium, similar to those on the frontal

side but facing more distinctly proximally and with a more pointed apex.

REMARKS. — Siphonicytara glabra resembles S. vittata sp. nov. in colony form and in the arrangement of

zooids. It differs in having a mostly glabrous zooidal surface, more numerous frontal avicularia, and more

proximally directed abfrontal avicularia.

Siphonicytara vittata sp. nov.

Figs 109-111

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 08, 435 m. — Stn DW 33, 675 m. — Stn DW 36,

650 m. — Stn DW 44, 440 m. — Stn DW 51, 700 m.

Chalcal 2: stn DW 76, 470 m.

TYPES. — Holotype : a preserved rooted colony from BlOCAL Stn DW 44, 22°47.30'S, 167°14.30'E, 440 m,

MNHN-Bry 19990.

48

D. P. GORDON & J.-L. D’HONDT

Paratypes : MNHN-Bry 20119, MNHN-Bry 20123, and MNHN-Bry 20128, all from Biocal Stn DW 08,

20°34.35'S, 166°53.90'E, 435 m; MNHN-Bry 20113, from the same locality as the holotype; and MNHN-Bry

20129, from Chalcal 2 Stn DW 76, 23°40.50’S, 167°45.20,E, 470 m.

Description. — Colony erect, bifurcating at intervals of 2. 5-9. 6 mm, the stems 0.50-1.32 mm diameter (not

including zooidal peristomes), with a distinct abfrontal surface that is slightly flattened; basally, the colony is

anchored by rootlets that issue from pores on the abfrontal surface only. Zooids mostly 4-serial, in longitudinal

series, becoming 6-serial in the thickest stems, 0.68-0.98 mm long and -0.30-0. 49 mm wide, the calcareous

surface textured with light irregular ridging. Frontally, the peristomial orifices (internal diameter 0.10-0.15 mm)

and ascopores occur between parallel sinuous ridges that are transversely continuous and typically without

connecting longitudinal ridges; the arrangement of peristomes is such that they occur in regular series both

longitudinally and obliquely ; peristomes of the zooids flanking the abfrontal surface are longer and curve round to

face more or less frontally. Conspicuous areolar pores occur in a collective furrow on each side proximolateral to

every peristome; although slightly offset, the vitta-like furrows off all the zooids in a vertical series are more or

less aligned. Ascopore circular, not hooded, with an areolar pore on each side, the centre-to-centre distance from

ascopore to peristomial orifice 0.31-0.42 mm. Avicularia very rare frontally. Abfrontally, there is an irregular

zigzag ridge to which the frontal transverse ridges join; between that ridge and each adjacent peristome is a small

suboval avicularium -1.23 mm long, slightly elevated, with the rostrum directed laterally outwards, the palate

facing obliquely laterally or proximolaterally towards the stem axis.

DISTRIBUTION. — New Caledonia: near Lifou, and south of lie des Pins, 435-700 m.

Remarks. — Siphonicytara vittata resembles S. formosa Harmer, 1957, which, however, can achieve a larger

size, has longitudinal ridges frontally (i.e., ridges completely surround the peristomes and ascopores), and

proportionately larger abfrontal avicularia.

Siphonicytara granulosa sp. nov.

Figs 112-113

Material EXAMINED. — New Caledonia. Biogeocal: stn CP 232, 760-790 m.

Type. — Holotype : a stem fragment 10 mm long, from BIOGEOCAL Stn CP 232, 21°33.81'S, 166°27.07'E,

760-790 m, MNHN-Bry 20118. No separate paratypes.

DESCRIPTION. — Colony erect, cylindrical and slightly curving, the unique holotype stem 0.73-0.94 mm

diameter (not including peristomes); no discernible abfrontal surface. Zooids 1.02-1.28 mm long and -0.50-

0.62 mm wide, the calcareous surface granular; arranged in 5 longitudinal series. Peristomes (internal diameter

0.15-0.17 mm) proportionately longer than in the other New Caledonian species, in a subhexagonal area bounded

by low ridges, with 4-6 areolar cavities proximolaterally. Ascopore completely bounded by a ridge, not hooded,

with 1-2 areolar pores on each side, the centre-to-centre- distance from ascopore to peristomial orifice 0.47-

0.64 mm. Avicularia sporadic, generally occurring singly on a ridge adjacent to a peristome, -0.12 mm long, the

rostrum directed towards the peristome, the palate obliquely proximofrontally.

Distribution. — New Caledonia: off the coast near the town of Thio, 760-790 m.

Remarks. — Siphonicytara granulosa is distinguished from S. armata, which it most resembles, by its more

slender colony form, proportionately longer peristomes, and fewer avicularia.

Siphonicytara mosaica sp. nov.

Figs 114-117

Material EXAMINED. — New Caledonia. Biogeocal: stn CP 260, 1820-1980 m. — Stn CP 265, 1760-1870 m

Musorstom 6: stn CP 465, 480 m.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

49

Types. — Holotype : branching fragment from Biogeocal Stn CP 265, 21°04.09'S, 167°00.40'E, 1760-

1870 m, MNHN-Bry 20127.

Paratypes : MNHN-Bry 20115, from Biogeocal Stn CP 260, 21°00.00'S, 167°58.34’E, 1820-1980 m;

MNHN-Bry 20126, from MUSORSTOM 6 Stn CP 465, 21°03.55’S, 167°32.25’E, 480 m.

DESCRIPTION. — Colony erect, dichotomously branching, the stems delicate, only 0.37-0.49 mm diameter.

Zooids 2-4-serial, 1.00-1.24 mm long and -0.22-0.30 mm wide, the peristomes facing laterally or frontally such

that there is an abfrontal surface characterised by the absence of peristomes but no other features; zooidal surface

distinctively sculptured by a mosaic of polygonal ridges that continue onto the peristome. Stems regularly

annulate, the transverse ridges separating peristomes from ascopores being continuous; no longitudinal ridges.

Peristomes (internal diameter 0.11-0.13 mm) somewhat projecting, flanked on each side by a linear row of

4-5 areolae. Ascopore circular, not hooded, set immediately proximal to a transverse ridge and with a relatively

extensive zooidal surface proximal to it; centre-to-centre distance from ascopore to peristomial orifice 0.37-

0.43 mm. No avicularia.

Distribution. — New Caledonia: South Loyalty Basin, 480-1980 m.

REMARKS. — This is the most delicate of all known Siphonicytara species. In its annular transverse ridges it

resembles S. symetrica David & Pouyet, 1986 from south of Madagascar, which differs, however, in its larger size

and in having avicularia and a smooth zooidal surface.

Superfamily MAMILLOPOROIDEA Canu & Bassler, 1927

Family ASCOSIIDAE Jullien, 1883*

Diagnosis. — Colony discoidal to conical, the basal surface concave. Zooids recumbent to suberect, radiating

from the central ancestrular region, the frontal shield imperforate. Orifice with a broad poster delineated by a pair of

condyles. No oral spines. Avicularia, if present, typically adjacent to the orifice; the crossbar complete. Ovicell, if

present, not unusually large, imperforate, closed by the zooidal operculum. Basal surface of colony composed of

the basal or proximobasal parts of autozooids.

REMARKS. — Ascosia Jullien, 1883 is usually included in the family Mamilloporidae Canu & Bassler. The

type genus of that family, Mamillopora Smitt, 1873, while also having discoidal or domed colonies, differs in its

minutely perforated enlarged ovicells and dimorphic orifices. JULLlEN's (1883: page 526, as Ascosidae)

introduction of the family Ascosiidae appears to have been completely overlooked by later authors, otherwise it

would almost certainly have continued in use. For example, SlLEN (1947) argued that the related genus Fedora

Jullien, 1882 should be excluded from the Mamilloporidae but did not suggest an alternative family. Similarly,

LAGAAIJ (1963) declined to suggest a family for Fedora. Mamillopora , by contrast with Ascosia , has enlarged,

densely perforated ovicells. In our opinion, the Ascosiidae should be recognised as a separate family of the

superfamily Mamilloporoidea, introduced by D'HONDT (1985). Other putative mamilloporid genera should

possibly be included in the Ascosiidae but, until there has been a general restudy and redescription of all related

genera, we prefer not to comment further.

Genus FEDORA Jullien, 1883

Type Species. — Fedora edwardsi Jullien, 1883.

*This work, though with a publication date of 1882, was not published until 1883. According to Zoological Record

for that year (page Moll. 97), Jullien’s paper was “Read at the Meeting of 26th December, 1881, but not yet received in

print at Berlin before August, 1883, and the plates have not yet been seen by the Recorder [Prof. E. von Martens]".

50

D. F. GORDON & J.-L. D’HONDT

Fedora platydiscus sp. nov.

Figs 118-120

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn DR 117, 97-92 m.

Type. — Holotype : unique infertile colony from MUSORSTOM 3 Stn DR 117, 12°31.3'N, 120°39.5'E, 97-

92 m, MNHN-Bry 20035.

Description. — Colony discoidal, flattened, 3.60 mm maximum diameter. Zooids recumbent, radiating

outwards from the central ancestrular region; 0.45-0.72 mm long and 0.20-0.47 mm wide, the frontal shield

densely tubercular, the tubercles themselves with minute papillae. Orifice elongate, almost twice as long (0.1 1-

0.17 mm) as wide, the broad rounded poster delimited from the anter by a pair of small angled condyles; orifice

laterally flanked by tubercles. Articulated oral spines absent. Avicularia absent. Ovicells not present. Ancestrular

opesia (or broken frontal shield?) 0.18 mm long, surrounded by 8 periancestrular zooids. Basal surface of colony

somewhat nodular, with convexities indicating the basal surfaces of individual zooids. Zooidal budding taking place

from a distobasal pore-chamber on each zooid.

Distribution. — Philippines: Mindoro Strait, 92-97 m.

Remarks. — We are not certain about the generic placement of the present species, which may represent a

new genus. The two previously known living species of Fedora [Myriozoum ovum Smitt, 1873 is unlikely to be

conspecific; cf. Sil£n 1947] have more or less conical colonies, but Lagaaij (1963) noted some variation in

F. nodosa Silen from ovoid and spindle shapes to hemispherical and that colonies started life as encrustations on

small objects. Fedora is the nearest genus in terms of overall morphological characters, especially in the

tuberculate frontal shield, lack of ovicells, and the distobasal pore-chambers. Silen (1947) called the latter ‘special

chambers’ and speculated on their purpose. LAGAAIJ (1963), observing an apparent rootlet emerging from one

chamber, concluded that they give rise to anchoring rhizoids. This is possible, but they must soon become

obliterated by the new autozooids that develop around them. In the flattened colony form, F. platydiscus is more

reminiscent of Ascosia , but in that genus the zooids are suberect, with ovicells and proximobasal budding, and lack

the distobasal pore-chambers.

The name of the new species is derived from the Greek platys , flat, and diskos , plate. It should be noted in

passing that the spelling of the type species of Fedora is given twice by JULLIEN (1883) as Edwarsi (sic), above

the formal description and in the table on page 527, but as Edwardsi in the plate captions on page 529. We

conclude, therefore, that Edwarsi is a lapsus calami for edwardsi , presumably named for Henri MILNE EDWARDS.

Family CLEIDOCHASMATIDAE Cheetham & Sandberg, 1964

Remarks. — We here include this family in the superfamily Mamilloporoidea on the basis of the zooidal

frontal shield and the shape of the orifice. Although the ovicell, when present, in mamilloporids and ascosiids is

closed by the zooidal operculum and the colony form is largely different, a relationship with cleidochasmatids is

suggested by the genus Fedorella Silen, 1947. This monotypic genus has a similar colony form to Fedora but a

typically cleidochasmatid ovicell (i.e., minutely porous and tuberculate and not closed by the zooidal operculum).

[Puzzlingly, COOK and Lagaaij (1976: page 321) listed Fedorella among genera with ‘frontal’ zooidal astogeny

(i.e., like Conescharellina ), but Sil£n's (1947) description and illustrations indicate otherwise.) D’HONDT (1981)

also noted a similarity between the orifices in Characodoma Maplestone, 1900 (a senior synonym of Cleidochasma

Harmer, 1957) and Fedora. As pointed out by COOK and BOCK (1996), Characodoma can be erect and conical from

basally encrusting colonies (see also Harmer, 1957).

Soule et al. (1991) revised Cleidochasma sensu lato , introducing a number of new genera of Cleido-

chasmatidae. Several of these have phidoloporid orifices (with a beaded distal arch) and ovicells (with a labellum),

however, and should be excluded from the Cleidochasmatidae, as suggested by GORDON (1993b).

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

51

Genus CHARACODOMA Maplestone, 1900

TYPE Species. — Schizoporella excubans Waters, 1881, by synonymy with Characodoma halli Maplestone,

1900 (see COOK & Bock, 1996).

Characodoma areolata (Canu & Bassler, 1929)

Figs 121-122

Gemelliporella areolata Canu & Bassler, 1929: 309, pi. 34, figs 5-6.

Cleidochasma areolatum - Harmer, 1957: 1046, pi. 71, figs 19-21.

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn DR 117, 97-92 m.

DISTRIBUTION. — Philippines, 42-969 m; Indonesia (Makassar Strait), 59-320 m.

Remarks. — Characodoma areolata is represented in the collection by a single specimen. It is a compressed

pisiform colony that appears to have encrusted a tiny piece of substratum that is now largely enclosed by it. It

closely matches Canu and BaSSLER's (1929: pi. 34, fig. 6) illustration, which shows occasional small flattened

kenozooidal chambers (possibly loci of frontal budding) associated with some zooids, also present in the

MUSORSTOM specimen. The zooids (0.50-0.70 mm long and 0.37-0.51 mm wide) and orifice (0.16-0.19 mm

long) overlap in range with those of CANU and BaSSLER's specimens or are slightly larger; the avicularium is

0.20-0.25 mm long, rounded at both ends, and with a thin crossbar that may frequently be broken, giving the

appearance of mandibular pivots. No zooid had two avicularia on the same side of the orifice, but one zooid has a

small avicularium on the opposite side of the orifice from the large one (as seems also indicated in Canu &

BaSSLER’s pi. 34, fig. 5).

Characodoma biavicularia (Canu & Bassler, 1929)

Figs 124-126

Gemellipora biavicularia Canu & Bassler, 1929: 312. pi. 34, fig. 7.

Material EXAMINED. — Philippines. Musorstom 3: stn DR 117, 97-92 m. Also USNM 8067, holotype of

Gemellipora biavicularia Canu & Bassler, 1929, "Albatross" Stn 5179.

DESCRIPTION. — Colony encrusting to erect, the latter up to 6.2 mm long, circular to lensoid or almost

bilamellar in cross section, 5-6 zooids across (1.98 mm) at the widest part. Zooids 0.56-0.75 mm long, 0.33-

0.58 mm wide. Frontal shield evenly granular-tubercular, with up to 10 or more areolar pores along each margin.

Orifice distinctly cleidridiate, 0.16-0.19 mm long, tapering proximally. Avicularia of two sizes, the larger lateral-

oral, 0.15-0.19 mm long, directed proximally towards the centre of the trontal shield; the rostrum almost an

equilateral triangle, as is the proximal part of the mandible that lies across the rostrum, the remainder of the

mandible being long and setiform; its total length 0.39 mm. The smaller avicularium suboral, not present on

every zooid and may be wanting over an extensive area of the colony; orientated transversely, the tiny semicircular

mandible 0.04 mm long. Ovicell globular, densely and evenly perforated.

Distribution. — Philippines, 38-104 m.

REMARKS. — The MUSORSTOM material comprises one encrusting and three erect specimens. One of the latter

(uncoated) is well preserved, with intact opercula and mandibles. The erect specimens closely resemble Canu and

BaSSLER's (1929) sole illustration, showing two sizes of avicularia, the larger one lateral-oral, the smaller one

suboral. Zooidal dimensions in the present material are larger than cited by Canu and BASSLER, but the species

appears very variable. In the present material the emplacement of the larger of the two avicularia varies from

laterally adjacent to the orifice to a position subjacent to the sinus. The type specimen from the Smithsonian

52

D. P. GORDON & J.-L. D’HONDT

Institution appears superficially different in overall zooidal morphology but it is likely this arises from the degree

of secondary calcification. Harmer (1957) recorded putative C. biaviculciria, with delicate erect rooted colonies, in

" Siboga " samples, but his illustrations indicate a different species in our opinion. Contrary to Canu and

Bassler's description he shows the suboral avicularium as the larger of the two and with an acute mandible.

Characodoma glabra sp. nov.

Fig. 123

Material EXAMINED. — Philippines. Musorstom 3: stn DR 117, 97-92 m.

Type. — Holotype : colony encrusting a small piece of shell from MUSORSTOM 3 Stn DR 117, 12°31.3'N,

120°39.5'E, 97-92 m, MNHN-Bry 20050. No separate paratypes.

Description. — Colony encrusting. Zooids 0.37-0.56 mm long and 0.20-0.64 mm wide, each separated from

adjacent zooids by a thin marginal groove; frontal shield smooth with sparse granulations and small sparse

marginal areolar pores. Orifice 0.1 1-0. 13 mm long, the deep rounded poster almost one third the orificial length,

separated from the larger anter by lateral indentations with tiny condylar points. Avicularia 0.066-0.075 mm long,

paired or, more usually, single, lateral to the orifice and directed proximofrontally, the crossbar very delicate, the

rostrum acute. Ovicell relatively small (0.17-0.21 mm wide), the endooecium relatively thick, its outer surface

with small spiky tubercles and small sparse perforations; ectooecium membranous.

Distribution. — Philippines, Mindoro Strait, 92-97 m.

Remarks. — This species is distinguished by the mostly smooth frontal shield, the size and arrangement of

the avicularia, and the relatively small, coarse-surfaced ovicell.

Characodoma parva sp. nov.

Fig. 127

Material EXAMINED. — Philippines. Musorstom 3: stn DR 1 17, 97-92 m.

Type. — The sole colony from Musorstom 3 Stn DR 117, 12°31.3'N, 120°39.5'E, 97-92 m, MNHN-Bry

2005 1 . No separate paratypes.

DESCRIPTION. — Colony small, compressed-pisiform in shape. Zooids very small, 0.26-0.34 mm long and

0.28-0.34 mm wide, the frontal shield coarsely surfaced, with blunt tubercles and relatively large areolae. Orificial

poster one-fourth the total orificial length of 0.09-0.1 1 mm, widely V-shaped, delimited from the larger anter by

small condyles. Avicularia paired or single, comprising a suboral one set transversely, the rostral and opesial rims

rounded; and frequently and additional one adjacent to the orifice, the acute rostrum directed obliquely proximally

toward the lateral margin; the crossbars in both kinds of avicularia relatively stout. Ovicell 0.16-0.21 mm wide,

the endooecium with the numerous small tubercles and pores evenly distributed, the ectooecium membranous.

Distribution. — Philippines, Mindoro Strait, 92-97 m.

Remarks. — This species is distinguished by its small colonial and zooidal dimensions and the shape and

disposition of the avicularia.

Characodoma sp.

Fig. 128

Material examined. — Philippines. Musorstom 3: stn CP 102, 192 m.

Source : MNHN. Paris

BRYOZOA ASCOPHOR1NA FROM NEW CALEDONIA

53

Description. — Colony fragment encrusting. Zooids 0.47-0.62 mm long and 0.41-0.57 mm wide, the frontal

shield somewhat irregularly surfaced in the proximal half where there are small scattered areolar pores, and granular

all over. Orifice a little longer than wide, ovoid in shape, with only the small condylar points differentiating the

anter from the broad poster. No avicularia, but a pair of relatively areolar cavities either side of the orifice can

appear superficially like tiny avicularia under transmitted light. Ovicells not seen.

Distribution. — Philippines, 192 m.

REMARKS. — Only a tiny infertile fragment of this species occurred in the collection. Despite the broad

orificial poster, it appears to be a species of Characodoma.

Genus YRBOZOON Gordon, 1989b

Type Species. — Yrbozoon ringens Gordon, 1989b.

Yrbozoon ringens Gordon, 1989b

Figs 129-131

MATERIAL EXAMINED. — New Caledonia. BlOCAL: stn DW 66, 515 m.

Distribution. — Norfolk Ridge from New Zealand (South Maria Ridge, 1024-1049 m) to New Caledonia

(northern Norfolk Ridge, 515 m).

REMARKS. — Four small colony fragments, two ovicellate or partly so, represent only the second record of

this genus and species. They are identical to the sole New Zealand specimen except in the number of oral spines or

tubercles, being only two instead of 4-5.

GORDON (1989b) included Yrbozoon in the Celleporidae, in which it is very much on the fringe. Having now

seen several cleidochasmatids in the MUSORSTOM samples it is apparent that the ovicell of Yrbozoon is very

similar to that of Cleidochasma, comprising a reticulate (but not perforate, however) endooecium and membranous

ectooecium. The presence of oral spines in Yrbozoon is exceptional in the superfamily Mamilloporoidea, which

may indicate that the affinities of Yrbozoon lie elsewhere; it may be noted that some of the spines in Y. ringens

from New Zealand are merely long tubercles and the others appear to be non-articulated. Thus, even including

Yrbozoon in the Mamilloporoidea need not preclude ‘absence of articulated oral spines’ as a superfamily

characteristic.

Superfamily CELLEPOROIDEA Johnston, 1838

Family CELLEPORIDAE Johnston, 1838

Genus BUFFONELLARIA Canu & Bassler, 1917

TYPE Species. — Hippothoa divergens Smitt, 1873.

Buffonellaria erecta sp. nov.

Figs 132-133

Material EXAMINED. — New Caledonia. BIOCAL: stn DW 66, 515 m. — Stn CP 109, 495 m.

54

D. P. GORDON & J.-L. D’HONDT

Type. — Holotype : unique colony, 5.5 mm long, from Biocal Stn DW 66, 24°55.43'S, 168°21.67'E,

515 m, MNHN-Bry 20040. No separate paratypes.

Description. — Colony erect, rigid, branching shortly in 1 plane, the stem subcylindrical with zooids facing

all around, or partly encrusting the stem of an erect bryozoan and erect parts arising from this. Zooids somewhat

porcellanous, 0.50-0.74 mm long and 0.32-0.47 mm wide, the frontal shield imperforate, with tiny sparse areolar

pores along the margins. Orifice slightly sunken, wider (0.14-0.15 mm) than long, with a disproportionately small

and narrow U-shaped sinus with flattened adjacent orificial shoulders. A suboral avicularium present to 1 side of the

sinus, set transversely, with a crossbar, the acute rostrum directed laterofrontally. Ovicell initially recumbent,

becoming subimmersed, the tabulate area of endooecium large with marginal pores and short associated furrows, all

but the tabula becoming encroached upon by secondary calcification.

DISTRIBUTION. — New Caledonia, northern Norfolk Ridge, 495-515 m.

REMARKS. — This species is uniquely distinguished among living species of the genus by its erect habit; the

combination of small sinus, asymmetrical avicularium, and endooecial tabular pores are also distinctive. The

encrusting Philippine species B. indistincta Canu & Bassler, 1929 has a similar ovicell and suboral avicularium,

but has a broader, V-shaped sinus and 1-2 small lateral-oral avicularia additional to the transverse one.

Buffonellaria regenerata Powell, 1967

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 38, 360 m. — Stn DW 66, 515 m.

Description. — Colony thinly encrusting. Zooids hyaline, 0.38-0.70 mm long and 0.26-0.37 mm wide,

somewhat elongate-rectangular in outline, the frontal shield smooth, imperforate, with no marginal areolar pores,

but a small pore proximal to each avicularium. Orifice high-arched, longer (0.084-0.095 mm) than wide, the

median sinus very shallow and rounded, flanked by straight shoulders with a small condyle angled across each

corner. Avicularia paired, lateral-oral, the crossbar thin, the rostrum rounded, directed obliquely laterofrontally.

Ovicell recumbent, the endooecium smooth, exposed in the distal half of the ovicell, the proximal face of the

ovicell comprising a transverse band distal to the opening.

Distribution. — New Caledonia, northern Norfolk Ridge, 360-515 m. New Zealand, Kermadec Ridge and

Three Kings Islands to Cook Strait and Kahurangi Shoals, 10-240 m.

Remarks. — The present material is very similar to colonies previously described from the New Zealand

region. There are minor differences in the depth of the orificial sinus and degree of secondary calcification around

the ovicell, and a suboral ridge is normally present in the New Zealand material. Nevertheless, these characters are

subject to variation and we conclude that the New Caledonian specimens are not sufficiently distinct to warrant

subspecific recognition. Several zooids in the New Caledonian material have bore-holes in the frontal shield. These

are 0.037 mm diameter; the predator species is not known.

Genus OSTHIMOSIA Jullien, 1888

Type Species. — Cellepora eatonensis Busk, 1881.

Osthimosia sp.

Figs 134-136

Material EXAMINED. — New Caledonia. Musorstom 4: stn DW 187, 65-120 m

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

55

Description (Based on juvenile colonies only). — Colony pisiform, the zooids suberect, squat and barrel¬

shaped, about 0.28 mm long (0.47 mm including the peristome) and 0.26 mm wide. Frontal shield smooth, with a

few areolar pores along each margin. Primary orifice 0.094-0.113 mm wide, orbicular with a small rounded-V-

shaped sinus proximally; distinct condyles not present. Peristome encircling the primary orifice, rising to an

avicularian column on either side; each avicularium ovoid in shape, with a thin crossbar and a small subacute

triangular rostrum; the avicularium surface facing proximolaterally toward the orifice. No other avicularia. Ovicells

not present. Ancestrula initially tatiform, subsequently developing a peristome around the former opesia.

Distribution. — New Caledonia, northeast of the Belep Islands, 65-120 m.

REMARKS. — Two tiny infertile colonies, one ancestrulate, occurred on stem segments of the quadricellariid

bryozoan Nellia tenella (Lamarck, 1816). In zooidal characters they resemble specimens from the Kermadec Ridge

and New Zealand continental shelf attributed to Osthimosia bicornis (Busk, 1881) (GORDON, 1984, 1989a), but

which, because of orificial characters, may represent an undescribed species.

Genus LAGENIPORA Hincks, 1877

Type Species. — Celleporella lepralioides Norman, 1868.

Lagenipora sp.

Fig. 137

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 46, 775 m.

DESCRIPTION (Based on a juvenile colony only). — Colony encrusting, the zooids radiating somewhat linearly

from the ancestrular region. Zooids recumbent, -0.38 mm long (including peristome) and 0.25 mm wide, the

frontal shield smooth, with a few areolar pores around the margin. Primary orifice 0.06 mm wide, with a broad

rounded sinus and non-projecting simple condylar surfaces angled in the corners. Peristome long and tubular,

incorporating a median suboral avicularium tube opening at the peristomial rim which flares outwards.

1-2 additional small adventitious avicularia occur near the lateral margins of the frontal shield. Vicarious avicularia

and ovicells not seen.

Distribution. — New Caledonia, south of lie des Pins, 775 m.

REMARKS. — It is not possible to ascribe the single tiny colony (epizoic on a branch of a buguloidean

bryozoan) to a described species. It corresponds to none of the New Zealand region species, which all lack similar

adventitious avicularia on the frontal shield.

Genus GALEOPSIS Jullien in JULLIEN & Calvet, 1903

Type Species. — Galeopsis rabidus Jullien in Jullien & Calvet, 1903.

Galeopsis mimicus Gordon, 1989

Figs 138-140

Galeopsis mimicus Gordon, 1989a: 68, pi. 3 D-E.

Material EXAMINED. — New Zealand. New Zealand Oceanographic Institute, holotype H-513, paratype P-729.

New Caledonia. Biocal: stn DW 33, 675 m. — Stn DW 36, 650 m. — Stn CP 67, 500 m. — Stn CP 75, 825 m.

Chalcal 2: stn DW 76, 470 m.

Smib 4: stn DW 37, 540 m.

56

D F. GORDON & J.-L. D’HONDT

Distribution. — New Zealand, off the Westland coast, 297-520 m. New Caledonia, South Loyalty Basin to

northern Norfolk Ridge, 470-825 m.

REMARKS. — The New Caledonian material gives additional information about this species. Whereas in New

Zealand colonies occurred only on aboral spines of Spatangus multispinus Mortensen, the present colonies occur

on axes of small gorgonians. The shape of the orifice and sinus are identical throughout the range, as is that of the

ovicell, which has a characteristically long proximal face with a short labellum. A colony from BlOCAL

Stn CP 75 has a relatively large basal disc for this species (2.96 mm maximum diameter) and a colony from

Chalcal 2 Stn DW 76 has rare spatulate avicularia - these are 0.16-0.19 mm long, with a small, very short

opesia and a relatively broad round rostrum with an extensive palatal shelf.

Galeopsis pentagonus (d'Orbigny, 1842)

Vincularia pentagona d'Orbigny, 1842, pi. 10, figs 4-6; 1847: 21.

Spiroporina pentagona - BROWN, 1952: 213, figs 148, 149 ( cum syn.).

Galeopsis pentagonus - Gordon, 1984: 116, pi. 46, A-C (cum syn.); 1985: 178, fig. 29; 1989a: 69. — MOYANO, 1985:

90, pi. 1, figs 1-6 ( cum syn.).

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 66, 515 m.

Musorstom 6: stn CP 419, 283 m. — Stn DW 421, 245 m.

Smib 4: stn DW 38,510 m.

DISTRIBUTION. — New Caledonia, 245-515 m. New Zealand, widespread from 40-549 m (also Lower

Oligocene to Pleistocene). Tasmania (Late Miocene to present day). Juan Fernandez Island. Magellanic South

America, Falkland Islands (lies Malouines).

Remarks. — The New Caledonian colonies from Musorstom 6 Stn CP 419 (20°41.65’S) constitute the

northernmost record of this long-lived austral species.

Galeopsis lageniporoides sp. nov.

Figs 141-142

MATERIAL EXAMINED. — New Caledonia. Musorstom 6: stn DW 489, 700 m.

Types. — Holotype : a coated colony, now in two fragments, from MUSORSTOM 6 Stn DW 489, 20°48.87'S,

167°05.86'E, 700 m, MNHN-Bry 20041.

Paratype : an unnumbered colony in a separate well of the holotype slide, from the same locality as the

holotype.

Description. — Colony encrusting, linear, 2-3 zooids across and branching. Zooids 0.51-0.62 mm long and

0.26-0.28 mm wide, the frontal shield smooth and imperforate with small sparse areolar pores. Orifice suborbicular

with a broad shallow sinus delimited by blunt rounded condyles. Lateral-oral avicularia small, ovoid, with a

rounded rostrum directed laterally outwards or proximolaterally, the avicularia borne at the corners of a high

peristome that forms a bridge across the orifice leaving a large spiraminal opening; the proximofrontal face of the

peristomial bridge fairly broad. 1-2 additional avicularia are borne adventitiously near the lateral margins. Ovicell

recumbent, with a crescentic or triangular exposure of endooecium with tiny distal pores and associated grooves;

the proximal face of the ovicell fairly broad, with a labellum on the lower edge.

Distribution. — New Caledonia, north of Lifou, 700 m.

Remarks. — Neither of the two colonies in the collection is attached to a substratum, which appears to have

been a hydroid stem. The larger of the colonies, now in two pieces, was 5.15 mm long and 0.54-0.64 mm wide.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

57

Superficially, this species resembles a species of Lagenipora because of the high aviculiferous peristome. No

Lagenipora species, however, has an ovicellular labellum or a peristomial spiramen.

Genus RICHBUNEA nov.

Diagnosis. — Colony erect, branching, the zooids facing on one side. Primary orifice with broad sinus and

small condyles, concealed by a tall peristome with an irregular rim in which is set 1-2 columnar avicularia. Large

spatulate avicularia may occur frontally and abfrontally. Ovicells recumbent, becoming somewhat immersed in

secondary calcification, with a broad tabulate area.

Type Species. — Osthimosia incomposita Gordon, 1984.

REMARKS. — A new genus is established here for two species of erect celleporids that have long, irregularly

rimmed, aviculiferous peristomes and broadly tabulate ovicells. The type species from the Kermadec Ridge,

initially included in Osthimosia , was later transferred to Buchneria Harmer, 1957 (Gordon, 1985), with the

suggestion that it might represent a new genus. It has since been established that the type species of Buchneria ,

with an umbonuloid frontal shield, differs in several significant respects from O. incomposita (GORDON, 1989c),

and a new genus appears justified. Comparison is invited with Spigaleos Hayward, 1992, which also forms erect

branching colonies with the zooids facing on one side, but the type and only species, S. horneroides Waters, has

peristomial orifices flush with the colony surface, and median, sessile avicularia. Richbunea is an anagram of

Buchneria. At present, the new genus is known only from the Kermadec and Norfolk Ridges at depths of 135-

500 m.

Richbunea gracilis sp. nov.

Figs 143-144

MATERIAL EXAMINED. — New Caledonia. Biocal: stn CP 67, 500 m.

BiOGEOCAL: stn DW 307, 470-480 m.

TYPES. — Holotype : a single colony from BlOCAL Stn CP 67, 24°55.44'S, 168°21.55'E, 500 m, MNHN Bry-

20039.

DESCRIPTION. — Colony erect, slender, sparsely branching, the zooids in two longitudinal zigzagging series,

with a pair of orifices on one side (one near the branch axis, the other, slightly beyond it, projecting outward

laterally) alternating with a similar pair on the other side. Branch width between peristomes 0.46-0.60 mm. Zooids

c. 0.67-0.71 mm long and 0.23 mm wide, the surface smooth, with 2-5 small areolar pores along each margin.

Primary orifice suborbicular, with a broad shallow sinus separated from the anter by a small pair of rounded

condyles, concealed from frontal view by a tall peristome. Peristomial rim irregular, with several processes and a

proximal pseudosinus that descends as a sinus groove down the inside of the peristome. A small, columnar

avicularium projecting from the peristomial rim next to the pseudosinus, with a complete crossbar and acute

triangular rostrum directed frontally. No other avicularia. Ovicell prominent, recumbent, with a large frontal

endooecial tabula. Abfrontal surface smooth, with a thin median line, indicating zooidal lateral boundaries,

zigzagging along the branch axis. Zooids near the colony base have occluded orifices.

DISTRIBUTION. — New Caledonia between Ouvea and Lifou, and northern Norfolk Ridge, 470-500 m.

REMARKS. — Richbunea gracilis superficially resembles a species of Phidoloporidae, especially in its erect

habit and lodictyum- like peristomes, but it lacks a beaded primary orifice and the ovicell is distinctively celleporid.

It differs from the type species, R. incomposita comb, nov., in its slender colony form with fewer zooidal rows,

single columnar avicularium, and absence of spatulate avicularia.

58

D. P. GORDON & J.-L. D’HONDT

Family PHIDOLOPORIDAE Gabb & Horn, 1862

Genus LIFUELLA nov.

Type Species. — Lepralia multidentata Thornely, 1905.

Diagnosis. — Colony encrusting to calyciform. Zooids porcellanous, the frontal shield imperforate, with

small areolar pores or these lacking. Orifice with a broadly arcuate poster delimited from the anter by inwardly

directed condyles, the distal arch not beaded. Articulated oral spines present in both infertile and fertile zooids.

Adventitious avicularia, when present, frequently lateral-oral; the crossbar complete. Ovicell subimmersed, the

ectooecium imperforate, usually covered by a smooth layer of secondary calcification; the ovicellular entrance

usually widely open, not closed by the zooidal operculum.

REMARKS. — A new genus is established here for the largely tropical to warm-temperate species previously

attributed to Hippoporella. There has been some debate over the relationships between numerous smooth-shielded,

warm-water ascophorines with arcuate posters, the species being variously attributed to Hippoporella Canu, 1917,

Lepraliella Levinsen, 1917, Hippoporina Neviani, 1895 sensu CANU & Bassler, 1920, Cleidochasma Harmer,

1957, and Brodiella Uttley & Bullivant, 1972 (e.g., Harmer, 1957; Powell, 1967; Hayward & COOK, 1983;

Gordon, 1984; Cook, 1985). Hayward and Cook (1983: 104) commented concerning Hippoporella that "this

genus includes many tropical species ... that seem to have little in common with the boreal-arctic type species".

We agree, hence the establishment of Lifuella , named for the type locality of the following species. The putative

species of Hippoporella with a beaded oral arch should probably be included in Stephanollona Duvergier, 1921 (see

Gordon, 1994).

Lifuella calyciformis (Philipps, 1900)

Fig. 145

Lepralia calyciformis Philipps, 1900: 446, pi. 43, figs 9, 9a.

Hippoporella calyciformis - Harmer, 1957: 1097, pi. 73, figs 1-5. — Hayward, 1988: 322, pi. 10c.

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn DR 117, 97-92 m.

Distribution. — New Caledonia, Lifou. Torres Strait. Indonesia, 0-34 m. Philippines, 49-97 m. Mauritius.

Remarks. — The single specimen in the collection is infertile and the oral spines were detached, but it

otherwise closely matches the illustrations of Philipps (1900), Harmer (1957), and Hayward (1988), who

shows the ligula on the avicularian crossbar. The present specimen has the paired lateral-oral avicularia as shown

by Philipps and Hayward but these may also occur proximal to the orifice and a third avicularium may occur

distally.

Genus SCHEDOCLEIDOCHASMA Soule, Soule & Chaney, 1991

Type Species. — Schedocleidochasma porcellaniforme Soule, Soule & Chaney, 1991.

Schedocleidochasma sp.

Fig. 150

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 38, 360 m.

Philippines. Holotype specimen of Hippoporina planulata Canu & Bassler, 1929 (Smithsonian Institution,

Department of Paleobiology, registration no. 8086, from "Albatross" Stn 5147, 38 m. Sulu Archipelago).

Source MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

59

DESCRIPTION. — Colony encrusting. Zooids very small, 0.19-0.32 mm long and 0.13-0.28 mm wide, the

frontal shield glabrous, even or with low ridges, wholly imperforate except for a single tiny areolar pore near the

mid-proximal margin. Orifice longer (0.058-0.077 mm) than wide with a distinct rounded-U-shaped sinus with tiny

condylar processes at the entrance; 6 spines bases around the distal rim. No avicularia. Ovicells not seen.

Distribution. — New Caledonia, near lie des Pins, 360 m.

REMARKS. — Insofar as the sole colony in the collection is infertile, it resembles Hippoporina planulata Canu

& Bassler, 1929 (= Metacleidochcisma planulata comb. nov.). Both species have small, vitreous zooids that are

strongly reflective. The orifices of M. planulata , however, have beaded rims and lack adjacent oral spines.

[Metacleidochasma planulata , incidentally, is identical to M. ovale Soule, Soule & Chaney, 1991, from Hawaii

and thus a senior synonym]. Although not noted in the holotype specimen by Canu & Bassler (1929), an

avicularium was seen that is identical to thus illustrated in M. ovale.

Genus RHYNCHOZOON Hincks, 1895

TYPE Species. — Lepralia bispinosa Johnston, 1847.

Rhynchozoon tubulosum (Hincks, 1880)

Figs 146-147

Mucronella (?) tubulosa Hincks, 1880: 383, pi. 17, fig. 7.

Rhynchozoon tubulosum - Harmer, 1957: 1064, pi. 65, figs 16-19. — Gordon, 1984: 125, pi. 52, C-E. — Hayward,

1988: 336 (part). — d’Hondt, 1986: 703. — Ryland & Hayward, 1992: 294, fig. 32 c-e.

MATERIAL EXAMINED. — New Caledonia. Musorstom 4: stn DW 231, 75 m.

DISTRIBUTION. — Sri Lanka. Indonesia, 0-52 m. Philippines, 13-275 m. Queensland. New Caledonia,

Chesterfield Islands and lie des Pins, 15-75 m. Kermadec Ridge, Curtis and Raoul Islands, 18-60 m.

Remarks. — Ryland and Hayward (1992) have commented on the limitations of Harmer's (1957)

synonymy of this Indo-Pacific species.

Rhynchozoon ligulatum sp. nov.

Figs 148-149

MATERIAL EXAMINED. — New Caledonia. BlOCAL: stn DW 38, 360 m.

Musorstom 4: stn DW 223, 545-560 m.

Types. — Holotype : a dried colony from BlOCAL Stn DW 38, 22°59.74'S, 1 67° 1 5.3 1 'E, 360 m, MNHN Bry-

20020.

Paratypes : an alcohol-preserved colony from MUSORSTOM 4 Stn DW 223, 22°57.00'S, 167°30.00'E, 545-

560 m, MNHN Bry- 19998, and a gold-coated colony from the same station, MNHN Bry-20077.

DESCRIPTION. — Colony encrusting, the dorsal surface with short stout basal-wall processes supporting the

colony above the substratum in places. Zooids relatively large, 0.74-0.94 mm long and 0.59-0.86 mm wide, with

an undular-nodular surface. Primary orifice transversely elongate-oval, with a beaded distal rim and a small pair of

rounded condyles that separate the anter from the poster. Surrounding the orifice is a peristome composed of several

stout tubercles. Secondary calcification can reduce their relative height; otherwise, with lateral fusion (with some

gaps between) they can form a palisade around the orifice, concealing the suboral mucro. Zooidal surfaces with

abundant small oval avicularia; these have a crossbar with a stout ligula and an upturned rostral rim. Larger avicu¬

laria absent. Ovicells somewhat immersed in secondary calcification, each with a very broad, descending labellum.

60

D. P. GORDON & J.-L. D'HONDT

Distribution. — New Caledonia, southwest of lie des Pins, 360-560 m.

Remarks. — This distinctive species is easily recognisable from the abundant small ligulate avicularia.

Genus IODICTYUM Harmer, 1933

Type Species. — Retepora phoenicea Busk, 1884.

lodictyum bicuspidatum sp. nov.

Figs 151-154

MATERIAL EXAMINED. — New Caledonia. Musorstom 6: stn CP 419, 283 m.

TYPES. — Holotype: an ovicelled branch fragment, gold-coated, from MUSORSTOM 6 Stn CP 419, 20°41.65'S,

167°03.70’E, 283 m, MNHN-Bry 20074.

Paratype : colony fragments, from the same locality as the holotype, MNHN Bry-20183.

DESCRIPTION. — Colony non-pigmented, erect, branching more or less in one plane, exceeding 2 cm in

height. Not strictly fenestrate, but anastomoses occur. Branches mostly 2-3-serial, with branch widths ranging

from 0.43-2.13 mm. Primary orifice nearly circular, just a little wider than long, without beading of the rim or a

proximal sinus. Zooids 0.71-0.84 mm long (orifice to orifice) and c. 0.29-0.43 mm wide, mostly discrete,

especially in young branches, clearly outlined by ridges along the zooidal boundaries. Zooidal skeletal surface

smooth or faintly subgranular, with 1-2 pairs of marginal pores in the proximal half. Zooids lacking ovicells have

a tubular, somewhat projecting peristome that has blunt denticulations around the rim, and a median descending

groove that ends in a pseudosinus at the beginning of the shaft. Ovicelled zooids have incompletely tubular

peristomes, the distal rim being occupied by an ovicell that is initially prominent and subglobular but later appears

recumbent owing to secondary calcification. The frontal face of the ovicell is completely smooth apart from a faint

midfrontal depression. There is no labellum. Avicularia occasional, large, borne frontally on ovicelled or non-

ovicelled zooids in the median row(s). They occupy most of the frontal surface from which they project

prominently. The crossbar is complete but, unusually, the opesia behind the crossbar is narrow to the point of

obliteration. The rostrum is Ungulate in shape, slightly narrowing in the middle, before expanding and rising

distally in a pair of smooth rounded cusps. A palatal shelf occupies about the rostral area. The mandible exactly

fits the shape of the rostrum. Dorsal branch surface entirely lacking avicularia, but with transverse vibices.

Distribution. — New Caledonia, northwest of Lifou, 283 m.

Remarks. — lodictyum bicuspidatum is most easily distinguished by its large bicuspid (hence the species

name) frontal avicularia. Both this species and the following two species are unusual in being both white and

largely non-fenestrate. Harmer (1934), in the most comprehensive treatment of tropical erect phidoloporids to

date, described 1 1 species of lodictyum from Indonesian waters. Of these, only three species were white and only

one, pigmented, was open-branched.

A commensal organism, probably a hydrozoan, may be associated with this species. Some of the colony

fragments have slit-like gaps in the calcification along zooidal margins, indicating that stolons of the epizoite are

mostly covered over by the calcification of the bryozoan. Circular openings at zooidal corners, larger than areolar

pores, indicate where hydroid stems emerged.

lodictyum blandum sp. nov.

Figs 155-157

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 44, 440 m.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

61

TYPES. — Holotype: unique colony from Biocal Stn DW 44, 22°47.30’S, 167°14.30’E, 440 m. No separate

paratypes.

Description. — Colony reticulate, shaped overall like a convex clam shell, the zooids opening on the

concave face, 11 mm high; unpigmented. Zooids 2-3-serial in trabeculae 0.58-1.04 mm wide. Fenestrae ranging

from 0.59-2.60 mm long and 0.20-0.66 mm wide. Both faces of the colony and the zooidal surfaces glabrous,

relatively smoothly textured, and blandly lacking many distinctive features. Primary orifice lacking beading on

distal rim. Peristome with internal diameter of c. 0.094-0.113 mm, with peristomial sinus in proximal rim the

descends as a spiramen within the rim; some peristome rims with crenulations. Avicularia rare; occasional

relatively large avicularia occur proximolateral to the peristome, the long acute rostrum directed past the

pseudosinus. No fenestral avicularia. Ovicell fairly immersed, with a short labellum and lateral flanges, and a short

slit-like pore extending onto the labellum. Abfrontal surface of colony lightly textured, with thin ramifying

vibices and rare avicularia like the acute suboral ones frontally.

Distribution. — New Caledonia: west of lie des Pins, 440 m.

REMARKS. — This species is characterised by its particularly bland appearance, including the striking paucity

of polymorphs.

Iodictyum illinguum sp. nov.

Figs 158-160

MATERIAL EXAMINED. — New Caledonia. MUSORSTOM 6: stn DW 421, 245 m.

Types. — Holotype : an ovicelled colony, gold-coated, from MUSORSTOM 6 Stn DW 421, 20°26.27'S,

166°40.17'E, 245 m, MNHN Bry-20182.

Paratype : colony fragment, from the same locality as the holotype, MNHN Bry-20048.

Description. — Colony non-pigmented, generally hyaline, erect, branching more or less in one plane, the

only complete colony 1 cm in height. Branches mostly biserial, triserial in places, with branch widths ranging

from 0.33-1.10 mm. Primary orifice and peristomes as in /. bicuspidatum. Zooids 0.71-0.90 mm long (orifice to

orifice) and c. 0.33-0.41 mm wide, the frontal shield and boundaries (including the presence of a commensal

organism) appearing as in /. bicuspidatum. Ovicells also more or less identical, but with a longer and more

apparent median depression. Large avicularia occasional frontally, situated in a concavity of each zooid that bears

one, with a tapering triangular rostrum, and a short narrow opesia behind the crossbar. Similar avicularia are

commoner abfrontally, with generally one on each dorsal kenozooid between transverse vibices.

Distribution. — New Caledonia, northeast of Ouvea, 245 m.

REMARKS. — This species is remarkably similar to /. bicuspidatum in colonial and zooidal morphology, but

may be readily distinguished on the basis of the avicularia - those of /. bicuspidatum being raised, linguiform,

bicuspid, and lacking abfrontally, whereas those of /. illinguum are depressed, triangular, and present abfrontally.

The species name illinguum alludes to the lack of an ovicellular labellum.

Iodictyum trochus sp. nov.

Figs 161-164

Material EXAMINED. — New Caledonia. Biocal: stn DW 46, 775 m. — Stn DW 66, 515 m.

Musorstom 6: stn CP 419, 283 m.

TYPES. — Holotype: a relatively large flabellate colony fragment preserved in alcohol, from MUSORSTOM 6

Stn CP 419, 20°41.65'S, 167°03.70'E, 283 m, MNHN Bry-19985.

62

D. P. GORDON & J.-L. D'HONDT

Paratypes : colony fragments from the same locality as the holotype, MNHN Bry-20184; a colony fragment

from BIOCAL Stn DW 66, 24°55.43’S, 168°21.67'E, 515 m, MNHN Bry-20084.

Description. — Colony non-pigmented, attaining 28 mm height from a basal stem c. 2.0 mm diameter,

branching more or less in one plane, with anastomoses, in a flabellate manner. Branches 2-4 serial, with branch

widths ranging from 0.41-1.40 mm. Primary orifice orbicular with no beading. Zooids c. 0.59-0.66 mm long

(orifice to orifice) and c. 0.28-0.37 mm wide (internal chamber width c. 0.22 mm), with indistinct boundaries and a

lightly textured surface calcification. Each zooid has a single minute pore frontally, usually adjacent to an

occasional small oval avicularia that is set transversely. It has thin crossbar and orbicular opesia, and a

proportionately larger rostrum with a semicircular palatal shelf. Zooids have long, projecting peristomes with

circular orifices. These are striking in appearance, with a circlet of short denticulate ridges inside the rim. The

denticles from these ridges vary somewhat in the extent to which they project, but each is acute and directed at

angle towards the orifice interior. Beneath the medioproximal pair of denticles a sinus groove, not externally

obvious, descends the full length of the peristome to the primary orifice. A rare avicularium may be incorporated

into the peristomial rim; this is long and acute, curving with the rim on one side where denticles are lacking; the

crossbar is a ridge, with a sunken opesia on one side and an elongate palatal foramen on the other beyond which is

a palatal groove. Ovicells difficult to see; set near the bottom of the peristomial tube and largely concealed by

secondary calcification. Dorsal branch surface evenly and lightly rugose with smooth depressions and ridges and

occasional obliquely set vibices ridges; no abfrontal avicularia.

Distribution. — New Caledonia, northwest of Lifou, south of tie des Pins, and Norfolk Ridge, 283-775 m.

Remarks. — The radiate peristomial openings of this species are particularly striking and are alluded to in the

epithet trochus , from Greek trochos , a wheel. One other noteworthy characteristic of this species is the intimate

association of a commensal epibiont, which forms a ramifying stolon along the frontal axis of each branch.

The stolons are covered over by zooidal calcification but gaps in this are apparent, along with circular

openings, nearly the same size as the small oval avicularia, at zooidal margins. The association between

commensal begins immediately after larval settlement. The ancestrula is sac-shaped, with an opesia in the distal

half surrounded by a circlet of five short spines. Periancestrula zooids have ramifying calcified tubes closely

adherent to them.

A colony fragment from Biocal Stn DW 46 doubtfully belongs to this species. The peristome is more

expanded distally, the sinus is apparent at the rim, the frontal zooidal surface has polygonal depressions, and frontal

avicularia are not apparent.

lodictyum sp.

Figs 165-166

MATERIAL EXAMINED. — New Caledonia. MUSORSTOM 4: stn DW 151, 200 m.

DESCRIPTION. — Colony reticulate, pigmented. Zooids 2-3-serial in trabeculae 0.43-0.57 mm wide. Fenestrae

ranging from 1.24-1.36 mm long and 0.73-0.75 mm wide. Frontal shield relatively smooth with 3-5 areolar pores

distributed around its surface. Primary orifice with no beading of the distal rim; poster with a broad rounded sinus

medially, separated from the anter by a pair of stout condyles. Peristomial rim raised, lacking spines or

crenulations, with a conspicuous pseudosinus and short sinus groove flanked and defined inwardly by a pair of

stout tubercles. Avicularia of two kinds: a subcircular avicularium on the frontal shield of most zooids, and a larger

sublingulate avicularium on the zooids that occur at the proximal end of each fenestra; the avicularium broader

proximally, with a subparallel rostrum that has an extensive palate and is truncated at the tip. Ovicells not seen.

Frontal calcification conceals the ramifying stolons of symbiotic hydroids; the openings of the erect hydroid stems

visible at the edges of zooids as circular holes intermediate in size between areolar pores and the round avicularia.

Abfrontal surface of colony fairly smooth, with thin vibices. No avicularia on the inner sides of fenestrae, only the

lateral areolar pores of autozooids.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

63

Distribution. — New Caledonia, north of Belep Islands, 200 m.

Remarks. — This species was represented only by a small, infertile fragment, now in two pieces, one of

which is gold-coated. It appears to have been transported from shallower water. The overall colour is pale pink but

some peristomes give evidence that the colour in life was deep magenta.

It is probable that the species is new, but the fragment lacks ovicells and is too small to name formally.

Genus RETEPORELLA Busk, 1884

Type Species. — Reteporella flabellata Busk, 1884.

Remarks. — Reteporella is here regarded as encompassing the genus Sertella Jullien in JULLIEN & Calvet,

1903.

Reteporella concinnoides sp. nov.

Figs 167-171

MATERIAL EXAMINED. — New Caledonia. Biocal: stn CP 108, 335 m.

MUSORSTOM 4: stn DW 231, 75 m.

MUSORSTOM 6: stn DW 421, 245 m.

TYPES. — Holotype : an alcohol-preserved colony from BlOCAL Stn CP 108, 22°02.55'S, 167°05.68'E, 335 m,

MNHN Bry- 19983.

Paratypes : an alcohol-preserved colony, MNHN Bry- 19982, and same dried colony fragments, some gold-

coated, MNHN Bry-20193, from the same locality as the holotype. Some dried colony fragments, some gold-

coated, from MUSORSTOM 4 Stn DW 231, 22°33.70’S, 167°10.50'E, 75 m, MNHN Bry-20192.

DESCRIPTION. — Colony fenestrate, starting more or less evenly calyciform, then developing more on one side

while curving inwardly; the two largest colonies attaining a height of 28 mm and a lateral spread of 35 mm.

Within-colony variation is apparent between the two stations, with the deeper-water colonies (BlOCAL

Stn CP 108) having a more open construction than a colony from MUSORSTOM 4 Stn DW 231. Thus, two sets

of measurements follow, with those from the shallower station given first, separated by a slash from those for the

deeper station. Fenestral length: 0.69-1.53 mm, averaging 1.18 mm / 0.97-2.43 mm, averaging 2.03 mm.

Fenestral width: 0.26-0.74 mm / 0.41-1.33 mm. Trabecular width: 0.41-0.77 mm / 0.35-0.71 mm. Zooid length

(orifice to orifice): c. 0.24-0.31 mm / 0.35-0.45 mm. Zooidal boundaries mostly merge, even in developing

zooids, though frontal vibices occur occasionally; zooidal frontal calcification smooth though lightly textured,

with inconspicuous small sparse tubercles. Primary orifice beaded, the proximal rim obscured by a short broad

lyrula. Peristomial orifice with 6 oral spines (up to 8 in a specimen from MUSORSTOM 6 Stn DW 421) in

developing zooids; some of these remain even in well-calcified parts of branches. The proximal rim of the orifice

has 3-4 parallel ridges, mostly at the opesial end of the small avicularium that is on the frontal side of the

peristome. This is acute and generally slightly curved, adjacent to a conspicuous spiraminal pore. Two kinds of

avicularia occur frontally on zooids. An occasional large avicularium, originating adjacent to the small labial

avicularium, curves across the front of the zooid at an angle. This is acute with an acute mandible, and a long

narrow palatal foramen. More numerous small avicularia occur scattered on the front of zooids; these are narrow,

acute, and set at various angles. Ovicell generally immersed in secondary calcification, with a short fissure visible

frontally that does not extend onto the well-developed, parallel-sided labellum. Abfrontal surface of branches evenly

and lightly rugose with sparse vibices; clusters of sunken small acute avicularia occur abfrontally at each end of a

fenestra, as well along the inner face of every fenestra adjacent to 2-3 larger elongate avicularia that have

conspicuously toothed rostral margins and an irregularly shaped opesia.

Distribution. — New Caledonia, Sarcelle Passage and offshore from Yate, 75-335 m.

64

D. P. GORDON & J.-L. D'HONDT

Remarks. — Reteporella concinnoides is very similar to R. concinna (Gordon, 1984), in colonial, zooidal,

and peristomial morphology. The obvious distinguishing feature is the form of the large avicularia, those of

R. concinna having a conspicuously toothed rostrum that projects directly outwards without curving.

Reteporella defensa sp. nov.

Figs 172-175

MATERIAL EXAMINED. — New Caledonia. Biocal: stn DW 38, 360 m. — Stn CP 108, 335 m.

Chalcal 2: stn DW 76, 470 m.

Musorstom 6: stn CP 419, 283 m. — Stn DW 421, 245 m.

Types. — Holotype : a flabellate colony in the form of a horizontal crescent, attached to the substratum, from

MUSORSTOM 6 Stn DW 421, 20°26.27’S, 166°40.17'E, 245 m, MNHN Bry-19974, in alcohol.

Paratypes : four colonies from the same station, not shaped like the holotype but in the same container, from

the same locality, MNHN Bry- 19975- 19978. Dried colony fragments from Biocal Stn CP 108, 22°02.55'S,

167°05.68'E, 335 m, MNHN Bry-19979; from Chalcal 2 Stn DW 76, 23°40.50’S, 167°45.20’E, 470 m, MNHN

Bry-20078; and from MUSORSTOM 6 Stn CP 419, 20°41.65'S, 167°03.70'E, 283 m, MNHN Bry-20076 and

20180.

Description. — Colony dendroid; in most of the colonies in the collection it is in the form of a horizontal

fan, at right angles to the relatively short ‘trunk’ by which it is attached to the substratum, with the zooidal

orifices facing the substratum; only 1 colony grew straight out from the substratum; colony size up to 42 mm

high and 36 mm across. The colony has numerous major branches, all in one plane, joined by short cross-

connections that have avicularia but no feeding zooids; in the central part of a colony the branches are 0.59-

1.40 mm wide, the cross-connections 0.37-0.62 mm wide. Zooids c. 0.63-0.71 mm long (orifice to orifice), the

calcareous surface smooth or faintly textured and somewhat undular, with varices roughly indicating zooidal

boundaries. Oral spines 6 in developing zooids, with 0-3 being retained in more calcified zooids; these may be

simple or bifurcates, or strikingly cervicorn. The proximal rim of the peristome includes a transversely set oval

avicularium with a toothed rostrum; subjacent to the opesia is a labial pore. A pair of oral spines or spine bases

frequently flanks the avicularium, even in ovicelled zooids. Apart from a few small areolar pores, the frontal shield

is well-endowed with avicularia, which are of two kinds. On some zooids are 1-2 small oval avicularia like the

peristomial avicularium, usually elevated, with a denticulate rostral rim. Much more common is an elongate boat¬

shaped avicularium, with an extensive flat palate and raised rims, that occurs in a range of orientations and sizes

from quite small to very large, the latter lying diagonally across the entire frontal surface of the zooid; the opesia is

narrow and slit-like, the suboval/subtriangular palatal foramen occupies about one-third the length of the palate,

with a small pore beyond; the entire avicularium is raised just above the surface of the autozooid, with the low

palatal rims each terminating in an upturned point so that there is a gap distally where the rostral tip would be; the

mandible does not project beyond the palate. Ovicell cucullate, with a transverse or subcircular apical foramen in

the ectooecium, revealing the endooecial tabula beneath; the entire ovicell becoming soon encroached up by

secondary calcification from adjacent zooids. Abfrontal surface divided into irregular sectors by vibices; within each

sector are several small pores and small to medium-sized avicularia like the frontal boat-shaped ones. The presence

of an ectosymbiotic hydroid is evidenced by the calcified tubes (mostly concealed) and tube openings on the frontal

sides of branches.

Distribution. — New Caledonia, Ouvea and Lifou Islands, southeastern New Caledonia, and northern

Norfolk Ridge, 245-470 m.

Remarks. — This species is instantly recognisable by its abundant, distinctively shaped avicularia, and cross

branches lacking autozooids. Additionally, however, R. defensa is characterised by cervicorn oral spines and

cucullate ovicells with a tabula, both features being apparently unique in the genus.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

65

Reteporella ferox sp. nov.

Figs 176-178

MATERIAL EXAMINED. — New Caledonia. Musorstom 6: stn CP 419, 283 m.

Types. — Holotype: a single large colony from Musorstom 6 Stn CP 419, 20°41.65'S, 167°03.70’E,

283 m, MNHN Bry- 19981.

Paratype : coated fragment from the holotype colony, MNHN-Bry 2008 1 .

DESCRIPTION. — Colony erect, non-fenestrate, with a few main axes from which dichotomous branches arise,

attaining 27 mm height. Main stem maximally 1.3 mm across, the smallest branches biserial, 0.36 mm across.

Zooids lacking distinct boundaries between them, the overall surface more or less smooth with gentle irregularities

and small sparse areolar pores. Occasional larger round pores between zooids presumably belonging to an epibiotic

hydroid in life. Peristomial orifices typically with 5 spine bases (2 on one side, 3 on the other) or these obliterated

by secondary calcification; internal diameter of orifice 0.11-0.15 mm wide. Proximal rim with a small oval

avicularium medially, the cross-bar with tiny processes on each side, the rostral rim with a few denticulations; the

peristomial side of the avicularium with a small cusp. Small avicularia scattered sparsely over the frontal surface of

the colony, these with a narrowly tapering acute rostrum. Larger avicularia conspicuous near or at the branch

margins; these with the opesial area obliterated by calcification and the distal half (most of the rostrum) serrated

and down-curved. Ovicells appearing externally as bulges, generally completely covered by secondary calcification;

a small part of the endooecial exposure may be visible proximally. The abfrontal surface of the colony like the

frontal surface or a little more crinkly, with tiny scattered pores, and some of the small acute avicularia along the

margins.

Distribution. — New Caledonia, northeast of Lifou, 283 m.

Remarks. — Reteporella ferox is most easily distinguished by the distinctive large lateral avicularia with

toothed, down-curved rostra. Where these are common the branch margins appear quite jagged, hence the specific

name ferox , Latin, fierce.

Reteporella orstomia sp. nov.

Figs 179-180

Material examined. — New Caledonia. Musorstom 6: stn DW 431, 21 m.

Types. — Holotype : an alcohol-preserved colony from Musorstom 6 Stn DW 431, 20°22.25'S,

166°10.00'E, 21 m, MNHN Bry-19984.

Paratypes : dried colony fragments from the same locality as the holotype, MNHN Bry-20082 and 20185.

Description. — Colony erect, to 16 mm high, more or less open-branched but with narrow cross-connections

that contain few or no autozooids. Branch width near the originally calyciform base of the colony c. 0.93-

1.22 mm, reducing to 0.31-0.75 mm in the distal half. Cross-connections are 0.20-0.34 mm thick. Zooids 0.37-

0.47 mm long (orifice to orifice), outlined by thin vibices, with faintly textured, smooth frontal calcification in

which are a few small pores. Peristome relatively well developed in neanic zooids, especially laterally, and with 1-

2 short processes, the height of the peristome becoming reduced as secondary calcification thickens the frontal

shield; the proximal rim of the peristome has a median sinus that descends as a slit-like sinus tube. There is no

labial avicularium or pore (spiramen). Frontal avicularia of two kinds, both large, absent from some parts of

colonies but occurring in clusters in others. One kind projects frontally in the centre of the zooid, with an acute

triangular rostrum that is upturned at the tip; the opesia is short and narrow, the palatal foramen relatively large

and suboval. The other, slightly sunken, is on one side of the zooid and directed proximally; the opesia and palatal

are as in the other avicularium, but the rostrum is subspatulate, narrowing in the middle before expanding distally.

66

D. P. GORDON & J.-L. D'HONDT

where it is truncate with upturned tips on each side; its mandible is likewise abruptly truncate. There are no small

avicularia. Ovicell with a median longitudinal fissure that does not extend onto the labellum which is turned

inward somewhat. Abfrontal surface of branches lightly rugose, with thin vibices and scattered small pores, but no

avicularia.

Distribution. — New Caledonia, Beautemps-Beaupre Atoll north of Ouvea, 21 m.

Remarks. — This species is relatively nondescript, but may be distinguished reasonably easily by the

presence of two kinds of large frontal avicularium and the complete absence of abfrontal avicularia.

Reteporella sp.

Figs 181-184

Material EXAMINED. — New Caledonia. Biogeocal: stn DW 253, 310-315 m.

DESCRIPTION. — Sole colony fragment 4 mm long, with 2 broken bifurcations; branch width 0.62-0.85 mm.

Colony surface, including abfrontally, regularly dimpled. Zooidal boundaries completely obscured. Peristomes

slightly raised above frontal surface, somewhat pyriform with a smooth inner rim; interior diameter of peristome

0.094 mm; a median spiraminal pore delimited by a pair of tubercle-like processes that fuse at their tips; no sinus

groove or incorporated avicularium. Oral spines not seen. Tiny oval avicularia occur sparsely on the colony

surface, with a short stubby ligula on the rostral side of the crossbar. Ovicell mostly concealed by dimpled

secondary calcification, the ectooecial surface smooth with a median slit in its frontal face, the labellum truncate

with no lateral flanges. No other characters discernible.

Distribution. — New Caledonia, off east coast, NE of Thio, 310-315 m.

Remarks. — Owing to the small size of the colony fragment it is not possible to determine whether the

species is reticulate or open-branching. Its dimpled surface invites comparison with the genus Malleatia Jullien in

Jullien & Calvet, 1903, characterised by “ponctuations cupuliformes”. Apart from the surface sculpturing,

particular similarities with Malleatia rara include: mostly concealed ovicell with a short slit frontally and short

truncate labellum, apparent absence of oral spines, and small round adventitious avicularia. Details of the

peristome differ, however, in that M. rara has a median suboral avicularium, so that the labial spiramen tube is

offset to one side. In this, M. rara resembles Reteporella malleatia (Gordon, 1984) from the Kermadec Ridge,

which also has oral spines. Thanks to the courtesy of Dr Peter J. Hayward (University of Wales, Swansea), it is

possible to compare the type species of Reteporella (R. flabellata Busk, 1884) with Malleatia rara. His

unpublished scanning electron micrographs of R. flabellata show that it has an irregularly beaded distal rim like

that of M. rara , the form of the ovicell is more or less identical, and the frontal shields of both species are

umbonuloid (as in some other phidoloporids - pers. obs., DPG). Reteporella flabellata lacks a peristomial

avicularium, having instead a median or submedian spiraminal opening (so-called labial pore) like that in the

present specimen from BIOGEOCAL Stn DW 253. Overall, then, it is apparent that there is no consistent

correlation between the presence of frontal-shield dimpling and the characters of the primary orifice and peristome.

Both Malleatia rara and the present New Caledonian specimen can be included in Reteporella Busk.

Genus RETEPORELLINA Harmer, 1933

Type Species. — Retepora denticulata Busk, 1884.

Reteporellina cruciformis sp. nov.

Figs 185-187

Material EXAMINED. — Philippines. Musorstom 3: stn DR 117, 97-92 m.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

67

Types. — Holotype : parts of a dried single colony from Musorstom 3 Stn DR 117, 12°31.3’N, 120°29.5’E,

97-92 m, MNHN Bry-20080.

Description. — Colony erect, dichotomously branching, the longest fragment 8 mm high. Branches 0.43-

0.84 mm wide, the zooids in 3-4 longitudinal rows, 5 before bifurcations. Zooids 0.41-0.45 mm long (orifice to

orifice) and 0.20-0.22 mm wide, with a smooth surface and distinct boundaries. Zooidal peristome long and tubular

on non-ovicelled zooids, especially the lateral ones, with an spout-like rim. Primary orifice suborbicular, with a

beaded distal rim and a small pair of condyles. The inner rim of the peristomial orifice has a median pseudosinus,

that descends as a sinus groove down the inside of the spout where it is flanked by a pair of stout tubercles.

At a lower level within the peristome, on the distal face, is another pair of stout tubercles. When seen frontally,

the two pairs of tubercles, though at different levels and incompletely cruciform, appear to oppose one another.

Neanic parts of colonies may consist entirely of zooids lacking ovicells and frontal avicularia. In contrast, mature

parts of colonies have clusters of zooids (ovicelled and non-ovicelled) which each have a large, spatula-shaped,

labial avicularium. This tends to curve around in front of the peristomial rim and are directed obliquely forward

with the palate facing distally. The avicularian opesia is more or less obliterated; the palatal foramen is elongate-

oval, and the rostral rim is roundly subtruncate with the outer corners produced slightly, giving a bicuspid

appearance. A similar avicularium, more obviously bicuspid, occurs along the outer lateral margin of many zooids,

descending the branch with the rostrum pointing proximally, the palate facing laterofrontally. Ovicells recumbent,

becoming covered distally by secondary calcification, with a short rounded labellum beyond which the lateral

processes curve round almost to fuse proximally; the median fissure does not extend as far as the labellum.

Abfrontal side of branches with the lightly cobbled surface crossed by vibices. A moderately large avicularium

occurs below most branch axils, raised or flush with the surface.

Distribution. — Philippines, Mindoro Strait, 92-97 m.

Remarks. — Reteporellinci cruciformis resembles R. idmoneoides Harmer, 1934 and the following new

species in having projecting peristomes and large labial avicularia, but may be distinguished from both by the

cruciform arrangement of internal peristomial tubercles and the shape of the lateral avicularia.

Reteporellina spiramina sp. nov.

Figs 188-191

Material examined. — Philippines. Musorstom 3: stn CP 139, 240-267 m.

BlOCAL: stn DW 44, 440 m. — Stn DW 66, 515 m.

TYPES. — Holotype : the largest of five dried colony fragments from BlOCAL Stn DW 66, 24°55.43'S,

168°21.67'E, 515 m, MNHN Bry-20085.

Paratypes : dried colony fragments from MUSORSTOM 3 Stn CP 139, 11°53.0'N, 122°14.0'E, 240-267 m,

MNHN Bry-20189, and from BlOCAL Stn DW 44, 22°47.30'S, 167°14.30’E, 440 m, MNHN Bry-20047.

Description. — Colony erect, tending to have one main, curving stem from which dichotomous branches

arise, the longest fragment c. 1 1 mm. Branches 0.43-1.01 mm wide, the zooids in 2-3 (mostly 3) longitudinal

rows, 4 before bifurcations. Zooids 0.57-0.78 mm long (orifice to orifice) and c. 0.31 mm wide, with a lightly

granular-tubercular surface and distinct boundaries. Zooidal peristome projecting, especially on marginal zooids, the

rim irregular; the distal rim of the youngest zooids may bear 3 articulated spines whose bases are discernible in

many older zooids. In the middle of the proximal rim of the peristomial orifice is the entrance to a spiraminal tube

that descends through a narrow ridge down the inside of the peristome. A large labial avicularium, more or less

identical to that of the preceding species, occurs with the same orientation and position in front of the peristome of

some zooids, both ovicelled and non-ovicelled. A different-shaped large avicularium occurs along the outer lateral

margin of many zooids and below branch axils; this is elongate with gently tapering sides and a rounded rostral tip

directed proximally; there is a small palatal foramen by the crossbar, a smaller foramen near the rostral tip, with a

68

D. P. GORDON & J.-L. D’HONDT

palatal shelf in-between. On the side opposite the avicularium is a concavity of the same size that accommodates

the folded-back mandible. Ovicells recumbent, becoming covered laterally by secondary calcification, with a narrow

labellum onto which the long median fissure extends; lateral processes do not fuse proximally. Abfrontal branch

surface granular, with vibices.

DISTRIBUTION. — Philippines, south of Mindoro, 240-267 m. New Caledonia, lie des Pins and northern

Norfolk Ridge, 440-515 m.

Remarks. — The spiraminal tube and shape of the lateral avicularia distinguish R. spiramina from the

preceding species.

Reteporellina granulosa sp. nov.

Figs 192-193

MATERIAL EXAMINED. — New Caledonia. BIOCAL: stn DW 38, 360 m.

TYPES. — Holotype : a single, broken, branch fragment from BlOCAL Stn DW 38, 22°59.74'S, 167° 15.3 l'E,

360 m, MNHN Bry-20079.

Description. — Colony erect, branching dichotomously, the sole fragment (now in two pieces) 6 mm long.

Branch width 0.48-0.69 mm, the zooids in 2-3 longitudinal rows. Zooids 0.65-0.79 mm long (orifice to orifice),

the surface coarsely granular with thin vibices, the zooidal boundaries not clearly delimited. Zooidal peristome

projecting only moderately in marginal zooids, tending to be shortened and lost through secondary calcification, the

rim somewhat scalloped by short ridges just inside the rim. No labial pore or avicularium; a median sinus groove

occurs down the midline. A large avicularium occurs on the front of a number of zooids, ovicelled and non-

ovicelled; this is directed proximally or at a slight angle; the rostrum is elongate-triangular with an extensive

palate and small, narrow palatal foramen; the opesia is minute, beyond which there is a subtriangular concavity to

accommodate the folded-back mandible. A similar avicularium occurs on the abfrontal surface, behind many of the

peristomes. Ovicell recumbent, pyriform, becoming covered by secondary calcification; the median fissure is

confined almost entirely to the short, triangular labellum, beyond which the long lateral processes converge below.

Abfrontal surface coarsely granular with thin vibices.

Distribution. — New Caledonia, southwest of lie des Pins, 360 m.

Remarks. — This species is distinguished particularly by the characters of the ovicellular labellum and frontal

avicularium.

Reteporellina projecta sp. nov.

Figs 194-195

Material EXAMINED. — New Caledonia. Biogeocal: stn CP 290, 920-760 m.

Types. — Holotype : a very small colony fragment from BIOGEOCAL Stn CP 290, 20°36.9TS, 167°03.34'E,

760-920 m, MNHN Bry-20190.

DESCRIPTION. — Colony erect, branching, the zooids more or less in 2 series, with the interpolation of

additional zooids just before a bifurcation. Branch width 0.45-0.6 mm between peristomes which project

alternately to left and right of the branch axis. Zooidal width (best determined from the abfrontal side) 0.28-

0.33 mm. Peristomes of variable length, the median and near-median ones shorter than the lateral ones that extend

up to 0.50 mm from the branch surface. Frontal shield and peristomes with a light granular-tubercular texture.

Primary orifice lacking beading of the distal rim. Peristomial orifice bordered by 7 spines distally; on the

proximofrontal rim a labial suture descends to a labial pore (spiramen). Typically a small oval avicularium near the

base of the peristome on the frontal shield; another such avicularium can occur proximally on the shield; these

Source : MNHN . Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

69

can later become occluded. No other avicularia seen. Ovicells not present. Abfrontal surface textured like

the frontal side, separated from the frontal surface by a thin varix coursing down the outer side of each

lateral zooid; with occasional small round avicularia, these tending to be raised slightly above the surrounding

surface.

DISTRIBUTION. — New Caledonia, north of Lifou, 760-920 m.

Remarks. — Although the sole fragment is small and infertile it is very distinctive and easily recognisable

among the relatively small number of described species of Reteporellina. The distinctive features are peristomial,

i.e., strongly projecting, with seven distal spines, and a long labial suture.

Genus TRIPHYLLOZOON Canu & Bassler, 1917

Type Species. — Retepora monilifera MacGillivray, 1 860.

Triphyllozoon gracile sp. nov.

Figs 196-200

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn DR 117, 92-97 m.

Types. — Holotype : the largest stem in a welled slide of fragments from Musorstom 3 Stn DR 117,

12°31.3'N, 120°39.5'E, 92-97 m, MNHN-Bry 20049.

Paratypes: the remaining fragments in the same slide, from same locality as holotype, MNHN-Bry 20049.

DESCRIPTION. — Colony erect, branching, the branches slender, minimally 0.41 mm across, widening to

0.85 mm before a bifurcation. Zooids 2-3-serial, c. 0.45-0.52 mm long, the frontal shield sparsely and lightly

granular-tubercular, with no obvious boundaries and small sparse areolar pores. Distal rim of primary orifice

lacking beading. Peristomial orifice outwardly symmetrical, with a pair of small oval avicularia with denticulate

rostral rim flanking a median spiraminal pore; notwithstanding the outward symmetry, a sinus groove descends to

one side of an asymmetrical lyrula internally that has a small ala on one side only. Additional small oval avicularia

are scattered sparsely on the frontal surface of branches; occasional larger suboval avicularia with parallel sides

occur along the outer margins of the most lateral zooids; no spatulate or munitiform avicularia. Ovicells occurring

in vertical rows of 3 or more along the branch axis; trilobed stigma with short lateral arms, the median arm

ascending to the apex of the ovicell which is submucronate; proximal edge of ovicell truncate with no lateral

flanges. Abfrontal surface separated from the frontal surface by a varix coursing along the sides of the most lateral

zooids; the surface like that frontally, with numerous similar avicularia, especially the larger ones, occurring with

small pores along the outer edges.

REMARKS. — Because the fragments are broken it is not possible to affirm whether the species is open-

branching or loosely fenestrate. If it is fenestrate, it must have the largest fenestrae or open appearance of any

species. The symmetrical arrangement of peristomial avicularia is very distinctive, allowing for easy recognition of

the species.

Superfamily CONESCHARELLINOIDEA Levinsen, 1909

Family LEKYTHOPORIDAE Levinsen, 1909

Genus HARPAGO Gordon, 1989

Type Species. — Harpago minutus Gordon, 1989b.

70

D. P. GORDON & J.-L. D’HONDT

Harpago dissidens sp. nov.

Figs 201-204

MATERIAL EXAMINED. — New Caledonia. BiOCAL: stn DW 66, 515 m.

Types. — Holotype : a single colony from BiOCAL Stn DW 66, 24°55.43'S, 168°21.67'E, 515 m, MNHN-Bry

20036. No paratypes.

Description. — Sole colony somewhat conical in overall shape, the outline broken by the projecting

peristomes, 2.04 mm high and potentially (but for broken peristomes) 2.26 mm maximum diameter. Zooids more

or less whorled around the colony axis, the zooids in one whorl nearly alternating with those above and below.

Zooids somewhat inflated in appearance, tapering to an outwardly projecting tubular peristome. Frontal shield

smooth, glabrous, with 4-5 tiny areolar pores along each margin. Primary orifice 0.15 mm long and 0.19 mm

wide with a more-or-less V-shaped sinus flanked and defined by a stout .pair of condyles angled across the proximal

corners of the orifice. No peristomial or other avicularia. Ovicells large, sac-like, with an extensive smooth tabular

area bordered by a low rim and with a pair of pores at the peristomial end of the tabula. At the broad growing apex

of the colony is a small central area of ?extrazooidal calcification in which are tiny septular pores.

Distribution. — New Caledonian waters: northern Norfolk Ridge, 515 m.

Remarks. — It is unfortunate that the sole colony lacks the earliest-formed zooids for it is not possible to be

certain if it was encrusting or rooted. In its overall form it closely resembles Harpago minutus , but lacks the

dimpled calcification of that species and, more importantly, has an ovicellular tabula. This latter feature may be

adequate for generic distinction from Harpago.

Family BATOPORIDAE Neviani, 1900

Remarks. — The name Batoporidae Neviani, 1900 has priority over Orbituliporidae Canu & Bassler, 1923.

Genus PTOBOROA nov.

Type Species. — Batopora pulchrior Gordon, 1989a.

Diagnosis. — Colony conical to claviform, with a single axial tube proximally. Zooids more or less

alternating in whorls with short, laterally projecting peristomes. Primary orifice circular or nearly so, with a broad

rounded poster separated from the anter by a pair of angular condyles; with a pore incorporated into the peristomial

rim middistally. Adventitious avicularia tiny, suboval with a pair of mandibular pivots, not associated with the

peristome. Ovicell prominent, recumbent, the broad endooecium convex, smooth, the ectooecium laterally

calcified, frontally membranous.

Remarks. — When describing Batopora pulchrior , GORDON (1989a) suggested that a new genus might be

required for this and some other species that do not strictly conform to either Batopora Reuss or Lacrimula Cook.

A new genus is established here, therefore, for B. pulchrior , B. sp. (D’HONDT, 1986), and the following new

species. It differs from the type species of Batopora in being claviform and in lacking interzooidal kenozooids, and

from the type species of Lacrimula in lacking dimorphic orifices and perforated ovicells.

Ptoboroa is an anagram of Batopora.

Ptoboroa gelasina sp. nov.

Figs 205-207

MATERIAL EXAMINED. — New Caledonia. Biocal: stn KG 101, 1790 m.

Source : MNHN , Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

71

Types. — Holotype : unique colony from Biocal Stn KG 101, 21°26.51’S, 166°24.43'E, 1790 m, MNHN-

Bry 20038. No separate paratypes.

DESCRIPTION. — Sole colony claviform, comprising a club-like dilatation of autozooids tapering to a narrow

axial tube; colony length 1.28 mm, dilatation width 0.92 mm, internal diameter of axial tube 0.1 1 mm. Entire

colony surface dimpled, with reticulate patterning, zooidal boundaries not evident and areolar pores tiny, sporadic.

Primary orifice circular, the broad rounded poster occupying the proximal third, separated from the anter by a small

pair of condylar ridges angles proximally. Peristomial rim low, with a circular pore middistally. Adventitious

avicularia tiny, sparsely distributed, the rim nearly circular, the rostral and opesial areas separated by a pair of

mandibular pivots. Ovicells not present.

REMARKS. — The unique holotype specimen is very similar to the type species, Ptoboroa pulchrior

comb. nov. in most respects. One difference, apart from the dimpled colony surface, is that the axial tube

is simple and hollow, whereas in P. pulchrior it has internal radial partitions, the significance of which is

unknown.

Family CONESCHARELLINIDAE Levinsen, 1909

Genus CONESCHARELLINA d'Orbigny, 1852

TYPE Species. — Conescharellina angustata d'Orbigny, 1852.

Conescharellina breviconica Canu & Bassler, 1929

Figs 208-211, 214

Conescharellina breviconica Canu & Bassler, 1929: 491, pi. 69, figs 10-17, text-figs 206 G-H, 207 G.

MATERIAL EXAMINED. — Philippines. MUSORSTOM 3: stn CP 87, 191-197 m. — Stn CP 100, 189-199 m. —

•Stn CP 101, 194-196 m. — Stn CP 102, 192 m. — Stn CP 106, 640-668 m.

DESCRIPTION. — Colony conical, the cone becoming proportionately higher as the colony grows, such that in

young colonies the base exceeds the height by c. 15% and in old colonies by c. 10%; the largest colony 3.09 mm

diameter and 3.00 mm high. Zooidal orifices arranged quincuncially, their peristomes projecting only around the

periphery of the base where they appear as truncated cones. Colony surface with a light granular-tubercular texture,

the ridges between peristomes rounded, somewhat tubercular apically in small colonies. Zooidal dimensions as

measured at the flat colony base: c. 0.62 mm x 0.20 mm (small colony) to 0.77 mm x 0.40 mm (largest colony).

Primary orifice ranges from 0.10-1.66 mm x 0.084-0.13 (small colony) to 0.13-0.19 mm x 0.11-0.17 (largest

colony); shape suborbicular, the proximal margin broadly or roundly V-shaped, flanked by a small pair of condyles

that may be lacking in some colonies (presumably because of abrasion), set within a smooth thin peristomial rim.

Small sparse areolar pores occur between orifices. Avicularia suborbicular, occurring between orifices in

interzooidal areas, though not with consistent regularity, with a thin, near-median cross-bar that is often broken.

Colony base nearly flat, comprising an outer row of radially arranged autozooids surrounded a “cancellate”

(kenozooidal) area that is proportionately smaller (30% of diameter) in the smallest colonies than in large colonies

(42-56% of diameter).

Distribution. — Central Philippines between 10° and 14°N, 58-969 m.

Remarks. — This species has not been described since its introduction and so is redescribed here. Canu &

Bassler (1929) gave bottom temperatures for two of the ' Albatross " stations at which this species occurred,

citing 14.3°C at 216 m and 10.4°C at 969 m.

72

D. P. GORDON & J.-L. D'HONDT

Conescharellina catella Canu & Bassler, 1929

Figs 212-213, 215

Conescharellina catella Canu & Bassler, 1929: 485, pi. 68, figs 8-10, text-fig. 206 C-D. — HARMER, 1957: 732, pi. 47,

fig. 5, text-fig. 70 B.

Material EXAMINED. — Philippines. Musorstom 3: stn CP 100, 189-199 m. — Stn CP 102, 192 m. —

Stn DR 117, 92-97 m.

New Caledonia. Biogeocal: stn KG 316, 1660 m.

Distribution. — East of Korea, Sea of Japan, Philippines, Indonesia, 35-421 m; Loyalty Islands, 1660 m.

Remarks. — The present material matches very well the descriptions of Canu and Bassler (1929) and

Harmer (1957), except that the radial zooidal rows are slightly oblique in some of our specimens. Our material

typically also shows a slight ligulation of the avicularian cross-bar, not mentioned by the previous authors. The

largest colony in the collection (from MUSORSTOM 3 Stn CP 102) was 2.89 mm high and 2.64 mm basal

diameter. A colony from BIOGEOCAL Stn KG 316 occurred in much deeper water than previously recorded. It

resembles the other material except for the lack of avicularia between some of the orifices.

This species is one of several that have zooidal orifices opening along the longitudinal ridges (“costules”, in

the terminology of Canu and Bassler, 1929), separated by furrows in which there are avicularia. This species is

particularly characterised, however, by its offset ovicells that are produced to one side of the ridge and lie partly

across a furrow. This character is shared by other species, including C. striata Silen, 1947 and C. brevirostris

Sil6n, 1947. It is likely that ovicellular displacement is a generic-level character but, at this stage of our

knowledge, in the apparent absence of correlated characters, it is not possible to be certain. Ovicells are usually

lacking from conescharellinid colonies, but ovicellular displacement can be inferred from the position of the pore,

distal to the peristomial orifice, from which ovicells are produced, the pore also being displaced. Notwithstanding,

a colony from Biogeocal Stn KG 316 has some of these pores in a median position while others are displaced.

Conescharellina atalanta sp. nov.

Figs 216-218

Material examined. — New Caledonia. Musorstom 4: stn DW 149, 165 m. — Stn DW 150, 110 m.

Types. — Holotype: colony from Musorstom 4 Stn DW 149, 19°07.60’S, 163°22.70’E, 165 m, MNHN-Bry

20099, gold-coated.

Paratypes : 16 ethanol-preserved colonies from MUSORSTOM 4 Stn DW 150, 19°07.50’S, 163°22.10'E, 110 m,

MNHN-Bry 20019.

DESCRIPTION. — Colony a low cone, wider (2.59-3.45 mm) than high (1.43-2.28 mm), the zooidal orifices

distributed in subregular radial rows between which are avicularia. Orifices 0.066-0.104 mm diameter, distinctly

smaller towards the apex, with thin, fragile condyles defining a U-shaped sinus. Avicularia subcircular to ovoid

0.058-0.094 mm long, with thin, complete cross-bars; avicularia overlapping the autozooidal orifices in size such

that, in an eroded colony with orificial condyles and avicularian crossbars lacking, it may be difficult

discriminating between orifices and avicularia. Basal surface with an extensive cancellate area occupying 61-70 %

of the basal diameter, surrounded by a relatively narrow zone of autozooidal walls. Tiny oval avicularia 0.037 mm

long present inside the rims of several cancellae, the cross-bar complete.

Distribution. — New Caledonia, near Belep Islands, 1 10-165 m.

Remarks. The species name is derived from the Greek atalantos , equal in weight or equal to, alluding to

the near similarity in size of orifices and frontal avicularia.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

73

Key characters to look for in quickly discriminating this species from others of the genus are the overlap in

dimensions of zooidal orifices and frontal avicularia in combination with the large cancellate area basally.

Genus TROCHOSODON Canu & Bassler, 1927

Type Species. — Trochosodon linearis Canu & Bassler, 1927.

Trochosodon sp.

Figs 219-220

MATERIAL EXAMINED. — New Caledonia. Biogeocal: stn KG 275, 1959 m. — Stn CP 317, 1630-1620 m.

Description. — Colony small, comprising a small, somewhat flattened, central subdiscoidal area 1.11 mm

across (minimum colony diameter) composed primarily of 4 autozooids with concave peristomial openings;

between each pair of peristomes another, projecting, autozooid and peristome may occur, depending on the stage of

growth; width of zooidal projection at the point where it projects from the central area c. 0.37-0.43 mm. Entire

colony surface granular-tubercular. Primary orifice with a small U-shaped sinus delimited and flanked by a small

pair of condyles. Adapical surface with tiny rootlet pores and suborbicular avicularia, the latter 0.094 mm long

with complete cross-bars.

DISTRIBUTION. — New Caledonia: South Loyalty Basin, 1620-1959 m.

REMARKS. — This tiny species is probably new but additional material is needed before it can be properly

characterised.

Genus CRUCESCHARELLINA Silen, 1947

TYPE Species. — Crucescharellina japonica Silen, 1947.

Crucescharellina japonica Silen, 1947

Figs 221-223

Crucescharellina japonica Silen, 1947: 44, text-figs 28-31, pi 1, figs 11-12.

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn CP 106, 640-668 m.

Distribution. — Kyushu, Japan, 175 m. Central Philippines, 640-668 m.

REMARKS. — A single colony 7 mm maximum length occurred in the collection. This appears to be only

the second colony known of the species. It is well preserved and accords in every detail with SlLEN's (1947)

description and illustrations, except that the expanded ends of the branches are slightly less expanded than in

SlLEN’s specimen. The Japanese colony was obtained from a shallower depth, with a recorded bottom temperature

of 13.7°C.

Crucescharellina jugalis Gordon, 1989a, from New Zealand, differs primarily in lacking crescentic kenozooidal

openings (‘lunoecia’); the primary orifice is proportionately narrower, the avicularian cross-bars are slightly

thicker, and there are no elongate avicularia. Crucescharellina decussis (Canu & Bassler, 1929), Irom the

Philippines and Indonesia, has a more regular colony form with narrower branches, the mandibles of the large

avicularia are more spatulate, the peristomial margins can vary in shape, and lunoecia, if present, are rare.

74

D. P. GORDON & J.-L. D'HONDT

Crucescharellina aster sp. nov.

Figs 224-227

MATERIAL EXAMINED. — New Caledonia. BlOGEOCAL: stn KG 210, 1190 m. — Stn KG 211, 975 m. —

Stn CP 232, 760-790 m. — Stn CP 260, 1820-1980 m. — Stn KG 262, 1380 m. — Stn KG 267, 1935 m. — Stn CP 317,

1630-1620 m.

New Zealand. NZOI: stn F874, 1357 m. — Stn F879, 1267 m. — Stn U198, 1573 m.

Types. — Holotype: a large colony from Biogeocal Stn KG 267, 21°02.20'S, 166°58.76'E, 1935 m,

MNHN-Bry 20096.

Paratypes : two gold-coated colonies from BlOGEOCAL Stn CP 232, 21°33.81'S, 166°27.07'E, 760-790 m,

MNHN-Bry 20091. Three colonies from NZOI Stn F879, 37°25.50'S, 177°30.00'E, 1267 m, registration no.

P-1089.

Description. — Colony stellate, in the form of a 6-rayed star, the longest radius of which is 3.81 mm (tip of

"arm” to centre of colony); “arms” biserial, the zooids alternating; “arm” width at midlength 0.50-0.64 mm.

Colony surface granular-tubercular in the centre, smooth elsewhere. Zooid length 0.71-0.79 mm, measured from

peristome to peristome. Primary orifice and operculum suborbicular, c. 0.14 mm wide, with a median U-shaped

sinus defined and flanked by a pair of condyles. A median pore distally from which ovicells (not seen) presumably

bud. Small circular avicularia scattered sparsely on both colony surfaces, 0.037-0.047 mm diameter, with complete

cross-bars. Small rootlet pores of the same diameter present centrally on the “apical” surface.

Distribution. — New Caledonia, 760-1980 m. New Zealand, 1267-1573 m.

Remarks. — This is a very striking species. When large numbers occur in a sample the overall visual effect

is one of clusters of snowflakes. The regularly stellate colony form immediately distinguishes this species from

the others in the genus.

DISCUSSION

A total of 98 species of ascophorine bryozoans is herein recorded, mostly from the infraorder Lepraliomorpha

as defined by GORDON (1989a), of which 10 species occurred only at Philippine stations. The most speciose

families are: Phidoloporidae, 19 species; Smittinidae, 9 species; Celleporidae, 8 species; and Porinidae,

Siphonicytaridae and Conescharellinidae, 6 species each. The most speciose genera are Siphonicytara (6 species),

Characodoma , Iodictyum , and Reteporella (5 species each), and Haswelliporina and Reteporellina (4 species each).

Of the 84 species occurring in New Caledonian waters it is striking how many are endemic to the area - 57 species

(68%). Ten species are shared only with New Zealand (Parasmittina erecta , Oppiphorina epaxia , Haswelliporina

multiaviculata, Riscodopa parva , Gigantopora oropiscis , Onchoporoides moseleyi , Yrbozoon r ingens y Buffonellaria

regenerata , Galeopsis mimicus , Crucescharellina aster) (especially in deeper water and along the Kermadec Ridge),

4 species are shared only with the Philippines ( Hemismittoidea ennea , Haswelliporina quinaria , Reteporellina

spiramina , Conescharellina catella ), and 13 others have a wider Indo-Pacific distribution.

The diversity of Siphonicytara species is quite striking. Only 5 extant species of this distinctive genus have

been described previously: one Malagasy, one Philippine, the remainder Indonesian. The New Caledonian fauna

more than doubles the number of species.

ACKNOWLEDGEMENTS

Sincere thanks are due to the Smithsonian Institution, Washington, D.C. (J. THOMPSON), The Natural History

Museum, London (Mary SPENCER JONES), the Naturhistorisches Museum, Vienna (Dr Norbert VAvra), and the

South Australian Museum, Adelaide (the late Shane Parker) for loans of specimens that permitted comparisons

to be made with the New Caledonian and Philippine material.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

75

REFERENCES

Audouin, J.V., 1826. — Explication sommaire des planches de polypes de l’Egypte et de la Syrie, publics par Jules-

C6sar Savigny. Pp. 225-244. In : Description de I'Egypte, on recueil des observations et des recherches qui out ete

faites en Egypte pendant /’ Expedition de I’Armee frangaise ... Histoire naturelle. Tome 1, 4C partie. Imprimerie

Imp6riale, Paris. 339 p.

BASSLER, R.S., 1935. — Fossilium catalogus, I: Animalia, Pars 67 - Bryozoa. Generwn et genotyporum. Index et

bibliographica. W. Junk, Gravenhage. 229 p.

BASSLER, R.S., 1953. — Treatise on invertebrate paleontology. Part G. Bryozoa. Geological Society of America &

University of Kansas Press, New York, i-xiii + 253 p.

BOCK, P.E. & Cook, P.L., 1993. — Quadriscutella gen. nov. (Bryozoa, Cheilostomata) from the Tertiary and Recent of

Victoria and South Australia. Alcheringa , 17: 159-171.

BROWN, D.A., 1952. — The Tertiary Polyzoa of New Zealand. Trustees of the British Museum (Natural History), London,

vii-xii + 405 p.

Brown, D.A., 1958. — Fossil cheilostomatous Polyzoa from southwest Victoria. Memoirs of the Geological Survey of

Victoria, 20: 1-90.

Busk G , 1852. — An account of the Polyzoa, and sertularian zoophytes, collected in the voyage of the Rattlesnake, on

the coasts of Australia and the Louisiade Archipelago, &c. In: J. MacGillivray, Narrative of the voyage of H.M.S.

Rattlesnake, commanded by the late Captain Owen Stanley ... 1846-1850 ... T.W. Boone, London. Vol. 1: 343-402,

pi. 1 (Appendix No. IV).

BUSK, G., 1855. — Zoophytology. Quarterly Journal of Microscopical Science, 3: 320-324, pis 3-4.

BUSK, G., 1859. — A monograph of the fossil Polyzoa of the Crag. The Palscontographical Society, London. 136 p.

BUSK, G., 1881. — Descriptive catalogue of the species of Cellepora collected on the “Challenger’ Expedition. Journal of

the Linnean Society , Zoology, 15: 341-356.

Busk, G., 1884. — Report on the Polyzoa collected by H.M.S. Challenger during the years 1873-76. Part 1. - The

Cheilostomata. Report on the Scientific Results of the Voyage of H.M.S. Challenger, Zoology, 10 (30): i-xxiv,

1-216, pis 1-36.

Canu, F., 1913. — Etudes morphologiques sur trois nouvelles families de Bryozoaires. Bulletin de la Societe Geologique

de France, s6r. 4, 13: 132-147.

Canu, F., 1917. — Bryozoaires. Revue critique de Paleozoologie, 21: 29-37.

Canu, F. & BASSLER, R.S., 1917. — A synopsis of American Early Tertiary cheilostome Bryozoa. Bulletin of the United

States National Museum, 96: 1-87, pis 1-6.

Canu, F. & Bassler, R.S., 1920. — North American Early Tertiary Bryozoa. Bulletin of the United States National

Museum, 106: (Vol. 1) i-xx, 1-879, (Vol. 2) 1-162 pis.

Canu, F. & Bassler, R.S., 1923. — North American later Tertiary and Quaternary Bryozoa. Bulletin of the United States

National Museum, 125: i-viii, 1-302, pis 1-47.

Canu, F. & BASSLER, R.S., 1927. — Classification of the cheilostomatous Bryozoa. Proceedings of the United States

National Museum, 69 (14): 1-42, pi. 1.

Canu, F. & BASSLER, R.S., 1928. — Fossil and Recent Bryozoa of the Gulf of Mexico region. Proceedings of the United

States National Museum, 12 (14): 1-199, pis 1-34.

Canu, F. & Bassler, R.S., 1929. — Bryozoa of the Philippine region. Bulletin of the United States National Museum,

100: i-xi, 1-685, pis 1-94.

Canu, F. & Bassler, R.S., 1930. — The bryozoan fauna of the Galapagos Islands. Proceedings of the United States

National Museum, 76 (13): 1-78, pis 1-14.

Cheetham, A.H. & Sandberg, P.A., 1964. — Quaternary Bryozoa from Louisiana mudlumps. Journal of Paleontology,

38:1013-1046.

Cook, P.L., 1966. — Some “sand fauna” Polyzoa (Bryozoa) from eastern Africa and the northern Indian Ocean. Cahiers de

Biologie marine, 7: 207-223, pis 1-2.

76

D. P. GORDON & J.-L. D’HONDT

Cook, P.L., 1985. — Bryozoa from Ghana: a preliminary survey. Annales du Musee Royal de I'Afrique Centrale (Sciences

zoologiques), 238: 1-315.

COOK, P.L. & BOCK, P.E., 1996. — Characodoma Maplestone, a senior synonym of Cleidochasma Harmer

(Cleidochasmatidae). Pp. 81-88. In: D.P. Gordon, A.M. Smith, & J.A. Grant-Mackie (eds), Bnozoans in space and

time. National Institute of Water & Atmospheric Research, Wellington. 442 p.

Cook, P.L. & Chimonides, P.J.. 1981. — Morphology and systematics of some rooted cheilostome Bryozoa. Journal of

Natural History , 15: 97-134.

Cook, P.L. & Lagaaij, R.A., 1976. — Some Tertiary and Recent conescharelliniform Bryozoa. Bulletin of the British

Museum ( Natural History), Zoology , 29: 317-376, pis 1-8.

David, L. & Pouyet, S., 1986. — Bryozoaires abyssaux des campagnes Safari (Ocean Indien). Annales de I'Institut

oceanographique , n. s6r., 62: 141-191.

DUMONT, J.P.C., 1981. — A report on the cheilostome Bryozoa of the Sudanese Red Sea. Journal of Natural History , 15:

623-637.

Duvergier, J., 1921. — Note sur les Bryozoaires du N6ogfcne de T Aquitaine. Actes de la Societe Linneenne de Bordeaux ,

72: 145-181, pis 1-4.

Duvergier, I, 1924. — Deuxteme note sur les Bryozoaires du N6ogfcne de l’Aquitaine. Actes de la Societe Linneenne de

Bordeaux, 75: 145-190, pis 1-6.

Fransen, C.H.J.M., 1986. — Caribbean Bryozoa: Anasca and Ascophora Imperfecta of the inner bays of Curasao and

Bonaire. Studies on the Fauna of Curasao and other Caribbean Islands , 68: 1-119.

Gabb, W.M. & Horn, G.H., 1862. — The fossil Polyzoa of the Secondary and Tertiary Formations of North America.

Journal of the Academy of Natural Sciences of Philadelphia , 5: 1 1 1-179, pis 19-21.

GOLDSTEIN, J.R.Y., 1882. — Some new species of Bryozoa from the Marion Islands, with notes on Bicellaria grandis.

Transactions and Proceedings of the Royal Society of Victoria , 18: 39-46, pis 1-2.

Gordon, D.P., 1984. — The marine fauna of New Zealand: Bryozoa: Gymnolaemata from the Kcrmadec Ridge. Memoirs.

New Zealand Oceanographic Institute , 91: 1-198.

Gordon, D.P., 1985. — Additional species and records of gymnolaemate Bryozoa from the Kermadec region. NZOI

Records, 4: 159-183.

Gordon, D.P., 1988. — The bryozoan families Sclerodomidae, Bifaxariidae, and Urceoliporidae and a novel type of

frontal wall. New Zealand Journal of Zoology, 15: 249-290.

Gordon, D.P., 1989a. — The marine fauna of New Zealand: Bryozoa: Gymnolaemata (Cheilostomida Ascophorina) from

the western South Island continental shelf and slope. Memoirs. New Zealand Oceanographic Institute, 97: 1-158.

Gordon, D.P., 1989b. — New and little-known genera of cheilostome Bryozoa from the New Zealand region. Journal of

Natural History, 23: 1319-1339.

Gordon, D.P., 1989c. — New and little-known deep-sea taxa of umbonulomorph Bryozoa and their classification. New

Zealand Journal of Zoology , 16: 215-267.

GORDON, D.P., 1990. — The Tertiary bryozoan family Prostomariidae - morphology and relationships. Memoirs of the

Museum of Victoria , 50: 467-472.

Gordon, D.P., 1993a. — Bryozoa: The ascophorine infraorders Cribriomorpha, Hippothoomorpha and Umbonulo-

morpha. In: A. Crosnier (ed.), R6sultats des Campagnes Musorstom, Vol. 11. Memoires du Museum national

d’Histoire naturelle, 158. 299-347.

Gordon, D.P., 1993b. — Bryozoan frontal shields: studies on umbonulomorphs and impacts on classification.

Zoologica Scripta,22: 203-221.

Gordon, D.P., 1994. — Tertiary bryozoan genera in the present-day Australasian fauna - implications for classification

and biogeography. Invertebrate Taxonomy, 8: 283-298.

Gordon, D.P. & Braga, G., 1994. — Bryozoa: Living and fossil species of the catenicellid subfamilies Ditaxiporinae

Stach and Vasignyellinae nov. In: A. Crosnier (ed.), R6sultats des Campagnes Musorstom, Vol. 12. Memoires du

Museum national d’Histoire naturelle, 161: 55-85.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

77

Cordon D.P. & Hondt, J.-L. d\ 1991. — Bryozoa: The Miocene to Recent family Petalostegidae. Systematics,

affinities, biogeography. In A. CROSNIER (ed.), R6sultats des Campagnes MUSORSTOM, Vol. 8. Memoires du Museum

national d’Histoire naturelle, 151: 91-123.

Gordon, D.P. & VOIGT, E., 1996. — The kenozooidal origin of the ascophorine hypostegal coelom and associated frontal

shield: 89-107 In:: D.P. GORDON, A.M. Smith, & J.A. Grant-Mackie (eds), Bryozoans in space and time. National

Institute of Water & Atmospheric Research, Wellington. 442 p.

HARMER, S.F., 1902. — On the morphology of the Cheilostomata. Quarterly Journal of Microscopical Science , n. s., 46:

263-350, pis 15-18.

HARMER, S.F., 1926. — The Polyzoa of the Siboga Expedition. Part 2. Cheilostomata Anasca. Siboga-Expeditie , 28b:

i-viii, 181-501, pis 13-34.

Harmer, S.F., 1933. — The genera of Reteporidae. Proceedings of the Zoological Society, London, 1933: 615-627.

HARMER, S.F., 1934. — The Polyzoa of the Siboga Expedition. Part 3. Cheilostomata Ascophora. I. Family Reteporidae.

Siboga-Expeditie, 28c: i-vii, 503-640, pis 35-41.

HARMER, S.F., 1957. — The Polyzoa of the Siboga Expedition. Part 4. Cheilostomata Ascophora. II. Siboga-Expeditie,

28d: i-xv, 641-1147, pis 42-74.

HaSSALL, A.H., 1842. — Remarks on the genus Lepralia of Dr. Johnston with descriptions of six undescribed species.

Annals and Magazine of Natural History, 9: 407-414.

Haswell, W.A., 1881. — On some Polyzoa from the Queensland coast. Proceedings of the Linnean Society of New South

Wales, S: 33-44, pis 1-3.

Hayward, P.J., 1988. — Mauritian cheilostome Bryozoa. Journal of Zoology, London, 215: 269-356.

Hayward, P.J., 1991. — Systematic studies on some Antarctic and sub-Antarctic Ascophora (Bryozoa: Cheilostomata).

Zoological Journal of the Linnean Society, 101: 299-335.

Hayward, P.J., 1992. — Some Antarctic and sub-Antarctic species of Celleporidae (Bryozoa, Cheilostomata). Journal of

Zoology, London, 226: 283-310.

Hayward, P.J. & Cook, P.L., 1983. — The South African Museum’s Meiring Naude cruises. Part 13. Bryozoa II. Annals

of the South African Museum, 91: 1-161.

Hayward, P.J. & Thorpe, J.P., 1988. — A new family of cheilostome Bryozoa endemic to Antarctica. Zoological

Journal of the Linnean Society, 93 : 1-18.

Hayward, P.J. & Ryland, J.S., 1979. — British ascophoran bryozoans. Keys and notes for the identilication of the

species. Synopses of the British Fauna, n. s., 14: 1-312.

HlNCKS, T., 1877. _ On British Polyzoa, Part II. Classification. Annals and Magazine of Natural History', ser. 4, 20:

520-532.

HlNCKS, T., 1879. — On the classification of the British Polyzoa. Annals and Magazine of Natural History', ser. 5, 3:

153-164.

HlNCKS, T., 1880. — A history of the British marine Polyzoa. Van Voorst, London. Vol. 1, cxli, 601 p., Vol. 2, 83 pis.

Hincks, T., 1895. — Index [to Marine Polyzoa: Contributions towards a general history]. London, vi p. [issued

privately.]

Hondt J -L D’ 1981. — Bryozoaires Cheilostomes bathyaux et abyssaux provenant des campagnes oc6anographiques

am£ricaines (1969-1972) de ‘T Atlantis II”, du “Chain” et du “Knorr” (Woods Hole Oceanographic Institution).

Bulletin du Museum national d'Histoire naturelle, s6r. 4, 3, A (1): 5-71.

Hondt, J.-L. d\ 1985. — Contribution & la syst^matique des Bryozoaires Eurystomes. Apports nkents et nouvelles

propositions. Annales des Sciences naturelles, Zoologie s6r. 13, 7: 1-12.

HONDT, J.-L. D\ 1986. - Bryozoaires de Nouvelle-Caledonie et du Plateau des Chesterfield. Bulletin du Museum national

d’Histoire naturelle, ser. 4, 8, A (4): 697-756.

JELLY, E.C., 1889. — A synonymic catalogue of the recent marine Bryozoa. Dulau & Co., London, xv, 322 p.

JOHNSTON, G., 1838. — A history of the British zoophytes. W.H. Lizars, Edinburgh, London & Dublin, xii, 341 p.,

44 pis.

78

D. P. GORDON & J.-L. D'HONDT

Johnston, G., 1847. — A history • of the British zoophytes . 2nd edn. Van Voorst, London. Vol. 1, xvi, 488 p., Vol. 2,

74 pis.

Jullien, J., 1883. — Dragages du Travailleur. Bryozoaires. Espfcces dragu6es dans l’ocean Atlantique en 1881. Bulletin de

la Societe Zoologique de France , 7: 497-529, pis 13-17.

JULLIEN, J., 1888. — Bryozoaires. Mission scientifique du Cap Horn , 6 (Zoologie 3): 1-92, pis 1-15.

Jullien, J. & Calvet, L., 1903. — Bryozoaires provenant des campagnes de l’Hirondelle (1886-1888). Resultats des

Campagnes scientifiques accomplies par le Prince Albert /, 23: 1-188, pis 1-18.

Kirkpatrick, R., 1888. — Polyzoa of Mauritius. Annals and Magazine of Natural History , ser. 6, 1: 72-85, pis 7-10.

Keferstein, W., 1868. — Ueber die Batrachier Australiens. Archiv fur Naturgeschichte, 34: 253-290, pis 1-4.

Lagaaij, R., 1963. — New additions to the bryozoan fauna of the Gulf of Mexico. Publications of the Institute of Marine

Science, University of Texas, 9: 162-236.

Lamarck, J.B.P.A., 1815-22. — Histoire naturelle des animaux sans vertebres ... precedee d'une introduction offrant la

determination des caracteres essentiels de l 'animal, sa distinction du vegetal et des autres corps naturels, enfin,

l’ exposition des principes fondamentaux de la zoologie. Verdure, Paris. [In 7 vols, Bryozoa in vol. 2 (1816), 568 p.]

LAMOUROUX, J.V.F., 1821. — Exposition methodique des genres de I’ordre des polypiers, avec leur description et celles

des principales especes figurees dans 84 planches; les 63 premieres appartenant a V Histoire naturelle des Zoophytes

d’ Ellis et Solander. V. Agasse, Paris, viii, 115 p., 84 pis.

Levinsen, G.M.R., 1909. — Morphological and systematic studies on the cheilostomatous Bryozoa. National Forfatteres

Forlag, Copenhagen, i-vii + 431 p., 24 pis.

Levinsen, G.M.R., 1917. — Bryozoa. In: Danmark-Ekspeditionen til Grpnlands Nordpstkyst, 1906-1908, III.

Meddelelser orn Gr0nland, 43: 432-472, pis 19-24.

Livingstone, A. A., 1926. — Studies on Australian Bryozoa. No. 3. Report upon the Bryozoa collected on the Great

Barrier Reef, Queensland, in 1925, by W.E.J. Paradice, Lieutenant Surgeon on H.M.A.S. “Geranium”. Records of the

Australian Museum , 15: 79-99.

MacGillivray, J., 1842. — Catalogue of the marine zoophytes of the neighbourhood of Aberdeen. Annals and Magazine

of Natural History , 9: 462-469.

MacGillivray, P.H., 1860. — Notes on the cheilostomatous Polyzoa of Victoria and other parts of Australia.

Transactions of the Philosophical Institute of Victoria , 4: 159-168, pis 2-3.

MacGillivray, P.H., 1869. — Descriptions of some new genera and species of Australian Polyzoa; to which is added a

list of species found in Victoria. Transactions and Proceedings of the Royal Society of Victoria, 9: 126-148.

MacGillivray, P.H., 1887. — A catalogue of the marine Polyzoa of Victoria. Transactions and Proceedings of the Royal

Society of Victoria , 23: 187-224.

MacGillivray, P.H., 1880. — [Polyzoa.]. In: F. McCoy (ed.), Prodromus of the Zoology of Victoria. Vol. 1, Decade 4:

21-40, pis 35-38; Decade 5: 27-52, pis 41-50. Government Printer, Melbourne.

MacGillivray, P.H., 1883. — Descriptions of new, or little-known, Polyzoa. Part II. Transactions and Proceedings of

the Royal Society of Victoria , 19: 130-138, pis 1-3.

MacGillivray, P.H., 1895. — A monograph of the Tertiary Polyzoa of Victoria. Transactions of the Royal Society • of

Victoria , n. s., 4: 1-166, pis 1-22.

Maplestone, C.M., 1900. — Further descriptions of the Tertiary Polyzoa of Victoria. - Part 4. Proceedings of the Royal

Society of Victoria, n. s., 13: 1-9, pis 1-2.

Moyano, G.H.I., 1985. — Briozoos marinos chilenos V. Taxa nuevos o poco conocidos. Boletin de la Sociedad de

Biologia de Concepcion, 56: 79-114.

Moyano, G.H.I., 1991. — Bryozoa marinos chilenos VIII: Una smtesis zoogeogr£fica con consideraciones sistem£ticas

y la descripci6n de diez especies y dos g^neros nuevos. Gayana Zoologia, 55: 305-389.

Neviani, A., 1895. — Briozoi neozoici di alcune locality dTtalia. 1. Bolletino della Societa Romana per gli studi

Zoologici, 4:109-123.

Neviani, A., 1900. — Briozoi neogenici delle Calabrie. Palaeontographica italica, 6: 1 15-266, pis 16-19.

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

79

NORMAN, A.M., 1868. — Notes on some rare British Polyzoa, with descriptions of new species. Quarterly Journal of

Microscopical Science , n. s., 8: 212-222, pis 5-7.

Norman, A.M., 1876. — Crustacea, Tunicata, Polyzoa, Echinodermata, Actinozoa, Foraminifera, Polycystina, and

Spongida. Pp. 101-1 15. In: J.G. Jeffreys, Preliminary report of the biological results of a cruise in H.M.S. Valourous

to Davis Strait in 1875. Proceedings of the Royal Society of London , 25: 177-237.

Norman, A.M., 1903. — Notes on the natural history of East Finmark. Polyzoa. Annals and Magazine of Natural History ,

ser. 7, 11: 567-598, pis 1-13.

Okada, Y., 1921. — Notes of some Japanese chilostomatous Bryozoa. Annotationes zoologicae japonenses. 10: 19-32.

Okada, Y., 1929. — Cheilostomatous Bryozoa of Mutsu. Scientific Reports of the Tohoku University, ser. 4 (Biology),

4: 11-35.

ORBIGNY, A.D. d\ 1835-47. — Voyage dans I'Amerique meridionale ... execute pendant ... 1826-33 par A.D. d'Orhigny,

&c. P. Bertrand, Paris; V. Levrault, Strasbourg. 7 Tom. + Atlas. [Zoophytes in Tom. V, 4, pp. 7-28, pis 1-13. Text,

pp. 7-28, Livr. 89 (1847); pis 1. 3, 5, Livr. 56 (1841); pis 2, 6-8, Livr. 57 (1842); pis 4. 9, 10, 12, 13, Livr. 58

(1842); pi. 11. Livr. 59 (1842). Dates as given by Sherborn & Woodward, 1901.]

ORBIGNY, A.D. D\ 1851-54. — Paleontologie fran^aise. Descriptions des Mollusques et Rayonnes fossiles. Terrains

Cretaces, V. Bryozoaires. Victor Masson, Paris. [Pp. 1-188 (1851); pp. 185 bis-472 (1852); pp. 473-984 (1853);

pp. 985-1192 (1854); pis 600-800. Dates as given by Sherborn, 1899.]

Ortmann, A., 1890. — Die japanische Bryozoenfauna. Bericht iiber die von Herrn Dr. L. Doderlein in Jahre 1880-81

gemachten Sammlungen. Archiv fiir Naturgeschichte, 54: 1-74, pis 1-4.

OSBURN, R.C., 1952. — Bryozoa of the Pacific coast of America Part 2, Cheilostomata-Ascophora. Allan Hancock Pacific

Expeditions, 14: 271-611.

Pallas, P., 1766. — Elenchus zoophytorum sistens generum adumbrationes generaliores et specierum cognitarum

succinctas descriptiones cum selectis auctorus synonymis. Pelrum van Cleef, Hagae-Comitum. i-xxviii +451 p.

Parker, S.A. & Gordon, D.P., 1992. — A new genus of the bryozoan superfamily Schizoporelloidea, with remarks on

the validity of the family Lacernidae Jullien, 1888. Records of the South Australian Museum, 26: 67-71.

Philipps, E.G., 1900. — Report on the Polyzoa collected by Dr. Willey from the Loyalty Isles, New Guinea and New

Britain. Arthur Willey’s Zoological Results, 4: 357-530, pis 34-53.

Powell, N.A., 1967. — Polyzoa (Bryozoa) - Ascophora - from north New Zealand. Discovery Reports, 34: 199-393,

pis 1-17.

Rho, B.J. & Seo, J.E., 1986. — A systematic study on the marine bryozoans in Cheju-do. Korean Journal of Zoology,

29: 31-60.

Ridley, S.O., 1881. — Polyzoa. In: Account of the zoological collections made during the survey of H.M. S. Alert in the

Straits of Magellan and on the coast of Patagonia. Proceedings of the Zoological Society of London, 1881: 44-61,

pi. 6.

ROGICK, M.D., 1960. — Studies on marine Bryozoa. 13. Two new genera and new species from Antarctica. The Biological

Bulletin, 119: 479-493.

RYLAND, J.S. & Hayward, P.J., 1992. — Bryozoa from Heron Island, Great Barrier Reef. Memoirs of the Queensland

Museum, 32: 223-301.

Sherborn, C.D., 1899. — On the dates of the ‘‘Pal6ontologie Fran$aise” of d’Orbigny. Geological Magazine, ser. 4, 6:

223-225.

Sherborn, C.D. & Woodward, BB., 1901. — Notes on the dates of publication of the natural history portions of some

French voyages. - Part I. 'AmSrique meridionale’; ‘Indes orientales’; ‘Pole Sud’ (‘Astrolabe’ and ‘Z616e’); ‘La Bonite’;

‘La Coquille’; and TUranie et Physicienne’. Annals and Magazine of Natural History', ser. 7, 7: 388-392.

Sil£n, L., 1941. — Cheilostomata Anasca (Bryozoa) collected by Prof. Dr. Sixten Bock’s expedition to Japan and the

Bonin Islands 1914. Arkiv for Zoologi, 33, A (12): 1-130, pis 1-9.

Sil£n, L., 1947. — Conescharellinidae (Bryozoa Gymnolaemata) collected by Prof. Dr. Sixten Bock’s expedition to

Japan and the Bonin Islands 1914. Arkiv for Zoologi, 39, A (9): 1-61, pis 1-5.

80

D. P. GORDON & J.-L. D’HONDT

Smitt, F.A., 1873. — Floridan Bryozoa, collected by Count L.F. de Pourtales. Part II. Kongliga Svenska Vetenskaps-

Akademiens Handlingar, 11 (4): 1-83, pis 1-13.

Soule, D.F. & Soule, J.D., 1973. — Morphology and speciation of Hawaiian and eastern Pacific Smittinidae (Bryozoa,

Ectoprocta). Bulletin of the American Museum of Natural History , 152: 365-440.

Soule, J.D., Soule, D.F. & Chaney, H.W., 1991. — New tropical Pacific and Indian Ocean Cleidochasmatidae

(Cheilostomata: Ascophora). In: F.P. Bigey & J.-L. D'HONDT (eds), Bryozoaires actuels et fossiles: Bryozoa living and

fossil. Bulletin de la Societe des Sciences Naturelles de I’Ouest de la France , M£moire h. s. 1: 465-486.

STACH, L.W., 1935. — The genera of Catenicellidae. Proceedings of the Royal Society of Victoria , n. s., 47: 389-396.

StaCH, L.W., 1936. — Studies on Recent Petraliidae (Bryozoa). Records of the Australian Museum, 19: 355-379.

Stoliczka, F., 1865. — Palaontologie von Neu-Zeeland. Beitrage zur Kenntniss der fossilen Flora und Fauna der

Provinzen Auckland und Nelson. IV. Fossile Bryozoen aus dem tertiaren Griinsandsteine der Orakei-Bay bei Auckland.

Reise der Osterreichischen Fregatte Novara, Geologisher Theil, 1 (2): 89-158, pis 17-20.

Strand, E., 1928. — Miscellanea nomenclatdrica zoologica et paleontologica. Arkiv fur Naturgeschichte , 92, A (8):

36-37.

TENISON- WOODS, J.E., 1879. — On some Australian Tertiary fossil corals and Polyzoa. Journal and Proceedings of the

Royal Society of New South Wales, 12: 57-61, 1 pi.

Thomson, W.T.C., 1858. — On new genera and species of Polyzoa from the collection of Professor W.H. Harvey, Trinity

College, Dublin. Part 1. Proceedings of the Dublin University Zoological and Botanical Association, 1: 77-93.

Thornely, L.R., 1905. — Report on the Polyzoa collected by Professor Herdman, at Ceylon, in 1902. Report to the

Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, 4 (Suppl. Rep. 26): 107-130, pi. 1.

Uttley, G.H., 1956. — Remarks on the bryozoan genera Spiroporina Stoliczka and Haswellina Livingstone and their

genotypes. Transactions of the Royal Society of New Zealand, 84: 29-31.

Uttley, G.H. & Bullivant, J.S., 1972. — Biological results of the Chatham Islands 1954 Expedition. Part 7. Bryozoa

Cheilostomata. Memoirs. New Zealand Oceanographic Institute, 57: 1-61. [Bulletin of the New Zealand Department

of Scientific and Industrial Research, 139 (7).]

Vigneaux, M., 1949. — Revision des Bryozoaires n6og£nes du Bassin d’ Aquitaine et essai de classification. Memoires de

la Societe Geologique de France, n. s., 28: 1-153, pis 1-11.

Voigt, E. & Hillmer, G., 1983. — Oberkretazische Hippothoidae (Bryozoa Cheilostomata) aus dem Campanium von

Schweden und dem Maastrichtium der Niederlande. Mitteilungen aus dem Geologisch-Paldontologisches Institut der

Universitdt Hamburg, 54: 169-208.

Waters, A.W., 1881. — On fossil chilostomatous Bryozoa from south-west Victoria, Australia. Quarterly Journal of the

Geological Society of London, 37: 309-347, pis 14-18.

Waters, A.W., 1889. — Supplementary report on the Polyzoa collected by H.M.S. Challenger during the years 1873-

1876. Report of the Scientific Results of the Voyage of H.M.S. Challenger, Zoology, 31 (79): 1-41, pis 1-3.

Waters, A.W., 1899. — Bryozoa from Madeira. Journal of the Royal Microscopical Society, 1899: 6-16, pi. 3.

Waters, A.W., 1913. — The marine fauna of British East Africa and Zanzibar, from collections made by Cyril Crossland,

M.A., B.Sc., F.Z.S., in the years 1901-1902. Bryozoa-Cheilostomata. Proceedings of the Zoological Society,

London, 1913: 458-537, pis 64-73.

Winston, J.E., 1984. — Shallow-water bryozoans of Carrie Bow Cay, Belize. American Museum Novitates, 2799: 1-38.

Winston, J.E., 1986. — An annotated checklist of coral-associated bryozoans. American Museum Novitates, 2859:

1-39.

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

81

Fig. 1. — Indeterminate cribrilinid ancestrulae, partially overgrown by zooids of Buffonellodes crosnieri sp. nov. (x 47).

Biocal Stn DW 33.

FlGS 2-3. — Domosclerus sp.: 2, Lateral view of small colony (x 28). 3, Same (x 1 12). BlOGEOCAL Sin KG 210.

FIGS 4-6. — Bryosartor Isutilis Gordon & Braga: 4, Single broken segment (x 53). 5, Single zooid (x 140). 6. Costal

field of a different zooid (x 496). Biogeocal Stn CP 260.

Fig. 7. — Hippothoa calciophilia Gordon (x 153). Biocal Stn CP 84.

Source : MNHN, Paris

82

D. P. GORDON & J.-L. D'HONDT

Figs 8-9. - Xynexecha pulchra sp. nov.: 8, Base of colony, showing kenozooids and the beginning of an erect

cylindrical stem (x 94). 9, A single zooid, with its distinctive frontal-shield morphology and spout-like peristome

(x 179). BiOCALStnCP 108.

Figs 10-1 1. — Kladapheles gammadeka Gordon: 10, Branching colony fragment (x 25). 11, Oral view of orifices and an

avicularium (x 103). Biogeocal Stn DW 307.

Source : MNHN, Paris

FlG. 12. — Hippothyris caledonica sp. nov. (x 70). BlOCAL Stn DW 82.

Figs 13-14. — Parkermavella fidelis sp. nov.: 13, Ovicelled zooid (x 52). 14, Autozooidal orifice (x 265). MUSORSTOM 6

Stn CP 419.

FlG. 15. — Parkermavella minuta sp. nov.: Several zooids; ovicell at lower right (x 134). Biocal Stn DW 65.

Figs 16-17. — Smittoidea maunganuiensis multiporosa ssp. nov.: 16, Several zooids, one ovicelled (x 56). 17, Oral

view of avicularium (x 215). Musorstom 6 Stn CP 419.

Source : MNHN, Paris

84

D. F. GORDON & J.-L. D'HONDT

FIGS 18-19. — Hemismittoidea ennea sp. nov.: 18, Ovicelled zooids (x 98). 19, Primary orifice and lyrula of ovicelled

zooid (x 289). Biocal Stn DW 66.

Fig. 20. — Hemismittoidea 1 ennea: Part of infertile colony (x 98). MUSORSTOM 3 Stn DR 117.

Figs 21-22. — Parasmittina glabra sp. nov.: 21, Ovicelled zooid (x 137). 22, Primary orifice and lyrula (x 289). Biocal

Stn DW 66.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

85

Figs 23-24. — Parasmittina ? glabra: 23, Autozooids showing disposition of avicularia (x 103). 24, Primary orifice with

lyrula (x 347). Biocal Stn DW 66.

FIGS 25-26. — Parasmittina marginata (Canu & Bassler): 25, Autozooids showing characteristically curved aviculana

(x 88). 26, Ovicelled zooid (x 109). Musorstom 3 Stn DR 1 17.

Source :

86

D. P. GORDON & J.-L. D'HONDT

Fig. 27. — Parasmittina serrula Soule & Soule: Autozooids and ovicelled zooids (x 107). Musorstom 3 Stn DR 117.

Figs 28-29. — Parasmittina erecta sp. nov. 28. Branching fragment (x 19). 29, Group of zooids proximal to the lower

bifurcation in fig. 28 (x 76). Biocal Stn DW 66.

Figs 30-31. — Parasmittina sp.: 30, Oral view of autozooids (x 149). 31, Orifice (x 370). Musorstom 3 Stn DR 117.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

87

Figs 32-34. — Smittina asymmetrica sp. nov.: 32, Ovicelled zooid and autozooids (x 54). 33. Close-up of another

ovicell (x 97). 34, Orifice of ovicelled zooid showing a pair of spine bases (normally concealed), part of broken

ovicell (at right), lyrula, and suboral avicularium (x 221). Biocal Stn DW 66.

Figs 35-36. — Smittina abyssicola (Harmer): 35. Several autozooids (x 72). 36, Close-up of suboral avicularium with

ligulate cross-bar (x 304). Biocal Stn CP 84.

Source :

88

D. P. GORDON & J.-L. D’HONDT

Res 37-41. — ' Mawatarius secundus sp. nov.: 37, Branching fragment (x 26). 38. Peristomial region of an ovicelled

(ov) zooid (x 109). 39. Ancestrula region of colony (x 61). 40. Peristomial orifice (x 221). 41. Lyrula of primary

orifice (x 243). Biocal Stn DW 66. F 3

FlG_ MucroPetra^e^a philippinensis (Canu & Bassler): Primary orifices of two autozooids (x 54). Biocal

Stn KG 22.

Fig. 43. — Mucropeiraliella serraia (Livingstone): Partly eroded autozooids (x 47). MUSORSTOM 4 Stn DW 151.

Source : MNHN, Paris

BRYOZOA ASCOPHOR1NA FROM NEW CALEDONIA 89

Fig. 44. — Hippomenella avicularis (Livingstone): Several autozooids, two with spatulate avicularia (x 52).

Musorstom 4 Stn DW 231.

Figs 45-46. — Emballotheca rara sp. nov.: 45, Whole colony (x 13). 46, Autozooids and ovicell (x 74). Biocal

Stn DW 46.

Figs 47-48. — Calyptotheca sp.: 47, Tiny colony on a branch of the cheilostomate bryozoan Nellia tenella (Lamarck)

(x 47). 48, Same, close-up of some autozooids (x 156). MUSORSTOM 4 Stn DW 187.

Source : MNHN, Paris

90

D. P. GORDON & J.-L. D HONDT

Figs 49-52. — Tetraplaria orospinea sp. nov.: 49, Proximal end of a segment (x 34). 50, Autozooid and orifices with

spine bases (x 52). 51, Encrusting zooids at colony base (x 54). 52, Autozooidal orifice with spine bases (x 215).

Biogeocal Stn CP 232.

Figs 53-54. — Tetraplaria ventricosa (Haswell): 53, Segment of opposite and decussate zooids (x 75). 54, Orifice

(x 311). Musorstom 3 Stn DR 117.

Figs 55-56. — Tetraplaria sp.: 55, Segment (x 13). 56. Arrangement of zooids in segment (x 78). Biocal

Stn DW 38.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

91

Figs 57-59. —Phorioppnia cookae sp. nov.: 57, Part of erect stem (x 13). 58. Ovicelled zooid and skewed distal

autozooid (x 44). 59, Autozooidal orifice and profile of ovicell (x 105). Biogeocal Stn DW 253.

FIGS 60-61. — Phorioppnia nova sp. nov.: 60, Part of erect stem (x 13). 61, Close-up of zooids (x 56). Biocal

FlGS62D64. — Oppiphorina epaxia (Gordon): 62, Part of erect stem (x 13). 63, Autozooids and profiles ol two ovicells

(x 62). 64, Ovicelled orifice (x 145). Biocal Stn DW 66.

Source : MNHN, Paris

92

D. P. GORDON & J.-L. D'HONDT

Fig. 65. — Genus and species indet.. Pan of erect stem (x 101). Biogeocal Stn KG 275

Fl°B iocALStnCP75POr‘na mul,iavicula,a <Gordon): Prof'le of zooidal peristomes; spiramen tubes arrowed (x 78).

FlGStnDW ~"aswelliporina vaubani <d’Ho"d»: 67. Part of erect stem (x 18). 68. Stem apex (x 56). Musorstom 4

Fig. 69. Haswelliporina quinaria sp. nov.: Profile of peristome and spiramen tube (x 94). Musorstom 6 Stn CP 419,

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

93

Fig. 70. — Haswelliporina quinaria sp. nov.: Distal part of stem, the lowest whorl of zooids ovicelled (x 55).

Musorstom 6 Stn CP 419.

FIGS 71-73. — Haswelliporina Ivenusta (Harmer): 71, Part of erect stem (x 13). 72, Lateral view of peristome (x 106).

73, Frontal view of peristome, spiramen, and two sizes of avicularia (x 106). ESTASE 2 Stn DR 07.

Source : MNHN. Paris

94

D. P. GORDON & J.-L. D'HONDT

FlGo 74'I5,’ T Mosaic°P°rina uni serialis sp. nov.: 74. Portion of colony (x 27). 75, Individual zooid (x 86). BlOCAL

Mn DW 66.

FlGc 76nw ™ Semihaswellia umbrella sp. nov.: 76, Disposition of zooids (x 29). 77. Ovicell (ov) (x 56). Biocal

Mn D W Uo .

Source MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

95

Figs 78-81. — Actisecos regularis Canu & Bassler: 78, Whole colony (x 16). 79, Ancestrula and periancestrular zooids

(x 58). 80, Primary orifice and ovicell (x 131). 81, Laterobasal view of ovicells and distal pore-chambers (x 81).

Musorstom 3 Stn DR 117.

Fig. 82. — Gigantopora oropiscis sp. nov.: Part of erect stem, including ovicelled zooids (x 34). BIOCAL bin uw js.

Source : MNHN. Paris

l\G,-SL7oGi8r°POra or°Piscis SP "ov.: Ovicell (X 79). Biocal Stn DW 38

epiztfc SP,': 84' Sma" C°'°nieS °f MicroP<>rella sp. (right) and Puellina banner, Risted. (left)

86 F on?al Sh “ SP' n0V' (q V ) (X 67)' 85' Anc«>™la (lower right) and auto/ooids (x 141

re,lc“",ion of ps"‘dopores 11 285>- «• <« «4>:

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

97

Figs 89-90. — Buffonellodes crosnieri sp. nov.: 89, Zooids, two ovicelled (x 91). 90, Primary orifice and oral spine

bases (x 270). Biocal Stn DW 33. . / D.

FIGS 91-92. — Ipsibuffonella repens sp. nov.: 91, Biserially encrusting zooids, with ovicells (x 53). VZ, Primary

orifice with broad arcuate poster (x 1 17). BiOCAL Stn DW 33.

Source : MNHN, Paris

■HBK

98

D. P. GORDON & J.-L. D'HONDT

Figs 93-94. — Macrocamera erecta sp. nov.: 93, Distal end of erect stem with ovicell (left) seen in profile (x 58).

94, Ovicelled zooid and orifice with adjacent autozooids (x 83). Chalcal 2 Stn DW 78.

Figs 95-97. — Pseudoplatyglena mirabilis sp. nov.: 95, Part of erect stem, frontal side (x 36). 96, Abfrontal side of

stem with median ridge, adjacent areolar pores, and orifices (x 152). 97, Operculum and surrounding epithecal

membrane (x 251). MUSORSTOM 4 Stn DW 187.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

99

Figs 98-102. — Pseudoplatyglena mirabilis sp. nov.: 98. Proximal part of orifice, showing the broad arcuate poster and

buttressed crescentic bar (x 328). 99, Inner side of buttressed crescentic bar and frontal shield (x 432).

100. Longitudinal section through the buttressed bar and adjacent frontal shield (x 430). 101, Proximal end ot

segment from abfrontal side (x 109). 102. Cross section through the median abfrontal keel and areolar tubes (x 200).

Musorstom 4 Stn DW 187.

Source :

1 00 D. P. GORDON & J.-L. D’HONDT

Figs 103-105. — Siphonicytara armata sp. nov.: 103. Part of erect stem (x 26). 104. Surface features of stem (x 52).

105. Ascopore and avicularia (x 135). BlOCAL Stn DW 33.

Fig. 106. — Siphonicytara excentrica sp. nov.: Surface features of stem; notice complete cross-bar on most distal

avicularium, lacking in the other avicularia (x 52). BlOCAL Stn DW 38.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

101

FIGS 107-108. — Siphonicytara glabra sp. nov.: 107, Part of erect stem at bifurcation (x 49). 108. Longitudinal section

through peristome and ascopore (x 130). Biocal Stn DW 70.

FIGS 109-111. — Siphonicytara vittata sp. nov.: 109, Part of stem, frontal side (x 52). 110, Part of stem, abfrontal

side (x 36). Ill, Part of erect stem with two bifurcations (x 26). Biocal Stn DW 08.

Source : MNHN, Paris

102

D. P. GORDON & J.-L. D’HONDT

Figs 112-113. — Siphonicytara granulosa sp. nov.: 112, Part of erect stem (x 27). 113, Same, (x 54). Biogeocal

StnCP 232.

Figs 114-117. — Siphonicytara mosaica sp. nov.: 114, Part of erect stem with bifurcation, abfrontal side (x 52).

115, Part of erect stem, frontal side (x 52). 116, Apex of stem showing differentiation of zooid proximal to

developing transverse ridge (arrows) (x 109). 117, Apex of stem showing newly formed peristome, which lies on the

proximal frontal calcification of the differentiating distal zooid; areolae and sunken septular pores present on either

side of the peristome (x 93). Biogeocal Stn CP 265.

Source :

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

103

Figs 118-120. — Fedora platydiscus sp. nov.: 118, Whole colony (x 18). 119, Autozooids and orifices (x 131).

120. Zooids at colony margin showing distal pore-chambers (x 93). MUSORSTOM 3 Stn DR 117.

Figs 121-122. — Characodoma areolata (Canu & Bassler): 121. Ovicell (x 120). 122, Autozooids (x 87). MUSORSTOM 3

Stn DR 117.

Fig. 123. — Characodoma glabra sp. nov.: Autozooids and ovicell (x 1 18). MUSORSTOM 3 Stn DR 117.

Source : MNHN, Paris

104

D. P. GORDON & J.-L. D'HONDT

Figs 124-126. — Characodoma biavicularia (Canu & Bassler): 124, Part of erect colony (x 41). 125. Ovicell (x 144)

Musorstom 3 Stn DR 117. 126. From USNM type specimen 8067 (part) (x 69). "Albatross" Stn 5179.

Fig. 127. — Characodoma parva sp. nov.: Ovicelled and infertile zooids (x 131). Musorstom 3 Stn DR 117.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA 105

Fig. 128. — Characodoma sp.: Infertile zooids (x 69). MUSORSTOM 3 Stn CP 103.

FIGS 129-131. — Yrbozoon ringens Gordon: 129, Two autozooids, one with an operculum in place (x 60). 130, Profile

of fractured ovicell (x 111). 131, Ovicell, showing reticulated endooecium (x 134). Biocal Stn DW 66.

Figs 132-133. — Buffonellaria erecta sp. nov.: 132, Entire branching colony (x 18). 133, Orifices, avicularia, and

exposed ovicellular endooecium (x 124). Biocal Stn DW 66.

Source :

106

D. P. GORDON & J.-L. D'HONDT

Figs 134-136. — Osthimosia sp.: 134, Whole juvenile colony on a segment of Nellia tenella (Lamarck) (x 77).

135, Ancestrula and first daughter zooid, on N. tenella (x 134). 136, Primary orifice (x 506). Musorstom 4

Stn DW 187.

FIG. 137. — Lagenipora sp.: Very young colony (x 79). BiOCAL Stn DW 46.

Figs 138-140. — Galeopsis mitnicus Gordon: 138. Young colony with encrusting base (x 26). 139, Primary orifice

(x 252). Biocal Stn CP 75. 140, Spatulate avicularia (x 89). Chalcal 2 Stn DW 76.

Source :

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

107

Figs 141-142. — Galeopsis lageniporoides sp. nov.: 141, Part of linear encrusting lobe (x 49). 142, Ovicelled orifice

(x 195). Musorstom 6 Stn DW 489.

Figs 143-144. — Richbunea gracilis sp. nov.: 143, Part of erect branch (x 54). 144, Ovicell and adjacent peristomes

(x 92). Biocal Stn CP 67.

Source :

108

D. P. GORDON & J.-L. D HONDT

Fig. 145. — Lifuella calyciformis (Philipps): Autozooids (x 118). MUSORSTOM 3 Stn DR 117.

Figs 146-147. — Rhynchozoon tubulosum (Hincks): 146, Ovicelled zooids near growth margin (x 103). 147, Orifices,

tilted to show avicularia on peristomial side of mucrones (x 129). Musorstom 4 Stn DW 231.

Figs 148-149. — Rhynchozoon ligulatum sp. nov.: 148, Oral view of zooids near growth margin (x 65). 149, Primary

orifice (x 235). Biocal Stn DW 38.

Source :

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

109

Fig. 150. — Schedocleidochasma sp.: Autozooids (x 163). BlOCAL Stn DW 38.

Figs 151-154. — Iodictyum bicuspidatum sp. nov.: 151, Part of erect branch (x 34). 152, Autozooids (one ovicelled),

with profiles of large avicularia (x 68). 153, Ovicelled zooid with large avicularium (x 109). 154, Same, oral view

(x 125). Musorstom 6 Stn CP 419.

Source :

110

D. P. GORDON & J.-L. D’HONDT

Figs 155-157. — lodictyum blandum sp. nov.: 155, Part of fenestrate colony; an ovicell at upper left (x 23).

156,Ovicell and adjacent peristomial orifices (x 105). 157, Large lateral-oral avicularium (x 211). Biocal

Stn DW 44.

FIGS 158-159. — lodictyum illinguum sp. nov.: 158, Part of erect branch (x 54). 159, Lateral view of avicularia

(x 1 1 1). Musorstom 6 Stn DW 421.

Source :

BRYOZOA ASCOFHORINA FROM NEW CALEDONIA

1 1 1

FIG. 160. — Iodictyum illinguum sp. nov.: Ovicelled peristome (x 215). MUSORSTOM 6 Stn DW 421.

Figs 161-163. — Iodictyum trochus sp. nov.: 161, Distal end of erect branch (x 145). 162, Broken peristome, showing

sinus groove (top right) (x 129). 163, Rare peristomial avicularium (x 211). MUSORSTOM 6 Stn CP 419.

Source

112 D P. GORDON & J.-L. D'HONDT

Fig. 164. — Iodictyum Itrochus : Partially concealed ovicell (ov) (x 103). Biocal Stn DW 46.

Figs 165-166. — Iodictyum sp.: 165, Part of fenestrate colony showing two subfenestral avicularia (x 78).

166, Peristomial orifices and frontal avicularium (x 195). Musorstom 4 Stn DW 151.

Figs 167-168. — Reteporella concinnoides sp. nov.: 167, Part of trabecula with ovicell (x 141). 168, Intrafenestral

avicularium (x 125). Musorstom 4 Stn DW 231.

Source :

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

113

Figs 169-171. — Reteporella concinnoides sp. nov.: 169-170, Stages in skeletal morphogenesis of peristomes and

ovicells (x 334, x 239). 171, Small abfrontal avicularia (x 88). Musorstom 4 Stn DW 231.

Figs 172-173. — Reteporella defensa sp. nov.: 172, Part of fenestrate colony showing large avicularia (x 59). Biocal

Stn DW 38. 173, Ovicell and peristomial avicularium (x 156). Chalcal 2 Stn DW 76.

Source : MNHN. Paris

D. P. GORDON & J.-L. D HONDT

1 14

Figs 174-175. — Reteporella defensa sp. nov.: 174, Abfrontal side of trabecula (x 92). Biocal Stn CP 108. 175, Oral

view of peristomes and frontal avicularia (x 195). Biocal Stn DW 38.

Figs 176-177. — Reteporella ferox sp. nov.: 176, Part of erect branch (x 68). 177, Lateral view of peristome and

avicularium (x 268). Musorstom 6 Stn CP 419.

Source :

BRYOZOA ASCOFHORINA FROM NEW CALEDONIA

115

Fig. 178. — Reteporellci ferox sp. nov.: Ovicell and lateral avicularium (x 248). MUSORSTOM 6 Stn CP 419.

Figs 179-180. — Reteporella orstomia sp. nov.: 179. Trabecula of fenestrate colony (x 77). 180. Ovicells and frontal

avicularia (x 144). Musorstom 6 Stn DW 431.

FIGS 181-183. — Reteporella sp.: 181, Part of erect branch (x 27). 182, Ovicell with median slit visible (x 227).

183, Peristomial orifice and spiramen (x 304). BiOGEOCAL Stn DW 253.

Source :

116

D. P. GORDON & J.-L. D’HONDT

Fig. 184. — Reteporella sp. : Ovicelled orifice [ovicellular labellum (la) barely visible) and suboral avicularia (arrows)

(x 227). Biogeocal Stn DW 253.

Figs 185-187. — Reteporellina cruciformis sp. nov.: 185, Developing zooids at growth margin showing beaded rim of

primary orifice (above) and ovicell anlage (below) (x 227). 186, Part of erect branch (x 83). 187, Zooidal

peristomes with cusps (arrows) and lateral avicularium (x 173). Musorstom 3 Stn DR 117.

FIGS 188-190. — Reteporellina spiramina sp. nov.: 188, Part of erect branch; one ovicell present (x 33). MUSORSTOM 3

Stn CP 139. 189, Part of erect branch with lateral avicularia (x 28). 190, Lateral avicularium (x 144). Biocal

Stn DW 44.

Source :

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

1 17

Fig. 191. — Reteporellina spiramina sp. nov.: Ovicell and suboral avicularium (xl52). MUSORSTOM 3 Stn CP 139.

FlGS 192-193. — Reteporellina granulosa sp. nov.: 192, Part of erect branch (holotype) (x 28). 193, Ovicells and large

avicularia (x 103). Biocal Stn DW 38.

Figs 194-195. — Reteporellina projecta sp. nov.: 194, Part of erect branch (holotype) (x 32). 195, Peristomes and

avicularia (x 98). BiOGEOCAL Stn CP 290.

Source :

118

D. P. GORDON & J.-L. D'HONDT

figs 196-200. — Triphyllozoon gracile sp. nov.: 196. Part of erect branch (x 18). 197. Ovicells (x 103).

198. Ovicells (profile) and lateral avicularia (x 119). 199, Abfrontal side of branch bifurcation (x 62). 200, Ovicell

anlage and developing peristomes (x 206). Musorstom 3 Stn DR 117.

Source : MNHN, Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA 1 1 9

Figs 201-204. — Harpago dissidens sp. nov.: 201. Holotype colony, base broken (x 35). 202. Same, antapical view

(x 35). 203, Broken peristome with ovicell (x 155). 204. Primary orifice (x 217). Biocal Stn DW 66.

Figs 205-207. — Ptoboroci gelasina sp. nov.: 205. Holotype colony (x 66). 206. Primary orifice with distal ovicellular

pore (x 317). 207. Avicularium (x 457). Biocal Stn KG 101.

Source : MNHN, Paris

120

D P. GORDON & J.-L. D'HONDT

Figs 208-211. — Conescharellina breviconica Canu & Bassler: 208, Lateral view (x 19). 209, Apical view (x 29).

210, Basal view of older colony (x 19). Musorstom 3 Stn CP 87. 211, Basal view of small colony (x 29).

Musorstom 3 Stn CP 106.

Figs 212-213. — Conescharellina catella Canu & Bassler: 212, Lateral view (x 25). Musorstom 3 Stn DR 117.

213, Basal view (x 25). Musorstom 3 Stn CP 102.

Source : MNHN. Paris

BRYOZOA ASCOPHORINA FROM NEW CALEDONIA

121

FlG. 214. — Conescharellina breviconica Canu & Bassler: Orifices and avicularia (x 105). MUSORSTOM 3 Stn CP 87.

Fig. 215. — Conescharellina catella Canu & Bassler: Orifices and avicularia (x 125) Musorstom 3 Stn CP 100.

FIGS 216-218. — Conescharellina atalanta sp. nov.: 216, Apical view of colony (x 26). 217, Basal view (x 21).

Musorstom 4 Stn DW 150. 218, Orifices and avicularia (x 104). Musorstom 4 Stn DW 149.

Source : MNHN, Paris

122 D. P. GORDON & J.-L. D'HONDT

FIGS 219-220. — Trochosodon sp.: 219, Small colony (x 33). 220, Primary orifice and avicularium (x 215). BlOGEOCAL

Stn KG 275.

Figs 221-223. — Crucescharellina japonica Sil6n: 221, Whole colony (x 12). 222, Large and small avicularia and

lunoecia’ (x 101). 223, *Lunoecium\ avicularia, and primary orifice (x 212). Musorstom 3 Stn CP 106.

Source : MNHN, Paris

BRYOZOA ASCOFHOR1NA FROM NEW CALEDONIA

123

Figs 224-227. — Crucescharellina aster sp. nov.: 224, Whole colony, orificial side (x 13). 225, Whole colony,

aborificial side (x 13). 226, Centre of colony, orificial side (x 47). 227, Primary orifice and avicularium (x 149).

BiOGEOCAL Stn CP 232.

Source : MNHN, Paris

124

D. P. GORDON & J.-L. D'HONDT

FlGv>n28 23° Ichthyaria simplex sp. nov.: 228, portion of biserial stem with an ovicelled zooid (x 63)

229, autozooids with a putative ascopore arrowed (x 96). 230, proximal part of colony with rootlets (x 11)

blOGEOCAL Stn DW 313.

Figs 231-233. — Wrigiana strepsis gen. et sp. nov.: 231, part of branch showing disposition of zooids with polypides

in transparency (x 95). 232, part of stem with origin of lateral branch at lower left (x 95). 233, zooid with ovicell

and contained embryo (em) (x 88). Musorstom 4 Stn DW 185.

Source :

ESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 18 — RESULTATS DES CA

Cirripedia Thoracica: New ranges and species

of Verrucomorpha from the Indian

and Southwest Pacific Oceans

John S. BUCKERIDGE

UNITEC Institute of Technology

Private Bag 92095, Auckland, New Zealand

ABSTRACT

Verrucomorpha from deep sea collections made by several French cruises to New Caledonia, Loyalty Ridge, Vanuatu,

Wallis Island and Futuna Islands, Comoro Islands, and by the French-Indonesian cruise Karubar in Indonesian waters,

over the period 1985-1994, are investigated. Fourteen species of verrucid are described, including four new species.

Verruca jagoy Altiverruca jonesae , Brochiverruca crosnieri and Metaverruca maclaughlinae\ the bathymetric and geo¬

graphic ranges of verrucid taxa are extended, and it is confirmed that this is one of the most diverse verrucomorph faunas

known. The status of both Verruca and Metaverruca is considered, and a revised key to genera of the Verrucidae is given.

RESUME

Cirripedia Thoracica: Nouvelles repartitions et nouvelles especes de Verrucomorpha de

I'ocean Indien et du Sud-Ouest Pacifique.

Les Verrucomorphes, principalement bathyaux, r£colt6s entre 1985-1994, lors de diverses campagnes fran^aises au

large de la Nouvelle-CalSdonie et de la Ride des LoyautS, au Vanuatu, aux lies Wallis et Futuna, & la Grande-Comore, ainsi

que lors de la campagne franco-indon£sienne Karubar en Indon6sie orientale, sont 6tudi6s. Quatorze espfcces ont 6t6 trou¬

pes, parmi lesquelles quatre sont nouvelles pour la science : Verruca jago, Altiverruca jonesae , Brochiverruca crosnieri

and Metaverruca maclaughlinae. II se confirme que la faune des Verrucomorphes 6tudi6e ici est l'une des plus diversifi6e

connue dans une meme region. Les distributions bathymStriques et g6ographiques de plusieurs espfcces sont 6tendues. Le

statut des genres Verruca et Metaverruca est examine, et une nouvelle cl6 des genres de la famille Verrucidae est propos^e.

INTRODUCTION

The deep sea verrucomorph cirripede fauna of Indonesia, New Caledonia and the Wallis and Futuna Islands has

recently been described by BUCKERIDGE (1994), in which earlier work in the region is cited. This paper records

BUCKERIDGE, John S., 1997. — Cirripedia Thoracica: New ranges and species of Verrucomorpha from the Indian and

Southwest Pacific Oceans. In: A. Crosnier (ed.), RSsultats des Campagnes Musorstom, Volume 18. Mem. Mus. natn.

Hist, nat., 176: 125-149. Paris ISBN 2-85653-511-9.

126

J. S. BUCKERIDGE

previously unsorted material collected from New Caledonian waters by BIOCAL (1985), MUSORSTOM 6 (1989),

Bathus 1 (1993), BaTHUS 2 (1993), BATHUS 3 (1993), Bathus 4 (1994) and Halipro 1 (1994); from Vanuatu by

Musorstom 8 (1994); from the Wallis and Futuna Islands region by MUSORSTOM 7 (1992); from the Loyalty

Ridge by MUSORSTOM 6 (1989); from Grande-Comore (in the Indian Ocean) by Jago-Coelacanthe (1989) and

from Indonesian waters by Karubar (1991) expeditions.

In addition, the study has been expanded to include verrucid material collected by the Western Australian

Museum, Perth, Australia, from off northwestern Australia by R.V. "Soela" (1982, 1984). The decision to include

western Indian Ocean material was based upon the need to compare the current collection, and that of BUCKERIDGE

(1994), with Verruca macani Stubbings, 1936, a species which appeared similar to Metaverruca described in

BUCKERIDGE (1994), but which had previously been only partially described. A first description of the soft parts of

this species is provided here. The geographic region thus expanded, permitted a most intriguing new taxon from

the Comore Islands to be included. With the exception of unusual varieties, only new species, or parts of taxa not

previously illustrated, are Figured in this paper.

The Verrucidae are a family of asymmetrical sessile barnacles characterised by six calcareous plates. The shell is

made up of a carina and rostrum, plus a tergum and scutum, the latter two having moved from the operculum to

become "fixed" as part of the shell wall. The remaining plates, a tergum and scutum (hereafter termed movable

plates) make up the operculum. It appears to be totally random as to whether the "left" or "right" opercular plates

move to complete the shell wall (Darwin, 1854; BUCKERIDGE, 1994). However Newman (1989) has

demonstrated that in Neoverruca at least, the asymmetry is determined in the earliest juveniles by the side closest

to the substratum.

Types: Holotypes of all new species are held by the Museum national d'Histoire naturelle (prefix "MNHN-Ci")

in Paris, France. Paratypes are also held by the Museum national d'Histoire naturelle, and when numbers of

specimens available permit, by the National Museum of Natural History, Washington DC, United States of

America (prefix "USNM") and UNITEC Institute of Technology, Auckland, New Zealand (prefix "CAX").

In the lists of material examined the abbreviations for the gear used are: CP = beam trawl and DW = Waren

dredge. The specimens from Grande-Comore were collected by the German submersible Jago. The Western

Australian Museum material was collected by Engel trawl. Grid references for the Western Australian material have

been corrected to a single value, represented by degrees and minutes only.

LIST OF SPECIES

Genus VERRUCA Schumacher, 1817

Verruca albatrossiana Pilsbry, 1912

Verruca jago sp. nov.

Genus METAVERRUCA Pilsbry, 1916

Metaverruca defay eae Buckeridge, 1994

Metaverruca macani (Stubbings, 1936)

Metaverruca pacifica Buckeridge, 1994

Metaverruca plicata Buckeridge, 1994

Metaverruca recta (Aurivillius, 1898)

Metaverruca maclaughlinae sp. nov.

Genus ALTIVERRUCA Pilsbry, 1916

Altiverruca cristallina (Gruvel, 1907)

Altiverruca jonesae sp. nov.

Altiverruca nitida (Hoek, 1883)

Genus ROSTRATOVERRUCA Broch, 1922

Rostratoverruca intexta (Pilsbry, 1912)

Rostratoverruca kruegeri (Broch, 1922)

Genus BROCHIVERRUCA Zevina, 1993

Brochiverruca crosnieri sp. nov.

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

127

SYSTEMATIC ACCOUNT

Superorder THORACICA Darwin, 1854

Suborder VERRUCOMORPHA Pilsbry, 1916

DIAGNOSIS. — Sessile thoracican cirripedes with asymmetrical shell wall comprising fixed scutum, fixed

tergum, rostrum and carina, closed by movable scutum and movable tergum (Verrucidae), sometimes including

rostrolatus and carinolatus on movable side (Pro verrucidae), or basal whorls of imbricating plates (Neoverrucidae);

basis membranous or calcareous.

Family VERRUCIDAE Darwin, 1854

(amend. Newman & Hessler, 1989)

DIAGNOSIS. — Verrucomorphans with primary wall (carina, rostrum, fixed scutum, fixed tergum), in contact

with substratum; latera absent.

Discussion. — There are currently seven genera recognised within the Verrucidae. Initially, PlLSBRY (1916),

recognised four "sections", which were subsequently elevated to subgenera (e.g. Newman, Zullo & Withers,

1969; FOSTER, 1978). BROCH, (1922) added Rostratoverruca , and Zevina (1987, 1993) added Spongoverruca and

Brochiverruca respectively.

The presence of a myophore on the fixed scutum has been previously regarded by many workers as of

considerable value in distinguishing between genera, e.g. PlLSBRY (1916), Zevina (1987), BUCKERIDGE (1994),

FOSTER & BUCKERIDGE (1995). Indeed, PlLSBRY (1916) used this character to define his "Section A" Metaverruca ,

i.e.,

"The basal borders of the wall plates are inflexed, forming a wide basal ledge; the fixed scutum bears a

depending tongue-shaped adductor ridge or myophore; the apex of the rostrum is marginal; top flattened; sculpture

weak".

However, as Paulo YOUNG has pointed out (pers. comm.), PlLSBRY failed to stress that his Lepas stroemia

Muller, 1776, the type for "Section B" Verruca , also possesses a myophore. If this is taken into account, then

PlLSBRY's diagnosis for Verruca s.s. is less than satisfactory, i.e.,

"Top flattened, the plane of the movable plates not far from parallel with that of the base; radio-alar area

between fixed scutum and fixed tergum small or linear".

Of the points stressed in the above two diagnoses, it must be noted that:

1. If the apex of the rostrum is not marginal, the species cannot be Verruca, rather it is either Rostratoverruca

or Brochiverruca.

2. The basal ledge is a difficult character to use to distinguish Verruca and Metaverruca , as a number of Verruca

s.s. (see in PlLSBRY. 1916), have slightly thickened basal edges to the shell wall, e.g. Verruca alba Pilsbry,

Verruca calotheca Pilsbry, Verruca entobapta Pilsbry.

3. The radio-alar area between fixed scutum and fixed tergum does not appear to be particularly linear in some

Verruca s.s., e. g. V. entobapta.

It should also be noted that the angle of the operculum to the base may vary a little, particularly where the

nature of the substrate appears to have a strong influence. However, I firmly believe that Verruca and Metaverruca

are distinct. Key features that distinguish Metaverruca are the presence of all of the following characters:

— thickened basal ledge on the compartments (this is often so well developed that it is very difficult to

disarticulate the shell evenly along compartmental sutures).

— a distinctive D-shaped orifice, with opercular hinge straight.

— shell shape "box-like".

— operculum nearly parallel to base.

— a well developed myophore on the fixed scutum.

128

J.S. BUCKERIDGE

Although each of these characters may be found in Verruca , they are only found together in Metaverruca.

Clearly juveniles of Metaverruca will not necessarily possess all features, but it would be most cumbersome in a

diagnosis of a genus to include all stages of ontogeny. A revised key is provided below, which permits

differentiation between the genera.

In the past, numerous workers have spent considerable time in primary descriptions, detailing the number of

interlocking ridges between the carina and rostrum. Whilst there is some validity in considering these aspects, they

should only be taken as a guide, for in species like Metaverruca recta , (for which there is now a very large amount

of comparative material), one can distinguish a series (not necessarily solely ontogenetic) that shows a range from

one or two very flat ridges to more than five (see BUCKERIDGE 1994, fig. 13). The spacing of growth lines may

also be misleading (see under Metaverruca recta). However, one feature that does appear consistent, except in the

most juvenile of specimens, is the number of interlocking ribs (or ridges) on the movable opercula. These very

quickly reach an optimal number and are not known to add further ribs, even in very large specimens.

Morphology of the soft parts can be of considerable help in distinguishing species, particularly of Metaverruca.

However, most taxa described here may be identified on shell morphology alone. The nomenclature used in this

paper follows that of BUCKERIDGE, 1994, fig. 1. The use of a "+" in the cirral counts indicates that the cirrus is

incomplete, and "c.a." refers to caudal appendages.

Stratigraphic Range. — ?Middle, Upper Cretaceous to Recent, Europe, South America, Australasia.

Distribution. — Cosmopolitan in present seas.

Key to genera of the Verrucidae

1. Rostral apex marginal . 2

— Rostral apex removed from margin . . . 3

2. Operculum nearly vertical to base . 4

— Operculum nearly parallel to base . 5

3. Carinal apex marginal . Rostratoverruca

— Carinal apex removed from margin . Brochiverruca

4. Myophore present on fixed scutum . Cameraverruca

— Myophore absent on fixed scutum . 6

5. Shell box-like, with thickened, inflected basal margin and D-shaped aperture . Metaverruca

— Shell low conic, with non-linear hinge-line . Verruca

6. Not embedded in sponge . Altiverruca

Embedded in sponge . Spongoverruca

Genus VERRUCA Schumacher, 1817

Verruca "Section B"- Pilsbry, 1916: 23.

Verruca (Verruca) - Newman Zullo & Withers, 1969: R281. — FOSTER, 1978: 68.

Verruca - Zevina 1987: 1812. — Newman & HESSLER, 1989: 268.

Diagnosis. — Verrucids with shell form depressed; apices of rostrum and carina marginal; operculum parallel

to base.

Type Species. Lepas stroemia Muller, 1776. North Atlantic, northern Europe, Mediterranean, ?Red Sea,

intertidal - 500 m.

Species. — Twenty eight taxa are presently assigned to this genus, (f indicates extinct taxa): Verruca alaskana

Pilsbry, 1943; V. alba Pilsbry, 1907; V. alba caribbea Pilsbry, 1916; V. alba barbadensis Pilsbry, 1916;

Source :

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

129

V. albatrossiana Pilsbry, 1912; V. calotheca Pilsbry, 1907; V. calotheca flavidula Pilsbry , 1916; V. calotheca

heteropoma Pilsbry, 1916; V. calotheca niasiensis Nilsson-Cantell, 1929; V. cookei Pilsbry, 1927; V. floridana

Pilsbry, 1916; V. grimaldi Gruvel, 1920; V. laevigata (Sowerby, 1827); + V. nuciformis Buckeridge, 1983;

fV. prisca Bosquet, 1853; \V. pusilla Bosquet, 1857; fV. rocana Steinmann, 1921; V. sinuosa Foster &

Buckeridge, 1995; V. spengleri Darwin, 1854; V. stroemia (Muller, 1776); tV. tasmanica Buckeridge, 1983;

fV. tasmanica chatheca Buckeridge, 1983; V. trisulcata Gruvel, 1920; V. xanthia Pilsbry, 1916; V. xanthia

insculpta Pilsbry, 1916; V. entobapta Pilsbry, 1916; V. scrippsae Zullo, 1964; plus the new species described

here: V. jago sp. nov.

Remarks', t Verruca withersi Newman & Schram, 1980 (included in the listing for Verruca sensu stricto in

Buckeridge, 1994) has been re-evaluated and interpreted as "non-verrucid" in Buckeridge. 1996. The generic

status of a further group of verrucids, comprising V. aequalis Aurivillius, 1898; V. cornuta Aurivillius, 1898;

V. costata Aurivillius, 1898, is uncertain.

DISCUSSION. — The genus Verruca is a grouping of convenience. In this sense, the taxa within it are rather

discordant. It is clear that future revision will require the genus to be split, with perhaps only V. cookei, V. jago

sp. nov., V. laevigata , V. spengleri , V. stroemia , and some fossil taxa remaining in Verruca sensu stricto , the

remaining being placed in at least one new genus (see BUCKERIDGE, 1994).

Stratigraphic Range. — Cretaceous (Australia, New Zealand, Columbia, western Europe), Palaeogene

(New Zealand, Argentina, Chatham Islands) to Recent.

Distribution. — Pacific, North Atlantic, Mediterranean, Caribbean, ?Red Sea, Indian Ocean, intertidal -

620 m.

Verruca albatrossiana Pilsbry, 1912

Verruca albatrossiana Pilsbry, 1912: 292.

Verruca (Eu-Verruca) albatrossiana - BROCH, 1922: 290, figs 39-40.

Verruca grex Hoek, 1913: 142, pi. 11, figs 7-13, pi. 13, figs 11-13.

Verruca albatrossiana - BUCKERIDGE, 1994: 91, figs la-f.

MATERIAL EXAMINED. — Indonesia. Karubar: stn CP 17, 05° 1 5'S, 133°01'E, 439-459 m, 24.10.1991:

1 specimen on a cidaroid spine.

NW Australia. " Soela ": WAM 909-86, 13°33'S, 122°14'E, 494-496 m, 13.2.1984: 11 specimens, on cidaroid

spines.

Diagnosis. — Verruca with rostrum and carina low, strongly developed longitudinally to give shell oblique

appearance; fixed scutum considerably larger than fixed tergum, opercular plates displaced carinally.

Discussion. — This material conforms to that described in BUCKERIDGE (1994). This is the first record of the

species from northwestern Australia.

Distribution. — Indo-Pacific, 345-620 m.

Verruca jago sp. nov.

Fig. 1 a-m, 8 e-i.

Material EXAMINED. — Grande-Comore. JAGO-COELACANTHE: stn at ll°5rS, 43°20’E, 203 m, 1.12.1989:

4 specimens. — Stn at 11°50’S, 43°21’E, 196 m, 8.12.1989: 4 specimens.

Types. — Holotype : MNHN-Ci 2425, r-c diameter: 2.9 mm; height: 0.4 mm (from 1 1 °5 1 ’S, 43°20'E).

130

J.S. BUCKERIDGE

Paratypes: MNHN-Ci 2426, r-c diameter: 2.27 mm; height: 0.25 mm (1 specimen, from 1 1 °5 1 'S, 43°20'E);

MNHN-Ci 2431 (3 specimens, from 11°50’S, 43°2rE); CAX 108 (1 specimen, from 1 1°50’S, 43°21’E); USNM

282635 (2 specimens, from ll°5rS, 43°20’E).

DIAGNOSIS. — Small, flattened Verruca\ shell with regularly disposed, large punctae; movable tergum

quadrangular, 3 ribs; fixed scutum with large myophore; movable scutum triangular, with 3 ribs, occludent portion

lacking ribbing, internally with elevated depression for adductor muscle attachment; rami of cirrus I spatulate.

Description. — Verruca with shell white, translucent, low conic, weakly to moderately plicate, patterned with

fine circular tubes (punctae), which extend from exterior (infilled with chitinous material), to open to interior as

very small pores; punctae generally separated by at least own diameter and arranged as rows along leading edge of

sub-parallel, concentric growth lines; shell wall comprised of up to 7 calcareous lamellae (particularly clear in

basal view of rostrum of Ci 2431); operculum sub-parallel to base; rostrum and carina articulating with up to

2 ribs; basis membranous. Fixed scutum with large, well formed myophore for adductor muscle attachment; fixed

tergum low, internally with small horizontal ledge, although not as well developed as myophore in fixed scutum;

movable tergum quadrangular, with well defined apico-basal rib, plus 2 secondary ribs, interlocking with movable

scutum; articular margin slightly concave; movable scutum triangular, with 2 ribs interlocking with tergum and

further adjacent curved, apico-basal rib terminating at rostro-carinal suture, occludent portion lacking ribbing,

internally with elevated depression for adductor muscle attachment.

Mandible tridentate, lower angle pectinate, with 2 moderately large basal spines; first maxilla with 2 large and

2 medium upper spines, moderate to weakly formed notch, lower angle with 2 large and 3 medium spines; labrum

gently arched, with single large tooth on either side and at least 4 smaller, bidentate teeth centrally; palps with 3-4

terminal spines, not meeting across labrum.

Cirri possess the following number of segments (r-c being the rostro-carinal diameter):

r-c (mm) I II III IV V VI c.a.

2.27 7,6 5,9 12,14 12,14 13,14 13,13 5

Cirrus I with both rami heavily hirsute, moderately spatulate, cirri I and II with anterior rami shorter than

posterior rami, although in cirrus I with more segments.

Length of caudal appendages about twice that of basal segment of pedicel of cirrus VI; penis very large,

occupying significant part of mantle cavity, with numerous terminal setae and scattered tufts of long setae; eggs

present in holotype, larvae in mantle cavity of Ci 2431a.

DISCUSSION. — The interesting features of this species are the punctae, or pores, the horizontal ledge on the

tergum, the spatulate rami on the first cirri and the very large penis. Although the latter is likely a reflection of the

current sexual activity of the specimens studied, it was proportionally larger than any other I have viewed in the

Verrucidae. The presence of a ledge on the inner surface of the fixed tergum, although not unique ( Verruca stroemia

has one), is rare. In addition, some specimens of Verruca jago sp. nov. are remarkably flat. Paratype Ci 2426, from

which the soft parts were drawn, had a shell height of only 0.25 mm.

The most distinctive character about this species is the small punctae that are regularly disposed over the entire

shell. The shell is translucent in many specimens, and with a stereomicroscope, it is possible to focus through the

punctae, and to perceive them as pores. The punctae narrow as they approach the inner side of the wall, and open to

the interior. Although under light microscopy, most punctae appear to be closed to the exterior, examination under

the scanning electron microscope (following the desiccation and electrolytic coating process) shows that soft tissue

extends to the exterior (fig. 8g-i). In regions of higher corrasion, such as around the apices of plates, the punctae

form slightly raised granules. In light of this, it is suggested that the tissue (probably chitinous) has some

mitigating effect on erosion rates.

The inner surface of the plates of some other species within Verruca are characterised by chitin filled pores (e.g.

Verruca laevigata , Verruca stroemia ). However the pores in V jago sp. nov. are at least twice the diameter of those

observed in any others, and differ further by being clearly visible on the exterior surface, a feature generally

restricted to of worn, or juvenile specimens of other species.

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

131

Fig. 1. — Verruca jago sp. nov.: a, holotype, complete shell (dorsal view), MNHN-Ci 2425; b, fixed scutum (right,

dorsal view), showing lunulitiform myophore for adductor muscle attachment; c, fixed tergum (right, dorsal view),

showing internal ledge for muscle attachment; d, movable scutum (left, interior); e, movable scutum (left, exterior);

f, movable tergum (left, interior); g, movable tergum (left exterior); h, cirrus I; i, cirrus II; j, basal portion of cirrus VI,

with penis and caudal appendage attached; k, mandible; I, first maxilla; m, labrum with palps. All material from ll°5rS,

43°20'E.

132

J. S. BUCKERIDGE

Specimen Ci 2426 still retains a small fragment of substrate, which appears to be an unweathered, fine grained,

intermediate siliceous volcanic rock.

V. jago sp. nov. falls within a group of "shallow water" verrucids, including V. cookei , V. laevigata ,

V. spengleri and V. stroeniia. V. jago may be distinguished from V. stroemia by its weaker external ribbing,

a less projecting lower lobe on the first maxilla, a labrum with bidentate teeth and well separated palps. It differs

from V. cookei by having a weakly hirsute central portion on the first maxilla, and much shorter caudal

appendages. In V. jago the movable opercula are much narrower and the shell is more plicate than in V. laevigata.

The movable scutum of V. spengleri is quite distinct, being both narrower, and possessing a more deeply

excavated adductor muscle pit.

Etymology. — An acronym for the submersible used during the expedition: Jago.

Distribution. — Grande-Comore Island, 196-203 m.

Genus ALTIVERRUCA Pilsbry, 1916

Verruca "Section D": Altiverruca Pilsbry, 1916: 40.

Verruca (Altiverruca) - FOSTER, 1978: 68.

Altiverruca - Zevina, 1987: 1813.

Diagnosis. — Verrucids with erect form, bases of plates not inflected; operculum close to vertical; myophore

absent.

Type Species. — Verruca hoeki Pilsbry, 1907: 1 13. West Indies, 907-1060 m.

Species. — The 37 taxa presently assigned to this genus include: Altiverruca angustiterga Zevina, 1987;

A. aves (Zevina, 1975); A. bicornuta (Pilsbry, 1916); A. beringiana Zevina, 1992; A. cassis (Hoek, 1913);

A. casula (Hoek, 1913); A. crenata (Aurivillius, 1898); A. cristallina (Gruvel, 1907); A. cristallina laevis

(Broch, 1922); A. darwini (Pilsbry, 1907); A. erecta (Gruvel, 1900); A. galapagosa Zevina, 1987; A. galkini

Zevina, 1990; A. gibbosa (Hoek, 1883); A. gibbosa somaliensis (Nilsson-Cantell, 1929); A. gira

(Zevina, 1987); A. hoeki (Pilsbry, 1907); A. incerta (Hoek, 1883); A. laeviscuta Buckeridge, 1994; A. longa

Zevina, 1988; A. longicarinata (Gruvel, 1900); A. mitra (Hoek, 1907); A. mollae Zevina, 1990; A. navicula

(Hoek, 1913); A. nitida (Hoek, 1883); A. obliqua (Hoek, 1883); A. plana (Gruvel, 1907); A. quadrangularis

(Hoek, 1883); A. radiata (Gruvel, 1901); A. rathburniana (Pilsbry, 1916); A. sculpturata Zevina, 1987;

A. sub lima Zevina, 1987; A. sulcata (Hoek, 1883); A. tchesunovi Zevina, 1988; A. vitrea Zevina, 1988;

A. vertica Foster & Buckeridge, 1995; plus the new species described here: A. jonesae sp. nov.

Stratigraphic Range. — Recent.

Distribution. — Cosmopolitan, 233-4950 m.

Altiverruca cristallina (Gruvel, 1907)

Verruca cristallina Gruvel, 1907, pi. 1, figs 9-10.

Verruca cristallina laevis Broch, 1922, fig. 41 a-d.

Verruca (Altiverruca) cristallina - ROSELL, 1989: 24, pi. 6. figs d-i. — ROSELL, 1991: 34.

Altiverruca cristallina - BUCKERIDGE, 1994: 93, figs 2a-h.

Material EXAMINED. — Vanuatu. Musorstom 8: stn CP 1109, 14°52’S, 1 67° 1 8'E, 1550-1620 m 8 10 1994

21 specimens, on rocks. — Stn CP 1110, 14°49'S, 167°15,E, 1360 m. 8.10.1994: 3 specimens, on rock.

Diagnosis. —Altiverruca with carina and rostrum interlocking with up to 6 ribs; movable tergum with 4 to 5

articulating ribs, at least two articulating with scutum; caudal appendages short.

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

133

Discussion. — This material conforms to that described in BUCKER1DGE (1994). This collection has extended

the known geographic range for this species to Vanuatu waters.

Distribution. — Tropical Pacific, 233-2340 m.

Altiverruca jonesae sp. nov.

Figs 2 a-f, 3 a-g.

MATERIAL EXAMINED. — Vanuatu. Musorstom 8: Stn CP nil, 14°5rs. 167°14'E. 1210-1250 m, 8.10.1994:

1 specimen, on an hexactinellid sponge.

TYPE. — Holotype : MNHN-Ci 2430, rostro-carinal diameter: 7.1 mm; height: 7.7 mm (from Stn CP 1111,

14°5TS, 167°14'E).

FIG. 2. — Altiverruca jonesae sp. nov.: a, holotype, complete shell (dorsal view), MNHN-Ci 2430; b. movable scutum

(right, interior); c, movable scutum (right, exterior); d, complete shell (fixed opercula view); e, movable tergum

(right, interior); f, movable tergum (right exterior). All material from Stn CP 1111.

134

J.S. BUCKERIDGE

Diagnosis. — Moderate sized Altiverruca with surface ornamented by short rounded spines; movable tergum

and scutum possessing 3 diagonal ribs; movable tergum with concave occludent margin and incurved apex.

Description. — Shell white, with short, flattened and rounded spines aligned along regularly spaced growth

lines; rostrum and carina interlock with 4 to 5 ribs; apex of rostrum and carina produced.

Fixed scutum and fixed tergum with apices extending beyond opercular plates; fixed scutum folded over on

upper margin to form surface confluent with plane of operculum; fixed tergum strongly flexed towards fixed

scutum, with strong, sharply rounded primary diagonal rib, and weak minor secondary rib running along lower part

of scutal margin.

Movable scutum triangular, apico-basal ridge prominent, with 2 secondary ribs, tergal margin concave; interior

with elevated, well developed depression for adductor muscle attachment. Movable tergum with concave occludent

margin and incurved apex; upper angle between occludent margin and scutal margin protuberant; 1 primary apico-

basal ridge and 2 secondary ribs present on scutal side, carinal side without ribs.

Mandible tridentate, lower angle pectinate with 6 small spines; first maxilla with 1 large and 1 small upper

spine, notch with 3 very small spines, lower angle with 1 large spine and 2 smaller spines, hirsute; labrum

broadly arched, finely denticulate along entire margin, palps narrow, clearly separated across labrum.

Cirri I and II with anterior rami approximately half length of posterior, cirrus III more like cirrus IV than II;

penis thin with terminal setae. Cirri with following number of segments, where cirri are incomplete, maximum

for both sides considered (r-c being rostro-carinal diameter) :

r-c (mm) I II III IV V VI c.a.

7.1 7,10 7,12 19,23 19+, 25 21+, 23+ 25+, 28+ 8

Intermediate segments of cirrus VI with 2 pairs of setae, distal pair much larger than proximal; length of caudal

appendages approximately 1.2x length of basal segment of pedicel of cirrus VI.

Fig. 3. — Altiverruca jonesae sp. nov.: a, holotype, cirrus I, MNHN-Ci 2430; b. cirrus II; c, cirrus III; d, basal portion of

cirrus VI, with penis and caudal appendage attached; e, labrum with palps, setae shown on left palp only; f, first

maxilla; g, mandible. All material from Stn CP 1111.

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

135

DISCUSSION. — This is a most elegant species. Unfortunately only one specimen has been collected; however

the spinose nature of the shell ornamentation, the deep adductor muscle depression on the movable scutum and the

strongly incurved apex and shape of the movable tergum are sufficient to warrant specific differentiation. Shell

ornamentation is considered significant with this species, as this is clearly neither a function of substrate nor over¬

crowding. Verruca jonesae resembles, but may be distinguished from, both Altiverruca gibbosa (Hoek, 1883), and

Altiverruca sulcata (Hoek, 1883), by the ornate ornamentation, a more defined second rib on the movable scutum,

the incurved apex of the movable tergum and a much shorter anterior ramus on cirrus I.

It may be distinguished from a similar north Pacific species, Altiverruca sculpturata Zevina 1987, by a

narrower movable scutum, a more protruded fixed tergal apex, and more spinose shell ornamentation.

Etymology. — Named for Diana Jones, Curator of Crustacea, Western Australian Museum, in recognition

of her extensive work with the cirripedes of the region.

Distribution. — Vanuatu, 1210-1250 m.

Altiverruca nitida (Hoek, 1883)

Fig 8 a-b.

Verruca nitida Hoek, 1883: 138, figs 6-7. — Gruvel, 1905: 177, fig. 194. — Nilsson-Cantell, 1927: 778.

Altiverruca nitida - BUCKERIDGE, 1994: 101, fig 6 a-g.

Material EXAMINED. — New Caledonia. Biocal: stn CP 29, 23°08'S. 166°40’E, 1100 m, 29.8.1985:

5 specimens, on pebbles. — Stn CP 30, 23°09'S, 166°4rE, 1140 m, 9.8.1985: 6 specimens, on pebbles, shell. —

Stn DW 51, 23°05'S, 167°44’E, 700 m, 31.8.1985: 1 specimen.

Bathus 1: stn CP 651, 21041'S, 166°40'E, 1080-1180 m, 11.3.1993: 1 specimen.

Vanuatu. Musorstom 8: stn CP 1074, 15°48'S, 167°24’E, 775-798 m, 4.10.1994: 1 specimen, on scaphopod shell.

— Stn CP 1125, 15°57'S, 166°38’E, 1160-1220 m, 10.10.1994: 4 specimens, on pebbles. — Stn CP 1129, 16°00’S,

166°39'E, 1014-1050 m, 10.10.1994: 8 specimens on pebbles.

Diagnosis. — Altiverruca with carina and rostrum interlocking with 2 ribs; movable scutum with 2 ribs

articulating with 1 strong diagonal rib of movable tergum; caudal appendages long.

DISCUSSION. — This material conforms to that described in BUCKERIDGE (1994). The specimen from station

CP 1074 has a less flared shell than most, but this is attributed to the substrate.

Distribution. — Tropical western Pacific, 650 to 2040 m.

Genus BROCHIVERRUCA Zevina, 1993

DIAGNOSIS. — Verrucids with apex (umbo) of both carina and rostrum removed from margin of plates; fixed

scutum with or without myophore.

Type Species. — Brochiverruca margulisae Zevina, 1993: 10. Mozambique Channel, Indian Ocean, 935-

950 m.

SPECIES. — Four species are here assigned to this genus: Brochiverruca margulisae Zevina, 1993; B . dens

(Broch, 1932); B. polystriata Buckeridge, 1994; and the new species described here: B. crosnieri sp. nov.

Stratigraphic Range. — Recent.

Distribution. — Western Pacific and Western Indian Ocean, 270-950 m.

DISCUSSION. — The distinctive nature of Brochiverruca was first recognised by BROCH (1932). He suggested

that the patelliform carina might be sufficient cause to set V. dens apart from Rostratoverruca. However, he left

136

J. S. BUCKERIDGE

this "to future investigators to settle, whether Verruca dens must be considered type of a separate subgenus or not".

In light of this reference, and the abundance of V. dens , it is to be regretted that ZEVINA chose not to use Verruca

(Rostratoverruca?) dens as her type.

Brochiverruca crosnieri sp. nov. extends the bathymetric range of the genus into shallower water. Prior to this

record, the genus was unknown from depths of less than 348 m. Brochiverruca is unlike other verrucids in that the

species appear to have strong substrate (= host) preferences. Of the Brochiverruca species that I have studied, all are

consistently attached to the following cnidarians:

Brochiverruca dens (Broch) scleractinian Madrepora

Brochiverruca polystriata Buckeridge scleractinian Ellanospammia and Madrepora

Brochiverruca crosnieri sp. nov. antipatharian antipatharian sp. indet.

Unfortunately, Zevina states only that the type, B. margulisae Zevina, represented by 4 specimens, is attached

to "coral".

Brochiverruca crosnieri sp. nov.

Fig 4 a-h

Material EXAMINED. — Loyalty Ridge. Musorstom 6: stn CP 401, 20°42’S, 167°00*E, 270 m, 14.2.1989:

1 specimen, on antipatharian.

Type. — Holotype : MNHN-Ci 2424, r-c diameter: 2.2 mm; height: 2.4 mm (from Stn CP 401, 20°42'S,

167°00'E).

Diagnosis. — Shell white, porcelanous, very finely sculptured with delicate longitudinal striae crossed by fine

growth lines; rostrum and carina each with apex separated from upper margin by slightly more than l/3rd distance

from basal margin; interior of movable tergum with articular node on scutal margin; fixed scutum with ledge for

adductor muscle attachment.

DESCRIPTION. — Shell white, porcelanous, molariform; exterior very finely sculptured, with delicate

longitudinal striae crossed by fine growth lines.

Rostrum and carina each with apex protruded and incurved distally, separated from upper margin by slightly

more than one third distance from basal margin; margin between both plates characterised by 5 flat and non-

prominent, articulating ribs.

Fixed scutum with ledge internally for adductor muscle attachment; fixed tergal edge confluent with fixed

tergum and with no clear articulating ribs, fixed tergal apex most elevated part of shell.

Movable tergum of low relief, quadrangular, with 1 narrow, flat medial rib; with extended upper angle of

articular margin articulating with movable scutum; internally with articular node on scutal margin.

Movable scutum quadrangular, less than half width of movable tergum, with weak apico-basal ridge; apex

incurved towards movable tergum; interior with raised central portion excavated centrally for adductor muscle

attachment; tergal margin of raised area with rounded embayment (fig. 4b).

Mandible tridentate, lower angle pectinate with about 7 small spines; first maxilla with 2 larger spines above

notch and 3 spines on lower part, centrally hirsute, 3 tufts of setae on upper surface. Labrum gently arcuate,

margin finely denticulate. Penis about twice length of caudal appendages, terminally hirsute. Cirri I and II with

flattened anterior rami, cirrus III more like cirrus IV than II; intermediate segments of cirrus VI with 3 pairs of

setae. Cirri with following number of segments, (r-c being rostro-carinal diameter) :

r-c (mm) I II III IV V VI c.a.

2-2 6, 8 5, 10 14, 16 9+, 17 16, 16+ 17, 23 5

Caudal appendages about half as long as basal segment of pedicel of cirrus VI.

DISCUSSION. This species is related to Brochiverruca dens (Broch) and Brochiverruca polystriata Buckeridge.

It may be distinguished from B. dens by its significantly smoother shell, with longitudinal ornamentation

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

137

Fig. 4. — Brochiverruca crosnieri sp. nov.: a, holotype, complete shell (dorsal view), MNHN-Ci 2424; b, movable

scutum and movable tergum (interior); c, cirrus I; d, cirrus II; e, intermediate segment of cirrus VI; f, basal portion of

cirrus VI, with penis and caudal appendage attached; g, mandible; h, first maxilla. All material from Stn CP 401.

restricted to rare striae; and a broader movable scutum and finely denticulate labrum. It differs from B. polystriata in

its shell ornamentation, and the possession of only one rib on the movable tergum.

ETYMOLOGY. — Named for Alain CROSNIER, ORSTOM (Institut Fran^ais de Recherche Scientifique pour le

Developpement en Cooperation), Paris, who was instrumental in much of the collecting and sorting, and who

provided sustained support and friendship during my sojourn in Paris.

Distribution. — Loyalty Ridge, 270 m.

138

J. S. BUCKERIDGE

Genus META VERRUCA Pilsbry, 1916

Verruca "Section A": Metaverruca Pilsbry, 1916: 21.

Verruca ( Metaverruca ) - Nevvnan. ZULLO & Withers, 1969: R283 —FOSTER, 1978: 68.

Metaverruca - Zevina, 1987: 1812.

DIAGNOSIS (Amended). — Verrucids with box-like shell; apices of carina and rostrum marginal; fixed scutum

with myophore; operculum parallel to base, with straight basal margin, such that aperture is "D-shaped"; base of

shell wall inflected internally, thickened.

Type Species. — Verruca recta Aurivillius, 1898: 195 (Subsequent Designation). The Azores, 1 135 m.

Species. — Thirteen species are presently assigned to this genus: Metaverruca corrugata (Broch, 1922);

M. defayeae Buckeridge, 1994; M. lepista (Zevina, 1987); M. maedni (Stubbings, 1936); M. norfolkensis

Buckeridge, 1994; M. paciftca Buckeridge, 1994; M. pallida Zevina, 1990; M. plicata Buckeridge, 1994; M. recta

(Aurivillius, 1898) (for full synonymy and comments see BUCKERIDGE, 1994); M. reunioni Foster & Buckeridge,

1995; M. seriola (Zevina, 1987); M. tcirasovi Zevina, 1971; plus the new species described here, M. maclaughlinae

sp. nov.

Stratigraphic Range. — Lower Miocene (New Zealand) to Recent.

Distribution. — Cosmopolitan, 167- 4100 m.

Metaverruca defayeae Buckeridge, 1994

Metaverruca defayeae Buckeridge, ly94: 109, fig. 19 a-g.

Material EXAMINED. — Vanuatu. Musorstom 8: stn DW 989. 1 9° 1 3'S, 1 69°20’E, 650-669 m, 23.9.94:

2 specimens, on rock.

Diagnosis. — A large Metaverruca with movable tergum and scutum having 4 articular ribs; exterior smooth

with closely spaced growth lines; mandible quinquedentate, caudal appendages short, less than length of basal

segment of pedicel of cirrus VI.

DISCUSSION. — This material conforms to that described by BUCKERIDGE (1994).

Distribution. — Western Pacific, 370-710 m.

Metaverruca macani (Stubbings, 1936)

Fig 5 a-i.

Verruca macani Stubbings, 1936: 39, figs 17 a-b.

Material examined. — Zanzibar. John Murray Expedition 1933-1934: stn 122, 05°22'S. 39°23’E, 762 m,

1935: 1 specimen, slightly damaged.

Type. — Holotype : Natural History Museum (London) Cat. No. NHM 1938.1.20.154, (unique specimen).

Diagnosis (Amended). — Metaverruca with movable tergum quadrangular, movable scutum and tergum each

with 4 articular ridges; caudal appendages short.

DESCRIPTION (Amended and updated to include internal anatomy). — Medium sized Metaverruca with shell

white, low conic, sides steep; operculum sub-parallel to base; exterior generally smooth, with relatively closely

spaced concentric growth lines; rostrum and carina articulating with 3 ribs; basis membranous. Fixed scutum with

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

139

well formed myophore for adductor muscle attachment; movable tergum quadrangular, with well defined apico-basal

rib, plus 3 secondary ribs, interlocking with movable scutum; articular margin slightly concave; movable scutum

triangular, with 3 ribs interlocking with tergum and further adjacent curved, apico-basal rib terminating at rostro-

carinal suture; occludent portion lacking ribbing; internally with elevated and well formed depression for adductor

muscle attachment; both opercular plates together form almost straight rostro-carinal hinge.

Fig. 5. — Metaverruca macani (Stubbings, 1936): a, holotype, complete shell (dorsal view), BM(NH) Cat. No.

1938.1.20.154; b, movable tergum (right, exterior); c, movable scutum (right, exterior); d, movable scutum (right,

interior); e, cirrus 1; f, cirrus II; g, mandible; h, first maxilla; i, intermediate segment of cirrus VI. All material from

Stn 122.

Mandible tridentate, lower angle pectinate; first maxilla with 2 large upper spines, moderate notch with 1 small

spine, lower angle hirsute, with 4 large spines.

Cirri possess the following number of segments (r-c being the rostro-carinal diameter) :

r-c (mm) I II III IV V VI c.a.

6.4 13,12 12,12 18,18 24,26 24,28 35,36 7

13,15 13,14

Counts for cirri I and II for both sides of animal.

Caudal appendages very short, length less than that of basal segment of pedicel of cirrus VI; penis short.

140

J.S. BUCKERIDGE

DISCUSSION. — Metaverruca macani has not been recorded since the original collection made by the John

Murray Expedition. This species shows some similarity to Metaverruca pacifica Buckeridge. Comments on the

differences between the two species are provided here-in, under the discussion of that species.

Distribution. — Off Zanzibar, 726 m.

Metaverruca maclaughlinae sp. nov.

Fig 6 a-k.

Material EXAMINED. — Vanuatu. Musorstom 8: stn CP 1080, 15°57'S, 167°27'E, 799-850 m. 5.10.1994:

1 specimen, on scaphopod shell.

Type. — Holotype : MNHN-Ci 2429, r-c diameter: 6.7 mm; height: 2.9 mm (1 specimen, from Stn CP 1080,

15°57'S, 167°27'E).

DIAGNOSIS. — Medium Metaverruca with moderately rugose external ribbing; movable tergum and scutum

each having 4 articular ribs; cirrus I with sub equal rami, cirrus 11 with anterior ramus less than half length of

posterior; caudal appendages almost 4 times that of basal segment of pedicel of cirrus VI; penis vestigial.

Description. — Medium sized Metaverruca with shell white, moderate to low conic, sides steep; operculum

sub-parallel to base, orifice D-shaped; exterior characterised by rugose, irregular, vertical ribbing, with relatively

closely spaced concentric growth lines and ridges, secondary ribs develop in interstices of primary ribs as shell

diameter increases; rostrum and carina each articulating with 3 ribs; basis membranous.

Fixed scutum quadrangular, with large, tongue-shaped myophore for adductor muscle attachment; articulating

with fixed tergum with 1 rib and with rostrum with 2 ribs; fixed tergum of similar width to fixed scutum,

articulating with carina with 1 rib; basal margin of shell slightly thickened and inflected.

Movable tergum quadrangular, having well defined apico-basal rib, and 3 secondary ribs, interlocking with

movable scutum; movable scutum triangular, with 3 ribs interlocking with tergum and further adjacent curved,

apico-basal rib terminating and protruding slightly at rostro-carinal suture; internally with moderately weak

depression for adductor muscle attachment developed close to occludent margin; both opercular plates together form

almost straight rostro-carinal hinge.

Mandible tridentate, lower angle pectinate, with 5 stout spines and numerous small setae; first maxilla with

2 prominent upper spines and 1 smaller adjacent spine, lower angle with 4 long and 2 shorter spines. Labrum

moderately deeply arched, basally slightly protuberant, denticulate along entire margin, palps well spaced,

moderately sharp; cirrus I with subequal rami, cirrus II with anterior ramus less than half length of posterior;

intermediate segments of cirrus VI with long upper, medium middle and very small lower pairs of setae.

Holotype cirri possess following number of segments (r-c being the rostro-carinal diameter) :

r-c (mm) 1 II III IV V VI c.a.

67 15,17 9,18 18,21 26,29 28,30 30.29 23

Caudal appendages very long, almost 4 times that of basal segment of pedicel of cirrus VI; penis vestigial.

DISCUSSION. When first observed. Metaverruca maclaughlinae appeared similar to Metaverruca defayae; with

the primary external difference, a rugose surface, being attributed to the type of substrate. The growth of this

verrucid was indeed tortuous, as the diameter of the specimen exceeded that of the scaphopod shell on which it

grew, such that the resulting exposed base required to be sealed with sand grains. However, M. maclaughlinae has

quite distinctive internal anatomy, the caudal appendages are proportionately longer than in any other known

Metaverruca, and the cirri are also unusual, with cirrus I being longer than cirrus II.

Metaverruca maclaughlinae may be distinguished from Metaverruca pacifica Buckeridge by its movable tergum,

which possesses a longer occludent margin, the movable scutum, which lacks longitudinal striae or ribs on the

occludent side, the shorter anterior ramus of cirrus II and the presence of spines in the notch of the first maxilla. It

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

141

differs from M. defay ae primarily by its more rugose external ornamentation and the very much longer caudal

appendages.

Fig. 6. — Metaverruca maclaughlinae sp. nov.: a, holotype, complete shell (dorsal view), MNHN-Ci 2429; b. movable

scutum (left, interior); c, movable scutum (left, exterior); d, movable tergum (left, interior); e, movable tergum (left

exterior); f, basal portion of cirrus VI, with penis and caudal appendage attached; g, cirrus I; h, cirrus II; i, labrum

with palps, setae shown on right palp only; j, first maxilla; k, mandible. All material from Stn CP 1080.

142

J.S. BUCKERIDGE

Etymology. — Named for Dr Patsy McLaughlin, Sedro Woolley, Washington, United States of America,

in recognition of her previous work on cirripedes, and her support and knowledge during my Paris sojourn.

Distribution. — Vanuatu, 799-850 m.

Metaverruca pacifica Buckeridge, 1994

Fig 7 a.

Metaverruca pacifica Buckeridge, 1994: 1 12, figs 1 1 a-g, 16 c-d.

Material EXAMINED. —New Caledonia. MUSORSTOM 6: stn DW 485, 21°23’S, 167°59'E, 346-360 m.

23.2.1989: 1 specimen on a stylasterine coral.

Bathus 1: stn CP 663, 20°58'S, 165°38’E, 730-780 m, 13.3.1993: 3 specimens, on pumice. — Stn CP 708, 21°43’S,

166°38'E, 550-580 m, 19.3.1993: 1 specimen, on gastropod. — Stn CP 709, 21°41'S, 166°37’E, 650-800 m,

19.3.1993: 8 specimens, on pumice.

Bathus 2: stn CP 743, 22°35'S, 166°26’E, 713-950 m, 14.5.1993: 3 specimens, on pumice. — Stn CP 755, 22°2rS,

166°13'E, 495 m, 16.5.1993: 3 specimens, on basalt. — Stn CP 771, 22°09'S, 166°0rE, 610-800 m, 18.5.1993:

1 specimen, on a daphnelline gastropod.

Bathus 3: stn CP 832, 23°03’S, 166°53'E, 650-669 m, 30.11.1993: 13 specimens, on pumice. — Stn CP 850,

21°43'S, 166°39'E, 541-580 m, 19.3.1994: 1 specimen, on pumice. — Stn DW 776, 24°44'S, 170°08'E, 770-830 m,

24.11.1993: 2 specimens, on a gastropod. — Stn CP 821, 23°19'S, 167°58’E, 864-880 m, 29.11.1993: 1 specimen, on a

gastropod.

Halipro 1: stn CP 854, 21°40’S, 166°38'E, 650-780 m, 19.3.1994: 7 specimens, on pebbles and a gastropod. —

Stn CP 867, 21°26’S, 166°18'E, 720-950 m, 22.3.1994: 4 specimens, on a gastropod.

Bathus 4: stn CP 913, 18°08'S, 163°86'E, 777-820 m, 5.8.1994: 5 specimens, on a gastropod shell and detritus.

Loyalty Ridge. MUSORSTOM 6: stn CP 467, 21°05'S, 167°32’E, 575 m, 21.2.1989: 1 specimen.

Wallis-Futuna. MUSORSTOM 7: stn CP 622, 12°34'S, 178°11'W, 1280-1300 m, 28.5.1992: 1 specimen, on a

cidaroid spine.

NW Australia. " Soela stn 6-84, 18°43'S, 166°35'E, 610-612 m, 7.4.1982: 1 specimen, on a scalpellid. —

Stn 1001-86, 18°43'S, 116035'E, 600 m, 7.4.1982: 2 specimens. — Stn 253-95, 18°00'S, 118°10,E, 436 m, 24.2.1984:

28 specimens, on a gastropod.

a b

1 mm 1 mm

Fig. 7 a. — Meiaverruca pacifica Buckeridge, 1994. Complete shell (dorsal view), variety, with distortion resulting from

basal restriction following attachment to the apex of a gastropod. Material from Stn CP 821.

Fig. 7 b. — Meiaverruca recta (Aurivillius, 1898). Complete shell (dorsal view), MNHN-Ci 2423, variety with moderately

rugose exterior and close, well defined growth lines. Material from Stn CP 1 125.

Source :

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

143

DIAGNOSIS. — Metaverruca with movable tergum and scutum having 4 articular ribs; exterior generally

smooth with widely spaced growth lines; caudal appendages more than twice length of basal segment of pedicel of

cirrus VI.

DISCUSSION. — The new material extends both the depth and geographic ranges of M. pacifica as provided in

BUCKERIDGE (1994).

Since the original description of this species, I have reviewed the Verrucidae, and briefly considered that Verruca

macani Stubbings, 1936 may be very similar to this species. Although there is only a single, incomplete

specimen of V. macani known, I was able to view the type in the Natural History Museum, London, and

concluded that the two species are quite distinct. On shell morphology alone, V. macani (here described as a

Metaverruca ), differs in having a more quadrangular movable tergum, the movable scutum lacks apico-basal ridges

and the rostrum does not possess a distinct ribbed zone arcing back to the base of the movable scutum.

A number of specimens of M. pacifica have been recovered from gastropod shells inhabited by parapagurid

hermit crabs (e.g. DW 776, CP 821, CP 867, CP 913). In most instances, M. pacifica is attached at the posterior

of the gastropod shell, and with the somewhat restricted attachment site this provides, the verrucid is unusually

contorted (figure 7a).

Distribution. — From waters off New Caledonia, Loyalty Ridge, Wallis and Futuna Islands, north west

Australia, 346-1300 m.

Metaverruca plicata Buckeridge, 1994

Metaverruca plicata Buckeridge, 1994: 114, fig. 12 a-i.

MATERIAL EXAMINED. — New Caledonia. Bathus 3: stn DW 786, 23°54'S, 169°49’E, 699-715 m. 25.11.1993:

1 specimen, on a scleractinian.

Vanuatu. MUSORSTOM 8: stn DW 989, 19°13'S, 169°20'E, 650-669 m, 23.9.1994: 1 specimen, on rock. —

Stn CP 1125, 15°57'S, 166°38'E, 1160-1220 m, 10.10.1994: 1 specimen, on pebble.

DIAGNOSIS. — A medium to large Metaverruca with strong external ribbing; movable tergum and scutum each

having 3 articular ribs; movable scutum moderately narrow; caudal appendages long, about two and half times

length of basal segment of pedicel of cirrus VI.

DISCUSSION. — These records extend the geographic distribution of M. plicata to the waters off New Caledonia

and Vanuatu. The bathymetric range is doubled. A specimen from CP 1125 conforms to the holotype with the

exception that it has slightly less well developed external ribbing.

Distribution. — South west Pacific, 520-1220 m.

Metaverruca recta (Aurivillius, 1898)

Fig. 7 b.

Verruca recta Aurivillius, 1898: 195.

Verruca sculpta Aurivillius, 1898: 197.

Verruca linearis Gruvel, 1900: 243; 1902: 107, pi. 5 figs 11-12 .

Verruca magna Gruvel, 1901: 261; 1902: 109, pi. 5 figs 1-2 .

Verruca halotheca Pilsbry, 1907: 188, pi. 12 figs 9, 10.

Verruca capsula Hoek, 1913: 130, pi. 12 figs 1-3, pi. 13 figs 1-4.

Verruca coraliophilia Pilsbry, 1916: 21, pi. 1 figs 1-5.

Verruca cookei - ROSELL, 1989: 299, pi. 11 figs r,s,u,v; 1991: 33. Non Verruca cookei Pilsbry. 1927.

Metaverruca recta - BUCKERIDGE, 1994: 1 16, fig 13 a-f.

Material EXAMINED. — New Caledonia. MUSORSTOM 5: stn DW 302, 22°10S, 159°23'E, 346-360 m, 12.10.

1986: 1 specimen, on a turritellid.

144

J.S. BUCKERIDGE

Bathus 1: stn CP 651, 21°41’S, 166°40'E, 1080-1180 m, 11.3.1993: 1 specimen, on pumice. — Stn CP 660,

21°10'S, 165°53'E, 786-800 m, 13.3.1993: 3 specimens, on pumice.

Bathus 2: stn DW 734, 23°00'S, 166°5rE, 766-775 m, 13.5.1993: 1 specimen, on pumice.

Bathus 3: stn DW 778, 24°43'S, 170°07’E, 750-760 m, 24.11.1993: 1 specimen. — Stn CP 842, 23°05’S, 166°48'E,

830 m, 1.12.1993: 1 specimen, on pumice.

Halipro 1: stn CP 858, 21°42'S, 166°41'E, 1000-1120 m, 20.3.1994: 1 specimen, on shell fragment.

Indonesia. Karubar: stn DW14, 05°18'S, 132°38'E, 245-246 m, 24.10.1991: 1 specimen, on pebble.

Grande-Comore. Jago-Coelacanthe: ll°5rS, 43°20'E, 203 m, 1.12.1989: 1 specimen.

Vanuatu. Musorstom 8: stn CP 991, 1 8°5 1 ’S, 168°52’E, 936-910 m, 24.9.1994: 2 specimens, attached to pumice

rock. — Stn CP 956, 20°33,S, 169°35’E, 1175-1210 m, 21.9.1994: 5 specimens, on pumice. — Stn DW 969, 20°18'S,

169°53'E, 252-280 m, 21.9.1994: 1 specimen, on a brachiopod. — Stn DW 977, 19°24'S, 169°28'E, 410-505 m,

22.9.1994: 2 specimens, on pebble. — Stn DW 986, 19°20'S, 169°31'E, 602-648 m, 23.9.1994: 3 specimens, on

pebbles. — Stn CP 990, 18°51'S, 168°50'E, 980-990 m, 24.9.1994: 10 specimens, on pumice. — Stn CP 991, 1 8°5 1 'S,

168°52’E, 936-910 m, 24.9.1994: 8 specimens, on rock. — Stn CP 1008, 18°53’S, 168°52’E, 919-1000 m, 25.9.1994:

3 specimens, on pumice. — Stn CP 1025, 17°49'S, 168°39'E, 385-410 m, 28.9.1994: 1 specimen, on coral. —

Stn CP 1036, 18°0rS, 168°48'E, 920-950 m, 29.9.1994: 6 specimens, on pumice. — Stn CP 1074, 15°48’S, 167°24’E,

775-798 m, 4.10.1994: 35 specimens, on basalt. — Stn CP 1076, 15°53'S, 167°42’E, 1100-1191 m, 4.10.1994:

11 specimens, on pebbles. — Stn CP 1080, 15°57'S, 167°27'E, 799-850 m, 5.10.1994: 5 specimens, on rock, shell

debris. — Stn CP 1095, 15°07'S, 167°irE, 304-320 m, 6.10.1994: 1 specimen, on rock. — Stn DW 1097, 15°05'S,

167°10'E, 281-288 m, 7.10.1994: 5 specimens, on rock. — Stn CP 1110, 14°49’S, 167°15'E, 1360 m, 8.10.1994:

2 specimens, on rock. — Stn DW 1113, 14°53'S, 167°06'E, 700-736 m, 8.10.1994: 5 specimens, on rock. —

Stn CP 1114, 14°52'S, 167°03’E, 647 m, 8.10.1994: I specimen, on rock. — Stn CP 1125, 15°57'S, 166°38’E, 1160-

1220 m, 10.10.1994: 6 specimens, on pebbles. — Stn CP 1126, 15°58'S, 166°39’E, 1210-1260 m, 11.10.1994:

7 specimens, on pebbles. — Stn CP 1129, 16°00'S, 166°39'E, 1014-1050 m, 10.10.1994: 9 specimens, on pebbles. —

Stn CP 1131, 15°38'S, 167°03'E, 140-175 m, 11.10.1994: 5 specimens, on scleractinian. — Stn CP 1137, 15°4rS,

167°02'E, 360-371 m, 11.10.1994: 2 specimens, on gastropod shell.

DIAGNOSIS. — Metaverruca with movable tergum and scutum each having 3 articular ribs; exterior generally

smooth with widely spaced growth lines; caudal appendages short.

DISCUSSION. — Specimen Ci 2423 (fig. 7b) from Vanuatu appeared at first to be a new taxon. Although the

movable opercula are of similar dimension and ribbing to M. recta , the growth lines are much more closely spaced,

and the exterior of the compartments is rather rugose. However, the mandible and first maxilla are found to fall

within the M. recta range, the labrum is broadly curved and finely denticulate, and the cirri possess the following

number of segments (r-c being the rostro-carinal diameter):

r-c (mm) I II III IV V VI c.a.

7.9 13,13 10,15 19,21 24,27 25,28 28,29 9

The caudal appendages are slightly shorter than the basal segment of the pedicel of cirrus VI; and intermediate

segments of cirrus VI possess 3 pairs of spines each (as in V recta).

I am cautious about using the spacing of growth lines as a criterion for specific differentiation, and in this case

consider that they merely reflect a growth rate variable from the norm.

M. recta is the most cosmopolitan of verrucids, with material being recorded from all seas. It is also the largest

verrucid known, with r-c diameters of over 14.5 mm being recorded (BUCKERIDGE, 1994).

The three articular ribs between movable scutum and tergum, the generally inornate exterior, the fixed scutum

myophore, and the short caudal appendages distinguish this from all other verrucids. This material conforms to that

described in BUCKERIDGE (1994), and extends depth range, the upper limit changing from 160 to 140 m.

Distribution. — Cosmopolitan, 140-2110 m. Miocene (New Zealand, BUCKERIDGE, 1983).

Genus ROSTRATOVERRUCA Broch, 1922

Verruca "Sectio Rostrato-verruca " Broch, 1922: 298.

Verruca (Rostratoverruca) - FOSTER, 1978: 68.

Rostratoverruca - Zevina, 1987: 1813.

Source :

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

145

DIAGNOSIS. — Verrucids with apex of rostrum removed from margin; fixed scutum without myophore;

operculum sub-parallel to base.

TYPE Species. — Verruca nexa Darwin, 1854: 522. West Indies, 60 m.

Species. — The following 8 species are assigned to this genus: Rostratoverruca conchula (Hoek, 1913), Timor

Sea; R. intexta (Pilsbry, 1912), Indian Ocean - Western Pacific; R. koehleri (Gruvel, 1907), Bay of Bengal;

R. kruegeri (Broch, 1922), Western Pacific; R. murrayi (Stubbings, 1936), Zanzibar; R. nexa (Darwin, 1854),

West Indies; R. sewelli (Stubbings, 1936), Zanzibar; R. malevichi Zevina, 1988, South Pacific.

Stratigraphic Range. — Recent.

Distribution. — West Indies and Indo-West Pacific, 60-3250 m.

Rostratoverruca intexta (Pilsbry, 1912)

Fig 8 c-d.

Verruca intexta Pilsbry, 1912: 292; 1916: 47. — NlLSSON-CANTELL, 1927: 774; 1929: 468, fig. 3. — Stubbings, 1940:

389.

Verruca conchula Hoek, 1913: 146, figs 14-15.

Verruca (Rostratoverruca) intexta - ROSELL, 1989: 26, pi. 7 f-g; 1991: 33.

Rostratoverruca intexta - BUCKERIDGE, 1994: 119, fig 14 a-f.

Material EXAMINED. — New Caledonia. Halipro 1: stn CP 850, 21°43'S, 166°39’E, 541-580 m, 19.3.1994:

4 specimens, on sponge. — Stn CP 850, 21°43’S, 166°39’E, 541-580 m, 19.3.1994: 4 specimens, on sponge.

Loyalty Ridge. Musorstom 6: stn CP 455, 21°00’S, 167°26'E, 260 m, 20.2.1989: 1 specimen, on octocoral.

Indonesia. Karubar: stn CP 25, 05°30'S, 132°52'E, 336-346 m, 26.10.1991: 1 specimen, on sponge. —

Stn CP 62, 09°01'S, 132°42'E, 246-253 m, 1.11.1991: 3 specimens, on octocoral. — Stn CP 69, 08°42’S, 131°53’E,

356-368 m, 2.11.1991: 4 specimens, on coral.

Vanuatu. MUSORSTOM 8: stn CP 1028, 17°54'S, 168°40'E, 624-668 m, 28.9.1994: 2 specimens, on cidaroid spine.

NW Australia. "Soela": stn 116-84, 15°44'S, 120°39'E, 430-446 m, 7.4.1982: 1 specimen, on a scalpellid. —

Stn 1001-86, 18°43’S, 116°35’E, 610-612 m, 7.4.1982: 2 specimens. — Stn 254-95, 18°49’S, 120°40'E, 396-400 m,

10.2.1984: 2 specimens, on hexactinellid sponge. — Stn 909-86, 13°33’S, 122°14’E, 494-496 m, 13.2.1984:

2 specimens, on cidaroid spine. — Stn 967-85, 16°56'S, 119°54'E, 436 m, 22.2.1984: 3 specimens, on hexactinellid

sponge spicules. — Stn 968-85, 16°56’S, 1 1 9°53’E, 436 m, 22.2.1984: 2 specimens, on hexactinellid. — Stn 253-95,

16°56'S, 1 19°55'E, 436 m, 22.2.1984: 1 specimen, on hexactinellid. — Stn 966-85, 18°00*S, 1 1 8°10E. 436 m,

24.2.1984: 1 specimen, on gastropod.

DIAGNOSIS. — Rostratoverruca with movable tergum and scutum each with 3 articular ribs; operculum sub¬

parallel to base.

DISCUSSION. — Although this material is consistent with that described in BUCKERIDGE (1994), some

specimens provide evidence of the colour of the shell, which in the samples from station CP 455 is a distinct

mottled, reddish-brown on white. In BUCKERIDGE (1994), the shell is described as "cream-white". The lack of

colour or patterning in many collections may be attributed to the preservatives causing colour fade. One specimen

was found growing within the coral Madrepora oculata. The verrucid had evidently reached adult size without being

overgrown by the coral, indeed, the action of the cirri appear to have either prevented deposition, or eroded the coral

skeleton near the verrucid orifice (fig. 8d).

DISTRIBUTION. — Indo-Pacific, 194-1002 m.

Rostratoverruca kruegeri (Broch, 1922)

Verruca Kriigeri Broch, 1922: 295, figs 43-44.

Verruca ( Rostratoverruca ) kriigeri - BROCH, 1932: 46.

Rostratoverruca kruegeri - BUCKERIDGE, 1994: 121, fig 15 a-f.

146

J.S. BUCKERIDGE

Fig. 8. — Verrucidae: Scanning Electron Microscope Photographs,

a-b, Altiverruca nitida (Hoek, 1883): a, complete shell (dorsal view, x 15); b, complete shell (fixed opercula view, x 15).

Material from Stn CP 30.

c-d, Rostratoverruca intexta (Pilsbry, 1912): c, complete shell (dorsal view, x 19). Material from Stn CP 69; d, shell

growing within the coral Madrepora oculata , note vertical grooves in the coral adjacent to the verrucid orifice,

possibly caused by cirral movement (dorsal view, x 15). Material from Stn CP 455.

e-i, Verruca jago sp. nov.: e, complete shell (carino-rostral view, x 37); f, complete shell (dorsal view, x 37); g, detail of

operculum from f (x 95); h, detail of porous surface from g, note desiccated organic material in centre of some pores

(x 500); i, detail of pore from h (x 4000). Material from 11°51’S, 43°20'E.

Material EXAMINED. — Vanuatu. Musorstom 8: stn CP 973, 1 9°2 1 *S, 169°27’E, 460-480 m, 22.9.1994:

311 specimens, on cidaroid spines. — Stn CP 974, 19°21'S, 169°28*E, 492-520 m, 22.9.1994: 66 specimens, on

Source :

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

147

cidaroid spines. — Stn CP 975, 19°23'S, 169°28'E, 536-566 m, 22.9.1994: 4 specimens, on cidaroid spines. —

Stn CP 1107, 15°05'S, 167°15'E, 397-402 m, 7.10.1994: 105 specimens, on cidaroid spines. — Stn DW 1108, 15°04'S,

167°15'E, 425-455 m, 7.10.1994: 65 specimens, on cidaroid spines.

DIAGNOSIS. — Rostratoverruca with fixed scutum and tergum possessing weak longitudinal ribbing; movable

tergum and scutum each with 4 articular ribs.

DISCUSSION. — This material is consistent with that described in BUCKERIDGE (1994). In addition, it extends

the bathymetric range from 290 to 566 m.

Distribution. — Western Pacific, 233-566 m.

DISCUSSION

Distribution. — The material studied in both this paper and in Buckeridge (1994) was collected from

depths ranging between 223 and 3680 m. This study extends the bathymetric ranges for the Indo-Southwest Pacific

region, as provided in BUCKERIDGE (1994). The following provides a broad appreciation of the current known

bathymetric distribution of vemicid genera for this region:

*only 2 specimens of Altiverruca were recovered from depths less than 510 m.

In the following table, data from this paper and BUCKERIDGE (1994) are combined to provide an overview of

the geographic distribution of verrucid species in the Indo-Pacific region. Verrucid genera listed in the table

(horizontally) are, respectively: Verruca, Altiverruca , Came rave rruca, Brochiverruca , Metaverruca , and Rostrato¬

verruca

Material recovered from waters off:

Philippines

New Caledonia

Vanuatu

Wallis and Futuna

Chesterfield

Loyalty Ridge

Indonesia

Northwest Australia

Grande-Comore

©

*

a

©

I §

u

l-s

a o

ccs:

© —

2 -s

©

©. -©

2 .2

•Cl

•c c

© ©

© ©

t!

§ ©

s

©

©. ©.

OC

©

©

. X . -----XXX

-XX - XXX - - - X - - XX XXX-

- - X- X - -X - - - - xx-xxxxx

- - - - - - - X - - - - X - - X - X - -

. XX . X-X--

. XX--X--XXXX-

x . X---X . XXX

X . X--X-

-x . ------ X - -

Source

148

J.S. BUCKERIDGE

ENDEMISM. — A number of verrucid taxa are known from only one location, often by a unique specimen. It is

reassuring then that some of the rarer taxa, described in Buckeridge (1994), after further collecting, have had their

geographic ranges extended, e.g. Metaverruca defayeae , Metaverruca plicata. The areas with highest biodiversity are

also characterised by high verrucid endemism: Vanuatu has two, Loyalty Ridge one, and New Caledonia two

endemic species. These three localities are situated in a transition area between tropical and subtropical zones; in

addition, the ocean bathymetry is characterised by numerous highs of volcanic origin, in all providing ideal

opportunities for speciation. Although the abundance of taxa recovered from waters off New Caledonia and Vanuatu

mark this region as one of extraordinary verrucid diversity, I am also confident that this reflects the extensive and

comprehensive collecting programme undertaken in the area by Musorstom.

Substrate. — Verrucids are generally not particular in their choice of substrate, the most common being the

abundant benthic debris (generally small pebbles) found at many stations, although broken shell material has also

been utilised. Both Metaverruca recta and Metaverruca pacifica have been found living on gastropod shells inhabited

by parapagurid crabs. These two large verrucids, like many cirripedes, are commensals, and are likely to have

benefited from the feeding habits of their hosts. Some verrucids appear to be host specific, in particular

Brochiverruca species, which are always found attached to cnidarians, and Rostratoverruca kruegeri , always found

attached to cidaroid spines.

ACKNOWLEDGMENTS

The author wishes to thank Alain CROSNIER. ORSTOM (Institut Francis de Recherche scientifique pour le

Developpement en Cooperation), for the invitation to work on the MUSORSTOM collections, for providing access

to the material, and for providing a stimulating working environment. Patsy McLaughlin, also working at the

Museum national d'Histoire naturelle, provided thoughtful comments and contributed to a good working

environment. I also thank Professor Bill Newman, of Scripps Institution of Oceanography for valuable comments

on the manuscript, Paulo Young, of Departemento de Invertebrados, Museu Nacional, Rio de Janeiro, Brasil, for

alerting me to problems with the previous diagnoses for verrucid genera, Danielle Defaye. Museum national

d'Histoire naturelle, Paris, for providing assistance with microscopy, Helmut ZIBROWIUS, Station Marine

d'Endoume, Marseille, who identified the coral substrates of both Rostratoverruca intexta and Brochiverruca

crosnieri, and Paul COLLINS, Natural History Museum, who made the holotype of Metaverruca macani available

for study.

REFERENCES

AURIVILLRJS, C.W.S., 1898. — Cirrhip&des nouveaux provenant des campagnes scientifiques de SAS le Prince de Monaco

Bulletin de la Societe zoologique de France , 23: 189-199.

BROCH, H 1922. — Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. No. 10: Studies on Pacific Cirripeds.

Videnskabelige Meddelesler fra Dansk Nalurhistorisk Forening i Kjdbenhavn, 73: 215-358.

BROCH, H 1932. - Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. No. 56: Indomalayan Cirripedia.

Videnskabelige Meddelesler fra Dansk Nalurhistorisk Forening i Kjdbenhavn, 91: 1-146.

Buckeridge J.S., 1983. — Fossil barnacles (Cirripedia: Thoracica) of New Zealand and Australia. New Zealand

geological Survey Paleontological Bulletin, 50: 1-151.

Buckeridge, J.S., 1994. — Cirripedia Thoracica: Verrucomorpha of New Caledonia, Indonesia, Wallis and Futuna Islands

In. A. CROSNIER (ed.), Resultats des Campagnes MUSORSTOM. Volume 12. Memoires d„ Museum national d’Histoire

naturelle , 4: 87-126.

Darwim c., 1854 — A monograph on the sub-class Cirripedia, with figures of all species. The Balanidae, the Verrucidae

Ray Society, London. 684 pp„ pis 1-30.

Darwin. C., 1855. — A monograph on the fossil Balanidae and Verrucidae of Great Britain. Palaeontological Society

Monograph, London (1854). 44 pp., pis 1-2. y

VERRUCOMORPHA OF THE INDIAN AND SOUTH WEST PACIFIC OCEANS

149

°f N™ B“‘“ »» *— -r#

F°STER, B A. & BUCKERIDGE, J. S„ 1995 — Barnacles (Cirripedia: Thoracica) of Seas of Reunion Island. Bulletin du

Museum national d Histoire naturelle , 16, Section A, 1994 (1995), 4e S6r., No 2-4: 345-382.

_SUr 165 efPi,C,eS nouvelles appartenant au genre Verruca provenant de la campagne du Talisman

Bulletin du Museum national d Histoire naturelle , 6: 242-244.

GRnarUL;W/e,l69025^263a8nOSeS * qUe'qUeS n0UVelleS de CirrhipMes. Bulletin du Museum national d'Histoire

GTCE0V M- 1902' — CinifipMcs. In: Expeditions scientifiques du Travailleur et du Talisman pendant les anndes 1880-

1583. Masson et Cie, Pans. 178 pp.

Gruvel, A.. 1905. — Monographic des Cirrhip&des ou Thecostraces. Masson et Cie. Paris. 472 pp.

GRrEuin” l907' ~CirrhiPfedes °Percules de l'lndian Museum de Calcutta. Memoirs of the Asiatic Society of Bengal,

Hoek, P.P.C 1881 - Report on the Cirripedia collected by H.M.S. Challenger. Report on the Scientific Results of the

Voyage of the H.M.S. Challenger during the Years 1873-1876. Zool., 8 (25): 1-169, pis 1-13.

H ° Zoo logie'' 3 1907 ~ Cirr‘Pedia' ln: R^ultats du Voyage du S.Y. Belgica en 1897-1899. Rapports scientifiques.

H° 3*ibP P129-27>51 3 ~~ ThC Cirriped'a °f ,he Sib0ga ExPedilion- B. Cirripedia Sessilia. Siboga-Expeditie monograph.

Newman, W.A., 1989.

Neoverruca , from an

— Juvenile ontogeny and metamorphosis in the most primitive living sessile barnacle,

abyssal hydrothermal spring. Bulletin of Marine Science, 45 (2): 467-477.

Newman, W.A. & Hessler, R.R., 1989. — A new abyssal hydrothermal verrucomorphan (Cirripedia: Thoracica): The

most primitive living sessile barnacle. Transactions of the San Diego Society of Natural History, 21 (16): 259-273.

Newman^ W.A Zullo, V.A. & Withers, T.H., 1969. - Cirripedia: 206-295. In: Treatise on Invertebrate Paleontology

Part R, Arthropoda 4(1). Geological Society of America, Boulder. 398 pp.

Nilsson-Cantell C.A., 1927. — Some barnacles in the British Museum (Nat. Hist.). Proceedings of the Zoological

Society of London, 3: 743-790.

PlLSBRY, H.A., 1907. — Hawaiian Cirripedia. Bulletin of the Bureau of Fisheries, Washington, 26: 181-190.

Pilsbry, H A., 1912. — Diagnoses of new barnacles from the Philippine Archipelago and China Sea. Proceedings of the

United States National Museum, 42: 291-294.

PILSBRY H. A., 1916. —The Sessile barnacles (Cirripedia) contained in the collections of the U.S. National Museum:

Including a Monograph of the American species. Bulletin of the United States National Museum, 93: 1-366.

Rosell, N.C., 1989. — Thoracic cirripeds from the Musorstom 2 Expedition. In: R6suhats des Campagnes Musorstom

Volume 5. Memoires du Museum national d'Histoire naturelle. (A), 144: 9-35.

Rosell N.C., 1991. — Crustacea Cirripedia Thoracica: Musorstom 3 Philippines collection. In: A Crosnier (ed.)

Resultats des Campagnes MUSORSTOM. Volume 9. Memoires du Museum national d'Histoire naturelle, (A), 152: 9-61.

Schumacher, H.C.F., 1817. — Essai d'un nouveau systdme des habitations des vers testaces. Copenhagen. 287 pp„

STUBBINGS, H.G., 1936. — Cirripedia. Scientific Reports of the John Murray Expedition, 1933-34, 4 (1): 1-70.

ST“oH.G.' 194°- — Cirripedia, additional part. Scientific Reports of the John Murray Expedition, 1933-34, 7 (3):

ZEVr: Gc® ,’ 1987' ~ Deep-sea Verrucomorpha (Cirripedia. Thoracica) of the Pacific. Zoologichesky Zhumal.66:

Zevina, G.B., 1993. — The new genus and the new species of Verrucomorpha (Cirripedia). Zoologichesky Zhurnal, 72: 9-

Source : MNHN. Paris

RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULTATS DES C

Crustacea Isopoda Serolidae: Acutiserolis cidaris

and Caecoserolis novaecaledoniae , two new species

from the Coral Sea

Gary C. B. POORE

Department of Natural Sciences

Museum of Victoria

71 Victoria Crescent, Abbotsford

Victoria 3067, Australia

&

Angelika BRANDT

Zoologisches Institut und Museum

Universitaet Hambourg

Martin-Luther-King-Platz 3

20146 Hamburg, Deutschland

ABSTRACT

Two new species of Serolidae are described from deep waters in the Coral Sea : Acutiserolis cidaris sp. nov. from 891-

1491 m depth and Caecoserolis novaecaledoniae sp. nov. from 500-2750 m depth. A. cidaris is compared with

A. bromleyana , illustrated and briefly diagnosed, from the Southern Ocean. A. bromleyana may be a variable species

widely distributed in the Southern Ocean and more than one similar species exist in deep seas in lower latitudes. Both

genera are rediagnosed.

RESUME

Crustacea Isopoda Serolidae: Acutiserolis cidaris et Caecoserolis novaecaledoniae , deux

especes nouvelles de la mer du Corail.

Deux nouvelles esptces de Serolidae, provenant des eaux profondes de la mer de Corail, sont decrites : Acutiserolis

cidaris sp. nov., rtcokte & des profondeurs comprises entre 891-1491 m et Caecoserolis novaecaledoniae sp. nov.

trouvte entre 500-2750 m de profondeur. A. cidaris , trouvee dans 1'octan Austral, est comparee avec A. bromleyana,

britvement dtcrite et illustrte ici. A. bromleyana pourrait etre une esptce variable, largement distribute dans 1’octan

Austral, tandis que plusieurs esptces voisines existent en eau profonde aux basses latitudes. Les deux genres considers ici

sont redtfinis.

POORE, G.C.B. & Brandt, A., 1997. — Crustacea: Isopoda Serolidae: Acutiserolis cidaris and Caecoserolis

novaecaledoniae, two new species from the Coral Sea . In : A. Crosnier (ed.), Rtsultats des Campagnes Musorstom.

Volume 18. Mem . Mus . natn. Hist . nat., 176 : 151-168. Paris ISBN 2-85653-511-9.

152

G.C.B. POORE & A. BRANDT

INTRODUCTION

Following extensive earlier reviews (Nordenstam, 1933; Sheppard, 1933) the history of the taxonomy of

the isopod crustacean family Serolidae Dana, 1853 was reviewed by BRANDT (1988, 1991) and WAGELE (1994)

who established new genera and subgenera to bring the total described to 20. BRANDT (1991) and WAGELE (1994)

both proposed phylogenies of the genera based on morphological characters.

French expeditions near New Caledonia and Australian expeditions along the eastern coast of Australia have

collected serolids in deep water and this contribution describes two new species from tropical waters. The First

species is similar to Acutiserolis bromleyana Willemoes-Suhm originally described from Antarctic waters and

reported from New Zealand by HURLEY (1961). Adults of our material were much smaller than adults of the

Antarctic species and differed morphologically from it and the New Zealand specimens. Investigation leads us to

believe that it was one of several similar species and that A. bromleyana itself may be variable. The new species is

described and its variability briefly discussed.

The second species was placed only with difficulty in any of the genera diagnosed by WAGELE (1994) and in

the phylogenies proposed by him and by BraNDT (1991) but is assigned to Caecoserolis Wagele, 1994.

METHODS

The material comes from two sources : deep dredging during cruises near New Caledonia organised by

ORSTOM and the Museum national d'Histoire naturelle, Paris; and dredging in northeastern Australia by James

Cook University, Townsville, Australia. It is deposited in the Museum national d'Histoire naturelle, Paris

(MHNH), the Museum of Tropical Queensland, Townsville, Australia (MTQ), and the Museum of Victoria,

Melbourne, Australia (NMV).

All illustrations are of left limbs unless otherwise noted, labelled as follows : Al, A2, antennae 1, 2;

MD, mandible; MDp, mandibular palp; MX 1, 2, maxillae 1, 2; MP, maxilliped; P1-P7, pereopods 1-7;

PL1-PL5, pleopods 1-5: U, uropod; 1, left; r, right.

SYSTEMATIC ACCOUNT

Genus ACUTISEROLIS Brandt, 1988

Acutiserolis Brandt, 1988 : 21; 1991 : 131, 139.

Serolis (Acutiserolis) - WAGELE, 1994 : 53, 60.

Diagnosis. — Body broad, weakly-sculptured, slate-grey, lateral margin strongly divided from head to

pleonites; pleotelson with weak dorsal lateral submarginal ridges. Eyes reniform, large, lateral. Pereonites 5-7 and

pleonite 1 fused. Pereonal sternite 1 visible, sternites 2-4 invisible (ventral coxal plates meeting in midline),

sternites 5-7 fused. Pleonal sternites 1-3 with 3-comered medial plate, not sexually dimorphic.

Coxal plates with interacting keys, otherwise not in contact distal to bases of pereopods and acutely projecting

and tapering; coxal sutures 2-4 visible dorsally; coxa 6 strongly exceeding pleonal epimera 2 and 3 posteriorly.

Pleonites 2 and 3 with epimera narrow and strongly produced posteriorly.

Mandible with incisor process untoothed; left lacinia mobilis three-quarters width of incisor process, right

toothed and narrower. Maxilla 2 with middle and outer lobes as long as inner, each with 2 distal setae. Maxilliped

with free coxa, epipod and lamella on basis; palp article 2 cordiform.

Pereopod 1 with alternating spiniform and plate-like setae. Pereopod 2 of male with propodus palm having

U-shaped marginal row of spiniform setae; unguis small and terminal; pereopod without felt of fine hairs.

Pereopod 7 of male with propodus little broader than in female, setose and with felt of fine hairs; dactylus simple.

NEW SPECIES OF SEROLIDAE FROM THE CORAL SEA

153

Pleopod 2 of male with endopod tapering. Pleopod 4 endopod not bilobed. Uropod biramous, peduncle short;

inserting half-way along pleotelson.

Oostegites present on pereopods 1-4 of female.

REMARKS. — Brandt (1988) diagnosed the genus and listed six species of Acutiserolis. We present an

expanded diagnosis. In 1991 she discussed its relationship to other genera, differentiating it from them with a

single autapomorphy, extreme posterior elongation of the coxal plates and pleonal epimera. WAGELE (1994) added

a seventh species while reducing the taxon to a subgenus of Serolis Leach. WAGELE placed Acutiserolis on a clade

in his dendrogram with Serolis , sharing two synapomorphies : a bilobed endopod on pleopod 4 and enlarged

posterior coxal plates. While S. paradoxa Leach, type species of Serolis s.s. has a bilobed endopod (POORE, 1987),

none of the species of Acutiserolis seen by us does. Nor do species of Serolis s.s. have the extremely elongated

coxal plates of Acutiserolis. For these reasons we do not accept the subgeneric status of Acutiserolis.

Species of Acutiserolis differ from each other primarily in adult size, sculpture and setation of the male

pereopods. Material available to us from New Zealand and from deep water off southern Australia contains at least

three undescribed species. Beddard (1884b) and Hurley (1961) referred material from the Tasman Sea and the

vicinity of New Zealand to A. bromleyana but only Beddard noted differences from the syntypes from Antarctic

waters. He figured a male from New Zealand which may be the species figured by Hurley. A second species

occurs off the southeastern coast of Australia but is not dealt with in this paper.

Here, we describe the third species which comes from the Coral Sea and the northern part of the Tasman Sea as

new. It is certainly not the species mentioned by other authors.

We also comment on A. bromleyana and material from Antarctic waters which may belong to this species.

Acutiserolis bromleyana (Willemoes-Suhm, 1874)

Figs 1-2

Serolis bromleyana Willemoes-Suhm, 1874 : XIX; 1876 : 591. — Beddard, 1884a : 331; 1884b : 53-57, pi. 4 (except

figs 3, 6). — Sheppard, 1933 : 280, 329-330.

Acutiserolis bromleyana - Brandt, 1988 : 17, 21.

Serolis ( Acutiserolis ) bromleyana - WAGELE, 1994 : 53.

not Serolis bromleyana - HURLEY, 1957 : 13; 1961 : 228-229, pi. 1.

MATERIAL EXAMINED. — Southern Ocean (Indian Ocean sector). "Challenger" : stn 156, 62°26'S,

95°44’E, 3614 m, 26 February 1874, diatom ooze bottom : 6 53 mm, and 9 45 mm, syntypes (BMNH 1889.4.27.20).

DIAGNOSIS OF MALE. — Greatest width of body (between tips of coxae 4) 105% of body length (rostrum to end

of pleotelson). Head with broad diverging anterolateral lobes lateral to bases of antennae, acutely produced

anterolaterally; continuous sharp transverse ridge at base of antennae; with pair of low elevations between anterior

margins of eyes and medial and pair of posterolateral elevations along posterior margin.

Pereonite 1 with anterior margin (medial to small triangular projection) concave; lateral margin shallowly

sinuous; posterolateral corner produced as an acute triangular plate, convex posteriorly; sharp oblique ridge runs

from near eye to meet an angled submarginal ridge, sharply defined between oblique ridge and anterior projection

and more obscure on posterolateral plate. Pereonites 2-4 articulating, with coxal plates marked off by dorsal

sutures; pereonites 5-7 fused, with groove between 5 and 6 dorsally; all pereonites with small spine on mid-dorsal

margin. Posterolateral angles of the coxal plates 2-6 longer than and reaching much further posteriorly than those

of preceding segments; coxal plate 4 reaching to base of pleotelson, 5 reaching 70% of length of pleotelson,

6 exceeding total body length by 38%). Pereonite 7 without coxal plates. Pleonite 1 not visible dorsally.

Pleonites 2 and 3 with narrow elongate diverging epimera, apically notched; (epimera 2 both broken); epimeron 3

only slightly exceeding pleotelson. Pleotelson 34% of total length, 1.2 times as wide as long, tapering to rounded

posterolateral corners and concave posterior margin; with long slight medial keel and obscure triangular elevations

near angle at base of uropods; a narrow medial pit at base of keel.

154

G.C.B. POORE & A. BRANDT

Pereopod 2 without fine setae; propodus cylindrical proximally, with heel about halfway along, greatest width

half of length, palm with 18 spiniform setae in irregular U-shaped marginal row plus 1 spiniform seta in middle of

oblique concave palm; dactylus strongly curved and overlapping heel of palm, unguis absent (or broken).

Fig. 1. Acutiserolis bromleyana (Willemoes-Suhm). Male syntype, BMNH 1889 4 V 20 •

from damaged specimen), scale bar = 10 mm; b, detail of head and pereonite 1.

a. habitus (reconstructed

\ ru ~ t gn™S dealS With lhe 0Vera11 shaPe and scu'P[ure of lhe body and the male

rnmhP r2' ** '° d,fferentiate ,hls sPecies fr°m the new species described herein and others. The

onThe nrnL r mtenT SP'ne ^ ^ ^ Submarginal ridge on pereonite 1. and a short concave palm

on the propodus of the male pereopod 2 serve to diagnose the species.

Source :

NEW SPECIES OF SEROLIDAE FROM THE CORAL SEA

155

Fig. 2. — Acutiserolis bromleyana (Willemoes-Suhm). Female syntype, BMNH 1889.4.27.20 : maxillae 1, 2,

maxilliped, pereopod 4, pleopods 1, 4, 5, uropod. — Male syntype, BMNH 1889.4.27.20 : pereopod 2 propodus and

dactylus, palm in face view; pleopod 2 (appendix masculina lost).

The syntypes are damaged. Our figures of the other limbs are incomplete but show only very slight differences

in proportions and setation from the new species.

Besides the two syntypes from the Southern Ocean, the Challenger collected other similar specimens, listed by

Beddard (1884b), and some of these remain in the Natural History Museum, London. These were dredged from

156

G.C.B. POORE & A. BRANDT

off the coast of New South Wales, Australia, and off New Zealand, after WiLLEMOES-SUHM wrote his description

(while the " Challenger " was in Sydney, April 1874) and are therefore not syntypes. According to BEDDARD

(1884b) these differed in features which we now regard as of specific importance. He illustrated (1884a : pi. 4

figs 3, 6) a male from New Zealand without the anterior spine on pereonite 1, curved coxal plates 6 and 7, and

setose male pereopod 2 with long palm, all characters differentiating it from the syntypes. D.E. Hurley too had

difficulty with the New Zealand material and recognised three forms with different depth ranges (in litt. to GCBP,

16 May 1984).

Acutiserolis sp.

Fig. 3

Material EXAMINED. — Southern Ocean (West Scotia Basin) : 57°12'S, 62°45'W to 57°14’S, 62°51’W,

3733-3806 m, 29 July 1962, USN "Eltanin" : 6 40 mm (NMV J 1 1 639, part of larger collection : USNM 12391 1).

Remarks. A male from the West Scotia Basin, on almost the opposite side of Antarctica from the type

locality, differs from the types of A. bromleyana in size and at least two morphological characters. Although this

male is smaller than other species, adult male features are generally seen in individuals over quite a size range. The

more pronounced, erect submarginal oblique ridge on pereonite 1 might be interpreted as a local variation.

However, the shape of the male pereopod 1 differs substantially from that of the syntypes. Its heel is more

proximal, the palm more elongate, and there are many more (30) and finer setae along its margins. MENZIES

(1962) described A. macdonnellae and A. maryannae from the South Atlantic but his material and figures are too

poor to reconcile with this specimen. Circumpolar variation or speciation in Acutiserolis remains to be

investigated before the status of this specimen can be determined. WAGELE (1986) concluded after studying

variability in Ceratoserolis trilobitoides (Eights, 1833) that this is a polymorphic circum-Antarctic species. The

same may be true for A. bromleyana.

Source :

NEW SPECIES OF SEROLIDAE FROM THE CORAL SEA

157

Acutiserolis cidaris sp. nov.

Figs 4-6

TYPE MATERIAL. — Coral Sea, Australia, near Townsville. Cidaris 1 (all beam trawl samples) : stn 31-1,

1 7° 1 2. 1 5'S, 147°10.80'E, 1489-1491 m, 12 May 1986 : 1 6 holotype, 25.1 mm (MTQ W13598). Paratypes : 5 6 26.9-

28.1 mm; 2 ovigerous 9 26.9 mm; 1 juvenile 9 (NMV J27642). — Stn 1-4, 18°08.69’S, 147°33.97’E, 966-962 m,

6 May 1986 : 6 22.1 mm (MTQ W13599). — Stn 8-1, 18°07.82'S, 148015.39*E, 1115-1119 m, 7 May 1986 : 3 6 21.2-

24.0 mm. 2 juvenile 9 15.4-22.4 mm (MTQ W13600). — Stn 11-4, 18°10.06’S, 148°32.44’E, 1121-1123 m, 8 May

1986 : 2 9 20.4-20.6 mm, juvenile 9 16.9 mm (MTQ W13601). — Stn 13-1, 17°58.49’S, 148°38.40'E, 1040-1059 m, 8

May 1986 : 6 6 20.1-21.8 mm, 1 ovigerous 9 21.0 mm. 1 juvenile 16.7 mm (MTQ W13602). — Stn 50-3, 18°01.69'S,

147°20.53'E, 918-891 m, 17 May 1986 : 3 6 21.0- 22.1 mm, 1 juvenile 9 15.7 mm (MTQ W13603).

NON-TYPE MATERIAL — Chesterfield Islands. MUSORSTOM 5 : stn 324, 21°15.01'S, 157°51.33'E, 970 m,

22 October 1986 : 1 6 19.8 mm (MNHN-Is 3068); 3 6 22-24 mm. and 3 9(1 ovigerous) 21-23 mm (MNHN-Is 3069).

Description OF male holotype. — Colour slaty-grey. Greatest width of body (between tips of coxae 4) 93%

of body length (rostrum to end of pleotelson). Head with broad diverging anterolateral lobes lateral to bases of

antennae, acutely produced anterolaterally; continuous sharp transverse ridge at base of antennae; with broad low

anteromedial elevation between eyes, posterolaterally rugose.

Fig. 4. — Acutiserolis cidaris sp. nov. Male holotype, MTQ W 13598 : a, habitus (reconstructed from damaged specimen),

scale bar = 10 mm; b, detail of head and pereonite 1; c. ventral view of pereonites 4-7 and pleonites 1-3.

Pereonite 1 with anterolateral margin convex and upturned over anterior quarter, straight posteriorly;

posterolateral comer produced as an acute triangular plate, concave posteriorly; a sharp oblique ridge runs from near

eye to meet a low pocked submarginal ridge, broad anteriorly and narrower on posterolateral plate. Pereonites 2-4

158

G.C.B. POORE & A. BRANDT

articulating, with coxal plates marked off by dorsal sutures; pereonites 5-7 fused, with groove between 5 and 6

dorsally. Posterolateral angles of the coxal plates 2-6 longer than and reaching much further posteriorly than those

of preceding segments; coxal plate 4 reaching to base of pleotelson, 5 reaching 85% of length of pleotelson;

6 broken on all specimens, exceeding total body length by over 60%. Pereonite 7 without coxal plates.

Ventral coxal plates 2-4 meeting in midline, smooth; sternites 5-7 fused, with paired small penes posteriorly;

coxal plates 5-7 fused, 7 visible only ventrally. Pleonite 1 not visible dorsally. Pleonites 2 and 3 with narrow

elongate diverging epimera, apically notched; epimeron 2 90% of length of pleotelson and epimeron 3 exceeding

pleotelson by 57% of its length. Pleonite 1 with 3-cornered sternal plate with median ridge between bases of

pleopods; pleonites 2 and 3 similar but less well developed. Pleotelson 35% of total length, little wider than long,

posterolateral corners rounded and posterior margin convex; with long elevated medial keel and obscure triangular

elevations near angle at base of uropods; a narrow medial pit at base of keel.

Source :

NEW SPECIES OF SEROLIDAE FROM THE CORAL SEA

159

Antenna 1 with peduncular articles 1 and 2 of similar width, article 2 about twice as long as 1, article 3 almost

as long as first two together; flagellum of over 50 articles; flagellar article 1 longest, one-quarter length of last

peduncle article, articles 1-2 without aesthetascs, remainder with 1 long aesthetasc and 2-3 simple setae. Antenna 2

with short first peduncular article, second longer than third, with few proximal short setules, third with medial and

lateral setae, fourth and fifth with tufts of lateral and dorsal setules, fifth 1 .3 times as long as fourth; flagellum of

at least 19 articles, each with group of 4 distolateral simple setae and 1 on opposite side.

Mandibles : Left lacinia mobilis a broad blade as wide as incisor, spine (spine row rudiment) simple and

straight. Right lacinia mobilis with 1 strong tooth and denticles, spine simple and straight. Mandibular palp

second article 1.6 times as long as first, with 35 short setae along distal part of lateral margin, third article

lanceolate, with row of 50 setae, last 3 longer. Maxilla 1 lateral lobe with 1 1 strong apical teeth; medial lobe with

1 short apical seta in weak notch. Maxilla 2 inner lobe with 19 slender setae, median and outer lobes each with

2 setae. Maxilliped : coxa and epipod lateral to it separated by suture; basis not separated by suture from lateral

almost-semicircular lamella; basis without facial setae proximal to palp; endite with oblique distal margin bearing

2 short spiniform setae; palp with short first article, second article without lateral row of setae, with 2 rows of

mesial setae with hiatus between, third article without lateral row of setae but with distal setae.

Pereopod 1 basis to merus without setae; carpus without setae on posterior margin, with 2 spiniform setae on

distal corner; propodus greatest width 50% of length, curved palm with row of 45 alternating flattened and

spiniform setae, each with apical projection, and with row of short lateral setae submarginally; dactylus evenly

curved and tapering, unguis weakly differentiated, overlapping carpus. Pereopod 2 with slender basis; ischium 75%

length of basis; merus and carpus subequal in length, merus with strong anterodistal setae; propodus with

prominent heel, greatest width 52% of length, palm with 20 strong flagellated setae in U-shaped marginal row plus

1 simple seta in middle of flat palm; dactylus strongly curved on to heel of palm, unguis small. Pereopods 3-6

similar. Basis, ischium and merus progressively shorter; carpus longer than merus, setose on lower margin;

propodus shorter than carpus, setose on lower margin and with long setae on distal margin; dactylus very slender

and tapering. Pereopod 7 about 0.6 length of pereopod 6; merus-propodus with mat of fine setae on lower margin;

carpus and propodus with strong setae on distal margins; propodus width 0.3 length; dactylus about 0.4 length of

propodus, closing back on palm, more curved than in pereopods 3-6.

Pleopod 1 peduncle subtriangular, with strong medial lobe (not figured) bearing 3 setulate setae; endopod

almost semicircular, with 25 marginal plumose setae; exopod larger, with 46 marginal plumose setae. Pleopod 2

peduncle more elongate than in pleopod 1, with narrow medial lobe bearing 2 setulate setae; endopod narrower than

in pleopod 1, with 23 marginal plumose setae, with apical slender appendix masculina, 4 times as long as

endopod; exopod larger, broader, with 53 marginal plumose setae. Pleopod 3 peduncle narrower than in pleopod 2,

with medial lobe bearing 2 setulate setae; endopod rounded, triangular, with 29 marginal plumose setae; exopod

almost semicircular, with 52 marginal plumose setae. Pleopod 4 exopod operculiform, chitinized, 2-articulate,

with lateral row of about 130 short marginal plumose setae; endopod smaller, without setae. Pleopod 5 rami with

transverse suture; exopod with 10 distal plumose setae; endopod as long as exopod, without setae. Uropod peduncle

triangular; exopod 0.85 length of endopod, distally rounded, with 10 widely spaced marginal setae; endopod

3.2 times as long as wide, distally rounded, with 16 distomedial setae.

Etymology. — For Cidaris, the name of the series of cruises organised by James Cook University of North

Queensland in the Coral Sea.

Distribution. — Coral Sea, 891-1491 m depth.

Remarks. — A. cidaris differs from A. bromleyana mainly in the dorsal sculpture of pereonite 1, shape and

setation of male pereopods 1 and 7, and size. The low submarginal ridge on pereonite 1 is characteristic of the

species. It does not end anteriorly in the marginal spine seen in A. bromleyana. The male syntype of

A. bromleyana is 53 mm long whereas adult males of A. cidaris range in size from 19.8 to only 28.1 mm.

Ovigerous females are 15.4 to 26.9 mm and have similar body proportions to males. The species also differs from

the other species of Acutiserolis [listed by BRANDT (1991) and WAGELE (1994) but only partly illustrated] :

Acutiserolis macdonnellae (Menzies, 1962) and A. maryannae (Menzies, 1962) have a small anterior triangular

160

G.C.B. POORE & A. BRANDT

projection on pereonite 1 (like A. bromleyana)\ A. neaera (Beddard, 1884), A. gracilis (Beddard, 1884) and

A. spinosa (Kussakin, 1967) have strong sculpture on the pleotelson and slightly different relative lengths of the

coxal plates and epimera. Serolis margaretae Menzies, 1962, included by both BRANDT (1991) and WAGELE (1994)

in Acutiserolis , lacks the extreme elongation of coxal plates and epimera so may belong in another genus but, at

8.4 mm long, may not be an adult specimen.

Fig. 6. — Acutiserolis cidaris sp. nov. Male holotype, MTQ W13598 (only representative setae shown on pleopods) :

a, ventral view of head and pereonites 1-2.

Source :

NEW SPECIES OF SEROLIDAE FROM THE CORAL SEA

161

Genus CAECOSEROLIS Wagele, 1994

Caecoserolis Wiigele, 1994 : 10-11, 50, 55-57.

Diagnosis. — Body ovoid, smooth, slate-grey, margin continuous from head to coxa 6; pleotelson with dorsal

lateral submarginal ridges. Eyes minute, round, lateral. Pereonites 5-7 and pleonite 1 fused. Pereonal sternite I

visible, sternites 2-4 invisible (ventral coxal plates meeting in midline), sternites 5-7 fused. Pleonal sternites 1-3

with vertical posterior ridge, not sexually dimorphic.

Coxal plates without interacting keys, closely fitting; coxal sutures 2-4 visible dorsally; coxa 6 as long as or

exceeding pleonal epimera 2 and 3 posteriorly. Pleonites 2 and 3 with epimera laterally flattened, narrow in dorsal

view.

Mandible with incisor process untoothed; left lacinia mobilis three-quarters width of incisor process, right

toothed and narrower. Maxilla 2 with middle and outer lobes shorter than inner, each with 2 distal setae. Maxilliped

with free coxa, epipod and lamella on basis; palp article 2 cordiform.

Pereopod 1 with alternating spiniform and plate-like setae plus submarginal lateral row along palm. Pcreopod 2

of male with propodus palm having margin of spiniform setae and lateral row of longer setae; unguis small and

terminal; pereopod without felt of fine hairs. Pcreopod 7 of male with propodus little broader than in female,

setose but without felt of fine hairs; dactylus simple.

Pleopod 2 of male with endopod tapering. Plcopod 4 endopod not bilobed. Uropod biramous, peduncle short;

inserting half-way along pleotelson.

Oostegites present on pereopods 1-4 of female.

Remarks. — The new species Caecoserolis novaecaledoniae is difficult to place in any genus as recently

defined. Brandt (1991) defined a genus-group, expanded by WAGELE (1994), of three genera, Atlantoserolis

Wiigele, 1994, Glabroserolis Menzies, 1962 and Caecoserolis Wiigele, 1994, which share synapomorphies of being

blind and being widest at pereonite 2. They were also said to have a “stalked” endopod on the male pleopod 2

which bears the appendix masculina. Wagele (1994) believed this to be a synapomorphy of his genus-group B

(which includes these genera). The new species has a similar shape and smooth dorsal surface to members of these

three genera but has small eyes. It does not have the stalked endopod but nor do Caecoserolis brinki (Kensley,

1978) or C. apheles (Schotte, 1992), species presently assigned to Caecoserolis. This genus was reported by

WAGELE (1994) to differ from Atlantoserolis and Glabroserolis in not having a shortened uropodal exopod but it is

very short in C. apheles.

The genera have been weakly differentiated in the past, not all species agreeing well with the diagnoses of the

genera to which they have been assigned. For many characters such as those used by Harrison and Poore (1984)

and POORE (1987) (coxal keys, mandible, maxilla 2, maxilliped, male pereopod 2 palm setation and unguis, felt of

setae on pereopods 2 and 7, pleonal sternites) the condition is known for few serolid genera. We present an

expanded diagnosis and. for the time being, we believe the new species is best assigned to Caecoserolis.

An alternative might be to place the new species in one of WAGELE’s genus-group C whose synapomorphy is

a 3-lobed ridge between the eyes. Similar structures are seen outside this genus-group, for example in Serolina

Poore, 1987, which may cast doubt on its monophyly. All but one genus of this group posses paired spines and

raised triangular fields on the pleotelson viewed as synapomorphies by Wagele (1994). If this sculpture is

homologous with the dorsolateral ridges seen in the new species and it is a member of genus-group C, the species

may be a member of Serolella Pfeffer, 1891 but its members have large reniform eyes and diverging coxal plates.

Caecoserolis novaecaledoniae sp. nov.

Figs 7-10

TYPE MATERIAL. — New Caledonia. Biocal : stn CP 62, 24°19'S, 167°49'E, 1395-1410 m, 2 September 1985 :

holotype 6 16.9 mm (MNHN-Is 4081). Paratypes : 1 ovigerous $ 18.5 mm, 1 juvenile $ 13.7 mm, damaged post-manca.

162

G.C.B. POORE & A. BRANDT

10.0 mm (MNHN-Is 4082). — Stn CP 58, 23°56’S, I66°41'E, 2660-2750 m, 1 September 1985 : manca, 7.8 mm (MNHN-

Is 4080). — Stn CP 69, 23°52'S, 167°58’E, 1220-1225 m, 3 September 1985 : 1 6 16.7 mm, 1 ovigerous 9 18.5 mm

(NMV J27644).

NON-TYPE MATERIAL. — Chesterfield Islands. Corail 2 : stn DE 13, 21C02.77’S, 160°55.00’E. 700-705 m,

21 July 1988 : 1 juvenile 9 12.2 mm (MNHN-ls 4083);

Musorstom 5 : stn 386, 20°56.21’S, 160°51.12'E, 770-755 m, 22 October 1986 : male, 13.0 mm (MNHN-Is 3067).

Norfolk Island : 29°46.6'S, 167°58.9'E to 29°42.0’S. 168°02.0'E. 500 m, Sigsbee trawl, J.E. Watson on R. V.

"Dimitry Mendeleev ", 1 January 1976 ; 1 6 11.4 mm (NMV J6796). — 30°31.0*S. 161°54.2*E to 30°19.4’S, 161°40.6'E,

1210 m,' 29 December 1976 : 1 6 1 1.7 mm (NMV J7763).

DESCRIPTION OF male holotype. — Greatest width of body (between tips of coxae 2) 78% of body length

(rostrum to end of pleotelson). Head with broad diverging anterolateral lobes lateral to bases of antennae, bluntly

produced anterolaterally; continuous obscure transverse ridge at base of antennae; eye minute, circular, submarginal

near widest part of head.

Fig. 7. — Caecoserolis novaecaledoniae. Male holotype, MNHN-Is 4081 ; habitus, scale bar = 5 mm.

Pereonite 1 with anterolateral margin continuously convex; posterolateral corner tightly overlapping coxa 2;

smooth except for slight mediolateral roughening. Pereonites 2-4 articulating, with coxal plates marked off by

Source :

NEW SPECIES OF SEROLIDAE FROM THE CORAL SEA

163

dorsal sutures; pereonites 5-7 fused mid-dorsally. Posterolateral angles of the coxal plates 2-6 barely protruding

posterolaterally, longer than and reaching further posteriorly than those of preceding segments; coxal plate 4

reaching to base of pleotelson, 5 reaching 30% of length of pleotelson, 6 reaching 80% of length of pleotelson.

Pereonite 7 without coxal plates. Ventral coxal plates 2-4 meeting, swollen and sculptured in mid-line;

sternites 5-7 fused and visible, with minute fused paired penes; ventral coxal plates 6-7 incompletely separated.

MX2

Fig. 8. — Caecoserolis novaecciledoniae. Male holotype, MNHN-Is 4081 : a. habitus in lateral view, b, ventral view of

pereonites 2-7, pleonites 1-3.

Pleonite 1 not visible dorsally. Pleonites 2 and 3 with narrow epimera, both 50% of length of pleotelson.

Pleonites 1-3 similar, with sternal plate broad, prominent, sculptured, angled sharply posterolaterally and at

midpoint.

164

G.C.B. POORE & A. BRANDT

Fig. 9. — Caecoserolis novaecaledonicie. Male holotype, MNHN-ls 4081 (p5 dactylus missing).

Source : MNHN, Paris

NEW SPECIES OF SEROLIDAE FROM THE CORAL SEA

165

Pleotelson 42% ot total length, 92% as wide as long, lateral margins curved, posterior margin rounded and

upturned; with obscure mid-dorsal ridge over proximal half, sharp dorsolateral ridge sharp and evenly curved over

three-quarters of length, widest at about midpoint of pleotelson; sides almost vertical over proximal half and

bulging over shallow concavity.

Antenna 1 with peduncular article 2 slightly narrower and shorter than article 1. article 3 little longer than

first two together; flagellum of about 60 articles; flagellar article 1 longest, one-sixth length of last peduncle

article, articles 1-4 without aesthctascs, remainder with 1 long aesthetasc and 2-3 simple setae. Antenna 2 with

short first peduncular article, second longer than third, with few proximal short setules, fourth and fifth with tufts

of setae, fifth 1.2 times as long as fourth; flagellum of more than 25 articles, each with group of distolateral

simple setae.

Mandibles asymmetrical, right lacinia mobilis narrower than on left. Left lacinia mobilis a broad blade

0.8 width of incisor, spine (spine row rudiment) simple and straight. Right lacinia mobilis irregularly toothed,

spine simple and straight. Mandibular palp second article 1.3 times as long as first, with 30 short setae along

distal part of lateral margin, third article lanceolate, with row of 37 setae, last 4 longer. Maxilla 1 lateral lobe with

1 1 strong apical teeth; medial lobe stalked, with 1 short apical seta. Maxilla 2 inner lobe with 16 slender setae,

median and outer lobes each with 2 setae. Maxilliped: coxa and epipod lateral to it separated by suture; basis

separated by suture from lateral rounded lamella; basis with facial setae near mesial face; endite with transverse

distal margin bearing 2 spiniform setae; palp with short first article, second article with lateral row of setae, with

2 rows of medial setae with hiatus between, third article with lateral row of setae and with distal setae.

Pereopod 1 basis to mcrus without setae; carpus with about 10 peg-like short setae on transverse palm and

2 spiniform setae distolaterally; propodus widest at 50% of length, curved palm with row of 35 alternating fan¬

shaped setae and spiniform setae, each with apical projection, and submarginal lateral row of short simple setae;

dactylus evenly curved and tapering, unguis differentiated. Pereopod 2 with basis longer than next 2 articles;

ischium with 1 anterodistal seta; mcrus and carpus subequal in length, merus with strong anterodistal setae; carpus

with setae along posterior margin; propodus with obtuse heel, greatest width 40% of length, palm with lateral row

of 3 long tapered setae, scattered smaller setae distally, lateral and mesial rows each of 6 peg-like spiniform setae;

dactylus curved, not reaching heel of palm, unguis not differentiated. Pereopods 3-6 similar, except distal articles

longer in posterior limbs. Basis, ischium and merus progressively shorter; carpus longer than merus, setose on

lower margin; propodus shorter than carpus, setose on lower margin and with long setae on distal margin; dactylus

very slender, 0.4 length of propodus, tapering. Pereopod 7 about three-quarters length of pereopod 6; setation

similar; dactylus one-third length of propodus, less slender than in pereopods 3-6.

Pleopod I peduncle subtriangular, with strong medial lobe bearing 3 setulate setae; endopod almost oval, with

24 marginal plumose setae; exopod larger, with 46 marginal plumose setae. Pleopod 2 peduncle more elongate

than in pleopod 1, with narrow medial lobe bearing 2 setulate setae; endopod narrower than in pleopod 1, with

18 marginal plumose setae, with apical slender appendix masculina, 3.6 times as long as endopod; exopod larger,

broader, with 48 marginal plumose setae. Pleopod 3 peduncle narrower than in pleopod 2, with medial lobe bearing

2 setulate setae; endopod rounded, with 28 marginal plumose setae; exopod almost semicircular, with 48 marginal

plumose setae. Pleopod 4 exopod operculiform, chitinized, 2-articulate, with lateral row of about 130 short

marginal plumose setae; endopod smaller, without setae. Pleopod 5 rami weakly 2-articulate; exopod with 3 distal

plumose setae; endopod as long as exopod, without setae.

Uropod attached just beyond midpoint of pleotelson, reaching three-quarters of its length; with subtriangular

peduncle; exopod 0.75 length of endopod, distally rounded, with 7 widely spaced marginal setae; exopod 2.9 times

as long as wide, distally rounded, with 8 distal setae.

Female. — Differs from male : oostegites present on pereopods 1-4; pereopod 2 similar to pereopods 3-6;

pereopod 7 dactylus very slender as in pereopods 3-6.

Etymology. — For New Caledonia.

Distribution. — Coral Sea, northern Tasman Sea; 500-2750 m depth.

166

G.C.B. POORE & A. BRANDT

FlG. 10. — Caecoserolis novaecaledoniae. Male holotype, MNHN-Is 4081 (only represeniative setae shown on

pleopods).

Source : MNHN. Paris

NEW SPECIES OF SEROLIDAE FROM THE CORAL SEA

167

REMARKS. — The shape and sculpture of the pleotelson of the two males far south from the type locality, in

the vicinity of Norfolk Island (30°S), differ slightly from that of the type material. The pleotelson is more angled

laterally at the ends of the pleonal epimera than in the holotype. The dorsolateral ridge is more sharply angled,

proximal to the ends of the pleonal epimera, whereas in the holotype this ridge is curved and widest further

posteriorly. These differences point to possible widespread polymorphism in Serolidae (Wagele, 1986).

Variability of this species over a wider geographic range remains to be investigated.

ACKNOWLEDGEMENTS

We are very grateful to Alain CROSNIER, Paris, Bertrand Richer de FORGES, Noumea, and Peter Arnold.

Townsville, for making this material available. We thank Ms M. Lowe for the loan of material from the Natural

History Museum, London, and J.-W. WAGELE, Bielefeld, for comments on an early draft, and Niel BRUCE for

comments on a later one. We thank Kate THOMPSON, Melbourne who prepared the habitus Figures and inked all the

others.

REFERENCES

Beddard, F.E., 1884a. — Preliminary notice of the Isopoda collected during the voyage of H.M.S. 'Challenger' - Part 1.

Serolis. Proceedings of the Zoological Society of London , 23 . 330-341.

Beddard, F.E., 1884b. — Report on the Isopoda collected by H.M.S. Challenger during the years 1873-76. Part I. - The

genus Serolis. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873-76.

Zoology , 11 : 1-85, pis I-X.

Brandt, A., 1988. — Antarctic Serolidae and Cirolanidae (Crustacea: Isopoda): New genera, new species, and

redescription. Koeltz Scientific Books, Konigstein. 143 pp. Theses Zoologicae Volume 10.

Brandt, A., 1991. — Zur Besiedlungsgeschichte des antarktischen Schelfes am Beispiel der Isopoda (Crustacea,

Malacostraca). Berichte zur Polarforschung, 98 : 1-240.

Harrison, K. & Poore, G.C.B., 1984. — Serolis (Crustacea, Isopoda, Serolidae) from Australia with a new species from

Victoria. Memoirs of the Museum of Victoria , 45 : 13-31.

HURLEY, D.E., 1957. — Some Amphipoda, Isopoda and Tanaidacea from Cook Strait. Zoology Publications from Victoria

University College, 21 : 1-20.

HURLEY, D.E., 1961. — The distribution of the isopod crustacean Serolis bromleyana Suhm with discussion of an

associated deep water community. Bulletin, New Zealand Department of Scientific and Industrial Research , 139 : 225-

233.

Nordenstam, A., 1933. — Marine Isopoda of the families Serolidae, Idotheidae. Pseudidotheidae, Arcturidae, Parasellidae

and Stenetriidae mainly from the South Atlantic. Further Zoological Results of the Swedish Antarctic Expedition,

1901-1903 ,3 : 1-284, 2 pis, errata.

Menzies, R.J., 1962. — The isopods of abyssal depths in the Atlantic Ocean. Verna Research Series, 1 : 79-206.

POORE, G.C.B., 1987. — Serolina, a new genus for Serolis minuta Beddard (Crustacea: Isopoda: Serolidae) with

descriptions of eight new species from Australia. Memoirs of the National Museum of Victoria, 48 : 141-189.

Sheppard, E.M., 1933. — Isopoda Crustacea Part I. The family Serolidae. Discovery Reports, 7 : 253-362.

WAGELE, J.-W., 1986. — Polymorphism and distribution of Ceratoserolis trilobitoides (Eights, 1833) (Crustacea,

Isopoda) in the Weddell Sea and synonymy with C. cornuta (Studer, 1879). Polar Biology, 6 : 127-137.

WAGELE, J.-W., 1994. — Notes on Antarctic and South American Serolidae (Crustacea. Isopoda) with remarks on the

phylogenetic biogeography and a description of new genera. Zoologische Jahrbiicher, Abteilung fur Systematik,

121 : 3-69.

168

G.C.B. POORE & A. BRANDT

WillemOes-Suhm, R. von. 1874. — Von der Challenger-Expedition. Briefe an C. Th. E. von Siebold. II. Zeitschrift fur

Wissenschaftliche Zoologie, 24 : IX-XXIII.

WillemOes-Suhm. R. von. 1876. — Preliminary report to Professor Wyville Thomson, F.R.S., Director of the Civilian

Scientific Staff, on Crustacea observed during the cruise of H.M.S. "Challenger" in the Southern Sea. Proceedings of

the Royal Society of London, 24 : 585-592.

Source :

■SULTATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULT ATS DES CA

Crustacea Decapoda : Pseudopandalus curvirostris ,

genre et espece nouveaux (Pandalidae)

de Nouvelle-Caledonie

Alain CROSNIER

Chercheur ORSTOM

Museum national d’Histoire naturelle

Laboratoire de Biologic des Invertebres marins et Malacologie

55 rue Buffon, 75231 PARIS Cedex 05

RESUME

Un genre et une espece nouveaux appartenant a la famille des Pandalidae, Pseudopandalus curvirostris, sont decrits et

figures. Proche du genre Pandalus , cette espece a etc recoltee en Nouvelle-Caledonie, ti plusieurs reprises, entre 420 et

570 m de profondeur.

ABSTRACT

Crustacea Decapoda : Pseudopandalus curvirostris , new genus and species (Pandalidae) off New

Caledonia.

A new genus and species, Pseudopandalus curvirostris, belonging in the family Pandalidae are described and figured.

Close to the genus Pandalus, this species has been collected, several times, off New Caledonia, at depths between 420 and

570 m.

INTRODUCTION

Lors de diverses campagnes autour de la Nouvelle-Caledonie, il a ete recolte, a dix reprises, des specimens d'une

crevette d'assez grande taille, qui s'est revelee appartenir a un genre et a une espece nouveaux que nous decrivons

dans les pages qui suivent.

Le materiel est depose au Museum national d'Histoire naturelle (MNHN), a Paris, h l'exception d'un male et

d’une femelle paratypes, envoyes au National Museum of Natural History (USNM), a Washington.

Crosnier, A., 1997. — Crustacea Decapoda : Pseudopandalus curvirostris . genre et espece nouveaux (Pandalidae) de

Nouvelle-Caledonie. In : A. Crosnier (ed.), Resultat des Campagnes MUSORSTOM, Volume 18. Mem. Mus. natn. Hist .

nat., 176 : 169-176. Paris ISBN 2-85653-511-9.

170

A. CROSNIER

La longueur de la carapace (LC) est mesuree du fond de I'orbite a la partie dorsale du bord posterieur de la

carapace; c'est elle qui est mentionnee dans la liste du materiel examine. La longueur totale correspond a la distance

separant la pointe du rostre de l'extremite du telson.

Genre PSEUDOPANDALUS nov.

DESCRIPTION . — Rostre fort et long, depassanl tres largement le scaphoccrite, non articule avec la carapace,

avec des dents toutes mobiles sur son bord dorsal (a l'exception de la dent subdistale) et des dents toutes fixes sur

son bord ventral. Dents postrostrales toutes mobiles. Carapace sans carencs longitudinales, avec une epine

antennaire, une epine pterygostomienne et une dent supra-orbitaire. Abdomen avec les pleurons des trois premiers

segments arrondis et ceux des quatrieme et cinquieme se terminant en pointe aigue.

Antennules dune longueur inferieure a celle du corps. Yeux tres developpes, pigmentes, avec une cornee

beaucoup plus large que le pedoncule oculaire et un ocelle dorsal tres bien marque. Stylocerite tres allonge et acere.

Lobe posterieur du scaphognathite court et arrondi, sans tres longues soies. Deuxiemes maxillipedes avec un article

terminal beaucoup plus large que long. Troisiemes maxillipedes avec un exopodite et une legere expansion de

l'antepenultieme article vers son milieu. Epipodites et arthrobranchies presents sur les quatre premieres paires de

pereiopodes. Premiers pereiopodes avec une pince minuscule et une legere expansion ventrale de l'ischion dans sa

moitie dislale. Seconds pereiopodes de tailles inegales (le droit ctant nettemenl plus court et plus fort que la

gauche) avec un carpe multiarticule et une pince bien developpee. Troisiemes, quatriemes et cinquiemes

pereiopodes forts. Ischion des troisiemes et quatriemes pereiopodes portant un forte epine laterale externe mobile,

en plus de l'epine de la face inferieure.

Espece Type. — Pseudopandalus curvirostris sp. nov., par monotypie.

Etymologie. — Du grec pseudos , mensonge, accole au nom de genre Pandalus afin de rappeler combien le

nouveau genre se situe pres du precedent.

Position. — Parmi les genres de la famille des Pandalidae (voir Christoffersen, 1989; Holthuis, 1993, en

particular), le genre que nous decrivons est proche du groupe d'especes Australopandcilus , Pandalina ,

Dichelopandal u s , Pandalus et Pandalopsis. Parmi ces especes, c'est de Pandalus Leach, 1814, qu'il semble le plus

proche. Comme lui, il presente un rostre tres long, ayant toutes ses dents dorsales mobiles (a l'exception de la

subdistale), une expansion de l'antepenultieme article des troisiemes maxillipedes et de l’ischion des premiers

pereiopodes faiblement developpee, des antennules dont la longueur n'excede pas celle du corps, une carapace avec

une epine antennaire et une epine pterygostomienne, des mandibules avec un processus incisif et un palpe de trois

articles, des pereiopodes assez forts, dont les deuxiemes sont de taille inegale avec un carpe pluriarticule, la

presence d'arthrobranchies et d'epipodites sur les quatre premieres paires de pereiopodes.

II en differe par :

— le lobe posterieur du scaphognathite court et arrondi (au lieu d'etre allonge et en pointe), sans tres longues

soies;

— la presence d'un exopodite (de petite taille) sur les troisiemes maxillipedes;

— le stylocerite tres long et acere (au lieu d'etre court et tronque).

Pseudopandalus partage, par ailleurs, de nombreux caracteres avec le genre Plesionika : carpe des deuxiemes

pereiopodes avec de nombreux articles, pas de carene longitudinale sur la carapace, roste long et fixe, yeux bien

developpes, avec une cornee plus large que le pedoncule oculaire, troisiemes maxilipedes avec un exopodite,

epipodes sur les quatre premieres paires de pereiopodes; stylocerite termine en pointe, lobe posterieur du

scaphognathite arrondi. Dans ce genre, il s'apparente au sous-genre Nothocaris (voir Burukovsky, 1981; Chace,

1985) par ses deuxiemes perdiopodes inegaux.

Mais il en differe par :

— le rostre qui ne porte sur son bord superieur que des dents mobiles (a l'exception de la subdistale);

Source :

PSEUDOPANDALUS GEN. NOV.

171

— le stylocerite tres long (beaucoup plus que chez Plesionika );

— le troisieme segment abdominal avec une dent dorsale recourbee, caracteristique;

— le faible developpement de l'exopodite des troisiemes maxillipedes.

Fig. 1. — Pseudopcindalus curvirostris gen. nov. sp. nov., a-d. 8 21,3 mm, holotype, Nouvelle-Caledonie, Smib 8,

st. DW 195 : a, partie anterieure du corps, vue laterale : b, abdomen, vue lat^rale; c, rostre; d, base du rostre, ail,

pedoncule antennulaire et scaphocerite, vue de dessus. — e-g. 9 23.0 mm, paratype, Nouvelle-Calddonie, Smib 8,

st. DW 193 : e, scaphocerite droit, vue de dessus; f. telson et uropodes gauches, vue de dessus; g, extremity du

telson, vue de dessus; h. stylocerite gauche vu de dessus.

Les figures a et b sont au meme grossissement.

172

A. CROSNIER

On remarquera, enfin, que la presence d'un lobe orbitaire semble etre un caractere bien particulier a

Pseudopandalus parmi les Pandalidae.

Pseudopandalus carvirostris sp. nov.

Fig. 1-3

Materiel EXAMINE. — Nouvelle-Caledonie. Biocal : st. DW 46, 22°53,05’S. 167°1708’E, 570-610 m,

30.08.1985 : 1 9 17,6 mm (MNHN-Na 13247). — St. CP 47, 22°53,04’S. 167°16,77’E, 550 m, 30.08.1985 : 1 9

23,9 mm (MNHN-Na 13248).

MUSORSTOM 4 : st. CP 216, 22°59,5'S, 167°22,0’E, 490-515 m, 29.09.1985 : 12 ex. 11,5 & 24,7 mm (MNHN-Na

13249).

Smib 2 : st. DW 13, 22°52,4'S, 167°13,4'E, 427-454 m. 18.09.1986 : 1 9 13,6 mm (MNHN-Na 13250).

Chalcal 2 : st. CC 2, 24°55,48’S, 168°21,29'S, 500-610 m, 28.10.1986 : 1 6 13,7 mm; 1 9 7,4 mm.

photographies (MNHN-Na 13251). — St. DW 76, 23o40,5'S. 167°45,2'E,470 m, 30. 1 0; 1 986 : I 9 8,6 mm,

photographic (MNHN-Na 13252).

Fig. 2. — Pseudopandalus curvirostris gen. nov. sp. nov., 9 13,0 mm. paratype, NouveIle-Cal6donie, Smib 8,

st. DW 193. Pieces buccales gauches, face exlerne : a, mandibule; b. premiere maxille; c, deuxieme maxille;

d. premier maxillip&de; e, deuxieme maxillipfede.

Toutes les figures sont au meme grossissement.

Source :

PSEUDOPANDALUS GEN. NOV.

173

Smib 3 : st. CP 4, 24°54,0'S, 168°22,5’E. 530 m. 20.05.1987 : 1 6 20,1 mm (MNHN-Na 13253), 1 9 20,4 mm.

photographiee (MNHN-Na 13254). — St. DW 24. 22°58,7'S, 167°21,rE. 535 m. 24.05.1987 : 1 6 19,5 mm; 3 9 8,7 &

14.6 mm (MNHN-Na 13255).

Smib 4 : st. DW 62, 23°00.4’S, 167°21,8’E. 540 m, 10.03.1989 : 2 d 17,0 & 21,4 mm; 1 9 14.5 mm (MNHN-Na

13257); 1 6 20,3 mm, photographic (MNHN-Na 13258).

AZTkQUE : st. 7, 23°37,5'S, 167042,1'E, 425-500 m, 14.02.1990 : 1 d 21,3 mm (MNHN-Na 13259).

Beryx 11 : st. CP 07, 24°55'S, 168°21'E, 510-550 m, 15.10.1992 : 1 9 14,8 mm (MNHN-Na 13260). — St. CP 08.

24°54’S, 168°21'E. 540-570 m, 15.10.1992 : 1 juv. 8,3 mm; 2 6 14,6 ct 17,9 mm (MNHN-Na 13261). — St. CP 32,

23°38'S, 1 67°43'E, 420-460 m, 18.10.1992 : 1 6 16,3 mm, 1 9 ov. 20.7 mm (USNM). — St. CP 53, 23°48'S,

168°17'E, 540-950 m. 21.10.1992 : 2 9 12,6 et 22,3 mm; 1 9 ov. 25.0 mm (MNHN-Na 13262).

Smib 8 : st. DW 193, 22°58,7’S, 168°20,rE. 500-508 m, 1.02.1993 : 1 6 17.1 mm; 3 9 13.6 a 14,8 mm; 1 9 ov.

22.6 mm (MNHN-Na 13263); 1* 9 23,0 mm (MNHN-Na 13264). — St. DW 195, 22°58,9'S, 167°20,2'E, 508-514 m,

1.02.1993 : 1 6 21,3 mm (MNHN-Na 13265); 4 d 13.3 a 21.9 mm; 1 9 14,5 mm (MNHN-Na 13266).

Smib 10 : st. DW 202, 24°55’S, 168°22’E, 525-513 m, 10.01.1995 : 1 d 10,5 mm; 1 9 15.9 mm (MNHN-Na 13267).

TYPES. — Le male dont la carapace mesure 21,3 mm, capture a la station DW 195 de Smib 8 (MNHN-B

13265), est I’holotype. Tous les autres specimens mentionncs dans le "Materiel examine" sont des paratypes.

Description. — Le rostre est fort, tres recourbe dorsalcment et long (plus de 1,5 fois la longueur de la

carapace chez l'adulte). Son bord dorsal est inerme sur la plus grande partie de sa longueur et porte, dans sa partie

basale, 2 dents suivies de 4 postrostrales, soit 6 au total, toutes mobiles; on observe, en outre, une dent fixe

subdistale; les dents mobiles sont de taille sensiblement egale, a l'exception de la proximale qui est legerement

plus petite que les autres et tres proche de la suivanle; les autres sont legerement et egalement espacees les unes des

autres. Son bord ventral porte 1 1 dents fixes (sans compter l'extremite du rostre), reparties sur toute sa longueur;

les trois dents basales sont tres longues, tres recourbecs dorsalcment, et tres proches les unes des autres; les

suivantes diminuent rapidcmcnt de taille, leur espacement croit, puis diminue vers l'extremite du rostre.

La carapace est lisse a l'ceil nu et se revele couverte de tres fines corrugations transversales sous la loupe

binoculaire; elle est depourvue de toute Crete postroslrale mais presente une forte dent antennaire, une dent

pterygostomienne moins developpee que la dent precedente. L'orbile presente, dans sa partie superieure, juste au

dessus du pedoncule oculaire, un petit lobe.

Les yeux sont gros avcc une cornee tres coloree et un ocelle bien marque.

Les antennules ont un pedoncule qui couvre la moitie du scaphocerite; leur plus long flagelle est environ egal a

la moitie de la longueur du corps (rostre non compris).

Les scaphocerites sont relativement etroits (rapport L/l voisin de 4,1); leur epine distale externe n'atteint pas

tout a fait le niveau de l'extremite de l'ecaille.

Les stylocerites, en forme de longue pointe, sont tres developpes; ils s'etendent sur les 85/100 environ du

scaphocerite et sont etroits (rapport L/l voisin de 8) et aceres.

Les pieces buccales sont representees sur les figures 2 a-f et 3 a. On notera le palpe a trois articles des

mandibules qui sont pourvues d'une partie molaire et d'une partie incisive; sur les maxilies, le scaphognathite court

et arrondi et, sur les troisiemes maxillipedes, la forme de 1'ischion qui presente une moitie distale legerement

expansee. Les troisiemes maxillipedes atteignent l'extremite des scaphocerites.

La repartition des branchies, des epipodites et des exopodites est donnee dans le tableau ci-dessous :

174

A. CROSNIER

L'exopodite des troisiemes maxillipedes est bien visible mais de petite taille.

Les premiers pereiopodes attcignent l'extremite du stylocerite. L'ischion porte, sur la moitie distale de sa face

ventrale, une dizaine de spinules mobiles.

Vnw P.s*ud°Pan*alus curv.rostris gen. nov. sp. nov.. a. 2 13.0 mm, paratype, Nouvelle-Caledonie, Smib

193 • Mxp3 gauche. — b-k, 2 23.0 mm. paratype, ibidem : b, PI droit, avec au dessus le merus grossi; c-

Tfrn daC,yl? £ P' S0US divcrs aspec,s; f‘ P2 droit; K- p2 gauche; h. P3 droit; i. partie distale c

propode et dactyle de P3 droit; j, P4 droit; k. P5 droit. V

Les figures b, f-g, h, j-k sont au meme grossissement.

Source :

PSEUDOPANDALUS GEN. NOV

175

Le deuxieme pereiopode gauche depasse le scaphocerite d'une longueur egale a la moitie de celle de ce dernier.

Son carpe est divise en nombreux articles; suivant les specimens nous cn avons compte de 55 a 61; sur le merus,

egalement divise, nous avons compte de 25 a 28 article environ, repartis sur toute sa longueur; sur l’ischion, on

distingue egalement 3 ou 4 traces de division dans sa parlie distale. La pince est bien developpee.

Le deuxieme pereiopode droit est nettement plus court que le gauche; il atteint presque I’exlremite du scaphocerite.

Son carpe est divise en de moins nombreux articles quc le deuxieme pereiopode gauche : nous en avons compte de

19 a 22 suivant les specimens; le merus n'est divise que dans sa partic distale; les divisions ne sont pas toujours

nettes et 1’on distingue 5 ou 6 articles; aucune trace de division ne s'observe sur l’ischion. La pince est bien

developpee comnie dans le deuxieme pereiopode gauche.

Les troisiemes, quatriemes et cinquiemes pereiopodes sont forts et d'une longueur moderee (les troisiemes

depassent le scaphocerite par leur dacyle et le dixieme environ de leur propode, les quatriemes atteignent l'extremite

du scaphocerite, les cinquiemes les neuf dixiemes du scaphocerite). De grosses epines mobiles s'obscrvent sur l'is¬

chion, le merus et le carpe; elles sont au nombre, rcspectivement, de 2, 4 ou 5 et 2 sur les troisiemes pereiopodes,

de 2, 6 et 2 sur les quatriemes et de 1, 5 ou 6 et 2 sur les cinquiemes. Les dactyles sont fort, recourbes, et garnis,

sur leur bord posterieur, de 6 epines mobiles dont la taille croit de la base a l'extremite du dactyle.

L'abdomen a les pleurons de ses trois premiers segments arrondis, tandis que ceux des quatrieme et cinquieme

segments presentent un denticule acere a la jonction dcs bords ventral et posterieur. La face dorsale du troisieme

segment porte, vers son centre, unc tres forte excroissance en forme d'epine recourbee vers l'arriere. Le sixieme

segment a ses faces laterales qui se prolongent par un lobe lermine en pointe; ses bords ventraux portent une petite

epine distale; le rapport de sa longueur (mesuree du condyle d’articulation avec le cinquieme a la base du telson) a sa

hauteur est tres voisin de deux. Le telson qui depasse tres legerement, par ses epines terminates, la rame interne des

uropodes, est environ 1,3 fois plus long quc le sixieme segment abdominal; il porte. a son extremite, une paire de

petites epines, encadree par une paire de tres fortes epines et est orne, dorsolateralement, de quatre paires d'epines

dont la derniere est subdistale.

Coloration - L'allure generate est celle d'un animal rouge avec de larges bandes transversales blanches.

La carapace presente une large bande transversale blanche au niveau des deux dernieres dents postrostrales; une

autre bande couvre la partie tout a fait posterieure de la carapace; entre ces deux bandes, se trouve une tache, de

forme irreguliere, souvent incurvee vers 1'arriere, qui part du bord dorsal et s'etend sur le tiers superieur environ de

la carapace; la partie anterieure de la carapace, enfin, entre les epines antennaire et pterygostomienne, est egalement

blanche.

Le rostre est rouge dans sa partie anterieure puis, en arriere, presente une zone blanche qui se recolore plus ou

moins dans sa parlie basale. Les dents rostrales dorsales et les dents postrostrales sont rouges, a l'exception de la

derniere, ou des deux dernieres dents postrostrales qui sont blanches.

L'abdomen se caracterise par une large bande tres blanche, tres marquee, qui part de la forte epine recourbee,

dorsale, du troisieme segment abdominal qu’elle englobe, et qui s’etend, cn diagonale, sur la partie posterieure du

pleuron du second segment abdominal. Les segments 1, 4 et 5 out, d’une maniere generate, leur partie anterieure

decoloree et leur partie posterieure rouge. Le sixieme segment a sa partie dorsale coloree et le reste plus ou moins

blanchatre. Le telson et les uropodes sont rouges avec une bande blanche transversale sub-basale.

Les pereiopodes presentent des alternances de longues zones blanches et rouges. Les pleopodes sont rouges ou

blancs suivant les specimens.

ETYMOLOGIE. — Du Latin curvus , courbe, et rostrum , rostre, afin de rappeler la forme tres incurvee du rostre.

Taille. — Les plus grands specimens observes sont une femelle ovigere dont la longueur totale est de

122 mm (LC = 25,0 mm) et un male male mesurant 123 mm (LC = 24,7 mm). La plus petite femelle ovigere

observee mesure 101 mm (LC = 20,7 mm).

Les ceufs sont tres nombreux et petits (diametre voisin de 0,2 mm).

Distribution. — Nouvelle-Caledonie, entre 420-570 m. Une recolte a ete faite lors d'un chalutage effectue

entre 540 et 950 m de profondeur, mais, compte tenu des autres recoltes. il est vraisemblable que la capture a ete

faite lors du debut du chalutage.

176

A. CROSNIER

Fig. 4. — Pseudopandalus curvirostris gen. nov., sp. nov., 9 paratype 20,4 mm, Smib 3, st. CP 4, 24°54,0'S,

168°22,5'E, 530 m (MNHN-Na 13254). Photo P. Tirard.

REMERCIEMENTS

Patrick Lehodey, Bertrand Richer de Forges et Cecilc Debitus, tous de l'ORSTOM, ont recolte 1c materiel

etudie ici. Maurice Gaillard, ancien dessinateur du Museum national d'Histoire naturelle, maintenant en retraite,

a effectue les dessins qui illustrent cette note. Nous sommes heureux de les remercier tous ici.

Par ailleurs, nous avons une gratitude toute particuliere envers le Dr Tomoyuki KOMAI (Natural History

Museum and Institute, Chiba, Japon), dont les commentaires nous ont ete precieux.

REFERENCES

Burukovsky, R. N., 1981. — Opredelitel krevetok roda Plesionika Bate. 1888 (Decapoda, Natantia, Pandalidae) i svodka

ikh geograficheskogo rasprostraneniya [C\6 pour les crevettes du genre Plesionika Bate, 1888 (Decapoda Natantia

Pandalidae) et distribution g^ographique dcs especes de ce genre]. Byulleten Moskovskogo Obschchestva Ispytatelei

Prirody, Otdel Biologicheskii , 86 (4) : 42-53, 2 fig.

Chace, F. A. Jr., 1985. — The Caridean Shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907-

1910, Part 3: Families Thalassocarididae and Pandalidae. Smithsonian Contributions to Zoology , (411) : i-iv

+ 1-143, fig. 1-62.

Christoffersen, M. L., 1989. — Phylogeny and classification of the Pandaloidea (Crustacea, Caridea). Cladistics , 5 :

259-274.

Holthuis, L. B., 1993. — The recent genera of the Caridean and Stenopodidcan shrimps (Crustacea Decapoda): with an

appendix on the order Amphionidacea. C.H.J.M. Fransen & C. van Achterberg eds, Leiden, Nationaal Natuur-

historisch Museum : 1-328, fig. 1-312.

Komai, T., 1994. — Deep-sea shrimps of the genus Pandalopsis (Decapoda: Caridea: Pandalidae) from the Pacific coast of

Eastern Hokkaido, Japan, with the description of two new species. Journal of Crustacean Biology, 14 (3) : 538-559,

fig. 1-10.

ULTATS DES CAMPAGNES MUSORSTOM. VOLUME 18 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 18 — RESULTATS DES CAN

Crustacea Decapoda: Chelonika macrochela , a new

genus and new species of pandalid shrimp (Caridea)

from New Caledonian waters

Charles H.J.M. FRANSEN

Naiionaal Natuurhistorisch Museum

P.O. Box 9517, 2300 RA Leiden

The Netherlands

ABSTRACT

A new genus and species of Fandalidae, Chelonika macrochela , is described and figured. The species was collected in

New Caledonian waters, at a depth between 144 and 155 m.

RESUME

Crustacea Decapoda : Chelonika macrochela , genre et espece nouveaux appurtenant a la

famille des Pandalidae (Caridea) et provenant des eaux neo-caledoniennes.

Un genre et une espece nouveaux dune crevette appartenant a la famille des Pandalidae, Chelonika macrochela, sont

decrits. L'espfece a €i 6 r6colt£e en Nouvelle-Caltklonie, ^ une profondeur comprise entre 144 et 155 m.

INTRODUCTION

Among the caridean shrimp collected by the MUSORSTOM cruises in New Caledonian waters, one specimen

of a pandalid shrimp with a large chela on the second pereiopod was found. Such large chelae on the second

pereiopods are not known within the family Pandalidae. For this reason, a new genus is erected.

Genus CHELONIKA nov.

Definition. — Rostrum long, slender, immovably connected to the carapace, with immovable dorsal and

ventral teeth. Carapace without longitudinal carinae, with antennal spine and pterygostomian tooth. Abdomen with

Fransen, C. H. J. M., 1997. — Crustacea Decapoda: Chelonika macrochela, a new genus and new species of pandalid

shrimp (Caridea) from New Caledonian waters. In : A. Crosnier (ed.), Resultats des Campagnes Musorstom, Volume 18.

Mem. Mas. natn . Hist, nat., 176 : 177-185. Paris ISBN 22-85653-511-9.

178

C. H. J M. FRANSEN

pleura of first four somites rounded, pleura of fifth somite acutely pointed. Eyes well developed, pigmented, cornea

broader than eyestalk; dorsal ocellus present. Posterior lobe of scaphognathite rounded. Second maxilliped with

terminal segment wider than long. Third maxilliped with exopod. Epipods and arthrobranchs present on first four

pairs of pereiopods. First pair of pereiopods without chelae; dactylus formed by a small, distal spine. Second

pereiopod with multiarticulate carpus; chela large. Ambulatory pereiopods long and slender; dactyli with spines on

flexor margin.

TYPESPECrES. — Chelonika macrochela sp. nov., by monotypy.

ETYMOLOGY. — Combination of the greek w'ord chela, meaning pincer, and the last part of the pandalid genus

Plesionika. Gender : feminine.

Fig. 1. — Chelonika macrochela sp. nov., holotype, 9 cl. 7.3 mm. New Caledonia (MNHN-B 12966).

Systematic position. — The present specimen falls within the diagnosis of the family Pandalidae Haworth,

1825, given by Chace (1985 : 9). At present, 19 genera are recognized (following Chaces (1985) synonymy of

Parapandalus with Plesionika) : Anachlorocurtis Hayashi. 1975; Austropandalus Holthuis, 1952; Bitias Fransen,

1990; Chlorocuriis Kemp, 1925; Chlordtocella Balss, 1914; Chlorotocus A. Milne Edwards, 1882;

Dichelopandalus Caullery, 1896; Dorodotes Bate, 1888; Heterocarpus A. Milne Edwards, 1881; Miropandalu 's

Bruce, 1983; Notopandalus Yaldwyn, 1960; Pandalina Caiman, 1899; Pandalopsis Bate, 1888; Pandalus Leach,

1814; Pantomus A. Milne Edwards. 1883; Peripandalus de Man, 1917; Plesionika Bate, 1888; Procletes Bate.

1888; and Stylopandalus Coutiere, 1905. In Holthuis' account of the recent caridean genera (Holthuis, 1993 :

261-263). the present specimen keys out with the genus Plesionika Bate. 1888. CHRISTOFFERSON (1989)

performed a manual cladistic analysis on 21 genera of the presumably monophyletic taxon Pandaloidea, which

consists of both the families Pandalidae and Thalassocarididae Bate, 1888. Christofferson suspects the genus

Plesionika , which comprises about 84 species, to be paraphyletic or polyphyletic. It could well be that Chelonika

will turn out to be an ingroup of the 'Plesionika' assemblage. More definite conclusions on the systematic

position of Chelonika awaits a phylogenetic analysis of the 'Plesionika' assemblage.

CHELONIKA GEN. NOV

179

Almost certainly an apomorphic character in Chelonika is the presence of the extremely large chela in the

second pereiopod. This character does not occur in any of the 19 genera of the family Pandalidae. The non-chelate

form of first pereiopod does not occur in Plesionika. In Plesionika the dactylus is more developed, emerging

subdistally to form a 'chela' with the distal tip of the propodus. Nor is the short squarish stylocerite known in

Plesionika , most species of which have the stylocerite about as long as the basal segment of the antennular

peduncle and the distal part acute.

Chelonika macrochela sp. nov.

Fig. 1-7

Material EXAMINED. — New Caledonia. Bathus 2 : st. DW 753, 22°21.5’S 166°14,6'E; 144-155 m. 15.05.

1993, B. Richer de Forges coll. : 1 9 cl. 7.3 mm (MNHN-B 12966).

Types. — The unique female is the holotype.

DESCRIPTION. — Rostrum slightly overreaching antennal scale, sabre-like, immovably connected to carapace;

dorsal lamina slightly elevated proximally; 10 dorsal fixed teeth present; proximal 7 teeth evenly spaced.

3 proximal teeth postorbital; distal third of lamina with 3 small teeth. Ventral lamina with 5 small teeth in distal

2/5. Antennal spine robust. Pterygostomian tooth distinct.

Fig. 2. — Chelonika macrochela sp. nov., holotype. 9 cl. 7.3 mm. New Caledonia (MNHN-B 12966) : A. anterior

region, dorsal aspect; B. anterior region, lateral aspect.

180

C. H. J. M. FRANSEN

Abdomen with third somite rounded posteriorly, unarmed, without median carina. Pleura of anterior four

somites rounded, fifth with sharp posteroventral tooth and slightly convex ventral margin. Sixth abdominal somite

about 2.5 times as long as fifth. Telson about 0.8 times as long as sixth abdominal somite; with four pairs of

dorsolateral spines; posterior margin of telson with three pairs of spines, lateral pair as long as dorsolateral spines,

intermediate pair longest, median pair about half length of intermediate pair. Uropods as long as telson. Exopod

with movable distolateral spine.

Tegumental scales not found.

Eye pyriform. Ocellus well developed.

Basal segment of antcnnular peduncle without ventromesial tooth. Stylocerite squarish, almost half as long as

basal segment; distolateral angle acute; lateral margin slightly convex. Penultimate and ultimate segment slightly

longer than wide; penultimate segment with 4-5 dorso-distal spines. Flagella long, slender and uniramous; basal

third of outer flagellum with aesthetascs.

Scaphocerite twice as long as antennal peduncle, 5 times as long as median width; distolateral tooth reaching

distal margin of lamina; lateral margin sinuous. Basal segment with strong distoventral tooth. Penultimate and

ultimate segments each about twice as long as wide.

Fig. 3. - Chelonika macrochela new species, holotypc, 2 cl. 7.3 mm. New Caledonia (MNHN-B 12966) : A caudal fan-

B, antennular peduncle (left), ventral aspect; C. antennal peduncle (left), ventral aspect.

Source :

181

Incisor process of left mandible with

6 teeth, right mandible with 5 teeth;

ventralmost tooth much larger than

others; molar process ends in 4 blunt

teeth and an area with series of short

setae. Mandibular palp three-segmented;

basal segment slender; penultimate

segment short, widening distally;

ultimate segment almost twice as long

as penultimate segment, with many

long, slender setae.

First maxilla with lower and upper

endites well developed; upper endite

broad, with two rows of strong spines;

lower endite distally rounded, with many

simple setae; palp bilobed, with each

lobe ending in one large seta.

Second maxilla with lower endite

somewhat reduced; upper endite bilobed.

of which upper lobe best developed; palp

well developed; scaphognathite with

posterior lobe rounded.

First maxilliped with endites of basis and coxa separated; palp slender, three-segmented; exopod with well

developed caridean lobe; epipod bilobed.

Second maxilliped with more or less triangular ultimate segment, bearing many bristle-like setae and one

robust proximal spine; penultimate segment oblong, with row of median, robust spines; carpal segment as long as

wide; meral segment nearly twice as long as its distal width; basis and ischium fused; exopod well developed; well-

developed epipod with podobranch.

Third maxilliped long and slender, reaching distal end of penultimate segment, just beyond rostrum; ultimate

segment 2/3 length of penultimate segment, with many rows of short setae and 4 distal spines; penultimate

segment with several rows of setae; antepenultimate segment 1.6 times as long as penultimate segment,

proximally strongly curved, distally with strong ventro-lateral spine; exopod well developed; lamellar and strap-like

parts of epipod well developed; 2 arthrobranchs present.

Branchial formula as in table 1. Epipods of second, third and fourth pereiopods reduced. Setobranchs not

developed.

CHELONIKA GEN. NOV

FlG. 4. — Chelonika macrochela sp. nov., holotypc. 9 cl. 7.3 mm. New

Caledonia (MNHN-B 12966) : A, mandible (left), ventral aspect;

B. first maxilla (left), ventral aspect.

Table 1. — Branchial formula of Chelonika macrochela sp. nov.

First pereiopods with tip of merus reaching just beyond rostrum, unarmed, not chelate; merus 3.5 times as

long as ischium; carpus slightly shorter than merus; propodus 0.7 times as long as carpus, with setiferous

cleaning organ on proximal part; dactylus very small, spiniform, surrounded by several groups of long setae.

182

C. H. J. M. FRANSEN

Fig. 5. — Chelonika macrochela sp. nov„ holotype, 2 cl. 7.3 mm. New Caledonia (MNHN-B 12966) : A, second maxilla

(left), ventral aspect; B, first maxilliped (left), ventral aspect; C. first maxilliped (left), dorsal aspect (setae omitted

except for palp); D, second maxilliped (left), ventral aspect!

Source

CHEL0N1KA GEN. NOV

183

Fig. 6. — Chelonika macrochela sp. nov., holotype, 9 cl. 7.3 mm. New Caledonia (MNHN-B 12966) : A. third

maxilliped (left), ventral aspect; B, same, detail of tip; C. first pereiopod (left); D-E. same, detail of tip.

184

C.H.J.M. FRANSEN

Fig

' ,?VT Cxhelonika macroch?la sp. nov., holotype, 9 cl. 7.3 mm. New Caledonia (MNHN-B 12966) : A, third pereiopod

(lelt). dactylus; B. branchiae (left); C. second pereiopod (left); D, same, chela; E. same, basal part of basis-ischium,

median aspect; F, first pleopod (left).

Source :

CHELONIKA GEN. NOV.

185

Left second pereiopod with tip of merus reaching just beyond rostrum, unarmed, with well-developed, large

chela; ischium 3.3 times as long as merus, with proximal ventral lamina with ca. 13 flat, curved setae. Carpus

with 59 articulations, about 3.3 times as long as merus. Chela 0.3 times length of carpus. Palm swollen, slightly

shorter than fingers. Fingers slender, with entire cutting edges and strongly hooked tips. Right second pereiopod

missing, except for basis and coxa which are of the same size as those of the left second pereiopod.

Third, fourth and fifth pereiopods similar. Third pereiopod reaching distal part of merus, just beyond tip of

rostrum, fourth falling just short of tip, and fifth reaching to 0.75 of rostrum length. Dactylus short, with 3 spines

on flexor margin of carpus; distalmost robust, with oblique distal edge. Propodus about 0.62 times length of

carpus, slender, with short spines along ventral margin. Carpus 0.80 times length of merus, with very short spines

along ventral margin; merus with ventrolateral spines; spination of merus and ischium as in table 2.

Table 2. — Spination of ischium and merus of third, fourth and fifth pereiopods of Chelonika macrochela sp. nov.

Endopod of first pleopod small. 1/3 of exopod length, triangular; lateral margin setose, median margin with

protuberance.

ETYMOLOGY. — A combination of the greek words macro, meaning large, and chela, meaning pincer.

ACKNOWLEDGEMENTS

A. CROSNIER and B. Richer deForges (Institut Frangais de Recherche scientifique pour le Developpement en

Cooperation, ORSTOM) kindly loaned me the unique specimen from which this new genus and species is

described. I gratefully acknowledge Dr. L. B. Holthuis for helpful comments, and for critically reading the

manuscript.

REFERENCES

Chace, Jr., F. A., 1985. — The Caridean Shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition. 1907-

1910, Part 3: Families Thalassocarididae and Pandalidae. Smithsonian Contributions to Zoology , (411) : i-iv + 1-

143, figs 1-62.

Christoffersen, M. L., 1989. — Phytogeny and Classification of the Pandaloidea (Crustacea, Caridea). Cladistics , 5 :

259-274.

Holthuis, L. B. 1993. — The recent genera of the Caridean and Stenopodidean shrimps (Crustacea Decapoda): with an

appendix on the order Amphionidacea. C.H.J.M. Fransen & C. van Achterberg eds, Leiden, Nationaal

Natuurhistorisch Museum : 1-328. figs 1-312.

Source : MNHN. Paris

ULTATS DES CAMPAGNES MUSORSTOM. VOLUME 18 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULTATS DES CA1

Crustacea Decapoda: Deep-sea shrimps of the genus

Plesionika Bate, 1888 (Pandalidae)

from French Polynesia, with descriptions

of five new species

Tin-Yam CHAN

Institute of Marine Biology, National Taiwan Ocean University

2 Pei-Ning Road. Kcelung, Taiwan. R.O.C.

&

Alain CROSNIER

Laboratoire de Biologie des Invertebrcs marins et Malacologie

Museum national d'Histoire naturelle

55 rue Button, 75005 Paris

ABSTRACT

Recent extensive deep-sea trappings in French Polynesian waters produced 21 species of shrimps belonging to the

caridean genus Plesionika Bate, 1888. Seven of these have been described in detail before (Chan & Crosnier, 1991). The

present report mainly deals with the remaining 14 species and has many interesting findings. P. ocellus (Bate, 1888) is

actually a senior synonym of P. chacei Hayashi, 1986 and the name P. sindoi (Rathbun, 1906) has to be revived.

P. macropoda Chace, 1939 and P. williamsi Forest, 1964 are distinct species and both occur, sometimes even

sympatrically, in the Atlantic and Indo-Pacific. Abundant material belonging to the poorly known species P. nesisi

(Burukovsky, 1986) is found in French Polynesia, and this species is closely related to P. laevis (A. Milne Edwards,

1883) and P. fenneri Crosnier, 1986. Furthermore, P. alexandri (A. Milne Edwards, 1883) and P. unicarinaius (Borradaile.

1915) also belong to the "P. laevis" group which is intermediate between the typical Plesionika and Heterocarpus. The

exact differences between P. reflexa Chace, 1985 and P. ensis (A. Milne Edwards, 1881) are not clear, and it appears that

the posterior spine on the abdomen somite III can be recurved or straight in the specimens from the Indo-Pacific but is

always straight in the Atlantic material. Five new species were found in French Polynesia: P. erythrocyclus sp. nov.,

which is near to P. rostricrescentis (Bate, 1888) and also distributed in Taiwan; P. payeni sp. nov.. which is somewhat

affiliated to P. ocellus ; P. picta sp. nov., which is distinct in lacking antennal spine and without epipod on the

pereiopod IV; P. poupini sp. nov., which is very similar to P. carsini Crosnier, 1986; and P. protati sp. nov., which is

closely related to P. sindoi. The other species reported from French Polynesia are P. edwardsii (Brandt. 1851), P. martia

(A. Milne Edwards. 1883), P. semilaevis Bate, 1888 and P. spinidorsalis (Rathbun. 1906). Moreover, two probable

postlarvae of P. fenneri Crosnier, 1986 were found and additional information is provided for this species.

Chan. T.-Y. & Crosnier, A., 1997. — Crustacea Decapoda: Deep-sea shrimps of the genus Plesionika Bate, 1888

(Pandalidae) from French Polynesia, with descriptions of five new species. In : A. CROSNIER (ed.), Resultats des

Campagnes MUSORSTOM, Volume 18. Mem. Mus. natn. Hist, nat., 176: 187-234. Paris ISBN 2-85653-51 1-9.

188

T.-Y. CHAN & A. CROSNIER

RESUME

Crustacea Decapoda : Crevettes d'eau profonde du genre Plesionika Bate, 1888 (Pandalidae) de

la Polynesie fran^aise. Description de cinq especes nouvelles.

De nombreuses peches au easier en eau profonde, faites en Polynesie fran?aise durant ces dernieres annees. ont permis

de capturcr 21 especes de crevettes carides appartenant au genre Plesionika Bate. 1888. Sept d'entre elles ont deia

decrites ou 6tudi6es (Chan & Crosnier, 1991). Le present travail considere les quatorze restantes et a permis plusieurs

mises au point. C'est ainsi qu'il a ete trouve que P. ocellus (Bate. 1888) et P. chacei Hayashi, 1986, sont synonymes et

que P. sindoi (Rathbun. 1906) doit etre ressuscite. P. macropoda Chace, 1939. et P. williams. Forest. 1964, sont bien

distinctes et se trouvent loutes deux, parfois en sympatrie, dans I'Atlantique el dans l'lndo-Pacifiquc. De nombreux

specimens de P. nesisi (Burukovsky. 1986) ont ete recoltes en Polynesie fran9aise et cette espece se revble tres proche de

P. laevis (A. Milne Edwards. 1883) et P. fenneri Crosnier. 1986, de meme que P. alexandri (A. Milne edwards, 1883) et

P. umcannatus (Borradaile, 1915), ces especes formant un groupe qui presente des caracteres intermediaires entre la plu-

part des Plesionika et les Heterocarpus. Les differences exactes entre P. reflexa Chace, 1985. et P. ensis (A. Milne

Edwards. 1881) n'ont pu etre eclaircies de maniere satisfaisante; il apparait que lepine postdrieure du troisieme segment

abdominal peut etre recourbee ou droite dans l'lndo-Pacifique. mais est toujours droite dans I'Atlantique. Cinq nouvelles

f Pfces sonl decrites de la Polynesie frangaise : P. erythrocyclus sp. nov. qui est proche de P. rostricrescentis (Bate.

1888) et qui se trouve egalement a Taiwan. P. payeni sp. nov. qui prdsente des affinity avec P. ocellus. P. picta

sp. nov. qui se caracterise par I'absence d une epine antennaire et d'un dpipode sur les quatriemes pereiopodes. P. poupini

sp. nov. qui est tres semblable a P. carsini Crosnier, 1986, et P. protati sp. nov. qui est tres proche de P sindoi

Lcsautres especes trouvccs en Polynesie frangaise sont P. edwardsii (Brandt. 1851), P. mania (A. Milne Edwards. 1883).

P.semilaevis Rm. 1888, et P. spinidorsalis (Rathbun, 1906). Enfin. deux postlarves, appartenant vraisemblablement

il P.Jennen Crosnier, 1986, ont ete trouvdes et des informations additionnelles sont fournies pour cette espece.

INTRODUCTION

Species of the genus Plesionika Bate. 1888 are common caridean shrimps found in deep waters. Abundant

material of this genus was also obtained during the recent extensive deep sea trapping in French Polynesian waters

conducted by the Service Mixte de Surveillance Radiologiquc et Biologique [SMSRB] under the leadership of

J. POUPIN (Poupin, 1988, 1996; POUPIN et al., 1990; POUP1N & RICHER DE FORGES, 1991).

Until now, only 2 main taxonomic works on the French Polynesian Plesionika have been published

(Crosnier, 1986a, Chan & Crosnier, 1991). They considered 7 species reported in details. The other species of

the genus found in this area were not easy to identify and a lot of problems were encountered by several authors

(Crosnier. 1986a, Poupin, 1988, 1996; Poupin et al.. 1990; Poupin & Richer deforces, 1991) The present

study attempts to clarify these problems and reports all the Plesionika species known to date from French

Polynesia. Altogether 21 species are recorded and five are described here as new to Science.

List of the Plesionika species found in French Polynesia

Most ol the species of Plesionika can be fairly readily divided into groups showing strong affinities. The

o owing species groups are used here in order to make comparisons and discussions of the various species clearer,

ihey are listed in the order that, tor various reasons, they are studied in the following pages.

"P. carsini Crosnier, 1986" group

P. carsini Crosnier, 1986

P. poupini sp. nov.

"P. edwardsii (Brandt, 1851)" group

P. edwardsii (Brandt, 1851)

'p • ensis (A. Milne Edwards, 1881)" group

P reflexa Chace, 1985

"P‘ laevis (A. Milne Edwards, 1883)" group

P. fenneri Crosnier, 1986

P. nesisi (Burukovsky, 1986)

'P. macropoda Chace, 1939" group

P. macropoda Chace, 1939

P. williamsi Forest, 1964

p '• mania (A. Milne Edwards, 1883)" group

P. mania (A. Milne Edwards, 1883)

P. semilaevis Bate. 1888

Source

PLESIONIKA FROM FRENCH POLYNESIA

189

"P. narval (Fabricius, 1787)" group

P. narval (Fabricius, 1787)

P. spinipes Bate, 1888

P. flavicauda Chan & Crosnier, 1991

P. rnbrior Chan & Crosnier, 1991

P. curvata Chan & Crosnier, 1991

" P • rostricrescentis ( Bate, 1888)" group

P. erythrocyclus sp. nov.

"P. sindoi (Rathbun. 1906)" group

P. sindoi (Rathbun, 1906)

P. protar i sp. nov.

Affinities not clear

P. payeni sp. nov.

P. picta sp. nov.

P. spinidorsalis (Rathbun, 1906)

As shown in previous studies on this genus (e.g. CHAN & Yu. 1991: Chan & CROSNIER. 1991), knowledge

of the coloration and morphological variations from many localities, including the types and topotypic material, is

often essential in resolving intricate taxonomic problems. Since P. narval (Fabricius, 1787), P. spinipes Bate,

1888, P.fenneri Crosnier. 1986. P. carsini Crosnier, 1986. P. flavicauda Chan & Crosnier. 1991, P. rnbrior Chan

& Crosnier, 1991 and P. curvata Chan & Crosnier, 1991, have previously been described in detail, they are not

studied further in the present work, except P.fenneri for which two postlarvae are found and look rather different

from the adults.

MATERIAL AND METHODS

Unless otherwise stated, specimens from French Polynesia were mainly collected by trapping aboard the

R. V. "Marara”, of the SMSRB (detailed station data of "Marara" can be found in POUPIN, 1996) and deposited at

the Museum national d'Histoire naturelle, Paris (MNHN). Specimens from the National Taiwan Ocean University,

Keelung (NTOU), the National Museum of Natural History. Washington, D.C. (USNM), The Natural History

Museum, London (NHM) and the Zoological Museum of the Moscow Slate University (MSU) were also

examined.

In the lists of material examined, the names of the vessels are in both italics and quotation marks. The names

of the cruises are printed in capital letters.

The measurements provided are of carapace length (cl), which is measured dorsally from the orbital margin to

the posterior margin of the carapace. The colour photographs provided for the species were taken by J. POUPIN of

the SMSRB, except the figure 35 taken by the first author.

The synonymy provided for each species is mainly restricted to previous French Polynesian records and

important works (e.g. original description and major taxonomic revisions related to the species).

SYSTEMATIC ACCOUNT

Plesionika poupini sp. nov.

Figs 1-2, 22

Plesionika carsini Crosnier, 1986a: 369 (in part). —Poupin & RICHER DP. Forges, 1991: 211 (in part) [non Crosnier,

1986].

Plesionika aff. carsini - Poupin el al., 1990, pi. 3h [non Crosnier, 1986].

Plesionika sp. nov. 3 - POUPIN, 1996, pi. 4g.

Material EXAMINED. — Specimens photographed. French Polynesia. SMSRB (J. Poupin coll.): Tuamotu

Islands. Fangataufa. Stn 438, 22°12.3’S, 138°46.6'W, 410 m, 14.11.1994: 4 6 20.3-23.8 mm. I ovig. 9

22.8 mm (MNHN-Na 13074). — Maria. Stn 37, 22°00.0’S, 136°12.0'W, 470 m. 24.11.1987, 2 6 23.5 and 25.8 mm,

2 9 16.2 and 23.9 mm (MNHN-Na 12519. transferred to USNM). — Sin 72, 22°00.0'S, 136°17.0'W, 430 m,

10.06.1988, 1 6 24.5 mm, 1 9 18.3 mm (MNHN-Na 13080). — Sin 241, 22°00.9'S. 136°I2.5'W, 380 m, 30.05.

1990: 2 9 21.1 and 22.6 mm (MNHN-Na 13079). —Rimatara. Stn 151, 22°38.2'S. 152°49.7'W, 260 m, 11.03.1989:

190

T.-Y. CHAN & A. CROSNIER

1 d 22.3 mm (MNHN-Na 13078). — Rurutu. Sin 149. 22°27.2'S, 151°23.3'W, 520 m. 10.03.1989: 2 d 15 6 and

21.6 mm (MNHN-Na 13077).

Tubuai Islands. Raevavae. Sin 99, 23°55.0'S. 147°40.0'W, 450 m. 23.08.1988: 1 ovig. 2 26.6 mm (MNHN-

Na 13122).

Specimens not photographed. French Polynesia. S.MSRB (J.-L. Carsin and J. PouPIN coll.): Tuamotu

Islands. Mururoa. 350-600 m, 1984: 1 ovig. 2 22.2 mm (MNHN-Na 7718). — Sin 53. 21°53.7'S, I38°57.3'W.

510 m. 11.12.1987: 1 d 21.1 mm. 1 2 21.1 mm (MNHN-Na 12520). — Stn 164, 21°47.3'S. I38°55.6'W. 360 m

21,06.1989: 2 8 14.2 & 21.2 mm (MNHN-Na 13072). — Makemtt. Sin 68, 16°36.0'S, 143°33.0'W 510 m

4.06.1988: I 8 24.0 mm (MNHN-Na 13073). — Sin 317, 21°53.8'S, I39°1.6'W. 500 m. 19.10 1990’ 1 d ’0 7 mm

(MNHN-Na 13075).

(MNHNN* 13076^ RaeVaVae Sln 100' 23°55 0's- 147°40.0'W. 530 m. 24.08.1988: 1 d 19.6 mm. I 2 18.4 mm

Below we give Ihe list of ihe specimens belonging in P. carsini examined for comparison with P. poupini.

Specimens photographed. French Polynesia. SMSRB (J. Poupin coll.): Gambier Stn 311 23°04 0'S

I35°01.6'W, 470 m, 11.10.1990: 1 ovig. 2 28.2 mm (MNHN). ’

Tuamotu Islands. Fangataufa. Sin 438, 22°12.3'S. I38°46.6'W, 410 m, 14.11 1994- 5 ovig 2 22 3-

27.3 mm. 3 2 17.3-26 mm (MNHN). — Maria. Sin 72, 22°00,0'S, I36°17.0'W, 430 m. 10.06 1988' 1 d 25 7 mm

[M""y -M""‘roa. Sin 39, 2I°49.3'S, !38°46.5'W, 470 m, 2.12.1987: 5 d 18.8-24.1 mm, I 2 17.8 mm (MNHN-

Na 12518. transferred to USNM). — Tureia. Sin 36. 20°45.0’S, 138°31.0'W, 490 m, 24 1 1 1987 2 d 15 6 and

20.8 mm, 1 ovig. 2 22.4 mm, 1 2 13.4 mm (MNHN-Na 12523).

Taiwan. Northeastern coast . Tai-Chi , l-Lan Counly, commercial trawler, about 300 m. 9.05.1989: 1 d

CNTOU). — 1 1 .07. 1989: 1 d 21.5 mm (NTOU, transferred to MNHN); 1 d 21.1 mm, I ovig. 2 23.1 mm

(NTOU). — 1.09.1989: 2 d 20.0 and 21.2 mm (NTOU). — 5.03.1993: 1 ovig. 2 25.4 mm (NTOU). — Su-Aou l-Lan

County, commercial trawler, about 300 m, 17.06.1993: 1 2 15.7 mm (NTOU).

Specimens not photographed. French Polynesia. SMSRB (J.-L. Carsin and J. Poupin coll.): Tuamotu

Islands. Make, no Jtn 68, 16°36.0'S. 143°33.0'W, 510 m, 4.06.1988: 1 d 24.4 mm (MNHN). - Mururoa. 450 m.

1984. l ovtg. 2 24.7 mm, holotype (MNHN-Na 7714); 1 ovig. 2 23.5 mm. I 2 20.2 mm, paratypes (MNHN-Na

77 7 ; ov,g' 2 23'4mm paralype 'MNHN-Na 7716). - 350-600 m, 1984: 1 d 15.7 mm, allotype (MNHN-Na

(MNHN °V’8' 2 22 mm ,MNHN‘Na 77I9)' ~ Stn 164‘ 21°47.3'S, 138°55.6'W. 360 m. 21.06.1989 I 2 16 2 mm

TYPES. The type series consists of the specimens photographed. The holotype is the ovigerous female (cl =

26.6 mm) collected in the Tubuai Islands at Raevavae (MNHN-Na 13122). The other specimens photographed are

paratypes. ° r

lAGNOSis. — Rostrum 1.28-1 .61 (avg. 1.4) times length of carapace, only slightly recurved, overreaching the

scaphocerite by hall or 2/5 of its length, armed with 3-5 dorsal and 1 1-14 ventral fixed teeth. The dorsal teeth are

on the basal 3/5 of the rostrum, the ventral along whole length. Five to 8 small post-rostral teeth, usually all

%6J^h°l0tyPe <MNHN’Na ,3I22)’ French P°'~ ^buai.

Source :

PLESIONIKA FROM FRENCH POLYNESIA

191

movable. Eye sub-pyriform, maximum diameter about 1/5 carapace length, ocellus very clear. Stylocerile acute,

just overreaching distal margin of the basal antennular segment. Scaphocerite nearly 2/3 length of carapace and

about 4 times as long as wide, distolateral tooth falling slightly short of distal margin of blade*! Maxilliped III and

pereiopods I-IV with an epipod. Penultimate segment of maxilliped III as long as distal one. Pereiopods 2

subequal, with 19-21 carpal articles. Pereiopods III overreaching the scaphocerite by lengths of dactyl, propodus,

carpus and about 1/7 of merus; dactyl short, about 1/15 as long as propodus. Abdomen without posteromesial

tooth or median dorsal carina on 3rd somite, 4th and 5th pleuron with a postcroventral tooth, 6th somite 1.65 to

1.75 times as long as maximum height and 1.55 to 1.65 longer than 5th; telson 1.45 to 1.55 longer than 6th

somite, with 5 pairs of dorsolateral spinules, including pair adjacent to lateral pair of posterior spines.

Remarks. — As in previous papers (Chan & Yu, 1991; Chan & Crosnier, 1991), coloration is here

employed as an important character for the identifications of the species of Plesionikci. However, ihe use of

coloration poses a difficult dilemma in the case of P. carsini Crosnier. 1986. As pointed out by Poupin et al.

(1990), two completely different patterns are present in the material of "P. carsini" from French Polynesia and

these sometimes even occur in the same catch. One form has the body reddish with red and white stripes, only the

antennular flagella arc banded with red and white, and a large red spot is present on the dorsum of abdominal somite

III (fig. 21, hereafter referred to as the "striped" form). The eggs of this form are bright blue. The other form is

orange, with many conspicuous white spots evenly distributed over the entire body while both the antennular and

antennal flagella are banded with red and white (fig. 22. hereafter referred to as the "spotted" form). The eggs of this

form appear to be mud-green, but this may due to the eggs being near to hatching. Fresh material of these two

forms can be readily separated on board ship when they just come out of the water. However, their coloration

quickly fades after preservation and they become almost indistinguishable morphologically.

Table 1. — Comparison of the morphological characters between Plesionika carsini Crosnier, 1986 (striped form) and

P. poupini sp. nov. (spotted form).

Only data from specimens photographed in colour were used. Numbers in brackets represent the average values.

Our best efforts to find morphological differences between these two forms were mostly unsuccessful. Of the

54 French Polynesian specimens available for the present study, 37 are accompanied by photographs showing

their fresh coloration. Amongst them, 17 specimens belong to the spotted form, while the other 20 are of the

striped form. Samples of both forms contain specimens of all sizes and sexes, including ovigerous females.

However, only 28 (14 specimens for each form) are almost intact and have a complete rostrum. Their

morphological characteristics, including the shape of the dactyli of the posterior pereiopods, are essentially

identical, except that the rostrum is slightly longer in the spotted form (Table 1). When rostral length is plotted

against carapace length, the two forms fall quite well in two separate categories (fig. 2a). Figure 2a shows that, for

specimens of similar sizes, the rostrum of the spotted form is always longer than that of the striped form. For

example, the rostrum of adults is less than 1.3 times carapace length in the striped form, but usually more than

1.3 times as long as the carapace in the spotted form. Although the rostrum is relatively longer in small

specimens (< 18 mm cl), it is 1.45 times the carapace length in the spotted form and less than 1.4 times as long

as the carapace in the striped form.

192

T.-Y. CHAN & A. CROSNIER

14

□

□

□

. ' +

P'' +

aP

P'*'

+ +

□'* +

x +

+

+

□

O. • ' '

+

16

- *+

18

20

22 24 26 28 cl (mm)

Fig. 2a. — Rostrum length/carapace length of the specimens assigned to Plesionika carsini Crosnier, 1986 (striped form)

and P. poupini sp. nov. (spotted form) from specimens with colour photographs (unless otherwise stated

the specimens are from French Polynesia).

+, P. carsini with photo (stripped form, n= 14). — □, P. poupini with photo (spotted form, n= 13). _

't;, P. carsini trom Taiwan (n = 4). — ■, P. poupini with photo (holotype).

14

□

16

!8

20

22

24

26

28 cl (mm)

FVb- _ RosIrum length/carapace lengih of ihe specimens assigned 10 Plesionika carsini Crosnier, 1986 and

P. poupini sp, nov. in the present study (unless otherwise stated, the specimens are from French Polynesia).

+. P. carsini with photo (n=l4). — *, P. carsini from Taiwan (n = 4). — ▲, P, carsini without photo (holotype).

~tPp CarS"" Wlth°ut photo (allotype) - •. P. carsini without photo (paratypes). — x. P. carsini without photo.

□ , P. poupini with photo (n— 13). — ■. P. poupini with photo (holotype). — 0, P. poupini without photo.

Source :

P LESION I KA FROM FRENCH POLYNESIA

193

The fact that preserved material of the two forms can only be separated by a slight difference in the proportional

length of the rostrum is not entirely satisfactory. The greatest difficulty is that the rostrum is often incomplete in

the specimens of Plesionika, making positive identification impossible. However, the patterns of the two forms

are so different that it is hard to believe that all the specimens could belong to the same species. It is significant

that only the striped form is found in Taiwan (the Taiwanese material is somewhat redder, but this may due to the

fact that the specimens were already dead when taken from fishing pots). The general characters of the Taiwanese

material are very similar to those of the French Polynesian striped form (Table 1 ); only the dactylus of pereiopod

III is somewhat longer (but this may be a general phenomena for species of this genus with a wide geographical

distribution). Of the four Taiwanese specimens with a complete rostrum, three fall in the group of the French

Polynesian striped form in fig. 2a. The fourth Taiwanese specimen (cl = 21.1 mm) with a relatively longer

rostrum, also has an unusual high number of ventral rostral teeth (19), and is probably abnormal. Since no

specimen with intermediate coloration has been found, and the difference in coloration appears to correlate quite

well with the length of the rostrum, the striped and spotted forms are considered to have distinct specific status.

Hence the rostral length of the types of P. carsini , as well as those specimens not accompanied by colour

photographs, was plotted against carapace length to determine which forms they belong to (fig. 2b). All the types

of P. carsini fall in the category of the striped form (only one non-type specimen [MNHN-Na 77 1 8 1 listed in

CROSNIER, 1986 falls in the range of the spotted form). Therefore, the name P. carsini should be used for the

striped form and a new name, P. poupini sp. nov., is proposed for the spotted form.

Size. — Largest male and female 25.8 and 26.6 mm cl, respectively. Smallest ovigerous female 22.2 mm cl.

ETYMOLOGY. — This species is named after Joseph POUPIN of the SMSRB. Without his invaluable efforts in

collecting many intact specimens and taking colour photographs of the fresh material, the recognition of this new

species would probably have been impossible.

Distribution. — At present known only from French Polynesia, at depths of 280-600 m.

Plesionika edwardsii (Brandt, 1851)

Fig. 23

Pandalus (Pontophilus) Edwardsii Brandt, 1851: 122 [type-locality: Mediterranean].

Panda l us ( Parapandalus) longirosiris Borradaile, 1900: 413, pi. 37, fig. 10 a-h [type-locality: New Britain).

Plesionika edwardsii - POUPIN, 1988: 28; 1996: pi. 3 h. —Poupin el al ., 1990: 68, pi. 1 g. — Chan & Yu. 1991: 550,

figs 2, 3 b.

Material examined. — French Polynesia. SMSRB (B. Richer de Forges, J.-L. Carsin and J. Poupin coll.):

Tuamotu Islands . Mururoa. 250 m, 4.11.1978: 2 ex. (MNHN-Na 7236). -350-400 m. 4.04.1979: 5 ex. (MNHN-

Na 7239). — 1984: 9 ex. — 1984: 11 ex. — Stn 16, 21°50.0,S, 138°57.2’W, 390 m, 11.12.1986: 14 ex.

Tubuai Islands. Raevavae. Sin 100, 23°55.0'S, 147°40.0,W, 450 m. 24.08.1988: 1 ex. — Rapa. Stn 101,

27°36.0'S, 144°16.0'W, 280 m. 26.08.1988: 4ex. — Stn 430. 27°36.2’S, 144°16.3'W, 290 m, 17.08.1991: 5 ex. —

Tubuai: 400 m, 13.05.1979: 23 ex. (MNHN-Na 7254). — 200 m, 14.05.1979: 20 ex. [MNHN-Na 7243]. — 300 m: 88

ex. [MNHN-Na 7244]. — 300 m: 6 ex. (MNHN-Na 7238, transferred to USNM). — 300 m: 1 ex. (MNHN-Na 7241).

Society Islands. Tahiti (Vairao). October 1978: 5 ex. (MNHN-Na 7237).

Diagnosis. — Detailed description and colour pattern of this species can be found in Chan and Yu (1991;

also with full synonymy).

Size. — The largest male and female in the present French Polynesian material are 25.7 mm (about 28 mm in

Poupin et al., 1990) and 26.1 mm cl, respectively. The smallest ovigerous female is 20.7 mm cl.

Remarks. — This species has been recorded in French Polynesia by Poupin (1988. 1996) and POUPIN et al.

(1990). All (total number 194), but one of the French Polynesian specimens agree very well with the concept of

this species as defined by Chan & Yu (1991). The exceptional specimen (12.0 mm cl) has only one post-rostral

194

T.-Y. CHAN & A. CROSNIER

tooth on the carapace, but it corresponds to P. edwardsi in all other characters. Therefore, it is still considered that

the closely related P. crosnieri Chan & Yu, 1991 is not present in French Polynesian waters.

DISTRIBUTION. — Worldwide: Mediterranean, Atlantic and Indo-West Pacific, at depths of 50-680 m. com¬

monly 200-400 m.

Plesionika reflexa Chace, 1985

Figs 3. 24

Plesionika reflexa Chace, 1985: 108, fig. 49 (type-locality: the Philippines], — CROSNIER, 1986a: 362 — Poupin

1996. pi. 5 a.

Plesionika aff. ensis - Poupin, 1988: 28 [non A. Milne Edwards, 1881].

Plesionika ensis - Poupin el al„ 1990: 64. pi. I h [non A. Milne Edwards, 1881],

Material examined. — French Polynesia. SMSRB (J. Poupin and B. Richer de Forges coll.): Marquesas

Islands. Fatu Hiva. Stn 61, 10°27.0'S. 138°41.0’W. 430 in, 28.01.1988: 17 ex. 8.3-17.4 mm.

Tuamotu Islands. Maria. Stn 37. 22°00.0,S. I36°I2.0,W, 470 m, 24.11.1987: 3 ex. 11.7-13.1 mm (MNHN-Na

12524). — Mururoa. Stn 18. 21°49.9'S. I38°57.3'W, 500 m, 11.12.1986: 1 ex. 15.6 mm. — Stn 21. 2I°49.9'S.

1 38°57.3'W, 630 m, 16.06.1987: 10 ex. 14.4-18.4 mm. — Stn 39, 21°49.3'S, 138°46.5'W, 470 m 2 121987- 9 ex'

13. 7-17. 4mm (MNHN-Na 12522). — Stn 477, 21°51.3'S, 139°01.7'W, 800 m. 11.04.1995: 3 ex. 15.9-16 7 mm —

Tureia. Stn 36. 20°45.0'S. I38°31.0'W. 490 m. 22.1 1.1987: 3 ex. 8.8-16.6 mm (MNHN-Na 12529)

Tubuai Islands. Tubuai. 700 m. 14.05.1979: 45 ex. 10.3-20.5 mm (MNHN-Na 7263); 25 ex. 15.1-20.3 mm

(MNHN-Na 7262). — 800 m, 14.5.1979: 6 ex. 15.4-19.2 mm (MNHN-Na 7261). — Rimatara. Stn 109, 22°38.0'S,

152°50.0'W. 700 m. 5.09.1988: 18 ex. 14.1-18.0 mm (transferred to USNM).

Diagnosis- — A detailed description of this species can be found in Chace (1985), though there are still

many uncertainties concerning the relationship between P. reflexa and P. ensis (A. Milne Edwards, 1881).

Coloration. — The French Polynesian material is somewhat transparent with red tints on the carapace

(including appendages), anterior half of rostrum, posterior margins of abdominal somites, pleopods and tail-fan.

Eyes black. Eggs bluish.

Size. — For the French Polynesian material, the largest male and female are both about 20 mm cl, and

smallest ovigerous female measured 12 mm cl (Poupin etal., 1990, under the name "P. ensis").

Remarks. — As mentioned by CROSNIER (1986), the material from French Polynesia has the dactylus of

pcreiopod III much shorter (0.09-0.14, avg. 0.1 1 as long as propodus; fig. 3b) than those from the Philippines

(0.30-0.46 as long as propodus - Chace, 1985; two paratypes and two other Philippine specimens were also

examined). The short dactylus of the French Polynesian material is therefore more similar to that of P. ensis

(A. Milne Edwards. 1881), described from the Antilles (with a ratio of 0.1 1-0.21, avg. 0.14; 20 specimens from

the Antilles in MNHN examined; see also Chace, 1985). The coloration (including eggs) appears to be rather

similar lor the French Polynesian material (fig. 24) and P. ensis (fig. 25, also sec PAULM1ER, 1993. pi. 15 figs 1-

4, pi. 16 figs 1-3). However, the posterior spine of the abdominal somite III is very often curved upwards in the

French Polynesian material (87.1%, n = 140), but not such condition occurs in the Antilles specimens.

Fhe problem is complicated by the fact that specimens deposited at the MNHN, belonging to the "P. ensis"

group Irom various localities, all show different proportional length of the dactylus of perciopod III. The dactylus

is longer in the west African specimens [0.26-0.41, avg. 0.31, as long as propodus, 10 specimens (MNHN-

Na7865) from Gulf of Guinea examined; see also Chace, 1985] but shorter in the few specimens from Kiribati

and Seychelles examined [the ratios are 0.10 (n = 2) and 0.12 (n = 3), respectively], Chace (1985) also mentioned

such variation on the relative length of the dactylus for specimens from western Africa, Indonesia and Hawaii,

Furthermore, the dactylus of pereiopod III appears to be one third the length of the propodus in specimens from

eastern Austraha (Kensley etal., 1987), but only about one fifth to one fourth of the propodus in material from

the Arabian Sea (Suseelan & Mohamed. 1969) and Japan (Hayashi. 1986).

Source :

PLESIONIKA FROM FRENCH POLYNESIA

195

Fig. 3. — Plesionika reflexa Chace, 1985: a, ovig. 9 17.1 mm (MNHN-Na 7262), French Polynesia, Tubuai Islands,

Tubuai, 700 m: lateral view of abdominal somites II to V. — b, ovig. 9 18.7 mm (MNHN-Na 7262). French

Polynesia, Tubuai Islands, Tubuai, 700 m: propodus and dactylus of pereiopod III.

On the other hand, the posterior spine on the abdominal somite III of the material available from Seychelles,

Kiribati and New Caledonia is frequently curved upwards, but not in eastern Atlantic specimens. The tendency of

having a recurved posterior spine on abdominal somite III seems to be universal throughout the Indo-Pacific [e.g.

Arabian Sea (Suseelan & MOHAMED, 1969); Japan (Hayashi, 1986); northwestern Australia (Hanamura &

TAKEDA, 1987); eastern Australia (KENSLEY et al., I987)|. In the present study, it was also found that the rostrum

of the French Polynesian material is generally shorter and has fewer ventral teeth (rostrum 1 .6-2.5, avg. 2. 1 times

as long as carapace and bearing 28-42, avg. 35.7 ventral teeth, n = 27) than in P. ensis from the Atlantic (western

Atlantic: rostrum 2. 2-2. 8, avg. 2.5 times carapace length and bearing 35-48, avg. 42 ventral teeth, n = 16; eastern

Atlantic: rostrum 2. 2-2. 8, avg. 2.5 carapace length and having 39-50, avg. 44.4 ventral teeth, n = 7). However, the

few specimens from the Kiribati and Seychelles have a slightly shorter rostrum (both avg. 2.2 cl, n = 2 and 3

respectively) and fewer ventral rostral teeth (avg. 37 and 38.7 teeth, n = 2 and 3 respectively), while those from

New Caledonia have a longer rostrum (2.3-2.7cl, n = 3) but fewer ventral rostral teeth (35-38, n = 3)! Since the

rostrum of P. ensis/reflexa is often broken when caught, additional intact specimens from various localities will be

needed to determine whether the differences in the proportional length and armature of the rostrum are of any

taxonomic value. Nevertheless, the absence of any specimen with a recurved posterior spine on the abdominal

somite III in the Atlantic is sufficient reason not to synonymize P. reflexa with P. ensis (altogether, 91 Atlantic

specimens were examined in the present study and in Chace, 1985). For the time being, it may be more

convenient to use the name P. reflexa for the French Polynesian or perhaps all the Indo-Pacific material, though

the presence of some specimens from various Indo-Pacific localities with very straight posterior spine on the

abdominal somite III is rather problematic (fig. 3a). Recent discoveries that a great diversity of species is actually

contained in the "P. mania (A. Milne-Edwards, 1883)" (Chace, 1985) and ”P. natval (Fabricius, 1787)" (Chan &

CROSNIER, 1991) groups suggest that many more species may be also present in the material now identified as

P. ensis and P. reflexa. On the other hand, it is also possible that the P . ensis-P. reflexa problem may be similar

to the case of P. edwardsii-P. crosnieri , in which both species are distributed in the Indo-Pacific but only one of

them occurs in the Atlantic (Chan & Yu, 1991).

196

T.-Y. CHAN & A. CROSNFER

Distribution. — Probably widely distributed in the Indo-Pacific (see "Remarks"). The French Polynesian

material occurs at depths of 360-910 m, but mostly between 550-700 m (Poupin et al ., 1990, under the name

P. ensis ).

Plesionika fenneri Crosnier, 1986

Figs 4, 26-27

Plesionika chacei Crosnier, 1986a: 363, figs 1 a-b. 2 a-h [type-locality: French Polynesia]

nesiomka fenneri Crosnier. 1986b: 691 [nomen nov. for Plesionika chacei Crosnier. 19861. — Poupin 1988

1996, pi. 4 b. — Poupin et ai, 1990: 29. pi. I a. — Poupin & Richer de Forges, 1991- '’ll

Plesionika all. trispinus - Poupin et ai, 1990: 16 [non Squires & Barraguan. 1976].

Material EXAMINED. — French Polynesia. SMSRB (J. Poupin coll.): Specimens lisied in Crosnifr (1986)

Additional specimens: '

Tuamotu Islands. Maria. Sin 37. 22°00.0’S, I36°I2.0'W, 470 m. 24.11.1987: 7 ex (MNHN-Na 12317)

GWner. S,n 74 23°08.0'S. 134‘53.0’W. 660 m. 10.06.1988: I pos.larva 12.4 mm (transferred Vo USNM) '

^Tubua, Islands. Rurutu. S.n 146. 22°27.8'S, 151°22.8'W. 580 m, 9.03.1989: 1 pos.larva 12.9 mm (MNHN-Na

DIAGNOSIS, — This species was described in detail by Crosnier ( 1 986a. under the name "Plesionika chacei").

COLORATION - Body generally orange. Tip of rostrum, distal par. of scaphoceri.e, distal segment of

maxilliped III and antennular flagella reddish. Eyes dark brown. Internal organs scarlet, visible inside carapace.

Eggs metallic blue (Crosnier, 1986a), becoming mud green near hatching.

(POUPIN r7r//La 1990 )malC and fcmale 30 mm and 32 mm d’ resPec,ive|y- Smallest ovigerous female 20 mm cl

Remarks. - P. fenneri is one of the most abundant species of the genus in French Polynesia Its

charactenst.cs are rather constant, except that the rostrum is often rather straight in males (also see Crosnier

6a. fig. lb). The mam problem is posed by the two small specimens collected from Gambier and Rurutu and

preliminary identified by Poupin et ai (1990) as Plesionika aff. trispinus. These two specimens are white in

colour and look quite d.fferent from the other species of the genus (fig. 26). At first glance they show some

resemb'ances to P. p.cta sp. nov. but closer examination shows that they actually belong to a species of the

P. laevis group. However, they differ considerably from the typical P. fenneri (fig. 27) and P nesisi

Burukovsky ,1986) by the carapace being more flattened and rectangular. Their rostra are rather straight and bear a

laminate basal crest (fig. 4a). However, as discussed in the "Remarks" under P. nesisi. the shape of fhe rostrum is

very variable for the species of the "P. laevis" group. On the other hand, the presence of rudimentary exopods on

anterior four pereiopods and the exopod of the maxillipcd III being rather small and thin suggest that these two

e;:;rrsm izn early deve,°pmentai s,agc- Therefore’ ,hey are possibiy thc p«*i™ 0f e^ p

though1 5! " both specimens have one or two movable post-rostral teeth, the latter possibility is ruled out

, ' C3rapaCes a,s° bear a 'veak uPPer laIeral carina posteriorly. Moreover, the other characters of these two

postlarvae agree quite well with those of P. fenneri (see Crosnier, 1986a): rostrum 0.9- 1.0 as lone as carapace

Se6" 1 arn 10rral 1Ce,h; 5 p0St-r0Slral present on carapace, with posteriormo^t "ne orTo

movable, eyes (sunken in the present specimens) appear to be subspherical and lacking ocellus; orbital margin

regularly concave, upper part not continuous with lateral rostral carina; antennal spine rather small while

nfo mm" T *°T ^ Thcie ,W° pos,larvae are therefore tentatively identified as P fZeH More

Nevertheless om- ° ' ", ^ ^ the P-m StTf'catn

ertheless, if our identification is correct, these two specimens should be in the final postlarval stage, since

Source :

PLESIONIKA FROM FRENCH POLYNESIA

197

juvenile specimens of P. fenneri larger than 13.2 mm cl all lack exopods on the pereiopods and already look very

similar to the adults.

Distribution. — Endemic to French Polynesia, at depths of 260-820 m and mostly 500-700 m (Poupin

etaL, 1990).

Fig. 4. — Plesionika fenneri Crosnier, 1986, postlarva 12.9 mm (MNHN-Na 13125), French Polynesia, Tubuai Islands,

Rurutu, stn 146, 580 m: a, lateral view of carapace; b, lateral view of posterior abdomen; c. dorsal view of abdominal

somite III; d-e, propodus and dactylus of pereiopod III.

Plesionika nesisi (Burukovsky, 1986)

Figs 5-10, 28

Heterocarpus nesisi Burukovsky, 1986: 62, fig. 1 [type-locality: East Pacific sea-mount]. — Chace, 1989: 87.

Plesionika aff. laevis - Poupin et al., 1990: 72, pi. 2 a. [non A. Milne Edwards, 1883].

Plesionika nesisi - POUPIN, 1996, pi. 4e.

198

T.-Y. CHAN & A. CROSNIER

Fig. 5. — Plesionika nesisi (Burukovsky, 1986). Lateral view of carapace,

a. ovig. 9 26.7 mm (MNHN-Na 13118) with abdominal pleuron IV rounded, French Polynesia, Tuamotu Islands,

Fangataula, stn 85, 780 m . — b, ovig. 9 27.6 mm (MNHN-Na 13115) with abdominal pleuron IV pointed, Tuamotu

Islands, Mururoa, stn 477, 800 m. — c, 6 25.6 mm (MNHN-Na 13120) with abdominal pleuron IV rounded. Tuamotu

Islands, Makcmo, stn 112, 670 m. — d, 6 25.9 mm (MNHN-Na 13117) with abdominal pleuron IV pointed, Tuamotu

Islands, Fangataufa, stn 139, 720 m.

Source :

PLES10NIKA FROM FRENCH POLYNESIA

199

MATERIAL EXAMINED. — East Pacific seamount. 13°34'N, 120°33'W. 800 m: 8 26.0 mm. holotype (MSU).

French Polynesia. SMSRB (J. POUPIN coll.): Marquesas Islands . Hiva Oa. Stn 293. 9°47.3'S. 139°11.8’W.

800 m, 30.08.1990: 1 8 21.4 mm (MNHN-Na 13116. drawn).

Tuamotu Islands. Fangataufa. Stn 85, 22°13.0’S. 138°42.2'W. 780 m. 22.06.1988: 4 8 23.5-25.1 mm.

6 ovig. 9 23.2-29.3 mm (MNHN-Na 13043, 13046); 1 ovig. 9 26.7 mm (MNHN-Na 13118, drawn). — Stn 118,

22°16.8'S, 138°4I.8'W, 1050 m, 23.11.1988: 5 8 16.4-24.7 mm, 2 ovig. 9 21.7 and 24.1 mm, 3 9 19.0 & 20.3 mm

(MNHN-Na 13045). — Stn 139. 22°18.0*S. 138°46.0'W. 720 m, 24.02.1989: 32 8 18.5-26.8 mm, 2 ovig. 9 19.5 and

23.4 mm, 13 9 18.1-28.6 mm (MNHN-Na 13048); 1 8 25.9 mm (MNHN-Na 13117. drawn). — Make mo. Stn 112,

16°37.0'S, 143°32.0'W. 670 m, 12.11.1988: 10 8 23.2-27.0 mm, 1 9 24.2 mm; 1 8 25.6 mm (MNHN-Na 13120,

drawn). — Mururoa. Stn 127, 21°51.2'S, 138°47.3'W, 600 m, 29.11.1988: 3 8 22.1-28.1 mm, 4 9 24.7-27.7 mm

(transferred to USNM). — Stn 474. 21°47’S, 138°55.5’W, 800 m, 10.04.1995: 2 8 22.0 and 22.4 mm. 2 9 16.9 and

26.1 mm. — Stn 477, 21°51.3’S, 139°1.7’W. 800 m. 11.04.1995: 1 8 23.9 mm, 4 ovig. 9 24.1-27.7 mm, 1 9

23.6 mm; 1 ovig. 9 27.6 mm (MNHN-Na 13115, drawn).

Kiribati. Commission du Pacifique Sud , 600 m, April 1987: 6 ex. 15.8-24.9 mm; 1 9 28.3 mm (MNHN-Na 13119,

drawn). — 750 m. April 1987, 12 ex. 18.0-28.0 mm (MNHN-Na 12487, 12501); 1 9 18.0 mm (MNHN-Na 13121,

drawn).

Samoa Islands. "Coriolis": stn 92, Lallarokkhe. 800 m, 26.11.1977: 6 ex. 27.0-29.8 mm (MNHN-Na 13050).

Philippines. Musorstom 2: stn 56, 13°57.7’N, 119°56.3’E, 970 m, 28.11.1980: 1 8 18.6 mm, 2 9 25.7 and

31.1 mm.

FlG. 6. — Plesionika nesisi (Burukovsky, 1986). Lateral view of carapace: a. 8 21.4 mm (MNHN-Na 131 16), French

Polynesia, Marquesas Islands, Hiva Oa, stn 293, 900 m. — b. 9 18.0 mm (MNHN-Na 13121), Kiribati, 750 m;

c, 8 20.4 mm (MNHN-Na 12487), Seychelles, Cepros. sin 3-14. 850-800 m.

200

T.-Y. CHAN & A. CROSNIER

Seychelles Islands. Cepros: stn 3-14. 4°33'S, 56°27,E. 850-800 m. 22.10.1987: 1 6 20.4 mm (MNHN-Na

12487). — Stn 5-25, 5°48.5'S, 56°43.2'E, 700-750 m, 24.10.1987: 1 6 19.0 mm (MNHN-Na 12486).

Madagascar. " Vauban trawling 35. 12°49.5'S. 48°5.9'E. 760-810 m. 14.09.1972: I 6 22.8 mm, 1 9 20.7 mm.

Diagnosis. — Body rather robust. Rostrum about as long as carapace and armed along whole length with

5-8 (very rarely 3, 4 or 9) dorsal and 6- 1 1 ventral teeth, generally curved upwards in females but nearly horizontal

in males. Carapace with a low post-rostral crest, which bears 4-6 fixed post-rostral teeth and extends for about 4/5

of carapace length; antennal and branchiostegal spines very strong (branchiostegal spine relatively larger), both

with weak lateral carina behind; upper lateral carina starting at orbital margin, slightly above antennal spine, and

extending almost to posterior margin of carapace, weak, more or less interrupted at mid-length, and strongly curved

in the posterior part; lower lateral carina continuous with branchiostegal spine, low and soon vanishing

posteriorly; very weak hepatic groove present. Eyes quite large, slightly kidney-shaped and without distinct

ocellus. Orbital margin regularly concave. Stylocerite tapered anteriorly and reaching between middle and distal end

of second segment of antennular peduncle. Scaphocerite 3.4-4. 4 times as long as broad, with distolatcral tooth

more or less extending to distal margin of lamella, basicerite spine elongate and exceeding proximal end of lateral

margin of scaphocerite. Well developed epipods present on maxilliped III and anterior 4 pereiopods. Maxilliped III

bearing a long and thread-like cxopod, with distal segment 1.3- 1.5 times as long as penultimate segment.

Pereiopods II with 6-1 1 carpal articles on one side (usually right) and 18-21 carpal articles on the other side.

Pereiopod III extending beyond scaphocerite more or less by propodus and dactylus, with propodus 5.2-7.8 times as

long as dactylus; dactylus claw-like and bearing 2-4 spinules on posterior margin, accessory spine about half to

nearly as long as terminal spine and well separated from it. Abdominal somite III bearing a dorsal boss and,

generally, a pair of distinct submedian furrows, posterior margin bluntly angular. Abdominal pleuron V. and

sometimes also pleuron IV, with sharp elongated posteroventral denticles. Telson much longer than abdominal

somite VI; with lour pairs of dorsolateral spines and three pairs of terminal spines. Eggs subsphcrical. about

0.5 mm in diameter.

Fig. 7. — Plesionika nesisi (Burukovsky, 1986). Lateral view of carapace: a. 6 26.0 mm. hololype (MSU), East Pacific

sea-mount. 13°34'N. 120°33’W, 800 m; b. 2 28.3 mm (MNHN-Na 13119), Kiribati. 600 m.

Coloration. — Body, including appendages, generally reddish, colour slightly lighter at base of rostrum, as

well as area between posterior carapace and anterior abdomen. Eyes browish-black. Eggs brown.

Source :

P LESION I KA FROM FRENCH POLYNESIA

201

Size. _ Largest male and female about 32.5 mm and 31.5 mm cl, respectively (POUPIN et cil 1990). Smallest

ovigerous female 19.5 mm cl.

REMARKS. - Recent experimental trappings in deep waters m French Polynesia produced an abundant mater a

of the present form, which is very similar to P. laevis (A. Milne Edwards, 1883) (POUPIN^ rt/ ' ly ,0' Po,'F^'

1996). This form has a clear vertical zonation and is mainly found in waters deeper than 800 m (instead of 260-

820, mostly 500-700 m, for P. fenneri). A careful comparison of this deep water French Pol>'ne-',aiJ k'rm

P. laevis from the Caribbean (22 specimens with 1 1.1-39.4 mm cl m the MNHN including the badly damaged

juvenile holotype) shows that the former is quite different in having a redder body (see tigs -8-29) and al ay

bearing a weak but distinct upper lateral carina on the carapace. The French Polynesian term however exhibits

large variations in the shape of the rostrum and abdominal pleuron IV. The rostrum is generally strongly curve

upwards and exceeds the scaphoceritc by almost a quarter of its length (tig. 5a-b). but in some specimens (mostly

small males) the rostrum is relatively longer and almost horizontal (fig. 5c-d). Of the P °0 'spec.™™ .e*

posteroventral angle of the abdominal pleuron IV bears a sharp denticle in 66 specimens (fig. 8c), but is rounded in

20 others (fig. 8e), while in 14 specimens one side of the pleuron is rounded, while the other side bears a denticle.

The degree of development of the lateral carapace carinae and the abdominal boss is also very variable in these

specimens (e.g. tends to be weaker in small individuals, but this is not always size related; the same is also true

202

T.-Y. CHAN & A. CROSNIER

for the abdominal boss of P. laevis, that of the holotype being almost absent). The above characters, however,

appear to vary independently, and specimens of all sizes with intermediate characters can often be found in both

sexes (figs 5, 8-10). Moreover, no difference in coloration has been observed for these deep-water French

Polynesian specimens. It is, therefore, reasonable to consider that they all belong to the same species.

b

Fig. 9. Plesionika nesisi (Burukovsky, 1986). Dorsal view of abdominal somite III.

21 4 mr8fMNHN6Mn' nmTc Na J n ',8)' French Po,ynesia- Tuamotu Islands. Mururoa. stn 477, 800 m. — b. 6

(MNHN-Na 13116), French Polynesia. Marquesas Islands, Hiva Oa, stn 293. 800 m. — c. 9 18 0 mm

l?° m' 9 28 3 mm (MNHN-Na 13119), Kiribati, 600 m. — e, 6 20 4 mm

(MNHN-Na 12487), Seychelles, Cepros, stn 3-14, 850-800 m.

The discovery of P. fenneri brought up the problem of the generic affinity of the "P. laevis" group, which has

charactertstics intermediate between Plesionika and Heterocarpus A. Milne Edwards, 1881 (sec Burukovsky

86; CROSNIER, 1986a, 1988; CHACE, 1989). The present form no doubt belongs to this intermediate group!

Actually there are three more species which probably could also be assigned to the "P. laevis" group These are

Heterocarpus alexandri A. Milne Edwards. 1883 from the western Atlantic, H. unicarinatus Borradaile, 1915 from

Seychelles and H. nests, Burukovsky, 1986 from the eastern Pacific. These three species are all described from a

single specimen and very poorly known. The type-locality of H. nesisi is nearest to French Polynesia (i.e. from an

underwater seamount north-east of French Polynesia). A comparison of accurate drawings of the damaged holotype

o /. nests, (figs 7a 8a-b; prepared at MNHN some years ago) shows that it is almost identical with the French

Polynesian form. The rostrum of the holotype is missing in the MNHN drawings, but the original figure of

Source :

PLESIONIKA FROM FRENCH POLYNESIA

203

H nesisi provided by BURUKOVSKY (1986) shows thal it is also very similar to that of the French Polynesian

material, except for a slightly longer length and a greater number of dorsal rostral teeth.

FlG. 10. - Plesionika nesisi (Burukovsky. 1986). Propodus and dactylus of pereiopod 111.

a nvie 2 27 6 mm (MNHN-Na 13118). French Polynesia, Tuamotu Islands. Mururoa. sl" 477-j*')0 m: n8t]t sl^'

, ?' (MNHN Na 13116) French Polynesia, Marquesas Islands, Hiva Oa, stn 293, 900 m. right side.

cb*| H SlSSSSSirnni) KiS hOO i left side. -d. d 20.4 mm (MNHN-Na 12487). Seychelles, CBPROS.

stn 3-14, 850-800 m: left side.

For BORRADA.LE'S (1915) H. unicarinatus, photographs of the type were examined (also see CROSN.ER, 1988.

fig 7) The dried holotype has the rostrum broken and all the pereiopods missing. It is generally similar to he

Jfench Polynesian foJand has the upper lateral carina of the carapace medially interrupt* d However

abdominal boss is more pronounced in H. unicarinatus and there are only three post-rostral teeth present

carapace.

The type of H. alexandri was described in detail by Chace (1989). Although the rostrum of the type is broken

(apparently very straight) and most of the thoracic appendages are missing, the remaining parts are very similar to

those " the French Polynesian form. Nevertheless, the much smaller size of H. alexandn (holotype ov.gerou

female only 12.8 mm cl) suggests that it should be a distinct species. Photographs of the add, '°"a‘

mentioned by Chace (1989) from Bahama Islands, Baja California and Hawaii were examined m the present y .

r male lit Bahama Islands is in a similar condition as the holotype of H. alexandn (,,. rostrum broken and

most of the pereiopods missing) and with almost the same characteristics (their sizes are also sim.larL As

mentioned by Chace (1989), the female from Baja California .s much larger (about 3! .5 mm cl) and ^with he

abdominal boss rather weak. The general appearance of this specimen ,s more s,m,lar 10 ^

form (some of the French Polynesian specimens also have the transverse grooves on the anterior two abdominal

sornites°somewhat indistinct) except for having pereiopods II with fewer carpal articles.

specimen reported by Rathbun (1906) as H. alexandri, of which we were able to get good photos, looks very

different and is generally more similar to the "typical" forms of Plesionika or Pandalus. No trace of a dorsal boss is

pfent on I abdominal somite III and some of the post-rostral teeth appear to be movable. With I he a bsemee o

all thoracic appendage, in this Hawaiian specimen, there are even difficult, es dectd.ng

belongs to! Therefore, the exact identity of Rathbun's (1906) specimen may never be known. On the

204

T.-Y. CHAN & A. CROSNIER

it may be relevant to point out that the collection of MNHN includes a damaged eastern Atlantic specimen (Ibero-

Maroccan Gulf, Balgim, stn DW 96, 1250 m. adult female 11.8 mm cl, MNHN-Na 1 1919) very similar to

H. alexcindri , but with abdominal somite III smooth. This specimen probably represents an undescribed species of

the "P. laevis" group (Raso, 1996: 741).

• from figures and photographs of the holotype and the Baja California specimen.

Numbers in brackets represent the average values.

Table 2. — Characteristics of Plesionika nesisi (Burukovsky, 1996) from different localities.

In conclusion, the present French Polynesian form can probably be identified as H. nesisi. However, the

problem becomes more complicated when more, similar specimens from various Indo-Pacific localities with slight

differences are found in the MNHN collection (Table 2). The material from Kiribati has a slightly longer rostrum

(lig. 7b) and a stronger abdominal boss (tig. 9c-d), while the abdominal plcuron IV is more often rounded, and the

dactyli of posterior pereiopods are more slender, sometimes with a short accessory spine (fig. 10c). The few

specimens front Samoa, the Philippines and Madagascar also usually have the abdominal pleuron IV rounded, but

the degree ol development of the abdominal boss and the dactyli of posterior pereiopods is intermediate between

those of the French Polynesian and Kiribati material (figs 9-10). On the other hand, the two young males from the

Seychelles look somewhat different to the French Polynesian material, because they have a much longer, straight

rostrum (figs 6c, 9e). Nevertheless, this seems to be unimportant when considering the variations already exhibited

in the different Indo-Pacific populations, as mentioned above. Moreover, two similar-sized specimens from

Marquesas Islands (fig. 6a) and Kiribati (fig. 6b) have the rostrum quite similar to that of the Seychelles material

The two Seychelles specimens are very similar to the type of H. unicarinatus , which was collected from the same

locality and is of similar size and the same sex. The holotype of H. unicarinatus, however, bears only three post-

rostral teeth on the carapace, a condition that never occurs in all the specimens of this study (except for one French

I olynesian specimen with an abnormal rostrum). For the time being, it seems appropriate to identify all the

lrom„French Po|ynes|a- Kiribati, Samoa, the Philippines. Madagascar and the two Seychelles specimens

(MNHN) as W nesisi. More material from the Seychelles will be needed to clarify the relationships between

H. nesisi and H. unicarinatus.

ruJcei“g numbcr of species belonging to the "P. laevis" group may prompt some workers to follow

chaces (1989) suggestion and propose a separate genus for them. There is no doubt that these species form a

na ural group^ The complete absence of lateral carapace carina and abdominal boss in P. fenneri prevents the

inclusion of the P laevis" group in Heterocarpus. However, no satisfactory character can be found to separate

P. fenneri rom Plesionika as currently defined, which can easily accommodate H. nesisi, H. unicarinatus and

p alexandr' s‘nce some typical" Plesionika species also bear similar lateral carinae on the carapace (e.g

. cannata Holthuis, 1951 and P. polyacanthomerus Pequegnat, 1970, see also "Remarks" under P. williamsi).

Source

PLESIONIKA FROM FRENCH POLYNESIA

205

We therefore prefer to follow CROSNIER (1986a, 1988) in putting all the members of the "P. laevis " group in

Plesionika until this genus can be satisfactory subdivided.

Distribution. — Indo-Pacific from Madagascar to Seychelles, Philippines, Samoa, Kiribati, French

Polynesia and surrounding areas, perhaps also Baja California, at depths of 550-1080 m (POUPIN et al 1990).

Plesionika macropoda Chace, 1939

Figs 11-13, 30-31

Plesionika macropoda Chace, 1939: 37 [type-locality: Cuba]. — Monterrosa, 1988: 637, figs 4-5. — Poupin, 1994: 25

PlesionTa^Xacanthomerus Pequegnai. 1970: 97 (in pari). - Paulmier. 1993: 19. pis 19-20 [non Pequegnai. 19701.

[?1 Plesionika macropoda - SPRINGER & BULLIS, 1956: 12.

[?] Plesionika wiliiamsi - Lemaitre. 1984: 435 [non Forest. 1964],

Plesionika aff. wiliiamsi - Poupin el al., 1990: 16 (in pari) [non Forest, 1964).

Not Plesionika macropoda - Poupin. 1994: 25 (in part), pi. le [= P. williams , Forest. 1964],

MATERIAL EXAMINED. — Antilles. Guadeloupe, Cote sous le Vent. 400-450 m. 1985: 10 ex. 13.7-

22.5 mm; Id 18.1 mm (MNHN-Na 12847. drawn); 1 d 21.9 mm (MNHN-Na 13099. drawn)

"Polka": stn 4-3, 260 m, 3.10.1990: 1 spec 24.6 mm. - Sin C21 1, \6°\ 1.8! IN, 6\ 4)A n 5° m’ ^ ^pV

1 soec 17.1 mm. — Stn D38. 16°22.51'N, 61°49.13'W, 500 m. April 1993: 3 ex. 23.8-25.0 mm. — - Stn CPI,

15°50.57’N. 61°40.56'W, 310 m. May 1993: 3 ex. 16.1-23.0 mm. — Stn CP3, 16°0.irN, 61°21.16W, 400 m. May

1993: 4 ex. 19.6-23.1 mm {6 23.1 mm, MNHN-Na 13113. drawn).

Loyalty Islands. "Vauban": stn 43: I ex. 24.7 mm. - Stn 90. 200-600 m. 1.03.1978: 1 ex_ 26 4 mm.

French Polynesia. SMSRB (J. Poupin and J.-L. Carsin coll.): Marquesas Islands. La l on. Stn 5).

9°23.0'S, I40°09.0'W. 450 m. 26.01.1988: 1 d 19.1 mm [transferred to USNM[.

Tuamotu Islands. Maria. Stn 37, 22°00.0'S, 136°12.0'W, 470 m, 23.11 .1987: 1 9 19.1 mm (MNHN-Na 1 5 6.

drawn) - Gambler. Stn 238, 23°01.1'S, 134°59.0’W, 500 m, 27.05.1990: I d 14.6 mm, 1 8 19,3 mm. - Stn 311.

->3°04 5'S I35°0I 6'W, 470 m. 11.10.1990: 2 d 23.1 mm (MNHN-Na 13112. drawn) and 25.7 mm. — Mururoa.

1984: 1 8 18.6 mm. — Stn 382. 21°46.4'S. 138°53.5'W. 400 m, 9.03.1991: I ov.g, 2 26.9 mm. — Stn 476.

2 1 °5 1 3'S, 139°01.2'W, 470 m. 11.04.1995: 1 d 30.1 mm.

Tubuai Islands. Maria. Stn 421. 21°47.7'S, 154°43.4'W, 500 m. 7.08.1991: 1

^Society Islands. Huanine. Stn 405, 16°45.8 S. 151°03.8'W. 310 m. 6.05.1991:

13114, drawn); I ovig. 2 31.9 mm (MNHN-Na 13165. drawn).

Diagnosis. — Detailed descriptions of this species can be found in CHACE (1939), MONTERROSA (1988) and

Paulmier (1993, under the name P. polyacanthomerus).

6 28.9 mm [transferred to

I 9 27.7 mm (MNHN-Na

Coloration. — Body light pink, with a mosaic patterns of light red markings on carapace and abdomen.

Eyes dark brown. Pereiopods lighter in colour. Pleopods reddish. Eggs blue.

SIZE. — Largest male and largest female 30.2 mm and 31.9 mm cl. respectively. Smallest ovigerous female

22.5 mm cl.

Remarks. — This species has been largely neglected since its original description in 1939. Only recently has

it been redescribed and illustrated in detail by MONTERROSA (1988), PAULMIER [1993, under the name

P. polyacanthomerus (non Pequegnai. 1970)] and POUPIN (1994). It is extremely similar to P. wiliiamsi Forest.

1964, but almost none of the previous authors discussed their relationships. The present study shows that they do

represent distinct species and even occur sympatrically in the Antilles and French Polynesia (see "Remarks under

P wiliiamsi). The colour photograph of P. macropoda (non Chace. 1939) provided by Poupin ( 994, pi. le)

actually refers to P. wiliiamsi, while those of P. polyacanthomerus (non Pequegnai, 1970) given by PAULMIER

(1993, pis 19-20) represent true P. macropoda. It is not known whether the report ot P. wiliiamsi by Lemaitre

(1984) from the Bahamas and Gulf of Mexico truly refers to that species, since no details about the material was

provided.

206

T.-Y. CHAN & A. CROSNIER

Fig. 11. — Plesionika macropoda Chace. 1939. a-c, 6 23.1 mm (MNHN-Na 13113), Antilles, "Polka", stn CP3, 400 m:

a. lateral view of carapace; b, dorsal rostral and post-rostral teeth enlarged; c. dorsal view of abdominal somite 111.

— d. 6 23.1 mm (MNHN-Na 13112), French Polynesia, Tuamotu Islands, Gambier, stn 311. 470 m: lateral view of

carapace. — e-f, ovig. 9 31.9 mm (MNHN-Na 13165), French Polynesia, " Marara ", stn 405, 310 m: e, : lateral view

of carapace; f, dorsal rostral and post-rostral teeth enlarged. — g, 9 19.1 mm (MNHN-Na 12526), French Polynesia,

" Marara ", stn 37, 22°00.0'S, 136o12.0’W, 470 m, dorsal rostral and post-rostral teeth enlarged. —

h, 9 27.7 mm (MNHN-Na 13114), French Polynesia, Society Islands, Huanine, stn 405, 310 m: dorsal view of

abdominal somite III.

Source :

PLESIONlfCX FROM FRENCH POLYNESIA

207

Fig. 12 a-d. — Plesionika macropoda Chace, 1939. a-b. 6 23.1 mm (MNHN-Na 131 13), Antilles, " Polka ", stn CP3,

400 m: a. lateral view of abdomen and tailfan; b, dorsal view of tailfan. — c, 6 21.9 mm (MNHN-Na 13099),

Antilles, La Guadeloupe, cote sous le Vent, 400-450 m: lateral view of posterior part of abdomen. — d, <3 18,1 mm

(MNHN-Na 12847), ibidem: dorsal view of telson. — e, ovig. 9 31.9 mm (MNHN-Na 13165), French Polynesia,

"Marara", stn 405, 310 m: lateral view of fifth abdominal segment. — f-g, 6 23.1 mm (MNHN-Na 13112), French

Polynesia, Marara , sin 311. 470 m: f, lateral view of fifth abdominal segment; g, dorsal view of telson.

208

T.-Y. CHAN & A. CROSNIER

Fig 13 — Plesionika macropoda Chace, 1939. Dactylus and propodus of pereiopod 111. 2 27.7 mm (MNHN-Na 13114).

French Polynesia. Society Islands. Huanine, stn 405, 310 m: a-b. left side; c-d. right side. Figures a. c at the same

scales, the same for figures b. d.

Paulmier (1993) confused P. macropoda with P. polyacanthomerus Pequegnat, 1970, but an examination

of 16 topotypic specimens (including 8 paratypes: 68-A-13-4, 67-A-5-6B, 67-A-5-9A, 68-A-7-17B; all trans¬

ferred to MNHN from Texas A&M Oceanography Collection) of P. polyacanthomerus shows that they all bear

distinct lateral carinae on the carapace and are, in fact, most similar to P. carinata Holthuis, 1951 from the

western Atlantic (the exact relationship between these two species is not very clear), rather than to P. macropoda .

The general appearance of P. polyacanthomerus is generally similar to the "P. martia (A. Milne Edwards,

1883)" group in having the body more fragile and the rostral crest less pronounced, while P. macropoda , as well as

p. mltiamsi , are closer to the "P. edwardsii (Brandt, 1851)" group in having a robust body and a distinct rostral

crest. PEQUEGNAT (1970) mentioned that one of her specimens was quite different from the others in having only

9 widely spaced ventral rostral teeth. This specimen was not at our disposal, but it probably belongs to

P. macropoda , rather than P. polyacanthomerus. Similarly, it is not known whether the material reported by

SPRINGER and BULLIS (1956) from the Gulf of Mexico represents P. macropoda or P. polyacanthomerus.

The present specimens from French Polynesia and Loyalty Islands are very similar to the topotypic material

from the Antilles (table 3) but they show several slight differences concerning:

1) the number and the shape of the dorsal rostral and post-rostral teeth. In specimens from the Antilles, if only

the teeth entirely behind the orbit are counted, there are 4 (sometimes 3) post-rostral teeth, 2 or 3 (seldom 1,

exceptionally 4) of them being movable and 1 or 2 partly divided only at their base; in specimens from South

Pacific there are 2 or 3 (sometimes 1, once 4) post-rostral teeth, none or 1 (once 2 and once 3) being movable and

1 or 2 being partly divided at their base. On an other hand the unmovable teeth are less spiny and more squarish in

the specimens from South Pacific (fig. 1 1 b and f-g).

2) the posteroventral angle of abdominal pleuron V is usually produced in a sharp denticle in the Antillean

specimens; in the South Pacific specimens the angle is acute but seldom with a denticle (fig. 12 a, c and e-f).

3) the telson generally bears four pairs of dorsolateral spines in the Antillean material, compared to three pairs

in the South Pacific specimens (fig. 12 b, d, g).

4) the number of posterior spinules (including the accessory spine) on the dactyli of the posterior pcreiopods is

usually 2-3 in the South Pacific material but 3-4 (with basal spinules often small and very slender) in the

Antillean specimens.

5) the colour pattern of the abdomen. The mosaic stripes on the abdomen are somewhat broader in the Antillean

material (fig. 31) than in the South Pacific specimens (fig. 30).

So it would appear that the French Polynesian and Loyalty Islands specimens are different from the Antillean

ones but in the present state of our knowledge on the variations in the Plesionika species with a wide geographical

range, and taking into account the variations occasionally exhibited in a same individual (e.g. fig. 13) and those

occuring in the different populations of P. williamsi, it seems reasonable not to separate the South Pacific material

from the Antillean one, at least for the time being.

Distribution. — Western Atlantic and southern Pacific: Caribbean and in the south Pacific, only known with

certainty from the Loyalty Islands and French Polynesia, at depths of 260-600 m.

Source :

PLESIONIKA FROM FRENCH POLYNESIA

209

Plesionika williamsi Forest, 1964

Figs 14-15, 32-33

Plesionika williamsi Forest, 1964: 620, figs 1-4 [type-locality: Gulf of Guinea], — Crosnif.r & Forest, 1973: 211,

fig. 65e. — Poupin, 1996, pi, 5f.

[?] Plesionika williamsi - Lemaitre, 1984: 435. — Burukovsky, 1995: 124.

Plesionika aff. williamsi - Crosnier, 1986a: 373, fig. 4a-g. — Poupin el al, 1990: 16 (in part).

Plesionika crosnieri Burukovsky, 1992: 145, figs 1-4 [type-locality: Southeast Pacific seamount].

Plesionika alaini Burukovsky. 1993: 18 [nom. nov, for P. crosnieri Burukovsky. 1992],

Plesionika macropoda - Poupin, 1994: 25 (in part), pi, le [non Chace, 1939],

Material examined. — West Africa. Gulf of Guinea Guinean Trawling Survey: stn 28-8. 4°16'N,

2°09.5'W, 380-400 m. mud, trawling, 4.10.1963: 3 ex. [MNHN-Na 14376], — Stn 27-8. 4°39’N, 2°46'W, 300-400 m]

mud. 6.10.1963: 1 2 25.3 mm. holotype (MNHN-Na 2129) and 28 ex. (MNHN-Na 8052, 8039, 8054).

Senegal. "Pr. Bogucki": stn 11, 14°24'N, 17°34'W, trawling. 416 m. 26.05.1979: 2 ovig. 2 30.7 and 31.3 mm

(MNHN Na 3845). — "Laurent Amaro": sin 2, 14°21.7'N, 17°35.2‘W, 400-420 m. trawling. 12 6 1981 3 ex 30 7-

31.1 mm [MNHN-Na 4088],

Canary Islands. Tenerife. I ex. (MNHN-Na 2126).

Madeira. Balgim: stn CP 156, 36°20'N, 7°53'W, 1130-1140 m. 18.06.1984: 1 ex. 30.0 mm (MNHN-Nal4374)

Antilles. "Polka"-, stn Y 1 1 , 16°00.01'N, 6r45.82'W, 500 m, March 1993: 1 ovig. 2 29.5 mm (MNHN-Na 13106

drawn). — Stn EP1. 15°50.93'N, 61°41.39'W, 560 m. May 1993: 2 6 20.0 and 24.3 mm.

Madagascar. "Vauban": trawling 38, 12°50.0'S, 48°09.I'E, 580-585 m. 14,09.1972: 7 ex. (MNHN-Na 14375)

La Reunion. 350-500 m, 2.02.1974: 1 8 26.6 mm.

Seychelles. Cepros: stn 1.3. 4°08'S. 56°11.3’E. 580-500 m, 20.10.1987: 2 ex. 24.0 and 24.9 mm (MNHN-Na

14373). — Stn 3.16, 4°34.7'S. 56°25.6'E, 410-390 m, 22.10.1987: 2 ex. 25.6 and 25.7 mm (MNHN-Na 14377).

Kiribati. South Pacific Commission :: 400 m, April 1987: 2 ex. 18.0 and 30.5 mm (MNHN-Na 14378).

French Polynesia. SMSRB (J. Poupin and B. Richer DE Forges coll.): Marquesas Islands. Eiao Stn ->79

8°10.0'S, 139°42.0'W, 400 m. 18.08.1990: 3 2 16.1-27.4 mm. 1 ovig. 2 26.7 mm. — Sin 280. 8o10.3'S;

139°42,6'W, 600 m. 18.08.1990: 2 8 23.8 and 26.4 mm. — Tahuata. 430 m, 13.09.1987: I 2 22.8 mm. — Stn 299,

9°54.5'S, 139°08.2'W, 350 m. 1.09.1990: 1 ovig. 2 30.4 mm (MNHN-Na 13019, drawn). — Ua Pou. Stn 57. 9°23.0'S,

140°09.0'W, 590 m, 10.09.1987: 1 8 14.3 mm. — Stns 59-60, 9°23.0’S. I40°09.0'W, 450-520 m. 26.01.1988: II 8

14.3-23.3 mm.

Society Islands, lies sous le Vent. Bellingshausen. Stn 96. 15°48.0'S. 154°32.0'W. 580 m. 21.07.

1988: 3 8 16.7-17.2 mm, 2 2 13.6 and 21.2 mm [transferred to USNM], — Tahiti (Port Phaeton). October 1978' 2

<3 23.5 and 25.6 mm (MNHN-Na 7720, Na 7721).

Tuamatu Islands. Makemo. Stn 111, 16°40.0'S, I43°39.0'W, 500 m, 12.11.1988: I 8 18.9 mm 1 2

16.7 mm. — Fangataufa. Stn 235, 22°I4.6'S, 138°46.4'W, 480 m, 22.05.1990: 1 2 17.7 mm. — Marutea Sud.

Stn 402, 21 °30.8'S. 135°38.4'W, trapping, 380 m, 25.03.1991: I 8 30.7 mm. — Stn 473, 21°47.0'S, 138°55.5'W,

465 m, 10.04.1995: 1 8 25.4 mm. — Mururoa. 2I°52.7’S, 1 38°53.I'W, 450 m. 2.10.1996: 1 8 25.4 mm. 5 2 23.9-

31.8 mm, 1 ovig. 2 31.3 mm (transferred to NTOU).

Tubuai Islands. Stn 338, 22°28.6'S, I5I°22.4'W, 500 m. 27.11.1990: 1 8 25.8 mm [transferred to USNM].

Diagnosis. — Detailed descriptions of this species can be found in Forest (1964), Crosnier and Forest

(1973) and Crosnier (1986a).

Coloration. — Body light pink. Carapace and tip of rostrum reddish. Abdomen covered with broad red bands.

Eyes dark brown. Distal segments of pereiopods somewhat whitish. Eggs bluish.

Size. Largest male and largest female 30.7 and 33.4 mm cl, respectively. Smallest ovigerous female

26.7 mm cl.

Remarks. — The present form has been reported by Crosnier (1986a) from French Polynesia under the name

P. aff. williamsi. However, the differences mentioned by Crosnier (1986a) between the French Polynesian and

the topotypic material arc found to be rather variable in the specimens available for the present study. Nevertheless,

there appears to be a link between the shape of the posterior margin of the abdominal somite Ill and the length of

rostrum, as well as the number of rostral teeth. For those 39 specimens with the posterior margin of the

abdominal tergite III produced into a sharp angle (in 13 specimens) or even a small spine (in 26 specimens, fig.

210

T.-Y. CHAN & A. CROSNIER

14b), the rostrum is relatively shorter (1.4-2, avg. 1.7 carapace length; with small individuals generally having a

relatively longer rostrum) and has more dorsal teeth (9-1 1, 8 in only one specimen) but fewer ventral teeth (7-12,

fig. 14a). On the other hand, those specimens with the posterior margin of abdominal tergite III bluntly angular or

rounded (fig. 1 lh) generally have a longer rostrum (1. 8-2.1, avg. 1.9 carapace length) with fewer dorsal teeth (6-9,

1 1 in only one specimen) but more ventral teeth (9-14, fig. 1 Id-e). Furthermore, colour photographs of fresh

specimens from French Polynesia showed that the above morphological differences are always accompanied by the

presence or absence of red bands on the abdomen. Colour photographs of 10 red banded specimens are available

(fig. 32). The specimens used for six of them (i.e. stn 235, 299, 473 and Bclingshausen, Ua Pou. Mururoa) are

deposited at the Paris Museum; they all have pointed abdominal tergite III, as well as a shorter rostrum. On the

other hand, the 4 colour photographs of unbanded specimens (fig. 30), from stn 311, 383, 405 and 476, all

correspond to individuals with rounded abdominal tergite III and a longer rostrum. It is, therefore, highly probable

that two distinct species are present in the French Polynesian material.

Numbers in brackets represent the average values.

Table 3. — Comparisons of the rostrum between Plesionika macropoda Chace, 1939 (i.e. specimens with abdominal

somite III rounded) and P. williamsi Forest. 1964 (i.e. specimens with abdominal somite III pointed) from different

localities.

An examination of 32 specimens from the type series (including the holotype) of P. williamsi from the Gulf of

Guinea shows that they all have pointed abdominal somites III (also varying from a sharp angle to bearing a

distinct denticle, figs 14e-f). However, the rostrum of the types are considerably shorter (1.1- 1.5, avg. 1.3 carapace

length) than those of the French Polynesian material with a pointed abdominal tergite III, thought the number of

rostral teeth are very similar. When a range of material from western Africa to the Indian Ocean and southern

Pacific is compared, the differences in the proportional length of the rostrum exhibited in the various populations

seem to represent natural variations of a single species with a very wide geographical distribution (Table 3).

Although the specimens of the Indo-Pacific material with a longer rostrum show some resemblances with

P. carinata Holthuis, 1951 (with a type locality close to that of P. williamsi ), the size of P. carinata is much

smaller (usually less than 20 mm cl) and it always bears a distinct lateral carina on the carapace [specimens from

West Africa reported in CROSNIER and FOREST (1973) deposited at MNHN were examined]. In P. williamsi , only

rudimentary lateral carina is sometimes present on the carapace and mainly in very large specimens.

Slight differences in the number of spines on the telson, the shape of the dactyli of the posterior pereiopods and

abdominal pleuron V are also observed in the various populations. Although the telson has generally been

described as having four pairs of dorsolateral spines in P. williamsi (Forest, 1964; CROSNIER & FOREST, 1973),

sometimes three or five pairs of dorsolateral spines are present and the number of spines on the left and right sides

Source :

PLESIONIKA FROM FRENCH POLYNESIA

21 1

Fig. 14. — Plesionika williamsi Forest, 1964. a-b, ovig. 9 30.4 mm (MNHN-Na 13109), French Polynesia, Marquesas

Islands, Tahuato, stn 299, 350 m: a, lateral view of carapace ; b, dorsal view of abdominal somite 111. — c-d, ovig. 9

29.5 mm (MNHN-Na 13106), Antilles, " Polka ", stn Y 1 1 . 560 m: c, lateral view of carapace; d, dorsal view of

abdominal somite III. — e, 9 25.3 mm, holotype (MNHN-Na 2129), Gulf of Guinea, Guinean Trawling Survey,

stn 27-8, 300-400 m: dorsal view of abdominal somite III. — f. 9 29.0 mm (MNHN-Na 14376 part). Gulf of Guinea,

Guinean Trawling Survey, stn 28-8, 380-400 m: dorsal view of abdominal somite III.

are often unequal (see Crosnier, 1986a, figs 4d, g). Nevertheless, the South Pacific material usually has only

three pairs of dorsolateral spines on the telson. The number of spinules on the posterior border (including the

accessory spine) of the dactyli of posterior pereiopods varies from 1-4 (but mostly 2-3) and such variation can

sometimes occur in the same specimen (fig. 15). Similarly, the posterolateral angle of abdominal pleuron V may

be round to sharp, or produced into a distinct denticle, and the shape of the pleuron is sometimes different between

the left and right sides in the same individual. Therefore, the differences suggested by CROSNIER (1986a) for the

French Polynesian material appear to be unimportant.

212

T.-Y. CHAN & A. CROSNIER

Fig. 15. — Plesionika williamsi Forest, 1964. Dactylus and propodus of pereiopod Ill.

a. 9 25.3mm. holotype (MNHN-Na 2129). Gulf of Guinea, Guinean Trawling Survey, stn 27-8, 300-400 m: right side. —

b-c, ovig. 9 31.1 mm (MNHN-Na 4088 part). Senegal. "Laurent Amaro", stn 2, 400-420 m: b. left side; c, right side.

— d, ovig. 9 30.4 mm (MNHN-Na 13109), French Polynesia, Marquesas Islands, Tahuato, sin 299, 350 m: right side.

_ e. ovig. 9 29.5 mm (MNHN-Na 13106), Antilles, "Polka", stn Y 11, 560 m: left side.

On the other hand, P. macropodci Chace, 1939, described from the western Atlantic, is actually similar to

P. williamsi , as said above. The types of P. macropoda are described by CHACE (1939) as having a round

abdominal tergite III and a very long rostrum (nearly twice as long as carapace). The number of rostral teeth of the

types (7-8 dorsal teeth and 1 1 ventral teeth) also agrees well with those of the French Polynesian material with a

rounded abdominal somite III. The colour photographs of fresh material from the Antilles provided by PAULMIER

(1993, pis 19-20. under the name P. polyacanthomerus) and POUPIN (1994, pi. le) show a very similar pattern of

banded and unhanded forms to that found in specimens from French Polynesia. A re-examination of Poupin's

(1994) material revealed that the banded specimen (fig. 33) used for his illustration (POUPIN, 1994, pi. le;

specimen from Stn EP1, not CPI as indicated in the caption) also has a pointed abdominal tergite III and a shorter

rostrum (fig. 14c-d), while another specimen (from Stn 21 1, fig. 31) with colour photograph showing an unhanded

abdomen has the typical characteristics of P . macropoda (fig. I la-c). Therefore, it appears that two distinct forms

are also present in the Antilles.

After comparing many specimens and colour photographs of fresh material from various localities in both the

Indo-Pacific and Atlantic, we conclude that the name P. williamsi can probably be applied to the banded form with

a pointed abdominal tergite III and shorter rostrum. For the unbanded form with a rounded abdominal tergite III and

longer rostrum, the name P. macropoda seems to be appropriate for the time being, even if the differences observed

between the Atlantic and the South Pacific specimens leave open the question of distinguishing the specimens of

these two areas by two different specific names (see p. 208). Both species have a very wide geographical distri¬

bution and sometimes occur sympatrically (a situation which seems to be rather common in Plesionika for very

similar species). If the above conclusion is correct, P. crosnieri Burukovsky, 1992 (junior homonym of

P. crosnieri Chan & Yu, 1991) recently described from the southeastern Pacific, should be treated as a junior

Source :

PLESIONIKA FROM FRENCH POLYNESIA

213

synonym of P. williamsi, since they both have a shorter rostrum (about 1.5 times as long as carapace) and the

posterior margin of the abdominal tergite III in the form of a right angle instead of rounded. P. williamsi appears

to occur mainly in the eastern Atlantic and the Indo-Pacific, while P. macropoda is dominant in the western

Atlantic. This, together with the fact that only P. williamsi is found in the eastern Atlantic and the Indian Ocean,

may suggest that P. macropoda has only recently invaded the South Pacific from the Caribbean, through the

Panama Canal, and vice versa for P. williamsi, from the Pacific to the Antilles, though more extensive data are

needed for a more careful interpretation on the zoogeographical relationships of these two, closely related species.

DISTRIBUTION. — World-wide tropical and subtropical seas, but only known with certainty in the Atlantic

from the French Antilles to western Africa (Gulf of Guinea, Senegal, Canary and Madeira Islands); and in the Indo-

Pacific from Madagascar, La Reunion, Seychelles, Kiribati and French Polynesia to southeastern Pacific, near

Chile. At depths of 300-1 140 m, mostly less than 700 m.

Plesionika martia (A. Milne Edwards, 1883)

Pandalus martins A. Milne-Ed wards, 1883, pi. 21 [type-locality: Mediterranean].

Plesionika martia - Crosnier & Forest, 1973: 212, figs 63d, 64e, 66.

MATERIAL EXAMINED. — French Polynesia. SMSRB (J. Poupin coll.): Marquesas Islands . Nuku Hiva.

Stn 284, 9°06.0’S, 140°07.6’W, 650 m, 27.084990: 4 6 16.8-27.3 mm. 1 9 16.7 mm. — Ua Pou. Stn 57, 9°23.0'S,

140°09.0'W, 590 m, 10.09.1987: 1 9 22.2 mm [transferred to USNM].

Diagnosis. — The general appearance, as well as possible distribution and size of this species, can be found in

Crosnier and Forest (1973), though much confusions probably exists concerning the species of the "P. martia "

group.

Remarks. — The "P. martia " group is generally considered to be a difficult species complex (e.g. Crosnier

& Forest, 1973; Chace, 1985; Hanamura & Takeda, 1987; Kensley et al. , 1987). Since only 6 French

Polynesian specimens are available in this study, unaccompanied by any colour note, they are tentatively assigned

to this species according to the preliminary results obtained from a more detailed study of this group based mainly

on Taiwanese material (Chan & Yu, in prep.). The limited French Polynesian material has the post-rostral carina

sharp but not distinctly laminate, upper lobe of orbital margin nearly vertical, dorsal border of rostrum without any

tooth anterior to scaphocerite, basiccrite spine long and distinctly overreaching proximal end of lateral margin of

scaphocerite. Only one specimen (19.7 mm cl) still possesses a complete pereiopod IV and its propodus is longer

than the carapace length.

Plesionika semilaevis Bate, 1888

Plesionika semilaevis Bate. 1888: 644, pi. 113 fig. 3 (in part) [type-locality: the Philippines]. — CHACE, 1985: 113,

figs 51-54.

Material EXAMINED. — French Polynesia. SMSRB (B. RICHER DE FORGES coll.): Tubuai Islands.

Tubuai. 800 m, 14.05.1979: 2 6 16.1 and 17.8 mm.

Diagnosis. — The general characters and other information such as possible distribution and size of this

species can be referred to Chace (1985), though it should be borne in mind that the taxonomy of the "P. martia "

group is probably confused.

Remarks. — Only two French Polynesian specimens are available in this study and they lack a colour record.

Similar to the situation of the material identified as P. martia (A. Milne Edwards, 1883), these two specimens are

here tentatively assigned to P. semilaevis according to the preliminary results of a more detailed study of the

214

T.-Y. CHAN & A. CROSNIER

"P. mania" group (Chan & Yu, in prep.). The French Polynesian material has the eye kidney shaped, post-rostral

carina high and laminate; upper lobe of orbital margin strongly recurved backwards; basicerite spine very long and

far exceeding proximal end of lateral margin of scaphocerite. The rostrum is broken and all the posterior pereiopods

are incomplete in these two specimens.

Plesionika erythrocyclus sp. nov.

Figs 16, 34-35

Plesionika rostricrescentis - King, 1984: 178, fig. Pr. [non Bate, 1888].

Plesionika sp. nov. 1 - POUPIN, 1996, pi. 4a.

Material EXAMINED. — French Polynesia. SMSRB (J.-L. CARSIN and J. POUPIN coll.): Tuamotu Islands .

Mururoa. 350-600 m, 1984: 1 ovig. 9 10.7 mm (MNHN-Na 13123). — Stn 499, 21°47.6’S, 138°55.7'W, 200 m,

5.05.1996: 1 ovig. 9 11.6 mm (transferred to NTOU).

Tubuai Islands. Rurutu. Stn 356, 22°30.3'S, 151°21.7'W, 260 m, 10.12.1990: 1 ovig. 9 10.6 mm (transferred

to USNM).

Taiwan. North-eastern coast, Su-Aou, I-Lan County, commercial trawler, about 300 m, 22.05.1990: 1 9

10.3 mm (NTOU-P 1990-5-22).

Types. — The ovigerous female (10.7 mm cl) (MNHN-Na 13123) from Mururoa is the holotype. All the other

specimens from Polynesia are paratypes.

Description. — Body somewhat robust in appearance. Rostrum with lateral carina distinct on basal half and

not continuous with orbital margin, curving downward at base, but abruptly upturned after passing the eye, more

or less (0.9-1. 1) as long as carapace and just overreaching scaphocerite; dorsal border bearing only 2 small apical

teeth and 1-2 fixed basal teeth above eye; ventral border with 7-9 evenly distributed teeth between level of eye and

tip; post-rostral crest low and armed with 5-6 movable teeth (anteriormost one occasionally situated just above

orbital margin). Orbital margin regularly concave, with lower lobe somewhat truncate. Eye subspherical and

bearing distinct ocellus. Antennal spine large and elongate. Pterygostomian spine sharply pointed but small.

Styloceritc tapered anteriorly and extending to around basal or middle part of distal segment of antennular peduncle.

Scaphocerite 3. 9-4. 6 times as long as broad, with distolateral tooth more or less reaching distal margin of lamella.

Basicerite spine moderately long and extending to proximal end of outer margin of scaphocerite.

Maxilliped III bearing well developed epipod and a long exopod, with penultimate segment 0.6-0. 7 times as

long as distal segment; overreaching scaphocerite by 1/3- 1/2 of distal segment. Anterior four pereiopods all bearing

well developed epipods. Pereiopod I overreaching scaphocerite by I/2-4/5 propodus and entire dactylus. Pereiopod II

bearing 20-22 carpal articles on the right side and 76-83 carpal articles on the left side, longer one exceeding

scaphocerite by slightly more than lengths of dactylus, propodus and carpus. Pereiopod III exceeding scaphocerite

by 2/3-3/4 of propodus and entire dactylus; dactylus conical, short, 0.14-0.15 times as long as propodus, posterior

margin bearing 2-5 spinules, accessory spine about half to almost as long as, and abutting with, terminal spine.

Pereiopods IV and V similar to pereiopod III, with pereiopod IV overreaching scaphocerite by 1/3- 1/2 of propodus

and entire dactylus, and pereiopod V by slightly more than length of dactylus.

Abdominal somite III slightly arched dorsally, with posterior margin convex. Both abdominal pleura IV and V

each bearing a distinct posteroventral denticle. Telson 1.3- 1.5 times longer than abdominal somite VI, with three

pairs of dorsolateral and three pairs of terminal spines. Eggs suboval, about 0.4 mm in diameter.

Coloration. — Body covered with orange-red tints and a few irregular yellowish stripes, each abdominal

somite having a transverse white band near posterior border, somite I also bearing a broad transverse red band (the

yellow stripes and white bands quickly fade when the specimens are less fresh). Eyes black. Dorsal surface of

abdominal tergite III bearing a pair of large, white-margined, red circular spots, while that of tergite VI has a pair

of smaller red spots. Tip of rostrum reddish. Thoracic appendages, antennal and antennular flagella with red bands.

Exopods of uropods with two pairs of red dots along outer margin. Eggs a rather light orange.

PLESIONIKA FROM FRENCH POLYNESIA

215

Size. — Largest female 1 1.6 mm cl. Smallest ovigerous female 10.6 mm cl. Males unknown.

Tuamotu Islands, Mururoa, 350-600 m: a. lateral view of carapace; b, eye; c. lateral view of abdomen and tailfan;

d, dorsal view of telson; e, dorsal view of tip of telson; f, left pereiopod III; g, left dactyl of pereiopod III.

Figures a-d and f at the same scale.

216

T.-Y. CHAN & A. CROSNIER

REMARKS. — The present species is very close to P. rostricrescentis (Bate, 1888), described from New Guinea.

It differs from Bate's (1888) original description and figures in some important aspects. Apart from having a

much larger size (i.e. 18 mm cl), the type of P. rostricrescentis has the rostrum much longer (1.5 times as long as

the carapace and far exceeding scaphocerite) and with a very high basal crest (Bate, 1888: 654, table; pi. I 14,

fig. I, but in the text Bate mentioned that the rostrum is as long as the carapace). Furthermore, the stylocerite is

distinctly longer and reaches the distal end of the antennular peduncle in Bate’s (1888) New Guinean specimen.

The number of ventral rostral teeth in the type of P. rostricrescentis is 15, which is much higher than in the

present form (i.e. 7-9 teeth). However, the " Albatross " material from the Philippines and Indonesia identified as

P. rostricrescentis by Chace (1985) has only 9 or 10 ventral rostral teeth. But we should mention here that at

least two more forms belonging to the "P. rostricrescentis" group are present in the collections of MNHN and

NTOU from New Caledonia and Taiwan; CHACE’s (1985) description of the " Albatross " material is actually very

similar to one of the forms of the "P. rostricrescentis " group found in both New Caledonia and Taiwan, which

probably represents the true P. rostricrescentis. These probably true P. rostricrescentis from New Caledonia and

Taiwan have a very different coloration, bearing a large red circle on the lateral surface of the abdominal somite I,

but without any red spots on abdominal somite III. On the other hand, the material from Fiji and Tonga reported

by King (1984, fig. Pr) as "P. rostricrescentis" has the abdominal somite III bearing a large circular spot. Since

these specimens from Fiji and Tonga also have a short rostrum and a low post-rostral crest, they probably belong

to the same species as the present species, instead of P. rostricrescentis. Moreover, photographs of freshly

collected specimens belonging to the "P. rostricrescentis" group from New Caledonia also show the characteristic

red spot on the abdominal somite III. This indicates that the present form is probably also distributed in New

Caledonia. For the specimen identified as "P. rostricrescentis " by YOKOYA (1933) from southern Japan, no size or

description were given. In view of the confused taxonomy of the species of the genus Plesionika , particularly in

previous works, only a re-examination of YOKOYa's (1933) specimen can clarify the identity of the Japanese

material.

Although the true identity of P. rostricrescentis is still uncertain, several characters of the present form (e.g.

smaller size, low rostral crest, shorter rostrum, stylocerite, etc.) arc significantly different from typical

P. rostricrescentis (as well as related species, such as P. grahami Kensley, Tranter & Griffin, 1987), indicating

that it deserves specific status. The specimens from French Polynesia and Taiwan are generally very similar

(including coloration), except that the single female from Taiwan has the dactylus of pereiopod III bearing slightly

more posterior spinules and with the accessory spine shorter. Nevertheless, more specimens from various localities

may show that such differences in the dactylus are geographical variations, as in some of the other species of the

genus.

Etymology. — The Latin e/ythrocyclus refers to the conspicuous red spots on abdominal somite III in this

species.

Distribution. — Known from French Polynesia and Taiwan, probably also distributed in New Caledonia,

Fiji and Tonga, at depths of 200-600 m.

Plesionika sindoi (Rathbun, 1906)

Figs 17, 36-37

Pcindalus sindoi Rathbun, 1906: 915, pi. 21, fig. 4 [type-locality: Hawaii].

Plesionika Sindoi - DE Man, 1920: 126, pi. 11, fig. 27-27d, pi. 12, fig. 27c.

Plesionika ocellus - Chace, 1985: 90, fig. 40. — Toriyama el al. , 1990: 18, pi. 3a [non Bate, 1888].

Plesionika aff. ocellus - Poupin el al ., 1990: 16 [non Bate, 1888].

Plesionika sindoi - POUPIN, 1996, pi. 5c.

Not Plesionika sindoi - Balss, 1925: 279, figs 49-52, pi. 26 [= P. indica De Man, 1917].

Material EXAMINED. — Hawaii. "Albatross": sin 3998, vicinity of Kauai Island. 430-417 m, 14.06.1902: 2 6

10.3 and 13.1 mm, syntypes (USNM).

Source :

PLESIONIKA FROM FRENCH POLYNESIA

217

French Polynesia. SMSRB (J. Poupin coll.): Marquesas Islands. Fatu Hiva. Stn 306. 10o3M'S,

138°39.4'W, 250 m, 4.09.1990: 5 ovig. 9 7.1-7.9 mm (2 ex. transferred to NTOU); 1 ovig. 9 7.3 mm (MNHN-Na

13111, drawn).

Society Islands lies sous le Vent ( Bora Bora). Stn 97, 16°28.0'S, 151°47.0'W, 480 m. 23.07.1988: 2

6 9.5-10.4 mm, 2 ovig. 9 1 1.7 mm (MNHN-Na 13110, drawn) and 14.2 mm. 3 9 9.5-12.0 mm.

Tuamolu Islands. Fangataufa. Stn 85, 22°13.0’S. 138°42.2'W, 780 m, 22.06.1988: 1 6 11.2 mm. — Stn

438, 22°12.28'S, 138°46.64'W, 410 m, 14.11.1994: 1 ovig. 9 13.0 mm. — Stn 488. 22°14.4'S, I38°46.7'W, 510 m,

25.04.1995: 1 9 15.5 mm. — Gambler. Stn 238, 23°01.rS. 134°59.0’W, 500 m, 27.5.1990: 2 6 8.6-8.7 mm,

4 ovig. 9 10.2-11.5 mm. — Makemo. Stn 68, I6°36.0'S, I43°33.0'W. 510 m, 4.06.1988: 1 ovig. 9 13.5 mm. —

Maria. Stn 72, 22°00.0’S, 136°17.0'W, 430 m, 10.06.1988: 2 ovig. 9 9. 2-9. 8 mm. — Stn 241. 22°00.9'S.

136°12.5'W, 380 m, 30.05.1990: 1 6 11.3 mm. 5 9 10.2-12.6 mm. — Mururoa. Stn 39, 2I°49.3'S, 138°46.5'W.

470 m, 2.12.1987: 2 ovig. 9 15.1-16.0 mm (MNHN-Nal2525). — Stn 457, 21°53.9’S, 139°01.3'W. 450 m.

25.11.1994: 1 9 10.7 mm. — Tuanake. Stn 253, 16°37.3,S, 144°13.3’W. 450 m, 5.06.1990: 1 6 14.1 mm. 7 ovig.

9 11.4-14.3 mm, 2 9 10.2-12.1 mm (transferred to USNM).

Tubuai Islands. Maria. Stn 422, 21°47.9'S, 156°43.8,W, 680 m. 7.08.1991: 1 6 10.1 mm. — Raevavae.

Stn 100, 23°55.0'S, 147°40.0'W, 530 m, 24.08.1988: 2 6 11.5-12.9 mm, 1 9 11.8 mm. — Sms 105-106, 23°19.0'S,

142°22.0’W, 460-550 m. 31.08.1988: 3 6 13.9-14.7 mm. — Rurutu. Stns 107-108. 22°27.0'S, I51°23.0'W. 570 m,

3.09.1988: 6 6 10.8-12.6 mm, 1 ovig. 9 14.5 mm, 1 9 12.2 mm (2 ex. transferred to NTOU). — Sin 146, 22°27.8'S,

151°22.8'W, 580 m, 9.03.1989: 1 9 12.3 mm.

Diagnosis. — Detailed descriptions of this species can be found in DE Man (1920) and Chace (1985, under

the name Plesionika ocellus).

Coloration. — Body somewhat translucent, with some longitudinal red stripes (sometimes accompanied by

narrow white lines) running along entire body. Eyes dark brown. Anterior part of rostrum and antennular flagella

with alternating red and white bands. Antennal flagella orange-pink. Pereiopods pink to red. Posterior part of

abdominal somite VI reddish. Telson also with posterior half reddish, but basal part whitish. Eggs bluish.

Sizes. — The largest male and female in the present French Polynesian material are 14.7 and 16.0 mm cl

respectively. The smallest ovigcrous female is 7.1 mm cl.

Remarks. — The material of the "P. ocellus (sensu Chace, 1985)" group in French Polynesia appears to

have at least two forms. One form has red longitudinal stripes on the body and banded antennular flagella (fig. 36)

while the other form lacks bands or stripes on the antennules and body (fig. 38). Furthermore, the non-striped form

is rather small and always has long rostrum (average 1.9 times carapace length) and short abdominal somite VI

(average 2 times as long as height and 0.83 times as long as telson). The rostrum of the striped form is shorter

(1.1 -1.7, avg. 1.4 times as long as carapace), but its abdominal somite VI is longer (2. 1-2.6, avg. 2.3 times as

long as height and 0.9- 1.1, avg. 1.0 times as long as telson). All the French Polynesian specimens have the

dactylus of the posterior pereiopods very short, that of pereiopod III being 0.1-0.16 and 0.08-0.13 times as long as

the propodus in the striped and non-striped forms, respectively. Thus, they both differ from P. fimbriata Chace,

1985, which has long dactyli [one paratype of P. fimbriata from " Albatross " Stn 5391 (NTOU, USNM 205221 in

exchange) as well as many Taiwanese specimens (NTOU) of this species were examined).

The rostrum of the red-striped form is rather similar to that in the figure of P. ocellus provided by Bate

(1888). However, Bate's (1888) figure also shows a short abdominal somite VI. The types of P. ocellus (one

male 15.3 mm cl and one ovigerous female 18.5 mm cl, NHM 1888: 22) were re-examined. Both syntypes have

the right part of the carapace dissected and the thoracic appendages, as well as the rostrum, broken. Nevertheless,

the anterior part of the rostrum of the male and two pereiopods III with complete propodus and dactylus are present

in the jar. No pereiopod II was found, though Chace (1985) mentioned that B. Kensley had examined two of

them from the syntypes. The figure of P. ocellus provided by Bate (1888) is generally quite accurate: both

syntypes bear two small movable post-rostral teeth on the carapace, the dactylus of the pereiopod III is about 1/5

the length of the propodus, and the telson is almost 1.5 times longer than abdominal somite VI. Surprisingly, a

distinct posteroventral denticle is present on the abdominal pleuron IV in both syntypes. Therefore, the

characteristics of the types of P. ocellus conform well to P. chacei described by Hayashi (1986) from Japan [two

218

T.-Y. CHAN & A. CROSNIER

specimens of this species from Taiwan (NTOU) were compared). Thus, P. ocellus (Bate, 1888) must be

considered as a senior synonym of P. chacei Hayashi, 1986 (new synonymy).

Fig. I7 . — Plesionika sindoi (Raihbun, 1906). a. c, ovig. $ 11.7 mm (MNHN-Na 13110), French Polynesia, Society

Islands, Bora Bora, st. 97. 480 m: a, lateral view of carapace; c, lateral view of posterior abdomen and tailfan. —

b, d, ovig, 9 7.3 mm (MNHN-Na 13111), French Polynesia, Marquesas Islands, Fatu Hiva, stn 306, 250 m:

b, lateral view of carapace; d, lateral view of posterior part of abdomen and tailfan.

With respect to RATHBUN's (1906) P. sindoi, an examination of two male syntypes from Hawaii (USNM)

shows that they both lack posteroventral denticle on the abdominal pleuron IV and generally are similar to

CHACE's (1985) P. ocellus from the Philippines [9 ex., 8.3-12.9 mm cl, "Albatross" Stn 5519, 8°47'N,

123 31 15 E, 333 m, 9.8.1909, were examined (NTOU, USNM 221351 in exchange)] as well as to the Indonesian

material reported by DE Man (1920). Thus, the name P. sindoi should be revived and Chace's (1985) Philippines

material actually belongs in this species. For RATHBUN's (1906) "? P. ocellus", an examination of specimens

from ‘he lot used for her illustration [RATHBUN, 1906, pi. 21, fig. 1; "Albatross" Stn 3858. Hawaii. Pailolo

Channel, Mokuhooniki Islet, 35°N, 8°0I'E, 234-252 m, 9.04.1902, 2 males 14.0-14.7 mm cl, I ovig. female

1 1.3 mm cl (NTOU, USNM 30495 in exchange)] shows that they all have a posteroventral denticle on abdominal

Source :

PLESIONIKA FROM FRENCH POLYNESIA

219

pleuron IV and belong to the true P. ocellus, instead of P. fimbriata as considered by Chace (1985). Generally,

the body of P. ocellus is more robust than in P. sindoi , while that of P. fimbriata is somewhat intermediate. The

rostrum of the smaller male syntype of P. sindoi (i.e. the specimen of pi. 21, fig. 4 in RATHBUN, 1906) is 1.48

times carapace length and has four movable post-rostral teeth. The rostrum of the larger male is broken, but three

movable post-rostral teeth are present on the carapace. As for the posterior three pereiopods, only the right

pereiopod IV is still attached to the body in the larger male and its dactylus is 0.09 times as long as the propodus.

Only a loose pereiopod V was found in the jar and the dactylus is 0.06 times as long as the propodus. The

abdominal somites VI of the two syntypes are 2.2 and 2.4 times as long as high and 0.96 times as long as the

telson. Therefore, the striped form of the French Polynesian material with relatively shorter rostrum, but longer

abdominal somite VI, Fits well with the characteristics of P. sindoi, while the non-striped form proves to be new

and is described below as P. protati sp. nov.

The intensity of the red longitudinal stripes on the body appears to be rather variable in the present species (the

coloration of the Japanese material was figured by Toriyama et al. (1990, pi. 3a, under the name P. ocellus). In

large individuals, the stripes are broad and deep in colour but in small specimens they arc somewhat pale.

Furthermore, the rostrums of small ovigerous females (particularly those from Stn 306, figs 17b. 37) are rather

short, and their eggs are somewhat greenish. The shape of the accessory spines on the dactyli of the posterior

pereiopods also appears to be variable in the present species. In the types (only dactylus of pereiopod IV still

present) and small individuals of the French Polynesian material, the accessory spine is about half as long as, and

abuts with, the terminal spine. However, in large individuals of the French Polynesian material the accessory spine

is shorter and well separated from the terminal spine, as in Chace's (1985, fig. 40g) Philippines specimens.

Since specimens with intermediate characteristics can be found, they are all identified as the same species for the

lime being. On the other hand, the report of the present species from the Nicobar Islands by BALSS (1925) clearly

represents P. indica De Man, 1917, whereas that by Calm AN (1939) from the Gulf of Aden is probably correct.

The exact identities of the specimens reported as P. ocellus by ALCOCK (1901 ) from the Andaman Sea, by Parisi

(1919) from Japan, and by HANAMURA and Takeda (1987) from northwest Australia arc unclear.

DISTRIBUTION. — West and South Pacific: known with certainty from Japan, South China Sea, the

Philippines, Indonesia, Hawaii and French Polynesia. Probably also present in the Indian Ocean (see "Remarks"

above). At depths of 150-800 m, mostly between 300-600 m.

Plesionika protati sp. nov.

Figs 18, 38

Plesionika sp. nov. 4 - Poupin, 1996, pi. 4h.

MATERIAL EXAMINED. — French Polynesia. SMSRB (J. Poupin coll.): Marquesas Islands. Fatu Hiva.

Sin 303, 10°3 1 .4’S. 138°39.2’W, 210 m, 3.09.1990: 12 ovig. 9 7.8-12.5 mm, 4 9 7.9-10.3 mm (MNHN-Na 13182).

— Nuku Hiva (baie Anaho). Stn 495. 8°45.4’S, 140°03.7’W, 230 m. 11.02.1996: 5 ovig. 9 8.8-12.2 mm

(MNHN-Na 13274). — Tahuata. Stn 300, 9°54.5’S, 139°07.9'W, 190 m, 1.09.1990: 1 ovig. 9 9.5 mm (MNHN-Na

13271, drawn), 14 ovig. 9 7. 2-9. 9 mm, I 9 8.3 mm (2 ex. transferred to NTOU, others to USNM).

Types. — The ovigerous female (9.5 mm cl) from the Marquesas Islands (Tahuata) is the holotype (MNHN-Na

1327 1 ). The other specimens are paratypes.

Description. — Body of slender appearance. Rostrum long and far overreaching scaphocerite, about 1 .9 ( 1 .8-

2.2) times as long as carapace length, slightly S-shaped and armed throughout length with 8-12 dorsal and 6-10

ventral teeth. Three-4 post-rostral teeth present on carapace, 2-4 of them with basal sutures and slightly movable.

Orbital margin regularly concave, with lower lobe very slightly convex. Eye subspherical, bearing a distinct

ocellus. Both antennal and pterygostomian spines well developed, former elongated and much longer than latter.

Stylocerite tapered anteriorly and just exceeding basal segment of antennular peduncle. Scaphocerite 5. 3-6. 6 times

220

T.-Y. CHAN & A CROSNIER

as long as broad, with distolateral tooth overreaching distal margin of lamella. Basicerite spine moderately long,

more or less reaching proximal end of outer margin of scaphocerite.

Maxilliped III bearing well developed epipod and long exopod. with penultimate segment 1 .3-1.4 times as long

as distal segment, overreaching scaphocerite by 1/2- 1/3 of penultimate segment and entire length of distal segment.

Well developed epipods present on anterior four pereiopods. Pereiopod I overreaching scaphocerite by about 1/2

carpus and other distal segments. Pereiopods II subequal and bearing 20-23 carpal articles, exceeding scaphocerite

by 1/2-3/5 carpus and other distal segments. Pereiopod III exceeding scaphocerite by slightly more than lengths of

dactylus, propodus and carpus, with dactylus short and 0.10-0.16 times as long as propodus; dactylus elongate,

conical and lacking spinules on posterior border, with accessory spine half to almost as long as, and abutting,

terminal spine. Posterior two pereiopods similar to pereiopod III, with pereiopod IV exceeding scaphocerite by

about carpus and pereiopod V by 2/3-3/4 carpus as well as other distal segments.

Abdominal somite III slightly arched dorsally and with posterior margin convex. Posteroventral denticle present

on abdominal pleuron V, but absent on plcuron IV. Abdominal somite VI about twice (1.8-2. 4) as long as high.

Telson 1.0- 1.3 (avg. 1.2) times longer than abdominal somite VI, with three pairs of dorsolateral and three pairs of

terminal spines. Eggs suboval, about 0.5 mm in diameter.

Coloration. Body translucent and slightly red. Eyes dark brown. Antennular flagella white, antennal

flagella reddish. Tip of rostrum reddish. Pereiopods mainly white. Pleopods slightly reddish? with lateral margins

PLESIONIKA FROM FRENCH POLYNESIA

221

somewhat whitish. Distal margins of uropods yellow. Organs inside the carapace visible as reddish. Eggs bluish to

turquoise.

Size. — Largest female 12.5 mm cl. Smallest ovigcrous female 7.2 mm cl. Males unknown.

Remarks. — Other than having a smaller size and different coloration, the present form has a longer rostrum

and shorter abdominal somite VI than P. sindoi (Rathbun, 1906). The shortest rostrum recorded for the present

form is 1.8 times carapace length, which is just longer than the longest rostrum found in P. sindoi (1.7 limes

carapace length). The abdominal somite VI of P. sindoi is always more than 2.1 times as long as high and often as

long as or longer than the telson. However, in the present form the abdominal somite VI is always shorter than the

telson and with only four specimens having the abdominal somite VI more than 2.1 times as long as high. The

shape of the rostral teeth situated above the orbit is also somewhat different between the two forms: those of

P. sindoi have very wide bases (much wider than high) and the dorsal margin somewhat concave, whereas those of

P. protati are more slender (base narrower than high) and with the dorsal margin somewhat convex. Thus, it seems

justified to separate the present form from P. sindoi.

The specimens from northwestern Australia reported by Hanamura and Takeda (1987) as P . ocellus arc all

in very poor conditions (rostrum, all perciopods, and telson incomplete). It is not known whether they belong to

P. fimbriata Chace, 1985, P. sindoi or the present new species. Hanamura and Takeda (1986) mentioned that

the specimens have the abdominal somite VI about 1.8 times as long as high, but in their figure it is more than

twice as long as high!

Etymology. — This new species is named after Charles Protat, Captain of the fishing vessel " Marara'\ for

his excellent help in collecting most of the specimens for this study.

Distribution. — Only known with certainty from French Polynesia, at depths of 190-210 m.

Plesionika payeni sp. nov.

Figs 19, 39

Plesionika sp. nov. 5 - Poupin, 1996, pi. 5g.

Material EXAMINED. — French Polynesia. SMSRB (J. Poupin coll.): Tuamotu Islands. Mururoa.

Stn 475, 21°51.2'S, 139°00.6‘W, 250 m, 11.04.1995: 1 ovig. 9 8.2 mm (MNHN-Na 13272). — Stn 499, 21°47.6'S,

1 38°55.7'W, 200 m, 5.05.1996: 1 ovig. 9 8.4 mm (MNHN-Na 13273).

Types. — The ovigerous female (8.4 mm cl) (MNHN-Na 13273) collected at station 499 of the " Mararci ", off

Mururoa (Tuamotu Islands), is the holotype. The other specimen (station 475) is a paratype.

DESCRIPTION. — Rather small, with body slender. Rostrum with lateral carina in basal half, continuous with

orbital margin, slightly curving downwards in basal part, but distinctly curved upwards after passing antennular

peduncle, far exceeding scaphocerite and 1.6 times as long as carapace length; holotype with dorsal border,

(fig. 19e), bearing 7 teeth (including one subapical tooth), with distal teeth smaller and further apart; ventral border

armed with 6 teeth, posteriormost tooth large, situated just above second segment of antennular peduncle and far

away from other ventral teeth, which are restricted to distal 2/5 of rostrum and separated by slightly decreasing

space from base to tip; in the paratype (fig. 19a), the tip of the rostrum is broken and only 5 dorsal and 4 ventral

teeth can be seen. Post-rostral carina not particularly elevated, bearing 5 movable post-rostral teeth. Orbital margin

with feeble upper and lower lobes, middle part regularly concave. Eye subspherical and bearing a distinct ocellus.

Antennal and pterygostomian spines both well developed. Stylocerite sharply pointed and strongly curving

upwards, more or less extending to distal end of basal segment of antennular peduncle. Scaphocerite 4.4-5. 3 times

as long as broad, with distolateral tooth distinctly exceeding distal margin of lamella. Basicerite spine sharp but

short, just failing to reach proximal end of outer margin of scaphocerite.

222

T.-Y. CHAN & A. CROSNIER

Fig. 19. — Plesionika payeni sp. nov. a-d: ovig. 9 8.2 mm, paratype (MNHN-Na 13272), French Polynesia. Tuamotu

Islands, Mururoa, stn 475, 250 m: a, lateral view of carapace; b, lateral view of posterior part of abdomen and tailfan;

c-d, propodus and dactylus of pereiopod III. — e-f: ovig. 9 8.4 mm. holotype (MNHN-Na 13273), ibidem , st. 499,

200 m: e, rostrum; f, telson and uropods. Figures a-b at the same scale.

Maxilliped III bearing a small but distinct epipod and a long ramous exopod, with penultimate segment 1.0-

1.1 times as long as distal segment; overreaching scaphocerite by about 1/4 of penultimate segment and entire

distal segment. Anterior four pereiopods bearing small to rudimentary epipods; that on pereiopod I small but

distinct, while those of other pereiopods are progressively more reduced posteriorly, those of pereiopods III and IV

nearly absent. Pereiopod I overreaching scaphocerite by 1/3- 1/2 carpus as well as other distal segments. Pereiopods

II subequal, with 20-22 carpal articles, exceeding scaphocerite by lengths of dactylus, propodus and 1/2-2/3 of

carpus. Pereiopod III exceeding scaphocerite by lengths of dactylus, propodus, carpus and about 1/5 of merus;

dactylus conical and short, 0.05-0.06 as long as propodus, posterior margin bearing only one spinule near middle,

accessory spine about half as long as terminal spine and well separated from latter. Posterior two pereiopods

similar to pereiopod III, with pereiopod IV overreaching scaphocerite by entire carpus and pereiopod V by

4/5 carpus as well as other distal segments.

PLESIONIKA FROM FRENCH POLYNESIA

223

Abdominal somite III very slightly arched dorsally, with posterior margin convex. Abdominal pleura IV and V

both bearing a distinct posteroventral denticle. Abdominal somite VI 2. 4-2. 5 times as long as high. Telson very

slightly longer (1.04 times) than abdominal somite VI, with three pairs of dorsolateral and three pairs of terminal

spines. Eggs suboval, about 0.4 mm in diameter.

COLORATION. — Body yellowish, bearing a lateral white line and a longitudinal red stripe along dorsal

midline. A discontinuous, longitudinal red line also present laterally on carapace. Rostrum with anterior two thirds

reddish, posterior third transparent. Eyes dark brown. Outer margin of scaphocerite reddish. Antennular flagella

whitish; antennal flagella reddish. Pereiopods reddish, with dactyli whitish, basal segments somewhat yellowish.

Abdomen yellowish, but abdominal somite VI and tailfan entirely covered with broad, longitudinal red and white

stripes. Eggs light blue.

SIZE. — Only two specimens are known, both ovigerous females, of 8.2 and 8.4 mm cl.

Remarks. — This small and colourful species can be readily separated from most of the other species of the

genus by having reduced epipods on the maxilliped III and four anterior pereiopods. The other characters, such as

rostrum moderately long and bearing only a few teeth on both the dorsal and ventral borders, post-rostral teeth not

crest-like, pereiopods II subequal, both abdominal pleura IV and V bearing a distinct posteroventral denticle and

telson having three dorsolateral spines, align it somewhat with the true P. ocellus (Bate, 1888). Apart from the

fact that the epipods on the anterior four pereiopods are all well developed in P. ocellus , this French Polynesian

form is much smaller in size and has a very different coloration (e.g. fig. 39 and Hayashi, 1986, fig. 82).

Moreover, the arrangement of rostral teeth (particularly the posteriormost ventral tooth being separated very far

from the other teeth in the French Polynesian material), the length of abdominal somite VI, and the shape of the

orbital margin and dactyli of the posterior pereiopods, are very different between the two species.

The epipods of maxilliped III and the anterior four pereiopods in this French Polynesian form are all small and

progressively reduced posteriorly, with that of pereiopod II already very minute and those of pereiopods III and IV

almost absent. Therefore, the epipods of the pereiopods may easily be overlooked, as in the species of the

"P. edwardsii (Brandt, 1851)" group. Nevertheless, it can still be readily separated from those poorly known

species described as lacking epipods on the pereiopods [e.g. P. minor Caiman, 1939 and P. costelloi (Yaldwyn,

1971)] by the shape of the rostrum (e.g. length and arrangement of rostral teeth) and pereiopods (e.g. relative

length of left and right pereiopod II, shape of dactyli of posterior pereiopods). Hence there is little doubt that this

French Polynesian species is new to science.

Only two specimens of this new species were available in the present study. The holotype is intact (though

with some pereiopods detached) but the paratype has the tip of the rostrum broken (still carrying five dorsal and

four ventral teeth) and the telson broken in half. However the specimens are essentially identical in all other

respects.

Etymology. — This new species is named after Dr C. Pa YEN, head of SMSRB, for his constant help and

interest in the deep water fauna studies of French Polynesia.

Distribution. — At present only known from Tuamotu Islands (Mururoa) in French Polynesia, at depths of

200-250 m.

Plesionika picta sp. nov.

Figs 20, 40

Plesionika sp. nov. 2 - Poupin, 1996, pi. 4f.

MATERIAL EXAMINED. — French Polynesia. SMSRB (J. Poupin coll.). Marquesas Islands. Va Pou

(Vaiehu Ray). Stn 60, 9°23.0'S. 140°09.0’W, 520 m, 26.01.1988: 1 9 14.4 mm (MNHN-Na 13067).

224

T.-Y. CHAN & A. CROSNIER

Tuamotu Islands . Fangataufa . Stn 440, 22°14.06'S, 138°47.74'W, 650 m, 15.11.1994: 1 8 12.5 mm,

1 9 13.4 mm [transferred to NTOU]. — Gambier. Stn 74, 23°08.0’S, 134°53.0'W, 660 m, 10.06.1988: 1 8 12.3 mm

(MNHN-Na 13069).

Tubuai Islands. Maria. Stn 422, 21°47.9'S, 154°43.8'W, 680 m. 7.08.1991: 2 8 11.2 and 15.2 mm, 1 ovig. 9

14.8 mm; 19 1 1.1 mm [transferred to USNM]. — Rurutu. Stn 107-108, 22°27.0’S. 15r23.0'W, 570 m, 3.09.1988:

1 8 15.2 mm. 1 9 10.2 mm (MNHN-Na 13070). — Stn 146, 22°27.8'S, 151022.8*W, 580 m, 9.03.1989: 1 8 14.8 mm

(MNHN-Na 13124, drawn); 1 8 13.4 mm (MNHN-Na 13289. drawn); 2 8 13,4 and 15.5 mm, I 9 12.8 mm (MNHN-Na

13066). — Stn 399, 22°28.4’S, 151°23’W, 710 m, 27.11.1990: 2 8 12.3 and 15.8 mm (MNHN-Na 13068).

TYPES. — The male (14.8 mm cl) collected at station 146 of the " Marara ", off Rurutu (Tubuai Islands), is the

holotype (MNHN-Na 13124). The other specimens are paratypes.

Description. — Body somewhat robust in appearance. Rostrum with lateral carina very strong in basal half

and more or less continuous with orbital margin, basal half rather broad and nearly horizontal, while distal half is

narrow and gently to distinctly curved upwards, overreaching scaphocerite, and about 0.9 times (0.8-1. 1 ) as long as

carapace; dorsal border bearing 2-5 (mostly 3-4) teeth, dorsal teeth generally restricted to basal 3/5 of rostrum;

ventral border with 5-10 (mostly 8-9) evenly distributed teeth. Post-rostral carina elevated and somewhat lamellate

but not crest-like; 7-10 post-rostral teeth present, with anteriormost tooth (very rarely anterior two teeth) fixed,

while posterior 6-9 teeth are movable and bear a distinct basal suture. Orbital margin with upper lobe nearly

straight and dorsally slightly inclined forwards, middle part regularly concave, while lower lobe is slightly convex.

Eye roughly kidney-shaped, bearing a distinct ocellus. Antennal spine absent (replaced by a broad angle) or minute.

Pterygostomian spine small but sharply pointed. Stylocerite narrow and tapered distally, strongly curving upwards

and extending to distal end of basal segment or middle of second segment of antennular peduncle. Scaphocerite 3.1-

3.9 (avg. 3.6) times as long as broad, with distolateral tooth more or less reaching distal margin of lamella.

Basicerite spine moderately long and just overreaching proximal end of outer margin of scaphocerite.

Maxillipcd III bearing well developed epipod and long strip-like exopod, with penultimate segment 0.9-

1.0 times as long as distal segment; overreaching scaphocerite by more or less entire distal segment. Anterior two

pereiopods bearing well developed epipods, epipod on pereiopod III smaller and sometimes even rudimentary,

pereiopods IV almost always lacking epipod (see Remarks below) and pereiopods V always without. Pereiopod I

overreaching scaphocerite by lengths of dactylus, propodus and 1/3-1 of carpus. Pereiopods II subequal, bearing 15-

21 (av. 17) carpal articles, exceeding scaphocerite by lengths of dactylus, propodus and 2/5- 1/2 of carpus.

Pereiopod III exceeding scaphocerite by entire carpus and a small part of merus (up to 1/5) as well as other distal

segments; dactylus conical, short, 0.08-0.13 (av. 0.10) times as long as propodus, posterior margin usually

bearing one spinule, accessory spine abutting, and slightly shorter than, terminal spine. Posterior two pereiopods

similar to pereiopod III, with pereiopod IV overreaching scaphocerite from 2/3 carpus to a small part of merus and

pereiopod V by 1/2-4/5 carpus as well as other distal segments.

Abdominal somite III slightly arched dorsally, with posterior margin convex. Both abdominal pleura IV and V

bearing a distinct posteroventral denticle. Telson 1.3- 1.5 times longer than abdominal somite VI, with four pairs

of dorsolateral and three pairs of terminal spines. Eggs suboval, about 0.4 mm in diameter.

COLORATION. — Carapace reddish, with posterior margin white. Rostrum transparent, but with tip reddish.

Eyes dark brown. Thoracic appendages and antennal flagella reddish. Antennular flagella reddish, but with a white

band at middle. Abdomen and tailfan almost transparent, with alternated broad red and white bands on somites I to

V. Pleopods mostly transparent.

SIZE. — Largest female (ovigerous) 14.8 mm cl. and largest male 15.8 mm cl. Only the largest female is

ovigerous.

Remarks. — The present form differs from most other species of the genus in bearing epipods on the anterior

three pereiopods, but lacking them on the last two. At present, only P. intermedia Chace, 1985 from the

Philippines has such an arrangement of epipods on the pereiopods in the genus. However, almost all the major

characters are different between the present species and P. intermedia, which belongs to the "P. martia (A. Milne

Edwards, 1883)” group. The general appearance (ignoring the gill formula) of the present form (e.g. many movable

PLESIONIKA FROM FRENCH POLYNESIA

225

Fig. 20. — Plesionika picta sp. nov. 6 14.8 mm, holotype (MNHN-Na 13124), French Polynesia, Tubuai Islands, Rurutu,

stn 146, 580 m: a. lateral view of carapace; b, lateral view of posterior part of abdomen and tail fan; c-d, propodus and

dactylus of pereiopod III. — e, 6 13.4 mm, paratype (MNHN-Na 13289), ibidem , rostrum.

Figures a-b at the same scale.

post-rostral teeth, pereiopod II subequal, four distolateral spines on telson, etc.), indicates that it is more closely

related to the "P. carsini Crosnier, 1986" group. Nevertheless, the present species is probably unique in the genus

in having the antennal spine absent or very minute, and the scaphocerite somewhat oblique and not horizontal (i.e.

with inner side remarkably higher than the outer side). Further differences of the present form from P. carsini , as

well as from P. poupini sp. nov., are its much smaller size (ovigerous female only 14.8 mm cl), the rostrum and

pereiopods relatively shorter, the dactyli of the posterior pereiopods relatively longer while the scaphocerite is

226

T.-Y. CHAN & A. CROSNIER

relatively broader, and the slightly fewer ventral rostral, but more post-rostral, teeth. As this French Polynesian

form is very distinct (including coloration), and can readily be separated from all the other species known to date in

the genus, it is here described as new.

Although the present new species generally lacks an epipod on pereiopod IV and sometimes has that of

pereiopod III rudimentary, two specimens (the female from Ua-Pou and a male from Stn 422) have a small but

distinct epipod on the left pereiopod IV (right pereiopod IV completely lacks an epipod in both specimens). Since

all the other 14 specimens examined do not bear any epipod-like process on the pereiopod IV, the presence of a

small epipod on the pereiopod IV is likely to be an abnormality in this species. Furthermore, the rostrum of this

species is generally slightly shorter than the carapace, only one specimen having the rostrum just longer than the

carapace length. The distribution of the dorsal rostral teeth also appears to be highly variable in this species. The

dorsal rostral teeth are often restricted to about the basal 2/3 to 1/2 of the rostrum, but sometimes almost the entire

rostrum has dorsal rostral teeth (i.e. similar to the situation of the ventral rostral teeth). Nevertheless, the

distribution pattern of dorsal rostral teeth does not seem to be related to the length of the rostrum, or to the size

and sex of the specimens.

Etymology. — The name picta refers to the rather artistic colour pattern on the body of this species.

Distribution. — So far known only from French Polynesia, at depths of 520-710 m.

Plesionika spinidorsalis (Rathbun, 1906)

Fig. 41

Pandalus spinidorsalis Rathbun. 1906: 917, pi. 21, fig. 5 [type-locality: Hawaii].

Plesionika spinidorsalis - Chace, 1985: 132, figs 60-61. — POUPIN et al ., 1990: 16. — Poupin, 1996: pi. 5e.

Material EXAMINED. — French Polynesia. SMSRB (J. Poupin coll.): Society Islands. lies sous le

Vent (Bora Bora). Stn D32, 16°28.4’S, 151°47.5’W, 562 m, 23.06.1990: 2 ovig. 9 14.2 and 14.5 mm (transferred

to USNM). — Huahine . Stn 184, I6°44.5’S, I51°04.0'W. 540 m. 16.08.1989: 1 9 13.6 mm.

Tuamotu Islands. Gambier. Stn 74, 23°08.0'S, 134°53.0'W, 660 m. 10.06.1988: I ovig. 9 15.0 mm. —

Mururoa . Stn 127, 21°51.2’S, 138°47.3’W, 600 m. 29.11.1988: 1 ovig. 9 10.8 mm, 1 9 13.4 mm.

Tubuai Islands. Tubuai. Stn 145, 23°18.2'S, 149°27.rw. 720 m, 8.03.1989: 1 6 15.3 mm.

DIAGNOSIS. — A detailed description of this species can be found in CHACE (1985).

COLORATION. — Body nearly transparent with red internal organs visible inside carapace. Eyes dark brown.

Dactyl i of posterior pereiopods somewhat reddish. Eggs blue.

SIZE. — Largest size nearly 20 mm cl. The French Polynesian material ranges from 10.8 mm to 15.3 mm cl,

the smallest specimen being an ovigerous female.

Remarks. — The 7 specimens from French Polynesia agree very well with the description of this species

provided by Chace (1985), with the abdominal somite VI being 1.51-1.86 times as long as high. Only one

specimen (15.3 mm cl) has the rostrum intact, being 0.6 times as long as carapace length. The eggs of this species

are about 0.4 mm in diameter.

Distribution. — Western Pacific, recorded from Hawaii, French Polynesia, Indonesia, the Philippines and

probably also present in the South China Sea. At depths of 100-1250 m, mostly more than 400 m deep.

CONCLUSION

As a result of the scarcity of deep-water collections, no specimen of Plesionika had been collected from French

Polynesia until 1978. The SMSRB deep-trap fishing operations, conducted by B. RICHER DE FORGES and, on a

Source :

P LESION! KA FROM FRENCH POLYNESIA

227

larger scale, J. POUPIN, have permitted the capture of twenty-one species of Plesionika , ten of them new to

science. This is a remarkable result that shows how poor our knowledge of the bathyal fauna is, not only in

French Polynesia but in the Indo-West Pacific region as a whole. Moreover, this impression is highly confirmed

by the general results obtained during the MUSORSTOM campaigns.

As already noticed for most of the other deep-water groups, it appears that color photographs of live material

greatly help the study of the material.

Plesionika is a puzzling genus. It involves more than eighty species that can be grouped into several

homogeneous groups. It could be then more sensible to split it into several more homogeneous genera. This,

however, is practically impossible since certain species always present characters that are intermediate between the

intended new genera. No doubt, for example, that the " laevis group", related to the genus Heterocarpus, but with

some of the species lacking lateral carina on the carapace, will be greatly discussed in the future.

Another difficulty is that many species of Plesionika are common and have a wide geographical distri¬

bution. Morphological variations are observed among specimens collected from far different locations. It is

therefore difficult to decide if such variations arc specific or the result of geographical differences. In this work, the

reader will find some examples of these problems, particularly under Plesionika nesisi. P. macropoda and

P. williamsi.

With his remarkable monograph on the Pandalidae of the " Albatross " expedition, F.A. CHACE (1985) has

undoubtedly renewed the interest for the Indo-West Pacific Plesionika studies. The present study should also have

further development, especially if we consider the very abundant collections made in Taiwan and during the recent

MUSORSTOM campaigns around New Caledonia, Indonesia, Chesterfield islands. Vanuatu, and Wallis and Futuna

islands. The interest for these new collections is increased by numerous color photographs taken just after trawl

operations.

Preliminary examinations of this material show that, although more new species are found, others belong to

the new Polynesian species herein described. Thus, it would be speculative to conclude too quickly to a Polynesian

endemism, may be more apparent than real. More reliable conclusions on that point will have to wait until the

complete study of the recent MUSORSTOM collections, unfortunately not before some years.

ACKNOWLEDGMENTS

We sincerely thanks J. POUPIN of the Service Mixte de Surveillance Radiologique et Biologique for

enthusiastically collecting so many interesting specimens for the present study. His invaluable efforts in taking

colour photographs of the fresh material also proved to be very helpful.

Grateful acknowledgements are extended to F.A. CHACE, Jr. and R.B. Manning of the National Museum of

Natural History, Washington, D.C. for exchanging specimens of some Plesionika species and sending us on loan

the syntypes of P. sindoi\ and P. Clark of The Natural History Museum, London for sending us on loan the

syntypes of P. ocellus.

Special thanks are due to F. THEUREAU for preparing most of the drawings in this report and

J.-F. DEJOUANNET for mounting the plates of photos.

We also like to thank ORSTOM (Instilut fran^ais de Recherche scienlifique pour le Developpement en

Cooperation) for providing a short-term research grant to the first author to study at the Museum national

d’Histoire naturelle, Paris, thereby making the present work possible.

This study is also a contribution from a research grant on the Decapod Crustacea of Taiwan, supported by the

National Science Council, Taiwan. R.O.C.

REFERENCES

Alcock, A., 1901. — A descriptive catalogue of the Indian deep-sea Crustacea Decapoda Macrura and Anomala, in the

Indian Museum. Being a revised account of the deep-sea species collected by the Royal Indian Marine Survey Ship

Investigator. Indian Museum, Calcutta. 286 pp., 3 pis. Calcutta.

228

T.-Y CHAN & A. CROSNIER

BALSS, H., 1925. — Macrura der Dcutschen Tiefsee-Expedition, 2. Natantia, Teil A. Wissenschaftliche Ergebnisse der

Deutschen Tiefsee-Expedition auf dem Dampfer "Valdivia" 1898-1899 , 20 (5): 217-315, figs 1-75, pis 20-28.

Bate, C.S., 1888. — Report on the Crustacea Macrura collected by the Challenger during the years 1873-76. Report on

the Scientific Results of the Voyage of H.M.S . "Challenger" during the Years 1873-76,24. xc+942 pp., figs 1-76,

pis 1-157.

Borradaile, L.A.. 1900. — On the Stomatopoda and Macrura brought by Dr. Willey from the South Seas. In: A. Willey,

Zoological results based on materials from New Britain, New Guinea. Loyalty Islands and elsewhere, collected during

the years 1895, 1896 and 1897. 4: 395-428, pis 36-39. Cambridge.

Borradaile, L.A., 1915. — Notes on Caridea. Annals and Magazine of Natural History, ser. 8. 15: 205-213.

Brandt, F., 1851. — Krebse. In: A.T. Middendorff, Rcise in den aussersten Norden und Osten Sibiriens wahrend der

Jahre 1843 und 1844 mit allerhochster Genehmigung auf Veranstaltung der Kaiserlichen Akademie der Wissenschaftcn

zu St. Petersburg ausgefuhrt und in Verbindung mit vielen Gelehrten herausgegeben. 2(1): 77-148. pis 5-6.

St. -Petersburg.

Burukovsky. R.N., 1986. — A new shrimp species from the genus Heterocarpus (Crustacea: Decapoda: Pandalidae) and a

brief review of the genus. Byulleten Moskovskogo Obschchestva Ispytatelei Prirody Otdel Biologicheskii, 91 (5):

62-73. figs 1-4 [in Russian].

Burukovsky, R.N., 1992. — New species of the genus Plesionika (Crustacea, Decapoda, Pandalidae) from an underwater

rise in the Pacific Ocean. Zoologicheskij Zhurnal, 71 (7): 145-147, fig. 1 [in Russian).

BURUKOVSKY, R.N., 1993. — Plesionika alaini nom. nov., a new name for Plesionika crosnieri Burukovsky, 1992

(Crustacea Decapoda Pandalidae). Arthropoda Selecta, 2 (4): 18.

Burukovsky, R.N., 1995. — Two new species of shrimps of the genus Pasiphaea and new records of other shrimps.

Zoologicheskij Zhurnal, 14 (12): 121-126, figs 1-14 [in Russian].

Calm AN, W.T., 1939. — Crustacea: Caridea. The John Murray Expedition 1933-34 Scientific Reports. 6 (4): 183-224,

figs 1-8.

Chace, F.A. Jr., 1939. — Reports on the scientific results of the first Atlantis Expedition to the West Indies, under the

joint auspices of the University of Havana and Harvard University. Preliminary descriptions of one new genus and

seventeen new species of decapod and stomatopod Crustacea. Memorias de la Sociedad Cubana de Historia Natural, 13

(1): 31-54.

Chace, F.A. Jr.. 1985. — The Caridean Shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition. 1907-

1910. Part 3: Families Thalassocarididae and Pandalidae. Smithsonian Contribution to Zoology , (411): 1-143,

Figs 1-62.

Chace, F.A. Jr., 1989. — The holotype of Heterocarpus alexandri A. Milne-Edwards (Crustacea: Decapoda: Pandalidae).

Proceedings of the Biological Society of Washington , 102 (1): 84-88, fig. 1.

Chan., T.-Y. & CROSNIER, A., 1991. — Crustacea Decapoda : Studies of the Plesionika narval (Fabricius, 1787) group

(Pandalidae). with descriptions of six new species. In: A. CROSNIER (ed.). Resultats des Campagnes Musorstom, Vol.

9. Memoires du Museum national d'Histoire naturelle, (A), 152: 413-461, Figs 1-39.

Chan, T.-Y. & Yu. H.P., 1991. — Two similar species: Plesionika edwardsii (Brandt, 1851) and Plesionika crosnieri, new

species (Crustacea: Decapoda: Pandalidae). Proceedings of the Biological Socien of Washington, 104 (3): 545-555,

figs 1-3.

Crosnier. A.. 1986a. — Crevettes de la famille des Pandalidae recoltees durant ces demieres annees cn Polynesie

fran9aise. Description de Plesionika chacei et P. carsini spp. nov. Bulletin du Museum national d'Histoire naturelle,

Paris, 4eme serie. section A, 8 (2): 361-377, figs 1-4.

CROSNIER, A.. 1986b. — Plesionika fenneri, nouveau nom pour Plesionika chacei Crosnier. 1986. Bulletin du Museum

national d'Histoire naturelle, Paris, 4eme serie, section A, 8 (3): 691.

Crosnier, A., 1988. — Sur les Heterocarpus (Crustacea, Decapoda, Pandalidae) du sud-ouest de 1'ocean Indien. Remarqucs

sur d’autres espfcces ouest-pacifiques du genre et description de quatre taxa nouveaux. Bulletin du Museum national

d'Histoire naturelle, Paris, 4£me serie, section A, 10 (1): 57-103, Figs 1-7, pis 1-4.

Crosnier, A. & Forest, J., 1973. — Les crevettes profondes de l'Atlantique oriental tropical. Faune Tropicale. 19.

1-409, figs 1-121.

PLESI0N1KA FROM FRENCH POLYNESIA

229

Fabricius, J.C., 1787. — Mantissa Insectorum sistens eorum Species nuper detectas adiectis Characteribus genericis,

Differenliis specificis, Emendationibus, Observationibus. 1: xx + 348 pp. Hafniae.

Forest, J.. 1964. — Sur une crevette rccucillie au cours de la campagne de chalutage dans le golfe de Guince, Plesionika

williamsi sp. nov. Bulletin du Museum national d'Histoire naturelle, Paris, 2eme s£rie, 35 (6): 620-629, Figs 1-4.

Hanamura, Y. & Takeda, M., 1987. — Family Pandalidae (Crustacea, Decapoda, Caridea) collected by the RV "Soela"

from the Northwest Australian Shelf. Bulletin of the National Science Museum , Series A (Zoology). 13 (3): 103-121,

figs 1-5. Tokyo.

Hayashi, K.I., 1986. — Penaeoidea and Caridea. In: K. Baba, K.I. Hayashi, & M. Toriyama (eds). Decapod Crustaceans

from continental shelf and slope around Japan, 336 pp., 23 figs, 176 photos. Japan Fisheries Resource Conservation

Association. Tokyo.

Holthuis, L.B., 1951. — The Caridean Crustacea of the Tropical West Africa. In: A.F. Bruun (ed.). Atlantide-Report.

Scientific Results of the Danish Expedition to the Coast of Tropical West Africa 1945-1946, 2: 7-187, figs 1-34.

Kensley, B., Tranter. H.A. & Griffin. D.J.G., 1987. — Deepwater decapod Crustacea from Eastern Australia (Penaeidea

and Caridea). Records of the Australian Museum , 39: 263-331, figs 1-25, 1 pi. in colour.

King, M.G., 1984. — The species and depth distribution of deepwater caridean shrimps (Decapoda, Caridea) near some

southwest Pacific islands. Crustaceana. 47 (2): 174-191, figs 1-7.

Lemaitre, R., 1984. — Decapod crustaceans from Cay Sal Bank, Bahamas, with notes on their zoogeographical

affinities. Journal of Crustacean Biology, 4 (3): 425-447, figs 1-9.

Man, J.G. de, 1917. — Diagnoses of new species of Macrurous Decapod Crustacea from the Siboga-Expedition.

Zoologische Mededeelingen. Leiden, 3 (4): 279-284.

Man, J.G. DE, 1920. — The Decapoda of the Siboga-Expedition. Part IV. Families Pasiphaeidae, Stylodactylidae,

Hoplophoridae, Nematocarcinidae, Thalassocaridae, Pandalidae, Psalidopodidae, Gnathophyllidae, Processidae,

Glyphocrangonidae and Crangonidae. Siboga-Expeditie , monogr. 39 a3: 1-318, 25 pis.

Milne Edwards, A., 1881. — Description de quelques Crustaces Macroures provenant des grandcs profondeurs de la mer

des Antilles. Annales des Sciences Naturelles, Zoologie, ser. 6, 11 (4): 1-16.

Milne Edwards, A., 1883. — Recueil de figures de crustaces nouveaux ou peu connus. 3 pp.. 44 pis, Paris.

MONTEROSSA, O.E., 1988. — Heterocarpus cutressi, new species, and Plesionika macropoda Chace, 1939: two caridean

shrimps of the family Pandalidae (Crustacea: Decapoda) from Puerto-Rico and the U.S. Virgin islands. Proceedings of

the Biological Society of Washington, 101 (3): 633-639, figs 1-5.

Parisi, B., 1919. — Natantia, Part VII. In: I Decapodi giapponesi del Museo di Milano. Atti della Societd Italiana di

Scienze Natural i, 58: 59-99, Figs 1-8, pis 3-6.

Paulmier, G., 1993. — Crustaces profonds captures aux casiers aux Antilles fran^aiscs, 34 pp, 33 pis. IFREMER. Paris.

Pequegnat, L.H., 1970. — Deep-sea Caridean Shrimps with description of six new species. In: W.E. Pequegnat &

F.A. Chace, Jr. (eds). Contributions on the Biology of the Gulf of Mexico. Texas A & M University Oceanographic

Studies, 1: 59-123, figs 4.1-4.17.

Poupin. J., 1988. — Deep-water caridean shrimps on the steep slope of the Moruroa atoll. In: J.H. Choat et al. (eds).

Proceedings of the Sixth International Coral Reef Symposium, Townsville, Australia. 8th- 12th August 1988, 2: 27-

30, figs 1-6.

Poupin, J., 1994. — Faune marine profonde des Antilles fran^aises. Recoltes du navire Polka faites en 1993. Etudes et

Theses, ORSTOM, Paris, 79 pp., 1 Fig., 5 pis.

Poupin, J., 1996. — Atlas des Crustaces marins profonds de Polynesie Fran^aise. Recoltes du navire Marara

(1986/1996), 59 pp., 20 pis. SMSRB, Montlhery.

Poupin, J. & Richer deForges, B.. 1991. — New and rare crustaceans from French Polynesia (Crustacea, Decapoda). In:

P.J.F. Davie & R.H. Quinn (eds.). Proceedings of the 1990 International Crustacean Conference. Memoirs of the

Queensland Museum, 31: 211, Fig. 1.

Poupin, J., Tamarii, T. & Vandenboomgaerde, A., 1990. — Peches profondes aux casiers sur les pentes oceaniques des

Ties de Polynesie Frangaise. (N/O Marara- 1986/1 989). Notes et Documents d'Oceanographie du Centre ORSTOM de

Tahiti , (42): 1-97, figs 1-21, pis 1-3.

230

T.-Y. CHAN & A. CROSNIER

Raso. J.E.G., 1996. — Crustacea Decapoda (excl. Sergestidae) from ibero-moroccan waters. Results of BALGIM-84

Expedition. Bulletin of Marine Science. 5 8 (3): 730-752, fig. 1-5.

Rath BUN, M.J., 1906. — The Brachyura and Macrura of the Hawaiian Islands. Bulletin of the United States Fish

Commission , (1903), 23 (3): 827-930, figs 1-79, pis 1-24.

SPRINGER, S. & BULLIS, H.R., 1956. — Collections by the Oregon in the Gulf of Mexico. U.S. Fish and Widelife Service

Special Scientific Report. Fisheries , (196): 1-134.

SQUIRES, H.J. & Barragan, J.H., 1976. — A new species of Plesionika (Crustacea, Decapoda, Pandalidae) from the

Pacific coast of Colombia. Pacific Science. 30 (2): 1 13-1 17, figs 1-2.

SUSEELAN, C. & Mohamed, K.H., 1969. — On the occurrence of Plesionika ensis (A. Milne Edwards) (Pandalidae,

Crustacea) in the Arabian Sea with notes on its biology and fishery potentialities. Journal of the Marine Biological

Association of India. 10 (1): 88-94, figs 1-4.

Toriyama, M., Horikawa. H. & Kishida, S.. 1990. — Preliminary reports on ten rare Caridean Shrimps (Decapoda,

Caridea) from Tosa Bay and its adjacent waters. Bulletin of Nansei National Fisheries Research Institute, 23: 13-26,

fig. 1, pis 1-5 [in Japanese].

Yaldwyn. J.C., 1971. — Preliminary descriptions of a new genus and twelve new species of Natant Decapod Crustacea

from New Zealand. Records of the Dominion Museum , 7 (10): 85-94.

Yokoya, Y., 1933. — On the distribution of Decapod Crustaceans inhabiting the continental shelf around Japan, chiefly

based upon the materials collected by S.S. Soyo-Maru during the years 1923-1930. Journal of the College of

Agriculture, Tokyo Imperial University. 12 (1): 1-226, figs 1-71.

COLORED PHOTOGRAPHS

Fig. 21. — Plesionika carsini Crosnier, 1986. French Polynesia, Tuamotu Islands, Fangataufa, stn 438, 22°I2.3’S,

138°46.6'W, 410 m.

Fig. 22. — Plesionika poupini sp. nov., paratype, French Polynesia, Tuamotu Islands. Fangataufa, stn 438, 22°12.3’S,

138°46.6'W. 410 m.

Fig. 23. — Plesionika edwardsii (Brandt. 1851). French Polynesia, Tuamotu Islands, Mururoa. stn 452, 21°53,0’S,

139°02.9’W. 330 m.

Fig. 24. — Plesionika reflexa Chace, 1985. French Polynesia, Tubuai Islands, Rapa, stn 465, 27°35.9’S, I44°27.6’W,

765 m.

FlG. 25. — Plesionika ensis (A. Milne Edwards, 1881). Antilles, "Polka", stn DPI, 15°50.82'N, 61°40.75’ W, 460 m.

Fig. 26. — Plesionika fenneri Crosnier, 1986, juvenile. French Polynesia, Tubuai Islands. Rurutu, stn 146, 22°27.8’S,

151°22.8'W, 580 m.

FlG. 27. — Plesionika fenneri Crosnier, 1986. French Polynesia, Tuamotu Islands. Mururoa, stn 229, 2I°51.9’S.

139°2.2'W. 490 m.

Fig. 28. — Plesionika nesisi (Burukovsky, 1986). French Polynesia. Tuamotu Islands, Fangataufa, stn 85, 22°13.0'S,

138°42.2'W, 780 m.

Source :

PLESIONIKA FROM FRENCH POLYNESIA

231

Source : MNHN, Paris

232

T.-Y. CHAN & A. CROSNIER

Fig. 29. — Plesionika laevis (A. Milne Edwards, 1883). Antilles, "Polka", stn DPI, 15°50.82'N, 61°40.75‘ W, 460 m.

Fig. 30. — Plesionika macropoda Chace, 1939. French Polynesia, Society Islands, Huahine, stn 405, 16°45.8'S,

151°3.8'W, 310 m.

Fig. 31. — Plesionika macropoda Chace, 1939. Antilles, "Polka", stn C21 1, 6° 1 1 .8 1 ’N, 61049.1'W, 450 m.

Fig. 32. — Plesionika williamsi Forest, 1964. French Polynesia, Marquesas Islands, Ua Pou, stn 59-60, 9°23.0'S,

140°09.0'W, 450-520 m.

Fig. 33. — Plesionika williamsi Forest, 1964. Antilles, "Polka", stn EP1, 15°50.93'N, 61°41.39’W, 560 m.

Fig. 34. — Plesionika erythrocyclus sp. nov., paratype. French Polynesia, Tubuai Islands. Rurutu, stn 356, 22°30.3'S,

151 °21 .7’W, 260 m.

Fig. 35. — Plesionika erythrocyclus sp. nov., Taiwan, northeastern coast, about 300 m.

Fig. 36. — Plesionika sindoi (Rathbun, 1906). French Polynesia, Tuamotu Islands, Makemo, stn 250, 16°34.9’S,

143°27.2'W, 435 m.

Source :

PLESIONIKA FROM FRENCH POLYNESIA

233

Source : MNHN. Paris

234

T-Y CHAN & A. CROSNIER

Fig. 37. — Plesionika sindoi (Rathbun, 1906). French Polynesia, Marquesas Islands. Fatu Hiva, stn 306, I0°31.1'S,

138°39.4'W, 250 m.

Fig. 38. — Plesionika protati sp. nov., from the type series. French Polynesia, Marquesas Islands. Tahuata, stn 300,

9°54.5’S, 139°7.9’W, 190 m.

Fig. 39. — Plesionika payeni sp. nov., holotype. French Polynesia, Tuamotu Islands, Mururoa, stn 475, 21°51.2'S,

139°00.6'W, 250 m, 11.04.1995.

Fig. 40. — Plesionika picta sp. nov., paratype. French Polynesia, Tubuai Islands, Rurutu, stn 339, 22°28.4'S,

151°23.0’W, 710 m.

Fig. 41. — Plesionika spindorsalis (Rathbun, 1906). French Polynesia, Society Islands, Bora Bora, stn D32, 16°28.37'S,

151°47.52'W, 562 m.

Source : MNHN. Paris

SULTATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 18 — RESULT ATS DES CA

Crustacea Decapoda: Diacanthurus gen. nov.,

a new genus of hermit crabs (Paguridae)

with both Recent and fossil representation,

and the descriptions of two new species

Patsy a. McLaughlin

Shannon Point Marine Center. Western Washington University

1900 Shannon Point Road. Anacortes. WA 98221-4042. U.S.A.

&

Jacques FOREST

Museum national d'Histoire naturelle

Laboratoire de Zoologie (Arthropodes)

61 rue Buffon, 75005 Paris

ABSTRACT

The new genus, Diacanthurus , is proposed for a group of three Recent and one fossil species formerly assigned to the

heterogeneous genus Pagurus Fabricius. In addition to the transfer of Pagurus clifdenensis Hyden & Forest (fossil),

P. spinulimanus (Miers). P. rubricatus (Henderson), and P. ophthalmicus (Ortmann). two new species, Diacanthurus

eephyma sp. nov. from New Caledonia and Western Australia, and D. richeri sp. nov. from New Caledonia are assigned to

this new genus. Expanded diagnoses or descriptions and illustrations of all Recent species are provided.

RESUME

Crustacea Decapoda : Diacanthurus gen. nov., nouveau genre de Paguridae represente par des

formes actuelles et une forme fossile. Description de deux especes nouvelles.

Le nouveau genre Diacanthurus est propose pour un groupe comprenant trois especes actuelles et une espece fossile

precedemment attributes au genre heterogene Pagurus Fabricius. En plus de Pagurus clifdenensis Hyden & Forest (tossile),

P. spinulimanus (Miers). P. rubricatus (Henderson) et P. ophthalmicus (Ortmann), transfers au genre Diacanthurus .

McLaughlin, P.A. &, Forest, J., 1997. — Crustacea Decapoda: Diacanthurus gen. nov., a new genus of hermit crabs

(Paguridae) with both Recent and fossil representation, and the descriptions of two new species. In: A. Crosnilr (ed.),

Rtsultats des Campagnes MUSORSTOM, Volume 18. Mem. Mus. natn. Hist, nat., 176: 235-259. Paris ISBN: 2-85653-

511-9.

236

p. a. McLaughlin & j. forest

deux especes nouvelles sont d^crites: D. ecphyma sp. nov., de Nouvelle-Caledonie, des lies Chesterfield ct d'Australie

occidentale, et D. richeri . de Nouvelle-Caledonie. Des diagnoses developpees ou des descriptions, ainsi que des

illustrations, sont fournies pour toutes Ies espfeces actuelles.

Diacanthurus gen. nov. se distingue des autres genres de Paguridae par la structure particuli&re du telson : les deux lobes

posterieurs sont symetriques ou presque, en forme de croissants, avec un sommet aigu, el separes par une profonde

incision mediane dont Ies bords internes portent chacun une forte epine. D'autres traits morphologiques caractSrisent

encore le genre, comme le grand developpement de 1'article basilaire des pedoncules oculaires.

Les differences entre les especes portent notamment sur la forme et rornementation des chclipedes qui presentent un

remarquable gradient Svolutif suivant la sequence D. richeri , D. ophthalmicus , D. clifdenensis , D. spinulimanus ,

D. ecphyma et D. rubricatus. Ce gradient porte sur la largeur relative et l'asymetrie des mains, qui s'accroissent, alors que

les tubercules et dents de la face dorsale sont de plus en plus developpes.

En dehors des deux especes nouvelles, a distribution n£o-caledoniennc, le nouveau genre est represente dans plusieurs

regions de l'lndo-Ouest Pacifique : Nouvelle-Zelande ( D . spinulimanus et D. rubricatus), Japon (D. ophthalmicus ) et aussi

Australie occidentale (D. ecphyma).

Les distributions bathymelriques different suivant les especes. Celles-ci sont surtout rcpriSsentees entre 200 et

400 metres, a 1'exception de D. ophthalmicus, capture entre 100 et 200 metres, el de D. spinulimanus, le plus sou vent

recueilli par moins de 100 metres de profondeur.

INTRODUCTION

Hyden and FOREST (1980) described a new species of pagurid, Pagurus clifdenensis Hyden & Forest, from the

early Miocene of southern New Zealand. At the time, the authors noted the similarities between their fossil species

and two Recent New Zealand species, Pagurus spinulimanus (Miers, 1876) and Pagurus rubricatus (Henderson,

1888). Recognizing the distinctiveness of all three taxa, HYDEN and FOREST (1980) indicated that these species

would be transferred to a new genus when revisionary studies of the New Zealand pagurid fauna were completed.

Certain characters, not available in the fossil species, clearly set the two Recent species apart from other

Pagurus species, particularly the structure of the telson. It is, as the generic name Diacanthurus implies, composed

of two very prominent spines (Fig. la). This unique telson structure was also depicted by Miyake (1978) for the

Japanese species, Pagurus ophthalmicus (Ortmann, 1892). An examination of Ortmann's holotype by one of us

(PMcL) confirmed similarities indicative of a close phylogenetic relationship among the three Recent taxa. During

studies of the MUSORSTOM collections from New Caledonia, two additional and undcscribcd species have been

recognized that share all of the morphological characteristics of Diacanthurus as defined herein.

MATERIALS AND METHODS

The holotypes of Pagurus rubricatus and P. spinulimanus were borrowed from The Natural History Museum,

London (NHM); the holotype of P. ophthalmicus was borrowed from the Muscc Zoologique, Universite Louis

Pasteur, Strasbourg (MZUS). The MUSORSTOM collections are the property of the Museum national d'Histoire

naturelle, Paris (MNHN); the specimen from Western Australia is the property of the Western Australian Museum.

Perth (WAM). Supplemental materials of P. rubricatus and P. spinulimanus have come from the collections of the

New Zealand Oceanographic Institute (NZOI); National Museum of New Zealand, Wellington, (NMNZ), now the

Museum of New Zealand Te Papa Tongarewa (MoNZ), and include collections originally belonging to the

Zoology Department, Victoria University of Wellington (VUZ); Portobello Marine Laboratory, University of

Otago (PMLUO); and the personal collection of one of the authors (PMcL). Supplemental specimens of

P. ophthalmicus from the R/V Albatross Northwest Pacific Expedition are the property of the National Museum

of Natural History, Smithsonian Institution (USNM); additional specimens from Japan are from the collections of

Zoology, Crustacea, Natural History Museum and Institute, Chiba (CBM-ZC), and PMcL’s personal collection.

Specimens have, for the most part, been returned to their institutions of origin; however, some paratypes of the

new species have been deposited in the USNM. Most station data for the French bathyal faunal expeditions in New

Caledonia are documented in Richer de Forges (1990, 1993) and Richer de Forges and Chevillon (1996).

The abbreviations BS, DW, and CP refer to bottom station, Waren dredge and beam trawl, respectively. The

CRUSTACEA DECAPODA: DIACANTHURUS NEW GENUS OF HERMIT CRABS

237

measurement of shield length, given in parentheses ( ), and measured from the midpoint of the anterior margin of

the shield (broadly rounded rostral projection) to the midpoint of the posterior margin of the shield, provides an

indication of animal size. Measurements of the ocular peduncle refer exclusively to the length of the distal

segment, and are measured from the base to the distal margin (or apex) of the cornea along the lateral face; corneal

diameter represents maximum width of the cornea. Lengths of the dactyls and propodi of the ambulatory legs were

measured on the lateral faces from the dorsodistal margin (excluding the claw of the dactyl) to the dorsoproximal

margin. Terminology for the diagnoses and descriptions follows that of McLaughlin (1974, 1997). Two keys to

the species are presented. The first is based principally on characters of the chelipeds and ambulatory legs, which

may be lost in preserved specimens, thus rendering the key useless. The second, which is based essentially on

characters of the ocular peduncles and telson is less subject to the drawbacks than the first, is highly reliable for

large specimens, but may not be as applicable to small specimens. It is hoped that between the two, accurate

identifications of these generally quite similar species can be made.

Genus DIACANTHURUS nov.

Pa gurus Fabricius, 1775: 410 (in part).

Eupagurus Brandt, 1851: 105 (in part).

TYPE Species. — Eupagurus spinulimanus Miers, 1876. Gender masculine.

Diagnosis. — Eleven pairs of phyllobranchiate gills. Rostrum obsolete to broadly rounded. Ocular peduncles

well developed, intersegmental articulating membrane very prominent; ocular acicles widely separated, dorsal sur¬

face flattened or slightly convex, very broadly triangular, produced-distally and with strong terminal, marginal or

submarginal spine or spines. Antennal peduncles with supernumerary segmentation. Mandible with deeply concave

inner surface; ultimate segment of 3-segmented palp with 2 or 3 rows of short stiff bristles in distal half and 1 row

of finer setae proximally. Maxillule (Fig. lb) with internal endopodal lobe well developed, external lobe obsolete.

Maxilla (Fig. lc) with moderately broad, proximally subrectangular scaphognathite. First maxilliped (Fig. Id)

with basal portion of exopod somewhat bulbous. Third maxilliped (Fig. le) with crista dentata well developed and

1 accessory tooth.

Chelipeds unequal. Left cheliped with some degree of clockwise rotation of propodal-carpal articulation;

dorsolateral margin of chela weakly to strongly inflated proximally. Ambulatory legs with dactyls and propodi of

second and third pereopods similar. Fourth pereopods semichelate; propodal rasp consisting of several rows of

small corneous scales.

Males with paired gonopores, almost completely masked by ventral fringe of long stiff setae, no sexual tubes;

no paired pleopods, 3 unpaired, uniramous or very unequally biramous left pleopods (PI3-PI5). Females with paired

gonopores; no paired pleopods, 4 biramous left pleopods (p^-p^), first 3 egg-carrying, last as in males.

Abdomen with somites delimited by strong transverse fibrils. Tergite of sixth abdominal segment with deep

median transverse groove. Uropods strongly asymmetrical. Telson (Fig. la) with an indentation on the lateral

margins; posterior lobes generally symmetrical, each with primarily "half-moon" contour, blade-like lateral margin

and acute terminal angle, usually broadly separated and with distinct median cleft, inner margins each with

1 prominent spine in basal half.

Etymology. — From the Greek di meaning two, acantha meaning spine, and oura meaning tail, referring to

the distinctive telson characteristic of this genus.

Keys to the Recent species of Diacanthurus

1 . Carpi of third pereopods each with row of spines on dorsal margin . 2

— Carpi of third pereopods with only spine at dorsodistal angle (sometimes 1 additional

spinule proximally) . 4

238

p. a. McLaughlin & j. forest

2. Ocular peduncles short (Fig. 2d), stout, corneas noticeably dilated. Maximum width of

palm of right chela at least 1.25 (usually 1.5) times length of dorsomesial margin .

. D. rubricatus

— Ocular peduncles moderately long, not particularly stout, corneas not noticeably dilated.

Maximum width of palm of right chela only slightly greater than length of dorsomesial

margin . 3

3. Dactyl of right chela with row of short spines or spinulose tubercles on dorsomesial

surface; usually with 2 or 3 longitudinal rows of nearly coalesced, often blunt tubercles

on dorsal surface. Spines on carpi of third pereopods weakly developed .

. D. ophthalmicus

— Dactyl of right chela with row of long spines on dorsomesial surface; 2 or 3 longitudinal

rows of well-spaced, often acute tubercles on dorsal surface. Spines of carpi of third

pereopods strongly developed . D. eephyma sp. nov.

4. Ocular peduncles short (Fig. 2c), moderately stout; antennal peduncles reaching beyond

bases of corneas. Dorsal surfaces of palms of both chelas with numerous moderately

closely-spaced quite small tubercles or spinules . D. richeri sp. nov.

— Ocular peduncles long (Fig. 2e), moderately slender; antennal peduncles not usually

reaching to bases of corneas. Dorsal surfaces of palms of both chelae with relatively few,

widely-spaced irregular longitudinal rows of moderately large tubercles .

. D. spinulimanus

1 . Corneas slightly dilated. Ratio of peduncular length to corneal diameter between 3.5 and

4:1 . D. spinulimanus

— Corneas moderately or strongly dilated. Ratio of peduncular length to corneal diameter

less than 3:1 . 2

2. Ratio of peduncular length to corneal diameter between 2: 1 and 2.5:1 . 3

— Ratio of peduncular length to corneal diameter less than 2:1 . 4

3. Telson longer than broad. Palm of right chela strongly asymmetrical; strong spines on

mesial margin and on distal half of lateral margin . D. eephyma sp. nov.

— Telson as long as broad. Palm of right chela almost symmetrical; both margins with

regular spines of moderate size . D. ophthalmicus

4. Ocular acicles unidentate. Telson longer than broad. Maximum width of palm of right

chela more than 1.25 times length of dorsomesial margin. Antennal peduncles equal to or

slightly longer than ocular peduncles . D. rubricatus

— Ocular acicles usually bidentate (at least on 1 side). Telson as long as broad. Maximum

width of palm of right chela only slightly greater than length of dorsomesial margin.

Antennal peduncles shorter than ocular peduncles . D. richeri sp. nov.

Diacanthurus eephyma sp. nov.

Figs 2a, 3a-i

Material EXAMINED. — New Caledonia. Chalcal: stn D 65, 22°11.50'S, 159°15.40’E, 305 m, 27.07.1984:

1 6 (4.4 mm) (MNHN Pg 5479).

Smib 4: stn DW 43, 24°46.6'S, 168°08.8’E, 245 m, 08.03.1989: 2 8, 1 9 (3.6-5.6 mm) (USNM 276062).

Beryx 1 1: stn 18. 24°48'S, 168°09'E, 250-270 m, 16.10.1992: 1 9 (4.2 mm) (MNHN Pg 5478).

Source MNHN \ Paris

CRUSTACEA DECAPODA: DIACANTHURUS NEW GENUS OF HERMIT CRABS

239

Fig. 1. — a, Diacanthurus ophthalmicus (Ortmann. 1892), 9 holotype (6.2 mm) from Sagami Bay, Japan: telson. —

b-e, Diacanthurus spinulimanus (Miers, 1876), 8 (8.7 mm) from Paria Rock, Hawks Bay, New Zealand: b. maxillule

(external view); c, maxilla (external view); d first maxilliped (external view); e, third maxilliped (internal view).

Scales equal 1.0 mm (a-d) and 3.0 mm (e).

SMIB 8: stn DW 155, 24°45'S, 168°08'E, 257-262 m. 28.01. 1993: I 8 (5.4 mm) (MNHN Pg 5480). — Stn DW 159,

24°46'S, 168°08'E, 241-245 m. 28.01.1993: 1 8 (2.8 mm). — Stn DW 170, 23°4I-S. 168°01'E. 241-244 m, 29.01.

1993: 1 ov. 9 (5.8 mm) (MNHN Pg 5481).

Bathus 4: sin DW 904, 18°59'S, 163°1 I E. 461 m, 04.08.1994: I ov. 9 (8.9 mm) (MNHN Pg 5483). — Stn DW 924,

1 8°54'S, 163°24'E. 344-360 m, 07.08.1994: 2 ov. 9 (7.6, 7.9 mm) (MNHN Pg 5484). — Stn DW 925. 18°54'S.

163°23'E, 370-405 m. 07.08.1994: I 8. 2 9 (4.6-8.9 mm) (MNHN Pg 5485). — Stn DW 926, 18°56’S. I63°25'E. 325-

330 m. 07.08.1994: 1 9 (4.0 mm) (MNHN Pg 5486). — Stn CP 936, 19°03'S. I63°28'E, 258-252 m. 08.08.1994: 2 6

(8.3, 8.5 mm) (MNHN Pg 5487). — Stn CP 939, 18°58’S, I63°25'E, 304-320 m. 08.08.1994: 1 8 (9.2 mm) (MNHN Pg

5488). — DW 942, 19°04'S, 163°28'E, 270-264 m, 08.08.1994: 1 8 (4,9 mm) (MNHN Pg 5489).

Halical 1: stn DW 01, 18°56'S, 163°24'E, 380-400 m. 23.1 1.1994: 2 9 (6.8. 8.7 mm) (MNHN Pg 5477).

Source .

240

p.a. mclaughlin & j. forest

Smib 10: stn DW 208. 24°49'S. 168°09'E, 270 m. 10.01.1995: 1 6 (4.6 mm) (MNHN Pg 5482). — Stn DW 209,

24°49’S, 168°09,E, 329-560 m, 10.1.1995: I 6 , 1 9 (4.5. 4.8 mm) (USNM 276063).

Chesterfield Islands. MUSORSTOM 5: no other data: 1 6 , 1 ov. 9 (4.1. 5.1 mm) (MNHN Pg 5490). — Stn DW

263, 25°22’S, 159°47'E, 150-225 m, 08.10.1986: 3 6 . 1 ov. 9 (4.7-5. 9 mm) (USNM 276064). — Stn CP 275,

24°46.60'S, 159°40.30’E, 285 m, 09.10.1986: 1 6 (7.9 mm) (MNHN Pg. 5491 ).

Western Australia. H.M.A.S. Diamantina: DM. 1/72, Stn 32, 32°15'S. 1I5°07E, 210-212 m, 17.03.1972:

1 ov. $ (6.3 mm) (WAM 1708.86).

Non paratype. New Caledonia. Smib 8: stn DW 158, 24°46'S. 168°02'E, 262-290 m, 28.01.1993: 1 6 , no

appendages (5.0 mm) (MNHN Pg 5492).

Types. — The male (8.9 mm) (MNHN Pg 5485) from Bathus 4, station DW 925 is the holotype. All hut

one of the other specimens are paratypes.

Description. — Shield (Fig. 2a) width equal to length or slightly longer than broad, dorsal surface moderately

smooth, with row of tufts of stiff setae laterally delimiting gastric region. Rostrum obsolete, rostral region rarely

produced beyond level of obtusely rounded and unarmed or weakly spined lateral projections; anterior margin

between rostral and lateral projection regions concave; anterolateral margin also slightly concave, anterolateral

angle obtuse; posterior margin truncate, with slight median concavity.

Ocular peduncles moderately long, 0.80-0.95 length of shield, slightly broader at bases of corneas; maximum

corneal width included twice to 2.5 times in peduncular length; tuft of stiff setae in dorsal notched area of cornea,

and usually longitudinal row of sparse tufts of setae on dorsal surface. Ocular acicles triangular, terminating

subacutely but with very strong submarginal spine.

Antennular peduncles overreaching distal margin of cornea by 0.15 to 0.50 length of ultimate segment.

Ultimate segment with few long setae in quasi longitudinal row on dorsal surface. Penultimate segment with

few setae. Basal segment with acute spine laterally in distal third; row of stiff setae dorsally, distally and ventrally.

Antennal peduncle not reaching distal margin of cornea, but often reaching beyond corneal base. Fifth and

fourth segments with rather long stiff setae dorsally and ventrally. Third segment with very long slender spine at

vcntrodistal angle practically obscured by long stiff setae. Second segment with dorsolateral distal angle produced,

terminating in strong slender simple or bifid spine, reaching to or beyond proximal margin of fourth peduncular

segment, mesial margin with 1 to 3 acute spines and often 1 small spine just beneath terminal spine; dorsomesial

distal angle with prominent spine. First segment with 1 or 2 strong spines on ventral distal margin. Antennal

acicle long, slender, reaching nearly to distal margin of Fifth peduncular segment, terminating in unequally bifid

spine or simple spine with 1 smaller accessory spine. Antennal flagellum moderately long; 2 or 3 setae every 4-6

articles.

Right chelipcd (Fig. 3a) moderately long and slender; maximum width of palm only slightly longer than

dorsomesial margin. Dactyl slightly longer than palm, slightly to considerably overlapped by fixed finger, with

slender hiatus, at least in proximal half; dactyl subtriangular in cross-section, dorsal surface roundly elevated in

midline and covered with irregular rows of low blunt, but very distinct tubercles, dorsomesial margin with row of

rather widely-spaced small to moderately strong spines; cutting edge of dactyl with 3 distinct or partially fused

calcareous teeth proximally and several small to moderately large more distinct molar-like teeth in distal half,

terminating in calcareous claw; mesial face rounding into ventral surface and provided with tufts of long stiff setae.

Palm equal to or shorter than carpus; dorsomesial margin with row of small to large, often slender acute or

subacute spines, dorsomesial distal angle with cluster of low but distinct tubercles; dorsal surface of palm and fixed

finger covered, but not densely, with small conical spinules or subacute tubercles, usually not obscured by mat of

very short fine setae (Fig. 3b), frequently 1 sparse row of slightly stronger conical tubercles in midline of palm,

dorsoproximal margin sometimes with 1 or 2 moderate to very prominent spines medially; dorsolateral margin

with row of usually very much smaller spines on palm becoming considerably stronger on fixed finger; cutting

edge of fixed finger writh row of distinct or somewhat fused calcareous teeth proximally, several more distinct teeth

distally, terminating in calcareous claw; inner faces of dactyl and fixed finger each often with several additional

calcareous teeth; mesial face of palm relatively smooth or with low protuberances and moderately long and stiff

setae, ventral surface and lateral face both with rows of tufts of setae. Carpus about as long as merus; dorsomesial

margin with row of long, slender, very acute spines and adjacent second smaller and somewhat irregular row;

Source :

CRUSTACEA DECAPODA: DIACANJ'HURUS NEW GENUS OF HERMIT CRABS

241

Fig. 2. — Shield and cephalic appendages.

a. Diacanthurus ecphyma sp. nov., 6 (5.9 mm) from MUSORSTOM 5 Stn DW 263. — b. Diacanthurus ophthalmicus

(Ortmann, 1892), ov. 2 (6.4 mm) from Minabe. Wakayama, Japan. — c, Diacanthurus richeri sp. nov., 6 (5.9 mm)

SMIB 8 Stn DW 190. — d, Diacanthurus rubricatus (Henderson, 1888), $ (8.4 mm) from NZOl Stn D 121. -

e, Diacanthurus spinulimanus (Miers, 1876), 2 (12.0 mm) from Paria Rock, Hawks Bay. New Zealand.

Scales equal 5.0 mm.

242

p. a. McLaughlin & j forest

dorsodistal margin with spine mesially, surface also with mat of short fine setae, midline with row of much

smaller spines and transverse rows of longer setae extending laterally, dorsolateral margin not delimited; lateral face

with longitudinal rows of tufts of long setae, ventrolateral margin with row of spines; ventrodistal margin also

with few spines, ventromesial margin with tufts of long setae. Merus (Fig. 3c) with 1 very strong acute spine on

dorsodistal margin; short transverse rows of long stiff setae on mesial and lateral faces; ventrolateral margin with 3

to several slender spines, occasionally fused to form broad single trifid spine, ventral surface with transverse

irregular rows of moderately strong spines, ventromesial margin with long stiff setae distal ly, strong spines

proximally. Ischium with row of small spines on ventromesial margin.

Left cheliped (Fig. 3d) somewhat shorter than right, much less robust; propodal-carpal articulation rotated

clockwise approximately 30° from horizontal plane; dactyl and fixed finger nearly twice length of palm. Dactyl

with convex dorsal surface, single or irregularly double row of very small spinules in dorsal midline at least

proximally, row of tufts of stiff setae near cutting edge, row of longer stiff setae on dorsomesial margin; cutting

edge with row of very fine corneous teeth, terminating in prominent corneous claw; mesial and ventral surfaces

with longitudinal rows of tufts of stiff setae. Palm half or slightly less than half length of carpus; dorsal surface

convex in midline; row of prominent spines, sometimes interspersed with smaller spines or spinules, on elevated

dorsolateral margin, decreasing in size on convex margin of fixed finger; dorsal surface covered, but often not

densely, with regularly placed small conical tubercles, usually lower and Hatter on fixed finger; dorsomesial margin

of palm frequently with 1 or 2 small spines proximally, occasionally nearly complete row of widely-spaced small

spines; ventral surfaces of palm and fixed finger with tufts of long stiff setae. Carpus with row of strong spines on

both dorsomesial and dorsolateral margins, prominent spinose projection on distal margin overlapping palm;

mesial and lateral faces with numerous transverse rows of long stiff setae; ventrolateral margin with few spines at

least distally. Merus with strong spine at dorsodistal margin, transverse rows of long stiff setae dorsally, mesially

and laterally; ventrolateral and ventromesial margins each with 1 or 2, sometimes complete row of prominent,

slender acute spines; ventral surface usually spinulose or with several prominent spines and tufts of long stiff

setae. Ischium with row of smaller, subacute spines on ventromesial margin.

Ambulatory legs (Figs 3e-h) with right pair longer than left and slightly overreaching tip of right cheliped.

Dactyls moderately long, 1.35 to nearly twice length propodi, with slight distal twist and slight ventral curve;

dorsal surfaces each with nearly triple row of long spiniform bristles or corneous spines, extending slightly onto

mesial faces dorsally; mesial face with row of corneous spines ventrally and additional row on ventral surface;

dorsal and ventral surfaces also with tufts of long stiff setae. Propodi approximately same length as carpi, with

tufts of long stiff setae arising from low protuberances dorsally and ventrally. Carpi each with row of spines on

dorsal surface and few tufts of setae. Meri of third pereopods unarmed or with 1 spine on ventral margin; second

pereopods each with single or double row of spines on ventral margin; tufts of long setae dorsally and ventrally on

both. Ischia with 1 or 2 small spines (second) or unarmed (third). Sternite of third pereopods with anterior lobe

semisubcircular.

Male pleopods usually with endopod of third reduced or rudimentary, fourth vestigial, and Fifth absent. Telson

(Fig. 3i) with lateral angles of posterior lobes each produced as prominent spine; strongly concave inner margins

each with 1 well developed spine on either side of moderately wide median cleft, I or 2 tufts of setae occasionally

accompanied by small spine or spinule in distal half.

Color. — In preservative: Dactyl, fixed finger and dorsomesial surface of palm of right cheliped red-orange.

Dorsal surtaces ol dactyl and fixed finger of left cheliped with white spots on background of red-orange.

Ambulatory dactyls with splotches of reddish-orange on white background, and median longitudinal reddish-orange

stripe.

Etymology. — A noun in apposition, from the Greek, ekphyma , meaning an eruption of pimples, and

referring to the pimple-like appearance of the left chela.

Distribution. — New Caledonia and Chesterfield Islands; Western Australia; 212-461 m (at least).

Remarks. — Diacanthurus eephyma bears a strong resemblance to D. ophthalmicus , but is easily

distinguished from the latter species by: 1) the clearly distinct rows of subacute to blunt tubercles on the dorsal